< Digitized by the Internet Archive in 2017 with funding from IMLS LG-70-15-0138-15 https://archive.org/details/pharmaceuticalbo00youn_0 PHARMACEUTICAL BOTANY YOUNGKEN Dryopteris marginalis, one of the ferns whose rhizome and stipes constitute the drug, Aspidium. ( Frontispiece ) PHARMACEUTICAL BOTANY I BY HEBER. W. YOUNGKEN, Ph.G., A.M., M. S., Ph.D. HEAD OF THE DEPARTMENT OF BOTANY AND PHARMACOGNOSY IN THE PHILADELPHIA COLLEGE OF PHARMACY; MEMBER OF THE AMERICAN PHARMACEUTICAL ASSOCIATION, ACADEMY OF NATURAL SCIENCES OF PHILADEL- PHIA, BOTANICAL SOCIETY OF , AMERICA, BOTANICAL SOCIETY OF PENNSYLVANIA, AMERICAN ASSO- CIATION FOR THE ADVANCEMENT OF SCIENCE, ETC. SECOND EDITION, REVISED AND ENLARGED WITH 195 ILI nsTBATinN5 Li PHILADELPHIA P. BLAKISTON’S SON & CO. 1012 WALNUT STREET ' 3 H 1 — Copyright, 1918, by P. Blakiston’s Son & Co. “The use in this volume of certain portions of the text of the United States Pharmacopoeia is by virtue of permission received from the Board of Trustees of the United States Pharmacopoeial Convention. The said Board of Trustees is not responsible for any inaccuracy of quotation nor for any errors in the statement of quantities or percentage strengths.” T II K MAPLE PKESS TO K K I* A PREFACE TO THE SECOND EDITION The appearance of the United States Pharmacopoeia IX and the National Formulary IV, with the many changes in the lists and definitions of officially recognized vegetable drugs made it necessary to revise the former edition of this work. In the course of revision, the writer has taken cognizance of the growing importance of Botany in the curricula of pharmaceutical institutions and has accordingly expanded upon the subject matter of the former text. With the adoption for the first time by the new United States Pharmacopoeia of pharmacognic standards for numerous drugs, Pharmacognosy has risen to the forefront in this country as a science. While its proper comprehension requires laboratory instruction in chemistry, physics, and crystallography as well as botany, neverthe- less a rather extended foundation in structural botany stands out preeminently as the most needed requirement. The work has been for the most part remodeled. Chapter I deals with Fundamental Considerations. Chapter II is devoted to the life history of the Male Fern, a median type of plant, the considera- tion of which, after the students have received fundamental practice in the use of the microscope, the writer has found commendable, for it not only gives the beginner a working knowledge of structures and functions, the homologies and analogies of which will be met in the later study of forms of higher and lower domain, but holds their interest on account of its economic importance. The life history of a type of Gymnosperm, White Pine, is next taken up in Chapter III. Chapter IV considers the life history of an Angiosperm as well as codrdinates the resemblances and differences between Gymnosperms and Angiosperms. Chapters V, VI and VII are devoted respectively to Vegetable Cytology, Plant Tissues and Plant Organs and Organisms. Among the many additions to the topics included in these might be mentioned a treatise on Cell Formation and Reproduction including Indirect Nuclear Division, VI PREFACE TO SECOND EDITION twenty pages on Non-Protoplasmic Cell Contents, the considera- tion of Woods, Root Tubercles, the gross structure and histology of different types of leaves, broad histologic differences between Mono- cotvl and Dicotyl leaves, the histology of floral parts and the his- tology of types of fruits and seeds. Chapter VIII on Taxonomy has been increased by the addition of 144 pages. Several new fami- lies of drug-yielding plants have been added and the treatment of family characteristics has in the majority of instances been broad- ened. The habitats of drug-yielding plants have been added. In that portion of the tables dealing with the names of official drugs, those official in the National Formulary have been so designated by the abbreviation N. F., to distinguish them from others that may occur in the same portion of the table and which are official in the Pharmacopoeia. In keeping with the increased size of the book, many new illus- trations have been introduced. A number of these are original draw- ings, photographs and photomicrographs. To the authors of other works from which cuts were borrowed, the writer’s thanks are due. The writer in conclusion desires to thank Dr. John M. Macfarlane, head of the Botanical Department of the University of Pennsylvania, for valuable suggestions during the preparation of portions of the text. H. W. Y. PREFACE TO THE FIRST EDITION The aim has been to eliminate from this book all those topics that are of minor importance to the student and practitioner of Phar- macy. As a pharmacist and teacher, the writer feels that the botan- ical preparation for Pharmacognosy and Materia Medica, in those colleges where Botany is given for one year, should include mainly the structural and systematic aspects of the science. In the Medico- Chirurgical College, of Philadelphia, Botany is taught the first year, extending over a period of 155 hours. The author has introduced in this concise volume the important subject matter of his lectures given to first year students, and has omitted laboratory directions for the obvious reason that fixed subjects for laboratory study are unneces- sary. It is not a book on Pharmacognosy, however, since it does not describe how one drug differs from another of the same group in all of its details. The work is included in two parts. Part I is largely devoted to the morphology (gross and minute) and, to a less extent, the physiology of the Angiosperms. Part II deals with the taxonomy of plants, mainly but not wholly of medicinal value, together with the parts used and the names of the official and non-official drugs obtained from these. The author does not claim sole originality for the facts presented, but has consulted many sources of information, mention of which will be found in the bibliography of the text. Acknowledgment is here rnade to his esteemed friends, Dr. Fran- cis E. Stewart of the Medico-Chirurgical College and Dr. John M. Macfarlane of the University of Pennsylvania, for valuable assis- tance in the reading of the proofs and preparation of the index. H. W. Y. Philadelphia. CONTENTS CHAPTER I Fundamental Considerations DEPARTMENTS OF BOTANICAL INQUIRY.— i. Morphology Pages (Gross Anatomy, Histology, Cytology). 2. Physiology. 3. Taxonomy or Systematic Botany. 4. Ecology. 5. Phyto- pathology. 6. Phytogeography. 7. Phytopalaeontology. 8. Etiology. 9. Economic Botany and its subdivisions 1-2 PRINCIPLES OF CLASSIFICATION.— Natural System: species; genus; family; order; class; division; variety; race; hybrid 2-3 OUTLINE OF PLANT GROUPS 3-4 THE MICROSCOPE. — Definition. The simple microscope. The com- pound microscope: its construction and use. Rules for the care of the microscope 4-8 THE TECHNIQUE OF MAKING PERMANENT MOUNTS.— The mounting medium; Staining. Method for the Preparation of a Canada Balsam Mount. Method for the Staining and Mount- ing of Material in Paraffine Ribbons affixed to slide. Making sections. Kinds of sections . 8-1 1 MICROMETRY. — Unit of microscopical measurement. Standardi- zation of the Ocular Micrometer 11-12 CHAPTER II Life History of the Male Fern HISTORY OF THE SPOROPHYTE OR ASEXUAL GENERATION. — Gross structure of stem. Histology of mature stem. His- tology of growing apex. Histology of mature root. Histology of root apex. Continuity of crude sap flow. Histology of stipe. Histology of lamina. Comparative physiology of root, stem IX X CONTENTS and leaf. Gross structure and histology of the sori and sporan gia. Rupture of sporangium and spore dissemination. HIS TORY OF THE GAMETOPHYTE OR SEXUAL GENERA TION. — Origin of new sporophyte or diploid plant from fer tilized egg. Growth of seedling into mature sporophyte Alternation of generations CHAPTER III Life History of a Gymnosperm (Pinus Strobus) DESCRIPTION OF THE WHITE PINE TREE (SPOROPHYTE).— Staminate cones. Carpellate cones. DESCRIPTION OF THE GAMETOPHYTE GENERATION.— The Male Game- tophyte. The Female Gametophyte. Fertilization. Seed Formation and Distribution. GERMINATION OF THE SEED 24-31 CHAPTER IV Life History of an Angiosperm DESCRIPTION OF THE DOG’S TOOTH VIOLET.— Development of Female Gametophyte. Pollination and Fertilization. Maturation of the Pollen Grain and formation of the Male Gametophyte. Ripening of the Ovule to form the Seed and of the Ovary to form the Fruit. Germination of the Seed and development of the Mature Sporophyte 32-36 RESEMBLANCES BETWEEN GYMNOSPERMS AND ANGIO- SPERMS 36 FUNDAMENTAL DIFFERENCES BETWEEN GYMNOSPERMS AND ANGIOSPERMS 36-37 CHAPTER V Vegetable Cytology VEGETABLE CYTOLOGY.— Definition. The Plant Cell as the Fundamental Unit. A typical plant Cell 38-39 PROTOPLASM AND ITS PROPERTIES.— Structure. Metabolism. Reproduction 39 ~ 4 ° PROTOPLASMIC CELL CONTENTS. — Cytoplasm; nucleus; nucleolus; plastids (leucoplastids. chloroplastids, chromoplastids) . . . 40-41 Pages 13-23 CONTENTS XI Pages CELL FORMATION AND REPRODUCTION.— Asexual and sexual cells. Reproduction defined. Asexual Reproduction: Fission; Gemmation; Free Cell Formation; Rejuvenescence. Sexual Reproduction: Conjugation and Fertilization 41-43 INDIRECT NUCLEAR DIVISION 43~45 NON-PROTOPLASMIC CELL CONTENTS.— Sugars. Starch. Inulin. Hesperidin. Strophanthin. Salicin. Saponin. Coniferin. Digi- toxin. Characteristics of Glucosides. Alkaloids and their pro- perties. The alkaloids, Strychnine, Veratrine, Nicotine, Caffeine, Cocaine, Aconitine, Colchicine. Gluco-alkaloids. Asparagine. Calcium Oxalate. Cystoliths. Silica. Tannins. Proteins. Aleurone Grains. Mucilages and Gums. Fixed Oils and Fats. Volatile Oils. Resins. Oleoresins. Gum Resins. Balsams. Pigments. Latex. Enzymes. Classification of Enzymes 45-64 CELL WALLS. — Their formation and composition. Growth in area and thickness. Various kinds of cell walls and behavior of each to micro-chemic reagents 64-66 CHAPTER VI Plant Tissues PLANT TISSUES. — Tissue defined. The Tissues of Spermatophytes and Pteridophytes: Generative Tissues. List of Tissues . . . 67-68 MERISTEM. — Definition. Primary and secondary meristems; their distribution 68 PARENCHYMA. — Definition; ordinary parenchyma; assimilation parenchyma; conducting parenchyma; reserve parenchyma; their distribution and function 69 COLLENCHYMA. — Definition, function and distribution 70 SCLERENCHYMA. — Definition; stone cells; sclerenchyma fibers, wood fibers; bast fibers; their distribution 70-71 EPIDERMIS. — Definition; transpiration and water stomata; epidermal papillae; trichomes; scales; their distribution and functions . . 71-75 ENDODERMIS. — Definition, distribution and functions 75 CORK. — Definition; its derivation, function and distribution 75 LATICIFEROUS TISSUE. — The structure, origin, distribution of latex cells, laticiferous vessels and secretory cells. Latex 75—7 7 SIEVE (LEPTOME OR CRIBIFORM) TISSUE— Definition; distri- bution; function 77 — 78 XII CONTENTS Pages TRACHEARY TISSUE.— Tracheae: Definition, function and classifica- tion; Tracheids: Definition and function 78-80 MEDULLARY RAYS. — Primary and Secondary; their distribution and functions 81 FIBRO-VASCULAR BUNDLES. — Definition of the various types; the distribution of each type; Xylem and Phloem 81-8,} SECRETION SACS. — Definition and distribution 83 INTERCELLULAR AIR SPACES. — Definition ; schizogenous and lysi- genous air spaces; their relative size 83 SECRETION RESERVOIRS. — Definition; structure; internal glands; secretion canals 84 CLASSIFICATION OF TISSUES ACCORDING TO FUNCTION. . 84 CHAPTER VII Plant Organs and Organisms PLANT ORGANS AND ORGANISMS.— Organ and organism defined. Vegetative Organs: Roots, stems and leaves. Reproductive Organs: Flower, fruit and seed. Embryo and its parts; function of the cotyledon 85 THE ROOT. — Definition; functions; root hairs; root cap; generative tissues; differences between root and stem ' 85-87 CLASSIFICATION OF ROOTS AS TO FORM— Primary root; tap root; secondary roots; fibrous and fleshy roots; anomalous roots; adventitious roots; epiphytic roots; haustoria 87-88 CLASSIFICATION OF PLANTS ACCORDING TO DURATION OF ROOT. — Annual; Biennial; Perennial 88 ROOT HISTOLOGY. — A. Monocotyledons. B. Dicotyledons; Histology and Development of the California Privet root; Abnormal structure of Dicotyl roots; Histology of a Dicotyl Tuberous Root, Aconitum 88-93 ROOT TUBERCLES. — -Definition; occurrence on roots of Leguminosae, Myricaceae, etc.; their etiology 93-96 THE BUD. — Definition; plumule, scaly buds, naked buds; Classification of Buds According to Position on Stem: terminal bud; axillary or lateral bud ; adventitious bud ; accessory bud. Classi- fication of Buds According to Development: leaf bud; flower bud and mixed bud. Classification of Buds According to their Arrangement on the Stem: alternate; opposite; whorled . 96-97 CONTENTS Xlll Pages THE STEM. — Definition; direction of growth; functions; size; nodes and internodes; stem elongation; duration of stems; stem modifi- cations; above ground stems; herb and tree defined; under- ground stems; exogenous and endogenous stems 97-101 STEM HISTOLOGY/ — Annual Dicotyl; Perennial Dicotyl; Exceptional Types of Dicotyl Stems; Lenticels and Their Formation; Annual Thickening of Stems; Method of formation; “Annual . Ring.” Bark: Definition; zones; Periderm; Phelloderm; Histology of Cascara Sagrada bark. Wood: alburnum; dura- men; microscopic characteristics of Angiospermous and Gym- nospermous woods; Histology of Typical Herbaceous Mono- cotyl Stems; Histology of Typical Woody Monocotyl Stem . . 101-115 THE LEAF. — Definition; leaf functions: photosynthesis; assimilation; respiration; transpiration 116 TYPES OF LEAVES DEVELOPED IN ANGIOSPERMS— Cotyle- dons; Scale leaves; Foliage leaves; Bract leaves: bracts and bracteoles; Sepals; Petals; Microsporophylls; Megasporophyllsn6-ii7 ORIGIN AND DEVELOPMENT OF LEAVES.— Primordial leaf. Its formation 117 PHYLLOTAXY. — Definition; spiral law of leaf arrangement; alternate; opposite, decussate; whorled; fascicled; leaf rank 117-118 VERNATION. — Definition; inflexed or reclinate; conduplicate; con- volute; circinate; plicate; involute; revolute 118-119 THE COMPLETE LEAF. — Its parts; sessile; petiolate; exstipulate; stipulate 1 19-120 LEAF VENATION. — Furchate; parallel; reticulate; pinni- veined; palmately veined; anastomosing veins 120 LEAF INSERTION. — Definition; radical; cauline; ramal; perfoliate; amplexicaul; connate-perfoliate; equitant 120-122 FORMS OF LEAVES.- — Simple and Compound. ( a ) General Outline: ovate; linear; lanceolate; elliptical; oblong; inequilateral; or- bicular; peltate; filiform; oblanceolate; cuneate; spatulate; ensi- form; acerose; deltoid. ( b ) Apex: acute; acuminate; obtuse; truncate; mucronate; cuspidate; aristate; emarginate; retuse; obcordate. (c) Base: cordate; reniform; hastate; auriculate; sagittate. ( d ) Margin: entire, serrate; dentate; crenate; repand; sinuate; incised; runcinate; lobed; cleft; parted; divided. Forms of Compound Leaves 122-127 LEAF TEXTURE. — Membranous; succulent; coriaceous 127 XIV CONTENTS Pages LEAF COLOR. — Variations in color 127 LEAF SURFACE. — Glabrous; glaucous; pellucid-punctate; scabrous; pubescent; villose; sericious; hispid; tomentose; spinose; rugose; verrucose 127 DURATION OF LEAVES. — Persistent or evergreen; deciduous; cadu- cous; fugacious 127 GROS S STRUCTURE AND HISTOLOGY OF THE LEAF PETIOLE. — In Monocotyledons. In Dicotyledons. Pulvinus. Periclad- ium 127-129 STIPULES. — Definition; lateral; free-lateral; lateral-adnate; lateral- connate; lateral interpetiolar. Axillary; ochrea. Modified Stipules 1 29-130 THE LAMINA. — Definition. Mode of Development of the Lamina of Leaves: Dorsoventral; convergent; centric; bifacial; reversed; ob-dorsoventral. A. Dorsoventral: (a) Dorsoventral Umbro- phytic; ( b ) Dorsoventral Mesophytic; (c) Dorsoventral Xero- phytic; (d) Dorsoventral Hydrophytic. Gross Structure and Histology of Different Types of Dorsoventral Leaf Blades. Gross Structure and Histology of the following types: B. Convergent; C. Centric; D. Bifacial 130-^34 STRUCTURE AND DEVELOPMENT OF STOMATA 134-136 HISTOLOGIC DIFFERENCES BETWEEN LEAVES OF DICOTY- LEDONS AND MONOCOTYLEDONS 136 INFLORESCENCE. — Definition; Parts of Inflorescences; Determinate; Indeterminate; Mixed Forms of Indeterminate and Determi- nate Inflorescences 136-140 PR.EFLORATION.- — Convolute; involute; revolute; plicate; imbricate; valvate; vexillary; contorted 140-141 THE FLOWER. — Definition; floral parts; essential organs; complete; perfect; hermaphrodite; regular; symmetrical; imperfect; double; staminate; pistillate; neutral; connation; adnation . .141-143 THE RECEPTACLE, TORUS OR THALAMUS.— Definition; varia- tions in structure; anthophore; gonophore; gynophore; carpo- phore 143 THE PERIGONE. — Definition; dichlamydeous; monochlamydeous . . 143 THE CALYX. — Definition; parts; physical characteristics; connation; kinds and forms; persistence; adnation; sepaline spurs; sepaline stipules; sepaline position 143-T45 CONTENTS XV Pages THE COROLLA. — Definition; parts; physical characteristics; func- tions. Forms of the Corolla and Perianth 145-147 THE ANDRCECIUM OR STAMEN SYSTEM.— Definition; parts; terms denoting number of stamens in flower; insertion of sta- mens; proportions of stamens; connation of stamens; stamen color. Gross Structure and Histology of the Filament. Gross Structure and Histology of the Anther. Anther Dehiscence. Development of the Anther. Attachment of Anther. Pollen: description; forms 147-154 THE GYNCECIUM OR PISTIL SYSTEM.— Definition; Gymno- spermous and Angiospermous; parts; the pistil a modified leaf; carpel; dehiscence; apocarpous and syncarpous pistils; terms denoting the number of carpels entering into the formation of the pistil; compound pistils; ovules or megasori as trans- formed buds; position of ovules in ovary; Gymnospermous and Angiospermous ovules; structure of Angiospermous ovule; shape of ovule 154-156 THE PLACENTA. — Definition; types of placenta arrangement. . . . 156 THE STYLE. — Definition; style-arms; relation to carpels forming the gynoecium; variations from typical stylar development . . .156-157 THE STIGMA. — Definition; forms in wind- and animal-pollinated flow- ers; stigmatic papillae 157-158 POLLINATION. — Definition; Close and Cross Pollination; terms applied to plants pollinated by various agencies 158 MATURATION OF THE POLLEN GRAIN AND FORMATION OF MALE GAMETOPHYTE 158-159 MATURATION OF THE EMBRYO SAC AND FORMATION OF THE FEMALE GAMETOPHYTE ico FERTILIZATION IN ANGIOSPERMS. — Process; formation of embryo and endosperm THE FRUIT. — Definition; modifications 160 FRUIT STRUCTURE. — Pericarp; pseudocarp; anthocarp; epicarp; mesocarp; endocarp; sarcocarp; putamen; sutures; valves; dehiscence r 60-1 61 CLASSIFICATION OF FRUITS. — Simple; Aggregate; Multiple; dry dehiscent; dry indehiscent; fleshy indehiscent. Forms of Simple Fruits: capsular; schizocarpic; achenial; baccate; drup- aceous. Etaerio. Forms of Multiple Fruits: strobile or cone; sorosis; syconium; galbalus 161-166 XVI CONTENTS Pages HISTOLOGY OF A CAPSULE, VANILLA 166 HISTOLOGY OF A MERICARP, FCENICULUM 167-168 THE SEED. — Definition; structure; functions; appendages . . : . .168-169 MODE OF FORMATION OF DIFFERENT TYPES OF ALBUMEN. — Perisperraic; endospermic; perispermic and endospermic; exalbuminous and albuminous seeds 169-170 A MONOCOTYL SEED. — Its gross structure and histology 170-171 A MONOCOTYL SEEDLING. — Germination and structure 1 71-172 A DICOTYL SEED. — Its gross structure and histology 1 72-1 73 CHAPTER VIII Taxonomy THALLOPHYTA. — Definition. Characters of the Protophyta, Myxo- mycetes, Algae, Fungi and Lichenes. The Mounting and Staining of Bacteria. Life histories of representative types of Thallophytes. Plants of the group yielding drugs and economic products 174-230 BRYOPHYTA. — Definition. Characters of the Hepaticae and Musci. Life history of a typical true moss 231-236 PTERIDOPHYTA. — Definition. Characters of the Lycopodineae, Equisetineae and Filicineae. Life history of a typical fern. Plants of the group yielding official drugs 236-241 SPERMATOPHYTA (PHANEROGAMIA).— Definition. Charac- teristics of the Gymnosperms and of the Order and Family yielding important drugs and economic products. Table of official and unofficial drugs yielded by Gymnosperms including part of plants used, botanical origins and habitats. Character- istics of the Angiosperms and of the classes and families yield- ing drugs. Tables of official and unofficial drugs with the names of the plants, parts constituting the drugs and habitats . 241-355 BIBLIOGRAPHY 356 INDEX 357 PHARMACEUTICAL BOTANY CHAPTER I [FUNDAMENTAL CONSIDERATIONS [Botany is the Science which Treats of Plants DEPARTMENTS OF BOTANICAL INQUIRY 1. Morphology treats of the parts, or structures of plants. It is divided into: (a) Macromorphology or Gross Anatomy which deals with the ex- ternal characters of plants or their parts; ( b ) Micromorphology or Histology which considers the minute or microscopical structure of plants and plant tissues; and (c) Cytology which treats of plant cells and their contents. 2. Physiology deals with the study of the life processes or func- tions of plants. It explains how the various parts of plants perform their work of growth, reproduction, and the preparation of food for the support of animal life from substances not adapted to that use. 3. Taxonomy or Systematic Botany considers the classification or arrangement of plants in groups or ranks in accordance with their relationships to one another. 4. Ecology treats of plants and their parts in relation to their environment. 5. Phytopathology treats of diseases of plants. 6. Phytogeography or Plant Geography treats of the distribution of plants upon the earth. The center of distribution for each species of plant is the habitat or the original source from which it spreads, often over widely distant regions. When plants grow in their native countries they are said to be indigenous to those regions. 2 PHARMACEUTICAL BOTANY When they grow in a locality other than their original home they are said to be naturalized. 7. Phytopalaeontology or Geological Botany treats of plants of former ages of the earth’s history traceable in their fossil remains. 8. Etiology is the study of the causes of various phenomena ex- hibited by plants. 9. Economic or Applied Botany deals with the science from a prac- tical standpoint, showing the special adaptation of the vegetable kingdom to the needs of everyday life. It comprises a number of subdivisions, viz.: Agricultural Botany, Horticulture, Forestry, Plant Breeding, - and Pharmaceutical Botany. Pharmaceutical Botany considers plants or their parts with reference to their use as drugs. It interlocks very closely with other departments of botanical science. PRINCIPLES OF CLASSIFICATION The classification of plants is an attempt to express the exact kinship which is believed to exist among them. By grouping to- gether those plants which are in some respects similar and combining these groups with others, it is possible to form something like an orderly system of classification. Such a system based upon natural resemblances is called a natural system. In a natural system of classification every individual plant belongs to a species, every species to a genus, every genus to a family, every family to an order, every order to a class, every class to a division. In many instances species may be subdivided into varieties or races. The crossing of two varieties or species, rarely of two genera, gives rise to a hybrid. Thus, the species Papaver somniferum which yields the opium of the Pharmacopoeia belongs to the genus Papaver, being placed in this genus with other species wh'ch have one or more essential character- istics in common. The genera Papaver, Sanguinaria and Cheli- donium, while differing from each other in certain essential respects nevertheless agree in other particulars such as having latex, perfect flowers, capsular fruits, etc., and so are placed in the Papaveracece family. The Papaveracece family and the Fumariacece family are closely allied, the latter only differing fiom the former in having FUNDAMENTAL CONSIDERATIONS 3 irregular petals, usually diadelphous stamens and non-oily albumen and so both of these families are placed in the order Papaverales. The orders Papaverales, Geraniales, Sapindales , Rhamnales , etc., are related by a common structure namely, two seed leaves or cotyledons and so are grouped together under the class Dicotyledonece. The Dicotyledonece differ from the Monocotyledonece in that the latter group possess but one cotyledon; but both classes agree in having covered ovules and seeds, and so are placed in the subdivision Angiospernm. The Angiospermce differ from the Gymnos per nice in that the latter possess naked ovules and seeds but both of these subdivisions agree in producing real flowers and seeds. For these reasons they are placed in the division Spermatophyta of the Vege- table Kingdom. OUTLINE OF PLANT GROUPS I. Thallophyta 4. Fungi 3. Alga; 2. Myxomycetes 1. Protophyta Bacteria Cyanophycea? Acrasiales Phytomyxales Myxogastrales Chlorophycese Phaeophyceae Rhodophyceae Phycomycetes Ascomycetes Basidiomvcetes Fungi Imperfecti Crustaceous 5. Lichenes Marchantiales II. Bryophyta 1. Hepaticae Jungermanniales . Anthocerotales 2. Musci 4 PHARMACEUTICAL BOTANY III. Pteridophyta IV. Spermatophyta 1. Lycopodineae 2. Equisetineae 3. Filicineae 1. Gymnospermae 2. Angiospermae f Lycopodiales Selaginalles 1 Isoetales { Equisetales j Ophioglossales 1 Filicales Cycadales Ginkgoales Coniferales Gnetales J Monocotyledoneae 1 Dicotyledoneae THE MICROSCOPE A microscope is an optical instrument, consisting of a lens, or combination of lenses, for making an enlarged image of an object which is too minute to be viewed by the naked eye. Microscopes are of two kinds, viz.: simple and compound. THE SIMPLE MICROSCOPE This consists simply of a convex lens or several combined into a system and mounted in a suitable stand. A good example of a simple microscope is a reading glass. THE COMPOUND MICROSCOPE A. Its Construction: The principal parts of a compound microscope are: r. The base, generally horseshoe shaped, which rests on the table. 2. The pillar, an upright bar, which is attached to the base below and supports the rest of the instrument. 3. The stage, a horizontal shelf upon which is placed the prepara- tion or slide to be examined. The stage is perforated in the center for transmitting light reflected up by the mirror. On the stage are two clips for holding the glass slide. 4. The mirror, situated below the stage, by which the light is reflected upward through the opening in the stage. FUNDAMENTAL CONSIDERATIONS 5 Draw-tube Horse-shoe base Fig. i. — I llustrating the parts of a compound microscope. 5. The diaphragm, inserted in the opening of the stage or attached to its lower face, and used to regulate the amount of light reflected by the mirror. Ocular Prism washer Inclination joint Pillar Body tube Nosepiece Rack and pinion coarse adjustment Micrometer head of fine adjustment Arm Fine adjustment pillar Clips Objectives] Stage ' Upper iris diaphragm substage ring Condenser mounting lower iris diaphragm Condenser focusing screw Mirror — Mirror fork Mirror bar 6. The body tube, a cylinder which holds the draw tube and lenses and moves up and down perpendicularly above the opening in the stage. The tube is raised or lowered either by sliding it back and forth with a twisting movement or by a rack and pinion mechanism. The latter is called the coarse adjustment. 6 PHARMACEUTICAL BOTANY 7. The fine adjustment, a micrometer screw back of the tube, which, on being turned, produces a very small motion of the entire framework which holds the body tube. 8. The oculars or eyepieces which slip into the upper end of the draw tube. Each of these consists of two plano-convex lenses, the lower one being the larger and known as the field lens because it increases the field of vision. The upper or smaller lens is called the eye lens. It magnifies the image formed by the objective. Midway between the field and eye lens is a perforated diaphragm, the object of which is to cut out edge rays from the image. According to the system adopted by the maker, oculars are desig- nated by numbers, as 1, 2, 3, 4, etc., or by figures which represent focal lengths. 9. The objectives, which screw into the bottom of the body tube or nose piece. They consist of a system of two, three or four lenses, some of which are simple, others compounded of a convex crown lens and a concave flint lens. Objectives like oculars are usually designated by numbers or by figures, as etc., or in milli- meters, which represent focal lengths. The smaller the number or fraction representing the focal length of an objective, the greater is its magnifying power. Objectives are either dry lenses or itnmersion lenses. If an air space be present between the objective and the object, the lens is called a dry one; if a liquid is present between the objective and the object, the lens is called an immersion lens. If this liquid be oil the objective is called an oil immersion objective; if water, a water immersion objective. Some microscopes are fitted up with a nose piece, capable of carry- ing two or three objectives, which may be revolved into place at the lower end of the body tube. Others have a condenser which is employed to concentrate the light upon the object examined. B. Its Use: 1. Place the microscope on the table with the pillar nearest you. 2. Screw the objectives into the nose piece and slip an ocular into the upper end, if not already on instrument. Turn the lowest power objective into position. 3. Find the light by looking into the ocular (eye piece) and at the FUNDAMENTAL CONSIDERATIONS 7 same time turning the mirror at such an angle that it reflects light from the window or lamp up through the opening in the stage to the objective. Mirrors have two faces, a plane and a concave. Use the concave unless otherwise directed. 4. Regulate the quantity of light by the diaphragm. If too bright it must be cut off somewhat. The higher powers require brighter light than the lower. 5. Place the slide on the stage in a horizontal position with the object over the middle of the opening through which light is thrown from the mirror. 6. With the lower pow’er in position, move the coarse adjustment until either the object or small solid particles on the slide appear distinctly, which means that the lenses are in focus. The object, if not under the lens, may now be brought into the field by moving the slide back and forth very slowly while looking through the ocular. To improve the focus, slowly turn the fine adjustment screw. 7. To focus with the high-power objective, first find the object with the low power and arrange in the center of the field. Put clips on slide without moving it. Raise the body tube by means of the coarse adjustment. Then turn the high-power objective into posi- tion. (If two objectives only accompany your instrument, the high- power is the longer one.) Lower the body tube carrying the objec- tive until the objective front lens nearly touches the cover glass. A slight movement of the fine adjustment should show the object clearly. Never focus down with the high-power objective because of the danger of pressing it into the cover glass and ruining the delicately mounted lenses. 8. Accustom yourself to use both eyes indifferently and always keep both eyes open. If right handed, observe with the left eye, as it is more convenient in making drawings. RULES FOR THE CARE OF THE MICROSCOPE 1. In carrying the microscope to or from your table, grasp it firmly by the pillar and hold in an erect position, so that the ocular which is fitted loosely into the draw tube may not fall out and its lenses become damaged. 8 PHARMACEUTICAL BOTANY 2. Never allow the objective to touch the cover glass or the liquid in which the object is mounted. 3. Note whether the front lens of the objective is clean before attempting to use it. If soiled, breathe on the lens and gently wipe with an old, clean, soft handkerchief or lens paper. If the lens be soiled with balsam or some other sticky substance, moisten the handkerchief or lens paper with a drop of xylol, taking care to wipe it perfectly dry as soon as possible. 4. Do not let the objective remain long near corrosive liquids, such as strong solutions of iodine, corrosive sublimate, or mineral acids. Never examine objects lying in such fluids without putting on a cover glass. 5. Never lift the slide from the stage, but, after raising the objec- tive, slide it off the stage without upward movement. 6. Never allow the stand (microscope without lenses) to be wetted with such substances as alcohol, soap, etc., which dissolve lacquer. 7. Keep the microscope covered when not in use. THE TECHNIQUE OF MAKING PERMANENT MOUNTS 1. The Mounting Medium . — When a microscopic object is to be preserved permanently it must be kept from decaying and the fluid in which it is placed must be kept from evaporating. These condi- tions can be met by adding an antiseptic (acetic acid, formaldehyde) to the water used in mounting and carefully sealing the cover glass. As a rule, a better way is to use a mounting medium that will not evaporate, e.g., glycerine or Canada balsam. Both these fluids have a high refractive index and so render the objects penetrated by them more transparent. This quality is generally an advantage, but for objects already almost transparent it is quite the reverse. Glycerine has the disadvantage of always remaining soft, so that the mount may at any time be spoiled by careless handling. Canada balsam, on the other hand, slowly becomes solid, so that the mount is ex- posed to no accident short of actual breakage. Balsam has the dis- advantage of being non-miscible with water, so that before it can be used the object must be carefully dehydrated. Even after this is done, and the object lying in absolute alcohol, an oil must be used as an intermediate agent between alcohol and balsam. FUNDAMENTAL CONSIDERATIONS 9 2. Staining . — For two reasons it is generally better to stain plant tissues before mounting. Transparent tissues may become almost invisible in glycerine or balsam, and different tissues take a stain differently. This being the case it becomes possible to stain one tissue and not another, or one tissue with one stain and another in the same seel ion with a different stain, so that the different parts may be brought out like areas on a colored map. The most common stains are hsematoxylin derived from logwood, and various anilin stains — safranin, fuchsin, eosin, iodine green, methyl-green, etc. METHOD FOR THE PREPARATION OF A CANADA BALSAM MOUNT 1. Stain object with 0.5 per cent, solution of safranin in 50 per cent, alcohol for from three to five minutes. 2. Wash out excess of stain and further dehydrate with 70 per cent, alcohol. 3. Stain with 0.5 per cent, solution of methyl-green in 70 per cent, alcohol for twenty seconds or longer, depending upon the nature of the material. 4. Dehydrate and wash out excess of stain with 95 per cent, alcohol for two minutes. 5. Further dehydrate by placing material in absolute alcohol for one minute. 6. Clear in cedar oil. 7. Mount in Canada balsam. 8. Label slide. METHOD FOR THE STAINING AND MOUNTING OF MATERIAL IN PARAFFINE RIBBONS AFFIXED TO SLIDE 1. Gently heat the dry slides with paraffine ribbons adhering to the fixative, high above the Bunsen flame (with the ribbon side up). 2. Place the slide upright in a well of xylol or turpentine. The xylol or turpentine will dissolve the melted paraffine in a minute or two. 3. Take the slide out of the well, wipe off the under side and allow a stream of 95 per cent, alcohol to run over the upper side from a pipette. IO PHARMACEUTICAL BOTANY 4. Place the slide upright in a well of safranin for from four to twenty-four hours. 5. Take the slide out of the safranin well and extract excess of stain with 50 per cent, alcohol. 6. Place the slide in a well of gentian violet or methyl-green for a second or more. The time varies for different objects and can only be determined by trial. 7. Rinse slide with 70 per cent, alcohol from pipette. 8. Pour absolute alcohol over sections, follow with a few drops of clove oil, replace clove oil with cedar oil. 9. Mount in balsam. MAKING OF SECTIONS Free-hand. Sectioning. — Free-hand sections are usually satisfactory for the general examination of roots, stems, leaves, barks and many fruits and seeds. Material which is fresh may be sectioned at once, but dry material should be well soaked in warm water before using. Very hard material like heartwoods, the shells of nuts and seeds, may be softened in solution of caustic potash or ammonia water and then washed free of alkali before sectioning. The object to be sectioned is held between the thumb and finger of the left hand, or, if small and of soft texture, it must be placed between two flat pieces of elder pith. The upper surface of the razor is wetted with water or 50 per cent, alcohol. The razor, which should be real sharp, is held in the right hand and is drawn across the object with the edge toward the student and the blade sliding on the forefinger of the left hand. The sections should be cut as thin as possible. As soon as a number of sections have been cut, they can be transferred to a vessel of water with a camel’s hair brush. Sectioning in Paraffine or Celloidin. — When it is necessary to study the microscopic structure of very delicate plant parts, the material should be imbedded in paraffine or celloidin, subsequently hardened, and sectioned by means of a sliding or rotary microtome. In your course this work will be carried on by the laboratory staff. KINDS OF SECTIONS 1. A transverse or cross-section is one made horizontally through the object, hence its plane lies at right angles to the long axis. FUNDAMENTAL CONSIDERATIONS 1 1 2. A radial-longitudinal section is one which is made parallel to the long axis of the object in such a way that it lies in plane of the radius. 3. A tangential-longitudinal section is one made parallel to a plane tangent to the cylinder. MICROMETRY The unit of length used in microscopic measurement is the micron (n) which is one-thousandth part of a millimeter (0.001 mm.) or one twenty-five thousandth part of an inch. In measuring microscopic objects it is necessary to make use of a micrometer of some kind. That pretty generally used is the ocular micrometer. It is a circle of glass suitable for insertion within the ocular with a scale etched on its surface. The scale is divided to tenths of a millimeter (0.1 mm.) or the entire surface of the glass may be etched with squares (0.1 mm.), the net micrometer. STANDARDIZATION OF OCULAR MICROMETER The value of each division of the ocular micrometer scale must be ascertained for each optical combination (ocular, objective, and tube length) by the aid of a stage micrometer. The stage micrometer is a slide with a scale engraved on it divided to hundredths of a millimeter (0.01 mm.), in some cases to tenths of a millimeter (0.1 mm.), every tenth line being made longer than intervening ones, to facilitate counting. Method: 1. Insert the ocular micrometer within the tube of the ocular by placing it on the diaphragm of the ocular, and adjust the stage micrometer by placing it on the stage of the microscope. 2. Focus the scale of the stage micrometer accurately so that the lines of the two micrometers will appear in the same plane. Make the lines on the two micrometers parallel each other. This can often be done by turning the ocular to the right or left while looking into the microscope. 3. Make two of the lines on the ocular micrometer coincide with two on the stage micrometer. Note the number of included divisions. 12 PHARMACEUTICAL BOTANY 4. Note the known value for each division of the stage micrometer scale which may either be etched on the stage micrometer or indi- cated on a label found pasted upon it. If the value indicated is 0.01 mm. (/"loo mm.) then each division of the stage micrometer scale has a value of 10 microns; if 0.1 mm. (}^o mm.), 100 microns. 5. Multiply the number of included divisions of the stage microm- eter scale by the value in microns given for each division and divide the result by the number of included divisions of the ocular microm- eter scale. The quotient represents the value of each division of the ocular micrometer scale. 6. Note the optical combination (number of ocular, objective and tube length) used and keep a record of it with the calculated microm- eter value. Repeat for each of the combinations. To measure an object by this method read off the number of divisions it occupies of the ocular micrometer scale, and express the result in microns by looking up the recorded value for the optical combination used. CHAPTER II LIFE HISTORY OF THE MALE FERN [DRYOPTERIS (ASPIDIUM OR NEPHRO DIUM ) FILIX-MAS] The Male Fern along with the Marginal Fern ( Dryopteris margin- alis) have long been known to the pharmaceutical and medical pro- fessions as the source of the drug Aspidium, a most valuable remedy for the expulsion of tapeworm. The parts of these plants employed are the rhizome and stipes which are collected in autumn, freed of the roots and dead portions and dried at a temperature not exceeding 7o°C. HISTORY OF THE SPOROPHYTE OR ASEXUAL GENERATION Gross Structure of Stem. — -The main axis of Dryopteris Filix-mas is the creeping underground stem or rhizome which is oblique or ascending in habit. It gives off numerous roots from its lower and posterior portions and fronds from its upper and anterior portions. Behind the fronds of the present year are to be noted the persistent stalk bases of fronds of previous seasons. Lateral buds are frequently to be noted connected with these. The roots are slender and brown with semi-transparent apices. They are inserted on the bases of the fronds, close to their junction with the stem. The growing end of the rhizome is called the anterior extremity and is marked by the presence of an apical bud overarched by young fronds. The opposite end is known as the posterior extremity and in the living plant is constantly decaying as the anterior portion elongates. Histology of Mature Stem. — Passing from periphery toward the center the following structures are to be observed: i. Epidermis, a protective outer covering tissue, composed of a single layer of brownish polyhedral cells from which are given off scaly hairs. 13 14 PHARMACEUTICAL BOTANY 2. Outer cortex (hypodermis), a zone of several layers of thick- walled lignified cells separating the epidermis from the inner cortex beneath. Its main function is to support the epidermis. Fig. 2 . — Dryopteris filix-mas — Plant and section through pinnule and sorus. {Sayre.) 3. Inner cortex of several layers of more or less isodiametric cells with thin cellulose walls and containing stored starch surrounded by a protoplasmic investment. These cells conduct sap by osmosis and LIFE HISTORY OF THE MALE FERN 15 store food. Between the cells are to be noted intercellular air spaces, many of which contain internal glandular hairs. 4. Fundamental tissue, resembling the last in aspect and function. 5. Vascular Bundles. — These are of two kinds, viz.: stem bundles and leaf-trace bundles. Both are of elliptical outline and are em- bedded in the parenchyma forming the broad central matrix. The stem bundles are comparatively broad and form a continuous net- work with good-sized meshes, each mesh being opposite the point of insertion of one of the leaves. Tn transverse section these bundles are seen to be usually ten in number and arranged in an interrupted circle within the fundamental tissue. The leaf-trace bundles are comparatively narrow and are observed to come off of the stem- bundles and pass out through the cortex into the leaves (fronds). When each bundle is examined under a high-power magnification it is seen to be composed of: (a) an endodermis or bundle sheath , a single layer of cells with yellowish walls and granular contents; ( b ) a pericambium or phloem sheath of one to three layers of delicate thin-walled cells, rich in protoplasm; (c) a phloem , a broad zone of tissue formed of phloem cells, with thin cellulose walls and proto- plasmic contents, which convey sugar in solution from the leaves to the roots and broader sieve tubes which appear polygonal in trans- verse section and whose function is that of conveying soluble pro- teins in the same direction; (d) a xylem (wood) formed of thin-walled xylem cells which store food and scalariform tubes or tracheids which copduct crude sap (water with mineral salts in solution) from the roots to the leaves (fronds). Since the xylem is surrounded by the phloem, the fibro-vascular bundle is of the concentric type. Strictly speaking the endodermis and pericambium are accessory regions sur- rounding, but not part of the bundle proper. Histology of Growing Apex. — When the bases of the leaves of the current year, the circinate leaves of the following year and the large mass of brown scales have been removed from around the apical bud of a well-grown plant, the following structures may readily be ob- served with a hand lens: 1. The apical cone ( punctum vegetationis) , a rounded papilla, which occupies a terminal position in the apical region. 2. The young fronds, arranged around the apical cone. i6 PHARMACEUTICAL BOTANY Upon removing the extreme apex of the apical cone with a sharp razor, mounting in dilute glycerine or water and examining under low power, it will be noted that a large pyramidal cell occupies the center of the apical cone. This is the apical cell. The cells sur- rounding it have been derived by segmentation (cell-division) from it, by means of walls parallel to its three sides; they are termed seg- ment cells and in turn undergo further division and redivision to originate the entire stem tissue and leaf tissue. Step by step the tissue cells become modified into epidermal, cortical, bundle and fundamental cells. Histology of Mature Root. -Transverse sections cut some distance above the apex will present the following structures for examination: 1. Epidermis, of epidermal cells whose outer walls are brown. Some of these cells have grown out as root hairs which surround soil particles and absorb water with mineral salts in solution. 2. Cortex, of many layers of cortical parenchyme cells with brown walls. The outer layers of cells of this region are thin- walled while the extreme inner ones are lignified and form a sclerenchymatous ring which surrounds the 3. Endodermis, a single layer of cells tangentially elongated. 4. Pericambium (Pericycle), usually of two layers of thin- walled cells containing protoplasm and large nuclei. This region surrounds the 5. Radial fibro-vascular bundle, consisting of two phloem patches of phloem cells and sieve tubes on either side of two radial xylem arms of xylem cells, spiral tracheae and scalariform tubes. 6. Lateral rootlets which take origin in the pericambium. Histology of Root Apex. -Microscopic examination shows this region to be composed of soft, pale, growing cells ending in the tri- angular apex-cell of the root. From the free base of the apex cell segment cells are cut off as calyptrogen cells. These by dividing form the root cap. The root cap or calyptra consists of a mass of loosely attached cells which forms a protective covering around the tip of the root. From the inner sloping sides of the apex cell the segment cells give origin to the dermatogen, which by repeated division of its cells origi- nates the epidermis (outer protective covering of the root), the peri- LIFE HISTORY OF THE MALE FERN 17 blem, originating cortex, and the plerome originating the bundle and related tissue. Continuity of Crude Sap Flow. — The crude sap (water with mineral salts in solution) penetiates the thin walls of the root hairs by osmosis and passes into the interior of hairs, thence into the root xylem and through this to stem xylem, thence through stem xylem into the leaves. Histology of Stipe (Petiole). — This in transverse section passing from periphery toward the center presents the following structural characteristics: 1. Epidermis, a single layer of epidermal cells with dark brown outer walls. 2. Outer cortex (hypodermis), a wide band of small cells with ligni- fied walls. 3. Inner cortex, similar to inner cortex of stem but devoid of leaf- trace bundle. 4. Fundamental parenchyma, similar to same region of stem in which are embedded a number of concentric fibro-vascular bundles arranged in an interrupted circle. Each of these shows a central xylem mass surrounded by an outer phloem mass. Each bundle is enveloped by a pericambium and an endodermis or bundle sheath. Histology of Lamina. — In transverse and surface sections the lamina or blade shows the following structural details: 1. Upper epidermis of wavy- walled, slightly chlorophvlloid, flat upper epidermal cells devoid of stomata but with rather thick cuticle. 2. Mesophyll of irregular shaped chlorophylloid cells containing abundant chloroplasts. Intercellular-air-spaces are found between various cells which are larger in the lower than the upper region. Internal glandular hairs are frequently to be discerned in many of these spaces. 3. Concentric vascular bundles or laminar veins that distribute sap to, and carry sap from the mesophyll. These are seen to be embedded in the mesophyll. The xylem portion of each bundle is nearest to the upper surface of the leaf and so the bundles approach the collateral type. 4. Endodermis, a continuous layer of mesophyll cells which sur- rounds each bundle and binds it in place. 2 i8 PHARMACEUTICAL BOTANY 5 . Lower epidermis of wavy-walled, flattened, chlorophylloid cells with thin cuticle and many stomata (breathing pores). Each stoma is surrounded by a pair of crescent-shaped guard cells which regulate its opening and closing. The upper and lower epidermis are con- tinuous around the laminar margin. Comparative Physiology of Root, Stem and Leaf (Frond). — The primary function of the roots of the Male Fern is that of absorption of water with mineral salts in solution. The secondary function is that of support for the stem, the tertiary, that of storing food- stuffs to tide the plant over the season when vegetative activities are lessened. Water is the most essential of all materials absorbed by vegetable organisms. It is found in the soil surrounding the soil particles with certain mineral salts dissolved in it. The delicate root-hairs with thin cellulose walls, protoplasmic lining and sap denser than the soil water, are firmly adherent to these particles. The soil water diffuses through these walls by osmosis and comes into relation with the ectoplasm, a delicate protoplasmic membrane which has the power of selecting what it wants and rejecting what it does not need. In this way only such solutes as are of value to the plant are admitted. The water with mineral salts in solution, once within the root-hair protoplast, is called “crude sap.” This passes through the hair into the cortical parenchyma cells which are in contact with the spiral ducts and scalariform tracheids. It passes from one cortex cell to another by osmosis and, under considerable root pressure, is forced into the spiral and scalariform tubes of the xylem. Therein it is conveyed upward by root pressure through the tracheids of the stem bundles into those of the leaves and finally osmoses into the leaf parenchyme cells (mesophyll). Carbon dioxide, (C0 2 ), from the air enters the leaf through the stomata. From the stomata it moves through the intercellular air spaces to the mesophyll cells which line these, whence it is absorbed. Within the mesophyll cells are found small chloroplasts composed of protoplasm and chlorophyll. The kinetic energy of the sun’s rays is absorbed by the chlorophyll which is thus energized to break up the C0 2 and H 2 0 into their component elements C, H and O, and rearrange them in such a way as to ultimately form sugar or starch. This process is called photosynthesis. According to von Baeyer, LIFE HISTORY OF THE MALE FERN 19 C0 2 is split into C and 0 2 , the C being retained, the 0 2 given off. The nascent C is linked with H 2 0 to form CH 2 0 (formic-aldehyde); six molecules of this are then united to form grape sugar (C 6 Hi 2 0 6 ). The formation of starch may be expressed by the following equation: 6C0 2 + 5H 2 0 = C 6 H 10 O 5 + 60 2 . A portion of the grape sugar is removed from solution by the chloroplast and converted into starch which is stored up within it; another portion is used to nourish the protoplasm of the cell. But the greater portion of sugar manu- factured descends in solution through the phloem cells of the bundles of the veins, mid-rib and stipe to the stem or roots where it is re- moved from solution by the action of the leucoplasts which convert it into reserve starch. Sugar and starch, however, are not the only food materials manufactured in the leaf. Proteins are likewise formed. These are composed of carbon, hydrogen, oxygen, nitrogen, sulphur and sometimes phosphorus. They are formed from grape sugar with the addition of nitrogen and the other elements by the living protoplasm. The source of nitrogen, sulphur and phos- phorus is the mineral salts which are found in the crude sap. These proteins descend through the sieve tubes of the veins, midrib and petiole, to the stem and roots nourishing all of these parts with protein material. Gross Structure and Histology of the Sori and Sporangia. — The sporangia or spore cases are found clustered together in circular groups on the under surface of the pinnules nearer the mid-vein than the margin. Each group of sporangia is covered with a membranous expansion of the epidermis called the indusium. The whole is called a sorus (Fig. 2 ) (pi. sori) and contains many sporangia. Each sporan- gium is composed of: (a) the stalk of considerable length and usually comprising three rows of cells, outgrowths of the epidermis of the pinmile; and ( b ) the head, sub-globular and hollow, consisting for the most part of a covering of thin walled, flattened cells within which will be noted a marginal ring of cells with walls having U-shaped thickenings and called the annulus. Within the sporangium are found the spores. Each spore is a single cell composed of an outer brown wall with band-like markings called an exosporium, an inner thinner wall or endosporium, and within this a mass of protoplasm containing a nucleus. 20 PHARMACEUTICAL BOTANY Rupture of Sporangium and Spore Dissemination. — As was pre- viously indicated, each sporangial head has a row of cells with U-shaped thickenings around the margin called an annulus. As the sporangium matures the water escapes from the cells, pulling them together and holding the annulus like a bent spring. The thinner walled cells at the side of the spore case opposite the annulus, unable to stand the strain are consequently torn, the annulus straightens and a wide rent is made in the sporangium. The annulus then recoils and hurls the spores out of the sporangium. This closes the sporo- phyte generation. Fig. 3. — Sporangia of an undetermined species of fern; li, lip-cells; an, annulus; st, stalk; sp, mature spores. Each of the four nuclei in the upper cells of the stalk is in the terminal cell of one of the four rows of cells that compose the stalk. (Gager.) History of the Gametophyte or Sexual Generation. — The fern spore, falling upon a moist surface, germinates, producing a delicate green septate filament called a protonema. One end of this structure shows larger cells which, by the formation of oblique walls, cut out an apical cell of somewhat triangular shape. This is the growing point of what eventually becomes a mature green heart-shaped body called the “prothallium” or “prothallus.” The prothallium, about the size of an infant’s finger nail, develops on its under surface anther- idia, or male sexual organs, archegonia, or female sexual organs, and rhizoids or hair-like absorptive structures. The antheridia appear three to five weeks after spore germination. They are hemispherical in shape and are situated among the rhizoids toward the posterior end. Each antheridium consists of a three-celled wall which com- pletely surrounds the spermatocytes or mother-cells of the sper- matozoids. Within each spermatocyte the protoplasm arranges itself ,IFE HISTORY OF THE MALE FERN 21 Fig. 4. — A, under surface of a fern prothallium showing archegonia (/), antheridia (m) and rhizoids (r): B, immature archegonium showing binucleate neck canal cell ( n.c.c .), ventral canal cell (v.c.c.), and egg (e); C, mature archegonium showing sperms (sp.) moving through neck canal (n.c.) toward ovum ( e ), and venter ( v ). All highly magnified. 22 PHARMACEUTICAL BOTANY in a spiral fashion forming a spermatozoid, a spiral, many ciliated, male sexual cell. From two to four weeks after the maturation of the antheridia, the arcliegonia make their appearance toward the indented apex of the lower prothallial surface as outgrowths of the prothallial cushion. Since they appear later than the antheridia they are not likely to be fertilized by spermatozoids from the anther- idia of the same prothallium. Each archegonium is composed of a venter, neck, neck canal-cells, ventral canal-cell, and ovum or egg-cell. The neck is composed of cells arranged in four rows, forming a cylin- der, one layer of cells thick. This protrudes from the surface of Fig. 5. — A, median longitudinal section through immature antheridium, and cell of prothallium showing prothallial cell ( p ), and antheridial wall surrounding a number of spermatocytes; B, similar section through mature antheridium and cell of prothallium showing fully developed spermatozoids ( sp .) enclosed by wall of antheridium. Both highly magnified. the prothallium and encloses the neck canal-cells and ventral canal- cell. The ovum is embedded in the prothallial cushion just beneath the ventral canal-cell. Upon the maturation of the archegonium, the canal-cells are transformed into a mucilaginous substance which fills a canal extending from the outside opening (mouth) to the ovum. During wet weather the mature antheridial wall bursts open and the many ciliated spermatozoids escape into the water. These mov- ing in the water are drawn by the chemotactic malic acid to the mouths of the archegonia of another prothallus and, passing down the canal of each of these, gather around the ovum. One, probably the best adapted, fuses with the ovum and fertilizes it forming an oospore or fertilized egg. LIFE HISTORY OF THE MALE FERN 2 3 Origin of New Sporophyte or Diploid Plant from Fertilized Egg. — The fertilized egg now rapidly divides and redivides to form octant cells. The octant cells further divide to produce anteriorly a stem rudiment (one cell), first leaf (two cells), second leaf (one cell) and, posteriorly, root rudiment (one cell), foot rudiment (two cells) and hair rudiments (one cell). Growth of Seedling into Mature Sporophyte. — The foot rudiment develops into the foot which obtains nourishment from the prothal- lium upon which the young sporophyte is for a time parasitic. The root rudiment becomes the first root which grows downward into the soil. The stem and leaves turn upward. In a few weeks the prothallus decays and the sporophyte is established as an independ- ent plant. More roots and leaves (fronds) are developed and ere long continued growth results in the formation of a mature sporo- phyte which presents for examination: (i) a subterranean stem bear- ing several roots; and (2) aerial fronds, each of which consists of a stipe or petiole and a lamina or blade, divisible into pinna or lobes and pinnules, upon which last sori are developed. Alternation of Generations. — It will be observed that in the life cycle of the Male Fern there occur two distinct generations, one, a sporophyte or asexual generation which begins with the oospore and ends with the dispersion of asexual spores; a second, the gametophyte or sexual generation beginning with the protonemal outgrowth of the spore and ending with the fertilization of the egg to form an oospore. The sporophyte gives rise to the gametophyte which in turn gives origin to the sporophyte. CHAPTER III LIFE HISTORY OF A GYMNOSPERM (PINUS STROBUS) The White Pine frequently called the Weymouth Pine ( Pinus Strobus), one of the principal timber trees of the Northern States and Canada, is also of value in pharmacy and medicine. The inner bark of its trunk and branches is used because of its valuable ex- pectorant properties and is official in the N. F. IV. DESCRIPTION OF THE WHITE PINE TREE (MATURE SPOROPHYTE) From an underground spreading root system there arises an erect aerial trunk or stem that extends from the ground to the apex of the tree, ending in a terminal bud. The trunk rarely exceeds 3 feet in diameter and 125 feet in height and is averagely iK to 3 feet in diameter and 50 to 90 feet high. At a varying distance above the soil, depending upon environal conditions as well as the age of plant, whorls of lateral branches (three to seven in a whorl) are seen ema- nating from the trunk in horizontal fashion at various levels up to near the apex. These become, under conditions prevalent when the tree is grown in the open, gradually shorter until the summit is reached, giving to the crown or upper part of the tree the appearance of a pyramid. These branches give rise to other branches which agree with the lateral branches in bearing commonly only scale-like leaves as well as ending in terminal buds. Another kind of branch, however, is found which is always shorter than the scaly branches. This type of branch is called a “spur shoot ” and arises from the former branches. The spur shoots bear the needles or foliage leaves which are light green when young and bluish-green, soft, flexible, 2 l 2 to 5 inches long when mature. The “needles” occur in tufts ( fascicles ) of five, are triangular in cross-section, have finely serrate ( saw-toothed ) edges and are surrounded at the base by a deciduous 24 LIFE HISTORY OF A GYMNOSPERM 25 sheath. These foliage leaves persist until the end of their second year when they are shed with the spur shoot which bears them. The white pine, like most of its allies among the Coniferae, bears cones. These structures are of two kinds, viz.: staminate and car- pellate. Both kinds are produced on the same tree. Fig. 6. — -Transverse section of pine stem in third year of growth showing xylem (a), cork (6), cortex (c). secretion reservoir (d), cambium ( e ), phloem (/), and pith (g). (Photomicrograph.) Staminate Cones. — The yellow, ovate, staminate cones appear about May and are clustered at the base of the new growth of the current season. Each consists of a main axis (modified branch) which bears spirals of scales (micro sporophylls or stamens). On the under surface of each scale are the spore-cases (micros por angia) , which develop the microspores (pollen grains). Each pollen grain when mature consists of a central fertile cell and a pair of air-sacs 26 PHARMACEUTICAL BOTANY Fig. 7. — Transverse section of white pine needle (leaf) showing epidermis (a), infolded parenchyma cells of mesophyll ( b , b'), oil reservoir (c), endodermis ( d ). and clear cellular area surrounding fibrovascular tissue in center (e). Fig. 8. — Staminate cones of the Austrian pine ( Pinus austriaca) . Below, before shedding pollen; above, after shedding. (Gager.) LIFE HISTORY OF A GYMNOSPERM 27 or wings, one on either side of the fertile cell. The purpose of the latter is to give greater buoyancy in the air to the microspore. Carpellate Cones.