- YOUNGKEN

Alexander Tschirch, Emeritus Professor of Pharmacognosy in the University of Bern,
whose brilliant investigations and numerous contributions in the fields of pharmaceuti-

cal botany and pharmacognosy during the present century have added much to our
knowledge of drugs.

(Frontispiece.)

A College
Textbook of

Pharmaceutical Botany

With Pharmaceutical and Medical
Applications, especially adapted

for the use of Students of Pharm-
acy and Economic Botany and
as a reference for Pharmacists,
Chemists and Students in Struc-

tural and Systematic Botany.
By
Heber W. Youngken, PH.M., PH.D., Sc.D.

Professor of Botany and Pharm-

acognosy in the Massachusetts
College of Pharmacy. Member of
the Committee of Revision of the
Pharmacopoeia of the United
States, Botanical Editor of the
United States Dispensatory; Au-
thor of “A Textbook of Pharm-

acognosy.””

Sixth Edition
Thoroughly Revised

S Ricksecsi’s s Son. and Co., Inc.
Philadelphia

AL.
MISSOURI! BOTANICAL -
GARDEN LIBRARY o

CopyricuT, 1938,

THE USE IN THIS VOLUME OF CERTAIN PORTIONS OF THE TEXT OF
THE UNITED STATES PHARMACOPOEIA, ELEVENTH EDITION, IS BY

BOARD OF TRUSTEES IS NOT RESPONSIBLE FOR ANY INACCURACY _
OF QUOTATIONS, NOR FOR ANY ERRORS IN THE STATEMENT OF ©
QUANTITIES OR PERCENTAGE OF STRENGTHS. en

NATIONAL FORMULARY, SIXTH EDITION, IN THIS VOLUME HAS
BEEN GRANTED BY THE COMMITTEE ON PUBLICATIONS BY

Preface to the Sixth Edition

Since the last edition of ‘Pharmaceutical Botany’? was
published, knowledge in plant science has advanced rapidly
and a new edition of the United States Pharmacopoeia and of
the National Formulary have appeared. For these reasons
another revision has been called for.

In preparing this revision the writer has endeavored to keep
in mind the requirements of varied courses in botany given to
students of pharmacy in departments of botany of both the
university schools and the independent pharmaceutical institu-
tions. In both types of schools botany should be taught with
cultural and professional objectives, and the pharmacy students
provided with the proper pharmaceutical slant which need not
detract from adequate treatment of the purely academic phases
of the subject.

A revision has, accordingly, been prepared which presents a
broad general treatment of botany with particular stress upon
those phases of this science such as histology, cell contents, tax-
onomy, and medicinal plants which are believed to be most
helpful in preparing students to cope with botanical problems
in pharmacognosy and in the practice of pharmacy.

In the course of this revision the writer has not only made
changes in the text so as to bring the subject matter up to date
but has reorganized some of the chapters, provided some better
illustrations and added 121 new figures.

The old chapter on Cytology has been deleted and in its
place new chapters have been substituted on “The Living Cell”
‘and on ‘“Non-protoplasmic Cell Contents.” Important ad-
ditions have been made to the subject of the cell and its contents

including an amplification of the subject matter on mitosis and
vii

vill PREFACE TO THE SIXTH EDITION

new material on plasmodesma, waxes, plant hormones and
vitamins.

A new chapter has been added on “Genetics and Evolution”
in keeping with the needs of some of the courses in botany.

The chapters on “The Microscope’ and on “Histological
Technique” have been transferred from their former positions
in the main body of the text to the back of the book where they
now appear as Appendix I and Appendix II respectively. New
material has been incorporated under both of these captions |
including modern methods of fixation, staining, imbedding and —
mounting plant materials for microscopic examination. &

The chapter on “Plant Tissues” has been lengthened and |
improved by the introduction of much new material on the
structure and physiology of meristems, endodermis, tracheary
tissue, laticiferous tissue, xylem, phloem, the stele, bundles,
leaf and branch traces, gaps and nectaries.