- — The young carpellate cones appear in May or early June as pinkish-purple structures arranged in solitary fashion or in small groups, lateral along the new growth. Each terminates Fig. 9. — Scotch pine ( Pinus sylvestris) . A-D, stages in the development of the carpellate cone, and its carpotropic movements. E, very young carpellate cone much enlarged; F, ventral, G, dorsal views of a scale from E; 1, ovuliferous scale; 2, ovule (in longitudinal section); 3, pollen chamber and micropyle leading to the apex of the nucellus (megasporangium); 4, integument of the ovule; G, 1, tip of ovuliferous scale; 5, bract; 4, integument; H, longitudinal section at right angles to the surface of the ovuliferous scale (diagrammatic); 6, megaspore; 7, pollen chamber; I, longitudinal section of a mature cone; 6, ovule; J, scale from a mature cone; 6, seed; w, wing of seed; K, dissection of mature seed; h, hard seed coat; c, dry membraneous remains of the nucellus, here folded back to show the endosperm and embryo; e, embryo; p, remains of nucellus; L, embryo; c, cotyle- dons; e, hypocotyl; r, root-end. (Gager.) a lateral axillary branch. A carpellate cone is composed of a main axis which bears spirals of scales, by some termed megasporophylls {carpels). Each scale is composed of an ovuliferous scale bearing two ovules or megasori and a bract. Each megasorus contains a nucellus or megasporangium which is surrounded by an integument 28 PHARMACEUTICAL BOTANY except at the apex where an opening, the micropyle is evident. The micropyle is the gateway to the pollen chamber which lies below it. Within the nucellus occurs a megaspore or embryo sac. DESCRIPTION OF THE GAMETOPHYTE GENERATION The Gametophyte generation of the White Pine begins with the development of the male and female gametophytes and terminates with the fertilization of the egg. The Male Gametophyte. — The male gametophyte commences to form in the mature pollen grain before the pollen is shed. A series of three nuclear divisions takes place which cut off two small prothal- lial cells (traces of one of which may be seen pushed up against the wall of the fertile cell of the pollen grain), a tube nucleus and a generative cell. At this stage the pollen is shed and some of it is Fig. io. — The white pine (Pinus Strobus). Sections through mature pollen grains; at the left the remnants of two prothallial cells can be seen, while at the right all signs of the first cell have disappeared. Pollen collected June 9, 1898. X about 600. (Gager, after Margaret C. Ferguson.) carried by air currents to the carpellate cones where it sifts in be- tween the ovule-bearing scales and accumulates at the scale bases. This is pollination. A number of the pollen grains are drawn close to the nucellus of the ovule by the drying up of the viscid fluid which fills the pollen chamber. In this fluid they germinate form- ing pollen tubes. The transfer of pollen grains from the pollen sac to the pollen chamber and consequent germination therein is called pollination. The Female Gametophyte.- — If the embryo sac be examined at about the time of pollination, it will be found to consist of a single cell containing a single nucleus surrounded by cytoplasm. Very shortly afterward, however, the nucleus divides repeatedly to form a LIFE HISTORY OF A GYMNOSPERM 2 9 large number of nuclei which are scattered throughout the cyto- plasm. Each nucleus accumulates around itself a portion of the cytoplasm and ultimately cell walls are laid down and the entire embryo sac contains endosperm (prothallial) tissue. Toward the micropylar end of the endosperm { proihallus ) originate several archegonia. Each archegonium consists of a much-reduced neck of four cells and an egg {ovum) which lies embedded in the prothallus which forms a narrow layer of cells around it called the jacket. The con- tents of the mature embryo sac constitutes the female gametophyte. Fig. 11. — White pine (Pinus Strobus). At left, megasporangium with mega- spore in the center; above, pollen grains in the micropyle and pollen chamber. At right, pollen grains beginning to germinate; the cells of the integument have X enlarged and closed the micropyle. {Gager, after Margaret C. Ferguson.) Fertilization.- — About a year after pollination the pollen tubes, lying within the pollen chamber show signs of renewed activity. The tube nucleus passes to the tip of the tube. The generative-cell divides to form a body and a stalk-cell which pass into the tube. The body-cell later forms two sperm nuclei. While these changes are taking place the tube is penetrating the nucellus and growing toward the embryo sac with its contained female gametophyte. It finally enters it, passing between the neck-cells of the archegonium. The tip of the tube then breaks and the entire tube contents is emptied into the egg. One of the sperm nuclei fuses with the egg nucleus and fertilizes it forming an oospore. 30 PHARMACEUTICAL BOTANY Seed Formation and Distribution.- — The oospore undergoes re- peated divisions and forms the embryo or young sporophyte plant and a suspensor to which it is attached. The embryo is nourished by a portion of the prothallus but the greater part of the prothallus forms the endosperm tissue of the seed surrounding the embryo. The thin nucellus persists as an endosperm covering. The integu- ment becomes modified to form the hard protective seed coat. A Fig. 12. — White pine (Pinus Strobus). Vertical section through theupper part of an ovule, shortly before fertilization, s.n., sperm- nuclei; st. c., stalk-cell; t.n. tube-nucleus; arch, archegonium; e.n., egg-nucleus. (Gager, after Margaret C. Ferguson.) portion of the scale of the cone directly above and adjacent to the ovule forms a membranous wing which separates from the scale as part of the seed. By this time (about two years after pollination) the scales of the cone, now quite woody, separate, the seeds are shaken out, and many are carried for a considerable distance by winds. LIFE HISTORY OF A GYMNOSPERM 3 r Germination of the Seed.- — Under favorable conditions, the seeds absorb water and germinate in the spring following their dispersal. The hypocotyl of the embryo appears first, arching upward and downward, and, straightening out, draws the green cotyledons with it which spread out toward the light while it grows into the soil to form the tap root and in time the remainder of the root system. Thus the seedling sporophyte is formed which in time develops into the mature White Pine tree. CHAPTER IV LIFE HISTORY OF AN ANGIOSPERM (ERYTHRONIUM AMERICANUM) This attractive little plant, commonly called the Dog’s Tooth Violet but related to the Lily, is found in the hollows of woods and may be seen in flower during the month of April in the Middle Atlantic States. It consists of an underground stem bearing scales (modified leaves) which is termed a bulb. From the lower surface of the bulb are given off numerous slender rootlets which penetrate the soil and from the upper surface, a pair of oblong lance-shaped leaves of pale green color mottled with purple and white and later, a flower stalk {scape) which bears upon its summit a single yellow nodding flower which is often marked with purple stripes. The flower consists of a torus or receptacle which will be observed as the upper swollen end of the flower stalk (scape). Inserted upon it, passing from periphery toward the center, will be noted four whorls of floral leaves which in order are calyx, corolla, andrcecium and gynce- cium. The calyx is composed of three lance-shaped and recurved yellow parts called sepals; the corolla of three similarly looking parts called petals which alternate in position with the sepals. Both of these whorls are collectively called the perianth or floral envelope. The androecium or male system of organs is composed of two whorls or circles of structures called microsporophylls or stamens. There are three stamens in each whorl. The outer whorl of stamens will be found opposite the sepals while the inner will be observed opposite the petals. Each stamen (microsporophyll) con- sists of an awl-shaped stalk or filament bearing upon its summit an oblong-linear body called an anther. The anther consists of two lobes called microsori. Each lobe or microsorus contains two anther sacs or microsporangia in which when mature are to be found micro- spores or pollen grains. In the center of the flower will be noted 32 LIFE HISTORY OF AN ANGIOSPERM 33 the gynoecium or female system of organs. This, upon dissection, will be found to consist of three fused carpellary leaves termed mega- sporophylls (carpels) forming a somewhat flask-shaped structure called a pistil. The swollen basal portion of the pistil is called the ovary; the stalk which arises from it is called the style and the knob- like viscid summit of the style is termed the stigma. Development of the Female Gamelophyte Through the Maturation of the Embryo Sac. — In its immature condition the embryo sac ( mega- spore ) contains a mass of protoplasm surrounding a nucleus. This nucleus undergoes three divisions forming as a result eight nuclei which ultimately arrange themselves within the protoplasm of the embryo sac as follows: three of them occupy a position at the apex, the lower nucleus of the group being that of the egg or ovum , the other two nuclei being the synergids or assisting nuclei; at the oppo- site end of the sac three nuclei known as antipodals take their posi- tion; the two remaining nuclei called polar nuclei take up a position near the center of the embryo sac. In this condition the contents of the embryo sac constitutes the female gamelophyte. POLLINATION AND FERTILIZATION The mature pollen grains are discharged from the ripened anther through the splitting open of its wall. They are transferred to the stigma of the pistil of another Erythronium flower through the agency of insects. Here they germinate, each putting forth a tube (pollen tube). The pollen tubes, carrying within it two sperm nuclei and a tube nucleus embedded in protoplasm, penetrate through the style canal until they reach the micropyles of various ovules. Each enters and passes through a micropyle, then piercing the nucellus, grows toward the embryo sac. The tip of the tube fuses with the end of the embryo sac and the two sperm nuclei are discharged into the sac. One of these sperm nuclei passes between the synergids and fuses with the nucleus of the egg to form an oospore. By this time the tube nucleus has disintegrated. The oospore by repeated divisions develops into as many as four embryos or young sporophyte plants. Only one of these, however, persists, The polar nuclei fuse to form the endosperm nucleus which soon undergoes rapid division into a large number of nuclei scattered 3 34 PHARMACEUTICAL BOTANY about through the protoplasm of the embryo sac. Later cell walls are laid down and endosperm is formed. The endosperm cells soon become fdled with abundant starch which is later to be utilized by the embryo during germination. Microscopical examination of sections of the ovary will reveal it to be composed of three chambers called locules within each of which are to be noted several inverted ovules. Each of these ovules is developed upon a nourishing tissue termed “ placenta ” which con- nects the ovules to the inner angle of the wall of the locule. The ovule is composed of a central prominent megasporangium or nucellus which is almost completely invested by two upgrown integuments or coverings. The opening between the tips of the inner integument is called the micro pyle (little gate). This is the gateway for the entrance of the pollen tube on its way to the nucellus. It is also the exit door for the hypocotyl of the embryo after the fertilized and ripened ovule becomes a seed. Within the nucellus, if the sec- tions examined have been properly fixed, will be found a megaspore or embryo sac. MATURATION OF THE POLLEN GRAIN AND FORMATION OF THE MALE GAMETOPHYTE The pollen grains (microspores) within the anther sacs all arise from a number of tetrads (groups of four) which are formed by the division and redivision of pollen mother-cells preceding them. Each pollen grain, after the tetrads have separated into their components, consists of an outer firm wall or exosporium, an inner wall or endo- sporium, within which will be found the region called the fovilla, which is nothing other than a mass of protoplasm containing a nucleus. Before the pollen is shed from the anther its protoplasmic contents undergo a series of changes leading up to the development of the male gametophyte. The nucleus and protoplasm enveloping it divide to form two cells, one a generative-cell containing a genera- tive nucleus, the other a tube-cell containing a tube nucleus. The generative nucleus then divides to form two sperm nuclei and the partition wall between the two cells disappears. In this condition the protoplasmic contents of the pollen grain constitute the male gametophyte. LIFE HISTORY OF AN ANGIOSPERM 35 RIPENING OF THE OVULE TO FORM THE SEED AND OF THE OVARY TO FORM THE FRUIT While the embryo and endosperm are being formed, the ovule enlarges and its integuments become modified to form a hard horny seed coat which encloses the endosperm surrounding the embryo. The ovary, containing the ovules, has by this time ripened to form a three-valved loculicidal capsule enclosing the seeds. GERMINATION OF THE SEED AND DEVELOPMENT OF THE MATURE SPOROPHYTE The seeds are fully developed by June or July when the capsule or fruit splits open to discharge them. They fall to the ground and he dormant until the following spring when they germinate or com- mence to grow. Each seed absorbs water from the ground which stimulates the ferment amylase, contained in the endosperm cells, to break up the insoluble starch into soluble sugar which passes into solution and diffuses into the cells of the embryo where the proto- plasm changes it into additional protoplasm and so the embryo in- creases in size, therefore, grows. The pressure of the swollen endo- sperm and growing embryo becomes so great that the seed coat bursts; the hypocotyl emerges first, dragging the cylindrical cotyle- don out of the seed coat and epicotyl with it. The hypocotyl elon- gates and extends itself into the soil where it develops a root near its tip. The tip enlarges through the storage of starch, manufac- tured by the green cotyledon and becomes a bulb. The bulb soon develops within it a plumule, the cotyledon withers, and the young plant (seedling) passes the following winter in this condition. During the next spring the plumule develops into a foliage leaf and the bulb gives rise from its base to several slender elongated runners, which, at their tips develop runner bulbs. These runner bulbs the third year give origin to another set of runners similar to those formed during the second year which also develop runner bulbs at their tips. A foliage leaf is also formed by each. The following spring (spring of fourth year) one of these bulbs develops into a mature sporophyte plant bearing a single flower at the summit of its elongated scape. 36 PHARMACEUTICAL BOTANY RESEMBLANCES BETWEEN GYMNOSPERMS AND ANGIOSPERMS 1. In both are developed those structures in which there is no homologue, e.g., flowers. 2. In both the flowers develop at least two sets of leaves (either on one or two plants of the same species) called sporophylla or sporophyll leaves, the stamens and carpels. The stamens or stam- inal leaves are also termed microsporophylls. The carpels or carpellate leaves are also known as megasporophylls. 3. Both groups produce microspores or pollen grains and mega- spores or embryo sacs. 4. In both are developed on the evident generation, the plant or sporophyte and the gametophyte, the latter concealed within the megaspore of the sporophyte. 5. Both develop seeds with one or two seed coats. 6. In both groups there is developed from the fertilized egg an embryo which lies within the cavity of the megaspore. 7. In both there exists a root and a stem pericambium. 8. Both produce collateral vascular bundles. Very rarely do we meet with concentric bundles in the stem or leaf of Angiosperms. FUNDAMENTAL DIFFERENCES BETWEEN GYMNOSPERMS AND ANGIOSPERMS 1. The flowers of Gymnosperms are often monoecious or dioecious but very rarely hermaphrodite as in Welwitchia, whereas those of Angiosperms are usually hermaphrodite, rather rarely monoecious, still more rarely dioecious. 2. In the Gymnosperms the sporophylls are usually inserted either spirally or in whorls around a distinctly elongated axis, whereas in Angiosperms the sporophylls are condensed to short whorls or spirals set around a shortened axis, the floral axis or recep- tacle, torus or thalamus, or, as in the more modified Angiosperms, the floral axis may even become hollow. 3. In Gymnosperms the microsporophylls or stamens are usually sessile, whereas in Angiosperms the microsporophylls are nearly always stalked. Rarely do we find sessile anthers among Angio- GYMNOSPERMS VS. ANGIOSPERMS 37 sperms, an instance of this being seen in Mistletoe (Viscum) where the anthers are set on the staminal leaf. 4. In Gymnosperms there is a traceable prothallus or gametophyte plant that later becomes the so-called “endosperm” of the gymno- sperm, whereas in Angiosperms no recognizable prothallus has been proven to exist. 5. The stored food tissue in Gymnosperm seeds is prothallial tis- sue loaded with starch, etc., whereas in Angiosperm seeds the stored food tissue (endosperm) is a special formation after fertilization. 6. Angiosperms bear naked ovules and seeds while Gymnosperms bear covered ones. 7. In Gymnosperms there are distinct recognizable archegonia formed on or imbedded in the prothallus, whereas in Angiosperms there are no distinct archegonia, only an isolated egg or eggs. 8. In Gymnosperms there are not infrequently found several embryos from one fertilized egg. This condition is called poly- embryony. Polyembryony is unknown in Angiosperms, only a false polyembryony being noticed. 9. In Gymnosperms the secondary xylem (wood) tissue of roots, stems and leaves consists either of punctated or scalariform cells, whereas in Angiosperms the secondary wood tissue may be varied in structural aspect. CHAPTER V VEGETABLE CYTOLOGY Vegetable Cytology treats of plant cells and their contents. THE PLANT CELL AS THE FUNDAMENTAL UNIT Schleiden, in 1838, showed the cell to be the unit of plant structure. The bodies of all plants are composed of one or more of these funda- mental units. Each cell consists of a mass of protoplasm which may or may not have a cell wall surrounding it. While most plant cells contain a nucleus and some contain a number of nuclei, the cells of the blue-green algae and most of the bacteria have been found to lack definitely organized structures of this kind but rather contain chromatin within their protoplasm in a more or less diffuse or loosely aggregated condition. A TYPICAL PLANT CELL If we peel off a portion of the thin colorless skin or epidermis from the outer convex surface of an onion bulb scale, mount in water and examine under the microscope, we find it to be composed of a large number of similar cells which are separated from one another by means of lines, the bounding cell walls. Under high power each of these cells will exhibit the following characteristics: An outer wall, highly refractile in nature and composed of cellulose which surrounds the living matter or protoplasm. This wall is not living itself but is formed by the living matter of the cell. Some- where within the protoplasm will be noted a denser-looking body. This is the nucleus. Within the nucleus will be seen one or more smaller highly refractile and definitely circumscribed bodies, the nucleolus or nucleoli. The protoplasm of the cell outside of the nucleus is called the “cytoplasm.” It will be seen to be clear and 38 VEGETABLE CYTOLOGY 39 granular-looking. Within the cytoplasm will be observed a number of clear spaces. These are vacuoles and because they are filled with cell sap (water with nutrient substances in solution) are called “ sap vacuoles Protoplasm is in intimate relation to water. The reaction of the cytoplasm to a bounding film of water between it and the cell wall forms the outer plasma membrane or ectoplasm, a clear homogeneous outer band of cytoplasm; the reaction of cytoplasm to the water within the sap vacuoles forms the vacuolar membranes; the reaction of the dense protoplasm of the nucleus to the water in the cytoplasm around it forms the nuclear membrane. Upon mounting another portion of epidermis in iodine solution, removing the excess of stain and adding a drop of sulphuric acid and then examining under high power, we note that the cell walls of cellulose are stained a deep blue. A yellow line is evident in the middle of each cell wall and separates each cell from its bounding cells. This fine is the middle lamella which is composed largely of calcium pedate. PROTOPLASM AND ITS PROPERTIES Protoplasm, or living matter, is the more or less semi-fluid, viscid, foamy, and granular substance in which life resides. It is the “physical basis of life.” The peculiar properties which distinguish protoplasm from non- living matter are as follows: 1. Structure. — Protoplasm invariably exhibits structure. No por- tion of it, however small, has been found to be homogeneous. Each advance in microscopical technique reveals new complexities. The protoplasm of a single cell, far from being a single unit, must rather be looked upon as a microcosm. 2. Metabolism. — Perhaps the most significant peculiarity of living matter is found in its instability and the chemical changes which continually go on within it. It is constantly wasting away, and as constantly being built up. These losses and gains are not upon the exterior surface, but throughout its mass. Its growth and renewal are by intussusception, or the taking in of new particles and storing them between those already present. A bit of protoplasm may re- 40 PHARMACEUTICAL BOTANY tain its identity while all the matter of which it is composed is changed over and over. It is like a whirlpool or wave in a river which remains the same while the water of which it is composed changes continually. Metabolism has been aptly defined by Huxley as the whirlpool character of the organism. 3. Reproduction.- — Protoplasm also shows a very remarkable ability to increase and to give off detached portions which retain the infinitely complex peculiarities and properties of the original. The process, moreover, may be continued indefinitely. Other physiological characteristics might be added, but the above are mentioned as the most satisfactory criteria by which living may be distinguished from non-living matter. PROTOPLASMIC CELL CONTENTS Protoplasm consists of four well-differentiated portions: (a) Cytoplasm, or the foamy, often granular matrix of protoplasm outside of the nucleus. (b) Nucleus or Nucleoplasm, a denser region of protoplasm con- taining chromatin, a substance staining heavily with certain basic dyes. ( c ) Nucleolus, a small body of dense protoplasm within the nucleus. (d) Plastids, composed of plastid plasm, small discoid, spheroidal, ellipsoidal or ribbon-shaped bodies scattered about in the cytoplasm. Sometimes, as in the cells of lower plants like the Spirogyra, plastids are large and are then called chromatophores. According to the position of the cells in which plastids occur and the work they perform, they differ in color, viz.: Leucoplastids are colorless plastids found in the underground portions of a plant and also in seeds, and other regions given up to the storage of starch. Their function is to build up reserve starch from sugar and other carbohydrates as well as to change the reserve starch back into sugar when it is needed for the growth of the plant. They are only evident after properly fixing and staining cells con- taining them. Chloroplastids are plastids found in cells exposed to light and con- tain the green pigment, chlorophyll. VEGETABLE CYTOLOGY 41 Chromoplastids are plastids found in cells independent of their relation to light or darkness and contain a yellow, orange or red pigment called chromophyll. Fig. 13. — A, embryonic cells from onion root tip; d, plasmatic membrane; c, cy- toplasm; a, nuclear membrane enclosing the thread-like nuclear reticulum; b, nucleolus; e , plastids (black dots scattered about). B, older cells farther back from the root tip. The cytoplasm is becoming vacuolate; /, vacuole. C, a cell from the epidermis of the mid-rib of Tradescantia zebrina, in its natural condi- tion on the right, and plasmolyzed by a salt solution on the left; g, space left by the recedence of the cytoplasm from the wall; the plasma membrane can now be seen as a delicate membrane bounding the shrunken protoplast. All highly mag- nified. (Stevens.) CELL FORMATION AND REPRODUCTION The cells of plants have all been derived from preexisting cells. In the bacteria and many other low forms of plant life, the division of the cell always results in reproduction; in higher forms, however, it merely increases the size of the individual and so is a phenomenon of growth. ( 42 PHARMACEUTICAL BOTANY There are two kinds of cells formed by plants, viz.: asexual and sexual. Both of these are endowed with the possibilities of repro- duction, although the former are frequently limited to the process of growth. Reproduction is the power possessed by an organism of giving rise to new individuals. This may take place through the agency of either asexual or sexual cells and is accordingly asexual or sexual in character. Whenever a union of cells or their protoplasmic con- tents takes place the process is called “sexual reproduction;” if, however, there is a mere separation of a cell or cells from an indi- vidual which later form a new organism, the process is termed “asexual or vegetative reproduction.” There are four modes of asexual reproduction, viz.: Fission, Gemmation, Free Cell Formation and Rejuvenescence. Fission. — This is the separation of a cell into two equal halves, each of which may grow to the size of the original parent cell from which it was derived. Fission is seen in the reproduction of bac- teria, growth of many algae and the formation of tissues of higher plants. Gemmation or Budding. — This is the method of reproduction common among yeasts. The cell forms a protuberance called a bud which increases in size until it equals the size of the cell which formed it and then becomes detached, although frequently not until it has developed other buds and these still others. Free Cell Formation. — This is a type of reproduction in which the nucleus and protoplasm become separated into two or more masses each of which forms a cell wall about itself. Seen in formation of ascospores within ascus of Ascomycetes and spores within spore cases of molds. Rejuvenescence. — In this mode of reproduction the protoplasm of the cell becomes rounded out, escapes by rupture of the cell wall, forms cilia and moves about as a zoospore. Later it looses its cilia, develops a cell wall and passes into a resting condition. Under favorable circumstances it grows into a new organism. It is found in CEdogonium, Ectocarpus, etc. There are two kinds of sexual reproduction, viz. : Conjugation and Fertilization. In both of these the sexual cells called gametes or VEGETABLE CYTOLOGY 43 their nuclei come together and their protoplasm blends to form a new cell. This is the common method seen in higher plants. Conjugation. — A union of two gametes, alike in character, the product being a zygote or zygospore. This method of reproduction is seen in the molds, Spirogyra, Zygnema and Diatoms. Fertilization. — A union of two unlike gametes or their nuclei, the product being an oospore. One gamete, the male sexual cell, is smaller and active while the other, the female sexual cell is larger and passive. This process is seen among the higher and many of the lower plants.' INDIRECT NUCLEAR DIVISION (MITOSIS OR KARYOKINESIS) This is the general method of division seen in the formation of tissues of higher plants. The process begins in the nucleus and ends with the formation of a cell wall dividing the new-formed cells. When we examine a cell in its resting stage we find the nucleus more or less spherical in shape, surrounded by a nuclear membrane and containing a nuclear network, nuclear sap and one or more nucleoli. The nuclear network consists of a colorless network of linin adhering to which are numerous minute granules called chro- matin which take the stain of a basic dye. Surrounding the nucleus is the cytoplasm. As the cell commences to divide, the nucleus elongates and the linin threads of the nuclear reticulum shorten, drawing the chro- matin granules together into a thickened twisted chromatic thread. This thread splits transversely and thus becomes divided into a number of rods termed chromosomes. Each of these then splits into two longitudinal halves that may be termed the daughter-chromo- somes. They lie within the nuclear cavity united by delicate threads. There now begins a phenomenon concerned with the cytoplasm which is primarily a process of spindle formation. The granular cytoplasm accumulates at the poles of the elongated nucleus forming the cytoplasmic caps. Presently it begins to show a fibrillar struc- ture, the threads extending outward around the periphery of the nucleus forming an umbrella-like arrangement of fibers from both 44 PHARMACEUTICAL BOTANY Fig. 14. — Semi-diagrammatic representation of nuclear and cell-division, a, resting cell ready to begin division; b, the nuclear reticulum is assuming the form of a thickened thread, and the cytoplasm at opposite poles is becoming thread- like to form the spindle fibers; c, the nuclear thread has divided longitudinally through the middle, and the spindle fibers have become more definite; d, the nu- clear membrane and the nucleolus have disappeared, and the nuclear thread has become segmented into chromosomes which are assembling at the equator of the cell. All of the phases of division thus far are called prophases, e, the metaphase, where the longitudinal halves of the chromosomes are being drawn apart pre- paratory to their journey toward the opposite poles; /, the anaphase, or move- ment of the chromosomes toward the poles, is about completed, connecting fibers extend from pole to pole; g, telophase where the chromosomes have begun to spin out in the form of a nuclear reticulum. The connecting fibers have begun to thicken in the equatorial plane; h, the connecting fibers have spread out and come into contact with the wall of the mother cell in the equatorial plane, and the thickening of the fibers throughout this plane has made a complete cell plate within which the dividing wall will be produced; i, a nuclear membrane has been formed about each daughter nucleus, and the dividing cell-wall is completed. The two daughter cells are now ready to grow to the size of the parent cell in a, when the daughter nuclei will appear as does the nucleus there. All highly mag- nified. (Stevens.) VEGETABLE CYTOLOGY 45 cytoplasmic caps. With the formation of fibers comes a breaking down of the nuclear membrane and in consequence the fibers enter the nuclear cavity and organize the spindle. Some of the fibers become attached to the split chromosomes and push, draw or pull them to the equatorial plate, halfway between the poles. Mean- while the nucleolus disappears. As the chromosomes line up at the equatorial plate their daughter halves are drawn apart in V-shaped fashion. The split extends and eventually one daughter-chromo- some is drawn to one pole and the remaining half to the other. At the respective poles the daughter chromosomes form a dense com- pact knot. A cell membrane, composed of material contributed largely through the shrinking of the spindle fibers, is now formed through the middle of the spindle. This soon splits to form a thin vacuole lying between the two membranes (the plasma membranes). Presently there appears within the vacuole and between the mem- branes a carbohydrate substance. On either side of this deposit the plasma membranes form a cellulose membrane. The flattened vacuole extends toward the periphery and ultimately a complete cell wall is formed. The dense compact knots of chromosomes at the poles of the spindle, that are to form the daughter-nuclei, now begin to expand and clear mesh-like spaces to appear between the expanding threads. As this process advances the chromosome substance becomes dis- tributed throughout the nuclear cavity. It is soon possible to dis- tinguish the chromatin from the linin. Eventually an irregular network of linin carrying chromatin granules is formed through the area of the nucleus. A nuclear membrane also is formed and the nucleolus reappears. The spindle fibers disappear. The daughter- nuclei increase in size and each daughter-cell formed by this process now assumes the resting stage. NON-PROTOPLASMIC CELL CONTENTS i. Sugars. — Sugars comprise a group of crystalline substances found in the cell sap of many plants either free or in combination with glucosides. They may be divided into two main groups: monosaccharoses and disaccharoses. To the former belong simple 46 PHARMACEUTICAL BOTANY sugars containing two to nine atoms of carbon, which are known respectively as bioses, trioses, tetroses, pentoses, hexoses, etc. Of these the hexoses (C 6 Hi20 6 ) are the most important and of wide distribution. Examples of the hexoses found in drug plants are: (a) dextrose (grape sugar), found in the leaves, stems, fruits, sprout- ing grains and nectaries of flowers of nearly all plants; ( b ) fructose (levulose or fruit-sugar), commonly associated with dextrose; (c) d-mannose, found in the saccharine exudation of the Manna Ash ( Fraxinus Ornus ); and ( d ) sorbinose, found in ripe Mountain Ash berries. Upon evaporating the sap or treating the parts containing these principles with alcohol they can be crystallized out. Fliickiger’s Micro-chemic Test for the determination of different kinds of sugars: Dissolve a small portion of copper tartrate in a drop of sodium hydrate on a glass slide; in this place the section and put on the cover slip. If fructose is present cuprous oxide crystals will at once be formed without warming. If grape sugar is also present a gentle warming will produce another crop of reddish-yellow crystals. If dextrin be present continued heating will still further augment the number of crystals. Cane sugar and mannite, on the other hand, will respond negatively to this test. The zymase of yeasts is capable of fermenting dextrose, levulose and d-mannose forming carbon dioxide and alcohol. Sorbinose is claimed to be non-fermentable. The disaccharoses having the chemical formula of C12H22O11 in- clude sucrose, maltose, trehalose, melibiose, touranose and agavose. Of these sucrose is the most important. It is found in the stems of sugar cane, sorghum, corn and Mexican grass; in many fleshy roots notably the sugar beet; in the sap of the sugar maple and various palms including Cocos nucifera, Phoenix sylvestris, Arenga saccharif- era; in various fruits as apples, cherries, figs, etc., in the nectaries of certain flowers ; in honey ; and in a number of seeds. It crystallizes in monoclinic prisms or pyramids. When sections of plant parts containing cane sugar are placed for a few seconds in a saturated solution of copper sulphate, then quickly rinsed in water, trans- ferred to a solution of 1 part of KOH in 1 part of water, and heated to boiling, the cells containing the sugar take on a sky-blue color. Inverlase of the yeast reduces cane sugar to dextrose and levulose VEGETABLE CYTOLOGY 47 and zymase of the same plant ferments these forming carbon dioxide and alcohol. Maltose is found in the germinating grains of barley and other cereals as a product of the action of the ferment diastase on starch. It reduces Fehling’s solution forming cuprous oxide but one-third less with equal weights. Trehalose or mycose is found in ergot, Boletus edulis, the Oriental Trehala and various other fungi. Melibiose is formed with fructose upon hydrolyzing the trisac- charose melitose which occurs in the molasses of sugar manufacture and in Australian manna. Touranose is produced upon hydrolyzing melizitose, a trisaccha- rose which occurs in Persian manna, and Agavose is found in the cell sap of the American Century Plant, Agave americana. 2 . Starch. — Starch is a carbohydrate having the chemical formula of (C 6 Hio 0 5 )„ which is generally found as the first visible product of photosynthesis in most green plants. It is found in the chloro- plasts and chromatophores of green parts in the form of minute granules. This kind of starch is known as Assimilation Starch. Assimilation starch is dissolved during darkness within the chloro- plasts by the action of ferments and passes into solution as a glucose which is conveyed downward to those parts of the plant requiring food. In its descent some of it is stored up in medullary ray cells, and in various parts of the xylem, phloem, pith and cortex in the form of small grains. Considerable, however, is carried down to the underground parts such as rhizomes, tubers, corms, bulbs or roots where the leucoplasts store it in the form of larger-sized grains called Reserve Starch. This type of starch is generally characteristic for the plant in which it is found. It constitutes stored-up food for the plant during that period of the year when the vegetative processes are more or less dormant. Structure and Composition of Starch. — Starch grains vary in shape from spheroidal to oval to chonchoidal to polygonal. They are composed of layers of soluble carbohydrate material and probably other substances called u lamellce” separated from each other by a colloidal substance resembling a mucilage in its behavior toward 48 PHARMACEUTICAL BOTANY aniline dyes. They contain a more or less distinct highly refractile point of origin or growth called the “ hilum,” which also takes the stain of an aniline dye. The layers of carbohydrate material stain variously, blue, indigo, purple, etc., with different strengths of iodine solutions. Each grain is covered with a stainable elastic membrane. Fig. 15. — Cell of Pellionia Daveauana, showing starch-grains. The black, crescent-shaped body on the end of each grain is the leucoplast. Greatly enlarged. (Gager.) Starch grains may be grouped, according to the condition in which they are found in the cells of storage regions into three kinds, viz.: simple starch grains, compound starch grains and fill starch grains. Simple starch grains are such as occur singly. Compound starch grains occur in groups of two, three, four, five, six or more and are designated as two, three, four, five, six, etc., compound, according to the number of grains making up the group. Fill starch grains are small grains filling up the spaces between the larger grains in storage cells. These are common in commercial starches. Method of Examining Reserve Starches.- — Many of the reserve starches are used commercially, such as potato, corn, rice, maranta, oat, wheat, sago, tapioca, etc., and it frequently becomes necessary for the microscopist to determine their purity or their presence in a VEGETABLE CYTOLOGY 49 sample of food or drug. The following characteristics should be noted in determining the identity or source of the starch. 1. The shape of the grain. 2. Whether simple or compound or both; if compound, the number or range in numbers of grains composing it. 3. The size of the grain in microns. 4. The position of the hilum, if distinct; whether central or excen- tric (outside of the center). 5. The shape of the hilum and the degree to which it is often fissured. 6. The nature of the lamellae, whether distinct or indistinct; if distinct whether concentric (surrounding the hilum) or eccentric (apparently ending in the margin and not surrounding the hilum), or both, as in potato starch. 7. The color of the grains when stained with dilute iodine solu- tions; whether indigo, blue, purple, red or yellowish-red, etc. 8. The appearance under polarized light. 9. The temperature at which the paste is formed. to. The consistency of the paste. Characteristics of Important Commercial Starches Potato Starch ( Solatium tuberosum) Lamellie: concentric and eccentric. Mostly simple, conchoidal or ellip- Polarization cross very distinct; soidal, with occasional spheroidal and two- to three-compound grains. beautiful play of colors with selenite plate. Size: 5 to 125/1 Maranta Starch ( Maranla Hilum: circular at smaller end of grain. arundinacea ) Fig. 16. — Potato Starch. ( X 250.) (Sayre.) Fig. 17. — Maranta Starch. (+250.) (Sayre.) 4 PHARMACEUTICAL BOTANY SO Size: io to 6 sm- Hilum: a transverse or crescent- shaped cleft in center or near broad end of grain. Lamellae: usually indistinct. Polarization cross very distinct; fine play of colors with selenite plate. Com Starch ( Zea Mays ) Polygonal to rounded, io to 35 ix. Most grains over 15/1 in diameter. Simple. Fig. 18. — Corn Starch. ( X 500.) {Sayre.) Hilum: circular or a two- to five- rayed cleft. Lamellae: indistinct. Polarization cross distinct but no marked play of color with selenite plate. Rice Starch (Oryza sativa ) Polygonal. 2 to 10 n in diameter. Simple or two- to many-compound. Fig. 19. — Rice Starch. ( X 250.) (Sayre.) Hilum: usually indistinct, occasion- ally a central cleft. Lamellae: indistinct. Polarization cross distinct but no play of colors with selenite plate. Wheat Starch ( Triticum sativum ) Lenticular grains appearing elliptical shaped on edge view; simple. 28 to 50^ in diameter. Hilum: central, appearing as dot, usually indistinct. Fig. 20. — Wheat Starch. ( X 250.) (Sayre.) Lamellae: indistinct. Polarization cross indistinct; no play of colors with selenite plate. Rye Starch ( Secale cereale ) Grains having a similar shape to those of wheat starch but many larger; simple. 20 to 6oju in diameter. Hilum: a star-shaped central cleft. Lamellae: concentric. Polarization cross distinct. Buckwheat Starch (Fagopynnn esculentum) Grains simple and compound. Simple grains polygonal or rounded polygonal. Compound grains more or less rod- shaped. 2 to 1 5 m in diameter. Hilum: central. Lamellae: indistinct. Polarization cross distinct. VEGETABLE CYTOLOGY 51 Cassava Starch ( Manihot utilissima ) Grains rounded, truncated on one side. Simple or two- to three- or four- to eight-compound. 6 to 35 m in diameter. Hilum: central, circular or triangu- lar with radiating clefts frequently. Lamellae: indistinct. Polarization cross prominent. Leguminous Starch Ellipsoidal or reniform-shaped simple grains. 50 to 100M in length. Hilum: central, elongated with branching clefts. Lamellae: distinct, concentric. Polarization crosses shaped thus,^ Canna Starch ( Canna edulis and other species of Canna) Broadly elliptical, flattened, with beak or obtuse angle at one end. 50 to 1 3 5 m in length. Hilum: excentric near narrower end. Lamellae: concentric and excentric. Polarization cross very distinct; fine plav of colors with selenite plate. 3. Inulin. — Inulin is a carbohydrate isomeric with starch which has the chemical formula of C12H20O10. It is found dissolved in the cell sap of many plants, especially those of the Composite. If pieces of a plant part containing this substance be placed directly in alcohol for at least a week, then sectioned and mounted in alcohol, sphaero- crystals of inulin will be seen applied to the walls of the cells. When these sections are treated with a 25 per cent, solution of alpha naphthol and 2 or 3 drops of strong H2SO4, the sphaerocrystals will dissolve with a violet color. Fehling’s solution is not reduced by inulin. 4. Hesperidin. — Hesperidin is a glucoside having the chemical formula of C21H26O12. Like inulin it occurs in solution within the cell sap. It is found in abundance in the Rutaceae family but occurs in many other plants. If sections of alcoholic material containing this substance such as Buchu leaves or unripe orange peel, are mounted in alcohol and examined, sphaerocrystals will be seen. If these are then treated with a drop of alpha naphthol solution and 2 or 3 drops of strong H2SO4, they dissolve with a yellow color. The same coloration is evident when 5 per cent, solution of KOH is substituted for the alpha naphthol and H2SO4. 5. Strophanthin. — This is a glucoside occurring in the cell sap of the endosperm of Strophanthus Kombe, S. hispidus and other species of Strophanthus. If sections of fresh Strophanthus seeds are mounted in a drop of water and then transferred to a drop of con- 52 PHARMACEUTICAL BOTANY centrated H2SO4, the cells containing strophanthin will assume a bright green color. 6. Salicin. — Salicin is a glucoside occurring in the cell sap of the bark and leaves of the Willows and Poplars. Sections of these mounted in concentrated H2SO4 will show a red coloration in the cells containing this substance. If water be added a red powder is thrown down. 7. Saponin, another glucoside, found in Soap Bark, Senega, Saponaria and other drugs also takes a red color with strong H2SO4. 8. Coniferin is a glucoside, occurring in the cell sap of the spruce, pine, and other plants of the Coniferce. If sections containing it are first treated with a solution of phenol and then with sulphuric acid, the cells containing it take on a deep blue color. 8. Digitoxin, a glucoside found in the leaves of Digitalis purpurea, is colored green with hydrochloric acid. The glucosides are very numerous. Those listed above represent but a few examples. They arise in the cell sap of plants containing them as products of constructive metabolism (anabolism) and are thought by many to have the function of protecting plants against the ravages of animals. Some are known to serve as reserve food. All glucosides are characterized by the property of being split up into glucose and other substances when acted upon by a ferment, dilute acids or alkalies. 9. Alkaloids. — Chemically, these are basic carbonaceous amines which like glucosides are products of metabolism. Their method of formation in plants is uncertain. Some hold that they are kata- bolic products, resulting from the breaking down of tissues, while others believe them anabolic in character. They undoubtedly serve as defensive agents in plants containing them on account of their bitter taste and poisonous properties. Properties of Alkaloids Alkaloids are invariably found in combination with acids forming salts which dissolve in water or alcohol. They are composed of carbon, hydrogen and nitrogen. Some contain oxygen. They are precipitated from saline solutions by the addition of alkalies. They VEGETABLE CYTOLOGY S 3 are mostly colorless and crystallizable. They can be precipitated by one or more of the following alkaloidal reagents: tannic acid, gold chloride, phospho-molybdic acid, picric acid and potassio- mercuric iodide. Examples of Alkaloids Strychnine. — This alkaloid, with a chemical formula of C21H22N2- O2, occurs in the seeds of Strychnos nux vomica, Strychnos Ignatii and other species of Strychnos. When sections of strychnine con- taining seeds, previously treated with petroleum ether and absolute alcohol, are mounted in a solution of 1 Gm. ammonium vanadate in 100 mils of sulphuric acid, they take on a violet-red color which later changes to brown. Veratrine. — This alkaloid, with a composition of C37H53NO11, is found in various parenchyma cells of Veratrum album. If sections of the rhizome or roots are mounted in 2 drops of water and a drop of concentrated H2SO4 and examined microscopically on a glass slide, the cell contents and walls of the cells which contain this sub- stance first take a yellow color which soon changes to an orange-red and then to a violet. Nicotine. — This is a volatile alkaloid having the formula of C10H14N2 which is found in the Nicotiana genus of the Nightshade family. Sections of tobacco leaves or stems mounted in dilute Lugol’s solution will show first a carmine-red color and then a red- dish-brown precipitate which in time loses its color. Caffeine. — This alkaloid, with a formula of C8H10N4O2 + H 2 0 , occurs in Thea, Coffea, Cola, Sterculea, Ilex and Neea. If thin sec- tions containing it are placed on a glass slide in 2 or 3 drops of con- centrated hydrochloric acid and gently heated and then 2 or 3 drops of gold chloride solution are added, the sections then pushed to the side and the liquid allowed to evaporate, slender yellowish branch- ing needles of caffeine gold chloride will be seen to separate. Cocaine.-^This narcotic alkaloid, having the formula C17H21NO4, is found in the leaves of Erythroxylon Coca and E. Truxillense. If sections of these leaves are prepared in the same manner as indicated for those containing Caffeine, but platinum chloride solution substi- 54 PHARMACEUTICAL BOTANY tuted for that of gold chloride, large feathers or plumes of cocaine- chloro-platinate will be seen separating. Aconitine 1C33H43NO12) is found in various parts of Aconitum Napellus. It is particularly abundant in the tuberous root of this plant. If sections of aconite root are treated on a glass slide with solution of potassium permanganate, a red precipitate of aconitine permanganate will appear in the cells containing this alkaloid. Colchicine (C22H 2 5N0 6 ).- — -This alkaloid occurs in the corm and seeds of Colchicum aulumnale. It is very abundant in the cells surrounding the fibro-vascular bundles of the corm. If a section of either corm or seed be treated with a mixture of 1 part of H2SO4 and 3 parts of H2O, the cells containing colchicine will be colored yellow. If a crystal of KNO3 then be added the color will change to a brownish-violet. 10. Gluco -alkaloids. — These are compounds intermediate in nature between alkaloids and glucosides having characteristics of each. To this group belongs solanine (C28H47NO11) which is found in Solanum nigrum, Solarium Dulcamara, Solarium carolinense and other species of the Solanacece. When sections of those plant parts which contain this constituent are mounted in a solution of 1 part of ammonium vanadate in 1000 parts of a mixture of 49 parts of sulphuric acid with 18 parts of water, the cells containing solanin take on a yellow color which changes successively to orange, various shades of red, blue-violet, grayish-blue and then disappears. n. Asparagine (C4H 8 N 2 + H 2 0 ). — This is an amino compound of crystalline nature which occurs widely in the plant kingdom. It has been found in certain of the slime molds and fungi, in the roots of Althcea officinalis and Atropa belladonna, in young shoots of Asparagus, in the seeds of Castanea dentata, in the tubers of Solanum tuberosum and varieties of Dahlia, and is known to play an important part in metabolism. Stevens claims that proteids are reduced for the most part to asparagine during seed germination. 1 If thick sec- tions are cut from a plant part containing this substance and mounted in alcohol, rhombohedral crystals of asparagin in the form of plates will be deposited upon the evaporation of the alcohol. If to these a few drops of a saturated solution of asparagine are added 1 Stevens’ Plant Anatomy, 3d Edit., p. 189. VEGETABLE CYTOLOGY 55 the crystals already formed will increase in size. To get satisfactory results the saturated solution must be of the same temperature as the mount. 12. Calcium Oxalate. — This substance occurs in many plants always in the form of crystals. It is apparently formed by the reac- tion of salts of calcium, which have found their way into the cell sap from the soil, with oxalic acid which is manufactured by the plant. Calcium oxalate crystals dissolve readily in mineral acids without effervescence. They are insoluble in acetic acid or water. These crystals are classified according to form and belong either to the monoclinic or tetragonal system. Crystals belonging to the Monoclinic System and Examples of Drugs Containing them: 1. Solitary — Hyoscyamus, Acer Spicalum, Viburnum Prunifolium. 2. Rosette Aggregates — Althaea, Gossypii Cortex, Stramonium, Granatum, Rheum, Foeniculum. 3. Columnar (Styloids) — Quillaja. 4. Raphides — Convallaria, Sarsaparilla, Veratrum, Scilla, Phytolacca. 5. Micro-crystals (Crystal sand) — Belladonnae Radix, Cinchona, Stramonium. 6. Crystal Fibers — Cascara Sagrada, Prunus Virginiana, Glycyrrhiza. 7. Membrane Crystals — Aurantii Dulcis Cortex, Limonis Cortex, Condurango. Solitary crystals, usually in the form of rhombohedra, occasionally in twin crystals, occur as sharp angular bodies each one often com- pletely filling up the lumen of a cell. Rosette aggregates consist of numerous small prisms or pyramids, or hemihedral crystals arranged around a central axis, appearing like a rosette or star. Columnar crystals or styloids are elongated prisms. Raphides are groups of acicular or needle-shaped crystals which occur in long thin-walled cells containing mucilage. They are more frequently found in Monocotyledons than in any other plant group. Micro-crystals (sphenoidal micro-crystals or crystal sand) are minute arrow-shaped or deltoid forms completely filling the 56 PHARMACEUTICAL BOTANY parenchyme cells in which they occur and giving these a grayish- black appearance. Crystal fibers are longitudinal rows of superimposed parenchyme cells each of which contains a single monoclinic prism or rosette aggregate. Crystal fibers are found adjacent to sclerenchyme fibers such as bast or woody fibers. Membrane crystals are monoclinic prisms, each of which is sur- rounded by a wall or membrane. In the process of formation a crystal first is formed in the cell sap and then numerous oil globules make their appearance in the protoplasm surrounding it; later some of the walls of the cell grow around the crystal and completely envelop it. 13. Cystoliths. — Cystoliths are clustered bodies formed by the thickening of the cell wall at a certain point and subsequent in- growth which latter forms a cellulose skeleton consisting of a stalk and body. Silica is subsequently deposited on the stalk while calcium carbonate is piled up on the body in layers forming an irreg- ular spheroidal or ellipsoidal deposit. These structures are abun- dantly found in the plants of the Nettle Family and constitute a leading peculiarity of the same. Hair cystoliths differ from the average type in that they are devoid of a stalk. Such are seen in the non-glandular hairs of Cannabis saliva. The calcium carbonate incrustation of a cystolith dissolves with effervescence on the addition of a mineral or organic acid. 14. Silica. — Silica (Si 0 2 ) occurs in a number of plants either as an incrustation in the cell wall as in Diatoms, the Equisetinea and Graminece or more rarely in the form “silica bodies” such as are found in certain Palms, Orchids and Tristicha. It is insoluble in all the acids except hydroflouric. It may be obtained in pure form by placing tissue containing it in a drop or two of concentrated sulphuric acid and after a time treating with successively stronger solutions of chromic acid (starting with 25 per cent.) and then wash- ing with water and alcohol. 15. Tannins. Tannins are amorphous substances occurring in plants having an astringent taste, and turning dark blue or green with iron salts. They occur in greatest quantity in the bark of VEGETABLE CYTOLOGY 57 exogens, and in gall formations. They are soluble in water, alcohol, glycerine, and a mixture of alcohol and ether. They are almost insoluble in absolute ether and chloroform. They give insoluble precipitates with organic bases such as alkaloids and with most of the salts of the heavy metals. According to their behavior with solution of iron chloride or other soluble iron salts two kinds of tannic acid are recognized: (a) a form of tannic acid giving a blue color, as that which is found in Rhus, Castanea, Granatum, Galla, etc.; ( b ) another tannic acid producing a green coloration, as that found in Krameria, Kino, Mangrove bark, Quercus, Catechu, etc. If sections are placed in a 7 per cent, solution of copper acetate for a week or more, then placed on a slide in 0.5 per cent, aqueous solution of ferric chloride, and after a while washed with water and mounted in glycerin, an insoluble brownish precipitate will be pro- duced in those cells containing tannin. 16. Proteins. — Proteins are complex nitrogenous substances forming the most important of the reserve foods of plants. They are found in all the living and many of the dead cells of plants although most abundant in seeds. Protoplasm, itself, is composed largely of these substances. They all contain carbon, hydrogen, oxygen, nitrogen and sulphur, and many contain in addition phosphorus. They are formed by the addition of nitrogen, sulphur and fre- quently phosphorus to elements of grape sugar. The nitrogen, sulphur and phosphorous elements are obtained from nitrates, sulphates and phosphates which are dissolved in the water taken in through the roots. The names of proteins recorded may be found by the hundreds. These are grouped into chemical classes, the most important of which from the standpoint of their occurrence in plants are the globulins, albumens, glulelins, nucleins, and gliadins. Of these the globulins are found most extensively. Globulins are insoluble in water but soluble in sodium chloride solutions. They do not coagulate upon the application of heat. Albumens are soluble in water and coagulate with heat. Glutelins are insoluble in water, sodium chloride solution and st rong alcohol. 58 PHARMACEUTICAL BOTANY Gliadins are nearly or wholly insoluble in water but soluble in 70 to 90 per cent, alcohol. Nucleins are insoluble in water but soluble in alkaline solutions. The following tests are of value in determining the presence of proteins. Lugol’s solution stains proteins yellow or brown. Concentrated nitric acid stains proteins yellow. This color becomes deeper upon the addition of ammonia water. Million’s reagent stains proteins a brick-red. Concentrated solution of nickel sulphate colors proteins yellow or blue. If sections are placed for an hour or two in a solution of 1 Gm. of sodium phospho-molybdate in 90 Gm. of distilled water and 5 Gm. of nitric acid, the proteid substances appear as yellowish granules. The globulins (phytoglobulins) frequently occur in bodies called “aleurone grains.” ALEURONE GRAINS Aleurone grains are small bodies found in seeds particularly those containing oil, and like starch grains often are characteristic of the genus or species. Each aleurone grain consists of a ground sub- stance (composed of amorphous proteid matter soluble in water, dilute alkali or acid), in which are usually embedded one or more phyto-globulins (insoluble in cold water, but soluble in less than 1 per cent, solution of an alkali, in dilute HCland acetic acid), one or more transparent globular globoids composed of Ca and Mg phosphate (insoluble in water and dilute potash solution but soluble in 1 per cent, acetic acid solution), and frequently a crystal of calcium oxalate, the whole being enclosed by a protoplasmic membrane (soluble in water). The proteins insoluble in the cell-sap water are made soluble for translocation by means of proteolytic enzymes which change them into proteoses and peptones. 17. Mucilages and gums are those substances occurring in plants which are soluble in water or swell in it, and which are precipitated by alcohol. VEGETABLE CYTOLOGY 59 Mucilage is formed in plants in several ways, viz.; either as a product of the protoplasm, as a disorganization product of some of the carbohydrates, as a secondary thickening or addition to the cell wall, or as a metamorphosis of it. In the first two cases the mucilage is called cell-content mucilage; in the last two, membrane mucilage. Mucilage is stored as reserve food in the tubers of Salep and many other Orchids and also in the seeds of some species of the Leguminosae. Cell-content mucilage has been found in the leaves of Aloe, the rhizomes of Triticum, the bulb scales of Squill and Onion and in certain cells of many other Monocotyledons especially those containing raphides. Membrane mucilage has been observed in Barosma, Ulmus, Althaea , Linum, Astragalus, and Acacia species, in the Blue-green Algae, and many of the Brown and Red Algae. When mucilage is collected in the form of an exudate from shrubs and trees it constitutes what is termed a gum. Many of these gums are used in pharmacy, medicine and the arts. The three most im- portant from a pharmaceutical standpoint are: Acacia, yielded by Acacia Senegal and other species of Acacia; Tragacanth, yielded by Astragalus gummifer and other Asiatic species of Astragalus; and Cherry Gum, obtained from Prunus Cerasus and its varieties. Mucilage may be demonstrated in plant tissues containing it by placing sections of these in a deep blue solution of methylene-blue in equal parts of alcohol, glycerin and water on a glass slide, allowing them to remain in the solution for several minutes, then draining off the stain and mounting in glycerin. Those cells containing muci- lage will exhibit bluish contents. 1 8. Fixed Oils and Fats. — These are fatty acid-esters of glycerin which are found in the vacuoles of cells or formed within the cell walls from which they may be liberated as globules upon treating sections vvith chloral hydrate or sulphuric acid or heating them. They are quite soluble in ether, chloroform, benzol, acetone and volatile oils but' insoluble in water and with the exception of castor oil insoluble in alcohol. They are readily distinguished from the volatile oils in that they leave a greasy stain upon paper which does not disappear. Fixed oils and fats take a brownish to black color 6o PHARMACEUTICAL BOTANY with osmic acid, a red color with alkannin or Sudan III and a blue color with cyanin. In Vaucheria, the Diatoms and a few of the other Thallophytes, fixed oil is formed in the chromatophores in- stead of starch as the first visible product of photosynthesis. In higher plants it is generally found in storage regions such as the parenchyma seeds, fruits and the medullary ray cells and paren- chyma of barks, roots and rhizomes. 19. Volatile Oils. — These are volatile odoriferous principles found in various parts of numerous plants which arise either as a direct prod- uct of the protoplasm or through a decomposition of a layer of the cell wall which Tschirch designates a “ resinogenous layer.” They are readily distilled from plants together with watery vapor, are slightly soluble in water but very soluble in fixed oils, ether, chloro- form, glacial acetic acid, naphtha, alcohol, benzin and benzol. They leave a spot on paper which, however, soon disappears. They respond to osmic acid, alkannin, Sudan III, and cyanin stains similar to the fixed oils and fats. Volatile oils may be grouped into four classes: A. Pinenes or Terpenes, containing carbon and hydrogen and having the formula of Ci 0 H 16 . Examples: Oil of Turpentine and various other volatile oils occurring in coniferous plants. B. Oxygenated oils, containing carbon, hydrogen and oxygen. Examples: Oil of cassia and other cinnamons. C. Nitrogenated oils, containing carbon, hydrogen and oxygen with nitrogen (from HCN). Example: Oil of Bitter Almonds. D. Sulphurated oils, containing carbon, hydrogen and sulphur. Example: Volatile oil of mustard. 20. Resins, Oleoresins, Gum Resins, and Balsams. — These sub- stances represent products of metabolism in many plants which are formed either normally as Turpentine, Asafcetida, Mastiche, etc., or as a result of pathological processes through injury to the plant tissues as Styrax, Benzoin, Balsam of Tolu and Peru, etc. They occur usually in special cavities such as secretion cells, glands, or secretion reservoirs. Resins are insoluble in water but mostly soluble in alcohol. They combine with alkalies to form soap. Many of them are oxidized oils of plants. Examples: Guaiacum, Resina. VEGETABLE CYTOLOGY 6l Oleoresins are mixtures of oil and resin. Examples: Terebin- thina, Terebinthina Canadensis. Gum resins are natural compounds of resin, gum and oil. Ex- amples: Asafoetida, Myrrha, Cambogia. Balsams are mixtures of resins with cinnamic or benzoic acid or both and generally a volatile oil. Examples: Balsamum Tolu- tanum, Styrax, Balsamum Peruvianum. If sections of a resin containing plant part are placed in a saturated aqueous solution of copper acetate for a week or two and mounted in dilute glycerin, the resin will be stained an emerald green. 21. Pigments. — These are substances which give color to various plant parts in which they are found. They occur either in special protoplasmic structures as chloroplasts, chromoplasts or chroma- tophores or, dissolved in the cell sap. Of the pigments named the following will be considered: Chlorophyll, Xanthophyll, Chromo- phyll, Etiolin, Anthocyanin, Phycocyanin, Phycophaein and Phy- coerythrin. Chlorophyll is the yellowish-green pigment found in the chloro- plastids or chromatophores of leaves or other green parts of plants. Its composition is not definitely known although it yields products similar to the haemoglobin of the blood when decomposed. Iron is known to be essential to its formation. If an equal portion of xylene be added to a fresh alcoholic solution of chlorophyll and the mixture shaken, The chlorophyll in solution will break up into a yellowish and greenish portion. The greenish portion dissolves in the xylene which rises forming the upper stratum while the yellowish portion dissolves in the alcohol forming the lower stratum. To this isolated greenish portion of chlorophyll has been given the name of “chlorophyllin” while the yellowish portion has been designated “xanthophyll.” Chlorophyllin when examined spectroscopically produces absorp- tion bands in the red, orange, yellow and green of the spectrum, the broadest and most distinct band being in the red. Chromophyll also called “ xanthophyll ” and “carotin” is the yel- low or orange pigment found in chromoplastids. By some the term carotin is limited to the orange pigment found in the carrot. Sulphuric acid forms a blue color with chromophyll. 62 PHARMACEUTICAL BOTANY Etiolin is a pale yellow pigment which appears when green plants are kept for some time in darkness. It is probably identical with xanthophyll. Anthocyanins are applied to the blue, purple and red pigments which occur in the cell sap. The character of the color is claimed to be due to the alkalinity or acidity of the cell sap. Phycocyanin is the blue pigment found in the blue-green algae, associated with chlorophyll. It is soluble in water. Phycophaein is the brown pigment found in the brown algae. Phycoerythrin is the red pigment found in many of the red algae. The last two are always associated with chlorophyll but frequently conceal it. 22. Latex. — This is an emulsion of varying composition and color found in special passages as latex cells and laticiferous vessels of many plants. It may contain starch, sugar, proteid, oil, enzymes, tannins, alkaloids, gum, resins, caoutchouc and mineral salts. The color may be absent as in Oleander; whitish as in Asclepias, Papaver, Hevea, and Apocynum; yellowish to orange as in Celandine, or red as in Sanguinaria. Chlor-zinc-iodine solution imparts to latex a wine red color. The latex of the following plants is of value to pharmacy and the arts: Papaver somniferum and its variety album which yields Opium. That from the unripe capsules is alone used for this drug. Palaquium Gutla which yields Gutta Percha. Hevea species, Ficus elaslica, Landolphia species, Castilloa elastica, Hancornia speciosa, Forsteronia species, Funlumia elastica and F. africana, Manihot species, Clitandra species and various species of Euphorbia furnish most of the Rubber of commerce. Lactuca virosa and other species of Lactuca yield the drug Lactu- carium. 23. Enzymes. — An enzyme or ferment (according to Hepburn) is a soluble organic compound of biologic origin functioning as a thermolabile catalyst in solution. Ostwald has defined a catalyst as an agent which alters the rate of a reaction without itself entering into the final product, or which does not appear to take any imme- diate part in the reaction, remains unaltered at the end of the reac- VEGETABLE CYTOLOGY 63 tion and can be recovered again from the reaction product un- altered in quantity and quality. The biologic catalysts (enzymes) differ from the inorganic catalysts in that they are sensitive to heat and light. According to Haas and Hill they are destroyed at ioo°C. and most of them cannot be heated safely above 6o°C. Enzymes are soluble in water, glycerin or dilute saline solutions. They are stimulated to activity by substances known as “ activators ” and their activity is checked by other substances called “paralyzers.” Fre- quently the paralyzers consist of products of enzyme action. Cold inhibits and warmth accelerates enzyme action. Moisture must always be present for enzymic activity. CLASSIFICATION OF ENZYMES A. According to Difusibility through Cell Wall. Endocellular : Those that cannot diffuse out of the cell. Example : Zymase of Yeast. Extracellular: Those that can diffuse out of the cell. Example: Inver tase of Yeast. B. According to Kind of Substances Acted upon and Transformed. 1. Carbohydrate enzymes: Diastase found in the germinating seeds of barley and other grains and in Aspergillus oryzce, etc., converts starch to maltose and dextrin. Invertase, secreted by yeasts, and found in younger parts of higher plants, transforms cane sugar, producing dextrose and levulose. Maltase, found in malt and Saccharomyces octosporus, transforms maltose to dextrose. Trehalase, found in Polyporus, hydrolyzes trehalose to dextrose. Cytase, found in Nux Vomica seeds, in barley, dates, etc., decom- poses hemicellulose and cellulose to galactose and mannose. Lactase, found in Kephir grains, hydrolyzes lactose to dextrose and galactose. Inulase, found in Compositaceous plants, transforms inulin to levulose. Zymase, found in yeast, hydrolyzes glucose (dextrose and levulose) to alcohol and carbon dioxide. 64 PHARMACEUTICAL BOTANY 2. Fat and Oil Ferment : Lipase splits up fats and oils into fatty acids and glycerin. It is found in various mildews, molds and numerous oily seeds and other fatty-oil storage regions of higher plants. 3. Proteinaceous Ferments : Pepsin converts proteids into proteoses and peptones. Trypsin, found in yeast, Boletus edulis, Amanita species, etc., resolves proteins to peptones and amino-acids. Bromelin, found in the fruit of the Pineapple and Papayin (Pa- pain), found in the latex of the fruit of the Papaw, act similarly to trypsin. Nepenthin, found in the pitchers of Nepenthes species, acts simi- larly to pepsin. 4. Glucoside Ferments : Emulsin (synaptase), found in the seeds of the Bitter Almond, Cherry Laurel leaves, in the barks of the Wild Black Cherry and Choke Cherry and in other Rosaceous plant parts, in Manihot utilissima, Polygala species, etc., hydrolyzes the glucoside present (either amygdalin or 1-mandelonitrile glucoside) to hydrocyanic acid, benzaldehyde and glucose. Myrosin (myronase), found in the seeds of Brassica nigra and other members of the Cruciferce, converts the glucoside, Sinigrin, into allyl- iso-sulphocyanide and glucose. Rhamnase, found in Rhamnus Frangula and probably other species of Rhamnus, hydrolyzes the glucoside frangulin to rhamnose and emodin. Gaultherase, found in Gaultheria procumbens and other Erica- ceous plants, resolves the glucoside, gaultherin, to methyl-salicylate and glucose. CELL WALLS The cell walls of plants make up the plant skeleton. They are all formed by the living contents of the cells (protoplasts) during cell- divisions. In most plants the cell wall when first formed consists of cellulose, (CbHjoOs)*, a carbohydrate, or closely allied substances. It may remain of such composition or become modified to meet certain VEGETABLE CYTOLOGY 65 functions required of it. Thus, in the case of outer covering cells as epidermis and cork, whose function is that of protecting the under- lying plant units, the walls become infdtrated with cutin and suberin, waxy-like substances, which make them impermeable to water and gases as well as protect them against easy crushing. Again, in the case of stone cells and sclerenchyme fibers whose function is that of giving strength and support to the regions wherein found, the walls become infiltrated with lignin which increases their strength, hard- ness, and in the case of sclerenchyme fibers, their elasticity also. Moreover, in the case of the cells comprising the testa or outer seed coat of the pumpkin, squash, mustard and flax, etc., whose function is that of imbibing quantities of water, the walls undergo a muci- laginous modification. Growth in Area and Thickness. — The cell wall when first formed is limited in both extent and thickness. As the protoplast within enlarges new particles are placed within the wall by the process called intussusception. This increases its area. New particles, also, are deposited on its surface which gradually increases its thickness. The latter process is known as growth by apposition. 5 66 PHARMACEUTICAL BOTANY Various Kinds of Cell Walls and Behavior of Each to Micro-Chemic Reagents Nature of wall Where found Reagent and behavior toward same Cellulose Parenchyme cells, tri- chomes such as cotton, etc. Cuoxam dissolves it. Chlorzinc- iodine solution imparts a blue or violet color. Iodine solution followed by sulphuric acid colors it blue. ■ Lignocellulose (Lignified wall). Woody parts of plants, such as stem cells, bast fibers, wood fibers, etc. Phloroglucin with HC 1 imparts a red color except to bast fibers of flax. Corallin-soda solution imparts pink color. Aniline sulphate with H2SO4 colors it a golden-yellow. Chlorzinc- iodine imparts a yellow color. Reserve cellulose Found in certain seeds such as nux vomica, ivory nut, date, cof- fee, etc. As for cellulose. Mucilaginous modification of cellulose. In various parts of plants. Alcoholic or glycerin solution of meth- ylene-blue imparts a blue color. Suberized walls. . In cork, wounded areas of plants, endo- dermis. Alcoholic extract of chlorophyll in the dark imparts a green color. Alcannin and Sudan III impart a red colora- tion. Converted into yellowish drop- lets and granular masses upon heating with a strong solution of KOH. Cutinized walls. . Forming outer walls of many epidermal cells. * As for suberized walls. Callus of sieve plates. Silicified walls. . . Plates of sieve tubes. Epidermis of Equi- setaceae, Grammes, etc.; Diatoms. Corallin-soda solution imparts pink color. Soluble in hydrofluoric acid. CHAPTER VI PLANT TISSUES A tissue is an aggregation of cells of common source, structure and function in intimate union. THE TISSUES OF SPERMATOPHYTES AND PTERIDOPHYTES The tissues of seed plants and pteridophytes are all derived from a fertilized egg (oospore) which has undergone repeated divisions. At first either an apical cell arises or a mass of cells is formed which are essentially alike, but gradually we find that a division of labor has become operative setting aside many different groups of cells, each group of which has its particular role to perform in the economy of the whole. Each group of cells similar in source, structure and function is called a tissue. The tissues found in higher plants range from those whose component cells are more or less rounded, in a rapid state of division, and whose thin cellulose cell walls enclose a mass of protoplasm devoid of vacuoles or with exceeding small ones to those whose cells through various physical and chemical factors become compressed, elongated, and highly modified in respect to their contents and walls. As was shown by Hanstein , 1 the embryo of Angiosperms, while still constituted of only a few cells in the process of division, becomes differentiated into three layers of cells which differ in their arrange- ment and direction of division; these were called by him, Dermatogen , Periblem and Plerome. In roots a fourth layer of cells is sometimes evident at the apex. This was termed by Janczewski 2 the Calyptro- gen layer. These primary layers or groups of cells are called primary meristems or generative tissues. They are composed of more or less 1 Hanstein, “Die Scheitelzellgruppe im Vegetationspunkt der Phanerogamen,” Bonn, 1868. 2 Am. Sci. Nat. 5 s6rie, tom. xx. 67 68 PHARMACEUTICAL BOTANY rounded cells having delicate cell walls of cellulose which enclose protoplasm and nucleus and wherever found in living embryos are in a rapid state of division. The generative tissues are found in the growing apices of plant organs such as root, stem and leaf apex. By the division and redivi- sions of their cells they give rise to the mature or adult tissues of plants. 1. Dermatogen originates epidermal tissue and derivative struc- tures such as stomata, non-glandular and glandular hairs, glands, and cork cambium. 2. Periblem originates cortex tissue, chlorophylloid cells (chlor- enchyma) colloid cells (collenchyma), strengthening cells (scleren- chyma), crystal cells (raphiderchyma) latex cells (lacterchyma), endodermis and cork cambium. 3. Plerome originates fibro-vascular bundles, fundamental tissue, pericambium and cambium. According to structure the following tissues are found in various forms of higher plants: 1. Meristem 7. Cork 2. Parenchyma 8. Laticiferous tissue 3- Collenchyma 9. Cribiform or sieve tissue 4- Sclerenchyma 10. Tracheary tissue 5- Epidermis n. Medullary rays 6. Endodermis MERISTEM Meristem, frequently called embryonic tissue, is undifferentiated tissue composed of cells in the state of rapid division. It is found in the growing apices of roots, stems and leaves and is in these regions called primary meristem, since it is the first meristem to appear. Such meristem gives rise to the permanent or mature tissues of plants and retains the power of independent growth and capacity for division as long as the plant part survives which con- tains it. Meristem is also found in other regions of plant organs such as the cambium, cork cambium and pericambium and is there called secondary meristem. Secondary meristem loses with its de- velopment the power of division and independent growth. PLANT TISSUES 69 PARENCHYMA Parenchyma or Fundamental Tissue is the soft tissue of plants consisting of cells about equal in length, breadth and thickness (isodiametric) with thin cellulose cell walls enclosing protoplasm and a nucleus and frequently substances of a non-protoplasmic nature. There are four generally recognized types of parenchyma, viz. : O rd inary Parenchyma (S of t Groun d Tissue , F undamen tal Tissue ) . — Next to the meristem this is the least modified of all plant tissues. It is composed of thin-walled cells, commonly polyhedral or sphe- roidal in form and often of approximately the same length, breadth, and thickness (isodiametric), the cell walls are composed of cellu- lose which is usually unmodified. Protoplasm and a nucleus are always present, but in old cells are only seen as a thin layer pushed up against the cell wall. Ordinary Parenchyma may be seen composing the soft tissues of roots, stems, and barks. Assimilation Parenchyma (Chlorophyll or Chromophyll Paren- chyma, Chlorenchyma).— This form of parenchyma tissue is found in foliage leaves, floral leaves, in the outer region of young green stems and fruits. Its cells are thin walled and vary in shape from more or less isodiametric to irregular and elongated forms. The cells always contain chloroplasts or plastids in whose pores may be found some other coloring substance. Conducting Parenchyma. — This type of parenchyma functions in the rapid translocation of food materials to distant regions in the plant. It includes the wood parenchyma cells of the xylem which convey a portion of the crude sap (water with mineral salts in solution) and the phloem parenchyma (soft bast) which transports the elaborated sap (carbohydrate and proteid material in solution). Conducting parenchyma cells differ from those of ordinary paren- chyma in being usually more elongated and in conducting soluble food materials with greater celerity. Reserve Parenchyma. — This resembles ordinary parenchyma in many particulars of structure but differs from it mainly by its cells being filled with starch, protein crystals, or oil globules. It is usually found in seeds, fleshy roots, or underground stems such as tubers, corms, and bulbs. 7 ° PHARMACEUTICAL BOTANY Collenchyma. — This form of tissue is characterized by its cells being prismatic, more elongated than ordinary parenchyma, and thickened in their angles with a colloidal substance. The cells, like those of parenchyma tissue contain protoplasm and a nucleus, and frequently chloroplasts. Collenchyma is generally found underneath the epidermis, and gives strength to that tissue. It is frequently observed forming the “ribs” of stems and fruits of the Fig. 21. — Stone cells from different sources, i. From coffee; 2, 3, and 4, from stem of clove; 5 and 6, from tea leaf; 7, 8, and 9, from powdered star-anise seed. ( Stevens , after Moeller.) Umbelliferce and “ribs” of stems of the Labiates. In many leaves it has been found as the supporting and strengthening tissue between the stronger veins and the epidermis. Sclerenchyma or stony tissue comprises a variety of supporting elements having thickened cell walls composed of lignocellulose. When first formed these cells resemble those of ordinary parenchyma in having walls of pure cellulose, but later lignin becomes deposited on the inner surface of the walls in one or more layers. (Occasion- ally as in the rhizomes of Ginger no lignin is deposited on the walls PLANT TISSUES 71 of the sclerenchyma fibers). When sclerenchyma is composed of cells which are more or less isodiametric or moderately elongated, with thickened lignified walls and conspicuous pores, its elements are called Stone Cells. Stone cells are distributed in fruits, seeds and barks of many plants, rarely in woods. They have been found forming the gritty particles in the “flesh” of certain fruits as the Pear, the endocarp or stone region of drupaceous fruits as the Olive, Peach, Cubeb, Pepper, etc., the hard portions of seed coats as in Physostigma, Walnuts, etc. Each stone cell presents for examina- tion a cell wall of cellulose with one or several layers of lignin on its inner surface which surround a central lumen. The latter is in communication with radial pore canals leading outward to the middle lamella. Longitudinal pore canals are also evident. When sclerenchyma is composed of cells which are greatly elon- gated and more or less obtusely or taper ended, its component ele- ments are termed Sclerenchyma fibers. These fibers are frequently spindle-shaped, contain air and exhibit oblique slits in their walls. They are either polygonal, rectangular or somewhat rounded in trans- verse section. They occur in various parts of roots, stems, leaves, fruits and seeds as supporting elements. When sclerenchyma fibers occur in the xylem region of fibro-vascular bundles they are termed Wood Fibers; when they appear in the phloem region, Bast Fibers. EPIDERMIS Epidermis is the outer covering tissue of a plant and is protective in function. Its cells may be brick-shaped, polygonal, equilateral or wavy in outline. Their outer walls are frequently cutinized (infil- trated with a waxy-like substance called cutin). Among the epider- mal cells of leaves and young green stems may be found numerous pores or stomata (sing, stoma) surrounded by pairs of crescent- shaped cells, called guard cells. • The stomata are in direct commun- ication with air chambers beneath them which in turn are in com- munication with intercellular spaces of the tissue beneath. The function of the stomata is to give off watery vapor and take in or give off carbon dioxide, water and oxygen. In addition to stomata some leaves possess groups of water stomata which differ from trans- 72 PHARMACEUTICAL BOTANY piration stomata in that they always remain open, are circular in out- line, give off water in droplets directly, and lie over a quantity of small-celled glandular material which is in connection with one or more fibro-vascular bundles. Examples: Leaves of Crassula, Saxi- fraga and Ficus. The epidermis of leaves, stems, fruits, and seeds of many plants frequently give rise to outgrowths in the form of papillae, hairs and scales. Epidermal papilla are short protuberances of epidermal Fig. 22. — Upper epidermis of Comptonia aspleni folialeat (surface view) show- ing epidermal cells and two non-glandular trichomes. cells. They may be seen to advantage on the upper epidermis of the ligulate corolla of various species of Chrysanthemum, on the lower epidermis of the foliage leaves of species of Erythroxylon and upon the upper epidermis of the petals of the Pansy ( Viola tricolor). Epi- dermal hairs or trichomes are more elongated outgrowths of one or more epidermal cells. They may be unicellular (Cotton) or multicellular, non-glandular (simple) or glandular. The non- glandular hairs may be of various shapes, viz.: clavate (club-shaped) as on Rhus glabra fruits; stellate (or star-shaped) as on Deutzia leaves; candelabra-shaped, as on Mullein leaves; filiform as on PLANT TISSUES 73 Hyoscyamus, Belladonna and Digitalis leaves; hooked, as on stems of Phaseolus multi florus or Hops; barbed, as on the stems of Loasa species; or tufted, as found on the leaves of Marrubium vulgare. They may be simple as in Cotton, etc., or branched as in Hyoscyamus muticus. The glandular hairs comprise those whose terminal cell or cells are modified into a more or less globular gland for gummy, resinous or Fig. 23. — Lower epidermis of Comptonia asplenifolia leaf (surface view) showing stomata (s) scattered amongst epidermal cells. To the left is to be noted the globular head of a glandular hair. (Photomicrograph). oily deposits. They are generally composed of a stalk and a head region although rarely the stalk may be absent. The stalk may be unicellular, bicellular or uniseriate (consisting of a series of super- imposed cells). The head varies from a one- to many-celled struc- ture. The drug Lupulin consists of the glandular hairs separated from the strobiles of HumuJus lupulus. Scales are flat outgrowths of the epidermis composed of one or several layers of cells. They occur attached to the stipes of Aspid- 74 PHARMACEUTICAL BOTANY ium, Osmunda and other ferns where they are called “chaff scales.” They are also found on a number of higher plants. Plant hairs are adapted to many different purposes. They may absorb nourishment in the form of moisture and mineral matter Fig. 25. Fig. 24. Fig. 24. — 1, Epidermis of oak leaf; 2, epidermis of Iris leaf, both viewed from the surface; 3, group of cells from petal of Viola tricolor; 4, two epidermal cells in cross-section showing thickened outer wall differentiated into three layers, namely, an outer cuticle, cutinized layer (shaded), and an inner cellulose layer; S and 6, epidermal outgrowths in the form of scales and hairs. (1, 2, 6 after Stevens, 3 after Strasburger, 4 after Sachs, and 5 after de Bary.) Fig. 25. — Different forms of epidermal outgrowths. 1, Hooked hair from Phaseolus multiflorus; 2, climbing hair from stem of Humulus Lupulus; 3, rod- like wax coating from the stem of Saccharum officinarum; 4, climbing hair of Loasa hispida; 5, stinging hair of Urtica urens. {Fig- 3 after de Bary; the re- mainder from Haberlandl.) in solution, e.g., root hairs. Those which serve as a protection to the plant may be barbed and silicified, rendering them unfit for animal food, or, as in the nettle, charged with an irritating fluid, penetrating the skin when touched, injecting the poison into the wound. A dense covering of hairs also prevents the ravages of insects and the clogging PLANT TISSUES 75 of the stomata by an accumulation of dust. They fill an important office in the dispersion of seeds and fruits, as with their aid such seeds as those of the milkweed and Apocynum are readily scattered by the wind. The reproductive organs of many Cryptogams are modified hairs, as the sporangia of Ferns. ENDODERMIS Endodermis is the “starch sheath” layer of cells, constituting the innermost layer of the cortex. In Angiospermous stems it usually resembles the other parenchyme layers of cortex as to struc- tural characteristics, save that it frequently contains more starch. In fern stems and roots of Angiosperms, however, its cells are clearly distinguished from the other cells of the primary cortex by their elongated form and suberized radial walls. In the roots of Mexican Sarsaparilla the inner as well as the radial walls are suberized; in those of the Honduras variety, inner, radial and outer walls all show suberization. Endodermal tissue is devoid of intercellular air spaces. Its cells contain protoplasm and nucleus. Its functions seem to be to give protection to the stele (tissues within it) and to reduce permeability between primary cortex and stele. CORK Cork or suberous tissue is composed of cells of tabular shape, whose walls possess suberized layers. Its cells are mostly filled with air containing a yellow or brownish substance. It is derived from the phellogen or cork cambium which cuts off cork cells outwardly. Cork tissue is devoid of intercellular-air-spaces. It forms a protective covering to the roots of secondary growth, stems (after the first season) of Dicotyledons and Gymnosperms, and wounds of stems and branches. Living cork cells contain protoplasm and cell sap while dead cork cells are filled with air. LATICIFEROUS TISSUE This form of tissue comprises either latex cells, laticiferous vessels, or secretory cells differing from each other in origin and method of development. Latex cells are elongated tubes which take their 76 PHARMACEUTICAL BOTANY origin from meristematic cells of the embryo. Elongating with the growth of the plant, they branch in various directions and traverse Fig. 26. — Laticiferous vessels from the cortex of root of Scorozonora hispanica. A . As seen under low power, and B, a smaller portion under high power. (Stevens, after Sachs.) at maturity all of its organs. Such cells are abundant in the following families: Apocynacece, Asclepiadacece, Urticacece and Euphorbiacea. PLANT TISSUES 77 Laticiferous vessels are long simple or branching tubes, which owe their origin to chains of superimposed cells whose transverse walls have early become absorbed, the lumina of the cells then becoming filled with latex. They are found in various parts of roots, stems, and leaves. When branched the branches connect with those of other tubes forming anastomosing network. These vessels occur in the following families: Composites, Papa- veraceee, Campanulacece, Convolvula- cece, Euphorbiacece, Aracece, Oleacece, Geramacece, and Musacece. Secretory cells with a latex-like content are probably of secondary origin in plants. They resemble in many respects latex cells and are seen in various species of the Celastracece, Urticacece, Tiliacece, and Oleacece families. All laticiferous elements contain a colorless, milky-white, or otherwise colored emulsion of gum-resins, fat, wax, caoutchouc and in some cases, alkaloids, tannins, salts, ferments, etc. This emulsion is called “latex.” SIEVE (LEPTOME OR CRIBIFORM) TISSUE This tissue found in the phloem (rarely in the xylem) region of fibro- Fig. 27.- — Stages in the develop- ment of sieve tubes, companion cells, and phloem parenchyma. A, a and b, Two rows of plerome cells; in c and d, a has divided longitudi- nally and c is to become companion cells; d, a sieve tube, and b, phloem parenchyma. B, c, Companion cells, and d, a beginning sieve tube from c and d, respectively in A. The cross- walls in d are pitted; b, phloem parenchyma grown larger vascular bundles consists of superim- than in A. C, The same as B with , , . j , , t n 1 the pits in the cross-walls of the posed, elongated, tubular cells whose sieve tubes become perforations, longitudinal walls are thin and com- and the nuclei gone from the cells , r ,,, , , , composingthe tube. (From posed of cellulose and whose trans- stevens.) verse walls, called “ sieve plates ” are perforated, permitting of the passage of proteids from one cell to another. Occasionally sieve plates are formed on the longitudinal walls. Sieve tubes are usually accompanied by companion cells ex- 78 PHARMACEUTICAL BOTANY cepting in Pteridophytes and Gymnosperms. Both companion cells and sieve tubes arise by the division of the same mother-cell. The companion cells may be distinguished from the sieve tubes by their . , abundant protoplasmic contents, and also by the fact that they retain their nuclei after complete maturation. Be- sides sieve tubes, companion cells, and bast fibers, parenchyma cells are often found in the phloem. TRACHEARY TISSUE Fig. 28. — Vascular ele- ments. A , annular tracheal tube; B, spiral trachea tube; C, reticulated tracheal tube; D, pitted tracheal tube; E, cross-section through plate of sieve tube, and adjoining companion cell; F, length- wise section of sieve tube; G, portions of two companion cells. (A, B, C, D, Robbins; E, F, and G, after Stras- burger.) The tracheary tissue of plants comprises two kinds of elements, the trachea (ducts or vessels) and tracheids. Both of these conduct crude sap (water with mineral salts in solution). The trachea are very long tubes of a cylindrical or prismatic shape which are formed by the disintegra- tion of the transverse walls between cer- tain groups of superimposed cells, during the growth of the plant. The tubes frequently retain some of their transverse walls. The longitudinal walls of these tubes are of varying thickness, usually, however, thinner than those of woody fibers. The thickness is due to an infil- tration of lignin upon the original cel- lulose wall. The walls show character- istic thickenings on their inner surfaces. Tracheae are classified according to their markings as follows: Annular, with ring-like thickenings. Spiral, with spiral thickenings. Reticulate, with reticulate thickenings. Porous or pitted with spherical or oblique slit pores. Annulo-spiral, with both ring and spiral thickenings. Scalariform, with ladder-like thickenings. 79 " PLANT TISSUES D Fig. 29. — Stages in the development of the elements of the xylem. A, pro- gressive steps in the development of a tracheal tube. 1, Row of plerome or cambial cells that are to take part in the formation of a tube; 2, the same at a later stage enlarged in all dimensions; 3, the cells in 2 have grown larger, their cross-walls have been dissolved out, and the wall has become thickened and pitted; 4, the walls in 3 have become more thickened, the pits have an overhang- ing border, the walls have become lignified as indicated by the stippling, and finally the protoplasts have disappeared, and the tube is mature and dead. B, Stages in the formation of tracheids from plerome or cambial cells. The steps 8o PHARMACEUTICAL BOTANY Tracheids are undeveloped ducts having bordered pores and fre- quently scalariform thickenings. Like trachese their walls give the characteristic lignin reaction with phloroglucin and HC1. The Fig. 30. — Closed collateral bundle of stem of Zea mays. VG, Bundle sheath; L, intercellular space; A, ring from an annular tracheal tube; SP, spiral tracheal tube; M, pitted vessels; V, sieve tubes; 5 , companion cells; CP, crushed primary sieve tubes; F, thin-walled parenchyma of the ground or fundamental tissue. C From Sayre after Strasburger.) bordered pores exhibit a wall surrounding the pore which forms a dome-shaped protrusion into the cell. Like tracheae, also, tracheids convey water with mineral salts in solution. Tracheids and medul- lary rays make up most of the wood of Gymnosperms. are the same as in A, excepting that the cross- walls remain and become pitted. C, steps in the development of wood fibers from cambial cells. 1, Cambial cells; 2, the same growth larger in all dimensions with cells shoving past each other as they elongate; 3, a later stage with cells longer and more pointed and walls becoming thickened and pitted; 4, complete wood fibers with walls more thick- ened than in the previous stage and lignified, as shown by the stippling. The protoplasts in this last stage have disappeared and the fibers are dead. D, steps in the formation of wood parenchyma from cambial or procambial cells. 1, Group of cambial or plerome cells; 2, the same enlarged in all dimensions; 3, the same with walls thickened and pitted; 4 and 5 show the same stages as 2 and 3, but here the cells have enlarged radially or tangentially more than they have vertically. The walls of these cells are apt to become lignified, but the cells are longer lived than the wood fibers. ( From Stevens.) PLANT TISSUES MEDULLARY RAYS These are bands of parenchyma cells which extend radially from the cortex to the pith (primary medullary rays) or from a part of the xylem to a part of the phloem (secondary medullary rays). Their primary function is to supply the cambium and wood with elabo- rated sap formed in the leaves and conveyed away by the sieve tubes, Fig. 31. — Transverse section of a concentric bundle from the rhizome of Iris (a monocotyledon). Xylem surrounding the phloem, t, Tracheae; t 1 , proto- xylem; s, sieve tubes; g, companion cells of the internal phloem portion. ( From Sayre after Vines.) and phloem parenchyma and to supply the cambium and phloem with crude sap which passes up mainly through the tracheae and tracheids from the absorptive regions of the roots. They further- more serve as storage places for starch, alkaloids, resins, and other substances. Fibro-vascular Bundles are groups of fibers, vessels and cells cours- ing through the various organs of a plant and serving for conduction and support. According to the relative structural arrangement of their xylem and phloem masses they may be classed as follows: I. Closed collateral, consisting of a mass of xylem lying alongside of a mass of phloem, the xylem facing toward the center, the phloem 6 82 PHARMACEUTICAL BOTANY facing toward the exterior. Stems of most Monocotyledons and Horsetails. II. Open collateral, consisting of a mass of xylem facing toward the pith and a mass of phloem facing toward the exterior and sepa- rated from each other by a cambium. Stems and leaves of Dicoty- ledons and roots of Dicotyls and Gymnosperms of secondary growth. Fig. 32.- — Cross-section through a portion of a root of Acorus calamus. A. Cortical parenchyma; B, endodermis; C, pericycle; E, phloem; F, xylem. At F, Y, are large tracheal tubes, which were formed last, the narrow tubes near the periphery of the xylem being formed first. At the center of the root, within the circle of the vascular bundle, occur thin-walled parenchymatous pith cells. (From Sayre after Frank.) III. Bicollateral, characterized by a xylem mass being between an inner and an outer phloem mass. There are two layers of cam- bium cells, one between the xylem and inner phloem mass, the other between the xylem and outer phloem mass. Seen chiefly in stems and leaves of the Cucurbitacece and Solanacece. ' IV. Concentric, characterized by a central xylem mass surrounded by a phloem mass or vice versa. No cambium present. PLANT TISSUES 83 (a) Concentric, with xylem central in bundle. Seen in stems and leaves of nearly all ferns and the Lycopodiacece. ( b ) Concentric, with phloem central in bundle. Seen in stems and leaves of some Monocotyledons. Examples: Calamus and Conval- laria rhizomes. V. Radial, characterized by a number of xylem and phloem masses alternating with one another. Seen in the roots of all Spermalo- phytes and Pteridophytes. Xylem is that part of a fibro-vascular bundle that contains wood cells and fibers. It may also contain tracheae, tracheids, seldom sieve tubes. Phloem is that part of a fibro-vascular bundle that contains sieve tubes, phloem cells, and often bast libers. SECRETION SACS (SECRETION CELLS) These were formerly parenchyma cells which sooner or later lost their protoplasm and nucleus and became receptacles for oil, resin, oleoresin, mucilage or some other secretory substance. They are generally found in parenchyma regions of stems, roots, leaves, flower or fruit parts and frequently possess suberized walls. Good illustrations of these structures may be seen in Ginger and Calamus. INTERCELLULAR AIR SPACES Intercellular air spaces are cavities filled with air found between cells or groups of cells throughout the bodies of higher plants. Their function is to permit of the rapid movement of atmospheric gases through the entire plant body. They are formed either by the breaking down of the middle lamella of the cell walls, where several cells come together, and a later separation of the cells at these places ( Scliizogenous intercellular air spaces ), or by a breaking down and disappearance of cell walls common to groups of cells ( lysigenous intercellular air spaces). In terrestrial plants which live in middle regions ( mesophytes ) and in desert plants ( xeropkytes ) the intercellu- lar air spaces are averagely small and more or less angular. In plants of swamp or marsh habit they are medium-sized while in those which live entirely in the water ( hydrophytes ) they are of large size and more or less rounded. 8 4 PHARMACEUTICAL BOTANY SECRETION RESERVOIRS These structures are either found as globular or irregular spaces, as in Orange and Lemon Peel and Eucalyptus leaves, containing oil or oil and resin when they are called internal glands or, as tube-like spaces filled with hydrocarbon principles such as are found in Pine leaves and stems, when they sometimes receive the name of secretion canals. Occasionally they are named according to the nature of their contents — resin or oil canal or reservoir, etc. They are gen- erally lined with a layer of cells, usually more or less flattened, which are characterized by possessing large nuclei. To this layer has been assigned the name “epithelium.” Fig. 33. — Resin duct (secretion reservoir) in leaf of Pinus silvestris, in cross section at A , and in longitudinal section at B ; h, cavity surrounded by the secret- ing cells;/,/, sclerenchyma fibers surrounding and protecting the duct. ( Stevens , after Haberlandt.) Classification of Tissues According to Function.— According to their particular function, tissues may be classified as follows: I. Conducting Tissue II. Protective Tissues III. Mechanical Tissues ’ Parenchyme (fundamental tissue) Xylem cells (wood parenchyme) Tracheae (ducts) Phloem cells Sieve tubes . Companion cells I Epidermis (outer cell walls cutinized) l Cork (suberized tissue) Bast fibers Wood fibers Sclerenchyme fibers Stone cells CHAPTER VII PLANT ORGANS AND ORGANISMS An organ is a part of an organism made up of several tissues and capable of performing some special work. An organism is a living entity composed of different organs or parts with functions which are separate, but mutually dependent, and essential to the life of the individual. The organs of flowering plants are either Vegetative or Reproduc- tive. The vegetative organs of higher plants are roots, stems, and leaves. They are concerned in the absorption and elaboration of food materials either for tissue-building or storage. The reproductive organs of higher plants include those structures whose function it is to continue the species, viz.: th t flower , fruit and seed. The ripened seed is the product of reproductive processes, and the starting point in the life of all Spermatophytes. The living part of the seed is the embryo, which, when developed, consists of four parts, the caulicle, or rudimentary stem, the lower end of which is the be- ginning of the root, or radicle. At the upper extremity of the stem are one, two, or several thickened bodies, closely resembling leaves, known as cotyledons, and between these a small bud or plumule. The function of the cotyledon is to build up nourishment for the rudimentary plantlet until it develops true leaves of its own. THE ROOT The root is that part of the plant that grows into or toward the soil, that never develops leaves, rather rarley produces buds, and whose growing apex is covered by a cap. The functions of a root are absorption, storage and support. Its principal function is the absorption of nutriment and to this end it 85 86 PHARMACEUTICAL BOTANY generally has branches or rootlets covered with root-hairs which largely increase the absorbing surface. These root-hairs are of minute and simple structure, being merely elongations of the epidermis of the root back of the root cap into slender tubes with thin walls. Fig. 35- — Root-hairs, with soil-particles adhering. (Gager, after Sachs.) The tip of each rootlet is protected by a sheath- or scale-like cover- ing known as the root cap, which not only protects the delicate grow- ing point, but serves as a mechanical aid in pushing its way through the soil. The generative tissues in the region of the root cap are: plerome, producing fibro-vascular tissue; periblem, producing cortex; PLANT ORGANS AND ORGANISMS 87 dermatogen, producing epidermis; and calyptrogen, producing the root cap. Differences Between Root and Stem The Root The Stem I. Descending axis of plant. 1. Ascending axis of plant. 2. Growing point sub-apical. 2. Growing point apical. 3 - Contains no chlorophyll. 3. Chlorophyll sometimes present. 4 - Branches arranged irregularly. 4. Branches with mathematical regu- larity. 5 - Does not bear leaves or leaf rudi- 5. Bears leaves and modifications. ments. 6. Structure comparatively simple. 6. Structure better defined. Classification of Roots as to Form. — i. Primary or first root, a direct downward growth from the seed, which, if greatly in excess of the lateral roots, is called the main or tap root. Examples: Taraxa- cum, Radish. 2. Secondary roots are produced by the later growths of the stem, such as are covered with soil and supplied with moisture. Both primary and secondary roots may be either fibrous or fleshy. The grasses are good examples of plants having fibrous roots. Fleshy roots may be multiple, as those of the Dahlia, or may assume simple forms, as follows: Fusiform, or spindle-shaped, like that of the radish or parsnip. Napiform or turnip-shaped, somewhat globular and becoming abruptly slender then terminating in a conical tap root, as the roots of the turnip. Conical, having the largest diameter at the base then tapering, as in the Maple. 3. Anomalous roots are of irregular or unusual habits, subserving other purposes than the normal. 4. Adventitious roots are such as occur in abnormal places on the plant. Examples: Roots developing on Bryophyllum and Begonia leaves when placed in moist sand. 5. Epiphytic roots, the roots of epiphytes, common to tropical forests, for example, never reach the soil at all, but cling to the bark of trees and absorb nutriment from the air. Example: Roots of Vanilla. 88 PHARMACEUTICAL BOTANY 6. The roots of parasitic plants are known as Ilaustoria. These penetrate the bark of plants upon which they find lodgment, known as hosts, and absorb nutritious juices from them. The Mistletoe, Dodder and Geradia are typical parasites. Duration of Root. — Plants are classified according to the duration of the root, as follows: 1. Annual plants are herbs with roots containing no nourishment for future use. They complete their growth, producing flower, fruit and seed in a single season, then die. 2. Biennial plants develop but one set of organs the first year, and as in the beet and turnip, etc., a large amount of reserve food material is stored in the root for the support of the plant the following season when it flowers, fruits, and dies. 3. Perennial plants live indefinitely, as trees. Root Histology. - Monocotyledons. — The histology of mono- cotyledonous roots varies, depending upon relations to their sur- roundings, which may be aquatic, semi-aquatic, mesophytic, or xerophytic. In this connection we will discuss only the type of greatest pharmacognic importance, i.e., the mesophytic type as seen in its most typical form in the transverse section of Honduras Sarsaparilla root. Examining such a section from periphery toward the center, one notes the following: 1. Epidermis of a single layer of cells many of which give rise to root-hairs. 2. Hypodermis of two or three layers of cells whose walls are extremely thickened. 3. Cortex, consisting of a broad zone of parenchyme cells many of which contain starch grains. 4. Endodermis of one layer of endodermal cells whose walls are extremely thickened through the infiltration of suberin and lignin. 5. Pericambium of one or two layers of meristematic cells whose walls are extremely thin. 6. A radial fibro-vascnlar bundle of many alternating xylem and phloem patches and hence polyarch. The phloem tissue consists of phloem cells and sieve tubes. The xylem is composed of xylem cells, tracheae and wood fibers. Fig. 36. — Part of a transverse section of Honduras sarsaparilla root showing epidermis (e), root hair (ha), hypodermis (h), cortex (c), endodermis ( en ), peri- cambium (p), trachea of one of the numerous xylem patches (<), and pith (m). The phloem patches are the small oval cellular areas wedges in between the outer portions of adjacent xylem masses. (Photomicrograph.) xylem alternating with four phloem patches. These roots have an unlimited power of growth. A. Of Primary Growth. A trans-section of a dicotyl root in its young growth shows the following structure from periphery toward center: PLANT ORGANS AND ORGANISMS 89 7. Medulla or pith composed of parenchyme cells containing starch and often showing xylem patches cut off and enclosed within it. Dicotyledons. — The typical dicotyl root is a tetrarch one, four 9 ° PHARMACEUTICAL BOTANY 1. Epidermis with cutinized outer walls, the cells often elongating to form root-hairs. 2. Hypodermis. 3. Primary cortex with usually small intercellular spaces. 4. Endodermis, or innermost layer of cells of the cortex with lenticularly thickened radial walls. 5. Pericambium of one to two layers of actively growing cells which may produce side rootlets. 6. Radial fibro-vascular bundle of four, rarely two or three or five or six phloem patches alternating with as many xylem arms. Not uncommon to find bast or phloem fiber along outer face of each phloem patch. Xylem has spiral tracheae, internal to these a few pitted vessels, then, as root ages, more pitted vessels, also xylem cells and wood fibers make their appearance. B. Of Secondary Growth (Most official roots). At about six weeks one notes cells dividing by tangential walls in the inner curve of phloem patches. This is intrafascicular cambium. A single layer of flattened cells starts to cut off on its inner side a quantity of secondary xylem and pushes out the patches of bast fibers, adds a little secondary phloem on the outer side. Secondary xylem finally fills up the patches between the arms. The patches of bast fibers get flattened out. The pericambium has a tendency to start division into an inner and outer layer. The outer layer be- comes a cork cambium (phellogen) surrounding the bundle inside of the endodermis. It cuts off cork tissue on its outer face, hence all liquid material is prevented from filtering through and cortex includ- ing endodermis, as well as the epidermis, shrivel and dry up and separate off at the age of two to three months. The cork cambium (phellogen) may lay down secondary cortex internal to itself and external to the phloem. Patches of cells of the inner layer of pericambium divide rapidly and are called interfascicular cambium. These join the intrafasci- cular cambium to form a continuous cambium ring which then cuts off additional secondary xylem on its inner face and sec- ondary phloem on its outer face pushing inward the first-formed or protoxylem and outward the first-formed or protophloem. The medullary rays become deepened. PLANT ORGANS AND ORGANISMS 91 Thus, in a transverse section made through a portion of a Dicotyl root showing secondary growth , the following regions are noted pass- ing from periphery to center: 1. Cork 2. Cork cambium (phellogen) 3. Secondary cortex r 4. Protophloem 5. Secondary phloem 6. Cambium 7. Secondary xylem 8. Protoxylem w <& > t/i jQ E Strands of cells extending radially from the cortex to the center of the section separating each open fibro- vascular bundle from its neighbors. These are called medullary rays. Histology and Development of a Dicotyl Root (California Privet). — A. Make a permanent mount of a T. S. of the root of the California Privet ( Ligustrum Calif or nicum) cut just above the root cap, and note the fol- lowing structures, passing from pe- riphery toward the center: 1. Epidermis, composed of a layer of epidermal cells whose outer walls have been infiltrated with a substance called Cutin. Fig. 37. — Cross-section of a young root of Phaseolus mulli- Jlorus. A, pr, cortex; m, pith; x, stele or central cylinder — all tissue within the pericycle, in- clusive; g, primary xylem bun- dles; b, primary phloem bun- dles. B, cross-section of older portion of root; lettered as in A ; b', secondary phloem, k, cork. ( Stevens , after Vines.) 2. Hypodermis, a layer of somewhat thick walled cells just be- neath the epidermis. 3. Cortex, contposed of cortical parenchyme cells with small angular intercellular air spaces. 4. Endodermis, or innermost layer of cells of the cortex, whose radial walls are lenticularly thickened. 5. Pericambium, of a layer of actively growing meristematic cells, which has the power of producing lateral rootlets. 92 PHARMACEUTICAL BOTANY 6. Radial fibro-vascular bundle of five xylem arms alternating with as many phloem patches. Note the narrow spiral tracheae in the xylem patches. The section you have just studied illustrated in general the appear- ance of any Dicotyl root of primary growth. B. Mount permanently another T. S. cut through the same root a short distance above the first. Note that this is somewhat larger in diameter. Observe the root hairs starting from the epidermis; a broad cortex; a large clear and open looking endodermis; then pericambium; next, a central patch of xylem showing a faint pentarch relation. Pushed out are five phloem tracts. Each of these constitutes a mass of protophloem (first formed phloem). On the inner face of each phloem mass may be seen intrafascicular cambium. At the outer end of each xylem tract there has been cut off a patch of fine cambial cells (interfascic- ular cambium) which becomes joined to the intrafascicular cambium to develop secondary phloem on the outer face and secondary xylem on the inner face. C. Mount permanently a third T. S. out through the same root a short distance above the second. Note that this is still larger in diameter than the second. The pericambium has already divided into an inner and an outer layer. The outer layer has become the cork cambium, cutting off cork on its outer face beneath the endo- dermis. Cork being an impermeable barrier to water has prevented the nourishing sap from percolating through to the endodermis, cor- tex and epidermis. The epidermis has consequently begun to sluff off. Note that the cambium has begun to spread out into the form of a ring. More secondary xylem has been formed on its inner face and additional secondary phloem has appeared on its outer face. D. Make a permanent mount of a fourth T. S. cut through the same root some distance above the third. Note that the epidermis, primary cortex and endodermis have completely peeled off. Cork is found as the external bounding layer and underneath it, cork cambium. This cork cambium has developed secondary cortex on its inner face. The cambium has assumed a circular aspect. Just beneath the secondary cortex will be found flattened patches of protopliloem, and beneath these secondary phloem masses have PLANT ORGANS AND ORGANISMS 93 been formed through the activity of the cambium: The cambium has developed new or secondary xylem on its inner face which has pushed the first formed or protoxylem toward the center of the root. Abnormal Structure of Dicotyl Roots. — In certain Dicotyl roots as Amaranthus, Jalap, Pareira, and Phytolacca after the normal bundle system has been formed, there then develop successive cam- biums outside of these bundles, producing concentric series of open collateral bundles. Histology of a Dicotyl Tuberous Root (Aconitum).- — A transverse section made through the tuberous root of Aconitum Napellus near its middle shows a cork region of one or more layers of blackish or brownish cells; a broad cortex of two regions, viz.: an outer narrower and an inner broader zone. The narrower zone consists of from eight to fifteen layers of cortical parenchyme cells, interspersed among which are numerous irregular-shaped stone cells. Separating this zone from the broader one is an endodermis of a single layer of tan- gentially elongated endodermal cells. The broader zone consists of about twenty layers of parenchyme cells. Next, a five- to seven- angled cambium, within the angles of which and frequently scattered along the entire cambial line, occur collateral fibro-vascular bundles. In the center is found a broad five- to seven-rayed pith composed of parenchyme cells. The parenchyme cells of the cortical regions and pith contain single or two- to five-compound starch grains. Root Tubercles The roots of plants of the Leguminosoe, Myricacece as well as some species of Aristolochiacece and of the genera Alnus and Ceanothus are characterized by the appearance upon them of nodule-like swellings called root tubercles. In the case of the Leguminosae the causative factor is a species of bacteria named Pseudomonas radicicola. This is a motile rod-shaped organism which appears widely distributed in soils. It is apparently attracted to the root-hairs of leguminous plants by a chemotactic influence probably due to the secretions poured out by these structures. A number of these organisms pene- trate the walls of the root-hairs by enzymic action. Upon entering the hairs they form bacterial tubes which branch and rebranch and 94 PHARMACEUTICAL BOTANY extend into the middle cortex cells carrying the bacteria with them. Within the cortex cells the organisms multiply rapidly producing nest-like aggregations. Their presence here causes the formation of nodules or tubercles. Under oil-immersion magnification these bacteria are found to exhibit variously shaped involution forms called bacterioids. They remain within the cells of the medio-cortex region Fig. 38. — Root system of a legume showing tubercles. (Marshall.) gradually swelling up into zoogloea masses until finally their bodies break down into soluble nitrogenous substances which are partly absorbed and assimilated and partly stored as reserve nitrogenous food for the green leguminous plant. In the modern rotation of crops, plant growers plough under the leguminous crops or their nodule-producing roots which decay and enrich the soil with ample nitrogenous material to supply the next season’s crop of nitrogen-consuming plants. PLANT ORGANS AND ORGANISMS 95 The writer has found tubercles on Myrica cerifera, Myrica Car- oliniensis and Myrica Macfarlanei seedling primary roots of 5 to 6 months’ growth, and from thence onward on the secondary roots inserted on the hypocotyl axis, on nearly all the adventitious roots of subterranean branches and on the subterranean branches of Myrica cerifera, M. Caroliniensis, M. Gale, M. Macfarlanei, and Comptonia asplenifolia. The inciting organism has been isolated by him in pure culture according to Koch’s postulates and named Actinomyces Myricarum Youngken. Fig. 39. — Ps. radicicola. i, From Melilotus alba; 2 and 3, from Medicago saliva ; 4, from Vicia villosa. ( Marshall , after Harrison and Barlow from Lipman.) The tubercles occur either singly, as is frequently the case on subterranean branches, in small groups the size of a pea, or in larger coralloid loose or compact clusters which frequently attain the size of a black walnut. Each tubercle is a short cylindrical blunt-ended root-like structure which branches di- or trichotomously after attain- ing a certain length. The branches frequently rebranch at their tips which grow out into long thread-like structures from 1-3 cm. in length that may also branch and become entwined about the roots of other plants. The color of the youngest tubercles is a pinkish-gray brown. As the tubercles become older their color changes to brown, dark-brown and even black. (For a detailed description of the Myrica and Comptonia tubercles and their inciting organism, con- 9 6 PHARMACEUTICAL BOTANY suit, “The Comparative Morphology, Taxonomy and Distribution of the Myricaceas of the Eastern United States” by Youngken in U. of Pa. Library.) Fig. 40. — Tubercular clusters on underground stem and roots of Myrica Mac- farlanei observed by the author at North Wildwood, N. J., Jan. 31, 1915. THE BUD Buds are short young shoots with or without rudimentary leaves (bud scales) compactly arranged upon them. The plumule represents the first bud on the initial stem or caulicle. Scaly buds are such as have their outer leaf rudiments transformed into scales; there are often coated with a waxy or resinous substance without and a downy lining within, to protect them from sudden changes in climate. Buds of this character are common among shrubs and trees of temperate regions. Naked buds are those which are devoid of protective scales. They are common to herbaceous plants. PLANT ORGANS AND ORGANISMS 97 Classification of Buds According to Development. — i. A leaf bud is a young shortened shoot bearing a number of small leaves. It is capable of elongating into a branch which bears leaves. 2. A flower bad is a rudimentary shoot bearing one or more concealed and unexpanded young flowers. 5. A mixed bud is a young shoot bearing concealed unexpanded leaves and flowers. Classification of Buds According to Position on the Stem. — 1. A terminal bud is one which is located on the end of a stem (shoot). It is capable of elongating into a shoot which bears leaves or both leaves and flowers. 