The histological and physiological portions of the various
chapters dealing with Plant Organs have also been augmented i
with considerable new text and illustrative material. pe

The Glossary has been retained and somewhat enlarged as
has also the Classified List of Reference Works. A complete
Index is also provided, making the data desired by the reader
easily found. s

Many new drawings, photographs and photomicrographs
have been added in this revision which should prove helpful to
the student. A considerable number of these are original;
many have been borrowed. ae

The writer desires to express his gratitude to the various
publishers and authors from whom permission to use a number
of the cuts used in this edition was cheerfully granted. Acknowl-
edgment for these is given under the respective cuts in th
text. He is particularly indebted to Dr. A. F. Sievers and the
Bureau of Plant Industry for the loan of a number of prints on
medicinal plants from which cuts were prepared, to Instructor.
Richard W. St. Clair of the Massachusetts College of Pharmac
for valued assistance in photographing many of the original
figures, to Dr. F. J. Bacon of the Biology Department of Western

Reserve University who furnished photographs on Zamia plan

ee ee ee a

PREFACE TO THE SIXTH EDITION ix

and to Asst. Professor H. L. Reed of the Massachusetts College
of Pharmacy who contributed the prints from which Figs. 138,
139 and 140 were prepared.
Finally, the author wishes to express his appreciation to
P. Blakiston’s Son & Co. Inc. for doing its utmost in producing an
attractive and well bound book.
HEBER W. YOUNGKEN.

Bosron, Mass.,
June, 1938.

Preface to the Fifth Edition

Since the fourth edition of this volume was issued, there have
been notable advances in the development of pharmaceutical
botany. A new United States Pharmacopceia and a new
National Formulary have appeared. The Commonwealth
Fund Educational Research Committee has rendered its report
on the study of Pharmacy from the functional standpoint and
in it the findings of its botanical studies.

These events alone made some additions and alterations of
the text necessary. But the writer has gone further. A large
part of the text has been rearranged, rewritten and amplified
with a view toward improving the clarity of certain portions of
the subject matter, and extending its adaptability.

More than a hundred pages of new matter and one hundred
and twenty-four new figures comprising original drawings and
photographs and borrowed cuts from classic works have been
added, and some of the older illustrations have been replaced
by better ones.

The botanical findings of the Commonwealth Fund Educa-
tional Research Committee have been incorporated, although
the text is not limited to these.

Sufficient new information has been added to make the book
a distinct improvement over the former editions, and this should
make it better for use in professional pharmaceutical colleges
and for the courses in botany now prescribed for Pre-medical
students in academic colleges, as well as for students in Bachelor
of Science courses in colleges of pharmacy.

The table of contents clearly indicates the changes made in
the form of the text which now contains twenty-five chapters; an
increase of sixteen over the previous edition.

It is believed the increased number of chapters and their
new sequence will be found convenient for teachers in assigning
work alike for study and collateral reading.

It is hoped that the new edition will also be of value to
research students in various fields of botany, since it furnishes in
the classified list of reference works a summary of a large
amount of literature of the subject. HOW. Y.

x1

Preface to the Fourth Edition

The value of a thorough systematic training in Botany with
emphasis upon the structural and taxonomic aspects of this
science is now generally recognized as a prerequisite to the
proper comprehension of problems in pharmacognosy and
materia medica. But apart from the direct relation of such
training to these and other applied fields, Botany per se stands
out preeminently as a cultural subject and, when presented
with laboratory work, is of great value in developing, in students,
the power of observation.

In preparing the new edition of this text-book, the author has
primarily kept in mind the needs of the pharmaceutical student
but he has not been unmindful of its use by students of academic
colleges and accordingly has increased the scope of the work.

Fifty-three new pages of subject matter and 25 illustrations
have been added. Chapter I on “Fundamental Considera-
tions” has been augmented by the inclusion of a list of standard
micro-chemical reagents used in food and drug work together
with formule for their preparation. Chapter VII on “Plant
Organs and Organisms” has been lengthened by 23 pages of
new subject matter. Among the outstanding features of its
revision may be mentioned the introduction of considerable
more plant physiology as well as new drawings and photographs,
most of which are original.