2. An axillary or lateral bud is one which arises in the leaf axil. It is capable of giving rise to a side branch or to a flower. Occa- sionally axillary buds do not develop and are then called dormant buds. 3. An adventitious bud is one which occurs on some position of the stem other than at its apex or in the axil of a leaf. Such buds may be seen developing along the veins of a Begonia leaf or along the margin of a Bryophyllum leaf after these have been planted in moist soil for several days. 4. An accessory bud is an extra bud which forms in or near the leaf axil. Classification of Buds According to Their Arrangement on the Stem. 1. When a single bud is found at each joint or node of a stem, the buds are said to be alternate. 2. When two buds are found at a node they are opposite. 3. When several buds occur at a node they are whorled. THE STEM The stem is that part of the plant axis which bears leaves or modi- fications of leaves and its branches are usually arranged with mathe- matical regularity. Stems usually grow toward the light and so are heliotropic. The functions of a stem are to bear leaves or branches, connect roots with leaves, and conduct sap. When the stem rises above ground and is apparent, the plant is said to be caulescent. 7 PHARMACEUTICAL BOTANY 98 . When no stem is visible, but only flower or leaf stalks, the plant is said to be acaulescent. Stems vary in size from scarcely I25 inch in length, as in certain mosses, to a remarkable height of 400 feet or more. The giant Sequoia of California attains the height of 420 feet. Some of the Eucalyptus trees of Australia and Tasmania are reported to attain the height of 500 feet. Nodes and Intemodes. — The nodes are the joints of stems. They represent the parts of the stem from which leaves or branches arise. Internodes are the parts of stems between nodes. Direction of Stem Growth. — Generally the growth of the stem is erect. Very frequently it may be: Ascending, or rising obliquely upward. Example: Saw Palmetto. Reclining, or at first erect but afterward bending over and trailing upon the ground. Example: Raspberry. Procumbent, lying wholly upon the ground. Example : Pipsissewa. Decumbent, when the stem trails and the apex curves upward. Example: Vines of the Cucurbitacese. Repent, creeping upon the ground and rooting at the nodes, as the Strawberry. Stem Elongation. — At the tip of the stem there is found a group of very actively dividing cells (meristem) which is the growing point of the stem. All the tissues of the stem are derived from the cells of the growing point whose activity gives rise in time to three generative regions which are from without, inward: 1. Dermatogen, forming epidermis; 2. Periblem, forming the cortex; and 3. Plerome, forming the fibro-vascular elements and pith. Duration of Stems. Annual, the stem of an herb whose life terminates with the season. Example: Corn. Biennial, where the stem dies at the end of the second year. Example: Burdock. Perennial, when the stem lives for many years. Example: Oak. ^ Stem Modifications. — (1) twining, by elongation and marked circumnutation of young internodes as in Convolvulus, Dodder, PLANT ORGANS AND ORGANISMS 99 etc. (2) Tendriliform by thread-like modification and sensitivity to contact of a side branch as in Passion flower, Squash, etc. (3) Spiny, by checking and hardening of a branch that may then become defensive ecologically as in hawthorn, honey locust, etc. (4) Aerial tuberous, in which one or more internodes enlarge above ground and store reserve food as in pseudobulbs of orchids, Vitis gongylodes, etc. (5) Subterranean tuberous in which a subter- ranean stem or branch enlarges as a food-storing center: (a) annual type, tuber as in potato, etc., corm as in crocus, etc.; (b) perennial type, bulbs as in lily (scaly) and onion or hyacinth (tunicated). (6) Phylloid or leaf-like in w r hich flattening branch expansion occurs, when leaves become reduced in size as in Asparagus, Ruscus, etc. (7) Cactoid, in which reduced condensed branches or stems become swollen for water (and food) storage as in Cacti, Euphorbia sp., etc. Above-ground Stems. — A twining stem winds around a support, as the stem of a bean or Morning Glory. A culm is a jointed stem of the Grasses and Sedges. A climbing or scandent stem grows upward by attaching itself to some support by means of aerial rootlets, tendrils or petioles. Examples: Ivy, Grape, etc. The scape is a stem rising from the ground and bearing flowers but no leaves, as the dandelion, violet, or blood root. A tendril is a modification of some special organ, as of a leaf stipule or branch, capable of coiling spirally and used by a plant in climbing. Present in the Grape, Pea, etc. A spine or thorn is the indurated termination of a stem tapering to a point, as the thorns of the Honey Locust. Prickles are outgrowths of the epidermis and cortex and are seen in the roses. A stolon is a prostrate branch, the end of which, on coming in con- tact with the soil, takes root, so giving rise to a new plant. Examples: Currant and Raspberry. An herbaceous stem is one which is soft in texture and readily broken. Example: Convallaria. An undershrub or sujf'ruitcose stem is a stem of small size and woody only at the base. Examples: Bitter-sweet, Thyme, etc. IOO PHARMACEUTICAL BOTANY A shrubby or fruitcose stem is a woody stem larger than the pre- ceding and freely branching near the ground. Example: Lilac, etc. A trunk is the woody main stem of a tree. Herb and Tree A tree is a perennial woody plant of considerable size (20 feet or more in height) and having as the above-ground parts a trunk and a crown of leafy branches. An herb is a plant whose stem does not become woody and perma- nent, but dies, at least down to the ground, after flowering. Underground Stems. — A rhizome is a creeping underground stem, more or less scaly, sending off roots from its lower surface and stems from its upper. The rhizome grows horizontally, vertically or ob- liquely, bearing a terminal bud at its tip. Its upper surface is marked with the scars of the bases of aerial stems of previous years. Examples: Triticum, Rhubarb, etc. The tuber is a short and excessively thickened underground stem, borne usually at the end of a slender, creeping branch, and having numerous eyes or buds. Example: Tubers of the Potato. The corm is an underground stem excessively thickened and solid and characterized by the production of buds from the center of the upper surface and rootlets from the lower surface. Examples: Colchicum, Jack-in-the-Pulpit, etc. A bulb is a very short and scaly stem, producing roots from the lower face and leaves and flower from the upper. Tunicated bulbs are completely covered by broad scales which form concentric coatings. Examples: Onion, Squill, Daffodil. Scaly bulbs have narrow imbricated scales, the outer ones not en- closing the inner. Example: Lily. Tubers and corms are annual. Bulbs and Rhizomes are perennial. Exogenous and Endogenous Stems.- — Exogenous stems are typical of Gymnosperms and Dicotyledons and can increase materially in thickness due to presence of a cambium. Such stems show differen- tiation into an outer or cortical region and an inner or central cylinder region. Endogenous stems are typical of most Monocotyledons and cannot PLANT ORGANS AND ORGANISMS IOI increase materially in thickness due to absence of cambium. The limited increase in diameter that does take place is due to the en- largement of the cells of the primary tissues. Such stems show no differentiation into cortical and central regions. Histology of Annual Dicotyl Stem. — (In both annual and perennial dicotyledonous stems endodermis and pericambium are rarely seen Fig. 41. — Photomicrograph of cross-section of stem of Aristolochia sipho, where cambial activity is just beginning, a, Epidermis; b, collenchyma; c, thin- walled parenchyma of the cortex, the innermost cell layer of which is the starch sheath or endodermis; d, sclerenchyma ring of the pericycle; e, thin-walled paren- chyma of the pericycle; /, primary medullary ray; g, phloem; h,\x ylem; i, inter- fascicular cambium; j, medulla or pith. X 20. ( From Stevens.) since each has become so similar to cortex through passage of food, etc.) 1. Epidermis, cutinized, with hairs. 2. Cortex composed of three zones: an outer or exocortex, whose cells are thin walled and contain chloroplasts; a middle or medio- cortex, consisting of cells of indurated walls giving extreme pliability and strength, an inner or endocortex, a very broad zone of thin- and thicked-walled parenchyme cells. 3. The innermost layer of cells of the cortex called endodermis. (Not generally distinguishable.) 102 PHARMACEUTICAL BOTANY PLANT ORGANS AND ORGANISMS 103 4. Pericambium. (Not generally distinguishable.) 5. Fibro-vascular bundles of open collateral type arranged in a circle with primary medullary rays between the bundles. 6. Pith. 104 PHARMACEUTICAL BOTANY Growth of Perennial Dicotyl Stem and its Histology. — A perennial dicotyl stem in the first year does not differ is structure from an annual. By the close of the year a cork cambium (phellogen) has originated beside the epidermis. In origin of cork cambium — one of two methods: ( a ) either the epidermis may divide into an outer layer of cells that remains epidermis and an inner layer of cells that becomes cork cambium, or, ( b ) the outermost layer of cortex cells underneath the epidermis becomes active after being passive for one year, and lays down walls, the inner layer becoming cork cam- bium, the outer becoming a layer of cork. The cork cuts off water and food supplies from epidermis outside and so epidermis separates and falls off as a stringy layer. The cork cambium produces cork on its outer face and secondary cortex on its inner. Between the bundles certain cells of the primary medullary rays become very active and form interfascicular cambium which joins the cambium of the first-formed bundles (intrafascicular cambium) to form a complete cambium ring. By the rapid multiplication of these cambial cells new (secondary) xylem is cut off internally and new (secondary) phloem externally, pushing inward the first-formed, or protoxylem, and outward the first-formed, or protophloem, thus increasing the diameter of the stem. The primary medullary rays are deepened. Cambium may also give rise to secondary medullary rays. Sometimes, as in Grape Vines, Honeysuckles, and Asclepias, in- stead of cork cambium arising from outer cortex cells it may arise at any point in cortex. It is the origin of cork cambium at varying depths that causes extensive sheets of tissue to separate off. That is what gives the stringy appearance to the stems of climbers. At close of first year in Perennial Dicolyl Stem we note : Epidermis — development of dermatogen or periblem — in process of peeling oS, later on entirely absent. Cork tissue or periderm. Cork cambium or phellogen. Sometimes zone of thin-walled cells containing chloroplasts cut off by cork cambium on inner face and known as phelloderm. Cortex — in perennial stem cells of cortex may undergo modification into mucilage cells, into tannin receptacles, crystal cells, spiral cells, etc. PLANT ORGANS AND ORGANISMS 6. Fibro- vascular bundles of open collateral type which are now arranged into a compact circle, and between which are found primary and often secondary medullary rays. From without inward the following tissues make up f. v. bundles. Protophloem j Hard Bast — long tenacious bast fibers. Secondary Phloem [ Soft Bast — phloem cells and sieve tubes. Cambium — active layer giving rise to secondary phloem on outer and secondary xylem or inner face, and adding to depth of med. rays. Secondary xylem — wood fibers, pitted vessels, tracheids. Protoxylem — spiral tracheae. Pith. Fig. 44. — Portion of cross-section of four-year-old stem of Arislolochia sipho, as shown by the rings of growth in the wood. The lettters are the same as in Fig. 41 but new tissues have been added by the activity of the cambium; and a cork cam- bium has arisen from the outermost collenchyma cells and given rise to cork. The new tissues are; l, cork cambium; k, cork; g, secondary phloem from the cambium, and just outside this is older crushed phloem; n, secondary xylem produced by the cambium; m, secondary medullary ray made by the cambium (notice that this does not extend to the pith). Half of the pith is shown. Notice how it has been crushed almost out of existence. Compare Figs. 41 and 44, tissue for tissue, to find out what changes the primary tissues undergo with age, and to what extent new tissues are added. Photomicrograph X 20. ( From Stevens.) io6 PHARMACEUTICAL BOTANY EXCEPTIONAL TYPES OF DICOTYL STEMS In a number of Dicotyledons and Gymnosperms, the secondary growth in thickness of the stem and frequently of the root differs from that which is found in the vast majority of species and so is called exceptional or anomalous. In Phytolacca, etc., there first arises a ring of primary bundles with broad loose medullary rays. Then the stem cambium ceases its Fig. 45. — White birch ( Betula populifolia) . Portion of a branch showing the prominent lenticels. (Gager.) activity and, outside the bast of the bundles already formed in the pericambium or tissue developing from it, a new cambium starts to lay down another ring of bundles in rather irregular fashion. Then after developing a wavy ring of bundles and connecting tissue that cambium closes up. Still another cambium ring arises without this, and in a single season quite a number of these are found succes- sively arranged in concentric fashion. In Gelsemium , species of Solanacece, Cambrel ace a, Cucurbitacece, etc., there arises a cambium on the inner face of the xylem which PLANT ORGANS AND ORGANISMS 107 forms internal phloem (or intraxylary phloem ), thus giving rise to bicollateral bundles'. In Strychnos Nux Vomica internal phloem exactly as in Gelse- mium, etc., appears but in addition interxylary phloem is developed. In the wood region of this plant axis the cambium starts at a certain age to lay down patches of phloem which become wedged in between xylem tissue as interxylary phloem. Lenticels and Their Formation. — The epidermis in a great majority of cases produces stomata, apertures, surrounded by a pair of guard cells, which function as passages for gases and watery vapor from and to the active cells of the cortex beneath. Fig. 46. — Cross-section through a lenticel of Sambucus nigra. E, Epidermis; PH, phellogen; L, loosely disposed cells of the lenticel; PL, cambium of the lenticel; PS, phelloderm; C, cortical parenchyma containing chlorophyll. (From Sayre after Strasburger.) There very early originate in the region beneath the stomata loosely arranged cells from cork cambium which swell up during rain and rupture, forming convex fissures in the cork layer, called lenticels. The function of lenticels is similar to that of stomata, namely, to permit of aeration of delicate cells of the cortex beneath. Annual Thickening. — In all woody exogenous stems such as trees and shrubs the persistent cambium gives rise to secondary xylem thickening every spring, summer and autumn. Soon a great cylinder of xylem arises which constitutes the wood of the trunk and branches. In the spring, growth is more active, and large ducts with little woody fiber are produced while in summer and autumn io8 PHARMACEUTICAL BOTANY growth is lessened and small ducts and much mechanical woody fiber are formed. Thus the open, loosely arranged product of the spring growth abuts on the densely arranged product of the last summer and autumn growth and the sharp contrast marks the periods of growth. To the spring, summer and autumn regions of growth of each year is given the term of “annual ring.” By count- ing the number of these rings it is possible to estimate the age of the tree or branch. Fig. 47. — Part of a transverse section of a twig of the lime, four years old. m, Pith; ms, medullary sheath; x, secondary wood; Ph, phloem; 2, 3, 4, annual rings; c, cambium; pa, dilated outer ends of medullary rays; b, bast; pr, primary cortex; k, cork. ( From Sayre after Vines.) Bark. — Bark or bork is a term applied to all that portion of a woody exogenous plant axis outside of the cambium line. In pharmacognic work, bark is divided into three zones, these from without inward being: , 1. Outer Bark or Cork. 2. Middle Bark or Cortical Parenchyme. 3. Inner Bark or Phloem. Periderm. — Periderm is a name applied to all the tissue produced externally by the cork cambium ( P hello gen ). This term appears often in pharmacognic and materia medica texts. Phelloderm. — Phelloderm or secondary cortex is all that tissue produced by the cork cambium on its inner face. PLANT ORGANS AND ORGANISMS IO9 U Fig. 48. — Part of a cross-section through branch of Cytisus laburnum. (The branch was cut from the tree at the end of October.) Frbm A to £ the last annual ring of wood; from A to £ the spring growth with large tracheal tubes ( T , T, T)\ between B and C and D and D are wood-fibers; between C and D and D and E, wood parenchyma; from E to F, cambium; F to G. phloem portion; G to H, cortical parenchyma; M, medullary ray. Below A the last wood-fibers and wood parenchyma formed the previous year. ( From Sayre after Haberlandt.) I IO PHARMACEUTICAL BOTANY Histology of a Typical Bark, Cascara Sagrada. — In transverse section passing from outer to inner surface, the following structural characteristics are evident: 1 . Cork, or outer bark, composed of several layers of rectangular cork cells. The most external layers are dead and appear black because they are filled with air. The inner layers of this region are living and contain brownish contents. 2. Cork cambium (phellogen), a layer of delicate cells with pro- toplasmic contents in the process of division. 3. Cortex, or middle bark, consisting of two regions, viz.: an outer zone of two or three rows of brownish collenchyme cells, and an inner broader zone of tangentially elongated cortical parenchyme cells. Imbedded within this zone will be noted numerous groups of stone cells. 4. Phloem, or inner bark, a very broad zone composed of irregular- shaped elongated phloem masses separated from each other by me- dullary rays which converge in the outer phloem region. Each phloem mass consists of numerous sieve tubes and phloem cells, some of which latter contain spheroidal starch grains while others contain monoclinic prisms or rosette aggregates of calcium oxalate. Em- bedded within the phloem masses in tier-like fashion will be noted groups of bast fibers, each group of which is surrounded by a row of crystal fibers, individual cells of which can only be made out in this kind of a section. Each of these contains a monoclinic prism of calcium oxalate. The medullary rays possess brownish contents which take a red color with an alkaline solution. In radial longitudinal section a lengthwise view of the tissues will be seen. The medullary rays appear 15 to 25 cells in height and crossing at right angles to the other elements. The crystal fibers here will be seen to be composed of vertical rows of superimposed thin-walled cells each of which contains a monoclinic prism of calcium oxalate. The bast fibers appear elongated and taper ended and are associated with crystal fibers. In a tangential longitudinal section which has been cut through the phloem, the exact range in width of the medullary rays may be ascertained. In this bark the medullary rays are spindle-shaped in tangential view and one to four cells in width. PLANT ORGANS AND ORGANISMS III Wood.- — From a pharmacognic standpoint as well as that of the lumber trade, wood is all that portion of woody exogenous plant axes inside of the cambium line. In Dicotyl and Gymnosperm stems it therefore includes the xylem regions of the bundles, the st Fig. 49. — Photomicrograph of transverse section of Cascara Sagrada bark; k, cork; g, cork cambium; c, cortex; st, group of stone cells; bf, group of bast fibers; mr, medullary ray. xylem portions of the medullary rays and the pith, while in the roots of secondary growth of these plants it comprises the xylem portions of the bundles and the xylem medullary rays. As the cambium year after year adds new layers of wood to that 1X2 PHARMACEUTICAL BOTANY already present on its inner face, the conveying of sap and storing of starch, etc. is gradually relegated to the outer wood layers, since Pig. 50. — Diagrammatic representation of a block of pine wood highly mag- nified. a, Early growth; b, late growth; c, intercellular space; d, bordered pit in tangential wall of late growth; m, f and e, bordered pit in radial wall of early growth from different points of view; h, row of medullary cells for carrying food; g, row of medullary ray cells for carrying water; k, thin place in radial wall of ray cells that carry food. (From Stevens.) the inner layers, step by step, lose their protoplasmic contents and power of conducting sap and become filled with extractive, resinous PLANT ORGANS AND ORGANISMS IJ 3 and coloring matters. The outer whitish layers of wood which con- tain living cells, functioning in the vegetative processes of the plant, constitute the alburnum or sap-wood. The drug Quassia is a good example of this kind of wood. The inner dead colored layers con- stitute the duramen or heart-wood. Important examples of this kind of wood used in pharmacy are Lignum Guaiaci, Hasmatoxylum, and Santalum Album. Microscopic Characteristics of Angiospermous and Gymnosperm- ous Woods.— The wood of Angiosperms is characterized by the pres- ence of tracheae (vessels) with various markings on their walls, particularly by small pits in the walls of some of the tracheae, together with wood fibers, wood parenchyma and medullary rays. The wood of Gymnosperms is made up for the larger part of tracheids with bordered pits which latter are charac- terized in radial longitudinal section by the presence of two rings, one within the other. A single row of these is seen on the tracheid wall. Medullary rays, frequently diagnostic for different species and woody parenchyme cells, are also found. Histology of Typical Herbaceous Monocotyl Stems (Endogenous). — Passing from exterior toward center the following structures are seen: Fig. 51. — Photomicrograph of cross-section of very young cornstalk, where certain plerome strands have just gone over into vascular bun- dles. For comparison with Fig. 52. (Stevens.) 1. Epidermis whose cells are cutinized in their outer walls. 2. Hypodermis, generally collenchymatic. 3. Cortex. 4. Endodermis or innermost layer of cortex. 5. A large central zone of parenchyme matrix in which are found scattered fibro-vascular bundles of the closed collateral or rarely concentric type (amphivasal). In this latter type, which is typical of old monocotyl stems, the xylem grows completely around phloem so that phloem is found in the center and xylem without and sur- rounding it. 8 PHARMACEUTICAL BOTANY 114 Histology of a Typical Woody Monocotyl Stem. — The stem of the Greenbrier, a woody monocotyl, will here be considered. In trans- verse section passing from periphery toward the center the following structural details will be noted: 1. Epidermis, of a single layer of epidermal cells whose outer walls are strongly cutinized. Cutin is a wax-like substance which forms a protective coat to the epidermis, preventing the evaporation of Fig. 52. — Cross-section of cornstalk stem; a, epidermis; b, cortex and c, ground tissue. ( After Stevens.) water, the ingress of destructive parasites, and injury from insects. 2. A cortex, composed of about ten or twelve layers of thick- walled parenchyme cells, the outer two or three layers of which are termed hypodermis. 3. An endodermis , wavy in character and composed of endodermal cells whose brownish walls are strongly suberized. 4. A sclerenchymatous cylinder sheath composed of somewhat separated masses of sclerenchymatous fibers and undeveloped fibro- vascular bundles of the closed collateral type. 5. A central matrix of strongly thickened parenchyme cells in which are scattered, irregularly, numerous closed collateral bundles. Small starch grains will be found in the parenchyma cells. Examine a representative bundle, and note the two very large tracheae and PLANT ORGANS AND ORGANISMS 115 several smaller ones in the xylem portion of the bundle which faces toward the center of the section. In the outer or phloem portion of the bundle will be seen an area of soft small-celled sieve tubes and phloem parenchyme. The entire bundle is enclosed by a several Fig. 53. — Photomicrograph of a representative portion of Greenbrier stem showing epidermis ( e.p .), cortex (c), endodermis ( e.n.d .). cylinder sheath ( c.s .), sclerenchyme fibers of closed collateral bundle ( b ), fundamental parenchyma ( f.p .), trachea (/). X 22. layered ring of sclerenchyme fibers, which on the inner face are called wood fibers, on the outer, bast fibers. The wood fibers con- stitute the supporting elements of the xylem, while the bast fibers are the supporting elements of the phloem. ii6 PHARMACEUTICAL BOTANY THE LEAF The leaf is a usually flattened, rarely semi-centric, or centric-lateral expanse developed by the stem or by branches and in whose axil one or more branches arise. Leaves seldom develop buds over their surface or along their margin and in connection therewith roots. The capacity for bud development is restricted to three families, viz.: Crassulacese, Be- goniaceae and Gesneraceae. Leaf Functions. — The most essential function of plants is the con- version of inorganic into organic matter; this takes place ordinarily in the green parts, containing chlorophyll, and in these when exposed to sunlight. Foliage is an adaptation for increasing the extent of green surface. The functions of a leaf are photosynthesis, assimilation, respiration and transpiration. Photosynthesis is the process possessed by all green leaves or other green parts of plants of building up sugar, starch or other complex organic substances by means of chlorophyll and sunlight. This process takes place in nature, only during sunlight. CO2 is taken in and O given off. Assimilation is the process of converting food materials into proto- plasm. Respiration or breathing is the gaseous interchange whereby all living organisms take in oxygen and give off carbon dioxide. Transpiration is the giving off of watery vapor. Types of Leaves Developed in Angiosperms. — These may be tabulated as follows: 1. Cotyledons (the primitive or seed leaves). 2. Scale leaves. 3. Foliage leaves. 4. Bract leaves: (a) primary at base of inflorescence; ( b ) bracteo- lar leaves at base of individual flowers. 5. Sepals. 6. Petals. 7. Microsporophylls (stamens). 8. Megasporophylls (carpels). PLANT ORGANS AND ORGANISMS 1 1 7 Cotyledons. — Cotyledons are the first leaves to appear upon the ascending axis and are single in Monocotyledons, double in Dicoty- ledons. Occasionally, as in certain Maples, there may be three cotyledons shown. This is due to a splitting of one of the cotyledons. There exist no true cases of polycotyledony (development of many cotyledons) among Angiosperms as in Gymnosperms. In Mono- cotyledons the single cotyledon is a terminal structure and truly axial in relation to the hypocotyl and radicle. From a primitively Monocotyl-like ancestry Dicotyledons develop a second cotyledon on the Epicotyledonary node. Later, by a suppression of the second node the second cotyledon is brought to the level of the first. Scale Leaves. — Scale leaves are reduced foliage leaves. They are found on certain rhizomes, above ground stems, such as Dodder, etc., on bulbs, and forming the protective scales of scaly buds. Foliage Leaves. — These are the common green leaves so familiar to all. Bract leaves are modified leaves appearing on inflorescence axes. Sepals, petals, microsporophylls and megasporophylls are floral leaves and will be treated at length under the subject of the flower. Origin and Development of Leaves. — Leaves arise around the growing apex region of a stem or branch as lateral outgrowths, each consisting at first of a mass of cells called the primordial leaf. Through continued cell-division and differentiation of these cells in time the mature leaf is developed. The primordial leaf is formed by a portion of the dermatogen of the growing stem apex, which becomes epidermis, a portion of the periblem, producing mesophyll which grows into this, and a part of the plerome, which becomes vascular tissue within the mesophyll. In the sub-divisions of cells around the growing stem-apex, the primordial leaves (primordia) do not arise exactly at the same time. There is a tendency toward spiral arrangement. Phyllotaxy. — Phyllotaxy is the study of leaf arrangement upon the stem or branch, and this may be either alternate, opposite, whorled, or verticillate, or fascicled. It is a general law in the arrangement of leaves and of all other plant appendages that they are spirally disposed, or on a line which winds around the axis like the thread of a screw. The spiral line is formed by the union of ii8 PHARMACEUTICAL BOTANY two motions, the circular and the longitudinal, and its most common modification is the circle. In the alternate arrangement there is but one leaf produced at each node. Opposite, when a pair of leaves is developed at each node, on opposite sides of the stem. Example: Mints, Lilac. Decussate, when the leaves are arranged in pairs successively along the stem, at right angles to each other. Whorled or Verticillate, when three or more form a circle about the stem. Example: Canada Lily and Culver’s root. Fascicled or T ufted, when a cluster of leaves is borne from a single node, as in the Larch and Pine. The spiral arrangement is said to be two-ranked when the third leaf is over the first, as in all Grasses; three-ranked, when the fourth is over the first. Example: Sedges. The five-ranked arrangement is the most common, and in this the sixth leaf is directly over the first, two turns being made around the stem to reach it. Example: Cherry, Apple, Peach, Oak and Willow, etc. As the distance be- tween any two leaves is two-fifths of the circumference of the stem, the five-ranked arrangement is expressed by the fraction In the eight-ranked arrangement the ninth leaf stands over the first, and three turns are required to reach it, hence the fraction % ex- presses it. Of the series of fractions thus obtained, the numerator represents the number of turns to complete a cycle, or to reach the leaf which is directly over the first; the denominator, the number of perpendicular rows on the stem, or the number of leaves, counting along thd spiral, from any one to the one directly above it. Vernation.- — Prefoliation or Vernation relates to the way in which leaves are disposed in the bud. A study of the individual leaf enables us to distinguish the following forms. When the apex is bent inward toward the base, as in the leaf of the Tulip Tree, it is said to be inflexed or reclinate vernation; if doubled on the midrib so that convolute Fig. 54- — Three principal types of verna- tion. (Robbins.) PLANT ORGANS AND ORGANISMS 119 the two sides are brought together as in the oak, it is conduplicate; when rolled inward from one margin to the other, as in the Wild Cherry, it is convolute; when rolled from apex to base, as in Ferns, it is circinate; when folded or plaited, like a fan as in Ricinus, Maples, Aralias, etc., it is plicate; if rolled inward from each margin, as the leaf of the common Violet, involute; when rolled outward from each margin as Rumex, revolute. The inner surface is always that which will form the upper surface when expanded. Fig. 55. — Stereogram of leaf structure. Part of a veinlet is shown on the right. Intercellular spaces are shaded. ( From Stevens.) The Complete Leaf.- — The leaf when complete consists of three parts, lamina , petiole, and stipules. The lamina or blade is the expan- sion of the stem into a more or less delicate framework, made up of the branching vessels of the petiole. The petiole is the leaf stalk. The stipules are leaf-like appendages appearing at the base of the petiole. The leaf of the Tulip Poplar or Liriodendron affords a good exam- ple of a Complete Leaf. 120 PHARMACEUTICAL BOTANY Sometimes the lamina or blade is attached directly to the stem by its base and is then said to be sessile. If the petiole is present, petiolate. When leaf stipules are absent, the leaf is said to be exstipulate, when present, stipulate. The petiole is seldom cylindrical in form, but usually channelled on the upper side, flattened, or compressed. The stipules are always in pairs and closely resemble the leaf in structure. The blade of the leaf consists of the framework, made up of branch- ing vessels of the petiole, which are woody tubes pervading the soft tissue called mesophyll, or leaf parenchyme, and serve not only as supports but as veins to conduct nutritive fluids. Veins are absent in simple leaves such as many of the Mosses. Leaf Venation. — Furcate or Forked Venation is characteristic of many Ferns. Parallel Venation is typical of the Monocotyledons, as Palms, Lilies, Grasses, etc. Reticulate or Netted Veins characterize the Dicotyledons, as the Poplar or Oak. Pinni-veined or Feathered-veined leaves consist of a mid-vein with lateral veinlets extending from mid-vein to margin at frequent intervals and in a regular manner. Example: Calla. Palmately Veined leaves consist of a number of veins of nearly the same size, radiating from petiole to margin. Example: Maple leaf. Veins are said to be anastomosing when they subdivide and join each other, as the veins near the margin of Eucalyptus leaves. Leaf Insertion. — The point of attachment of the leaf to the stem is called the insertion. A leaf is: Radical, when inserted upon an underground stem. Cauline, when upon an aerial stem. Ramal, when attached directly to a branch. When the base of a sessile leaf is extended completely around the stem it is perfoliate, the stem appearing to pass through the blade. Example: Uvularia perfoliata or Mealy Bellwort. When a sessile leaf surrounds the stem more or less at the base, it is called clasping or amplexicaul. Example: Poppy (Papaver somniferum). PLANT ORGANS AND ORGANISMS 1 2 1 Fig. 56. — Leaf outlines: Linear (1); lanceolate (2); oblong (3); elliptical (4), ovate-lanceolate (5); oblanceolate (6); spatulate (7); obovate-lanceolate (8); orbicular (9); reniform (10); cuneate (11). 122 PHARMACEUTICAL BOTANY When the bases of two opposite leaves are so united as to form one piece, they are called connate-perfoliate, as Eupatorium or Boneset. Leaves are called equitant when they are all radical and succes- sively folded on each other, as the Iris. The Forms of Leaves. — Simple leaves are those having a single blade, either sessile or petiolate. Compound leaves are divided into two or more distinct subdivi- sions called leaflets, which may be either sessile or petiolate. Simple leaves and the separate blades of compound leaves are de- scribed as to general outline, apex, base, marginal indentations, sur- face and texture. (a) General Outline (form viewed as a whole without regard to indentations of margin). Dependent upon kind of venation. When the lower veins are longer and larger than the others, the leaf is Ovate, or Egg-shaped. Example: Parallel- veined leaves are usually linear, long and narrow of nearly equal breadth throughout (Linaria), or lanceolate, like the linear with the exception that the broadest part is a little below the center. Example: Long Buchu. Elliptical, somewhat longer than wide, with rounded ends and sides. Example: Leaf of Pear. Oblong, when longer than broad, margins parallel. Example: Matico. Inequilateral, margin longer on one side than the other, as the Hamamelis, Elm and Linden. Orbicular, circular in shape. Example: Nasturtium. Peltate, or shield-shaped, having the petiole inserted at the center of the lower surface of the lamina. Example: Podophyllum. Filiform, or thread-like, very long and narrow, as Asparagus leaves. Ovate, broadly elliptical. Example: Digitalis. Obovate, reversely ovate. Example: Short Buchu. Oblanceolate, reversely lanceolate. Example: Chimaphila. Cuneate, shaped like a wedge with the point backward. Spatulate, like a spatula, with narrow base and broad rounded apex. Example: Uva Ursi. Ensiform, when shaped like a sword. Example: Calamus. Acerose or acicular, tipped with a needle-like point, as Juniper. PLANT ORGANS AND ORGANISMS 123 Fig. 57. — Leaf bases (12-17); leaf apices (18-26); compound leaves (27-31). Cordate (12); auriculate (13); connate-perfoliate (14); sagittate (15); hastate (16); peltate (17). Acuminate (18); acute (19; obtuse (20); truncate (21); retuse (22); emarginate (23; cuspidate (24); mucronate (25); aristate (26). Imparipin- nate (27); paripinnate (28); bi-pinnate (29); decompound (30; palmately 5-foliate (31). 124 PHARMACEUTICAL BOTANY Deltoid, when the shape of the Greek letter A, as Chenopodium. ( b ) Apex of Leaf. — Acute, when the margins form an acute angle at the tip of the leaf. Example: Eriodictyon. Acuminate, when the point is longer and more tapering than the acute. Examples: Pellitory, Coffee. Obtuse, blunt or round. Example: Long Buchu. Truncate, abruptly obtuse, as if cut square off. (Tulip Poplar). Mucronate, terminating in a short, soft point. Example: Senna leaflets. Cuspidate, like the last, except that the point is long and rigid. Aristate, with the apex terminating in a bristle. Emarginate, notched. Example: Pilocarpus. Reluse, with a broad, shallow sinus at the apex. Example: Petal of Rosa gallica. Obcordate, inversely heart-shaped. Example: Oxalis. (c) Base oi Leaf. — Cordate, heart-shaped. Example: Lime. Reniform, kidney-shaped. Examples: Ground Ivy, Asarum. Hastate, or halbert-shaped, when the lobes point outward from the petiole. Example: Aristolochia Serpentaria. Auriculate, having ear-like appendages at the base. Example: Philodendron. Sagittate, arrow-shaped. Example: Bindweed. (d) Margin of Leaf. — Entire, when the margin is an even line. Example: Belladonna. Serrate, with sharp teeth which incline forward like the teeth of a hand-saw. Example: Peppermint. Dentate, or toothed, with outwardly projecting teeth. Chestnut. Crenate, or Scalloped, similar to the preceding forms, but with the teeth much rounded. Examples: Digitalis, Salvia. Repand, or Undulate, margin — a wavy line. Example: Hamamelis. Sinuate, when the margin is more distinctly sinuous than the last. (Stamonium.) Incised, cut by sharp, irregular incisions. Example: Hawthorn. Runcinate, the peculiar form of pinnately incised leaf observed in the Dandelion and some other Compositae in which the teeth are recurved. A Lobed leaf is one in which the indentations extend nearly to the PLANT ORGANS AND ORGANISMS 125 Fig. 58. — Leaf margins: Pinnately-lobed (32); pinnately-cleft (33) ; pinnately- parted (34); pinnately-divided (35); palmately tri-lobed (36); palmately tri-cleft (37): palmately 3-parted (38); palmately 3-divided (39); crenate (40); serrate (41); dentate (42); repand or undulate (43); sinuate-dentate (44). 126 PHARMACEUTICAL BOTANY mid-vein, or mid-rib, as it is usually called, the segments or sinuses, or both, being rounded. Example: Sassafras. Cleft is the same as lobed, except that the sinuses are deeper, and commonly acute. Example: Dandelion. A Parted leaf is one in which the incisions extend nearly to the mid-rib. Example: Geranium maculatum. In the Divided leaf the incisions extend to the mid-rib, but the segments are not stalked. Example: Watercress. If the venation is pinnate, the preceding forms may be described as pinnately incised, lobed, parted, or divided. If the venation is radiate, then the terms radiately or palmately lobed, incised, etc., are employed. The transition from Simple to Compound Leaves is a very gradual one, so that in many instances it is difficult to determine whether a given form is to be regarded as simple or compound. The number and arrangement of the parts of a compound leaf correspond with the mode of venation, and the same descriptive terms are applied to outline, margin, etc., as in simple leaves. Leaves are either pinnately or palmately compounded. The term pinnate is frequently given to the former while that of palmate is often assigned to the latter. They are said to be abruptly pinnate or paripinnate when the leaf is terminated by a pair of leaflets; odd pinnate or imparipinnate when it terminates with a single leaflet. When the leaflets are alternately large and small, the leaf is inter- ruptedly pinnate, as the Potato leaf. When the terminal leaflet is the largest, and the remaining ones diminish in size toward the base the form is known as lyrate, illustrated in the leaf of the Turnip. Palmately compound leaves have the leaflets attached to the apex of the petiole. When these are two in number the leaf is bifoliate, or binate; if three in number, trifoliate, or ternate; when four in number, quadrifoliate, etc. If each of the leaflets of a pal- mately compound leaf divides into three, the leaf is called biternate; if this form again divides, a triternate leaf results. Beyond this point the leaf is known as decompound. In the case of pinnately- compound leaves, when division progresses so as to separate what would be a leaflet into two or more, the leaf becomes bipinnate, as the compound leaves of Acacia Senegal or on the new wood of PLANT ORGANS AND ORGANISMS 127 Gleditschia; if these become again divided, as in many Acacia species, the leaf is termed tripinnate. Examples of decompound leaves seen in Cimicifuga and Parsley. Leaf Texture. — Leaves are described as: Membranous, when thin and pliable, as Coca. Succulent, when thick and fleshy, as Aloes, and Live Forever. Coriaceous, when thick and leathery, as Eucalyptus, Uva Ursi and Magnolia. Leaf Color. — Petaloid, when of some brilliant color different from the usual green, as the Coleus and Begonia, and other plants which are prized for the beauty of their foliage rather than their blossoms. Leaf Surface.— Any plant surface is: Glabrous, when perfectly smooth and free from hairs or protuber- ances. Example: Tulip. Glaucous, when covered with bloom, as the Cabbage leaf. Pellucid- punctate, when dotted with oil glands, as the leaves of the Orange family. Scabrous leaves have a rough surface with minute, hard points. Pubescent, covered with short, soft hairs. Example: Strawberry. Villose, covered with long and shaggy hairs. Example: Forget- me-not. Sericious, silky. Example: Silverleaf. Hispid, when covered with short, stiff hairs. Example: Borage. Tomentose, densely pubescent and felt-like, as the Mullein leaf. Spinose, beset with spines, as in the Thistle. Rugose, when wrinkled. Example: Sage. Verrucose, covered with protuberances or warts. Duration of Leaves. — Leaves vary as to their period of duration. They are: Persistent, or evergreen, if they remain green on the tree for a year or more. Deciduous, if unfolding in spring and falling in autumn. Caducous, or fugacious, if falling early in the season. Parts of Typical Leaf.- — The parts of a typical leaf are petiole or leaf stalk, lamina or blade, and stipules. Gross Structure and Histology of the Petiole.— The petiole in Monocotyledons is usually a broadened, sheathing basal structure which connects the lamina to the stem. Into this a set of closed 128 PHARMACEUTICAL BOTANY collateral vascular bundles of the stem extend, these showing xylem uppermost and phloem beneath; but in the Palmacece, Aracece, Dioscoreacece and Musaceee the petiole in part or throughout may be much thickened, strengthened and developed as a semi-cylindric or cylindric structure frequently showing, as in Palmacece , generally, two sets of bundles. In all of these the petiole shows distinct scat- tered closed collateral bundles embedded in parenchyme and sur- rounded by epidermis. In the Monocotyl genus Maranta a special swelling is found at the apex of the petiole which is termed a pulvinus. In Dicotyledons the petiole attains its most perfect development and here usually shows differentiation into a pulvinus or leaf cushion and stalk portion. The pulvinus is sensitive to environal stimuli and in some groups as Oxalidacece and Leguminosce a gradual increase in sensitivity up to a perfect response can be traced. Moreover, in these, if we start with the simpler less sensitive pulvini and pass by stages to the most complex we note that a special substance known as the aggregation body develops in the pulvinar cortex cells and that this substance undergoes rapid molecular change on stimulation of the leaf. The stalk portion of the petiole in Dicotyledons is usually plano-convex or nearly to quite circular in outline; rarely in certain families does it simulate Monocotyledons in becoming abruptly or gradually thinned or flattened or widened out so as to sheath round the stem. The most striking example of this is seen in the Umbel- liferce where the flattened sheathing leaf stalk is known as the peri- cladium. Such a structure is not peculiar to the Umbelliferce for in many Ranunculacece, etc., a similar sheathing development is ob- served. The stalk may bear the laminar tissue on its extremity. This is most commonly the rule, but when the plant is exposed to xerophytic conditions, as the Acacias of Australia, the stalk instead of being cylindric or sub-cylindric becomes flattened from side to side until there is produced a bifacial vertically placed petiole with a large green surface that wholly takes the place of the lamina. The petiolar structure in primitive types of Dicotyls resembles that seen in Monocotyls except that the bundles are more con- densed side by side. In these the petiole is somewhat dorsiventral, shows an external epidermis, a flattened cortex with a set of parallel vascular bundles each with xylem uppermost and phloem below. PLANT ORGANS AND ORGANISMS 129 From this we pass to another group in which the bundles form three-fourths of a circle and in which the upper bundles show incurv- ing orientation, to still another in which as in Nepenthes all of the bundles form nearly a cylinder. Finally in Ficus, Geranium, Podo- phyllum and other plants showing completely formed cylindric petioles, the bundles form a continuous ring enclosing pith and sur- rounded by cortex and epidermis as in Dicotyl stems. Stipules. — Stipules are lateral leafy or membranous outgrowths from the base of the petiole at its junction with the stem. They may be divided into two groups, viz.: lateral and axillary. The lateral group includes four types, namely, free lateral, lateral adnate, lateral connate and lateral interpetiolar. Free lateral stipules are seen in Leguminosce, Rosacea, Beeches, etc. They are free on either side of the petiole and supplied by vascular tissue from the petiolar bundle mass. In appearance and duration they may be either green, foliaceous and persistent or mem- branous to leathery, scale-like and caducous. Caducous scaly stip- ules only function as bud scales through the winter and fall in spring as the buds expand. Lateral adnate stipules are such as fuse with and are carried up with the petiole as wing-like appendages. This type is seen in the genus Rosa, in Clovers, etc. Lateral connate stipules are such as join and run up with the petiole to form a structure which is called a ligule. This structure is ‘common to the Graminaceae or Grass family. Lateral interpetiolar stipules are common to many species of the Rubiaceae. In the genus Cinchona the leaves are opposite and orig- inally had free lateral stipules which latter gradually fused with the stem, slid across it and adjacent stipules, then fused together to form a median structure on either side of the stem. The axillary group represent stipules which stand in the axil of the leaf with the stem. Such may be free axillary structures, arising as distinct processes or connate, when the two stipules unite at their margins and sheath the stem, as in many species of the Polygonaceae such as Buckwheat, Rhubarb, Yellow Dock, Knot Weeds, etc. The sheath formed is called an ochrea. Modified Stipules. — In some plants such as the Locust and several i3° PHARMACEUTICAL BOTANY other trees and shrubs of the Legume family, the stipules become modified for defensive purposes as spines or prickles. In the Sarsa- parilla-yielding plants and other species of the genus Smilax they undergo modification into tendrils which are useful in climbing. The Lamina. — This as was previously indicated represents an ex- pansion of the tissues of the petiole, but in sessile leaves is directly attached to the stem and so a direct stem outgrowth. Mode of Development of the Lamina of Leaves. — The lamina of leaves develops in one of six ways. 1. Normal or Dorsoventral. 2. Convergent. 3. Centric. 4. Bifacial. 5. Reversed. 6. Ob-dorsi-ventral. The first four will be considered. A. Dorsoventral (the commonest). (a) Dorsoventral Umbrophytic.- — Flattened from above downward. Plants with such leafblades tend to grow in the shade. (b) Dorsoventral Meso phytic. — Similar to the former, but plants usually grow directly in the open and exposed to sunlight and winds. (c) Dorsoventral Xerophytic. — Similar to former, but plants not only grow exposed, but exposed to hot desert conditions or to cold vigorous conditions. (, d ) Dorsoventral Hydro phytic. — All transitions between typiaal mesophytic forms to those of marshy places, to swamps and borders of streams and finally with leaves wholly emersed, the last a com- pletely hydrophytic type. Gross Structure and Histology of Different Types of Dorsoventral Leaf Blades. — 1. Umbrophytic. — Characterized by leaves mostly undivided and having the largest and most continuous leaf expanse. Usually the deepest green leaves we have, to enable the leaves to absorb scattered and reduced rays that pass in through high trees and shrub overhead. Their texture is usually thin and soft. In microscopic structure they are covered with a cutinized epidermis which has all the stomata on the lower surface. The mesophyll is fairly spongy, the spongy parenchyme having decided intercellular V s Fig. 59- — Photomicrograph of cross-section through a portion of the leaf of a xerophyte, Ficus elastica, showing upper epidermis ( u.e .), water storage tissue ( w.s .), cystolith suspended on stalk within a cystolith sac ( cys ), palisade paren- chyma ( p.p . ), spongy parenchyma ( s.p .), vein (»), lower epidermis ( l.e .), and stoma (s). (Highly magnified.) ture, they consist of an upper and lower epidermis, the upper epider- mis being the thicker of the two. The stomata are wholly or are mainly on the lower epidermis. Hairs are seldom seen. The pali- sade mesophyll is toward the upper surface, the spongy mesophyll toward the lower. The intercellular air spaces in the spongy paren- chyma are small. 3. Xerophytic. — Leaves characterized by a thick upper and lower cuticle and by having their numerous, small stomata restricted to PLANT ORGANS AND ORGANISMS 131 spaces.. The lower epidermis is more or less hairy. Examples: Dog’s Tooth Violet, Asters. 2. Mesophytic. — Leaves tend to subdivision, either to slight or moderate lobing, seldom to complete subdivision in pinnate or tripinnate fashion. Example: Dandelion. Tn microscopic struc- 132 PHARMACEUTICAL BOTANY the lower surface or present more or less equally on both surfaces, where they are sunken in depressions. They may be either firm, leathery, tough, fibrous, or may become swollen up in their mesophyll chiefly in their spongy parenchyme cells and store considerable mucilage. Examples: Yucca, Ficus, Aloe, Agave. Succulent forms like Aloe generally possess a thin but tenacious cuticle. 4. Hydrophytic. — All gradations are seen. In pond plants, such as the Water Lily, the leaves have long split petioles which bring the blade up to the surface of the water. The stomata are entirely Fig. 60. — Photomicrograph of dorsoventral leaf*of Myrica showing upper epi- dermis (u.e.), palisade parenchyma (p.p.), vein (v), spongy parenchyma (s.p.), lower epidermis (l.e.), glandular-hairs (g) and stoma (5). (Highly magnified.) on the upper surface. In Ranunculus, the lower leaves are cut up into filiform segments. These are devoid of stomata. Their meso- phyll is soft, open, and spongy. The epidermis is quite thin. The upper leaves are floating, trilobed, and have stomata only on their upper surface. In Utricularia, some of the filiform submerged leaves are modified into bladders which trap insect larvae and smaller Crustaceae. B. Convergent. — In Phormium tenax, the base of the blade is sheathing, it then converges and opens out above. In the various species of Iris the petiole is sheathing, the upper part being fused (mostly seen in monocotyls). C. Centric. — Succulent. — Nearly always associated with Xero- phytes. PLANT ORGANS AND ORGANISMS 133 Xerophytic .— Centric laminae are produced gradually by an en- croachment of the under on the upper surface, and the swelling of the whole. In a completely centric leaf of the succulent kind, like that of Sedum, the difference between the upper and lower surface is lost. Stomata are found scattered over the entire epidermis. The bundles o.r. ep. v p.p. are arranged in a circle, the mid-rib being in the center. A great deal of mucilage is found stored in the central cells. In a typical Xerophytic Centric leaf, like that of the Pine or Sanseviera cylindrica , the epidermis shows a thick cuticle; the stomata are sunken in cavi- ties of the epidermis; the epidermis and leaf tissue are strengthened by scleroid bands in the centric mesophyll. ep. p.p. Pig. 61. — Photomicrograph of a transverse section of a bifacial leaf of Eucalyp- tus globulus showing epidermis (ep.), palisade parenchyma (p.p.), toward both surfaces, spongy parenchyma ( s.p .), vein (v), and oil reservoir (o.r.) lined with secretory epithelium. (Highly magnified.) 134 PHARMACEUTICAL BOTANY D. Bifacial. — Leaves with laminae which stand edge on in relation to the sun’s rays. The best illustrations are seen among dicotyle- dons, such as Eucalyptus, Callistemnon, and other genera of Myrtaceae. Both surfaces are similar, having stomata about equal in number. The mesophyll is differentiated into a central spongy parenchyme containing bundles, and a zone of palisade cells on either side facing the surfaces. Structure and Development of Stomata.— Stomata are slit-like openings in the epidermis of leaves or young green stems surrounded by a pair of cells called guard cells whose sides opposite one another are concave. They form a communication between the intercellular- air-space (respiratory cavity) beneath them and the exterior. The slit-like opening taken with the guard cells, constitutes what is known as the stomatal apparatus. The epidermal cells which abut on the stomatal apparatus are called neighboring cells or subsidiary cells. These in many cases, as in species of Helleborus, Sambucus, Hyacinthus, Pceonia, Ferns, etc., are very similar to the other epidermal cells but in a large number of plants they differ in size, arrangement and shape from the other cells of the epidermis which do not abut upon the stomatal apparatus. In Senna they are two in number one larger than the other and arranged parallel to the guard cells of the stoma; in Coca a similar arrangement occurs but the cells are more even in size, nevertheless they lack the characteristic papillae found on the other epidermal cells; in Pilo- carpus they are usually four in number but quite narrow and more or less crescent-shaped; in Uva Ursi their number is usually seven to eight and their arrangement radial around the stomatal apparatus. On all dorsoventral leaves, the stomata arise more abundantly on the lower epidermis, less abundantly on the upper. Exceptions to this rule are due to the peculiar readaptation of the leaf to its sur- roundings. Thus, in the reversed types of leaves (twisted in a half circle) the stomata, formerly on the lower surface, have migrated to the upper surface which now has become the physiological lower surface. In Umbrophytic (shade) plants the stomata are either wholly on the lower surface or partly so with a number on the upper surface. Where the plants are Mesophytic and exposed to dense sunlight and PLANT ORGANS AND ORGANISMS 135 leaves remain dorsiventral, the stomata are on the lower surface; these stomata are large, if the surroundings are damp. If such plants live in dry soil and dry air, the stomata are of small size and numer- ous; if they dwell in dry soil in hot surroundings and dense light they are very small and frequently sunk. If the plants are Xerophytic and the leaves dorsoventral, the stomata are quite abundant, small, with narrow slit, and depressed below the level of the epidermis. There are live types of stomatal development, viz.: First Type. — Each primitive epidermal cell (or the majority, or only certain ones of the epidermis) at the close of the dermatogen stage, gradually lengthens and then cuts off a smaller from a larger cell. The smaller one is equilateral, has a very large nucleus, and is termed the Stoma Mother-cell; the larger, quadrangular, and called the Epidermal Daughter-cell. The latter, upon maturing, becomes a normal epidermal cell. A partition is laid down lengthwise through the Stoma Mother-cell dividing it into two stomatal daughter-cells. The wall laid down lengthwise splits and thus forms the orifice of the stoma; the cells on either side of the orifice are called Guard Cells. These, while at first flat and inoperative, soon become bulged and crescent-shaped. This mode of development is seen in Squill, Hyacinth, Daffodil, Sambucus, Silene, etc. Second Type. — After the cutting off of the stomal mother-cell there are cut off on either side portions of neighboring epidermal cells which form subsidiary cells to the stoma. This condition is seen in Gram- inacece, Cyperacece, Juncacece, in various species of Aloe, Musa and Proteacece. Third Type .- — Instead of two parallel subsidiary cells, four are cut off, as in Heliconia, in species of Tradescanlia, Araucaria, or four to five, as in Ficus elastica, or four to five or more, as in the Coniferce and Cycads. Fourth Type.- — Instead of only four subsidiary cells, each of these again subdivides by parallel walls, more rarely by radial walls, into eight radiating subsidiary cells, as in Maranta bicolor, Pothos argyraa, some of Proteacece, etc. Fifth Type . — The “stomal mother-cell” divides once or several times before becoming the true mother-cell of the stoma. As a result of the divisions there are also formed one or more subsidiary 13 6 PHARMACEUTICAL BOTANY cells. This mode of development is seen in theLabiat(e,Papilionace(E, Crucifertz , Solanacea, Crassulacece, Cactacecz, and Begoniacece, also in a number of ferns. Histologic Differences between Leaves of Dicotyledsuo and Monocotyledons. — The following may be cited as broad compara- tive histologic differences between Dicotyl and Monocotyl leaves: Dicotyl Leaves 1. Epidermal cells usually iso-dia- metric or sinuous. 2. The stomata are on the whole more numerous but smaller. 3. Non-glandular and glandular hairs frequent on upper but more fre- quent on lower surface, or both. 4. Leaf glands which excrete varied products are rather abundant. 5. Water stomata over the upper sur- face, more rarely over the lower surface, are frequent, especially along margins of leaves. 6. Palisade and spongy parenchyma in dicotyledons are more distinct and palisade parenchyma is denser. 7. The vascular bundles, in their in- trinsic elements, are more indur- ated but the accessory fibrous sheath is feebly developed. 8. A greater variety of accessory prod- ucts of assimilation are de- veloped. Monocotyl Leaves 1. Epidermal cells usually elongate and equilateral. 2. Stomata larger. 3. Hairs rare in Monocotyls. 4. Leaf glands rare and only seen as a rule on the sepals. 5. Water stomata absent or very rare. Present in some Aracetz. 6. Palisade and spongy parenchyma is less distinct and dense. 7. The vascular bundles, in their in- trinsic elements, are less indur- ated. The fibrous sheath is strongly developed. 