There has been an amplification of the subject matter of
roots, rhizomes, leaves, seeds and seedlings including data on
soil and water relation, diffusion, osmosis, osmosis applied to
root hairs, turgor, plasmolysis, essential and non-essential
elements absorbed by root hairs, transpiration and the histology
of a type of Monocotyl rhizome. New plates illustrating the
gross structure of Monocotyl and Dicotyl seeds and of seedlings
are introduced for the first time. Chapter IX on Ecology has

XL

PREFACE TO THE FOURTH EDITION

been lengthened by 20 pages and several illustrations on Carniv-
orous plants.

Every part of the work has been carefully revised and brought
uptodate. As in former editions, the index has been so planned
as to make the information contained in this book readily
accessible.

To the authors from whom cuts were borrowed, the writer’s
thanks are due.

Preface to the Third Edition

The author has first endeavored to present in a clear, syste-
matic way those fundamental principles of structural and
taxonomic botany which serve as a key to the approach of
pharmacognic problems. But he has not been unmindful that
the use of the work has extended to academic institutions and, ©
so, in this edition, has broadened the scope of the former text.
To this end about ninety additional pages of subject matter
have been introduced. Several old cuts have been removed.
Forty-three new ones have been inserted. Hypothetical dis-
cussions have been avoided which saves time for the reader.

The arrangement and plan of the chapters are similar to that
of the former edition, in order to adapt the work to several
methods of approach. Chapter I on “Fundamental Considera-
tions” has been augmented by treatises on Botanical Nomen-
clature, Paraffine and Celloidin Imbedding, Sectioning, Staining
and Mounting, Microtomes and other information dealing with
the preparation of materials for microscopic examination.

Ten pages have been added to Chapter V on Cytology.
Under ‘‘Protoplasm and its Properties,” six pages have been
written on the subject of Irritability and Irritable Reactions.
Under ‘“‘Non-Protoplasmic Cell Contents” several additional
commercial starches are discussed and two original plates on
starch grains added. Additional cuts on Collenchyma, Stone
Cells, Sclerenchyma Fibers, Trichomes and Fibrovascular
Bundles have been inserted in Chapter VI.

Nine additional pages of subject matter and illustrations have
been added to Chapter VII. Original figures of all of the
important types of fruits appear here for the first time.

Chapter VIII on ““Taxonomy”’ has been increased by seven
pages of new data, and the whole former text carefully revised.

Chapter IX on ‘‘Ecology”’ has been newly introduced as has
also a Glossary of Botanical Terms. The index has been so
planned as to make the information contained in this book
_ readily accessible.

To the authors of works from which cuts were borrowed the
writer’s thanks are due. Ev. X-

XV

Contents

PREFACE TO THE SIXTH EDITION.

PREFACE TO: THE. FIFTH EDITION 6 ins vos des
PREFACE TO THE FOURTH EDITION. ............
PREFACE TO THE THIRD EDITION. .........

CHAPTER I
Tue Scope AND IMPORTANCE OF BOTANY
EN TRODUCEION 250 = ae oe aan
DEPARTMENTS OF BOTANICAL INQUIRY.—1. Plant parte
ogy (Gross Anatomy, Histology, Cytology). 2. Plant Embry-
ology. 3. Plant Physiology. 4. Taxonomy of Plants or
Systematic Botany. 5. Plant Ecology. 6. Plant Genetics. 7.
Phytopathology. 8. Phytogeography. 9. Phytopalzontology.
10. Cryptogamic Botany. 11. Phanerogamic Botany. 12.
Algology. 13. Mycology. 14. Saad 15. Economic Bot-
any and its subdivisions. . . . reer dee ae ar a
THE IMPORTANCE COP BOPEANY 2PM sks ee a!