8. A comparatively small variety of accessory products of assimila- tion are developed. INFLORESCENCE Inflorescence or Anthotaxy. — A typical flower consists of four whorls of leaves modified for the purpose of reproduction, and com- pactly placed on a stem. The terms Inflorescence and Anthotaxy are applied to the arrangement of the flowers and their position on the stem, both of which are governed by the same law which determines the arrangement of leaves. For this reason flower buds are always either terminal or axillary. In either case the bud may develop a PLANT ORGANS AND ORGANISMS 137 solitary flower or a compound inflorescence consisting of several flowers. Determinate, cymose, descending, or centrifugal inflorescence is that form in which the flower bud is terminal, and thus determines or completes the growth of the stem. Example: Ricinus communis. Indeterminate, ascending, or centripetal inflorescence is that form in which the flower buds are axillary, while the terminal bud continues to develop and increase the growth of the stem indefinitely. Exam- ple: the Geranium. Mixed inflorescence is a combination of the other two forms. Example: Horse Chestnut. Fig. 62. — Types of indeterminate inflorescence. A, A raceme; B, a spike; C, a catkin; D, a corymb; E, an umbel. The flowers are represented by circles; the age of the flower is indicated by the size. ( From Hamaker.) Tht flower stalk is known as the peduncle, and its prolongation the rachis, or axis of the inflorescence. The flower stalk of a single flower of an inflorescence is called a pedicel. When borne without such support the flower is sessile. A peduncle rising from the ground is called a scape, previously mentioned under the subject of stems. The modified leaves found on peduncles are termed bracts. These vary much the same as leaf forms, are described in a similar manner, PHARMACEUTICAL BOTANY 138 and may be either green or colored. When collected in a whorl at the base of the peduncle they form an involucre, the parts of which are sometimes imbricated or overlapping, like shingles. This is gen- erally green, but sometimes petaloid, as in the Dogwood. The modified leaves found on pedi- cels are called bradeolar leaves. The Spat he is a large bract en- veloping the inflorescence and often colored, as in the Calla, or membranous, as in the Daffodil. Indeterminate Inflores- cences. — In the indeterminate or axillary anthotaxy, either flowers are produced from base to apex, those blossoming first which are lowest down on the rachis or from margin to center. The principal forms of this type are: Solitary Indeterminate. — The simplest form of inflorescence in which a single flower springs from the axil of a leaf. A number of these are generally developed on the same stem. Example: Periwinkle. Raceme, or simple flower- cluster in which the flowers on pedicels of nearly equal length are arranged along an 'axis. Examples: Convallaria, Cimi- cifuga, and Currant. Corymb, a short, broad cluster, differing from the raceme mainly in its shorter axis and longer lower pedicels, which give the cluster a flat appearance by bringing the individual florets to nearly the same level. Example: Cherry. Umbel, which resembles the raceme, but has a very short axis, and ax Pig. 63. — Photomicrograph of longi- tudinal section, through a staminate catkin of Comptonia asplenifolia X 10, showing catkin axis (ax), anther-lobe (a), and bract (b). PLANT ORGANS AND ORGANISMS 139 the nearly equal pedicels radiate from it like the rays of an umbrella. Many examples of this mode of inflorescence are seen in the family Umbelliferce, as indicated by the name, including Anise, Fennel and other drug-yielding official plants. A Spike is a cluster of flowers, sessile or nearly so, borne on an elongated axis. The Mullein and common Plantain afford good illustrations. The Catkin or Ament resembles the Spike, but differs in that it has scaly instead of herbaceous bracts, as the staminate flowers of the Oak, Hazel, Willow, Comptonia, etc. The Head or Capitulum is like a spike, except that it has the rachis shortened, so as to form a compact cluster of sessile flowers, as in the Dandelion, Marigold, Clover, and Burdock. The Strobile is a compact flower cluster with large scales concealing the flowers, as the inflorescence of the Hop. The Spadix is a thick, fleshy rachis with flowers closely sessile or embedded on it, usually with a spathe or sheathing bract. Example: Calla, Acorus Calamus, Arum triphyllum . The compound raceme, particularly if irregularly compounded, is called a panicle. Determinate Inflorescences. — Determinate Anthotaxy is one in which the first flower that opens is the terminal one on the axis, the other appearing in succession from apex to base or from center to margin. The principal varieties are: The Solitary Determinate, in which there is a single flower borne on the scape, as in the Anemone, or Windflower, and Hydrastis. The Cyme, a flower cluster resembling a corymb, except that the buds develop from center to circumference. Example: Elder. If the cyme be rounded, as in the Snowball, it is a globose cyme. A Scorpoid Cyme imitates a raceme, having the flowers pedicelled and arranged along a lengthened axis. A Glomerule is a cymose inflorescence of any sort which is con- densed into a head, as the so-called head of Cornus florida. A V erticillaster is a compact, cymose flower cluster which resembles a whorl, but really consists of two glomerules situated in the axils of opposite leaves. Clusters of this kind are seen in Catnip, Hore- hound, Peppermint and other plants of the Labiatce. 140 PHARMACEUTICAL BOTANY The raceme, corymb, umbel, etc., are frequently compounded. The compound raceme, or raceme with branched pedicels, is called a panicle. Examples: Yucca and paniculate inflorescence of the oat. A Thyrstis is a compact panicle, of a pyramidal or oblong shape. Examples: Lilac, Grape and Rhus glabra. A Mixed Anthotaxy is one in which the determinate and indeter- minate plans are combined, and illustrations of this are of frequent occurrence. Fig. 64. — Cymose inflorescences. F, A terminal flower; G, a simple cyme; H, a compound cyme. ( From Hamaker.) The order of flower development is termed ascending when, as in the raceme, the blossoms open first at the lower point on the axis and continue to the apex. Examples: WhiteLily, and many other plants of the same family. In the cyme the development is centrifugal, the central florets opening first, while in the corymb it is centripetal, or from margin to center. PREFLORATION Prefloration. — By prefloration is meant the arrangement of the floral envelopes in the bud. It is to the flower bud what vernation is to the leaf bud, the same descriptive terms being largely employed, as convolute, involute, revolute, plicate, imbricate, etc. In addition to those already defined, the following are important. PLANT ORGANS AND ORGANISMS 141 V ah ale Prefloration , in which the margins meet but do not over- lap. Of this variety the induplicate has its two margins rolled inward as in Clematis. In the reduplicate they are turned outward, as the sepals of Althaea. Vexillary, the variety shown in the corolla of the Pea, where the two lower petals are overlapped by two lateral ones, and the four in turn overlapped by the larger upper ones. Contorted, where one margin is invariably exterior and the other interior, giving the bud a twisted appearance, as in the Oleander and Phlox. THE FLOWER The flower is a shoot which has undergone a series of changes so as to serve as a means for the propagation of the individual. Fig. 65. — Diagrams of floral structures. A, Shows the relations of the floral parts inja hypogynous flower; B, the same in a perigynous flower; C, the same in an epigynous flower; D, a stamen; E, a simple pistil in longitudinal section; F, the same in cross-section; G, transitional forms between true petals (left) and true stamensi/right) ; H, slight union of two carpels to form a compound pistil; / and J, union of carpels more complete; K and L, cross-sections of compound pistils, of three carpels. In) 3 : a, stamen; b, petal; c, sepal; d, pistil; e, receptacle; /, pedicel. In D: a, anther cell; b, connective; c, filament. In E: a, stigma; b, style; c, ovules; d, ovary. ( From Hamaker.) A Typical or Complete Flower possesses four whorls of floral leaves arranged upon a more or less shortened stem axis called a receptacle, torus or thalamus. These whorls passing from periphery toward the 142 PHARMACEUTICAL BOTANY center are: calyx , composed of parts called sepals; corolla , composed of parts termed petals; andr cerium, composed of parts called stamens or microsporopylls; and gyncecium, composed of one or more parts termed carpels or megasporophylls. The stamens and carpels constitute the essential organs, and a flower is said to be Perfect when these are present and functional. A Hermaphrodite flower is one which possesses both stamens and carpels which may or may not be functionally active. In some cases the stamens may alone be functional while in others the carpels only may function. A Regular Flower possesses parts of each whorl of the same shape and size, as the flower of Veratrum. It is Symmetrical when the parts of each whorl are of the same number, or multiples of the same number. An Imperfect Flower shows one set of essential organs wanting. When either petals or sepals, or both, are present in more than the usual number, the flower is said to be “ double,” as the cultivated Rose and Carnation. The doubling of flowers is brought about through cultivation and is due either to the transformation of sta- mens (as in cases cited), and occasionally of carpels into petals, to a division of the petals, or to the formation of a new series of petals. If the pistils are present and stamens wanting, the flower is called pistillate, or female; if it possesses stamens but no pistil, it is described as staminate, or male; if both are absent, neutral, as marginal flowers of Viburnum. Some plants, as the Begonias and Castor oil bear both staminate and pistillate flowers, and are called Monoecious. When the staminate and pistillate flowers are borne on different plants of the same species, they are termed Dioecious, as the Sassafras and Willow. When staminate, pistillate and hermaphrodite flowers are all borne on one plant, as on the Maple trees, they are polygamous. Connation and Adnation. — In the development of the flowers of primitive species of flowering plants, the parts of each whorl are dis- joined or separate from each other. In many higher types, however, the parts of the same whorl frequently become partly or completely united laterally. This condition is termed connation, coalescence, cohesion or syngenesis. Illustrations of this may be seen in Bella- donna, Stramonium and Uva Ursi flowers, where the petals have PLANT ORGANS AND ORGANISMS 143 joined laterally to form gamopetalous corollas. When the one or more parts of different whorls are united, as of stamens with petals (. Rhammus ) or stamens with carpels ( Apocynum ) the union is called adnation or adhesion. The Receptacle. — The Receptacle , Torus or Thalamus is a more or less shortened axis (branch) which bears the floral leaves. It is usually flat or convex, but may be conical and fleshy as in the Straw- berry, concave as in the Rose and Fig or show a disc-like modifica- tion as in the Orange. The internodes of the receptacle in many species lengthen and separate various whorls. When the lengthen- ing of the internode occurs between calyx and corolla as in Lychnis the structure resulting is called an anthophore; if between corolla and androecium as in Passiflora, a gonophore; if between androecium and gynoecium as in Geum, a gynophore. If the flowers of the Umbel- liferce the receptacle elongates between the carpels producing the structure called a carpophore. The Perigone.- — The perigone or perianth is the floral envelope consisting of calyx and corolla (when present). When both whorls, i.e., calyx and corolla, are present the flower is said to be dichlamydeous; if only calyx is present, monochlamydeous. The Calyx. — The Calyx is the outer whorl of modified leaves. Its parts are called Sepals, and may be distinct (Chorisepalous, from a Greek word meaning disjoined) or more or less united (Gamosep- alous). They are usually green — foliaceous or leaf-like — but may be brilliantly colored, hence the term petaloid (like the petals) is applied. Examples: Tulip, Larkspur, Columbine and Aconite. In a gamosepalous calyx, when the union of sepals is incomplete, the united portion is called the tube, the free portion, the limb, the orifice of the tube, the throat. In form the calyx may be regular or irregular; regular, if its parts are evenly developed, and irregular if its parts differ in size and shape. The more common forms are tubular, resembling a tube; rotate, or wheel-shape; campanulate, or bell-shaped; urceolate, or urn-shape; hypocrateriform, or salver-shape; bilabiate, or two-lipped; corres- ponding to the different forms of corolla, under which examples illustrating each will be given. The calyx usually remains after the corolla and stamens have 144 PHARMACEUTICAL BOTANY fallen, sometimes even until the fruit matures — in either case it is said to be persistent. If it falls with the corolla and stamens, it is de- cidous, and if when the flower opens, caducous, as in the Poppy and May-apple. It often more or less envelops the ovary or base of the pistil, and it is important, in plant analysis, to note the presence or absence of such a condition, which is indicated in a description by the terms inferior, or non-adherent (hypogynous), when free from the ovary and inserted upon the receptacle beneath it (the most simple and primitive position); half-superior, or half-adherent (perigynous), when it partially envelops the ovary, as in the Cherry; superior or adherent (epigynous), when it completely envelops it, as in the Colocynth, etc. Sepaline Spurs. — Occasionally some or all of the sepals may become pouched and at length spurred as nectar receptacles or as receptacles for other parts that are nectariferous. Thus, in Cru- cifers we occasionally see a slight pouching of the two lateral sepals. These act as nectar pouches for the nectar secreted by the knobs or girdles surrounding the short lateral stamens. These become deep pouches in Lunaria while in others the pouches become elongated spurs. Again, in Delphinium, the posterior sepal forms an elongated spur into which pass the two spurred nectariferous petals. In Aconitum the same sepal, instead of being spurred, forms an enlarged hood-like body (galea) arching over the flower like a helmet; into this pass the two hammer-shaped nectariferous petals. Sepaline Stipules.- — These structures are well developed and easily traceable in the more primitive herbaceous members of the Rose family. Thus in Potentilla, Fragaria, Geum, etc., in addition to the normal calyx of five sepals, there is a supplementary epicalyx also of five parts. The five lobes of the epicalyx may be as large or larger than the sepals or smaller up to the disappearing point. Upon examining a few flowers of Potentilla or Fragaria, it will be observed that not infrequently one, sometimes two lobes of the epicalyx are bifid, or deeply cleft, or separated completely into two parts. The explanation is that the five sepals after evolving in the flower bud form at their bases two lateral swellings or sepaline stipules, which, as they grow, fuse in adjacent pairs, one stipule of one sepal joining with the adjacent stipule of another sepal to form five lobes. \ PLANT ORGANS AND ORGANISMS 145 Sepaline Position.- — As already noted the most simple and primi- tive position for the sepals in relation to the floral parts is hypogy- nous, in which the sepals are inserted directly into the enlarged floral axis (receptacle) below the petals, stamens and carpels. But in the more primitive herbaceous Rosacea, Leguminosa, etc., the floral axis forms a saucer-like transverse expansion which pushes out the sepals, petals and stamens on its edge. Thus originates the perigy- nous insertion of the sepals. In not a few higher Rosacea, Saxifraga- cea, Crassulacea, etc., the saucer-like floral axis becomes deepened and contracted into a cup-shaped structure (Cherry, Peach, Almond, Plum, etc.), and on the edge of this cup the sepals as well as the petals and stamens are inserted at different levels. Finally, in the Apple, Pear, Quince, etc., the greatly hollowed-out receptacle assumes a vase-shaped form and closes over the top of the ovary, at the same time lifting the sepals, petals, and stamens above the ovary. Here the sepals are epigynous. The Corolla. — The Corolla is the inner floral envelope, usually delicate in texture, and showing more or less brilliant colors and combinations of color. Its parts are called Petals, and when the calyx closely resembles the corolla in structure and coloring they are together called the Perianth. The purpose of these envelopes is to protect the reproductive organs within, and also to aid in the fertilization of the flower, as their bright colors, fragrance and sac- charine secretions serve to attract pollen-carrying insects. Forms of the Corolla and Perianth. — When the petals are not united with each other, the corolla is said to be Choripetalous, Apo- petalous or P oly petalous . When more or less united, they are Gamo- petalous, often called Synpetalous. When the distinct petals are four in number, and arranged in the form of a cross, the corolla is called Cruciform. Example: Mustard and other plants belonging to the order Cruciferae. The Papilionaceous corolla is so called because of a fancied resem- blance to a butterfly. The irregularity in this form is very striking, and the petals bear special names: the largest one is the vexillum, or standard; the two beneath it the ala, or wings; the two anterior, the carina or keel. Examples: Locust, Pea, and Clover. Orchidaceous flowers are of peculiar irregularity, combining calyx 10 146 PHARMACEUTICAL BOTANY and corolla. The petal in front of stamen and stigma, which differs from the others in form and secretes nectar, is called the Labellum. Examples: Cypripedium and other Orchids. When calyx and corolla each consist of three parts closely resem- bling each other in form and color, as in the Tulip and Lily, the flower is called Liliaceous. The Ligulate or Strap-shaped corolla is nearly confined to the family Compositse. It is usually tubular at the base, the remainder resembling a single petal. Examples : Marigold, and Arnica Flowers. Labiate, or Bilabiate, having two lips, the upper composed of two petals, the lower one of three. This form of corolla gives the name to the Labiatce, while in the family Leguminosoe this arrangement is sometimes reversed. The corolla may be either ringent, or gaping, as in Sage, or personate, when the throat is nearly closed by a projec- tion of the lower lip, as in Snapdragon. Rotate, Wheel-shaped, when the tube is short and the divisions of the limb radiate from it like the spokes of a wheel. Example: The Potato blossom. Crateriform, Saucer-shaped, like the last, except that the margin is turned upward or cupped. Example: Kalmia latifolia (Mt. Laurel). Hypocrateriform, or Salver-shaped (more correctly, hypocrateri- morphous), when the tube is long and slender, as in Phlox or Trail- ing Arbutus and abruptly expands into a flat limb. The name is derived from that of the ancient Salver, or hypocraterium with the stem or handle beneath. When of nearly cylindrical form, the corolla is Tubular, as in the Honeysuckle, and Stramonium. Funnel-form (Infundibuliform), such as the corolla of the common Morning Glory, a tube gradually enlarging from the base upward into an expanded border or limb. Campanulate, or Bell-shaped, a tube whose length is not more than twice the breadth, and which expands gradually from base to apex. Examples: Canterbury Bell, Harebell. Urceolate, or urn-shaped, when the tube is globose in shape and the limb at right angles to its axis, as in the official Uva Ursi, Chima- phila and Gaultheria. PLANT ORGANS AND ORGANISMS 147 Caryophyllaceous, when the corolla consists of five petals, each with a long slender claw expanding abruptly at its summit into a broad limb. Examples: Carnation and other members of the Pink family. The Andrcecium or Stamen System. — The andr cerium is the single or double whorl of male organs situated within or above the corolla. It is composed of stamens or micros porophylls. A complete stamen consists of a more or less slender stalk portion called & filament and a terminal appendage called the anther or micro- sorns. The anther is generally vertically halved by an upgrowth of the filament called the connective, dividing the anther into two lobes. Number of Stamens. — When few in number, stamens are said to be definite; when very numerous, and not readily counted, they are indefinite. The following terms are in common use to express their number: Monandrous, for a flower with but one stamen. Diandrous, with two stamens. Triandrous, with three. Tetrandrous , with four. Pentandrous, having five. Hexandrous, six. Polyandrous, an indefinite number. The most primitive flowers have numerous stamens but passing from these to those of more evolved families there occurs a gradual reduction from many to ten, as in Caryophyllacece, Leguminosce and some Aceracece, these being in two circles. In Malvacece, Umbelliferce and other Apopetalous families as well as many Sympetalae, the number five is typical. But in Scroplmlariacece, while five are devel- oped and fertile in Verbascum, four with a fifth staminode (sterile stamen) are found in the allied genus Celsia. In Pentstemon there are four didynamous fertile stamens and an equally long staminode. In Scrophularia the fifth staminode is reduced to a petaloid flap in the posterior part of the flower. In Linaria this exists only as a small knob at the base of the back part of the corolla and there secretes nectar. In most Scroplmlariacece the fifth stamen is entirely absent and the four stamens left are didynamous; but in Calceolaria two of these are rudimentary and thread-like, the other two alone 148 PHARMACEUTICAL BOTANY being well-developed and fertile. In Veronica three stamens are entirely absorbed and two only are left as fertile representatives. Insertion of Stamens. — As to insertion the stamens may be: Hypogynous , when inserted upon the receptacle below the base of the pistil. Perigynous, when inserted on the calyx or corolla above the base of and lateral to the pistil. Epigynous, when inserted above the ovary. Gynandrous, when inserted upon the pistil, as in Orchids and Aris- tolochia. Proportions of the Stamens. — The stamens may be of equal length; unequal, or of different length. Didynamous , when there are two pairs, one longer than the other. Example: Snapdragon. Tetradynamous , three pairs, two of the same length, the third shorter. Example: Mustard. Connation of Stamens. — Terms denoting connection between sta- mens are: Monad el phous (in one brotherhood), coalescence of the filaments into a tube. Example: Lobelia. Diadelphous (in two brotherhoods), coalescence into two sets. Example: Glycyrrhiza. Triadel phous, with filaments united into three sets. Example: St. John’s Wort. Polyadelphous, when there are several sets or branched bundles. Example: Orange. Syngenesious, when the anthers cohere. Example: Compositce. Color of Stamens. — In most species the color of these organs is seldom pronounced owing to their delicate structure. It varies from greenish-yellow to yellow to white, through pink, pinkish-red, red, purple, purple-blue to blue. It is yellow, for instance, in Sassa- fras, Cucumber and Golden Club; greenish-yellow, yellow to red in Maples; yellow-pink to pink and pinkish-red in some Mallows; in Azalea amena the filaments are crimson-purple and the anthers, pur- ple-blue; in the genus Scilla both filaments and anthers are blue. Gross Structure and Histology of the Filament. — The filament may be cylindric as in the Rose, awl-shaped as in Tulip, flat and with PLANT ORGANS AND ORGANISMS 149 a dilated base as in the Harebell, three-toothed as in Garlic, appen- diculate, when it bears an appendage as in Chcetostoma, Alyssum, etc. The filament is covered with a protective epidermis containing stomata. Beneath this is a soft, loose cellular tissue, the mesophyll, and in the center a small vascular bundle, the pathway of food from the floral axis to the anther. In some cases the single bundle may split into two or three bundle parts. Gross Structure and Histology of the Anther. — Each staminal leaf (microsporophyll) bears a special development or appendage as a rule on its extremity which is the anther or microsorus. This consists, fundamentally, of a median prolongation of the filament equal to the connective or placenta. This develops on either side a quantity of indusial tissue that grows out to form a covering substance that protects and carries two microsporangia on either side. An anther therefore consists of a median connective or placenta, producing on either side two anther lobes or indusial expansions. Each anther lobe encloses two pollen sacs or microsporangia which, in some cases, remain distinct up to the dehiscence (splitting open) of the anther. Thus in Butomus , the anthers show four pollen chambers up to the time of dehiscence. Again in various species of Lauracece, the anthers remain four lobed and dehisce by four recurved lids. But in the great majority of Angiosperms each pair of pollen sacs fuse before dehiscence, owing to the breaking down of the partition between them, and so, at that time, show two-celled anthers. Still more rarely the anthers may be two-celled in their young state and by the breaking down of the partition become one-celled, e.g., Malvaceae. Externally the mature anther is bounded by an exothecium or epi- dermis, often swollen, where lines of dehiscence occur, which may develop stomata, also hairs. Within it is a combined layer or set of one to often two or three, sometimes five or six cell layers (Agave, etc.) of indusial and sporangial cells, the endothecmm. The outermost one to three layers of this become spirally, annularly or stellately thickened to form the elastic tissue of the anther, which, by pressure against the delicate epidermis or exothecium, causes ultimate rupture of the anther wall. Within the innermost endothecial layer, bounding each sporangium, is the tapetum, a single-celled layer. This, near the time of dehiscence, undergoes breaking down or absorption by developing PHARMACEUTICAL BOTANY 150 pollen or microspore cells. Filling the cavities of the four sporangia are the mature pollen grains. The connective shows in or near its center a vascular bundle with xylem uppermost and phloem down- ward, surrounded by thin- walled cellular tissue, from which the indu- sial and sporangial substance has matured by extension. Anther Dehiscence. — This is the breaking open of the anther to discharge the pollen. When fully ripe the dividing partition between each pair of spor- angia usually becomes thinned, flattened and ultimately breaks down, Fig. 66. — Cross-section of a mature lily anther. The pairs of pollen chambers unite to form two pollen sacs, filled with pollen grains; s, modified epidermal cells at line of splitting. ( From a Text-book of Botany by Coulter, Barnes, and Cowles. Copyright by the American Book Company, Publishers.) while the elastic and resistant endothecium, steadily pushing against the more delicate and now shrinking exothecium causes rupture where endothecium is absent, namely along opposite lines of the anther wall. Thus arises a line of anther dehiscence called longi- tudinal anther dehiscence on either side of the anther sacs. In the division Solanece of the family Solanaceee which includes Belladonna, in some of the Ericacece as Rhododendron and Azalea, etc., the anthers dehisce by small apical pores from which the pollen is shed. This kind of dehiscence is called apical porous dehiscence. Again, in Lauracece and Berberidacece , the anthers dehisce by recurved valves. This is called valvular dehiscence. PLANT ORGANS AND ORGANISMS Moreover, in Malvacece the originally longitudinal anther is divided internally by a partition. It gradually swings on the filament so that eventually the anther is transverse and the partition becomes absorbed, thus becoming a one-celled anther with transverse dehis- cence in its mature state. Development of the Anther. — Each stamen originates as a knob- like swelling from the receptacle between the petals and carpels. This swelling represents mainly future soral (anther) tissue. The fila- ment develops later. When such a young sorus or anther is cut across and examined microscopically, it shows a mass of nearly simi- lar cellular tissue in which the first observable changes are the following: The surface dermatogen cells become somewhat flattened and regu- lar to form the future epidermis or exothecium of the anther. About the same time some cells, by more rapid division in the middle of the anther substance, give rise to the elements of the vascular bundle in the connective. Then, along four longitudinal tracts, rows of cells remain undivided or only divide slowly as they increase in size and around them cells divide and redivide to form the future endo- thecial and covering tissue to the four sporangia. Next, the four sporangial tracts of undivided cells cut off from their outer surfaces a layer of enveloping cells, the tapetum. This consists of richly proto- plasmic cells that form a covering to the spore mother-cells within. Each spore mother-cell undergoes division and redivision into four spore daughter-cells at the same time that reduction in the chroma- tin substance takes place in these cells. Thus originate tetrads (groups of four) of spore daughter-cells inside spore mother-cell wall. These continue to enlarge, press against the mother-cell wall which becomes converted into mucilage and each of the tetrad cells becomes in time a mature microspore or pollen grain. During this time the entire anther is growing in size, the cells of the endothecium in one or more layers becomes thickened by lignin deposits to form a mechanical endothecium; the tapetum gradually breaks down and appears only at length as an irregular layer around the maturing pollen cells. When the anther is finally ripe the parti- tion between each pair of microsporangia becomes narrowed, flat- tened and ruptured and thus numerous microspores or pollen grains 152 PHARMACEUTICAL BOTANY fill two cavities, one on either side of the connective. The micro- spores or pollen grains at first show only a thin clear cellulose layer, but from this, by a differentiation of the exterior film, the exospore layer becomes cut off. This becomes cuticular. The cellulose inner layer ( endospore ), remains unaltered. In the development of the exospore, one to several deficiencies are usually left in it through which the endospore may protrude later as the rudiment of the pol- len tube. Attachment of Anther. — The attachment of the anther to the fila- ment may be in one of several ways, as follows: Innate, attached at its base to the apex of the filament. Adnate, adherent throughout its length. Versatile, when the anther is attached near its center to the top of the filament, so that it swings freely. The adnate and versatile are introrse when they face inward, extrorse when they face outward. Pollen. — The pollen grains or microspores vary in form for differ- ent species and varieties and while they are averagely constant for these, nevertheless many exceptions have been recorded. The fol- lowing are some of the commoner forms: Four Spore Daughter -cells, hanging together as in the Cat Tail ( Typha ) forming a pollen grain. Elongated, simple pollen grains as in Zostera. Dumb-bell-shaped, as the pollen of the Pines. Triangular , as in the ( Enotheras . Echinate, as in the Malvaceae. Spherical, as in Geranium, Cinnamon and Sassafras. Lens-shaped, as in the Lily. Spinose, as in the Composite. Barrel-shaped, as in Polygala. Under the microscope the immature pollen grain generally consists of two membranes, an outer firmer one called the exospore , which may be variously marked and which possesses deficiencies in the form of “pores ”or “ clefts,” andan inner delicate cellulose membrane called the endospore, which surrounds a protoplasmic interior in which are imbedded a nucleus, oil droplets and frequently starch or protein. Pig. 67. — Various forms of pollen grains. Pollen from Typha latifolia (A), Zea mays ( B ), Ambrosia elatior (C), Lilium philadelphicum ( D ), Pinus (E), Ranunculus bulbosus ( F ), Car pinus caroliniana (G), Althcea rosea (H), Oenothera biennis, (/). All highly magnified. Drawing by Hogstad. I 54 PHARMACEUTICAL BOTANY Pollinia. — These are agglutinated pollen masses which are com- mon to the Orchidacece and Asclepiadacece. The pollen of many plants, notably certain species of Composite, Graminece and Rosacea, has been shown to be responsible for “Hay Fever.” At the present time serums, extracts* and vaccines are manufactured from pollen to be used in the treatment of this disease. The Gynoecium or Pistil System.- — This is the female system of organs of flowering plants. It may consist of one or more modified leaves called carpels. Each carpel or megasporophyll is a female organ of reproduction. In the Spruce, Pine, etc., it consists of an open leaf or scale which bears but does not enclose the ovules. In angiosperms it forms a closed sac which envelops and protects the ovules, and when complete is composed of three parts, the ovary or hollow portion at the base enclosing the ovules or rudimentary seeds, the stigma, or apical portion which receives the pollen grains, and the style, or connective which unites these two. The last is non- essential and when wanting the stigma is called sessile. The carpel clearly shows its relations to the leaf, though greatly changed in form. The lower portion of a leaf, when folded lengthwise with the margins incurved, represents the ovary; the infolded surface upon which the ovules are borne is the placenta, a prolongation of the tip of the leaf, the stigma, and the narrow intermediate portion, the style. A leaf thus transformed into an ovule-bearing organ is called a carpel. The carpels of the Columbine and Pea are made up of single carpels. In the latter the young peas occupy a double row along one of the sutures (seams) of the pod. This portion corre- sponds to the infolded edge of the leaf, and the pod splits open along this line, called the ventral suture. Dehiscence, or the natural opening of the carpel to let free the contained seeds, takes place also along the line which corresponds to the mid-rib of the leaf, the dorsal suture. The gynoecium or Pistil may consist of a number of separate carpels, as in the buttercup or Nymphaea flowers, when it is said to be apocarpous, or the carpels composing it may be united together to form a single structure, as in the flowers of Belladonna and Orange, when it is called syncarpous. If the pistil is composed of one carpel, it is called monocar pellary; PLANT ORGANS AND ORGANISMS 155 if two carpels enter into its formation, it is said to be dicarpellary; if three, tricar pellary; if many, poly car pellary. Compound Pistils are composed of carpels which have united to form them, and therefore their ovaries will usually have just as many cells (locules) as carpels. When each simple ovary has its placenta, or seed-bearing tissue, at the inner angle the resulting compound ovary has as many axile or central placentae as there are carpels, but all more or less consolidated into one. The partitions are called dissepiments and form part of the walls of the ovary. If, however, the carpels are joined by their edges, like the petals of a gamopetalous corolla, there will be but one cell, and the placenta will be parietal, or on the wall of the compound ovary. The ovules or megasori are transformed buds, destined to become seeds in the mature fruit. Their number varies from one to hun- dreds. In position, they are erect, growing upward from the base of the ovary, as in the Compositse; ascending, turning upward from the side of the ovary or cell; pendulous, like the last except that they turn downward; horizontal, when directed straight outward; suspended, hanging perpendicularly from the top of the ovary. In Gymnosperms the ovules are naked; in Angiosperms they are enclosed in a seed vessel. A complete angiospermous seed ovule which has not undergone maturation consists of a nucellus or body; two coats, the outer and inner integuments ; and a funiculus, or stalk. Within the nucellus is found the embryo sac or megaspore containing protoplasm and a nucleus. The coats do not completely envelop the nucellus, but an opening at the apex, called the foramen or micropyle admits the pollen tube. The vascular plexus near the point where the coats are attached to each other and to the nucellus is called the chalaza. The hilum marks the point where the funiculus is joined to the ovule, and if attached to the ovule through a part of its length, the adherent portion is called the raphe. The shape of the ovule may be ortho- tropous, or straight; campylotropous, bent or curved; amphitropous, partly inverted; and anatropous, inverted. The last two forms are most common. A campylotropous ovule is one whose body is bent so that the hilum and micropyle are approximated. PHARMACEUTICAL BOTANY 156 The Placenta . — The placenta is the nutritive tissue connecting the ovules with the wall of the ovary. The various types of pla- centa arrangement (placentation) are grouped according to their relative complexity as follows: (1) Basilar, (2) Sutural, (3) Parietal, (4) Central, (5) Free Central. Basilar placentation is well illustrated in the Polygonacece (Smart Weed, Rhubarb, etc.) in Piper and Juglans. Here, at the apex of the axis and in the center of the ovarian base, arises a single ovule from a small area of placental tissue. Sutural placentation is seen in the Leguminosce (Pea, Bean, etc.). Here each carpel has prolonged along its fused edges two cord-like placental twigs, from which start the funiculi or ovule stalks. Parietal placentation is seen in Gloxinia , Gesneria, Papaver, etc. Here we find two or more carpels joined and placental tissue running up along edges of the fused carpels bearing the ovules. Central or axile placentation is seen in Campanulacece (Lobelia), where the two, three, or more carpels have folded inward until they meet in the center and in the process have carried the originally parietal placenta with them. This then may form a central swelling bearing the ovules over the surface. Free Central placentation occurs perfectly in the Primulacece, Plantaginacece and a few other families. In this the carpels simply cover over or roof in a central placental pillar around which the ovules are scattered. Style. The style is the portion of the carpel which connects the stigma with the ovary. It is usually thread-like but may also be con- siderably thickened. It frequently divides into branches in its upper part. These are called style arms. As many style arms as carpels may be present. In the one-carpelled pistil of some Leguminosce , the usually bent-up style is the tapered prolongation of a single flower. Again, in the apocarpous carpels of many flowers of the Ranuncul- acece, each carpel bears a short to long stylar prolongation. When the carpels, however, are syncarpous the common styles tend to become more or less fused but usually show lobes, clefts or style arms at their extremities that indicate the number of carpels in each case which form the gynoecium. In some plants remarkable variations from the typical stylar PLANT ORGANS AND ORGANISMS 157 development may occur. Thus, in Viola, the end of the style is a swollen knob on the under surface of which is a concave stigma with a flap or peg. In the genus Canna the style is an elongate blade-like flattened body with a sub-terminal stigma. In forms of the Cam- panulacecE the style is closely covered with so-called collecting hairs. On these the anthers deposit their pollen at an early period before the flowers have opened. Later, when the flowers open, insects remove the pollen after which the collecting hairs wither. The stig- mas then curl apart to expose their viscid stigmatic hairs. In this instance there are two distinct and at separate times functioning hairs on the stylar prolongation, viz.: (a) collecting stylar hairs, functioning for pollen collection and distribution; and ( b ) stigmatic hairs for pol- len reception from another flower. In Vinca the style swells near its extremity into a broad circular stigma and then is prolonged into a short column bearing a tuft of hairs that prevents the entrance of insect thieves into the flower. In the genus Iris the common style breaks up at the insertion of the perianth into three wide petaloid style arms. Each of these bifurcates at its extremity. On the lower or outer face of this is a transverse flap that bears the stigmatic papillae. In Physostigma the style enlarges at its extremity into a flap-like swelling which bears a narrow stigmatic surface. Finally in Sarracenia the single style of the five-carpelled pistil enlarges above into a huge umbrella-like portion with five radiating ribs. At the extremity of each bifid end of each rib is a minute peg-like stig- matic surface. The Stigma. — This is usually a viscid papillose surface of greater or less expanse functioning for pollen reception. In wind-pollinated flowers such as the grasses, the stigmas are the numerous feathery hairs which cover the ends of the styles and intended to catch flying pollen grains. In animal-pollinated flowers, the stigmas are usually small restricted knobs, lines or depressions. The stigmatic papillae vary in size in different plants and even may vary on different individuals of the same species. Thus in the long styles of Primula, the stigmatic papillae are elongated columnar hair-like structures, whereas in the short styles of short-styled flowers the papillae are small knob-like cellular swellings. 158 PHARMACEUTICAL BOTANY POLLINATION Pollination is the transfer of pollen from anther to stigma and the consequent germination thereon. It is a necessary step to fertilization. When the pollen is transferred to the stigma of its own flower the process is called Close or Self-pollination; if to a stigma of another flower, Cross-pollination. If fertilization follows, these processes are termed respectively, Close or Self-fertilization and Cross-fertilization. Close-fertilization means in time ruination to the race and happily is prevented in many cases by (a) the stamens and pistils standing in extraordinary relation to each other, ( b ) by the anthers and pistils maturing at different times, (c) by the pollen in many cases germin- ating better on the stigma of another flower than its own. The agents which are responsible for cross-pollination are the wind, insects, water currents, small animals, and man. Wind- pollinated flowering plants are called Anemophilous; their pollen is dry and powdery, flowers inconspicuous and inodorous, as in the Pines, Wheat, Walnut, Hop, etc. Insect- pollinated plants are called Entomo philous . These, being dependent upon the visits of insects for fertilization, possess bril- liantly colored corollas, have fragrant odors, and secrete nectar, a sweet liquid very attractive to insects which are adapted to this work through the possession of a pollen-carrying apparatus. Example: Orchids. Plants pollinated through the agency of water currents are known as Hydrophilous. To this class belong such plants as live under water and which produce flowers at or near the surface of the same. Exam- ple: Sparganium. Animal-pollinated plants are called Zoo philous. Some plants like the Nasturtium and Honeysuckle are pollinated by humming birds. Before the pollen grain has been deposited upon the stigma and during its germination thereon, a series of events affecting both the pollen grain and the embryo sac occur which result in the ultimate formation of the male and female gametophytes. Maturation of the Pollen Grain and Formation of the Male Gametophyte. — The substance of the microspore (pollen grain) divides into two cells, the mother and tube cells of the future male PLANT ORGANS AND ORGANISMS 159 gametophyte. The nucleus of the mother-cell divides to form two sperm nuclei. Within the tube cell is found a tube nucleus embedded in protoplasm. Upon germinating the partition disappears and the thin endospore, carrying within it the protoplasm in which are embedded the tube nucleus and twos perm nuclei, penetrates through a deficiency of the exospore. The contents of the pollen grain at this stage is called the male gametophyte. Maturation of the Embryo Sac and Formation of the Female Gametophyte. — The nucleus of the megaspore or embryo sac under- goes division until eight daughter-nuclei are produced which are separated into the following groups: (а) Three of these nuclei occupy a position at the apex, the lower nucleus of the group being the egg or ovum, the other two nuclei being the synergids or assisting nuclei. (б) At the opposite end of the sac are three nuclei known as the antipodals which apparently have no special function. (c) The two remaining nuclei ( polar nuclei) form a group lying near the center of the embryo sac which unite to form a single endosperm nucleus from which, after fertilization, the endosperm or nourishing material is derived. This stage of the embryo sac con- stitutes the female gametophyte. Fertilization in Angiosperms. — After the pollen grain reaches the stigma, the viscid moisture of the stigma excites the outgrowth of the male gametophyte which bursts through the coats of the pollen grain forming a pollen tube. The pollen tube, carrying within its walls two sperm nuclei and one tube nucleus, penetrates through the loose cells of the style until it reaches the micropyle of the ovule, then piercing the nucellus, it enters the embryo sac. The tip of the tube breaks and one of the sperm nuclei unites with the egg to form the oospore. The oospore develops at once into an embryo or plantlet, which lies passive until the seed undergoes germination. The other sperm nucleus unites with the previously fused polar nuclei to form the endosperm nucleus which soon undergoes rapid division into a large number of nuclei that become scattered about through the pro- toplasm of the embryo sac. These accumulate protoplasm about them, cells walls are laid down, endosperm resulting. i6o PHARMACEUTICAL BOTANY THE FRUIT The fruit consists of the matured pistil (carpel or carpels) and contents, or ovarian portion thereof, but may include other organs of the flower which frequently are adnate to and ripen with it. Thus in the Apples, Pears and Quinces, the receptacle becomes thick and succulent, surrounds the carpels during the ripening process and forms the edible portion of these fruits. In Dandelion, Arnica, and many other members of the Composite, the modified calyx or pappus adheres to the ovary during its maturation into the fruit and renders the fruit buoyant. In Gaultheria the calyx becomes fleshy, sur- rounds the ovary, reddens, and forms the edible part of the fruit. In Physalis the calyx enlarges considerably and encloses the ovary in an inflated colored bladder. Involucres frequently persist around and mature with the fruits. These may be membranous as in Anthemis, Matricaria and other Composites, leathery and prickly as in the Chest- nuts, scaly woody cups (cupules) as in the Oaks, or foliaceous cups as in the Filberts. Occasionally, as in the Fig, Osage Orange, Mul- berry, etc., the fruit may consist of the ripened flower cluster or inflorescence. Fruit Structure The Pericarp, or seed vessel, is the ripened wall of the ovary, and in general the structure of the fruit wall resembles that of the ovary, but undergoes numerous modifications in the course of development. The number of cells of the ovary may increase or decrease, the external surface may change fron soft and hairy in the flower to hard, and become covered with sharp, stiff prickles, as in the Datura Stramonium or Jamestown weed. Transformations in consistence may take place and the texture of the wall of the ovary may become hard and bony as in the Filbert, leathery, as the rind of the Orange, or assume the forms seen in the Gourd, Peach, Grape, etc. Frequently the pericarp consists for the most part of other ele- ments than the ripened ovarian wall and is then termed a pseudocarp or anthocarp. The pericarp consists of three layers of different tex- ture, viz.: epicarp, mesocarp and endocarp. The epicarp is the outer layer. The mesocarp the middle, and the endocarp the inner PLANT ORGANS AND ORGANISMS 161 layer. When the mesocarp is fleshy, as in the Peach, it is called the sarcocarp. When the endocarp within the sarcocarp is hard, forming a shell or stone, this is termed a putamen. Sutures.- — The ventral suture is a line formed by the coherent edges of a carpellary leaf. The dorsal suture is the mid-rib of the carpel. Parietal sutures are lines or furrows frequently visible on the walls of fruits, formed by the ripening of a compound ovary. They occur between its dorsal sutures and indicate the places of union between adjacent septa or of two parietal placentas. Valves. — These are the parts into which the mature fruit separates to permit the escape of the seeds. Depending upon the number of these the fruit is said to be univalved, bivalved, trivalved, etc. Dehiscence.- This is the opening of the pericarp to allow the seeds to escape. Fruits are either Dehiscent or Indehiscent according as they open to discharge their seeds spontaneously when ripe (dehiscent), or decay, thus freeing the seeds, or retain their seeds, the embryo piercing the pericarp in germination {indehiscent) . Dehiscent fruits open reg- ularly, or normally, when the pericarp splits vertically through the whole or a part of its length, along sutures or lines of coalescence of contiguous carpels. Legumes usually dehisce by both sutures. Irregular or abnormal dehiscence has no reference to normal sutures, as where it is transverse or circumscissile, extending around the cap- sule either entirely or forming a hinged lid, as in Hyoscyamus, or detached. Dehiscence is called porous or apical when the seeds escape through pores at the apex, as in the Poppy; valvular, when valve-like orifices form in the wall of the capsule. Valvular dehiscence is septicidal, when the constituent carpels of a pericarp become disjoined, and then open along their ventral suture. Example: Colchicum; loculi- cidal, dehiscence into loculi, or cells, in which each component carpel splits down its dorsal suture, and the dissepiments remain intact. Example: Cardamon; septifragal dehiscence , a breaking away of the valves from the septa or partitions. Example: Orchids. Classification of Fruits (according to structure).- — Simple fruits result from the ripening of a single pistil in a flower. 11 162 PHARMACEUTICAL BOTANY Aggregate fruits are the product of all the carpel ripenings in one flower, the cluster of carpels being crowded on the ripened receptacle forming one mass, as in the Raspberry, Blackberry, and Strawberry. Multiple fruits are those which are the product of the ripening of a flower cluster instead of a single flower. Simple and Compound fruits are either Dry or Fleshy. The first may be divided into Dehiscent, those which split open when ripe; and Indehiscent, those which do not. Simple Fruits : [ I. Capsular (dehiscing). Dry j II. Schizocarpic (splitting). I III. Achenial (indehiscent). j IV. Baccate (berries), uccu ent ^ y p> rU p aceous (stone fruits). The capsular fruits include all of those, whether formed of one or more carpels, which burst open to let seeds escape. Schizocarpic or splitting fruits are those in which each carpel or each half carpel (in Labiatse) splits asunder from its neighbor and then falls to the ground. The split portion is one-seeded. Achenial fruits are dry, one-celled, one-seeded and indehiscent at the time of final ripening. Baccate fruits are such in which the endocarp always and the mesocarp usually becomes succulent and so the seeds lie in the pulp formed by the endocarp or endocarp and mesocarp combined. Drupaceous fruits are those in which the endocarp is always fibrous or stony in consistence, while the mesocarp is more or less succulent. The endocarp may become cuticularized as in the Apples. The mesocarp may form stone cells lying in the midst of soft parenchyme cells as in Pears; it may become hardened and thickened by lignin deposits to form fibers as in the Cocoanut, or it may become swollen and soft-succulent as in Peaches, Cherries, etc. I. Capsular Fruits. — These may be simple, when composed of one carpel as the follicle and legume, or compound, when composed of two or more carpels as the capsule, pyxis, regma, siliqua or silicule. The Follicle or pod is a dry, simple capsular fruit formed of a / PLANT ORGANS AND ORGANISMS 1 63 single carpel which dehisces by one suture. This is usually the ventral suture as in Aconite, Staphisagria, Larkspur and some other Ranunculaceee , but may be the dorsal suture as in Magnolia. A Legume is a dry simple capsular fruit formed of a single carpel and dehiscent by both ventral and dorsal sutures. Examples: Peas, Beans, etc. The legume is typical of most Leguminosce. A Capsule is a fruit formed of two or more carpels which dehisce longitudinally or by apical teeth or valves. Examples: Cardamon, Poppy, Iris, etc. A Pyxis or Pyxidium is a capsular fruit formed of two or more carpels that dehisce transversely. Examples: Hyoscyamus, Portu- laca. The upper portion forms a lid which fits upon the lower pot- like portion. Fig. 68. — Legume of the edible pea ( Pisum sativum), a, anther-, c, calyx; st, stigma. (Gager.) A Regma is a capsular fruit of two or more carpels that first splits into separate parts and then each of these dehisces. This type of fruit is typical of II lira crepitans (Sandbox), Pelargonium and Geranium. A Siliqua is a long slender one or two-celled capsule, often with a spurious membranous septum (when two-celled) and two persistent parietal placentae, the valves opening from below upward. Ex- amples: Chelidonium and Wallflower. A Silicule is a short siliqua in which the length is never much greater than the breadth. Example: Cochlearia. II. Schizocarpic Fruits. — A Carcerulus or Nutlet is the typical fruit of the Labiatae but is also seen in the Borraginaceae. The ovary 164 PHARMACEUTICAL BOTANY that has become four-celled at the time of flowering matures into four little pieces which split asunder lengthwise. Each split part is composed of one-half of a ripened carpel. A Cremocarp is the characteristic splitting fruit of the Umbel- liferse family. It consists of two inferior akenes or mericarps sepa- rated from each other by a forked stalk called a carpophore. These mericarps usually cling to the forks of the carpophore for a time after the cremocarp splits, but sooner or later fall. A Samara is a winged splitting fruit. It may be one-carpelled as in the Elm, Ash, Tulip Poplar and Wafer Ash or two-carpelled as in the Maples. A Lomentum or Loment is a splitting fruit that splits transversely into one-seeded portions. Seen in Cruciferce, in Entada scandens, Cathartocarpus Fistula, Desmodium, etc. of Leguminosoe. III. Achenial Fruits (all indehiscent). — The Akene is a dry one- chambered, indehiscent fruit, in which the pericarp is firm and may or may not be united with the seed, the style remaining in many cases as an agent of dissemination. The latter may be long and feathery as in Clematis or be hooked. Examples of akenes: Fruits of the Compositce, Anemone, etc. The Hip of the Roses consists of a number of akenes in a ripened concave receptacle. The Utricle is like the akene, except that the pericarp is loose and bladder-like. Example: Chenopodium. A Caryopsis or Grain is similar to an akene but differs from it by the pericarp being always fused with the seed coat. This fruit is more likely than any other to be mistaken for a seed. Examples: Wheat, Corn, Barley, Oats and other members of the Graminacece. A Nut or Gians is a one-celled, one-seeded fruit with a leathery or stony pericarp. Examples: Oaks, Beeches, Chestnuts, Alders and Palms. IV. Baccate Fruits (Succulent fruits in which the endocarp is always succulent and the mesocarp sometimes).- — The Berry is a small fleshy fruit with a thin membranous epicarp and a succulent interior in which the seeds are imbedded. Examples: Capsicum, Tomato, Belladonna, Grape, Currant, etc. An Uva is a berry from a superior ovary. Examples: Bella- donna, Egg-plant, Tomato. PLANT ORGANS AND ORGANISMS 165 A Bacca is a berry from an inferior ovary. Examples: Goose- berry, Honeysuckle, Currant. The Pepo or Gourd Fruit is a baccate fruit of large size which has developed from an inferior ovary. It is fleshy internally and has a tough or very hard rind. Examples: Fruits of the Cucurbitacece and the Banana. The Hesperidium is a large thick-skinned succulent fruit with seeds embedded in the pulp but from a superior ovary. Examples: Orange, Grape-fruit, Lemon, etc. In each of these there is to be noted a glandular leathery epicarp, a sub-leathery mesocarp and an endocarp in the form of separate carpels. From the endocarp hairs grow inward into the carpellary cavities and become filled with succulence. The seeds lie amid the hair cells. V. Drupaceous Fruits (Succulent fruits in which the mesocarp is more or less succulent, but the endocarp leathery or stony). — A Drupe is a one-celled, one-seeded drupaceous fruit such as the fruit of the Plum, Peach, Prune, Sabal, Rhus, Piper, Cherry, etc., whose endocarp or putamen is composed wholly of stone cells or stone cells and sclerenchyme fibers. The Pome is a fleshy drupaceous fruit, two or more celled with fibrous or stony endocarp, the chief bulk of which consists of the adherent torus. Quince, Apple and Pear are examples. The car- pels constitute the core, and the fleshy part is developed from the torus. Aggregate Fruits. — An Etaerio consists of a collection of little drupes on a torus of a single flower. Examples: Raspberry, Black- berry, etc. Multiple Fruits.- — The Syconium is a multiple fruit consisting of a succulent hollow torus enclosed within which are akene-like bodies, products of many flowers. Example: Fig. The Sorosis is represented by the Mulberry, Osage Orange, etc., the grains of which are not the ovaries of a single flower, as in the Blackberry, but belong to as many separate flowers. In the Pine- apple all the parts are blended into a fleshy, juicy, seedless mass, and the plant is propagated by cuttings. The Strobile or Cone is a scaly, multiple fruit consisting of a scale-bearing axis, each scale enclosing one or more seeds. The i66 PHARMACEUTICAL BOTANY name is applied to the fruit of the Hop, and also to the fruit of the Coniferce in which the naked seeds are borne on the upper surface of the woody scales. A Galbalus is a more or less globular multiple fruit formed of fleshy connate scales, as in Juniper. Histology of a Capsule, Vanilla. — The Vanilla fruit is a one-celled capsule formed by the union of three carpellary leaves and dehiscing by two unequal longitudinal valves. Microscopic Appearance of a Transverse Section. — Passing from periphery toward the center, the following structures present themselves: 1. Epicarp, consisting of epidermis and hypodermis. The epidermis consists of a layer of thick-walled epidermal cells whose outer walls show the presence of a thin yellow cuticle. Stomata are present in this tissue. The epidermal cells contain protoplasm and brownish bodies. Some also contain small prisms of calcium oxalate and a few, vanillin crystals. The hypodermis is composed of one to several layers of collenchymatic cells with dark-colored contents. Its cells are somewhat larger than those of the epidermis and thicker-walled. 2. Mesocarp, a broad region of somewhat loosely arranged large, thin-walled parenchyma cells becoming smaller in the inner zone of this region. Most of these cells contain brownish contents but some possess long raphides of calcium oxalate. If the section be mounted in phloroglucin solution (5 per cent.) and a drop of strong sulphuric acid is added, a carmine-red color will be observed showing the pres- ence of vanillin in this region. Several closed collateral bundles will be seen coursing through the mesocarp. 3. Endocarp, an irregular line of inner epidermal cells which is differentiated into two regions, the interplacental region and the placental region. The interplacental (inner) epidermis shows its cells elongated into numerous thin-walled glandular hairs which con- tain an abundance of balsam; the placental region covers the six bifid placentae which extend into the cavity of the capsule. Its (inner) epidermis is composed of mucilaginous cells. 4. Seeds. -These are minute blackish bodies attached to the pla- cental twigs of the placentae. Some of them may have been torn off in cutting the section. PLANT ORGANS AND ORGANISMS 167 Histology of a Typical Mericarp, Fceniculum. — This five-angled fruit, in transverse section, shows a concave commissural and convex dorsal surface. Passing from the surfaces toward the center we note: 1. Epicarp , or outer covering tissue, composed of colorless epider- mal cells and small stomata. The epidermal cells in cross-section appear rectangular, while in surface view they are both polygonal and rectangular. 2. Mesocarp, of several layers of thin-walled colorless isodia- metric cells, beneath which are two to several additional layers of thicker-walled cells with brownish walls. Through the angles or Fig. 69. — Photomicrograph of a transverse section of a mericarp of Foeni- culum vulgar e, showing epicarp (A), mesocarp (B), endocarp ( F ), vitta (C), endo- sperm of seed (D), carpophore (G) and fibro-vascular bundle in primary rib (£). (Highly magnified). rib portions of the mesocarp extend the fibro-vascular bundles. Between each fibro-vascular bundle and the tip of each rib will be found a zone of collenchyma cells. In the mesocarp between each two ribs on the dorsal side occurs a single vitta or oil tube which is lined with a layer of brownish polygonal cells. These vittas contain the official oil of fennel. Two vittae generally occur in the meso- carp of the commissural side although four are reported to have been found in this region of some fennel fruits. 3. Endocarp , a narrow layer of cells, transversely elongated, except over the regions of bundles where they may be seen elongated in several directions. i68 PHARMACEUTICAL BOTANY 4. Spermoderm or Seed Coat , consisting of a layer of somewhat broadened epidermal cells attached to the endocarp and several layers of collapsed cells which are only well defined in the region of the raphe. 5. Endosperm, a central mass of more or less polygonal cells con- taining aleurone grains and oil globules. Each aleurone grain con- tains a rosette aggregate of calcium oxalate and one or two globoids. 6. Embryo, embedded in the endosperm of the upper region of the seed. THE SEED A seed is a matured megasorus (ovule) borne by the sporophyte of a spermatophytic plant, enclosing a megaspore (embryo sac) within which a fertilized egg of the succeeding gametophyte generation has segmented to form a new sporophyte plant. The purpose of the seed is to insure the continuation and distribution of the species. Like the ovule, it consists of a nucellus or kernel enclosed by integu- ments, and the descriptive terms used are the same. The seed coats, corresponding to those of the ovule, are one or two in number. If but one seed coat is present it is termed the spermoderm. If two are present, the outer one is called the testa, and the inner one the tegmen. The testa, or outer seed shell, differs greatly in form and texture. If thick and hard, it is crustaceous; if smooth and glossy, it is polished; if roughened, it may be pitted, furrowed, hairy, reticu- late, etc. The testa may often present outgrowths or seed appendages whose functions are to make the seeds buoyant, whereby they may be dis- seminated by wind currents. Examples of these are seen in the Milk- weed, which has a tuft of hairs at one end of the seed called a Coma, and in the official Strophanthus, which has a long bristle-like appen- dage attached to one end of the seed and called an awn. The wart- like appendage at the hilum or micropyle, as in Castor Oil Seed, is called the Caruncle. The tegmen or inner coat surrounds the nucellus closely and is generally soft and delicate. A third integument, or accessory seed covering outside of the testa, PLANT ORGANS AND ORGANISMS 1 69 is occasionally present and is called the Aril. Example: Euonymus (succulent). When such an integument arises from the dilatation of the micro- pyle of the seed, as in the Nutmeg, it is known as an Arillode. The Nucellus or Kernel consists of tissue containing albumen, when this substance is present, and the embryo. Albumen is the name given the nutritive matter stored in the seed. The funiculus or seed stalk is usually absent in the official seeds. The scar left by its separation is called the hilum. When the funiculus is continued along the outer seed coat, it is called the raphe. MODE OF FORMATION OF DIFFERENT TYPES OF ALBUMEN If the egg-cell within the embryo sac segments and grows into the embryo and, stretching, fills up the cavity without food material laid down around it, it happens that the nutritive material lingers in the cells of the nucellus pressing around the embryo. This is called Peris per mic albumen. Seen in the Polygonaceoe. In by far the greater number of Angiosperms, the endosperm nu- cleus, after double fertilization, divides and redivides, giving rise to numerous nuclei imbedded in the protoplasm of the embryo sac, out- side of the developing embryo. Gathering protoplasm about them- selves and laying down cell walls they form the endosperm tissue outside of the embryo. Into this tissue food is passed constituting the Endospermic albumen. In the Marantaceoe, Piperacece, etc., nutritive material is passed into the nucellar cells causing them to swell up, while to one side a small patch of endosperm tissue accommodates a moderate amount of nourishing substance, thus resulting in the formation of abundant perisperm and a small reduced endosperm. Exalbuminous seeds are those in which the albumen is stored in the embryo during the growth of the seed. Such seeds show the fleshy embryo taking up all or nearly all the room within the seed coat. Examples: Physostigma, Amygdala, etc. Albuminous seeds are those in which the nourishment is not stored in the embryo until germination takes place. Such seeds show a larger nourishing tissue region and a smaller embryo region. Exam- ples: Nux Vomica, Myristica, Linum, etc. 170 PHARMACEUTICAL BOTANY Gross Structure of a Monocotyl Seed {With fruit wall attached), Indian Corn. — The ripened seed of Indian Corn is surrounded by a thin, tough pericarp which is firmly adherent to and inseparable from the Spermoderm or seed coat. Because of this fact, while in reality a fruit called a caryopsis or grain, this structure is sometimes erro- neously termed a seed. The mature grain of most varieties of Indian Corn is flattened and somewhat triangular in outline, the summit being broad and the base comparatively narrow. The summit is indented and often marked by a small point which represents a vestige of the style. The basal or “tip” region marks the part of the grain which was in- serted into the cob. Upon it may be found papery chaff, represent- ing parts of the pistillate spikelets. The groove noted on the broader surface indicates the position of the embryo. Histology of the Indian Corn Seed {With fruit wall attached). — If a longitudinal section be cut through the lesser diameter of a soaked grain, the following histologic characteristics will be observed: 1. The Pericarp or ripened ovarian wall which, alike with all other grains, adheres firmly to the wall of the seed forming a portion of the skin of the grain. The pericarp comprises an outer epicarp of elongated cells with thin cuticle, a mesocarp of thicker walled cells without, becoming thinner within and a layer of tube cells. 2. The Spermoderm or seed coat, a single layer of delicate elon- gated cells. 3. The Peris perm, another layer directly underneath the Spermo- derm, difficult to distinguish without special treatment, and repre- senting the ripened nucellar tissue of the ovule. 4. The Endosperm or nourishing tissue, consisting of: {a) The Aleurone Layer, for the most part a single row of cells, containing aleurone grains. Some of the cells may be seen to be divided by tangential partitions, {b) Starch Parenchyma, consisting of two regions: an outer horny zone composed of cells containing for the most part polygonal starch grains and oil droplets; and an inner mealy zone of cells with mostly rounded starch grains. 5. The Embryo, consisting of a single shield-shaped cotyledon adjoining the endosperm, the plumule or rudimentary bud at the end of the caulicle or rudimentary stem and the radicle or rudimen- PLANT ORGANS AND ORGANISMS I 7 I tary root, with its tip covered by a root cap. Continuous with the root cap is a root sheath or coleorhiza. The cotyledon or seed leaf consists of two parts: the scutellum which lies next to the endosperm, and is an organ of absorption ; and the sheathing portion which sur- rounds and protects the rest of the embryo. The embryo contains oil and proteids, but no starch. If a similar longitudinal section of a soaked grain be mounted in dilute iodine solution, the contents of the aleurone cells will be col- ored yellow indicating their proteid nature, while the starch grains will take on a blue to violet coloration. The endosperm will be observed taking up most of the room within the seed coat. The con- tents of its cells are not baled out to the embryo until after germina- tion begins. Indian Corn is therefore an albuminous seed. A MONOCOTYL SEEDLING Germination. — When any viable seed is planted in suitable soil, and furnished with oxygen and water and a certain degree of heat, ger- mination takes place. In the presence of moisture, etc., the seed swells, the ferments present within the cells of the endosperm then change the insoluble proteid, starch, and oil to soluble materials, which, in the case of Indian Corn, are absorbed in solution by the scutellum which bales this nourishment out to other parts of the growing embryo, there to be used in part in constructing new tissues, and in part to be consumed by oxidation or respiration. The process of respiration or breathing takes place when the plant takes in oxy- gen and gives off carbon dioxide. The oxygen oxidizes the tissues with an accompanying release of energy, which latter is necessary to life and growth. The combined pericarp and spermoderm bursts opposite the tip of the radicle, and the radicle, piercing through the cotyledonary sheath, protrudes. The cleft in the coat lengthens to the point opposite the tip of the plumule, which also protrudes after bursting through the cotyledonary sheath. The radicle, next, grows downward into the soil forming the primary root, and develops upon itself secondary or lateral roots, all of which give rise to root-hairs just above their root caps. Additional lateral roots emerge above the scutellar region 172 PHARMACEUTICAL BOTANY which ere long attain the size of the first or primary root. The cau- licle, carrying upon its tip the plumule, elongates and forms the stem; the leaves of the plumule spread out and turn green to function as foliage leaves. The perforated cotyledonary sheath grows out sur- rounding both the root and the stem for a portion of their length. By this time all or nearly all of the nourishment stored in the endosperm has been absorbed and assimilated by the young seedling and the coat and scutellum, left behind, gradually decay and disappear. The root-hairs absorb nourishment from the soil, the green leaves build up carbohydrates, prop-roots make their appearance at the first node (joint) above ground, and the seedling grows larger. Gross Structure of a Dicotyl Seed, Phaseolus lunatus (Lima Bean). — The Lima Bean Seed shows a flattened-ovate to somewhat reni- form outline. Externally it exhibits a polished seed coat which is perforated on its basal side by a minute pore called the micropyle or foramen. Just below this pore will be noted the hilum or scar which represents the point of detachment from the funiculus or stalk which connected the seed during its growth with the wall of the fruit. Upon soaking the seed in water, it is possible to remove the seed coat or spermoderm. This done, the embryo will be exposed. The two fleshy cotyledons are first seen. Upon spreading these out, convex sides down, the rest of the embryo, consisting of a thin leafy structure surrounding a bud and called the plumule, the caulicle or rudimen- tary stem and in line with the latter, the radicle, or rudimentary root, will be seen. Histology of the Lima Bean Seed. — In transverse sections, the following microscopic structures will be evident: i. Spermoderm of three regions, viz.: Palisade cells, Column cells, and Spongy Parenchyma. The palisade cell layer is composed of vertically elongated thick-walled cells which are covered on their outer faces by a clear glistening cuticle. The cells are 60 to 8oyu long and 1 2 to 2o/x wide. The column cells, found forming a layer directly beneath the palisade zone, are hour-glass-shaped and 25 to 35^ long by 14 to 35 ju wide. The spongy parenchyma forms a zone of several layers of thin- walled parenchyma cells, the cells of the outer and inner layers being considerably smaller than the middle layers. PLANT ORGANS AND ORGANISMS 17 3 2. Embryo, the two cotyledons of which make up the greatest bulk. These are composed of an epidermis covering over a region of meso- phyll. The mesophyll is constituted of moderately thick- walled cells which contain ellipsoidal and kidney-shaped starch grains up to 65/u in length. A conspicuous branching cleft will be seen in the larger grains. In the Lima Bean, the nourishment is stored in the embryo during the growth of the seed. It is, therefore, exalbuminous . CHAPTER VIII TAXONOMY DIVISION I.— THALLOPHYTA Plants, the greater number of which, consist of a thallus, a body undifferentiated into root, stem or leaf. The group nearest to the beginning of the plant kingdom presenting forms showing rudi- mentary structures which are modified through division of labor, dif- ferentiation, etc., in higher groups. SUBDIVISION I.— PROTOPHYTA (SCHIZOPHYTA) A large assemblage of “fission plants” comprising the bacteria and blue-green algte. In the simplest types no nucleus is present, but as we arise in scale through the bacteria and blue-green algae, there is to be observed an open granular, gradually growing to a crescentic, chromatin mass that may be called a nucleus. A common method of asexual reproduction is possessed by these plants whereby the cell cleaves or splits into two parts, each of which then becomes a separate and independent organism. I. SCHIZOMYCETES — BACTERIA Bacteria are minute, unicellular, colorless, rarely weakly red or green colored, non-nucleate vegetable organisms destitute of chloro- phyll. They serve as agents of decay and fermentation and are frequently employed in industrial processes. According to the vari- ous phenomena they produce, they may be classified as follows: (a) Zymogens producing fermentation; ( b ) Aerogens producing gas; (c) . Photogens producing light; (d) Chromogens producing color; ( e ) Saprogens, producing putrefaction; (/) Pathogens, producing disease. Physical Appearance of Bacterial Colonies and Individual Forms. Because of their minute size — a space the size of a pinhead may 174 TAXONOMY 175 hold eight billion of them — the student commences his study of bacterial growths in colonies or cultures, each kind possessing characteristics by which they may be distinguished and differ- entiated. Tihe individuals in the colony, depending upon the kind of bac- teria under examination, may be globular, rod-shaped, or spiral. Bacteria are classed according to form into the following families and genera. Fig. 70. — Types of micrococci. ( After Williams .) Family I. — Coccaceae. — Cells in their free condition globular, be- coming but slightly elongated before division. Cell-division in one, two or three directions of space. A. Cells without Flagella. 1. Division only in one direction of space forming an aggregation resembling a chain of beads — Streptococcus. 2. Division in two directions of space forming an aggregation resembling a cluster of grapes — Staphylococcus. 3. Division in three directions of space forming a package-shaped or cubical aggregation — Sarcina. B. Cells with Flagella. 1. Division in two directions of space — Planococcus. 2. Division in three directions of space — Planosarcina. Fig. 71. — Types of bacilli. ( After Williams.) Family II.— Bacteriaceae. — Cells longer than broad, generally two to six times, straight or only with an angular bend, never curved or 176 PHARMACEUTICAL BOTANY spiral, division only at right angles to axis or rod; with or without flagella and endospores. 1. Flagella and endospores absent — Bacterium. 2. Flagella and endospores present — Bacillus. Family III. — Spirillaceae. — Cells curved or spirally bent, generally motile through polar flagella. 1. Cells stiff, not flexile. (a) Cells without flagella — Spirosoma. ( b ) Cells with one, very rarely with two polar flagella — Micro- spira. ( c ) Cells with a bundle of polar flagella — Spirillum. 2. Cells flexile, spiral very close — Spirochaeta. Fig. 72. — Types of spirilla. ( After Williams .) Family IV. — Mycobacteriaceae.- — Cells short or long cylindrical or clavate-cuneate in form, without a sheath surrounding the chains of individuals, without endospores, with true dichotomous branching. A. In cultures possessing the characters of true bacteria. Growth on solid media smooth, flat, spreading. Rods with swollen ends, or cuneate or clavate forms — Corynebacterium. B. Cultures on solid media raised, folded or warty. Generally short slender rods, rarely short branched. Take the tubercle stain— Mycobacterium. Family V. — Chlamydobacteriacese. — Thread-like, composed of individual cells, surrounded by a sheath. Simple or with true branching. Ordinary vegetative growth by division in only one direction of space, i.e., at right angles to the longer axis. A. Cell contents without sulphur granules. 1. Filaments unbranched. (1 a ) Cell-division only in one direction of space. TAXONOMY 177 ( b ) Cell-division in gonidial formation in three directions of space — Streptothrix. *Marine forms with cells surrounded by a very delicate hardly discernible sheath — Phragmidiothrix. **Fresh-water forms with easily discernible sheath — - Crenothrix. 2. Filaments branched. B. Cell contents with sulphur granules — Thiothrix. Family VI. — ^Beggiatoaceae. — Thread-like, without a capsule, but with an undulating membrane. Cell contents show sulphur granules. A. Threads apparently not septated, septa only faintly visible with iodine staining. Colorless or faintly rose-colored — Beggiatoa. Sporulation. — A large number of bacteria possess the power of developing into a resting stage by a process known as sporulation or spore formation. Sporulation is regarded as a method of resisting unfavorable environment. This is illustrated by the anthrax bacilli which are readily killed in twenty minutes by a 10 per cent, solution of carbolic acid, and able, when in the spore condition, to resist the same disinfectant for a long period in a concentration of 50 per cent. And, while the vegetative forms show little more resistance against moist heat than the vegetative form of other bacteria, the spores will withstand the action of live steam for as long as ten to twelve minutes or more. Whenever the spores are brought into favorable condition for bacterial growth, as to temperature, moisture and nutrition, they return to the vegetative form and then are capable of multiplication by fission in the ordinary way. Reproduction.- — Bacteria multiply and reproduce themselves by cleavage or fission. A young individual increases in size up to the limits of the adult form, when by simple cleavage at right angles to the long axis, the cell divides into two individuals. Morphology Due to Cleavage. — According to limitations imposed by cleavage directors, the cocci assume a chain appearance, or a grape-like appearance, or an arrangement in packets or cubes having three diameters. This gives rise to the Staphylococcus (plural, staphylococci), from a Greek word referring to the shape of a bunch of grapes. 12 i7» PHARMACEUTICAL BOTANY Streptococcus (plural, streptococci), from a Greek word meaning chain-shaped. Sarcina, package-shaped or cubical. Form of Cell Groups after Cleavage. — The individual bacteria after cleavage may separate, or cohere. The amount of cohesion, together with the plane of cleavage, determines the various forms of the cell groups. Thus, among the cocci, diplo- or double forms may result giving rise to distinguishing morphological character- istics. Similarly among the bacilli characteristic forms result as single individuals and others which form chains of various lengths. Rapidity of Growth and Multiplication. — The rapidity with which bacteria grow and multiply is dependent upon species and environ- ment. The rapidity of the growth is surprising. Under favorable conditions they may elongate and divide every twenty or thirty minutes. If they should continue to reproduce at this rate for twenty-four hours a single individual would have 17 million de- scendants. If each^of these should continue to grow at the same rate, each would have in twenty-four hours more, 17 million offspring, and then the numbers would develop beyond conception. How- ever, such multiplication is not possible under natural or even artificial conditions, both on account of lack of nutritive material and because of the inhibition of the growth of the bacteria by their own products. If they did multiply at this rate in a few days there would be no room in the world but bacteria. Chemical Composition of Bacteria. — The quantitative chemical composition of bacteria is subject to wide variations, dependent upon the nutritive materials furnished them. About 80 to 85 per cent, of the bacterial body is water; proteid substances constitute about 50 to 80 per cent, of the dry residue. When these are extracted, there remain fats, in some cases wax, in some bacteria traces of cellulose appear, and the remainder consists of 1 to 2 per cent. ash. The proteids consist partly of nucleo-proteids, globulins, and protein substances differing materially from ordinary proteids. Toxic substances known as endotoxins to distinguish them from bacterial poisons secreted by certain bacteria during the process of growth, also occur. TAXONOMY 179 Some Bacteria Producing Diseases in Man or the Lower Animals Organism Disease Staphylococcus pyogenes aureus Boils, abscesses, carbuncles Streptococcus erysipelatis Erysipelas Micrococcus meningitidis Cerebrospinal meningitis Micrococcus gonorrhoea; Gonorrhoea Micrococcus melitensis Malta fever Micrococcus catarrhalis Catarrh Bacillus anthracis Anthrax Bacterium diphtheria; Diphtheria Bacillus typhosus Typhoid fever Bacterium influenzae Influenza Bacillus tetani Tetanus Bacillus leprae Leprosy Bacillus chauvei “Blackleg” of cattle Bacillus aerogenes capsulatus Emphysematous gangrene Bacterium tuberculosis Tuberculosis Bacterium mallei Glanders Streptococcus pneumoniae (Diplococcus pneumoniae) Pneumonia (croupous or . fibrinous pneumonia) Spirillum cholerae asiaticae Cholera Spirillum obermeieri Relapsing fever Strep tothrix (Actinomyces) bovis Actinomycosis in cattle Some Bacteria Producing Diseases in Plants Actinomyces Myricarum Tubercles upon and lesions within Myrica and Comp- tonia Bacterium tumefaciens Crown gall Bacterium savastanoi Olive knot Bacillus amylovorus Pear blight Pseudomonas juglandis Walnut blight Bacillus Solanacearum Wilt of Solanaceae Bacillus tracheiphilus Wilt of Cucurbits Pseudomonas Stewarti Wilt of Sweet Corn Mounting and Staining of Bacteria. — The mounting and staining of bacteria may be accomplished as follows : 1. Take the square or round cover slip, which has been previously cleaned, out of the alcohol pot. Dry it between filter paper. 2. Hold it in the bacteriolgic forceps, which is so constructed that a spring holds the cover slip firmly while an enlargement of the wire i8o PHARMACEUTICAL BOTANY handle permits the placing of the forceps on the table while the cul- ture material is obtained. 3. Place several drops of distilled water on the cover slip and add a loop full of the organism secured from the pure culture in a test tube as follows: 4. Remove the cotton plug by the third and fourth fingers of the left hand. 5. Hold the open test tube between the thumb and forefinger of the left hand. 6. By means of a previously flamed platinum needle, remove a little of the culture from the surface of the culture media. 7. Replace the cotton plug. 8. Add the culture media to the drop of distilled water on the cover slip and distribute this material by stirring. 9. Evaporate the water on the cover slip to dryness by holding it some distance above the Bunsen flame and slowly enough to prevent connection circles being formed by the material affixed to the cover. 10. Pass the cover glass three times through the Bunsen flame. 11. Apply the stain, which should remain long enough to stain the objects. 12. Wash off the stain with distilled water. 13. Dry the cover glass above the flame. 14. Apply a drop of balsam, turn the cover slip over and drop it on to the center of a glass slide previously provided and cleaned for this purpose. Gram’s Method. — This is a method of differential bleaching after a stain. The cover glass preparations or sections are passed from absolute alcohol into Ehrlich’s anilin gentian violet, where they remain one to three minutes, except tubercle bacilli preparations which remain commonly twelve to twenty-four hours. They are then placed for one to three minutes (occasionally five minutes) in iodine potassium iodide water (iodine crystals 1, potass, iodide 2, water 300), with or without washing lightly in alcohol. In this they remain one to three minutes. They are then placed in absolute alcohol until sufficiently bleached, after which they are cleared in clove oil and mounted in balsam. Certain organisms, when stained by this method give up the stain TAXONOMY 181 and are called “Gram negative;” others retain the color and are called “Gram positive.” Examples of the latter are B. diphtheriae, Bacillus anthracis, and Bacillus tetani. Stains. — One of the most useful bacteriologic stains is Ziehl's Carbol Fuchsin, prepared as follows: Fuchsin (basic), i. Absolute Alcohol, i. Carbolic Acid (5 per cent, aqueous solution), 100. The fuchsin should be dissolved first in the alcohol and then the two fluids mixed. Ehrlich’s Anilin Water Gentian Violet. — Anilin Oil Water, 75 parts. Sat. Sol. Gentian Violet in Alcohol, 25 parts. Anilin oil water is made by adding 2 mils anilin to 98 mils distilled water; shake violently. Filter through filter paper several times. Loffler’s Methylene-blue. — Sat. sol. Methylene-blue in Alcohol 30 mils Sol. KOH in distilled water (1:10,000) 100 mils Mix the solutions. Stain for “Acid Proof” Bacteria Including B. Tuberculosis. — 1. Flood the cover glass with Ziehl’s carbol fuchsin and boil over the flame for thirty seconds. 2. Wash and decolorize with a 2 per cent, solution of HC1 in 80 to 95 per cent, alcohol until the thinner portions of the film show no red color. 3. Wash in water. 4. Counter stain for contrast with Lofiler’s Methylene-blue. 5. Wash and examine. Van Ermengem’s Flagella Stain. — 1. Mordant: Osmic acid (2 per cent, aqueous solution) 50 Tannin (10 to 25 per cent, in water) 100 Four drops of glacial acetic acid may be added to this. 2. Silver Bath: Dissolve 0.25 to 0.5 per cent, nitrate of silver in distilled water in a clean bottle. 182 PHARMACEUTICAL BOTANY 3. Reducing and Strengthening Bath: Gallic acid 5 Tannin 3 Fused sodium acetate 10 Distilled water 350 The flamed cover glass is first covered with the mordant for one- half hour, or if in a thermostat at 5o°C. for five to ten minutes. The mordant is then carefully removed by thorough washing in water, alcohol and water. The cover (film side up) is now put into the silver bath (a few mils in a clean beaker or watch glass) for a few seconds, during which time it is gently agitated. Without rinsing it is next put into a few mils of the reducing solution and gently agitated until the fluid begins to blacken. It is then washed in water and examined. If not stained deeply enough the cover is returned to the silver bath. It is finally dried and mounted in balsam. All the dishes must be scrupulously clean. The fluids must not be contaminated by the fingers nor by dipping iron or steel instruments into them. 2. Cyanophyce^; Plants which are sometimes termed blue-green alga. They con- tain chlorophyll, a green pigment and phycocyanin, a blue pigment, a combination giving a blue-green aspect to the plants of this group. Found everywhere in fresh and salt water and also on damp logs, rocks, bark of trees, stonewalls, etc. Examples: Oscillatoria, Glce- ocapsa, and Nostoc. Gloeocapsa. — This blue-green alga is commonly found on old, damp flower pots in greenhouses and on damp rocks and walls near springs, where it forms slimy masses. Under the micro- scope a mount of Gloeocapsa will be seen to consist of isolated pro- toplasts and groups of protoplasts, surrounded by concentric gela- tinous envelopes. Each protoplast consists of a protoplasmic mass which contains blue and green pigments. No definitely organized nucleus is apparent but chromatin in the form of granules is scattered through the protoplasm. The whole is surrounded by a TAXONOMY 183 cell wall which undergoes mucilaginous modification producing thus the soft gelatinous envelopes which encircle parent-, daughter-, grand- daughter- and even great-granddaughter-cells. Fig. 73. — A, B, C, D, E, Gloeocapsa\ F, Oscillaloria showing a dead cell (d) which marks a place of separation into segments. (A), Gloeocapsa, parent cell composed of central protoplast containing scattered chromatin granules, sur- rounded by cell wall and 3 mucliaginous envelopes; ( B ), parent cell is shown elongated, the protoplast in process of division to form two daughter protoplasts; (C), daughter protoplasts, each surrounded by two gelatinous envelopes and both within the original parent envelopes; (D) the daughter protoplasts shown in C have just divided to form granddaughter protoplasts which have later separated, each forming envelopes of its own but all four encircled by the parent envelope. Oscillatoria. — Oscillatoria is a blue-green filamentous organism found abundantly on the surface of the mud of drains and ditches as well as in ponds where the water is foul. The filament is slender 184 PHARMACEUTICAL BOTANY and composed of compactly arranged disc-shaped cells which are all alike, excepting the terminal ones which appear rounded off. The filaments tend to be agglomerated in thick felts or gelatinous masses and each possesses peculiar oscil- lating and forward movements. At the time of reproduction the filament breaks up transversely into short segments, each of which, by fission occurring among its cells, grows into a new filament. Nostoc. — Nos toe occurs on the damp ground bordering streams or in slow bodies of water as greenish or brownish tough gelatinous masses varying in size from a pea to a hen’s egg. When one of these masses is dissected and examined microscopically, it is seen to contain, im- bedded in a gelatinous matrix, numerous serpentine filaments, composed of spherical or elliptical cells loosely attached to each other in chain-like fashion. Most of the cells are of the blue-green vegetative kind but there occur at intervals larger cells, often devoid of protoplasm which are termed heterocysts. Frequently the fila- ments break apart on either side of the heterocyst, setting free segments of cells which grow into new filaments. SUBDIVISION II— MYXOMYCETES, OR SLIME MOLDS Terrestrial or aquatic organisms, fre- quently classified as belonging to the animal kingdom and found commonly on decaying wood, leaves, or humous soil in forests. Their vegetative body consists of a naked, multinucleated mass of protoplasm called the plasmodium, which has a creeping and rolling amoeboid motion, putting out and retracting regions Fig. 74. — Nostoc. heterocyst. (h). a TAXONOMY 185 of its body called pseudo podia. The size of the plasmodium varies from a ten-cent piece to several square feet of surface. It is net-like, the net being of irregular dimensions. Like the amoeba the outer portion of the plasmodium is clear and watery and known as the ectoplasm, the inner portion is granular and called the endoplasm. Like the amoeba and unlike other plants, this slimy body engulfs solid food by means of its pseudopodia instead of admitting it in solution. It is extremely sensitive to light being negatively heliotropic, i.e., turning away from the sun’s rays. Fig. 75. — A, B, Comalricha nigra. A, Sporangium, natural size; B. capilli- tium, 20/1; C, E, Stemonitis fusca; C, sporangium, natural size; D and E, capilli- tia, 5/1, 20/1; F, H, Enerthema papillalum, F, unripe; G, mature sporangium, 1 o / r d Ii, capillitium, 20/1. (C, D. after nature. A. F, G, H. after Roslafinski; B, E, after de Bury in Die naturlichen Pflanzenfamilien I. 1, p. 26.) At the time of reproduction, the plasmodium creeps to the surface. The whole plasmodium then forms one or more fructifications. These fructifications vary from cushion-like masses ( celhallia ) to more elevated bodies in which the net-like structure of the plas- modium is preserved ( plasmodiocar ps ) to stalked sporangia (spore cases). All of the fructifications, however, produce spores. Dur- ing wet weather amoeboid protoplasts ( swarm spores ) escape from the spores, each developing a single cilium and moving actively about. In time the cilia disappear and these swarm spores coalesce in smaller then larger groups to form a plasmodium. i86 PHARMACEUTICAL BOTANY SUBDIVISION III. — ALG/E Low forms of thallophytes of terrestrial and aquatic distribution consisting for the most part of single cells or rows of single cells joined end to end to form filaments. The higher forms, however, possess structures, which might be compared to stems and leaves of higher plants although more rudimentary in structure. They contain chlorophyll or some other pigment, and so can use the CO2 and H 2 0 in the same manner as higher plants, e.g., in assimilating and providing for their own nutrition. Archegonia are absent in this group. Class I. — Chlorophyce^e, the Green Alg,e A large assemblage of terrestrial or aquatic plants many of which are vastly different from one another. In this group the cells are observed to possess distinct nuclei and chromatophores, the latter containing a green oil-like pigment called chlorophyll. Reproduc- tion is mostly by ciliated cells called zoospores but in some forms is accomplished by the fertilization of egg-cells and in others by simple division. 1. Order Protococcales or One-celled Green Algae. — This order contains nearly all of the one-celled green algae excepting the diatoms and desmids. Family Pleurococcaceae. — Pleurococcus vulgaris is a one-celled green alga, millions of which, living together in colonial fashion, con- stitute the so-called “green stain” that is common on the north sides of tree trunks, stone walls and fences. Each organism con- sists of a protoplast surrounded by a cell wall of cellulose. The protoplast contains a chromatophore, cytoplasm and nucleus. Reproduction takes place by the protoplast dividing into two equal parts and laying down a cell wall forming two daughter-protoplasts. These may again divide to form four granddaughter-protoplasts. Still another division may occur as a result of which eight great- granddaughter-protoplasts are formed which frequently adhere to one another forming colonies. 2. Order Confervales. — In this order are included a variety of green filamentous and membranous forms some of which show sexual reproduction. TAXONOMY l8 7 Family Ulothricaceae. — Ulothrix zonata, a typical representative of this family, is a filamentous organism found growing on stones around ponds, on rocks along the shores of lakes, in slow-moving streams, etc. Each filament is unbranched and consists of a row of short cells, one of the terminal cells, called the rhizoid cell, being elongated and serving as an attachment structure. Each cell con- sists of a cell wall of cellulose enclosing cytoplasm, a nucleus and a wide band-shaped green chromatophore, more or less cylindrical in shape. The chromatophore lies next to the cell wall and contains pyrenoids or starch-forming centers. The filament grows in length by the fission of its various component cells. After attaining a Fig. 76. — Vaucheria lerreslris. anth, antheridium (empty) ; o, oogonia. (Gager.) certain size it reproduces either asexually or sexually. Asexual reproduction takes place by certain cells becoming altered in their protoplasmic contents through division to form rounded or pear- shaped zoospores. Each zoospore contains a red pigment spot and bears four cilia (protoplasmic outgrowths). The zoospores escape into the water by lateral openings in the walls of cells containing them. They swim rapidly about, propelled by their cilia, and ere long attach themselves to various objects and grow into Ulothrix filaments. The sexual method of reproduction is effected through the production of many gametes, in cells of the filament, which re- semble the zoospores in shape but differ from them in being smaller and possessing but two cilia. These escape into the water, and, after swimming about for a short time come together in pairs and PHARMACEUTICAL BOTANY 1 88 fuse with one another. The product of the fusion of each pair of these like gametes is termed a zygospore. The zygospore swims about but finally comes to rest, remaining quiescent for a considerable length of time. It then enlarges and its protoplasmic content divides to form several zoospores which, escaping from the cell, swim about for a while and finally, attaching themselves to objects, grow into filamentous Ulothrix organisms. 3. Order Conjugales. — To this order belong the desmids and pond scums which are distinguished from other green algae by presenting no motile stages in their life histories. They are all of fresh-water habit and reproduce by conjugation. Family Desmidaceae. — The desmid family includes a number of genera of unicellular as well as filamentous green plants that present a variety of shapes. Each unicellular desmid is characterized by being composed of two like halves frequently separated by each other by a constriction called the isthmus. In each half there is a chromatophore containing pyrenoids. The nucleus is found in the isthmus. Reproduction is accomplished either asexually by fission or sexually by conjugation. Family Zygnemaceae. — This is a family of pond scums including the well-known genera, Spirogyra and Zygnema. Spirogyra or Brooksilk is a filamentous organism found suspended or floating in masses in quiet water. Each filament when examined microscopically will be found to consist of more or less elongated cylindrical cells arranged end to end, the terminal cells having rounded extremities. Each cell has a cell wall of cellulose within which is to be found a thin film of ectoplasm. One or more spirally shaped chromalophores will be seen directly within this area. Each chromatophore contains chlorophyll and a number of pyrenoids. In the center of the cell the nucleus is found. Fine strands of protoplasm hold it in place and run out to the ectoplasm. Under favorable circumstances the cells of Spirogyra increase rather rapidly in length. Abnormally long cells are not seen, however, because the elongating cells speedily divide, forming two daughter-cells. Under the best of conditions, division may occur every night. In this way the filaments are rapidly made longer. Sooner or later they break and in this way the plant multiplies. TAXONOMY 189 Spirogyra has also a process of sexual reproduction known as conjugation. This process occurs normally from March to June and July, but can be induced in the laboratory by allowing the water Fig. 77. — Spirogyra sp. A, terminal portion of vegetative filament; B, stages of scalariform conjucation; C, preparation for lateral conjugation; D, zygospores formed by lateral conjugation. (Gager.) in the vessel in which it is growing to slowly evaporate. Two fila- ments arrange themselves side by side, and the cells lying opposite each other undergo internal changes so as to form gametes or sexual 190 PHARMACEUTICAL BOTANY cells. Each protrudes a process or conjugation tube; these unite and the protoplasm from one cell passes over and coalesces with that in the cell opposite. The result of the process is a new cell called a zygospore or zygote. This is set free by decay of the walls of the old cell and falls to the bottom of the water, there to undergo a resting stage until favorable conditions for growth arise. 4. Order Diatomales.- — Family Diatomaceae. — This family com- prises several thousand species of unicellular plants called Diatoms which are found in fresh, brackish and salt water forming much of the diet of small animals. While unicellular they frequently are united in colonies. They all possess chromatophores containing chloro- phyll but this green pigment is often obscured by the presence also of a brown pigment. The most striking peculiarity of the group is the structure of the enclosing cell wall. This is in the form of a siliceous case consisting of two valves which fit into each other like the halves of a pill box. The valves, which are beautifully sculptured, are similar except that one is slightly larger than the other so as to fit over it. Diatoms vary in form being either circular, linear, elliptical, cylindrical rhomboidal, triangular or fan-shaped, etc. Some are borne on the ends of stalks, while others are held in gelatinous masses. Their siliceous skeleta are deposited constantly on the floor of ponds, rivers, lakes and seas, often in such abundance as to form Diatomaceous earths or Kieselguhrs (Siliceous Earths). Huge geological deposits of this material have been found in various parts of the world. The most remarkable for extent as well as for the number and beauty of the species contained in it is that at Richmond, Virginia. It is in many places 25 to 40 feet in depth and extends for many miles. Many of the diatomaceous earths are useful as absorbent and polish- ing powders. The United States Pharmacopoeia IX recognizes, under the name of Terra Silicea Purificata (Purified Siliceous Earth), a powder consisting of the frustules and fragments of diatoms which has been purified by boiling with diluted hydrochloric acid, washed and calcined. Diatoms exhibit two modes of reproduction, viz., fission and for- mation of an auxospore. The more common method is that of fission but this is peculiar for these plants. The cell-contents within TAXONOMY 191 the siliceous case separate into two distinct masses and the valves separate slightly from each other. As the two daughter-masses become more and more developed, the valves of the parent-cell are pushed more widely apart. Each of the two masses secretes for itself a new valve on the side opposite to the original valve. When the process is completed the girdle of the parent-diatom separates and the two daughter-diatoms thus become independent plants. Each of these possesses one of the parent valves and a second, which it has formed itself more or less parallel to the first. In a number of species, repeated fission results in the formation of succeedingly smaller and weaker individuals. This process, however, goes on only for a certain number of generations until the decrease of size has reached a limit for the species, when the plant is rejuvenated by the formation of an auxospore. This may be formed with or without the conjugation of two parent-protoplasts. In either case the auxospore resulting undergoes a resting stage after which it develops new valves. The newly formed diatom is then several times the size of the individual or individuals which con- tributed to its formation and is endowed with renewed vigor for growth and division. 5. Order Siphonales {Siphon Alga). — This group is characterized by the peculiarity that the organisms constituting it possess proto- plasm containing myriads of nuclei within a common filament or cell cavity not segmented by cell walls. The term canocyte has been given to such structures which consist of a many-nucleated mass of protoplasm surrounded by a cell wall. Some of the siphon algae reproduce by zoospore formation, others by conjugation as well as zoospore formation while Vancheria, the green felt, stands out alone in reproducing both by the formation of a single zoospore and by the production also of oogonia and antheridia with resultant fertilization. 6. Order Charales {The Stoneworts). — Family Characeae.— The highest group of algae, possessing forms which are differentiated into stems, leaves and rhizoids. Chara, a type of this family, is a submerged fresh-water plant which fastens itself to the muddy bottom of ponds, ditches and slow streams by means of slender filaments called rhizoids. From these 192 PHARMACEUTICAL BOTANY there arises a many noded (jointed) stem which bears whorls of slender green leaves at its nodes. Branches are also found issuing from some of the nodes which duplicate in appearance the main stem. Reproduction is either asexual or sexual. Asexual repro- duction is accomplished by means of tuber-like bodies borne on submerged parts or by special branches which form rhizoids on their lower nodes and later become separated from the parent plant. Sexual reproduction is effected through the formation of oogonia (female sex organs) and antheridia (male sex organs). These in some species are borne on the same plant; in others, on different plants. In all cases the sexual organs are produced at the nodes. The oogonium develops within itself a large ovum or egg. The antheridium produces within its wall numerous motile sperms. Upon the maturation of the antheridium the sperms are liberated into the water, and, propelled by their cilia, find their way to the oogonia which they enter, the one best adapted fusing with the egg in each case and fertilizing it. The resultant cell is called the oospore. This undergoes a resting stage and later germinates as a proembryo. The proembryo consists of a simple filament and a long rhizoidal cell. From this proembryo, the adult stem arises as a side branch. Class II. — Phaeophyce^e, tiie Brown Alg^: Mostly marine forms showing great diversity in the form of their vegetative bodies. They occur for the most part in salt water be- tween the high and low tide marks. Their bodies are usually fixed to some support in the water by means of a holdfast, and are often highly differentiated both as to form and tissues. Some reach hundreds of feet in length as, for example, Macrocyslis which grows in the Pacific Ocean off the coast of California. They all contain the brown pigment called phycophcein and the green pigment, chloro- phyll both of which are present in their chromatophores. A yel- lowish pigment called phycoxanthin has also been isolated from some of the species. Many of the kelps and rockweeds belonging to this class have long been sources of iodine, potash and sodium. A Filamentous Brown Alga, Ectocarpus Siliculosus. — Ecto- corpus occurs as tufts of branching filaments, each of which TAXONOMY I 93 is many-celled. These tufts are found on eelgrass or other algae as well as attached to pilings of wharves in salt water. It is a striking illustration of the simplest form of brown algae and serves to show the beginning of a more complex form of repro- duction than that observed in the forms studied up to this time. On examination of a filament we find it to consist of many cells joined end to end. A single cell has a cell wall of cellulose, just within the cell wall there is a layer of protoplasm. Going toward the center we find an irregular chromatophore containing a brown pigment called phyco- phaein. From certain cells of the filament spherical sporangia (spore cases) arise, which are unicellular. They contain numerous biciliate zoospores , which escape into the sea water, move about and later develop into new Ectocarpus plants. Along the filaments several branches will be seen. Some of these have undergone division into several cells and these again into still smaller cells until many-celled cham- bers have resulted, which are called plnrilocular sporangia. Each cell of a plurilocular sporangium contains a gamete or sexual cell, which resembles in many details a zoospore. When the sporangium matures these gametes are discharged into the salt water. They fuse together in pairs and form zygospores. Each zygospore under- 13 Pig. 78. — End of large branch of Fucus vesiculosus (natural size); e, re- ceptacle; b, air bladder. . i 9 4 PHARMACEUTICAL BOTANY goes a resting stage and upon the advent of favorable conditions develops into a new Ectocarpiis filament. Fucus Vesiculosus (The Bladder Wrack) . — This form, a brown alga, occurs as a flat thallus, which forks repeatedly, a type of branching called dichotomous. It grows near the surface of sea water, attached to rocks by means of a basal disc-shaped holdfast. In the upper branches of the thallus are to be found air bladders which are more or less spherical and usually in pairs. The tips of old branches become swollen and are termed receptacles. They are dotted over with minute cavities called conceptacles. Within these conceptacles the antheridia, or male sexual organs, and the archegonia , or female sexual organs, are produced. The conceptacles also contain numerous branching filaments called paraphyses, which arise from the cells lining the cavities. The antheridia are found as outgrowths of these paraphyses and produce sperms or male sexual cells. The oogonium is a large, globular, stalked cell and produces eight eggs, each of which is a female sexual cell. The eggs and sperm escape into the sea water. The eggs float and are surrounded by myriads of sperms. One sperm, only, gains an entrance, after which its nucleus fuses with that of the egg to form an oospore. The oospore at once develops into a new Fucus plant. Class III. — Rhodophyce^e, the Red Alg,e A greatly diversified group comprising the majority of marine algae but represented also by some fresh-water forms. The marine red algae are generally found at or just beyond the low water mark. Their vegetative bodies vary from simple branching filaments through all gradations to forms differentiated into branching stems, holdfasts and leaves. It has been observed that many of the higher types are composed of numerous filaments which are arranged so closely and connected so intimately by protoplasmic processes as to resemble the tissues of plants higher up. Their color may be red, purple, violet, or reddish-brown or even green and is due to the presence of phycoerythrin, a red pigment, which is found in the chromatophores with but frequently masking the chlorophyll. Chondrus crispus and Gigartina mamillosa yield the official drug TAXONOMY 195 Chondrus, Irish Moss or Carragheen. Both are purplish-red in color. Each consists of a dichotomously branched thallus the lower portion of which is differentiated as a stipe or stalk; the basal portion of which, called the holdfast, clings to the rock. The upper part is several times forked and its terminal segments appear notched or bilobed. Scattered here and there over the segments of the thallus will be noted sporangia which, when mature, contain tetraspores. In Chondrus crispus the sporangia are elliptical and embedded in the thallus near its surface, whereas in Gigartina they are ovate and project outward from the surface of the segments. Upon the ripen- I ing of these structures the spores are discharged into the sea water. These sooner or later germinate into new Chondrus or Gigartina organisms. The dried mucilaginous substance extracted from Gracilaria lichenoides, Gelidium and Gloiopeltis and other species of red algae growing in the sea along the eastern coast of Asia con- stitutes the drug Agar, a most valuable ingredient in culture media [ as well as a laxative. Rhodymenia palmata or Irish Dulse is a purplish-red, flat, mem- branous, palmately cleft or dichotomous red alga growing on the tissues of other algae along northern shores of the Atlantic between the low- and high-tide marks. SUBDIVISION IV.— FUNGI This great assemblage of thallophytes is characterized by the total absence of chlorophyll and so its members possess no independent power of manufacturing food materials such as starches, sugars, etc., from C0 2 and H 2 0. Consequently they are either parasites , depend- ing for their nourishment upon other living plants or animals, called hosts; or saprophytes, depending upon decaying animal or vegetable matter in solution. Some forms are able to live either as saprophytes or parasites while others are restricted to either the parasitic or saprophytic habit. The vegetative body of a fungus is known as a mycelium. It consists of interlacing and branching filaments called hyphce, which ramify through decaying matter or invade the tissues of living organisms and derive nourishment therefrom. In the case of parasites, the absorbing connections which are more or less 196 PHARMACEUTICAL BOTANY specialized and definite are called haustoria. In the higher forms the hyphae become consolidated into false tissues, and assume definite shapes according to the species. Of this character are the fructi- fying organs which constitute the above ground parts of Puff Balls, Cup Fungi, Mushrooms, etc. There are four classes of Fungi, viz.: Phycomycetes, Ascomycetes, Basidiomycetes and Fungi Imperfecti. Class I. — Phycomycetes, or Alga-like Fungi The Phycomycetes represent a small group of fungi showing close affinity with the green algae. Their mycelium is composed of coeno- cytic hyphae, which suggests a close relation with the Siphonales group of green algae. Their sexual organs are likewise similar in structure. Transverse septa appear upon the formation of repro- ductive organs separating these structures from the vegetative hyphae. Sub-class A. — Zygomycetes (Sexual apparatus shows isogamy) Order 1.- Mucorales, the black molds, mostly saprophytic. Ex- amples: Mucor Mucedo, Rhizopus nigricans , Thamnidium, Pilobolus. Rhizopus nigricans (Mucor stolonifer), commonly known as “ Black Mold” or “ Black Bread Mold,” is frequently found on bread, jellies, syrups, acetic pharmaceutical extracts and other substrata, where it forms a dense thready mycelium bearing numerous black tiny spore cases. The source of this mold is the spores which are found in the air or water with which the attacked substratum is in contact. Each of these upon germinating sprouts out and forms three kinds of hyphae, viz.: rhizoidal or submerged hyphae, spor- angiophores or aerial hyphae and stoloniferous hyphae. The branch- ing rhizoidal hyphae penetrate the substratum and secrete a dias- tatic ferment which changes the water insoluble carbohydrate materials into a soluble sugar which passes into solution and is absorbed by their walls. This, upon entering the hyphae, is converted into protoplasm, and so the mold increases in size. Sporangiophores or aerial hyphae arise vertically or obliquely from a bulged-out TAXONOMY 197 common base of the rhizoidal hyphae. Each of these when mature bears upon its summit a spheroidal sporangium containing numerous small brownish multinucleate spores called endospores. The wall Fig. 79. — Black mold ( Rhizopus nigricans). A, older plant; myc, mycelia; sph. sporangiophore; sp, sporangium; si, stoloniferous hypha produced by A, and giving rise at its tip to a new plant, B. Greatly enlarged. (Gager.) of the sporangium is beset with asperites of calcium oxalate. Spring- ing from the base of the sporangiophores or aerial hypha? one or more stoloniferous hypha? traverse a portion of the surface of the sub- Fig. 80 . — Rhizopus nigricans. A, Young sporangium, showing columella within; B, older sporangium, with the wall removed, showing ripe spores covering the columella; C, D, views of the collapsed columella after dissemination of the spores. (Gager.) stratum and their tips, coming in contact with the substratum, swell up forming an adhesive organ or a p pressor him which branches out below into a cluster of spreading submerged hyphae and above 198 PHARMACEUTICAL BOTANY into several aerial hyphae bearing sporangia. This method of growth proceeds until the entire surface of the nutritive medium is covered with a dense fluffy mycelium. Rhizopus reproduces by two methods. The most common one is that of internal cell formation. In this asexual method a transverse wall is laid downm the sporangiophore near its tip. The terminal cell thus formed swells up, becoming globular in shape and its protoplas- mic contents become changed to form numerous spores^within the wall of the sporangium or enlarged terminal cell of the sporangio- phore. The partition wall, separating the lumen of the sporangium from that of the sporangiophore, bulges into the sporangium as a dome- shaped structure which is termed the columella. Upon the ripening of the spores the wall of the spore case bursts liberating them. These falling upon moist nutrient substrata, germinate and ultimately form new Rhizopus plants. Under certain conditions Rhizopus reproduces sexually. Thicker and shorter club-shaped hyphae arise on opposite branches of the mycelium. A partition wall is TAXONOMY 199 laid down in each ol' these a short distance from its tip and the contents of each end cell then becomes a gamete or sexual cell. The apical cells of the tips of opposite hyphae then meet, a solution of the cell walls at the point of contact takes place and the gametes of both end cells fuse to form a zygospore. This enlarges and devel- ops a highly resistant wall. After a period of rest, upon coming into contact with a nutrient medium, it germinates into an elongated sporangiophore which develops a sporangium at its summit. Mucor mucedo, another closely allied species, found growing on old nuts, fleshy fruits, bread and horse manure, resembles Rhizopus nigricans in many respects but differs from it by the formation of sporangiophores singly instead of in clusters. Thamnidium differs from Rhizopus and Mucor in the development of two kinds of sporangia, microsporangia and megasporangia. The sporangiophore produces a terminal large megasporangium possess- ing a columella and whorls of side branches which bear smaller microsporangia in which the columella is frequently wanting. Sub-class B. — Oomycetes (Sexual apparatus heterogamous) Order 1. — Chytridiales. — Example: Synchytrium, a form para- sitic on seed plants and forming blister-like swellings. Order 2.- — Saprolegniales. — Water molds which attack fishes, frogs, water insects, and decaying plants and animals. Example: Saprolegnia. Order 3. — Peronosporales. — Mildews, destructive parasites, liv- ing in the tissues of their hosts and effecting pathologic changes. Example: Albugo , the blister blight, a white rust attacking members of the Cruciferce and Phytophthora, producing potato rot. Class II. — Ascomycetes, the Sac Fungi Mycelium composed of septate filaments and life history charac- terized by the appearance of a sac called an asciis in which ascos pores are formed. The largest class of fungi. Order 1.— Protoascales. — Plants with asci borne free or at the ends of hyphae, definite fruiting bodies being absent. Each ascus 200 PHARMACEUTICAL BOTANY usually develops four ascospores. To this order belong Exoascus, which is responsible for the abnormal development of tufted masses of branches on a number of trees and shrubs, and the yeasts (Sac- charomycetaceae) many of which produce fermentation. Yeasts are unicellular plants of spheroidal, oval, elliptical, pyriform or sausage shape which reproduce by budding. They occur either in the wild or cultivated condition and are generally found capable of breaking down some form of sugar into alcohol and carbon dioxide. According to the kind or kinds of sugar fermented Hansen in 1888 classified the yeasts as follows: 1. Species which ferment dextrose, maltose and saccharose: Saccharomyces cerevisice I, S. ellipsoideus I, S. ellipsoideus II, S. pastorianus I, S. pastorianns II, S. pastorianus III. 2. Species which ferment dextrose and saccharose, but notmaltose: Saccharomyces marxianus, S. exiguus, S. saturnus , S. Ludwigii. 3. Species which ferment dextrose, but neither saccharose nor maltose: Saccharomyces mali Duclauxii. 4. Species which ferment dextrose and maltose, but not saccharose: Saccharomyces n. sp. obtained from the stomach of the honey-bee. 5. Species which ferment neither maltose, dextrose nor saccharose: Saccharomyces anomalies var. belgicus, S. farinosus, S. hyalosporus, S. membranif aciens . The two most important yeasts in the fermentation industries are Saccharomyces cerevisice and Saccharomyces ellipsoideus. Saccharomyces cerevisice, commonly called Brewer’s Yeast, is a cultivated species with many strains. It is used extensively in the brewing and baking industries and in recent years has met with considerable esteem by the medical profession in the treatment of certain skin diseases. When examined under the microscope it is found to be somewhat spheroidal to ellipsoidal in outline, 8 to 12/z long, and 8 to 10/z broad. It consists of an outer cell wall of fungous cellulose enclosing cyto- plasm and a nucleus, the latter invisible without special staining. The cytoplasm is differentiated into a clear outer membrane lying directly within the cell wall and termed the ectoplasm and an inner granular region, the endoplasm. In the young condition of the yeast cell numerous glycogen vacuoles are found scattered more or TAXONOMY 201 less uniformly throughout this region but as the cell matures these coalesce, until, in a very old cell, a huge glycogen vacuole may be seen occupying most of the interior, with the cytoplasm and nucleus pushed up against the cell wall and forming there a very narrow layer. Fig. 82. — Yeast, Saccharomyces cerevisice, the variety known as brewers’ bot- tom yeast; a, spore formation; b, elongated cells. ( After Schneider, Pharmaceu- tical Bacteriology.) Yeast plants grow in dilute saccharine solutions containing dis- solved nitrogenous substances such as beerwort, Pasteur’s solution, grape juice, etc. Here they are constantly wasting away and as constantly being built up. The question may well arise: “How do they obtain the material necessary for growth and repair?” The answer, in a general way, is not difficult. The fluid in which they live is a solution of sugars and of nitrogenous and other matters. 202 PHARMACEUTICAL BOTANY The cell walls are readily permeable. Food substances diffuse through it into the cell, and by a series of changes (which, indeed, it is no easy matter to understand) are converted into new living substance. The waste products likewise diffuse readily outward. This method of nutrition is called saprophytic, and the yeast plant is said to be a saprophyte. A striking fact must be briefly mentioned in connection with the metabolism of yeast. Many organisms exercise a profound effect on the medium in which they live. Yeast causes a wholesale destruc- tion of sugar in the surrounding fluid. One of the decomposition products of sugar is alcohol. The alcohol of commerce is produced by the activity of this plant. Saccharomyces has its times of danger and stress when the cells perish in great numbers from cold, starvation, poisons, etc. If not too suddenly exposed, however, they are able to meet adverse con- ditions by eliminating most of their water, suspending physological processes, and becoming dormant. Sometimes they enter the rest- ing condition after a process of division, when each cell divides into four parts, each of which becomes nearly dry and is surrounded by a thick wall. Such cells are called ascospores, and their production serves both as a method of multiplying the plant and of tiding over adverse conditions. They can survive for a long time without food or water, and can endure higher temperatures than the active cells and almost any degree of cold. The dried cells and spores float in the air as dust and so accomplish a dispersal of the organism. Doubtless most of them never again meet suitable environment and so sooner or later perish. But some villi fall into favorable conditions and be able to multiply enormously again, and so the species is continued. The general method of reproduction in Saccharomyces is that of gemmation or budding. A small protuberance of the cell wall com- mences to form on the parent-cell. This grows larger and a portion of the cytoplasm and nuclear material pass into it. Eventually a daughter-bud, which may assume the size of the parent-cell, is formed. This generally adheres to the parent-cell and produces one or more granddaughter-buds which in turn may produce great-granddaugh- ter-buds before separation from the parent-cell takes place. TAXONOMY 203 There are two varieties of brewer’s yeast, viz.: Top yeasts and Bottom yeasts. Top yeasts grow on or near the surface of the liquid and produce rapid fermentation at summer temperatures causing great quantities of carbon dioxide to arise to the surface and thus forming the froth which is characteristic of ale, stout and porter. Bottom yeasts grow at about 4°C. at or near the bottom of the vat. They are used in the manufacture of lager beers. Fig. 83 . — Saccharomyces cerevisea. The form or variety known as brewers' top yeast. ( Oberhefe .) Compressed yeast (Cerevisue Fermentum Compressum) N. F. consists of the mosit, living cells of Saccharomyces cerevisice or of other species of Saccharomyces, combined with a starchy or absorbent base. Saccharomyces ellipsoideus is a wild species, several varieties of which are found growing on grapes especially in districts where wine is produced. It is termed the true wine yeast to distinguish it 204 PHARMACEUTICAL BOTANY from other wild species found in grape juice, like 5. apiculatus and 5. membranij aciens which exert a deleterious effect in wine produc- tion. Its cells are ellipsoidal, 6// long, occurring singly or in rows of several generations which are rather loosely joined. Order 2. — Pezizales or cup fungi. Examples: Peziza, Lachnea and Ascobolus. Parasitic or saprophytic plants, whose vegetative bodies consist of a mycelium ramifying through the substratum and whose above ground Fig. 84 . — Saccharomyces ellipsoideus. A common yeast found on grapes, jams, jellies, etc. Budding process is shown in many of the cells as also the vac- uoles. ( Schneider , Pharmaceutical Bacteriology.) fruiting bodies are sessile or stalked, cup or saucer-shaped structures termed apothecia (sing, apothecium) in which a fruiting membrane (hymenium) lines the concave upper surface. The asci are usually eight-spored and separated from each other by filamentous struc- tures called paraphyses. Order 3.