CHAPTER II
Tue GENERAL CHARACTERISTICS OF PLANTS
THE PLANT AS A LIVING ENTITY.—Organism, defined. Organ,
defined. Vegetative and reproductive processes of plants.
Parts of the plant body. Work of the root, stem, green leaves,
flowers, fruits and seeds. Tissues. Cells. Protoplasm.
GROUPS OF PLANTS.—The Thallophytes, Bryophytes, Pterido-
phytes and Spermatophytes, their etymology, leading sub-

groups and introduction to their habits. Cryptogams. —

Phanerogams. Origin and development of a seed plant.
Physiological division of labor. Differentiation. ..... .

CHAPTER III —
Tue Living CELL

THE CELL AS THE FUNDAMENTAL UNIT.—History of the cell

and protoplasm. (Hooke, Brown, Schleiden, Schwann, Von
Mohl, Corti, Treviranus, Dujardin). The Protoplast. A
typical colorless Plant Cell. Typical green cells. oe

ee Oe We is a a ee ee ON

2-4
4-5

6-12

12-16

XVill CONTENTS

THE AMG:BA. : Nee

THE RELATION OF PLANTS AND ANIMALS .

PROTOPLASM AND ITS PROPERTIES.—Structure. Metabolism.
Growth. Irritability. Tropisms, Taxies and Nasties. (Ther-
motropism, Chemotropism, Sitotropism, Oxytropism, Hydro-
tropism, Rheotropism, Heliotropism, Phototropism, Geotro-

pism, Galvanotropism, Thigmotropism). Reproduction.
Protoplasmic Movement .
PROTOPLASMIG CELL CONTENTS.—The protoplast. Cyto-

plasm; nucleus; nucleolus; plastids (proplastids, leucoplastids,
chloroplastids, isp Amaia ay cell membranes; chondriosomes.
Plasmodesma.

CELL FORMATION AND REPRODUCTION.—Asexual and sexual
cells. Reproduction defined. Asexual Reproduction: Fis-
sion; Gemmation; Free Cell Formation; Rejuvenescence or
Zoéspore Formation; Vegetative Multiplication. Sexual Repro-
duction: Conjugation and Fertilization. . ........

INDIRECT NUCLEAR DIVISION .
MEIOSIS (Reduction division)... . . ste we ee ee

CHAPTER IV

Lire Hisrory oF THE Mate Fern [AN INTERMEDIATE TYPE OF se :

MEDICINAL USE. PHASES INITS LIFECYCLE ..... ‘

HISTORY OF THE SPOROPHYTE OR ASEXUAL GENERATION.
—Gross structure of the stem. Gross structure of the frond.
Histology of mature stem. Histology of growing stem apex.
Histology of mature root. Histology of root apex. Con-
ngersl of crude sap flow. Histology of stipe. Histology of

a pinnule. Comparative physiology of root, stem and leaf.
Gross structure and histology of the sori and sporangia. eas
ture of sporangium and spore dissemination. . . .

ee ee rose

HISTORY OF THE GAMETOPHYTE OR SEXUAL GENERA-
TION.—Formation of protonema and _prothallus.
of new sporophyte or diploid plant from fertilized egg. Growth
of embryo into mature sporophyte. Alternation of generations

CHAPTER V
NoOn-PROTOPLASMIC CELL CONTENTS

PROTOPLAST INCLUSIONS.—Sugars. Starch (Assimilation, Tran- |

sitory and Reserve). Structure and Composition of Starch.

Method of Examining Reserve Starches. Characteristics of

Important Commercial Starches. Dextrin. Amylodextrin.
Inulin. Characteristics of | Glucosides. The glucosides

Hesperidin, Strophanthin, Salicin, Saponins, Coniferin and |
~ Digitoxin. Cyanogenetic glucosides. Pentosides. Alkaloids and —

Origen te:

PAGES
24-29

29-30

30-40

53

CONTENTS

their properties. The alkaloids, Strychnine, Brucine, Nicotine,
Caffeine, Cocaine, Aconitine, Morphine, Codeine and Col-
chicine. Gluco-alkaloids. Asparagine. Calcium Oxalate.
Micro-crystals. Cystoliths. Silica. Tannins. Proteins.
Aleurone Grains. Mucilages and Gums. Fixed Oils and Fats.
Waxes. Volatile Oils. Resins. Oleoresins. Gum _ Resins.
Balsams. Pigments (Chlorophyll, Carotin, Xanthophyll,
Etiolin, Anthocyanins, etc.) Latex. Enzymes. Classification
of Enzymes. Plant Hormones (Auxins). Vitamins... . .

CELL WALLS.—Their formation and composition. Growth in area

and thickness. Pores (Pits). Various kinds of cell walls and
behavior of each to micro-chemic reagents .

CHAPTER VI

PLANT Tissues

PLANT TISSUES.—Tissue defined. The Tissues of Spermatophytes
and Pteridophytes: Classification of tissues. Primary and
secondary tissues. Simple and complex tissues. Permanent

tissues. Primary Meristems. List of Tissues.

MERISTEM.—Definition. Primordial meristem. Primary meristems
(dermatogen, periblem and plerome) secondary meristems
(cambium and phellogen). Apical, recs and lateral
meristems. Their distribution . rea ee SOS ara

PARENCHYMA.—Definition; ordinary ae chlorenchyma or
assimilation parenchyma; conducting parenchyma; reserve
parenchyma; their structure, distribution and function. . .

COLLENCHYMA.—Definition, structure, function and distribution . .
SCLERENCHYMA.— Definition; stone cells; sclerenchyma fibers; wood
fibers; phloem- or bast-fibers; pericyclic fibers; cortical fibers;
their distribution 6 5 3 4 ee ae. PEL en
EPIDERMIS.—Definition; transpiration and water stomata; epidermal
papilla; trichomes; scales; their distribution and functions. .
ENDODERMIS.—Definition, structure, distribution and functions.
Caliparian pO. 6 5 rh Be aa ely oaks
CORK.—Definition; its derivation, function and distribution. Rhy-
dome. Oke 3 ee ee ba at ae
PERICYCLE.—Derivation, structure, position and function .
LATICIFEROUS TISSUE.—The structure, origin and distribution of
, latex cells, laticiferous vessels and coenocytic latex cells. Latex.
SIEVE (LEPTOME) OR CRIBIFORM TISSUE.—Definition; deriva-
tion; structure; distribution; Panchen. she cS My ela

-TRACHEARY TISSUE.—Trachez: Definition, origin, structure, func-
tion and classification; Tracheids: Definition, structure and
function. Distinction from trachee. . .-..-- -

xix
' PAGES

74-107

THT lO

«4144-114

. 114-116

. 16-417

117-118

s416-123

2 122-127.

© F27=129

= Lee

131

131-133

+ 133-135

7195-1)

XX ‘ CONTENTS

MEDULLARY RAYS.—Primary and Secondary; their structure, distri-
bution and functions. . . . . OP eae ty Eas wea Ei

VASCULAR AND FIBRO-VASCULAR BUNDLES.—Description of
the various types; the distribution of each type; Xylem and
Phloem. Leaf and branch traces. Recess

THE STELE.—Definition. Prostele. Siphonostele. Dictyostele.
Branch Traces. Leaf and Branch Gaps. ......-.--

SECRETION SACS.—Definition and distribution. . . . .....

INTERCELLULAR AIR SPACES.—Definition. Function. Schizo-
genous and lysigenous air spaces; their relative size. . .

INTERNAL SECRETION GLANDS AND RESERVOIRS.—Defini-
tion; structure; internal glands; secretion canals; vitte. Their
origin . pei mae

PC AE i oe ee eg ee ee oe pe Ralls ie i

CLASSIFICATION OF TISSUES ACCORDING TO FUNCTION .

CHAPTER VII
Tue Roor

THE ROOT.—Definition; functions; rootlets; root hairs; piliferous layer;
soil and water relation, diffusion, osmosis, osmosis applied to root

hairs, turgor, plasmolysis, absorption of nutrient salts, essential
elements and their importance, non-essential elements absorbed

by root hairs; regions of the root (root cap, embryonic region,

growing point, primary meristems or generative tissues, region

of elongation, region of maturation and mature region). Differ-

ences between root and stem . . .