- Plectascales, the blue and green molds. Examples: Aspergillus and Penicillium. Penicillium glaucum (green mold or mildew), a type of mildew, belonging to the Ascomycetes class of Fungi, forms sage-green crusts TAXONOMY 205 on bread, jellies, old boots, gloves, and various pharmaceutical preparations. It consists of a felt-like mass of interlaced tubular hyphae called a mycelium. From the mycelium numerous hyphae project into the air and bear a green powder, the spores. These hyphae are called aerial hyphce. Other hyphae grow down into the substratum and are called submerged hyphce. When a small portion of the mycelium is mounted in 10 per cent, alcohol and observed under the high-power objective, it will be noted Fig. 85. — Saucer-shaped fruit-bodies of Pcziza repanda. {Harshberger , from Photo by W. H . Walmsley.) that each hypha has a transparent wall and protoplasmic contents and is divided by transverse septa into a number of cells. Each cell contains protoplasm, which is differentiated into cytoplasm (cell protoplasm) and several nuclei. In the cytoplasm will be seen several large clear spaces. These are vacuoles and contain water with nutritive substances in solution, called cell sap. Each hypha with its branches is clearly distinct from every other one. The aerial hyphae bear brush-like branches, which become con- stricted on their ends into a moniliform aggregation of rounded 206 PHARMACEUTICAL BOTANY spores appearing like a row of beads. Each aerial hypha is com- posed of a vertical septate branch of the mycelium called the conidio- phore, branches of this, which are called secondary conidiophores, and chains of spores at the tips of sterigmata (cells bearing conidia) Fig. 86. — A, B, Lachnea scutelala. A, Habit; B, ascus with paraphysis; C, D, Lachnea hemisphcerica; C, habit; D, ascus with paraphysis; E, Sarcosphcera aren- osa habit; F, G, Sarcosphcera coronaria; F, ascus; G, habit; H, Sarcosphtzra areni- cola ascus with paraphysis. ( See Die nalilrlichen PJlanzenfamilien I, i, p. 181.) ( Harshberger .) which are called conidia or conidios pores. The conidia form the loose green powder characteristic of Penicillium. A number of species of Penicillium are useful in the arts. Peni- cillium roqueforti is the principal ripening agent of Roquefort, TAXONOMY 207 Gorgonzola and Stilton cheeses. It possesses blue-green globular conidia 4 to 5 m in diameter. Penicillium camemberti is the principal agent in the ripening of Camembert cheese. It possesses ellipsoidal bluish-green conidia 4.5 to 5. 5 m in diameter. Fig. 87. — Three aerial hyphae showing the characteristic brush-like branching and spore formation of Penicillium glaucum. This fungus si a true saprophyte and is never found on living fruits or vegetables, a, Conidiophore branching above into secondary conidiophores; b, sterigmata; c, conidiospores. ( Schneider .) Penicillium brevicaule grows on old moist paper and has been used to detect the presence of arsenic, for when grown in media contain- ing this element, it develops the compound, diethylarsine. It is yellowish-brown in color and its conidia are rough and spiny. 208 PHARMACEUTICAL BOTANY Penicillium expansum is often found on decaying apples where it produces brownish coremia. Aspergillus herbariorum . — This green mold also named Aspergillus glaucus and Eurotium Aspergillus glaucus is frequently found on fleshy drugs which have not been properly dried. It has also been observed on dried herbarium material, old extracts, on jams, jellies, Fig. 88. — Penicillium Roqueforti. a, Part of a conidiophore; b, c, other types of branching; d, young conidiophore, just branching, e, f, conidiiferous cells; g, h, j, diagrams of types of fructification, k, l, m, n, germinating spores. ( After Thom.) tobacco, cotton-seed meal, old leather, stale black bread, etc. Like Penicillium its vegetative body consists of a mycelium consisting of aerial and submerged hyphae. It differs from Penicillium, however, mainly in not possessing septated conidiophores and by the upper portion of the conidiophores being globular. Upon the globular ex- tremity of the conidiophores are placed numerous elongated sterig- mata which bear chains of grayish-green conidia. These are spher- taxc/nomy 209 ical and prickly and range from 7 to 30 /* in diameter. Under certain conditions closed brownish fruit bodies called perithecia are produced. These arise on the surface of the substratum from spirally coiled Fig. 89. — Penicillium Camemberli. a, Conidiophore with common type of branching with conidiospores; ( b ), a common less-branched form; c, d, f, diagrams of large fructifications; g, i, j, germinating conidiospores. ( From Bull. 82, Bureau of Animal Industry, also After Thom.) hyphae and when mature possess numerous asci, each of which con- tains five to eight ellipsoidal asco spores. Aspergillus oryzoe is a yellowish-green to brown mold which 14 210 PHARMACEUTICAL BOTANY secretes diastase, a valuable digestive ferment, having the power of converting starch into sugar and dextrin. For centuries the Japa- nese have employed this species in the preparation of rice mash for Sake, as well as in manufacture of Miso and Soja sauce. The spher- ical conidiospores are 6 to 7 /i in diameter and of a yellowish-green color. Aspergillus jumi gains is a pathogenic species which produces a disease in birds, horses, cattle and even though rarely in man that is Fig. 90 . — Penicillium brevicaule. a, Conidiophores and simple chains of con- idiospores; b, f, more complex conidial fructifications; c, two young chains of con- idiospores; d, e, echinulate conidiospores;, g, h, j, sketches of forms and habits of conidial fructifications; k, germinated conidiospores. {After Thom.) called aspergillosis. The organ most prone to infection by this organism is the lung, although the skin, cornea, ears and other parts are also subject to its parasitic influence. It produces short coni- diophores with sterigmata bearing long chains of rounded, colorless conidia 2.5 to 3/1 in diameter. Harshberger 1 cites the presence of perithecia in this organism which are nut-brown, globular, 250 to 350/1 in diameter, and inclose oval thin-skinned asci with eight red lenticular ascospores each of which has a diameter of 4 to 4.5/1. Aspergillus niger ( Sterigmatocystis niger ) develops dark brown 1 “Mycology and Plant Pathology,” p. 147. TAXONOMY 2 1 1 mycelial masses in which are to be noted slender conidiophores bear- ing handle-shaped, branched sterigmata that cut off from their tips Fig. 91 . — Aspergillus oryzce associated with yeasts in the making of the Japa- nese beverage Sak6. Vegetative hyphse (a) and spore-forming hyphffi (6, c, d) are shown. (Schneider, Pharmaceutical Bacteriology.) chains of rounded black-brown conidia 3.5 to 5/u in diameter. This fungus has been found to produce suppurative inflammation of the ex- 2X2 PHARMACEUTICAL BOTANY ternal and middle portions of the human ear. It is also a cause of cork disease, so often imparting a disagreeable taste to bottled beverages. Pig. 92. — Sterigmatocystis niger ( Aspergillus niger) showing conidiophores and conidiospore formation with stages in germination of spores. ( Harshberger , after Henri Coupin.) Order 4. — Tuberales, the truffles. Fungi whose septate mycelium is often connected with the roots of trees forming the structure TAXONOMY 213 known as mycorrhiza. Several species of the genus Tuber growing in woods of France, Germany and Italy produce tuberous subter- ranean bodies called Truffles, which are highly prized as a table delicacy by the inhabitants of these countries. Order 5. — Helvellales, the saddle fungi. Fleshy fungi entirely saprophytic, living attached to leaf mold or growing in humous soil or, in a few cases, on decaying wood. The fleshy fruiting bodies (ascocarps) are divided into stalk {stipe) and cap ( pileus ) portions. Fig. 93. — The morel, Morchella esculenla. (Gager, from photo by IV. A. Murrill.) The external surface of the cap is covered with a layer of asci and paraphyses which together constitute the ascigeral layer. To this group belong the Morels and the Earth Tongues. One of the Morels, Morchella esculenta, is frequently found in fire-swept woods. Its fruiting body consists of a hollow, externally ridged stipe, bearing upon its summit a fleshy pileus whose outer surface is honeycombed with ridges and depressions. The depres- sions are covered with an ascigeral layer composed of asci and paraphyses. This species is edible. Order 6. — Pyrenomycetales, the mildews and black fungi common as superficial parasites on various parts of plants. To the black 214 PHARMACEUTICAL BOTANY fungi division of this order the Ergot fungus, Claviceps purpurea, belongs. Life History of Claviceps Purpurea. — Through the agency of winds or insects the spores (ascospores or conidia) of this organism are brought to the young ovaries of the rye ( Secale cereale). They germinate into long filaments called hyphae, which, becoming entangled to form a mycelium, spread over the ovary, enter it super- ficially, secrete a ferment, and cause decomposition of its tissue and the resultant formation of a yellow-mucus substance called honey- dew, which surrounds chains of moniliform reproductive bodies known as conidia. The honey-dew attracts certain insects which disseminate the disease to other heads of grain. The mycelial threads penetrase deeper and deeper into the ovary and soon form a dense tissue which gradually consumes the entire substance of the ovary and hardens into a purple somewhat curved body called a sclerotium, or official ergot — the resting stage of the fungus, Claviceps. The ergot falls to the ground and in the following spring sprouts into several long stalked, globular heads called stromata or ascocarps. Each (fruiting) head or ascocarp has imbedded in its surface nu- merous flask-shaped invaginations called perithecia, from the bases of which several sacs or asci develop. Within each ascus are developed eight filiform spores ( ascospores ) which, when the ascus ruptures, are discharged and are carried by the wind to other fields of grain, there to begin over a new life cycle. Class III. — Basidiomycetes, or Basidia Fungi This large class of fungi, including the smuts, rusts, mushrooms, gill and tooth fungi, etc., is characterized by the occurrence of a basid- ium in the life history. A basidium is the swollen end of a hypha consisting of one or four cells and giving rise to branches called sterigmata, each of which cuts off at its tip a spore, called a basidiospore. In addition to the basidiospores, some forms also produce spores termed chlamyd os pores. TAXONOMY 215 Fig. 94. — A, Balansia claviceps on ear of Paspalum; B-L, Claviceps purpurea; B, sclerotium; C, sclerotium with S phacelia; D, cross-section of sphacelial layer; E, sprouting sclerotium; F, head of stroma from sclerotium; G, section of same; H, section of perithecium; J, ascus; K, germinating ascospore; C, conidiospores produced on mycelium. ( See Die natiirlichen PJlanzenfamilien I, 1, p. 371.) 2l6 PHARMACEUTICAL BOTANY Fig. 95. — Smut boil of U slilago zece on ear of corn, developed from one in fected kernel. ( After Jackson, F. S., Bull. 83, Del. Coll. Agric Exper. Slat. December , 1908.) TAXONOMY 217 Sub-class A. — Protobasidiomycetes (Basidium four-celled, each cell bearing a spore) Order 1.- — Ustilaginales, the smuts. Destructive parasites which attack the flowers of various cereals, occasionally other parts of these plants. Example: Ustilago Maydis, the corn smut. The basidio- spores in this group are borne on promycelia. Fig. 96. — Germination of the chlamydospores of corn smut ( Ustilago zece)\ 1, Various stages in germination from corn 3 days after being placed in water; 2, spores germinated in contact with air; 3, several days after spores were placed in yio per cent, acetic acid, formation of infection threads, a, Spores; b, promy- celia; c, basidiospores; d, infection threads; e, detached pieces of mycelia. ( After Bull. 57, Univ. III. Agric. Exper. Slat., March, 1900.) Ustilago Maydis (Ustilago Zeae) (Corn Smut). — Corn smut is a destructive parasite which for a long time was supposed to be con- fined to the Indian Corn, but which now is known to occur on Mexican Grass. It is the only smut useful in medicine. The 2x8 PHARMACEUTICAL BOTANY mycelium of the fungus extends through all parts of the infected host through the intercellular air spaces and produces large tumor- like masses on the ears, tassels, husk, leaves and stem. Each mass is filled with spores and covered with a tightly appressed membrane which has a whitish appearance like German silver. The spores are at first a dark olive-green, but on maturity are dark brown. They are sub-spherical and show prominent spines. They arise by the division of the septate mycelium into thick-walled echinulate resting spores called chlamydospores or brand spores. These spores fall to the ground and pass the winter. In the spring each germinates into a three- or four-celled filament called a promycelium, from the cells of which basidiospores arise. The basidiospores develop a mycelium which penetrates the seedling of the host plant. Order 2. — Uredinales, the rusts. Obligate parasites possessing a septated branched mycelium which ramifies through the inter- cellular-air-spaces of the host and sends haustoria into the cell cavities. The different stages of their life cycle are either restricted to one host or distributed between two or more hosts. An outline of the life history of the wheat rust will give an idea of the peculiari- ties of the group. ' The Wheat Rust (Puccinia Graminis). — If we examine the wheat plant just before harvest we will find on the stems and leaves some rust-red lines. The presence of the mycelium of the fungus in the intercellular spaces of the host does not kill the host directly or appear to stunt its growth, but the effect of the parasite on the host is seen when the grains mature. The grains are small and mushy, due to the fact that the nutrition of the host had been dis- turbed and the formation of starch in the grains inhibited. The mycelium is localized and gives rise underneath the epidermis to rounded egg-shaped spores attached to it by short pedicels. The spores are produced in such numbers that the space beneath is too confined. As the long epidermal cells of grasses run longitudinally, the pressure of the spore masses from within causes the epidermis to crack and its edges become turned back. Through the resultant cleft the summer spores or uredos pores are thrust out. These uredo- spores are orange-brown in color and covered with minute spines. The mass of them has been called a uredinium. These spores are TAXONOMY 219 detached from the pedicels and blown by the wind to healthy plants. After summer is over and dry weather comes on, an examination of stubble in the field (blades of grass and stems of wheat left carelessly), these rust-red lines are replaced by brownish-black spores called teleutospores (teliospores). A mass of these is known as a telium. The summer stage on wheat is known as Uredo linearis. The autumn stage on wheat is known as Puccinia graminis. Fig. 97. — Spore forms of wheat rust, Pucainia graminis. A, Section through barberry leaf showing pycnia on upper surface and aecia on lower; B, two uredinio- spores; C, germinating urediniospore ; D, teliosorus showing several teliospores; E, single two-celled teliospore; F, germinating teliospore with four-celled basidium and two basidiospores; G, basidiospore growing on barberry leaf. ( Harshberger , adapted from deBary.) The teleutospores are two-celled and have thick walls and per- sistent pedicels. They remain attached to the stubble until the following spring and then either one or both cells composing them produce an outgrowth known as a promycelium (nothing but a basidium divided transversely into four cells). Each cell of the basidium is capable of producing a branch, at the tip of which a basidiospore is formed. These basidiospores are blown to the Barberry (Berberis) and infect the leaves of this plant. The mycelium runs in the intercellular air spaces and causes the appear- 220 PHARMACEUTICAL BOTANY ance of a number of small depressions on the upper side of the leaf. These in section are a rich chocolate brown and known as sperma- gonia. In the center of a spermagonium are produced hyphse, which project out to its orifice and obstrict off minute spores called sper- macia. On the opposite side of the leaf cup-shaped depressions are formed, each with a limiting membrane (peridium). Within the cup-shaped depression thousands of spores are formed in chains closely packed together. These are the aecidiospores (aeciospores). The cluster cup is called an Ticidium (/Ecium). These secio- spores are conveyed to wheat and cause infection, thus completing the life cycle. Order 3. — Auricular iales. — The so-called “ear fungi” which occur on the bark of many plants, on wooden fences, etc., as auriculate growths which when young are jelly-like and brilliantly colored, when old, hard, grayish and considerably wrinkled. The ear- like fruiting body is known as the sporophore. Its internal surface is lined with a hymenium or fruiting body consisting of numerous four-celled basidia, each of which cuts off at its tip a basidiospore. Order 4. — Tremellales. — Saprophytes which live on decaying wood as moist, soft, quivering, gelatinous growths becoming later dry and horny. Sub-class B. — Autobasidiomycetes (Mostly fleshy forms characterized by one-celled basidia with generally four, occasionally six, eight or two sterigmata each of which cuts off a basidiospore at its tip.) Division a. Hymenomycetes (Hymenium or spore-bearing surface exposed) This division of Autobasidiomycete or higher basidionrycete fungi comprises the following orders: Dacromycetales, Exobasidiales, Thelephorales, Clavariales and Agaricales. Order 1.— Dacromycetales. — This order includes the “weeping fungi.” One of the most common is Dacromyces deliquescens which occurs as a gelatinous body of bright red color on dead wood. The basidiospores are formed during a wet period and the fungus swells up in the water forming a slimy mass. In addition to basidiospores TAXONOMY 221 the mycelium may break up into oidiospores, if the wet period is prolonged. In consisting of slimy gelatinous masses the “weeping fungi” approach the Tremellacea but are distinguished from them in the basidium being undivided in the former and divided in the latter. Order 2.- Exobasidiales. — This group is found growing parasitic- ally on shrubs especially those of the heath family. The mycelium Pig. 98. — Coral-like fruit-bodies of Clavaria flava. ( Harshberger , from Photo by W. II. Walmsley.) lives in the tissues of the stems, leaves, sepals and petals and pro- duces spongy ileshy yellowish or brownish galls which are popularly called “Azalea apples.” The galls are edible. They are covered with a hymenium. Order 3. Thelephorales, forms appearing on tree trunks as leathery incrustations or as bracts on the ground, old logs, etc. Order 4. — Clavariales, the coral or fairy club fungi. Fleshy coral or club-shaped forms, all of which are saprophytes found in woods 222 PHARMACEUTICAL BOTANY growing in bunches out of leaf mold. They are all edible and of a white, yellow or some other brilliant color. Order 5. — Agaricales, the mushroom or toadstool alliance. Alike with the other members of the Basidiomycetes, the plant body con- sists of the mycelium, ramifying through the substratum, but the part which rises above the surface (the Sporophore) is in most cases differentiated into a stalk-like body called a stipe bearing upon its summit a cap or pileus, the latter having special surfaces for the hymenium. Fig. 99. — Boletus felleus in three stages of development. (After Patterson, Flora W. and Charles, VeraK., Bull. 175, U . S. Dept. Agric., pi. xxxi, Apr. 29, I9I5-) Family I. — Hydnaceae, or tooth fungi. This group is charac- terized by the hymenium being placed over purple-like, spiny or long digitate projections of the pileus. Many of the species of the genus Hydnum are edible. Family II. — Polyporaceae, or pore fungi. The sporophores or fruiting bodies of these fungi are various. They may be entirely TAXONOMY 223 supinate with pores or shallow depressions on their upper surfaces ( Merulius ), or mushroom-like {Boletus), or of the nature of woody (. Fomes ) or fleshy {Fistulina) brackets. In all cases the hymenium or basidial layer lines the inner surface of pores. The sporophore of Polyporus officinalis when deprived of its outer rind and dried constitutes the official N. F. drug Agaricus. This species grows abundantly on various species of pines, spruces and larches. Family III. — Agaricacese, the gill family, in which the hymenium covers blade -like plates of the pileus, called gills, generally occurring on the under surface of the same. Examples: Agaricus campeslris, the common edible mushroom of fields; Amanita muscaria and Amanita phalloides, both of which are poisonous. Agaricus Campestris (Common Mushroom). — This plant is an edible gill fungus which grows in open, grassy fields during late sum- mer and early autumn. It is never found in the forest or on trees or fallen trunks, seldom in the mountains. The cultivated form grows in specially constructed houses made of boards. A corridor runs through these houses so that the mushroom beds can be easily reached. In the growth of mushrooms tons of horse manure are used. This is covered with loamy soil 1 Y , l and rravancore Habitat Eastern Asia Eastern Asia TAXONOMY 315 Canellacece Family . — Trees the bark of which contains aromatic principles. Leaves alternate, pellucid-punctate. Flowers regular, golden-yellow, and arranged in terminal or axillary cymes. Fruit a berry containing two to many seeds with oily and fleshy albumen. Official drug Part used Botanical origin Habitat Fig. 174 . — Garcinia hanburii — Branch. (Sayre.) Bixacea Family . — Tropical shrubs or trees. Leaves alternate, simple with minute or no stipules. Flowers hermaphrodite or uni- 3 l6 PHARMACEUTICAL BOTANY sexual, regular; stamens hvpogynous, mostly indefinite with anthers opening by slits, rarely by one or two apical pores ( Bi.xa ). Fruit fleshy or dry. Seeds with fleshy allrumen and sometimes covered with a fleshy arillus {Bi.xa Orellana). Unofficial drug Annatto Chaulmoogra oil Part used Coloring matter Fixed oil from seeds Botanical origin Bixa Orellana Gynocardia odorata Habitat Tropical America and Madagascar India Violacece or Violet Family. — Herbs or shrub. Stems upright, rarely creeping, spreading or acaulescent. Leaves either cauline or radical, stipulate, alternate, simple to pinnatifid or palmate. Flowers pentamerous, regular or irregular {Viola). Fruit a loculi- cidally dehiscent capsule {Viola) rarely baccate. Seeds albuminous. Unofficial drug Part used Botanical origin Habitat Viola Entire herb Viola tricolor Temperate regions Turner aceae or Damiana Family. — Tropical herbs, shrubs or trees. Leaves alternate, simple, petioled, exstipulate. Flowers perfect, regular, axillary, pentamerous with one-called ovary. Fruit a capsule with three valves. Seeds strophiolate with albuminous embryo. Official drug Part used Botanical origin I Turnera diffusa Damiana N.F. Leaves Turnera | aphrodisiaca Passifloracece or Passion Flower Family. — Herbaceous or woody vines climbing by tendrils. Leaves alternate, simple, entire, lobed or compound. Flowers perfect or imperfect, solitary; peduncles jointed at the flower; perianth petaloid with urceolate or tubular tube and four to five or eight to ten partite and two-seriate limb, the throat usually crowmed by one or more series of subulate filaments which are frequently colored; gynophore elongating supporting the stamens and pistil. Fruit a one-celled berry ( Passiflora ) or three- to five-valved dehiscent capsule containing numerous seeds. Official drug Part used Botanical origin Habitat Passiflora N.F. Entire herb Passiflora incarnata United States Habitat j Lower California 1 and Mexico TAXONOMY 3*7 Caricacece or Papaw Family. — A family of latex-containing trees composed of two genera indigenous to tropical America. Of chief pharmaceutic interest is the species Carica Papaya, the Papaw or Melon tree, the fruit of which yields Papain, a valuable digestive ferment. This plant is a tree about 20 feet high which bears at its summit a cluster of deeply lobed petiolate leaves and dioecious flow- ers. The fruit is a berry, the size of one’s head and contains an acrid milky juice from which papain can be precipitated by the addition of alcohol. Cistacece or Rock Rose Family. — Herbs or shrubs whose stem and branches are often glandular, pubescent or tomentose, with simple or stellate trichomes. Leaves simple, entire, the lower ones opposite, upper alternate. Flowers perfect, regular, terminal, and solitary or in cymes or unilateral racemes; sepals five, the two external ones often bractiform or wanting; petals five {H elianlhemum) rarely three or none { Lechea ); stamens hypogynous, indefinite; carpels three to five, ovary free, one-celled. Fruit a one-celled three- to five-valved capsule. Official drug Part used Botanical origin Habitat Helianthemum Herb Helianthemum Eastern United N.F. canadense States XX. Order Opuntiales. — Cactacece or Cactus Family. — Herba- ceous rarely arborescent ( Cereus giganteus) more or less succulent plants living in warm dry {Peireskia) usually desert situations, rarely becoming epiphytic and correspondingly modified. Stems accord- ingly varying from elongate, slightly enlarged, green ( Peireskia ) to flattened ( Cereus and Opuntia) to condensed [Echino cactus , Echino- cereus, etc.) to greatly condensed ( Mamillaria ). Leaves alternate, stipulate or exstipulate, enlarged and more or less fleshy (. Peireskia ) becoming reduced green and semicircular ( Opuntia ) or modified into spines or wholly absorbed. Flowers, regular, solitary or fascicled in axils of leaves; sepals five; petals similar to sepals, petaloid, small to much enlarged, in color varying from yellow to white or from yellow to yellowish-pink, pink, scarlet or crimson; stamens indefinite in- serted at varying levels in the throat of a greatly expanded upgrown receptacle; pistil generally tricarpellary; ovary inferior, often PHARMACEUTICAL BOTANY 318 deeply sunk in upgrown receptacular part; style thread-like, divided above into as many stigmas as carpels. Fruit a receptacular berry enclosing numerous small seeds. Seeds exalbuminous. Official drug Cactus Grandiflorus N.F. Part used Fresh succulent stems Botanical origin Habitat Cactus grandiflorus 1 _ ... ... , Tropical America (Cereus grandiflorus) j Fig. 175 . — Daphne mezereum — Fruiting branch and flowers. (Sayre.) XXI. Order Myrtales (Myrtiflorae). — Thymeleacece or Mezereum Family . — Shrubs ( Daphne Mezereum ) or low trees, usually of branch- ing habit, the stems developing long tenacious bast fibers. Leaves TAXONOMY 319 alternate, rarely, opposite, coriaceous, simple, varying from lanceolate to ovate. Inflorescence a condensed raceme or spike. Flowers perfect, polygamous or dioecious, small with calyx alone of the perianth parts developed. This is crimson-purple in Daphne Mezereum. Sepals usually fused to form a tube or cup-shaped perianth. Stamens usually eight in two rows of four longer and four shorter ( Daphne Mezereum ) inserted on the calyx tube. Pistil monocarpellary; ovary superior mostly one-celled with a single pendulous ovule. Fruit a nut, drupe or berry {Daphne). Official drug Part used Botanical origin Habitat | Daphne Mezereum J Mezereum Bark Daphne Gnidium f Europe and Asia [ Daphne Laureola J Fig. 176 . — Punica granalum — Branch with flowers. (Sayre.) Punicacece (. Lythracece ) or Pomegranate Family. — Herbs ( Cuphea ), shrubs ( Decadon ) or low trees {Punica). Leaves either alternate, opposite {Punica) or whorled, simple, usually lanceolate to ovate, entire often glandular and viscous. Inflorescence a raceme, spike, or condensed cyme. Flowers perfect usually regular but pass more or less to irregular, sometimes very irregular as in genus Cuphea; 3^6 PHARMACEUTICAL BOTANY sepals five to four, more or less fused below in themselves and with calyx tube; petals commonly five, often frilled or crumpled, inserted on the mouth of the calyx tube; stamens fifteen, ten or five in alter- nate rows of five each, inserted hypogynously or perigynously; pistil six-, five-, four-, two-, rarely one-carpeled with as many cavities in the ovary and numerous small ovules; style elongate with pointed or knobbed stigma. Flowers of Punica granatum are scarlet in color. Fruit a baccate capsule ( Punica granatum) or capsule dehiscing longi- tudinally or transversely. Seeds exalbuminous. Official drug Part used Botanical origin Granatum Bark Punica Granatum Unofficial Granati Fructus Rind of fruit Punica Granatum Cortex Myrtacece or Myrtle Family. — Rarely herbs {Carey a) mostly shrubs or trees, some being the tallest trees known ( Eucalyptus ). Stems often tend to develop cork in flakes which separate much as in the Buttonwoods. Leaves rarely alternate nearly always oppo- site, entire often glistening, subcoriaceous to coriaceous {Euca- lyptus, Pimento, etc.), frequently edge-on in position upon branches. Inflorescence cymose, at times forming scorpoid cymes becoming condensed into small fascicles, or each cyme condensing into a solitary flower. Flowers regular or very rarely irregular from the lop-sided development of the stamens. Symmetry rarely hexamerous, typically pentamerous, not infrequently reduced to tetramerous {Clove)-, sepals five, six or four, aposepalous, or synsepalous at base, superior, and inserted around the edge of an expanded, upgrown receptacular disc, varying from green and more or less expanded to short thick fleshy {Clove) or reduced to teeth {Eucalyptus) ; petals equal in number to the sepals, more or less petaloid and enlarged, rarely reduced and wanting, varying in color from green through greenish-yellow to white {Eugenia species) or from whitish to pink, scarlet, crimson, purple and blue, petals sometimes synpetalous and cup-like, detaching as the flower opens; stamens usually indefi- nite and epigynous, varying in the color of their filaments as do the Habitat India India TAXONOMY 3 21 21 322 PHARMACEUTICAL BOTANY petals; pistil rarely of ten to six carpels usually of five, not infre- quently as in Clove of four carpels; ovary inferior or semi-inferior, as many-celled as there are carpels and with central placentation ; Fig. 178 . — Eugenia aromalica. (Sayre.) style elongate; stigma undivided. Fruit either a hard, woody in- dehiscent nut (Brazil Nut), a capsule dehiscing at apex ( Eucalyptus ) or berry {Eugenia). Seeds exalbuminous. TAXONOMY 323 Official drug Part used Botanical origin Habitat Eucalyptus Leaves Eucalyptus globulus | Australia, Eucalyptol Organic oxide Eucalyptus globulus j 1 Tasmania Caryophyllus Flower buds Eugenia aromatica ] [ Molucca Islands Eugenol Aromatic phenol Eugenia aromatica j Pimenta N.F. Fruit Pimenta officinalis West Indies, Central America, Mexico Oleum Cajuputi Unofficial Volatile oil from leaves and twigs Melaleuca Leucadendron East Indies Myrcia Volatile oil and leaves Myrcia acris West Indies Eucalyptus Kino Inspissated juice Eucalyptus rostrata and other species Australia Combretacece or Myrobalans Family. — Mostly tropical shrubs and trees containing considerable tannin. Leaves exstipulate, alternate or opposite, simple, pinnately veined, entire or toothed. Inflorescence a raceme, spike or head. Flowers regular, perfect or imperfect. Fruit a drupe, frequently longitudinally winged containing a single seed. Unofficial drug Part used Botanical origin Habitat Combretum Leaves Combretum Sumatra sundaicum XXII. Order Umbellales or Umbelliflorae. — Araliacece or Gin- seng Family. — Herbs (. Panax quinquefolium, Hedera Helix, Aralia nudicaulis, etc.), undershrubs ( Aralia hispida, etc.), shrubs ( Fatsia horrida), or trees ( Aralia spinosa) with stems which are more or less hollow along internodes and solid at nodes. Leaves alternate, vary- ing from simple to trifoliate or to multipinnate (tropical Aralias) or passing by telescoping [into compound-palmate. Leaves serrate margined and along with stem they develop volatile oil, resin and gum contents in secretion reservoirs. Inflorescence varying from a raceme of umbels to a raceme and even to condensed racemose umbels. Flowers regular, generally pentamerous, small, generally inconspicuous, green, greenish-yellow to rarely white, usually hermaphrodite but sometimes polygamous or dioecious; sepals I . / 3 2 4 PHARMACEUTICAL BOTANY five, rarely four; petals five, rarely four, often greenish to greenish- yellow, occasionally white, seldom pink in color; stamens varying from indefinite to ten to commonly five, opposite sepals, and, like sepals, epigynous in insertion; anthers versatile; pistil occasionally fifteen- to ten-, usually five-carpellate; ovary as many celled with one or rarely two pendulous ovules in each cavity; styles distinct ending in knob-shaped stigmas. Fruit a berry. Seeds albuminous. Official drug Part used Botanical origin Habitat Aralia N.F. Unofficial Rhizome and roots Aralia racemosa Eastern United States and Canada Aralia Nudicaulis Rhizome Aralia nudicaulis Eastern United States and Canada Aralia Spinosa Bark Aralia spinosa Eastern United States Ginseng Root Panax quinquefolium North America Panax Repens Rhizome Panax repens Japan U mbelliferce or Parsley Pam ily. — Herbs, rarely shrubs, often rapid growth, and with upright fistular (hollow at internodes, solid at nodes) often grooved and ridged stems. Leaves alternate, compound and usually much divided, exstipulate, but with expanded sheathing and flattened leaf base (Pericladium), that ensheathes the stem. Inflorescence a simple or often compound umbel sur- rounded by an involucre of bracts or of bracteoles. Flowers small, pentamerous, with inferior ovary and superior floral parts. Sepals minute, tooth-like, inserted above inferior ovary, or absorbed. Petals small, usually yellow to white, rarely pink to purple, distinct, each with indexed tip. Stamens five, epigynous, inserted below a nectariferous, epigynous disc, incurved in bud. Carpels two fused into bicarpellate pistil. Ovary two-celled, with one pendulous ovule in each cell, ovarian wall traversed by oleoresin canals; styles two distinct above the nectar disc or stylopod. Fruit a dry, splitting fruit or cremocarp, that splits lengthwise into two mericarps which hang for a time by a forked carpophore. Seeds single in each mericarp, albuminous. TAXONOMY 325 Official drug Part used Botanical origin Habitat Anisum Ripe fruit Pimpinella Anisum Asia Minor, Egypt and Greece. Anethol N.F. 1 Methyl ether of para-pro- 1 phenyl phenol | Pimpinella Anisum Foeniculum vulgare Fceniculum Nearly ripe Foeniculum vulgare Mediterranean fruit region Sumbul Rhizome and Ferula Sumbul Turkestan roots Carum Fruit Carum Carvi Europe, Asia Conium N.F. Unripe fruit Conium maculatum Europe Asafcetida Gum resin Ferula foetida, F. Persia and Asafoetida, etc. Afghanistan Coriandrum Ripe fruit Coriandrum sativum Mediterranean and Caucasian regions Petroselinum Ripe fruit Petroselinum Southern Europe sativum Asia Minor Petroselini Radix Root Petroselinum Southern Europe, N.F. sativum Asia Minor Angelicse Fructus Ripe fruit Angelica Archangel- Northern Europe N.F. ica and other species of Angelica and Siberia Angelicse Radix Rhizome and Angelica atropur- United States and N.F. roots purea and other species of Angelica Canada Apii Fructus N.F. Ripe fruit Apium graveolens England Unofficial Imperatoria Root Imperatoria Europe Ostruthium Pimpinella Roots I 1 Pimpinella Saxifraga Pimpinella magna | Central Europe Ammoniacum Gum-resin Dorema Persia Ammoniacum Galbanum Gum-resin Ferula galbaniflua Persia and Afghanistan Levisticum Root Levisticum officinale Europe Cornacece or Dogwood Family. — Herbs ( Cornus canadensis, etc.), shrubs (1 Cornus sanguinea, etc.) or trees ( Cornus florida, Nyssa sylvatica, etc.). Leaves simple, alternate (Sour Gum), or opposite (Dogwoods). Inflorescence an umbel or head, the whole being 326 PHARMACEUTICAL BOTANY surrounded by an enlarged and more or less petaloid involucre. Flowers regular, rarely pentamerous, more frequently tetramerous; sepals usually four, small tooth-like or absorbed; petals usually four, small, greenish to yellowish to white ( Cornus florida), rarely pink or crimson; stamens four or five alternate to the petals and inserted with the sepals and petals epigynously around and between the nectar disc; pistil syncarpous, bicarpellate, rarely tricarpellate; ovary as many celled with one pendulous ovule in each cavity; style usually simple, ending in rounded or slightly bilobed stigma. Fruit a two-seeded drupe. Seeds albuminous. Official drug Part used Botanical origin Habitat Cornus N.F. Bark of root Cornus florida Eastern United States and Canada Sub-class b. — Sympetal.® (Gamopetal.® or Metachlamyde^e) A flivision of dicotyledonous plants in which the flowers possess both calyx and corolla, the latter with petals more or less united into one piece. I. Order Ericales. — Ericacece or Heath Family. — Sub-herbaceous (■ Chimaphila ), suffruticose {Erica), fruticose {Azaleas, Kalmias, etc.), rarely sub-arborescent {Arbutus unedo or Strawberry Tree) plants. Roots fibrous often saprophytically associated, rarely tuberous or more or less enlarged. Stem upright, ascending or creep- ing, more or less woody, rarely through saprophytic connection be- coming soft, annual and pale above ground {Monotropa uniflora ). Leaves alternate, simple, entire, exstipulate, rarely soft, delicate, herbaceous {Azaleas), usually leathery to wiry and evergreen, more rarely {Pterospora, Monotropa, etc.) becoming greenish-blue, bluish- yellow, yellowish-white to white and correspondingly saprophytic. Inflorescence typically a raceme {Pyrola, Andromeda, Gaylussacia, Erica, Arctostaphylos Uva Ursi, etc.) but raceme condensed into a racemose umbel {Azalea, etc.) or further reduced to a few flowers or, in the degraded saprophytic condition to one flower {Monotropa uniflora). Flowers regular, passing to irregular {Rhododendron) , pentamerous or tetramerous; sepals five to four rarely fewer, apo- to synsepalous, usually green, sometimes brightly petaloid; petals TAXONOMY 327 five more rarely four, slightly to deeply synpetalous, cup-shaped (Kalmia) to urceolate ( Arctostaphylos Andromeda , etc.), yellow to white or through yellow pink to scarlet to crimson to crimson- purple; stamens ten to eight in two circles of -five to four each, be- coming by absorption of inner circle five to four only, hypogynous, epipetalous or epigynous; anthers two-celled, dehiscing by apical pores ( Arctostaphylos ) or apical slits; pollen sometimes agglutinated into long viscous threads; pistil five- to four-, rarely six- to eight- car- peled, superior, rarely semi-inferior to inferior ( V accinece ) ; ovary as many celled as there are carpels; style elongated, filiform, usually five- to four lobed. Fruit a capsule (Trailing Arbutus), berry ( Vac - cinium ) or false drupe ( Gaultheria ). Seeds small, anatropous. 328 PHARMACEUTICAL BOTANY Official drug Part used Botanical origin Habitat ' United States, Chimaphila N.F. Leaves Chimaphila umbellata J ] i Canada, North- ( ern Europe and Asia Northern United Uva Ursi Leaves Arctostaphylos Uva | Ursi J States and 1 Canada, Europe and Asia Methylis Salicylas Unofficial Volatile oil Gaultheria procumbens United States and Canada Gautheria Leaves Gaultheria procumbens United States and Canada II. Order Ebenales. — SapotacecE or Star Apple Family. — Tropical shrubs or trees ( P alaquium ) characterized by the presence of lati- ciferous sacs in the pith and cortex of the stems and adjoining the veins of the leaves. Leaves alternate, exstipulate, evergreen and coriaceous. Flowers perfect, large and axillary. Fruit a berry (P alaquium) rarely a capsule ( Ponteria ). Official drug Gutta Percha N.F. Part used f Purified coagu- j lated milky [ exudate Botanical origin Various species of Palaquium Habitat Indo-China and East Indies Styracece or Benzoin Family. — Shrubs or low trees. Leaves alter- nate to opposite, entire, often acuminate. Flowers hermaphrodite, regular, rarely sub-irregular, either in condensed fascicles or solitary in the axils of the leaves; sepals and petals typically five each; corolla often white, rarely pinkish or yellowish; stamens many to four or two, perigynous or sub-hypogynous; pistil bicarpillary or four to five carpellate. Fruit either fleshy or dry, often winged and rarely as many-celled as there are carpels. Official drug Part used Botanical origin Habitat _ , . . / Styrax Benzoin and East Indies Benzoinum Balsamic resin 1 , [ other species III. Order Contortae. — Oleacea or Olive Family.- — Shrubs {For- sythia, Chionanthus , Syringa, etc.) or trees ( Fraxinus , Olea, etc.) with stems possessing close white wood, and slightly swollen or enlarged TAXONOMY 329 nodes. Leaves opposite, decussate, simple, rarely pinnately com- pound (Ash). Inflorescence dichesial or scorpioid cymes but tending constantly toward condensation and so in the Lilac, the inflorescence becomes a clustered raceme of cymes (thyrsus). Flowers regular, pentamerous or tetramerous; sepals small, green, rarely petaloid, synsepalous; petals synpetalous, elongated into a narrow tube, ex- panding above into a stellate limb; stamens very rarely five, rarely four to three, nearly always two, epipetalous and high set on corolla tube; pistil bicarpellate, rarely of three to four carpels; ovary two- celled with two to one pendulous ovules in each cavity. Fruit either a capsule (Lilac), drupe (Olive), berry (Privet) or a winged indehiscent akene (Ash). Seeds with moderate to scanty albumen becoming occasionally exalbuminous. Official drug Manna Oleum Olivae Chionanthus N.F. Fraxinus N.F. Part used Dried saccharine exudate Fixed oil Bark of root Bark Botanical origin Fraxinus Ornus Olea Europsea Chionanthus virginica Fraxinus Americana Habitat Southern Europe Southern Europe, Algeria, Asia Southern United States Northern United States and Canada Loganiacece or Logania Family. — Herbs ( Spigelia , etc.), woody vines ( Gelsemium , etc.) or trees (. Strychnos Nux Vomica , etc.) with a bitter juice usually containing alkaloids. Stem, rarely herbaceous usually woody, often long climbing and rope-like {Gelsemium) , usually with a bicollateral bundle system. Leaves opposite, stipulate or exstipulate. Inflorescence racemose or cynrose {Spigelia (scorpioid cyme) , Strychnos) , sometimes condensed into solitary, axillary flowers. Flowers perfect, usually regular; calyx gamosepalous; corolla gamo- petalous, hypogynous, rotate, campanulate or infundibuliform; sta- mens inserted on the corolla tube or throat and with thread-like filaments; ovary superior, two-celled; style elongate with bifid stigma; ovules numerous. Fruit usually a capsule, septicidally dehiscent {Gelsemium sempervirens) or loculicidally dehiscent {Spigelia mari- landica), sometimes a berry {Strychnos Nux Vomica) or drupe. Seeds numerous or solitary, sometimes winged. 330 PHARMACEUTICAL BOTANY Fig. 180 . — Strychnos nux vomica — Flowering branch and seeds. (Sayre.) Official drug Nux Vomica Gelsemium Spigelia Ignatia N.F. Part used Seeds Rhizome and roots Rhizome and roots Seeds Botanical origin Strychnos Nux Vomica Gelsemium sempervirens Spigelia marilandica Strychnos Ignatii Habitat East Indies Southern United States Southern United States Philippine Islands TAXONOMY .331 Gentianiacece or Gentian Family. — -Herbs often low-growing. Roots and short stems sometimes more or less thickened ( Gentiana lutea). Leaves opposite, decussate, entire, exstipulate. Inflorescence cymose Fig. 1 81 . — Gentiana lutea — Flowering head and dissected flower. (Sayre.) \ {Gentiana lutea) or condensing to a single solitary terminal flower {Gentiana verna , G. acaulis, etc.) Flowers regular, perfect, penta- merous or tetramerous, sepals five to four, green, more or less synse- 332 PHARMACEUTICAL BOTANY palous, not infrequently everted or reflexed, corolla of five, rarely four petals, more or less synpetalous, in shape passing from open- stellate as in Gentiana lutea through many stages of connation to long-tubed as in Gentiana acaulis; stamens five, epipetalous; pistil bicarpellate; ovary one-celled or incompletely two-celled; style more or less elongated with bilobed to divided stigma. Fruit a capsule. Seeds albuminous. Official drug Gentiana Chirata N. F. Menyanthes N.F. Centaurium N.F. Unofficial Sabatia Part used Rhizome and roots Entire plant Leaves Flowering plant Herb Botanical origin Gentiana lutea Swertia Chirayita Menyanthes trifoliata Erythraea Centaurium Sabatia angularis Habitat Europe and Asia Minor Northern India Europe and Asia Europe Eastern United States and Canada A pocynacece or Dog Bane Family: — Herbs, rarely shrubs not infre- quently clambering or climbing in habit ( Allamanda ). Stem and branches show bicollateral bundles. Stem, leaves and flowers have latex tubes which ramify through the cortex and mesophyll tissues. Leaves alternate, opposite or verticillate, simple, entire, deciduous or evergreen. Inflorescence cymose. Flowers regular, pentamer- ous, rarely tetramerous; sepals five, gamosepalous, green, rarely sub- petaloid to petaloid; petals five, slightly to deeply gamopetalous, in shape varying from open tubular, stellate, to elongate tubular to elongate funnel-shaped, in color varying from greenish-yellow to white or from yellow to yellow-red to crimson to crimson-purple to nearly purple-blue; stamens five, epipetalous; pistil usually bicarpel- late; ovary two-celled with central placentation; style more or less elongate with terminal brush of hairs, knobbed or multifid; stigma circular band or circular spur beneath terminal style swelling. Fruit two follicles ( Apocynum , etc.), a berry, drupe, or capsule. Seeds flattened, frequently hairy, albuminous. TAXONOMY 333 Fig. 182 .—Strophanthus hispidus — Branch and seed with comose awn. (Sayre.) Official drug Part used Botanical origin [ Strophanthus Habitat Strophanthus Seeds (deprived of awn) Kombe j Strophanthus l hispidus | Africa ) Strophanthin Glucoside Srophanthus Africa hispidus Apocynum N.F. Rhizome and Apocynum United States and roots cannabinum Canada Aspidosperma Bark Aspidosperma Central and South Quebracho bianco America 334 PHARMACEUTICAL BOTANY Asclepiadacece or Milkweed Family. — Herbs or shrubs containing a milky juice, many species yielding rubber. Leaves entire, more or less fleshy, sometimes verticillate. Inflorescence usually a dichesial or scorpioid cyme. Flowers regular, pentamerous; sepals wooly, small, synsepalous; petals five, rarely four, synpetalous, elon- gated into awls; the corolla varying in shape from stellate to cam- panulate and in color from pale green to yellow, to greenish-brown, chocolate, or from white to yellow, to scarlet, to crimson, to purple, to blue; stamens five, epipetalous, fused in relation forming a cylindrical swollen mass around the central pistil; filaments flattened and furnished with a crown having various appendages; anthers two- celled, each cell containing a pollen mass (pollinium), adhering to the glandular prominences of the stigma; pistil bicarpellate, superior. Fruit typically two dry follicles {Asclepias), rarely becoming succu- lent or bladdery. Seeds numerous, compressed, imbricate with a comose appendage. Official drug Part used Botanical origin Asclepias N.F. Roots Asclepias tuberosa Condurango N.F. Bark Marsdenia Condurango IV. Order Contort® or Polemoniales . — Convolvulacea or Morning- glory Family. — Frequently herbaceous, more rarely sub-woody, woody, perennial climbing plants with underground parts sometimes swollen into tuberous roots (Jalap, Sweet Potato, Wild Man of the Earth). Stems rarely short, upright or tufted, usually elongate and circumnutating in action. Vascular bundles frequently bi- collateral. Leaves alternate, simple, exstipulate, varying from cordate to cordate-sagittal, to broad reniform to reniform, palmately lobed to palmatifid to palmately compound ( Ipomcea shows all these transitions). Stem and leaves frequently contain a dull, viscous, watery to milky-white juice. Inflorescence a scorpioid cyme be- coming reduced in some forms to a solitary flower. Flowers penta- merous; sepals five, green, gamosepalous; corolla varying in shape from rotate to funnel-like with expanded mouth, in color from greenish-yellow to white or through yellowish-pink to scarlet, crimson, purple or blue; stamens five, often with the bases of the fila- Habitat United States Peru and Ecuador TAXONOMY 335 merits expanded; pistil bicarpellate; ovary two-celled, superior, often surrounded by a nectar girdle; style filiform with bilobed or bifid stigma. Fruit usually a capsule ( Exogonium , etc.), dehiscing septi- fragally, rarely a berry. Seeds scantily albuminous to exalbuminous. Official drug Jalapa Part used Tuberous root Scammoniae Radix Root Unofficial Male Jalap Root Tampico Jalap Root Wild Jalap Root Turpeth Root Root Botanical origin Exogonium Purga Convolvulus Scammonia Habitat Mexico 1 Asia Minor, j Greece, Syria Ipomcea orizabensis Mexico Ipomoea simulans Mexico Ipomoea pandurata United States Operculina Turpethum East Indies 336 PHARMACEUTICAL BOTANY Hydro phyllacece or Water Leaf Family.- — Annual, herbaceous, rarely perennial woody plants whose stems, branches, leaves and sepals are often viscous and glandular hairy. Leaves alternate, exstipulate, from simple linear to pinnatipartite to pinnate. Inflor- escence rarely expanded, usually scorpioid cymes. Flowers small to large, funnel-form in Eriodictyon calif ornicum; sepals five, green; petals five, regular; corolla varying from small stellate with slightly fused petals to large rotate, campanulate or tubular, in color varying from greenish-white or yellow to rarely white, often pink, purple or blue; stamens five, rarely with alternate staminodes; pistil bicar- pellate. Fruit a two-celled capsule dehiscing usually septicidally. Official drug Part used Botanical origin Habitat Eriodictyon Leaves Eriodictyon California and californicum New Mexico " Borraginacece or Borage Family.- — Herbaceous (Borraginece sub- family) or shrubby ( Heliotropece sub-family), plants forming a pri- mary root and a single or often branched shoots. Leaves often divisible into expanded, sometimes large basal and alternate scat- tered cauline leaves. Each of these simple, exstipulate, often hairy, rarely glabrous. Inflorescence a raceme of dichesial or scorpioid cymes, at times condensed into a dichesium of scorpioids or a simple scorpioid cyme. Flowers pentamerous, regular, passing to slight or marked irregularity ( Ecliium ) ; sepals five; green, slightly or deeply gamosepalous, often hairy; petals five, the corolla varying in shape from rotate with shallow tube ( Myosotis and Borage ), to tubular (Symphytum) , to funnel-shaped in most species, in color, all transi- tions frequently purple-blue to blue; stamens five; pistil bicarpellate, syncarpous, embryologically two-celled with two ovules in each cavity, but dorsal ingrowths divide ovary by time of flowering into four cells with one ovule in each cavity; style gynobasic. Fruit typically four-nutlets. Seeds solitary in each cavity and either scantily albuminous ( Heliotropece ) or exalbuminous ( Borraginece ). Unofficial Symphytum (Comfrey) Cynoglossum (Hound’s tongue) Part used Root Herb and root Botanical origin Habitat Symphytum officinale Europe and United States Cynoglossum United States officinale TAXONOMY 337 Solanacece or Nightshade Family . — Stem herbaceous rarely shrubby or arborescent, frequently with bicollateral bundles. Leaves alter- nate, exstipulate, entire, or more or less lobed, rarely compound; often glandular-hairy. Flowers in cymes; regular or rarely irregular (Petunia, Tobacco sps.), pentamerous, perfect, synphyllous; sepals green (rarely petaloid), rotate to tubular, usually persistent and accrescent; petals rotate (Solatium), to tubular (Atropa), to funnel- shaped (Tobacco), and so (i) open to all comers, or (2) to bees or 22 338 PHARMACEUTICAL BOTANY wasps, or (3) to butterflies, moths; color, greenish-yellow, or greenish-white, to white, to pink, crimson, purple, rarely blue; sta- mens five, epipetalous, hypogynous, along with style usually forming nectar glands. Filaments short to long, anthers dehiscing longi- tudinally or by apical pores; pistil bicarpellate, syncarpous, with or without nectar girdle, superior ovary, two-celled with central placentation, ovules numerous, style more or less elongate with bilobed or bifid stigma. Fruit, a capsule (Tobacco, Thornapple, Henbane) dehiscing longitudinally or transversely; ora berry (potato, egg-plant, tomato, red pepper). Seeds albuminous. TAXONOMY 339 Official drag Part used Botanical origin Habitat 1 Central and Belladonnas Folia Leaves Atropa Belladonna | Southern Europe Belladonnas Radix Root Atropa Belladonna J Asia Minor and Persia Stramonium Leaves Datura Stramonium and D. Tatula Asia and Tropical America Hyoscyamus Leaves and flower tops FTyoscyamus niger Europe, Asia Solanum N.F. Ripe fruit Solanum carolinense United States Capsicum Fruit Capsicum frutescens Tropical America Dulcamara N.F. Unofficial Twigs and stems Solanum Dulcamara Europe and Asia Duboisia Leaves Duboisia myoporoides Tabacum Leaves Nicotiana tabacum Tropical America Scopola Rhizome Scopola Carniolica Alps and Carpathian Mts. Manaca Root Brunfelsia Hopeana Tropical America Paprika Fruit Capsicum annuum | America ? culti- Pimiento varieties vated Fruit Variety of Capsicum annuum Spain 340 PHARMACEUTICAL BOTANY Scorphulariacece or Figwort Family. — Herbs ( Linaria , Verbascum, Gerardia, Digitalis, etc.), shrubs (shrubby Veronicas, etc.), rarely trees ( Paulownia imperialis). Stem, branches and leaves usually green and independently vegetating, but in Pedicularis, Gerardia, Euphrasia, Buchner a, Rhinanthus, tic., the stem, leaves, and branches are condensed from the development of a parasitic root habit. Stems cylindrical to frequently quadrangular, especially when leaves are opposite. Leaves alternate to opposite and decussate, simple, exstipulate, often hairy but becoming by drought or parasiticism reduced to scales or almost absorbed. Inflorescence a raceme of cymes ( Paulownia ) or a simple raceme {Foxglove, Linaria, etc.) or spike {Verbascum Thapsus ) or, if leaves are opposite, often a whorl of axillary flowers or solitary axillary flowers. Flowers rarely regu- lar, mostly irregular; calyx of five sepals condensed in Veronica to four through absorption of one sepal by fusion of two sepals; corolla of five to four petals, deeply synpetalous varying from rotate {Ver- bascum Blattaria, etc.) to irregular tubular to elongate, irregular bilabiate to funnel-shaped. In color corolla varies from greenish to greenish-yellow or white ( Scrophularia ) to pure white or from red to purple to blue {Veronica) . Stamens five, fertile, equal in length in a few Verbascum species or unequal in other Verbascum species to stamens four with a long sterile staminode (. Pentstemon ) to four didynamous stamens with a short petaloid staminode {Scrophu- laria) to four didynamous stamens with a minute often nectariferous staminode {Linaria), to frequenty four didynamous stamens only, the two lateral or two anterior stamens stronger and longer {An- tirrhinum) to two perfect stamens and two minute staminodes {Cal- ceolaria) to two stamens alone developed {Veronica). Pistil bicarpellate; ovary two-celled with central placentation; style terminal with bilobed stigma; ovules numerous, small. Fruit a two-celled and usually many-seeded capsule. Seeds richly al- buminous, anatropous or amphitropous. Official drug Digitalis Leptandra N.F. Part used Leaves Rhizome and roots Botanical origin Digitalis purpurea Veronica virginica Habitat Europe United States and Canada TAXONOMY 341 Fig. 187 .' — Digitalis purpurea var. gloxinaefolia. 