CLASSIFICATION OF ROOTS AS TO ORDER.—Primary root; tap
root; secondary roots; fibrous and fleshy roots; prop roots; adven-
titious roots; epiphytic roots; haustoria. ;

CLASSIFICATION OF ROOTS AS TO FORM. ee napiform,
conical fibrous, tortuous, nodose, moniliform, tuberous, fascicled

ROOT CONSISTENCY OR TEXTURE. et roots. Fleshy roots:
eRe tiie THEO ee te ae ot oe

CLASSIFICATION OF PLANTS ACCORDING TO DURATION
OF ROOT.—Annual; Biennial; Perennial . . .......

ROOT HISTOLOGY.—A, Monocotyledons: Histology of Similax root.
B. Dicotyledons: Histology of Dicotyl roots: a. young growth, B.
transitional growth and development of secondary structure.
Histology and Development of the California Privet root.
Abnormal structure of Dicotyl roots. shits of a Neots
Tuberous Root (Aconitum). Root branching.

ROOT TUBERCLES.—Definition; occurrence. | oie tubercles.

Rotation of crops. e® tubercles; their etiology. Nitrogen
CVE ko as 4

sh SR Aes act MEN eR a es eee ey ee pee ahi? ov og aes

PAGES

. 139-140

. 140-144

. 144-147

147

. 147-148

. 148-149 -

150
151

. 151-160

. 160-161

161-162

162

162-163

. 163-176

176-182

CONTENTS

MYCORRBIZA: 4c OG

CHAPTER VIII
Tue Bup

BUDS.—Definition; plumule, scaly buds, naked buds; Classification of
Buds According to Position on Stem: terminal bud; axillary or

lateral bud; adventitious bud; accessory bud; dormant bud;
sub-petiolar bud. Classification of Buds According to Develop-
ment: leaf bud; flower bud and mixed bud. Classification of

Buds According to their Arrangement on the Stem: alternate;
opposites whore. O66 Sr eg hae es ee

CHAPTER IX

Tue STEM
- THE STEM.—Definition; direction of growth; functions; size; nodes
and internodes; stem elongation; generative tissues; duration of
stems; forms of stems; stem modifications; above ground stems;
herb, shrub and tree defined; fall of leaves; underground stems
(rhizomes, bulbs, corms, tubers, suckers) exogenous and endo-

genous henaae Sia ee, eae eee eects ee Rn ay eae cag ON Ree Bae

STEM HISTOLOGY.—Histology of Annual Dicotyl Stem, Growth of
Perennial Dicotyl Stem and its Histology. Exceptional Types
of Dicotyl Stems; Lenticels and Their. Formation; Annual
Thickening of Stems; Method of formation; “Annual Ring.”
Bark: Definition; zones; Cork; Periderm; Phelloderm; Histology
of Cascara Sagrada bark. Wood: alburnum; duramen; tyloses;
microscopic characteristics of Angiospermous and Coniferous
woods. Histology of Typical Herbaceous Monocotyl Stems.
Histology of a Typical Monocotyl Rhizome. Histology of a
Typical Woody Monocotyl Stem. Histology of Grass Stems.
Secondary Growth in Monocotyl SOMA g 6 es pe oe

CHAPTER X
Tue Lear

THE LEAF.—Definition; the complete leaf and its structure; sessile,
petiolate, stipulate and exstipulate leaves; ieaf functions: photo-
synthesis; assimilation; r piration; transpiration. . - .. -
TYPES OF LEAVES DEVELOPED IN ANGIOSPERMS.—Cotyle-
dons; Scale leaves; Foliage leaves; Bract leaves: bracts and
bracteoles; Sepals; Petals; Microsporophylls; Megasporophylls .
ORIGIN AND DEVELOPMENT OF LEAVES.—Primordial leaf. Its
Bist oe ee is mies Nt at ty? Rot
-PHYLLOTAXY.—Definition; spiral law of leaf arrangement; alternate;
: opposite, decussate; whorled; fascicled; leaf rank. . . - - pat
- VERNATION.—Definition; jnflexed or reclinate; conduplicate; con-
volute; circinate; plicate; involute; revolute. . . +--+ +++

184-185

. 186-194

. 194-220

, Sel-227

XXil CONTENTS

LEAF VENATION.—Furcate; parallel; reticulate; pinnately-veined;
palmately-veined; anastomosing, tessellated and impressed veins

LEAF INSERTION.—Definition; radical; cauline; ramal; sian
amplexicaul; connate-perfoliate; equitant. . . ...-. - :

FORMS OF LEAVES.—Simple and Compound. Forms of the Lamina:
(a) General Outline; linear; filiform; ovate, obovate, rhom-
boidal; lanceolate; elliptical; oblong; equilateral; inequilateral;
reniform; orbicular; peltate; oblanceolate; cuneate; spatulate;
ensiform; subulate; acerose; falcate, deltoid. (4) Apex: acute;
acuminate; tapering; attenuate; obtuse; rounded; truncate;
mucronate; cuspidate; aristate; emarginate; retuse; obcordate.
(c) Base: cordate; reniform; hastate; auriculate; cuneate; peltate;
Sagittate; oblique; acute, tapering, attenuate, truncate; obtuse,
rounded, decurrent, etc. (d) Margin: entire; revolute; serrate;
serrulate; dentate; denticulate; crenate; crenulate; doubly ser-

_ rate, etc.; spinose, ciliate; sinuses, segments and lobes; repand;
sinuate; incised; runcinate; lobed; cleft; parted; divided; pin-
natifid; pinnatipartite; pinnatisect; palmatifid; palmatipartite;
palmatisect; pedate; interruptedly-pinnate; lyrate. Forms of
Compound Leaves . ere er

LEAP MODIVICA PIONS 23 tet Pe et

LEAF TEXTURE.—Membranous; succulent; coriaceous. .

LEAF COLOR.—Variations in color .

LEAF SURFACE.—Glabrous; glaucous; pellucid-punctate; papillose;
scabrous; pubescent; appressed-hairy; villose; sericious; hispid;
strigose; tomentose; sei rugose; verrucose; tuberculate;
bullate .

DURATION OF LEAVES. es | or evergreen; 5 ded cadu-
cous; fugacious

GROSS STRUCTURE AND HISTOLOGY ¢ OF THE LEAF PETI-
OLE.—In Monocotyledons. The ligule and pulvinus. In
Dicotyledons. Pulvinus. Aggregation Body. Pericladium.
Phyllode. Pe ee ee ere ee en ea TY

STIPULES.—Definition; eer free-lateral; lateral-adnate; lateral-
connate; lateral interpetiolar. The ligule. Axillary: free
axillary and connate. The ochrea. Modified Stipules. . .

THE LAMINA.—Definition. Mode of Development of the Lamina
of Leaves: Dorsiventral; convergent; centric; bifacial; reversed
ob-dorsiventral. A. Dorsiventral: (a2) Dorsiventral Umbro-
phytic; () Dorsiventral Mesophytic; (c) Dorsiventral Xero-
phytic; (¢) Dorsiventral Hydrophytic. Gross Structure and
Histology of Different Types of Dorsiventral Leaf Blades. Gross
Structure and Histology of the following types: B. Convergent;
C. Centric; D. Bifacial . Pe ee ee ae ee Y

STRUCTURE AND DEVELOPMENT OF STOMATA .

pra oak yc? Mee at “hag sand ay Sey See

PAGES

230-232

232

. 233-240
. 240-241

241
241

. 241-242

242

242-245

. 245-246

. 246-251
« 251-254

CONTENTS

HISTOLOGIC DIFFERENCES BETWEEN LEAVES OF DICOTY-
LEDONS AND MONOCOTYLEDONS ..

CHAPTER XI
THE INFLORESCENCE
_INFLORESCENCE.—Definition. Determinate, Indeterminate and
Mixed Inflorescences defined. Parts of Inflorescences: peduncle,
rachis, pedicel, bracts, involucre, bracteoles. Scape. Spathe.