342 PHARMACEUTICAL BOTANY Official drug Verbasci Flores N.F. Part used Corollas with stamens Verbasci Folia N.F". Leaves Botanical origin [ Verbascum I phlomoides | Verbascum l thapsiforme j Verbascum Thapsus j and other species of Verbascum Habitat ] | Europe and Asia ! Europe and Asia Pedalinacece or Sesame Family. — Tropical herbs often thickly covered with viscous hairs. Leaves soft, usually alternate, more rarely opposite, exstipulate. Flowers irregular, pentamerous. Fruit a capsule ( Sesamum , etc.) drupe or rarely a one-seeded indehiscent nut. Seeds exalbuminous usually. Official drug Part used Botanical origin Habitat Oleum Sesami Fixed oil Sesamum indicum Asia and Africa (cultivated varieties) Acanthacece or Acanthus Family. — Usually herbaceous ( Ruellia ), rarely sub-woody or woody plants, occasionally bushy in habit, con- taining cystoliths in the mesophyll or epidermal cells of the leaves and in the parenchyma of the roots and stems. Leaves opposite, more rarely whorled, entire, exstipulate. Inflorescence a raceme of condensed cymes becoming a simple raceme or spike, rarely con- densed into a solitary terminal inflorescence. Flowers hermaphro- dite, usually irregular; calyx five-cleft; corolla hypogynous, gamo- petalous, more or less bilabiate, funnel-form and composed of five sepals; stamens usually four ( Ruellia , etc.), occasionally reduced to two as in genus Dianthera, didynamous or diandrous, epipetalous; pistil bicarpellate; ovary two-celled, superior, with numerous cam- pvlotropal ovules; style terminal, filiform. Fruit a capsule contain- ing numerous curved seeds. The family is of pharmaceutic interest mainly because of Ruellia ciliosa, a pubescent perennial herb growing in the Eastern United States, whose rhizome and roots have fre- quently been admixed with or substituted for Spigelia. Labiatce (. Lamiacece ) • or Mint Family. — Herbs producing creeping runners that spread out and root at the nodes. Stems quadrangular, rarely cylindrical in outline. Leaves opposite, decussate, mainly petiolate; leaf margin nearly always serrate, dentate or crenate. TAXONOMY 343 Stems and leaves further characterized by the presence of glandular hairs containing aromatic volatile oil. These hairs consist of a short one-celled stalk and a head (gland) of six or eight cells. Inflo- rescence a raceme or spike of verticillasters (double dichesial cymes) or, as in the Ground Ivy, a reduced verticillaster. Flowers typically Fig. 188 . — Mentha piperita — Flowering branch. (Sayre.) pentamerous, rarely tetramerous; sepals five, synsepalous, ribbed and forming a tubular regular or irregular bilabiate (Scull cap, etc.) calyx whose upper lip is bifid and lower trifid; corolla of five to four gamopetalous petals, hypogynous, frequently two-lipped, the upper lip bifid, the lower trifid; stamens four, didynamous, rarely one pair alone fertile and the other pair reduced in some cases almost or 344 PHARMACEUTICAL BOTANY quite to disappearing point ( Salvia and Monarda ); pistil bicarpellate, embryologically two-celled with two ovules in each cavity, becoming at time of flowering four-celled with one ovule in each cavity. Style embryologically terminal but, upon opening of flower, deeply gyno- basic, elongate, slender with two stigmatic surfaces. Fruit four nutlets enclosing as many exalbuminous seeds. Official drug Part used Botanical origin Habitat Mentha Piperita Leaves and flowering tops Mentha piperita Europe Mentha Viridis Leaves and flowering tops Mentha spicata Europe Scutellaria N.F. Entire plant Scutellaria lateriflora United States and Canada Oleum Thymi Volatile oil from flowering plant Thymus vulgaris Southern Europe Oleum Rosmarini Volatile oil from fresh flowering tops Rosmarinus officinalis Mediterranean Basin Oleum Lavendulse Volatile oil from fresh flowering tops Lavendula vera Southern Europe Cataria N.F. Unofficial Leaves and flowering tops Nepeta Cataria Europe and Asia Salvia Leaves Salvia officinalis Southern Europe Marrubium Leaves and flowering tops Marrubium vulgare Europe and Asia Hedeoma Leaves and flowering tops Hedeoma pulegioides United States and Canada Herba Majoranae Leaves and flowering tops Origanum Majorana Mediterranean regions Collinsonia Rhizome and roots Collinsonia canadense United States Serphyllum Leaves and flowering tops Thymus Serphyllum Europe and Asia Melissa Leaves and flowering tops Melissa officinalis Southern Europe, Asia Minor Monarda Leaves and flowering tops Monarda punctata United States Origanum Leaves and flowering tops Origanum vulgare Europe, Asia and North Africa Hyssopus Leaves and flowering tops Hyssopus officinalis Southern Europe TAXONOMY 345 Verbenacece or Verbena Family. — Herbs ( Verbena sps.), shrubs ( Clarodendron ), occasionally trees (Black Mangrove of Florida) having characters which averagely agree with the Labiatce. They are distinguished from the Labiatce mainly by being rarely scented, by their ilowers having five stamens and a terminal style, by their fruits being either drupes or berries and by the fact that their seeds while being typically exalbuminous, yet occasionally are slightly albuminous. Official drug Part used Botanical origin Habitat Verbena N.F. Overground por- Verbena hastata United States tion in flower V. Order Rubiales. — Rubiacece or Madder Family. — Herbs ( Gal- ium , Mitchella, etc.), shrubs ( Cephalanthus , etc.), or trees ( Cin- chona species) with fibrous roots, sometimes, as in Cephaelis Ipecacu- anha, annularly enlarged. Roots, stems and to a less extent leaves rich in varied alkaloids, some of medicinal value. Leaves opposite, entire, stipulate and interpetiolate. Inflorescence a raceme of dichesial cymes occasionally condensing to scorpioids. Flowers perfect, often dimorphic, pentamerous or tetramerous; sepals five {Cinchona, etc.) but four in Galium, small, green, subtended with other flowers by one or two or more enlarged petaloid bracts; petals five {Cinchona, etc.) to four in Galium, stellate, varying from shallow rotate to elongate tubular or funnel-shaped with stellate limbs; stamens five to four, epipetalous; pistil nearly always bicarpellate, rarely of five to four carpels; ovary inferior, two-celled with central placentation; styles either distinct with knob-shaped stigmas or style elongate, filiform ending in bilobed stigmas. Fruit varied, a capsule in Cinchona, a berry in Coffee, a drupe, or frequently as in Galium dry and splitting into nutlets; seeds albuminous, each with a curved embryo. Official drug Part used Botanical origin Habitat Cafieina Feebly basic Coffea arabica Eastern Africa principle | Cinchona Ledgeri- 1 ana, C. Calisaya 1 Cinchona Bark | and hybrids of these withother Cinchona f South America 1 1 l species 1 346 PHARMACEUTICAL BOTANY Official drug Part used Cinchona Rubra Bark CoHea Tosta N.F. Roasted seeds Botanical origin f Cinchona succirubra 1 or its hybrids I Coffea arabica [ Coffea libcrica Habitat | South America } Eastern Africa Gambir Ipecacuanha Prepared ex- tract from de- coctions of leaves and twigs Ourouparia Gambir Root | Cephaelis j Ipecacuanha I Cephaelis acuminata I East Indies Brazil United States of Columbia Fig. 189 — Cephaelis ipecacuanha — Plant and dried root. (Sayre.) TAXONOMY 347 Capr'ifoliacecE or Honey Suckle Family. — Shrubs or rarely herbs. Leaves entire, opposite, exstipulate or with delicate, attenuate or filiform stipules. Inflorescence varying from a raceme of shortened cymes to a capitulum. Flowers varying from regular and small (. Sambucus , Viburnum, etc.) to increasingly large, slightly irregular and ultimately very irregular in some Loniceras and in a few Weige- las and allies; calyx pentamerous, superior; corolla superior, gamo- petalous, limb pentafid, small, in Viburnum and Sambucus to elon- gate, tubular or irregular infundibuliform in Loniceras; stamens five, inserted on tube of corolla and alternating with corolla seg- ments; filaments equal or didynamous (in irregular flowers); ovary inferior, rarely five- to three-celled, usually three- or frequently two- celled; style terminal. Fruit a berry ( Viburnum ) from an inferior ovary, several celled, occasionally becoming one-celled with several to rarely one seed, or fruit a capsule ( Diervilla , Weigelia). Seeds albuminous. Official drug Part used Botanical origin Habitat Sambucus N.F. Flowers Viburnum Prunifolium Bark Viburnum Opulus Bark N.F. VI. Order Campanulales. — Cucurbitacece or Gourd Family. — Her- baceous, very often annual ( Colocynth , etc.), more rarely perennial {Bryonia, etc.), sometimes shrubby plants, the perennial and shrubby forms perennating by swollen roots some of which are heavy and tuberous. Stems very usually grooved and ridged, often provided with roughened and barbed hairs. Tendrils are frequently produced in the axils of leaves from tendril axillary buds (Pumpkin, Colocynth, Watermelon, Cucumber, Bryony, etc.). Leaves varying from entire, simple, usually deltoid to triangular through stages of trilo- bate, pentalobate, deeply palmatifid to palmatipartite to seldom ap- proaching compound (Colocynth). Venation in nearly all cases pal- mate. Leaves thin, herbaceous, much expanded, often hairy. Vas- cular bundles of petioles, branches and stems, bicollateral. Inflo- 348 PHARMACEUTICAL BOTANY rescence either of loose cymes or more frequently of racemes or spikes or entire axillary inflorescence may become solitary axillary. Flow- ers pentamerous, very rarely tetramerous, monoecious ( Bryonia alba) or dioecious ( Bryonia dioica ); sepals five, gamosepalous, adnate to ovary; corolla of five, rarely four gamopetalous petals varying in size and shape from small to large campanulate or broadly cup-shaped (Cucumber) and in color from greenish-yellow to greenish white to pure yellow to yellowish- white to white; stamens typically five, epigynous, with anthers either joined by pairs or synantherous; carpels usually three; ovary inferior, one- to three-celled. Fruit a pepo (a berry from an inferior ovary with thick skin). Seeds flat and exalbuminous. Fig. 190. — Colocynth — Portion of vine and whole fruit. (Sayre.) Official drug Bryonia N.F. Colocynthis Pepo Elaterinum Unofficial Watermelon Seed Momordica (Bal- sam apple) Part used Root Pulp of fruit Seeds Principle from elaterium Botanical origin Habitat | Bryonia alba l Bryonia dioica J Citrullus Colocynthis Cucurbita Pepo (cultivated varieties) Ecballium Elaterium Europe Africa and Asia Tropics Mediterranean region Seeds Citrullus vulgaris Southern Asia Fruit Momordica East Indies Balsamina Campanulacece or Bluebell Family . — Herbs of annual or more com- monly perennial growth rarely sub-shrubby or sub-woody in habit, frequently with laticiferous tubes containing a milky juice. Stem upright or feeble and spreading. Leaves alternate, simple, "exstipu- TAXONOMY 349 late. Inflorescence primitively a racemose cyme condensing into a raceme, to a sub-capitulum and ultimately to a capitulum. Flowers regular, campanulate to campanulate-elongate to elongate and deeply cleft in petals; sepals five, only slightly synsepalous, epigynous; petals five, campanulate to campanulate-tubular to tubular elongate to tubular and deeply cleft; corolla varying in color from greenish- yellow to yellowish-white to white or again, from yellowish-purple (rarely through yellowish-pink or red) to purple to pure blue; stamens five, epigynous, usually free from corolla; nectary epigynous; pistil usually tricarpellary; ovary as many celled as number of carpels and with central placenta; style single elongate; stigmas as many as carpels. Fruit a capsule. Seeds albuminous. The plants contain inulin. LobeliacecB or Lobelia Family. — Herbs, with inulin and latex con- tents, corresponding with Campanulaceoe in their vegetative parts, but differing from that group by having irregular flowers (pale blue in Lobelia inflata), anthers always synantherous and pistil always bicarpellate with two-celled ovary and bilobed or bilabiate stigma. Official drug Part used Botanical origin Habitat Lobelia Leaves and Lobelia inflata United States and flowering tops Canada VII. Order Aggregate. — V alerianaceee or Valerian Family . — Herbaceous often low succulent plants with creeping rhizomes, fre- quently strongly scented and possessing stimulating properties. Leaves frequently dimorphic; radical fascicled; cauline opposite; petiole dilated, exstipulate. Inflorescence a raceme of dichesial or scorpioid cymes. Flowers more or less irregular; calyx absent as such but represented by a series of teeth that are incurved in the bud and flower and which expand later into a pappose crown and act in the fruit as a pappose disseminator; corolla pentamerous, gamo- petalous, varying from rotate synpetalous to irregular tubular with petals diversely united, in color varying from greenish-white to white or pink ( Valeriana officinalis ) to crimson; stamens three to two or one {Valerian), epipetalous; pistil syncarpous; ovary usually one-celled inferior; style filiform with three stigmatic surfaces. 35 ° PHARMACEUTICAL BOTANY TAXONOMY 35 1 Fruit an akene from inferior ovary crowned by a persistent ex- panded pappose calyx rudiment. Seeds anatropous, exalbuminous. Official drug Part used Botanical origin Habitat Valeriana Rhizome and Valeriana officinalis Europe and Asia roots Composites ( Asteracece ) or Daisy Family. — Herbs, rarely shrubs or trees, of annual or perennial habit, and with watery or milky juice. Fig. 192. — Capitulum of a composite Jerusalem artichoke ( Helianthus tuberosus). A, lengthwise section of capitulum, X 1 ; B, ray flower, X 6; C, disk flower, cut lengthwise, X 6. (A after Baillon, B and C, Robbins .) Inulin is present in cell sap of parenchyma. Leaves alternate, rarely opposite, simple to compound, exstipulate. Inflorescence a capitu- lum or a raceme of capitula, each capitulum surrounded by an in- 352 PHARMACEUTICAL BOTANY volucre or protective whorl of bracts, and composed of numerous florets that may be: (a) wholly regular, tubular and hermaphrodite (Thistle, etc.); or (b) central florets as in (a), but marginals strap- Fig. 193 . — Matricaria chamomilla — Branch and dissected flowers. (Sayre) shaped or ligulate and usua’ly pistillate (Daisy, Dahlia, etc.); or (c) florets all ligulate and hermaphrodite (Dandelion, Chicory, etc.); or (d) florets in part or in whole bilabiate (Mutisia, etc.). Flowers small (florets) closely crowded, pentamerous, shaped as above, with TAXONOMY 353 ovary inferior and other floral parts superior. Sepals rudimentary tooth-like (Sunflower), or reduced to a pappose or hairy rudiment above ovary that is functionless during flowering, but that expands in fruit as a hairy fruit disseminator (Dandelion, Thistle, etc.); or sepals wholly absorbed (Daisy). Petals synpetalous, tubular, ligu- late or rarely bilabiate, greenish-yellow to white, or through pink- crimson and purple to blue (chicory). Stamens five, epipetalous, filaments distinct, anthers united into an upright anther-box (so synantherous) into which pollen is shed before or during opening of each floret. Carpels two, syncarpous, ovary inferior, one-celled with single ovule; style simple, at first short', later elongating and by collecting hairs sweeping pollen to top of anther box, then dividing into two stigmatic surfaces with stigmatic hairs for pollen reception. Fruit an indehiscent achene often (Dandelion, Thistle) crowned by the pappose, calyx rudiment. Seed single, exalbuminous. Official drug Part used Botanical origin Habitat Lactucarium Dried milk juice Lactuca virosa Europe Arnica Flower heads Arnica montana Europe and northern Asia Matricaria Flower heads Matricaria Chamomilla Europe and western Asia Calendula N.F. Ligulate florets Calendula officinalis Mediterranean basin Senecio N.F, Overground parts Senecio aureus United States Absinthium N.F. Leaves and Artemisia United States and flowering tops Absinthium Canada Eupatorium N.F. Leaves and flowering tops Eupatorium perfoliatum North America Grindelia Leaves and Grindelia camporum Western North America flowering tops Grindelia cuneifolia ' Grindelia squarrosa Inula N.F. Rhizome and roots Inula Helenium Europe and Asia Taraxacum Rhizome and roots Taraxacum officinale Europe and Asia Echinacea N.F. Rhizome and Brauneria pallida Central United Pyrethrum roots Root States Anacyclus Pyrethrum Northern Africa and southern Europe 23 PHARMACEUTICAL BOTANY 354 Official drug Lappa N.F. Farfara N.F. Santoninum Part used Botanical origin Root Arctium Lappa and other species of Arctium Leaves Tussilago Farfara Inner anhydride Artemisia pauciflora of santonic acid Habitat Europe and northern Asia Europe Russian Turkestan Unofficial Anthemis Pyrethri Flores Flower heads Flower heads (unexpanded or partly expanded) Anthemis nobilis Chrysanthemum cinerariifolium Chrysanthemum roseum Chrysanthemum Marschallii Europe Dalmatia Herzegovina Western Asia Fig. 194. — Chicory ( Cichorium Inlybus). A, portion of flowering branch; B, basal leaf (runcinate-pinnatifid) ; C, median longitudinal section through a head, showing the insertion of the flowers; D, individual flower; E, fruit (ripened ovary), showing the persistent pappus (calyx) of short scales. (Gager.) TAXONOMY 355 Official drug Santonica Carthamus Achillea Tanacetum Gnaphalium Cichorium Oleum Erigerontis Part used Unexpanded flower heads Tubular florets Leaves and flowering tops Leaves and flowering tops Leaves and flowering tops Rhizome and roots Volatile oil Botanical origin Artemisia pauciflora Carthamus tinctorius Achillea millefolium Tanacetum vulgare Gnaphalium polycephalum Cichorium Intybus Erigeron canadensis Habitat Russian Turkestan India Europe and Asia Europe North America Europe North America BIBLIOGRAPHY The following works have been consulted, and credit is due to the authors of the same: United States Pharmacopoeia IX. National Formulary IV. Le Maout and Decaisne: “Descriptive and Analytical Botany.” C. Stuart Gager: “Fundamentals of Botany.” William C. Stevens: “Plant Anatomy,” 3d Ed. Henry Kraeher: “Applied and Economic Botany.” Heinrich A. DeBary: “Comparative Anatomy of the Phanerogams and Ferns.” John W. Harshberger: “Mycology and Plant Pathology.” Joseph Y. Bergen and Bradley M. Davis: “Principles of Botany.” Benjamin M. Duggar: “Plant Physiology.” K. Goebel: “Outlines of Classification and Special Morphology.” Henry H. Rusby: “Manual of Botany.” A. L. Winton: “Microscopy of Vegetable Foods.” Robert Bentley and Henry Trimen: “Medicinal Plants” (4 vols.). Strasburger, Noll, Schenck, Schimper: “Lehrbuch der Botanik.” A. Engler und K. Prantl: “Die Naturlichen Pflanzenfamilien.” Mary L. Creighton: “Elements of Botany and Pharmacognosy.” Edson S. Bastin: “College Botany.” Wilfred W. Robbins: “Botany of Crop Plants.” Lucius E. Sayre: “Organic Materia Medica and Pharmacognosy,” 4th Ed. Coulter, Barnes and Cowles: “A Text Book of Botany.” Albert Schneider: “Pharmaceutical Bacteriology.” Benjamin L. Robinson and Merrit L. Fernald: “Gray’s New Manual of Botany.” F. O. Bower: “Practical Botany.” Palladin: “Plant Physiology.” 356 INDEX Abies, 243 Abies balsamea, 244 excelsa, 244 Abrus precatorius, 292 Absinthium N. F., 353 Acacia, 59 Catechu, 292 Senegal, 291 Acajou Gum, 304 Acanthaceae, 342 Acer, 307 spicatum, 307 Aceraceae, 307 Achillea millefolium, 355 Aconiti Folia, 275 Aconitine, micro-chemic test for, 54 Aconitum Napellus, 273 Acorus calamus, 249, 250 Acrasiales, 3 Actaea, 273 Actinomyces Myricarum, 95, 261 Adhesion, 143 Adnation, 143 Adonis N. F., 275 vernalis, 275 jfScidium, 220 JEgle Marmelos, 299 Aerogens, 174 ^Ethallia, 185 Agar, 195 Agaricaceae, 223 Agaricales, 222 Agaricus, 223 campestris, 223 Fig. of, 224 Agathis loranthifolia, 244 Agave americana, 47 Agavose, 46 Aggregatae, 349 Aggregation-body, 128 Agropyron repens, 248 Ailanthus Family, 299 Akene, 164 Albugo, 199 Albumen, 169 endospermic, 169 mode of formation, 169 perispermic, 169 perispermic and endospermic, 169 Albumens, 57 Alburnum, 113 Alcannin, 66 Alectoria, 230 Aletris farinosa, 252 Aleurone grains, 58 Alga-like Fungi, 196 Algae, blue green, 182 brown, 192 green, 186 red, 194 siphon, 191 Alkaloids, definition of, 52 examples of, 53 properties of, 52, 53 Allamanda, 332 Allium, 252 sativum, 252 Allyl-iso-sulphocyanide, 64 Aloe, 145, 250, 252 ferox, 252 Perryi, 252 vera, 252 Alpinia officinarum, 255 3S7 INDEX 358 Althrea Flores, 313 Folia, 313 officinalis, 313 rosea, 313 Amanita muscaria, 224 Fig. of, 225 phalloides, 225 Fig. of, 226 Amber, 244 Ament, 139 Ammoniacum, 325 Amygdala, 169 Amara, 289 Amygdalin, 64 Amylum, 248 Anacardiaceae, 304 Anacardium, occidentale, 304 Anacyclus Pyrethrum, 353 Andreaeales, 234 Andrcecium, 147 Andromeda, 326, 327 Anemone Ludoviciana, 275 pratense, 275 Pulsatilla, 275 Anemonella, 273 Anethol, 325 Angelica Archangelica, 325 Fructus N. F., 325 Radix N. F., 325 Angiospermae, 245 Aniline sulphate, 66 Anisum, 325 Annatto, 316 Annual ring, 108 thickening, 107, 108 Annulus, 19, 20, 224, 240 Anthemis nobilis, 354 Anther, 147 adnate, 152 attachment of, 152 dehiscence, 150 development of, 151 Fig. of, 150 gross structure of, 149 histology of, 149 Anther, innate, 152 lobes, 149 syngenesious, 148 versatile, 152 Antheridia, 192, 194, 235 Anthoceros, 232 Anthocerotales, 232 Anthocyanins, 62 Anthophore, r43 Anthotaxy, 136, 139 Antipodals, 159 Antirrhinum, 340 Apetalas, 259 Apii Fructus, 325 graveolens, 325 Apocynaceae, 332 Apocynum, cannabinum, 333 Apothecia, 229 Apothecium, 204 Applied Botany, definition, 2 Apposition, growth by, 65 Appressorium, 197 Araceae, 249 Arales, 249 Aralia N. F., 324 hispida, 323 nudicaulis, 323, 324 racemosa, 324 spinosa, 323, 324 Araliaceae, 323 Arbutus unedo, 326 Archegonia, 235 Archegonium, defined, 231 Archichlamydeae, 258 Arctium Lappa, 354 Arctostaphylos, 326, 327 Uva Ursi, 327, 328 Areca catechu, 248 nut, 248 Arenga saccharifera, 46 Aril, 169 Arillode, 169 Arisasma triphyllum, 249 Aristolochia, Fig. of, 268 reticulata, 269 INDEX 359 Aristolochia, serpentaria, 269 sipho, 101, 105 Arnica, 353 montana, 353 Artemisia Absinthium, 353 pauciflora, 354, 355 Arum Family, 249 Asafcetida, 325 Asagraea officinalis, 252 Asarum N. F., 269 flower of, 268 Asci, 209, 214 Asclepiadacese, 334 Asclepias tuberosa, 334 Ascobolus, 204 Ascocarp, 213 Ascomycetes, 199 Ascospores, 199, 209 of claviceps, 214 Ascus, 199 Ash, 329 Asparagine, 54 Asparagus, 251 Aspergillus, 204 fumigatus, 210 glaucus, 208 herbariorum, 208 niger, 210, 21 1, 212 oryzae, 209, 210, 21 1 Aspidium. See Dryopteris. Aspidosperma quebracho-bianco, 333 Assimilation, defined, 116 parenchyma, 69 Asteraceae, 351 Astragalus gummifer, 291 Atriplex, 271 Aurantiaceas, 298 Aurantii Dulcis Cortex, 298 Amari Cortex, 298 Auriculariales, 220 Austrian Pine, Fig. of cones, 26 Autobasidiomycetes, 220 Auxospore, 190, 191 Awn, 168 Azalea, 150 Azalea, amena, 148 apples, 221 Azolla, 241 Bacca, 165 Bacilli, disease producing, 179 types of, 175 Bacillus, 176 Bacteria, 174 chemical composition of, 178 classification according to form, 175, 176, 177 disease producing, 179 forms of cell groups after cleav- age, 178 morphology due to cleavage, 177, 178 mounting and staining of, 179-182 physical appearance of, 174, 175 rapidity of multiplication, 178 reproduction of, 177 Bacteriaceae, 175 Bacterioids, 94 Bacterium, 176 Balansia claviceps, 215 Balsam Apple, 348 Balsamum Peruvianum, 291 Tolutanum, 291 Balsams, 60, 61 Baptisia tinctoria, 292 Barberry Family, 273 Bark, 108 cross-section through root- bark of Enonymus purpureus, 306 Barosma, 297 betulina, 298, 299 serratifolia, 299 Basidiomycetes, 214 Basidiospore, 214 Basidium, 214 Bast, fibers, 71 hard, 105 soft, 105 Bayberry Family, 261 Beech Family, 26s 3 6 ° INDEX Beet, 271 Beggiatoa, 177 Beggiatoaceae, 177 Belae Fructus, 299 Belladonnas Folia, 339 Radix, 339 Berberidaceae, 275 Berberis N. F., 275 Beta, 271 vulgaris, 272 var. Rapa, 248 Betula lenta, 265 Betulaceae, 265 Bifacial leaf blade, 134 Birch Family, 265 Birthwort Family, 268 Bistorta, 270 Bittersweet, 339 Bixa Orellana, 316 Bixaceae, 316 Black Mangrove, 345 Blackberry, 286 Black Mold, 196, 197 Bloodroot, 280 Blue Green Alga, 182 Bluebell Family, 348 Boletus, 222, 223 edulis, 47, 64 Borage Family, 336 Borraginaceae, 336 Boswellia carterii, 300 Botany, defined, 1 departments of inquiry, 1, 2 Bottom yeast, 201, 203 Bract, 137 Bracteolar leaf, 138 Brassica napus, 283 nigra, 64, 283 Brauneria pallida, 353 Brayera, 286 Brewer’s yeast, 200 Bridal wreath, 286 Bromelin, 64 Brooksilk, 188 Brown Algae, 192 Brunfelsia Hopeana, 339 Brunnichia, 269 Bryales, 234 Bryonia alba, 347, 348 dioica, 347, 348 Bryophyta, 231 outline of, 3 Buchnera, 340 Buckthorn Family, 307 Buds, 96 accessory, 97 adventitious, 97 alternate, 97 axillary, 97 classifications of, 97 flower, 97 leaf, 97 mixed, 97 naked, 96 scaly, 96 whorled, 97 Bulb, defined, 100 tunicated, 100 scaly, 100 Burseraceae, 299 Butomus, 149 Buttercup Family, 272 Cacao Praeparatum N. F., 331 Cactaceae, 317 Cactus Family, 317 grandiflorus, 289 Caffeine, 314, 345 micro-chemic test for, 53 Calamus, 249 Calamus Draco, 248 Calceolaria, 147, 340 Calcium oxalate, types of crystals, 55 56 Calendula officinalis, 353 Callus, 66 Callitris quadrivalvis, 244 Caltrop Family, 297 Calumba, 276 Calyptra, 16, 236 9 INDEX 361 Calyptrogen, 67 Calyx, bilabiate, 143 caducous, 144 campanulate, 143 chorisepalous, 143 deciduous, 144 defined, 143 epigynous, 144 gamosepalous, 143 half-superior, 144 hypocrateriform, 143 hypogynous, 144 inferior, 144 irregular, 143 perigynous, 144 persistent, 144 regular, 143 rotate, 143 superior, 144 Cambogia, 314 Cammeliaceae, 314 Campanulaceje, 348 placentation in, 156 Campanulales, 347 Camphora, 280 Cane sugar, micro-chemic test for, 46 Canella N. F., 315 Winterana, 315 Canellaceae, 315 Cannabis sativa, 56 var. indica, 267 Capitulum, 139 Caprifoliaceae, 347 Capsicum annuum, 339 frutescens, 339 Capsule, 163 histology of a, 166 Carcerulus, 163 Cardamomii Semen, 255 Carica Papaya, 317 Caricaceae, 317 Carina, 145 Carnauba wax, 248 Carotin, 61 Carpel, defined, 154 Carpophore, 143, 164 Carragheen, 195 Carthamus tinctorius, 355 Carum Carvi, 325 Caruncle, 168 Carya, 265 Caryophyllus, 323 Caryopsis, 164, 248 Cascara Sagrada, 307 Cascarilla, 303 Cassia acutifolia, 291, 292 angustifolia, 292 Fistula, 292 Cassia buds, 280 Castanea, 265 dentata, 54, 265 Castilloa, 62 Cataria N. F., 344 Catechu, 292 Cathartocarpus fistula, 292 Catkin, 139 Caulicle, 85 Caulophyllum, 275 thalictroides, 275 Cedrella, 301 Celandine, latex of, 62 Celastraceae, 304 Celastrus, 304, 305 Cell formation and reproduction, 41 resting stage of, 43 sap, 39 Cell walls, 64 behavior of, to micro-chemic re agents, 66 growth in area and thickness, 65 Cells, epidermal-daughter, 135 neighboring, 134 subsidiary, 134 stoma-mother, 135 Cellulose, 38, 66 fungous, 200 mucilaginous modification of, 66 reserve, 66 Celsia, 147 362 INDEX Centaurium, 332 Centrospermae, 270 Cephaelis acuminata, 346 Ipecacuanha, 345, 346 Cephalanthus, 345 Cereus giganteus, 317 grandiflorus, 318 Cerevisiae Fermentum Compressum, 203 Cetraria islandica, 230, 231 Chlaza, 155 Chamaelirium luteum, 252 Chara, 191, 192 Characeae, 191 Charales, 191 Chaulmoogra oil, 316 Chelidonium majus, 281 Chenopodiaceae, 270 Chenopodiales, 270 Chenopodium, 271, 272 anthelminticum, 272 Cherry, 286 Cherry gum, 59 Chicory, 352, 353, 354 Chimaphila, 326 umbellata, 327 Chinese Galls, 304 Chionanthus, 328 virginiana, 329 Chionanthus N. F., 329 Chlamydobacteriaceae, 1 76 Chlamydospores, 214, 218 Chlorenchyma, 69 Chlorophyceae, 186 Chlorophyll, 61 Chlorophyllin, 61 Chloroplastids, 40 Chlor-zinc-iodine, 66 Chondodendron tomentosum, 276 Chondrus crispus, 194^195 Choripetalae, 259 Chromatophores, 40 Chromophyll, 61 parenchyma, 69 Chromoplastids, 41 Chromatin, 40, 43 Chromogens, 174 Chromosomes, 43 Chrysanthemum cinerariifolium, 354 Marschalii, 354 roseum, 354 Chrysarobinum, 292 Cichorium, 355 Intybus, 334, 355 Cilium, 185 Cimicifuga, 275 racemosa, 275 Fig. of, 274 Cinchona, 345 Calisaya, 345 Ledgeriana, 345 Rubra, 346 succirubra, 346 Cinnamomum Burmanii, 280 Camphora, 280 Cassia, 280 Loureirii, 280 Saigonicum, 280 Zeylanicum, 279, 280 Cinquefoil, 286 Cistaceae, 317 Citrullus colocynthis, 348 vulgaris, 348 Citrus Aurantium, 297 amara, 298, 299 Bergamia, 299 sinensis, 298 medica Limomum, 298 acida, 299 Clarodendron, 345 Class, 2 Classification, principles of, 2, 3 Clavaria flava, 221 Clavariales, 221 Claviceps purpurea, life history of, 214, 215 Clematis, 272 Clitandra, 62 Close fertilization, prevention of, 158 Clove, 320, 322 INDEX 363 Clove Bark, 280 Club Mosses, 236 Coalescence, 142 Coca, 296 Coca Family, 296 Cocaine, 53, 54, 296 Coccaceae, 175 Coccoloba platyclada, 269 uvifera, 269 Cocculus, 276 Cocillana, 301 Cocoanut oil, 248 fruit, 248 Cocos nucifera, 46, 248 Coenocyte, 191 Colfea arabica, 345, 346 liberica, 346 Coffea Tosta N. F., 346 Coffee, 345 Cohesion, 142 Cola Family, 309 Cola acuminata, 31 1 Colchici Cormus, 251 Semen, 252 Colchicum autumnale, 251, 252 Colchicine, micro-chemic test for, 54 Colocynth, 347, 348 Collenchyma, 70 Collinsonia, 344 Columella, 198 Columnar crystals, 55 Coma, 168 Combretaceae, 323 Combretum sundaicum, 323 Comfrey, 336 Commiphora, 300 Commiphora myrrha, branch of, 301 Companion cells, 78 Compositae, 351 inulin in, 51 Compressed yeast, 203 Conceptacles of Fucu, S19 Conducting parenchyma, 69 Conducting tissues, 84 Condurango, 334 Cone, 165 Confervales, 186 Conidia, 206 Conidiospores, 206 Coniferae, 243 Coniferin, micro-chemic test for, 52 Coniferales, 242 Conifer, 242 Conium maculatum, 325 Conjugates, 188 Conjugation, 43 Connation, 142 Connective, 147, 149 Contortae, 328 Convallaria majalis, 253 Convallariae Radix, 252 Flores, 252 Convolvulaceae, 334 Convolvulus Scammonia, 335 Copaiba, 292 Copernicia cerifera, 248 Coptis N. F., 275 trifoliata, 275 Corallin-soda solution, 66 Coriandrum, 325 Cork, defined, 75 Cork cambium, origin in roots, 90 origin in stems, 104 Cork formation in roots, 90 in stems, 104 Corm, 100 Corn, inflorescence of, 247 smut, 217 Cornaceae, 325 Cornus N. F., 326 Cornus canadensis, 325 florida, 325, 326 sanguinea, 325 Corolla, defined, 145 apopetalous, 145 campanulate, 146 carophyllaceous, 147 choripetalous, 145 crateriform, 146 cruciform, 145 364 INDEX Corolla, gamopetalous, 145 hypocrateriform, 146 labiate, 146 ligulate, 146 infundibuliform, 146 papilionaceous, 145 ringent, 146 rotate, 146 Correa grandiflora, 298 Corymb, 138 Corynebacterium, 176 Coto, 280 Cotyledons, 85, 1 17 Cremocarp, 164 Crenothrix, 177 Cribiform tissue, 77 Crocus sativus, 252 Croton Eluteria, 303 tiglium, 303 Crude sap, 17, 78 Crystal fibers, 55, 56 Cubeba, 260 Cucumber, 347 Cucurbitaceae, 347 Cudbear, 230 Culm, 99 Cuoxam, 66 Cuphea, 319 Cupuliferae, 265 Curcuma N. F., 255 Curcuma longa, 255 Zedoaria, 255 Cutin, 1 14 Cyanophyceae, 182 Cycas revoluta, 242 Cydonia vulgaris, 289 Cydonium, 289 Cyme, 139 Cynoglossum officinale, 336 Cypripediae, 256 Cypripedium N. F., 257 hirsutum, 257 parviflorum, 257 Cystolith, 56 Cytisus scoparius, 292 Cytology, 1, 38 Cytoplasm, 38, 40 Cytoplasmic caps, 43 Dacromyces deliquescens, 220 Dacromycetaceae, 220 Dacromycetales, 220 Dahlia, 352 Daisy, 352, 353 Daisy Family, 351 Damiana, 316 Dammar, 244 Dandelion, 352, 353 Daphne Gnidium, 319 Laureola, 319 Mezereum, 318, 319 Date Palm, 248 Datura Stramonium, 339 Tatula, 339 Decadon, 319 Dehiscence, apical porous, 150 in fruits, 161 longitudinal anther, 150 transverse, 151 valvular, 150 Delphinium, T44 Ajacis, 275 consolida, 275 Staphisagria, 275 Delphinium N. F., 275 Dermatogen, 67, 68 Desmidacese, 188 Deutzia, 72 Dextrose, 46 Dianthera, 342 Diastase, 63 Diatomaceae, 190 Diatomales, 190 Diatoms, 190 Dicotyl Plant, morphology of type, 258 Dicotyl Seed, gross structure of, 172 Dicotyledoneae, characteristics of, 257, 258 Dictyophora duplicata, 228 Dicypellium carophyllatum, 280 INDEX 365 Diervilla, 347 Differences between Gymnosperms and Angiosperms, 36, 37 Digitalis, 340 purpurea, 340 var. gloxinaefolia, 341 Digitoxin, micro-chemic test for, 52 Dionaea, 283 Dioscorea, 252 villosa, 252 Dioscoreaceae, 252 Dittany, 298 Division, 2 Dog Bane Family, 332 Dogwood Family, 325 Dog’s Tooth Violet, life history of, 32- 36 Dorema Ammoniacum, 323 Dorsoventral Leaf blades, 130-132 Dragon’s Blood, 248 Drimys Winteri, 272 Drosera, 283, 284 longifolia, 284 intermedia, 284 rotundifolia, 284 Droseraceae, 283 Drupe, 165 Dryopteris Filix-mas, alternation of generations in, 23 comparative physiology of root, stem and leaf, 18 gross structure of stem, 13 sori and sporangia, 19 growth of seedling into mature sporophyte, 23 histology of growing apex, 15 lamina, 17 mature root, 16 stem, 13 r.oot apex, 16 sori and sporangia, 19 stipe, 17 history of gametophyte genera- tion, 20-22 sporophyte generation, 13-20 Dryopteris Filix-mas, origin of new sporophyte or diploid plant from fertilized egg, 23 vascular bundles of, 15 Dryopteris marginalis (frontispiece), 13 Duboisia, myoporoides, 339 Dulcamara, 339 Duramen, 113 Ear fungi, 220 Earth Tongues, 213 Ebenales, 328 Ecballium Elaterium, 348 Echinaceae, 353 Echinocactus, 317 Echinocereus, 317 Echium, 336 Ecology, 1 Economic Botany, 2 Ectocarpus siliculosus, 192 Ectoplasm, 18 Egg Plant, 338 Ehrlich’s Anilin Water Gentian Vio- let, 181 Elaeis guineensis, 248 Elastica, 303 Elaterinum, 348 Elettaria Cardamomum, 255 Elm Family, 265 Embryo, 85 Embryo-sac, 155 maturation of, 159 Emulsion, 64 Endocarp, 160 Endodermis, 75 Endospores, 197 Endosporium, 19 Endothecium, Enzyme, 62 Enzymes, 62, 63, 64 Epicalyx, 144 Epicarp, 160 Epidermis, 71, 72, 73 Epithelium, 84 366 INDEX Equatorial plate, 45 Equisetineae, 56, 237 Equisetum, 238 arvense, 239 Erica, 326 Ericaceae, 326 Ericales, 326 Erigeron canadensis, 355 Eriodictyon californicum, 336 Eriogonum, 269 Erythronium Americanum, life history of, 32-36 Erythroxylaceas, 296 Erythroxylon Coca, 296 Eschscholtzia, 280 Etaerio, 165 Etiolin, 62 Etiology, 2 Eucalyptol, 323 Eucalyptus, 323 globulus, 321, 323 Kino, 323 rostrata, 323 Eugenia aromatica, 322, 323 Eugenol, 323 Euonymus N. F., 305 Americanus, 304 atropurpureus, 305 Europaeus, 304 Eupatorium, 353 perfoliatum, 353 Euphorbia, 62 Pilulifera, 303 Euphorbiaceae, 303 Euphrasia, 340 Eurotium, 208 Exoascus, 200 Exobasidiales, 221 Exogonium Purga, 335 Exosporium, 19, 152 Exothecium, 149 Fabaceae, 289 Fagaceae, 265 Fagales, 265 Fagot Cassia, 280 Family, 2 Farfara, N. F., 334 Fats, 59, 60 Fatsia horrida, 323 Ferment, 62 Ferments, 62 carbohydrate, 63 fat and oil, 64 glucoside, 64 proteinaceous, 64 Fern, antheridium, 21 22 apical cell, 20 archegonium, 21 foot of, 241 prothallium, 21 sperms of, 22 Ferns, 238 true, 241 water, 241 Fern Palms, 242 Fertilization, 43 in angiosperms, 159 cross, 158 self, 158 Ferula, Asafoetida, 325 foetida, 325 galbaniflua, 325 Sumbul, 325 Fibro-vascular bundles, definition, 8 bicollateral, 82 closed collateral, 81, 82 concentric, 82 open, 82 radial, 83 Ficus N. F., 267 Carica, 267 elastica, 62 Filament, 147 gross structure of, 148, 149 histology of, 149 F'licales, 241 Filicineae, 238 Fission, 42 Fistulina, 223 INDEX 367 Fixed oils, 59, 60 Flax, 295 Floral diagrams, 251 Flower, complete, 141 diandrous, 147 dichlamydeous, 143 double, 142 hermaphrodite, 142 h'exandrous, 147 imperfect, 142 liliaceous, 146 monochlamydeous, 143 neutral, 142 orchidaceous, 145 perfect, 142 pentandrous, 147 pistillate, 142 polyandrous, 147 regular, 142 stalk, 137 staminate, 142 symmetrical, 142 Fceniculum, histology of mericarp, 167 vulgare, 325 Foenum graecum, 292 Foliage leaves, 117 Follicle, 162 Fomes, 223 Forestry, 2 Formation of male gametophyte, 158, 159 female gametophyte, 159 Forsythia, 328 Fothergilla, 285 Fovilla, 34 Foxglove, 340 Fragaria, 144 Frangula, 307 Fraxinus, 328 Americana, 329 Ornus, 46, 329 Free cell formation, 42 Frond, 239 Fructose, 46 Fruit, classification of, 161-166 Fruit, definition, 160 structure, 160 Fruits, achenial, 162 aggregate, 162 baccate, 162 capsular, 162 drupaceous, 162 multiple, 162 schizocarpic, 162 simple, 161 Fucus vesiculosus, description of, 194 Fig. of, 193 Fundamental considerations, 1-13 tissue, 69 Fungi, 195 Funtumia africana, 62 elastica, 62 Galanga, 255 Galbalus, defined, 165 Galbanum, 325 Galea, 144 Galega N. F., 292 officinalis, 292 Galium, 345 Galla, 265 Galls, Chinese, 304 Japanese, 304 Gambir, 346 Gamboge family, 314 Gamete, 42, 189, 193 Gametophyte, generation of Male Fern, 20-22 Gamopetalae, 326 Garcinia Hanburyi, 3x4, 315 Gasteromycetes, 226 Gaultherase, 64 Gaultheria, 327, 328 procumbens, 64, 328 Gaultherin, 64 Gaylussacia, 326 Geaster, 226 Gelidium, 195 Gelsemium, 329 sempervirens, 330 INDEX 368 Gemmae, 232 Gemmation, 42, 202 Gentiana, 332 acaulis, 331, 332 lutea, 331, 332 verna, 331 Gentianaceae, 331 Genus, 2 Geological Botany, defined, 2 Geraniales, 292 Geranium, 293, 294 maculatum, 294 Geranium Family, 292 Gerardia, 340 Germination, 171 . Gesneria, placentation in, 156 Geum, 143, 144 Gigartina mamillosa, 194, 195 Gills of mushrooms, 223 Ginger, absence of lignin in scleren- chyme fibers of, 70, 71 Family, 253 Fig. of plant, 254 Ginkgoales, 4 Ginseng Family, 323 Glands, internal, 84 Gians, 164 Gloeocapsa, 182, 183 Gliadins, 57, 58 Globoid, 58 Gloiopeltis, 195 Glomerule, 139 Gloxinia, 156 Gluco-alkaloids, 54 Glucosides, characteristics of, 52 Glutelins, 57 Glycyrrhiza, 292 glabra, 290, 292 glandulifera, 292 Gnaphalium polycephalum, 355 Gnetales, 4 Gonophore, 143 Goosefoot Family, 270 Gossypii Cortex N. F., 313 Gossypium, 31 1, 313 Gourd, 165 Gourd Family, 347 Gracilaria lichenoides, 195 Grain, 164 Graminales, 247 Gramineae, 247 silica in, 56 Gram’s method, 180 Granati Fructus Cortex, 320 * Granatum, 320 Grape Family, 307 Grass Family, 247 Greenbrier, 115 Green felt, 191 Grindelia camporum, 353 cuneifolia, 353 squarrosa, 353 Gross anatomy, defined, 1 Ground Ivy, 343 Gruinales, 292 Guaiaci Lignum, 297 Guaiacum officinale, 297 sanctum, 296, 297 Guarana, 306 Guard cells, 135 Guarea Rusbyi, 301 Gums, 58, 59 Gutta Percha, 62, 328 Guttiferae, 314 Gyncecium, 154 apocarpous, 154 syncarpous, 154 Gynophore, 143 Gymnosperm, life history of a, 24 Gymnospermae, 241 Haas and Hill, 63 Habitat, defined, 1 Haematoxylon, 291 Hagenia, abyssinica, 286 Hairs, barbed, 73 branched, 73 candelabra shaped, 72 clavate, 72 climbing, 74 INDEX 369 Hairs, glandular, 73 hooked, 73, 74 multicellular, 72 non-glandular, 72 root, 74, 86 stellate, 72 stinging, 74 unicellular, 72 Haloxylon, 271 Hamamelidaceae, 285 Hamamelidis Folia, 286 Hamamelis virginiana, 286, 287 Hancornia, 62 Hanstein, 67 Hard Bast, 105 Haustoria, 195 Hawthorn, 286 Head, defined, 139 Hedeoma pulegioides, 344 Hedera Helix, 323 Helianthemum, 317 canadense, 317 Helianthus, tuberosus, 351 Heliotropes, 336 Helleborus niger, 275 Helonias, 252 Helvellales, 213 Henbane, 338 Hepaticae, 231 Hepburn, 62 Herb, defined, 100 Herba Majorans, 344 Hesperidin, micro-chemic test for, 51 Hesperidium, 165 Heterocysts, 184 Hevea braziliensis, 303 Hibiscus Syriacus, 311 Hilum, 155, 169 Histology, defined, 1 of Aconitum, 93 of annual dicotyl stem, 101 of anther, 149, 150 of bark, no of California Privet root, 91-93 of Cascara Sagrada, no 24 Histology of dicotyl tuberous root, 93 of dicotyl roots, of primary growth, 89, 90 of secondary growth, 90, 91 of dicotyl stems, 101-108 of fern lamina, 17 root, 16, 17 sori and sporangia, 19 stem, 13-16 stipe, 17 of fruits, 166, 167, 168 of Greenbrier stem, 114, 115 of herbaceous monocotyl stem, 1 13 of leaves, 130-136 of monocotyl roots, 88, 89 of monocotyl stems, herbaceous, 113 woody, X14, 115 of perennial dicotyl stem, 104, 105 of seeds, 1 70-1 73 Holdfast, 194 Honey dew, 214 Honeysuckle Family, 346 Horsetails, 237 Host, 195 Hounds’ tongue, 336 Humulus lupulus, 267 Huxley, 40 Hybrid, 2 Hydnacese, 222 Hydnum, 222 Flydrangea, 284 arborescens, 285 Hydrastis canadensis, 273 Hydrophytes, 83 Hydrophyllaceae, 336 Hydropterales, 241 Hymenium, 224 Hymenomycetes, 220 , Hyoscyamus, 339 niger, 339 Flypha, 195 Hyphs, aerial, 205 submerged, 205 Hyssopus officinalis, 344 INDEX 37 ° Ignatia, 330 lllicium verum, 272 Indigenous, defined, 1 Indigo, 292 Indigofera tinctoria, 292 Indirect nuclear division, 43, 44 Indusium, 19, 240 Inflorescence, defined, 136 ascending, 137 centrifugal, 137 centripetal, 137 cymose, 137 determinate, 137 descending, 137 indeterminate, 137, 138 mixed, 137 Inflorescences, of Pine, 243 of Zea Mays, 247 Imperatoria Ostruthium, 325 Indehiscent fruit, 161 Indeterminate inflorescence, solitary, 138 Indian Corn, histology of seed, 170 Indian Turnip, 249 Integuments of angiospermous ovule, 155 Intercellular-air-spaces, lysigenous, 83 schizogenous, 83 Internal glands, 84 phloem, 107 Internode, defined, 98 Intussusception, 39, 65 Inula Helenium, 353 Inulase, 63 Inulin, 51 Invertase, 46, 47, 63 Involucre, 138 Ipecacuanha, 346 Ipo/ncea, 334 orizabensis, 335 pandurata, 335 simulans, 335 Iridaceae, 252 Iris, florentina, 252 germanica, 252 Iris, pallida, 252 style of, 157 versicolor, 252 Irish Dulse, 195 Moss, 195 Isoetaceae, 237 Isoetes, 237 Jalapa, 335 Male, 335 Tampico, 335 Wild, 335 Janczewski, 67 Japanese Galls, 304 Jateorhiza palmata, 276, 2 Jerusalem Artichoke, 351 Jungermanniales, 232 Juglandaceae, 264 Juglandales, 264 Juglans N. F., 265 cinerea, 265 Juniperus, 243 communis, 244 Oxycedrus, 244 Sabina, 244 Virginiana, 244 Juniperus N. F., 244 Kamala, 303 Karyokinesis, 43, 44 Kava, 260 Kieselguhr, 190 Kino, 292 Kochia, 271 Koenigia, 270 Kola, 31 1 Krameria argentea, 292 Ixina, 292 triandra, 292 Labellum, 146 Labiatae, 70, 342 Lachnea, 204, 206 Lactase, 63 INDEX 3 . 7 1 Lactuca, 62 Leaf base, hastate, 124 virosa, 353 reniform, r24 Lactucarium, 62, 353 sagittate, 124 Lamiaceas, 342 duration, 127 Lamina, mode of development of, 130 caducous, 127 Landolphia, 62 deciduous, 127 Lappa, 354 evergreen, 127 Larix Europaea, 244 persistent, 127 Larkspur, 272 insertion, i2o-r2 2 Lateral rootlets, 16 amplexicaul, 1 20 Latex, 62, 77 cauline, 120 cells, 75, 76 clasping, 120 Laticiferous tissue, 75 connate-perfoliate. vessels, 77 equitant, t2o Lauraceae, 278 perfoliate, 120 anthers of, 149 radical, 120 Laurus, 280 ramal, 120 Lavendula vera, 344 rank, 118 Leaf, color, 127 margin, X24, 125 complete, 119 crenate, 124 definition of, 116 cleft, t26 exstipulate, 120 dentate, 124 functions, 116 divided, 126 sessile, t2o entire, 124 stipulate, 120 incised, 124 Leaf apex, 124 lobed, 124 acuminate, 124 parted, 126 acute, 124 repand, 124 aristate, 124 runcinate, 124 cuspidate, 124 serrate, 124 emarginate, 124 outline, 122 mucronate, 124 acerose, 122 obcordate, 124 acicular, 122 obtuse, 124 cuneate, 122 retuse, 124 deltoid, 124 truncate, 124 elliptical, 122 arrangement, 117 ensiform, 122 alternate, 118 filiform, 122 decussate, 118 inequilateral, 122 fascicled, 118 linear, 122 opposite, xi8 lanceolate, 122 verticillate, 118 oblong, 122 base, 123, 124 oblanceolate, 122 auriculate, 124 orbicular, 122 cordate, 124 ovate, 122 INDEX 372 Leaf outline, peltate, 122 spatulate, 122 surface, 127 glabrous, 127 glaucous, 127 hispid, 127 pubescent, 127 pellucid-punctate, 127 rugose, 127 scabrous, 127 sericious, 127 spinose, 127 tomentose, 127 villose, 127 verrucose, 127 texture, 127 coriaceous, 127 membranous, 127 succulent, 127 venation, 120 anastomosing, 120 furchate, 120 palmate, 1 20 parallel, 120 pinni, 120 reticulate, 120 Leaves, bifacial, 130, 134 binate, 126 bipinnate, 123, 126 biternate, 126 bract, 1 1 7 centric, 130, 132, 133 bracteolar, 117 compound, 122 palmately, 126 pinnately, 126 convergent, 130, 132 decompound, 126, 127 development of lamina of, 130 dorso ventral, 130 hydrophytic, 130, 132 mesophytic, 130, 131 umbrophytic, 130 xerophytic, 130, 135 foliage, 1 17 Leaves, imparipinnate, 126 lyrate, 126 palmate, 126 paripinnate, 126 pinnate, 126 primordial, 117 scale, 1 17 stomata of, 134-136 types of Angiospermous, 117 xerophytic, 130, 133, 135 Lecanora, 230 Lechea, 317 Legume, 163 Leguminosae, 289 root tubercles of, 93, 94 Lenticels, 107 Leptandra, 340 Leptome, 77, 78 Leucoplastids, 40 Levisticum, 325 officinale, 325 Lichens, 229 Life history, of an angiosperm, 32-36 of aspidium, 13-23 of black mold, 196-199 of claviceps purpurea, 214 of fern, 13 of gymnosperm, 24-31 of moss, 235 of mushrooms, 223-226 Lignin, 70 Lignocellulose, 66 Ligule, 247 Lilac, 329 Liliaceae, 249 Liliales, 249 Lilium, 249 Lily Family, 249 Lima bean, histology of, 172 Limb, 143 Limonis Cortex, 299 Linaceae, 294 Linaria, 147, 340 Linden Family, 31 1 Linin, 43 INDEX 373 Linum, 169, 295 usitatissimum, 294, 295 Lipase, 64 Liquidambar orientalis, 285, 286 Litmus, 230 Liverworts, 231 Lobelia inflata, 349 Lobeliacese, 349 Loganiaceae, 329 Lomentum, 164 Lunaria, 144 Lunularia, Fig. of, 231 Lupulinum, 267 Lycoperdales, 226 Lycoperdon, 226, 227 Lycopodiaceae, 236 Lycopodineae, 236 Lycopodium clavatum, 236, 237 Lythraceae, 319 Macis, Macrocystis, 192 Macromorphology, 1 Magnoliaceae, 272 Mahogany Family, 300 Making of sections, 10 Male Fern, 13-23 Male Jalap, 335 Mallotus philippinensis, 303 Mallow Family, 31 1 Maltase, 62 Maltose, 46 Maltum, 248 Malva rotundifolia, 311 Malvaceae, 31 1 anthers of, 149 Malvales, 309 Mamillaria, 317 Manaca, 339 Mangels, 271 Manihot, 62 utilissima, 64, 303 Manna, 47 Maple Family, 307 Marchantia, 232 Marchantiales, 232 Marrubium vulgare, 344 Marsdenia Condurango, 334 Mastiche, 304 Matico, 260 Matricaria, 352 Chamomilla, 353 Maturation of pollen grain in Ery- thronium, 34 Maw seed, 281 Mechanical tissues, 84 Medullary rays, 81 Megaceros, 232 Megasori, 155 Megaspore, 155 Megasporophyll, 154 Melaleuca Leucadendron, 323 Melia, 301 Meliaceae, 300 Melibiose, 46 Melilotus, 292 officinalis, 292 Melissa officinalis, 344 Melochia, 309 Melon tree, 317 Membrane crystals, 55, 36 Menispermum, 275 canadense, 276 Mentha piperita, 343, 344 spicata, 344 viridis, 344 Menyanthes trifoliata, 332 Mericarp, 164 histology of a, 167 Meristem, 68 Merulius, 223 Mesocarp, 160 Mesophytes, 83 Metachlamydea?, 326 Method for mounting and staining bacteria, 179, 180 for preparation of Canada balsam mount, 9 for staining and mounting par- affine material, 9, 16 374 INDEX Methylene Blue, 66 Loffler’s, 181 Methylis Salicylas, 265, 328 Mexican Grass, 46, 217 Mezereum, 318, 319 Micrandra, 303 Micrococci, types of, 175 Micrococcus catarrhalis, 179 gonorrhoea;, 179 melitensis, 179 meningitidis, 179 Micro-crystals, 55, 56 Micrometer, ocular, n stage, 11 Micrometry, n Micromorphology, 1 Micron, 11 Micropyle, 155 Microscope, compound, 4, 5, 6, 7 defined, 4 figure of, s rules for care of, 7 simple, 4 Microscopic measurement, n Microsorus, 147 Microsporangia, 149 Microspore, 151 Microsporophylls, 147 Microspira, 176 Middle lamella, 39 Milkweed Family, 334 Milkwort Family, 302 Mimoseas, 289 Mint Family, 342 Mistletoe, 37 Mitchella, 34s Mitella, 284 Mitosis, 43, 44 Mode of development of lamina, 130 Mold, black, 196, 197 bread, 196 green, 204 Momordica Balsamina, 348 Monarda, 344 punctata, 344 Monkshood 272 Monocotyledoneae, 245 Monotropa, 326 Moonseed Family, 275 Moraceae, 266 Morchella esculenta, 213 Morning Glory Family, 334 Morphology, defined, 1 Mosses, 232 Mountain Ash, 46 Mucilage, 58 cell content, 59 membrane, 59 micro-chemic test for, 59 Mucor mucedo, 198, 199 stolonifer, 196 Mucorales, 196 Mucuna pruriens, 292 Muhlenbeckia, 269 Mulberry Family, 266 Musci, 232 Mushroom, 224 Mutisia, 352 Mycelium, 195, 205 Mycobacteriaceae, 176 Mycobacterium, 176 Mycorrhiza, 213 Myosotis, 336 Myrcia acris, 323 Myrica N. F., 264 Caroliniensis, 263 cerifera, 262, 263, 264 Gale, 264 Macfarlanei, 96 Myricaceae, characters of family, 261 root tubercles of, 93, 95, 96 Myricales, 261 Myrobalans Family, 323 Myristica, 169, 277 fragrans, 278 Myristicaceae, 277 Myronase, 64 Myrosin, 64 Myrrh Family, 299 Myrrha, 300 INDEX 375 Myrtaceae, 320 Myrtales, 318 Myrtiflorae, 318 Myrtle Family, 320 Myxogastrales, 3 Myxomycetes, 184 Natural system, 2 Naturalized, defined, 1 Negundo, 307 Nepenthes, 64 Nepenthin, 64 Nepeta, cataria, 344 Nephrodium. See Dryopteris. Nicotiana tabacum, 339 Nicotine, micro-chemic test for, 53 Nidulariales, 227 Nightshade, 339 Node, 98 Non-protoplasmic cell contents, 45-64 Nostoc, 184 Nucellus, 155, 169 Nuclear membrane, 39 Nuclei, assisting, 159 polar, 159 Nucleins, 57, 58 Nucleolus, 38 Nucleus, 40 • division, 43, 44 endosperm, 159 Nut, 164 Nutgall, Fig. of, 266 Nutlet, 163 Nux Vomica, 169, 330 Nyssa, 325 Ochrea, 129 Octant cells, 23 Ocrea, 269 Ocular micrometer, 1 1 CEdogonium, 42 Oil, cedarwood, 244 chaulmoogra, 316 cocoanut, 248 linseed, 295 palm, 248 Oil, rose geranium, 294 Oils, fixed, 59 nitrogenated, 60 oxygenated, 60 sulphurated, 60 volatile, 60 Olea Europaea, 329 Oleaceae, 328 Oleander, latex of, 62 Oleoresins, 60, 61 Olibanum, 300 Olive Family, 328 Oleum anisi, 272 auranlii florum, 299 betulae, 265 bergamottae, 299 cadinum, 244 cajeputi, 323 chenopodii, 272 cinnamomi, 280 erigerontis, 355 gossypii seminis, 313 juniperi, 244 lavendulae, 344 myristicae, 278 pini pumilionis, 244 rosmarini, 344 ricini, 303 santale, 268 sesami, 342 terebinthinae, 244 theobromatis, 31 1 thy mi, 344 tighi, 303 Oogonia, 191, 192 Oogonium, 194 Oomycetes, 199 Oosphore, 22, 67, 159, 192, 194 Operculina Turpethum, 335 Operculum, 236 Ophioglossales, 4 Opium, 62, 281 Optical combination, 12 Opuntia, 317 Opuntiales, 317 376 Orchid, floral organs of, 255 Orchidaceas, 256 Orchidete, 256 Order, 2 Ordinary Parenchyma, 69 Oregon balsam, 244 Organs, essential, 142 plant, 85 reproductive, 85 vegetable, 85 Origanum, 344 majorana, 344 vulgare, 344 Oscillatoria, 183, 184 Ostwald, 62 Ourouparia Gambir, 346 Outline of plant groups, 3 Ova, 240 Ovary, 154 Ovule, amphitropous, 155 anatropous, 135 campylotropous, 155 orthotropous, 155 shape of, 155 Ovules, 154 defined, 155 in angiosperms, 155 in gymnosperms, 155 position of, 155 Pachistima, 304, 303 Palaquium, 328 Palm oil, 248 Palmaceae, 248 Panax quinquefolium, 323, 324 repens, 324 Panicle, 139 Papaver, 136 somniferum, 281 Papaveraceae, 280 Papaverales, 280 Papaw, 64 Papaw Family, 317 Papilionaceae, 289, 290 Pappus, 160 INDEX Paprika, 339 Paraphyses, 194, 204, 235 Parasite, 193 Pareira, 276 Parenchyma, 69 assimilation, 69 conducting, 69 ordinary, 69 reserve, 69 Parietales, 314 Parmelia, 230 Parsley Family, 324 Passiflora N. F., 316 incarnata, 316 Passifloraceae, 316 Pathogens, 174 Paullinia Cupana, 306 Paulownia imperialis, 340 Pea Family, 289 Peach, 286 Pear, 286 Peat mosses, 233 Pedalinaceae, 342 Pedicularis, 340 Peduncle, 137 Peireskia, 317 Pelargonium, 293 capitatum, 294 odoratissimum, 294 Radula, 294 Pellionia, 48 Penicillium, 204 brevicaule, 207, 210 camemberti, 207, 209 expansum, 208 glaucum, 204, 207 roqueforti, 206, 209 Pentstemon, 147, 340 Peony, 273 Pepo, 165, 348 Pepper Family, 239 Pepsin, 64 Perianth, 143, 145 Periblem, 67, 68 Pericambium, 90 INDEX 377 Pericarp, 160 Pericladium, 128, 324 Pericycle, 16 Periderm, 108 Peridium, 220 Peridolum, 227 Perigone, 143 Perithecia, 209, 214 Permanent mounts, preparation of, 89 Peronosporales, 199 Petals, 145 Petiole, 1 2 7-1 29 Petroselini Radix, 325 Petroselinum, 325 Sativum, 325 Peziza, 204 repanda, 205 Pezizales, 204 Phaeophyceae, 192 Phallales, 228 Phanerogamia, 241 Pharmaceutical Botany, defined, 2 Phaseolus multifiorus, 73 Phelloderm, 108 Phloem, defined, 83 internal, 107 interxylary, 107 intraxylarv, 107 parenchyma, 69 Phloroglucin, 66 Phoenix, 248 sylvestris, 46 Photogens, 174 Photosynthesis, 18 defined, 116 Phragmidiothrix, 177 Phycocyanin, 62 Phycoerythrin, 62 Phycophaein, 62, 192 Phycomycetes, 196 Phycoxanthin, 192 Phyllotaxy, 117, 118 Physcia stellaris, 229 Physiology, defined, 1 Physostigma, 157, 169, 292 Physostigma, venenosum, 292, 293 Phytogeography, 1 Phytoglobulins, 58 Phytolacca, 272 decandra, 271 Phytopalaeontology, 2 Phytopathology, 1 Phytophthora, 199 Picea, 243 canadensis, 244 mariana, 244 rubra, 244 Picrasma excelsa, 299, 300 Pigments, 61 Pileus, 213, 222 Pilobolus, 196 Pilocarpus, 298 Jaborandi, 29S microphyllus, 298 Pimenta, 323 officinalis, 323 Pimiento, 339 Pimpinella, Anisum, 325 magna, 325 Saxifraga, 325 Pinaceai, 243 Pine Family, 243 Stem, 25 Pineapple, 64 Pinenes, 60 Pinites succinifer, 244 Pinna, 239 Pinnule, 239 Pinus, 243 alba, 244 maritima, 244 montana, 244 palustris, 244 sylvestris, 27, 84 strobus, 24, 244 Piperacete, 259 Piper angustifolium, 260 cubeba, 259 methysticum, 260 nigrum, 260 37 § Piscidia Erythrina, 292 Pistacia vera, 304 lentiscus, 304 Pistachio, 304 Pistil dicarpellary 155 monocarpellary, 134 polycarpellary, 155 tricarpellary, 155 system, 154 Pix Burgundica, 244 Canadensis, 244 Liquida, 244 Placenta, defined, 156 Placentation, 156 Planococcus, 175 Planosarcina, 175 Plant, acaulescent, 98 anemophilous, 158 annual, 88 biennial, 88 caulescent, -97 cell, 38 dioecious, 142 entomophilous, 158 geography, 1 hairs, 74, 75 hydrophilous, 158 indigenous, 1 monoecious, 142 naturalized, 2 organs, 85-173 perennial, 88 tissues, 67 zoophilous, 158 Plantaginacere, placentation in, 156 Plasma membranes, 39, 45 Plasmodiocarp, 185 Plasmodium, 184 Plastids, 40 Platystemon, 280 Plectascales, 204 Plerome, 67, 68 Pleurococcaceae, 186 Pleurococcus, 186 Plum, 286 INDEX Plumule, 85 Podophyllum, 275 peltatum, 275, 276 Polemoniales, 334 Pollen, 150, 151, 152, 153 maturation of, 158, 159 Pollination, 33, 34, 158 Pollinia, 154 Polyembryony, 37 Polygala, 64 lutea, 302 senega, 302, 303 Polygalaceae, 302 Polygonaceae, 156, 269 Polygonales, 269 Polygonatum, 249 Polygonum, 269 Bistorta, 270 Polypodiales, 241 Polyporacese, 222 Polyporus, enzyme in, 63 officinalis, 223 Polytrichum commune, 234, 235 Pome, 165 Pomegranate Family, 319 Poplar, 261 Poppy Family, 280 Populi Gemrme, 261 Populus balsamifera, 261 nigrum, 261 Pore Fungi, 222 Porella, 232 Potentilla, 144 silvestris, 289 Prefloration, 140, 141 Prefoliation, 118 Primulaceae, placentation in, 156 Principes, 248 Principles of classification, 2, 3 Privet, 329 Proembryo, 192 Promycelium, 218 Proportions of stamens, 148 Protective tissues, 84 Proteins, 19, 57, 58 INDEX 3 79 Prothallial cushion, 22 Prothallus, 240 Protoascales, 199 Protobasidiomycetes, 217 Protococcales, 186 Protonema, 20, 232, 235, 240 Protophloem, 90, 104 Protophyta, 174 Protoplasm, 38, 39 Protoplasmic cell contents, 40, 41 Protoxylem, 90, 104 Prunum N. F., 288 Prunus, 286 amygdalus car. amara, 289 cerasus, 59 domestica, 288, 289 serotina, 286 virginiana, 286 Pseudobulbs, 99 Pseudomonas radicicola, 95 Pseudopodia, 185 Pseudotsuga mucronata, 244 Ptelea, 298, 299 trifoliata, 299 Pteridophyta, 4, 236 Pterocarpus marsupium, 292 ' Puccinia Graminis, 218 Puff Balls, 227 Pulsatilla, 273 Pt»lvinus, 128 Pumpkin, 347 Punica Granatum, 320 Punicaceae, 319 Purified Siliceous Earth, 190 Putamen, 161 Pyrenoid, 188 Pyrenomycetes, 213 Pyrethri Flores, 354 Pyrethrum, 353 Pyrola, 326 Pyrus malus, 288 Pyxis, 163 Quassia amara, 299 Quercus N. F., 265 Quercus alba, 265 infectoria, 265, 266 occidentaiis, 265 Suber, 265 Quillaj aN. F., 286 Saponaria, 286, 288 Raceme, 138 Radicle, 85 Ranales, 272 Ranunculaceae, 272 Ranunculus, 273 Raphe, rss, 169 Raphides, 55 Receptacle, 143 Receptacles of Fucus, 194 Red Pepper, 338 Regma, 163 Rejuvenescence, 42 Reproduction, asexual, 42 defined, 42 sexual, 42 vegetative, 42 Resin, micro-chemic test for, 61 Resina, 244 Resins, 60 Resinogenous layer, 60 Reserve cellulose, 66 parenchyma, 69 starches, 48, 49 Respiration, 116 Rhamnales, 307 Rhamnase, 64 Rhamnose, 64 Rhamnus cathartica, 307 Frangula, 64, 307 # Purshiana, 307, 308 Rheum officinale, 270 palmatum, 270 Rhinanthus, 340 Rhizoids, 20, 191, 231, 235 Rhizome, 100 Rhizopus, 198 nigricans, 196 Rhododendron, 150, 326 INDEX 380 Rhodophycese, 194 Rhodymenia palmata, 195 Rhoedales, 280 Rhubarb, 270 Rhus, cotinus, 304 Glabra N. F., 304 japonica, 304 semialata, 304 toxicodendron, 304 typhina, 304 venenata, 304 Ribes, 284 Riccia, 232 Ricinus communis, 303 Rocella, 230 Rock Rose Family, 317 Root, defined, 85 adventitious, 87 anomalous, 87 cap, 86 classification as to form, 87, 88 conical, 87 distinction from stem, 87 duration of, 88 epiphytic, 87 functions, 85, 86 fusiform, 87 generative tissues of, 86, 87 hairs, 86 histology, 88-93 lateral, 16 primary, 87 secondary, 87 system, 94 tubercles, 93-96 Roots, Dicotyledon, abnormal struc- ture in, 93 ( of primary growth, 89-90 of secondary growth, 90-91 Monocotyledon, 88, 89 napiform, 87 nodule producing, 94 Roridula, 283 Rosa, canina, 289 centifolia, 289 Rosa, Gallica, 288 Rosaceae, 287 Rosales, 284 Rosette aggregates, 55 Rosmarinus officinalis, 344 Rubber, 62 Rubi Fructus, 288 Rubi ldaei Fructus, 288 Rubiacea?, 343 Rubiales, 345 Rubus N. F., 286 cuneifolius, 286 Idasus, 288 nigrobaccus, 286, 288 strigosus, 288 villosus, 286, 288 Ruellia ciliosa, 342 Rumex N. F., 270 Rumex, 269 acetosella, 269 crispus, 270 obtusifolius, 270 Rusts, 218 Ruta graveolens, 299 Rutacese, 297 hesperidin in, 51 Ruteas, 297 Sabal, 284 Sabbatia angularis, 332 Sabina, 244 Saccharomyces anomalus, 200 apiculatus, 204 cerevisiae, 200, 201 ellipsoideus, 203, 204 exiguus, 200 farinosus, 200 Ludwigii, 200 mali Duclauxii, 200 marxianus, 200 membranifaciens, 200, 204 pastorianus, 200 Saccharum, 272 officinarum, 248 Saddle Fungi, 213 INDEX 381 Salep, 59 Salicaceae, 260 Salicales, 260 Salicin, 262 micro-chemic test for, 52 Salix, 261 alba, 261 Salvia officinalis, 344 Salvina, 241 Samara, 164 Sambucus canadensis, 347 nigra, 347 Sandalwood Family, 268 Sandaraca, 244 Sanguinaria canadensis, 281 Santalaceas, 268 Santalales, 268 Santalum album, 268 Rubrum, 291 Santonica, 355 Santoninum, 354 Sap cell, 205 nuclear, 43 vacuoles, 39 Sapindaceae, 306 Sapindales, 304 Saponin, micro-chemic test for, 52 Sapotaceae Family, 328 Saprogens, 174 Saprolegniales, 199 Saprophyte, 195 Sarcina, 175, 178 Sarcocarp, 16 1 Sarcosphaera, 206 Sarracenia, style of, 157 Sarraceniales, 283 Sarsaparilla, 251 Sassafras medulla, 280 Sassafras variifolium, 280 Saw Palmetto Palm, Fig. of, 249 Saxifragaceae, 284 Saxifrage, 284 Scale leaves, 117 Scales, 73 Scammoniae Radix, 335 Scape, 99, 137 Schizomycetes, 174 Schizophyta, 174 Schleiden, 38 Scitaminales, 253 Sclerenchyma, 70 fibers, 71 Sclerotium, 214 Scoparius, 292 Scopola Carniolica, 339 Scorpoid cyme, 139 Scotch Pine, 27 Scouring Rushes, 237 Scrophulariaceae, 340 Scullcap, 343 Scutellaria lateriflora, 344 Secretion canals, 84 reservoirs, 84 sacs, 83 Secretory cells, 77 Sections, making of, 10 radial-longitudinal, 11 tangential-longitudinal, 1 1 transverse, 10 Seed, 168 albuminous, 169 Dicotyl, 172 exalbuminous, 169 germination, 35 Monocotyl, 170 Seedling, 171 Selaginella, 237 Martensii, 238 Selaginellaceae, 237 Semen Rapae, 283 Senecio N. F., 353 aureus, 353 Senna, 292 Sepaline, position, 145 spurs, 144 stipules, 144 Sepals, 143 epigynous, 145 hypogynous, 145 perigynous, 145 382 INDEX Serenoa serrulata, 248 Serpentaria, 26g Serphyllum, 344 Sesame Family, 342 Sesamum indicum, 342 Sieve, 77, 78 Silica, 56 Siliceous Earths, 190 Silicule, 163 Siliqua, 163 Simaruba amara, 299 officinalis, 299 Simarubaceae, 299 Sinapis alba, 283 nigra, 283 Siphonales, 191 Skunk Cabbage, 249 Slime Molds, 184 Smilax, 250 medica, 251 officinale, 251, 253 ornata, 251 Smuts, 217 c Soapwort Family, 306 Soft bast, 105 Solanaceae, 337 Solaneae, 150 Solanin, micro-chemic test for, 54 Solanum N. F., 339 carolinense, 339 Dulcamara, 339 Solitary crystals, 55 Sorbinose, 46 Soredia, 229 Sorghum, 46 Sorosis, 165 Sorus, 240 Sour Gum, 32's Soy Bean, 292 Spadix, 139 Spathe, 139 Species, 2 Spermacia, 220 Spermagonia, 220 Spermatocyte, 20 Spermatophyta, 241 outline'of, 4 Spermatozoid of fern, 21, 22 Spermoderm, 168 Sperms, 194 Sphagnales, 233 Sphagnum acutifolium, 233 squarrosum, 233 Spicebush Family, 278 Spigelia, 329 marilandica, 330 Spike, 139 Spinach, 271 Spine, 99 Spirilla, types of, 176 Spirillum, 176 cholerae asiaticae, 179 obermeieri, 179 Spirochasta, 176 Spirogyra, 188, 189 Spirosoma, 176 Sporangiophores, 196 Spore, 19 brand, 218 Sporogonium, 232, 236 Sporophore, 220, 222 Sporophyte, generation, 13-20 Spruce Gum, 244 Spurge Family, 303 Spur shoot, 24 Spurs, sepaline, 144 Staff Tree Family, 304 Stage micrometer, n Stain, for “acid proof” bacteria, 181 Van Ermengems flagella, i8t, 182 Staining, 9 of bacteria, 179-182 Stamen System, 147 Stamens, color of, 148 connation of, 148 definite, 147 diadelphous, 148 didynamous, 148 epigynous, 148 gynandrous, 148 INDEX 38,3 Stamens, histology of parts, 148-150 hypogynous, 148 indefinite, 148 insertion of, 148 monadelphous, 148 perigynous, 148 polyadelphous, 148 proportions of, 148 tetradynamous, 148 triadelphous, 148 Staphisagria, 275 Staphylococcus, 175 pyogenes aureus, 179 Star Apple Family, 328 Starch, 47 assimilation, 47 Buckwheat, 50 Canna, 51 Cassava, 51 characteristics of commercial kinds, 49, 50 Corn, 50 Leguminous, 51 < j Maranta, 49 Potato, 49 Rice, 50 Rye, 50 reserve, 47 structure and composition of, 49, 5 ° Wheat, 50 Stem, defined, 97 direction of growth, 98 distinction from root, 87 functions,- 97 generative tissues of, 98 modification of, 98, 99 Stems, above-ground, 99, 100 aerial-tuberous, 99 annual, 98 ascending, 98 biennial, 98 cactoid, 99 climbing, 99 decumbent, 98 Stems, duration of, 98 elongation of, 98 endogenous, 100, 101 exceptional types of dicotyl, 106 107 exogenous, 100 fruticose, 100 herbaceous, 99 histology of annual dicotyl, 101, 102 of perennial dicotyl, 104, 105 of herbaceous monocotyl, 113 of woody monocotyl, 114, 115 perennial, 98 phylloid, 99 procumbent, 98 reclining, 98 repent, 98 scandent, 99 spiny, 99 subterranean tuberous, 99 suffruticose, 99 tendriliform, 99 Sterculiaceae, 309, 31 1 Sterigmata, 31 1 Sterigmatocystis niger, 210, 21 1 Stevens, 54 Stigma, 154 defined, 157 of animal pollinated flowers, 157 of wind pollinated flowers, 157 papillae of, 157 Stillingia, 303 sylvatica, 303 Stink horn, 228 Stipe, 213, 222 Stipules, 1 19, 129 axillary, 129 caducous scaly, 129 lateral, 129 modified, 129, 130 sepaline, 144 Stomata, 71 structure and development of, 134-136 384 Stomata, transpiration, 71 water, 71 Stone cells, 71 Stramonium, 339 Strawberry, 286 Streptococcus, 175, 178 erysipelatis, 179 pneumoniae, 179 Streptothrix, 177 bovis, 179 Strobile, 139, 163 Stromata, 214 Strophanthin, micro-chemic test for, Si. 52 Strophanthus, 333 hispidus, 333 Kombe, 333 Strychnine, micro-chemic test for, 53 Strychnos Ignatii, 330 Nux Vomica, 330 Style, 154, 156 arms, 156 collecting hairs of, 157 Styraceae, 328 Styrax, 286 Benzoin, 328 Sub-hymenium, 224 Succinum, 244 Succus Citri, 299 Pomorum, 288 Sucrose, 46 Sudan III, 66 Sugar cane, 46 Sugars, 45 Fliickiger’s micro-chemic test for determination of various kinds of, 46 Sumbul, 325 Sutures, 161 Swarm spores, 185 Sweet Potato, 334 Swertia Chirayita, 332 Syconium, 165 Sympetalas, 326 Symphytum, 336 INDEX Symplocarpus fcetidus, 249 Synchytrium, 199 Synergids, 159 Syringa, 328 Systematic Botany, 174-355 defined, 1 Tabacum, 339 Tamarindus, 292 indica, 292 Tanacetum, 355 vulgare, 355 Tannin, 57 Tannins, 56, 57 Tapetum, 149, 151 Tapioca, 303 Taraxacum officinale, 353 Taxonomy, defined, 1 Taxus, 243 Tea Family, 314 Tegmen, 168 Tela contexta, 224 Teleutospores, 219 Tendril, 99 Terebinthina, 244 Canadensis, 244 Laricis, 244 Ternstrcemiaceae, 314 Terpenes, 60 Terra Silicea Purificata, 190 Testa, 168 Tetrads, 151 Tetraspores, 195 Thalamus, 143 Thallophyta, 174 outline of, 3, 4 Thallus, 174 . Thamnidium, 199 Thea sinensis, 314 Theaceae, 314 Thelephorales, 221 Theobroma Cacao, 310, 31 1 Thiothrix, 117 Thistle, 352, 353 Thorn, 99 INDEX 385 Thornapple, 338 Throat, 143 Thuja, 243, 244 occidentalis, 244 Thymeleaceae, 318 Thymus Serphyllum, 344 vulgaris, 344 Thyrsus, 140 Tilia, 3x1 Tiliaccae, 309, 31 1 Tissues, 67-84 Tobacco, 337, 338 Toluifera Balsamum, 291 Pereirae, 291 Tomato, 338 Tooth Fungi, 222 Tormentilla, 289 Torus, 143 Touranose, 47 Tracheae, 78, 79 Tracheary tissue, 78 Tracheids, 80 Tradescantia zebrina, 41 Tragacanth, 59 Tragacantha, 291 Trailing Arbutus, 327 Trama, 224 Transpiration, 116 Stomata, 72 Tree, 100 Trehalase, 63 Trehalose, 46 Tremellaceae, 221 Tremellales, 220 Trichomes, 72 Trifolium N. F., 292 pratense, 292 Trigonella Foenum-graecum, 292 Trillium, 252 Tristicha, 56 Triticum, 248 True Mosses, 234 Truffles, 212, 213 Trunk, 100 Trypsin, 64 2S Tschirch, 60 Tsuga canadensis, 244 Tube, 143 Tuber, 100, 213 Tuberaceae, 212, 213 Turnera aphrodisiaca, 316 diffusa, 316 Turneraceae, 316 Turpentine Bordeaux, 24 Turpeth Root, 335 Tussilago Farfara, 354 Ulmaceae, 265 Ulmus, 265 fulva, 265 Ulothricaceae, 187 Ulothrix zonata, 187 Umbel, 138, 139 Umbellales, 323 Umbelliferae, 70, 324 Umbelliflorae, 323 Uredinales, 218 Uredinium, 218 Uredo linearis, 219 Uredospore, 218 Urginea maritima, 252 Urticales, 265 Ustilaginales, 217 Ustilago Maydis, 217 Zeae, 216, 217 Utricle, 164 Uva, 164 Uva Ursi, 328 Vaccineae, 327 Vacuole glycogen, 201 Vacuoles sap, 39 Valeriana, 351 officinalis, 349, 350, 351 Valerianaceae, 349 Valves of fruits, 161 Vanilla, 257 planifolia, 257 histology of fruit, 166 Variety, 2 Vaucheria, 191 1 386 INDEX Vaucheria, terrestris, 187 Vavaea, 301 Veil, partial, 223 universal, 225 Venation, leaf, 120 Veratrina, 252 Veratrine, micro-chemic test for, 53 Veratrum, 251 viride, 251 Verbasci Flores, 342 Folia, 342 Verbascum, 147, 340 Blattaria, 340 Thapsiforme, 342 Thapsus, 342 Verbena N. F., 345 hastata, 345 Verbenaceae, 345 Vernation, 118, 119 Veronica, 148, 340 Verticillaster, 139 Viburnum Lentago, 347 Opulus, 347 Prunifolium, 347 Vinca, 157 Vinum Xericum, 309 Viola tricolor, 72, 316 Violaceae, 316 Viscum, 37 Vitaceae, 307 Vitis, 309 Volatile oils, 60 Vouacapoua Araroba, 292 Walnut Family, 264 Waltheria, 309 Water Leaf Family, 336 Watermelon, 347 seed, 348 Water stomata, 71, 72 Weigelia, 347 Welwitchia, 36 Weymouth Pine, 24 Wheat, Fig. of plant, 246 rust, 2x8 Wheat, starch, 50 White Pine, 24-31 Willow, 260 Winter’s Bark, 272 Witchhazel Family, 285 Wood, defined, 111 fibers, 71 heart-, 113 microscopic characters of angio spermous and gymnosper mous, 1 13 parenchyma, 69 pine, 112 sap-, 1 13 Xanthophyll, 61 Xanthoxyli Fructus N. F., 299 Xanthoxylum, 299 americanum, 299 Clava-Herculis, 299 Xerophytes, 83 Xylem, 79, 83 Yam Family, 252 Yeasts, bottom, 203 defined, 200 Hansen’s classification of, 200 top, 200 Yucca, 249 Zanthoxyleae, 298 Zea N. F., 248 Mays, 248 Zedoaria, 255 Ziehl’s Carbol-fuchsin, 181 Zingiber, 255 officinale, 254, 255 Zingiberaceae, 253 Zoogloea, 94 Zoospore, 42 Zoospores, 186, 187 Zygnemaceae, 188 Zygomycetes, 196 Zygophyllaceae, 297 Zygospore, 188, 190 Zymase, 46, 47, 63 ■ . . ■ i.