INDETERMINATE INFLORESCENCES.—Elongated and flat-topped.
Forms of: Solitary indeterminate; raceme; corymb; umbel;
compound umbel; spike; compound spike; catkin; head; strob-
ile; spacing DAMIR 3. Ss es i

DETERMINATE INFLORESCENCES.—Elongated and flat-topped.
Forms of: Solitary determinate; cyme; simple and dichasial
cymes; scorpioid cyme; glomerule; verticillaster . .

MIXED INFLORESCENCES.—Thyrsus; raceme of cymes; corymb
OR-GPIORS 65 oi a ee tg a ee es

CHAPTER XII -
THE FLOWER
THE FLOWER.—Definition; floral parts; essential organs; accessory
organs; complete; perfect; hermaphrodite; regular or actino-
morphic; irregular; zygomorphic; symmetrical; imperfect;
double; staminate; pistillate; neutral; hypogynous; perigynous;
Epigynous. Moncecious, dicecious and polygamous plants .
PREFLORATION: Definition. Convolute; involute; revolute; plicate;
imbricate; valvate; vexillary; contorted. ..........

CONNATION AND ADNATION.—Definitions and illustrations .

THE RECEPTACLE, TORUS OR THALAMUS.—Definition; varia-
tions in structure; anthophore; gonophore; gynophore;
COFDUNE cs ate dete ce ge ks yee sear Nags tens

THE PERIANTH.—Definition; dichlamydeous; monochlamydeous.

THE CALYX.—Definition; parts; physical characteristics; connation;
kinds and forms; persistence; adnation; sepaline nectaries and
spurs; sepaline stipules; the epicalyx; sepaline position .

THE COROLLA.—Definition; parts; physical characteristics; func-
tions. Forms of the Corolla and Perianth. Appendages of
Oe i ee ye ne tee ee

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. Another Dehiscence.

XXiil
PAGES

255

256-257

257-259

: 259-261

261

. 262-264

264-265
265

. 265-266

266

. 266-269

. 269-274

XXIV CONTENTS

PAGES
Development of the Anther. Attachment of Anther. Pollen;
description; forms. -Pollinia. Hay Fever pollens. . . . . 274-283

THE GYNC@CIUM 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 transformed
buds; position of ovules in ovary; Gymnospermous and Angio-
spermous ovules; structure of Angiospermous ovule; forms of
. did gcng, et ee et a ee mes amare Tae 283-287

THE PLACENTA.—Definition; types of placenta arrangement . . . . 287-288

THE STYLE.—Definition; style-arms; relation to carpels forming the
gynoecium; variations from typical stylar development . . . . 288-289

THE STIGMA.—Definition; forms in wind- and animal-pollinated

Rowers: stigmatic papillie s: o9o6). eh: Wag RE eG 290
POLLINATION.—Definition. Close and Cross Pollination. Pro-

tandrous and Protogynous flowers. Terms Byte to plants

pollinated by various agencies. Me ees sine Sy 7 . 290-292
MATURATION OF THE POLLEN GRAIN AND FORMATION OF

MALE GAMETOPHYTE (in Angiosperms) . ..... . . 292
MATURATION OF THE EMBRYO SAC AND FORMATION OF

THE FEMALE GAMETOPHYTE (in Angiosperms) . a 292
FERTILIZATION IN ANGIOSPERMS.—Process; formation of ee

and endosperm; double fertilization . . . . . . ae a A
DEVELOPMENT OF THE EMBRYO.—1. In Dicotyledons 2 ig Re

emnrreny period Se oh Se 295-296

CHAPTER XIII
Tue Fruit

THE FRUIT.—Definition; modifications. . . . . . . . SE rb is. 297
FRUIT STRUCTURE.—Pericarp; pseudocarp; anthocarp; epicarp;

mesocarp; endocarp; sarcocarp; putamen; sutures; valves;

dehiscent and indehiscent fruits; kinds of dehiscence . . . . . 297-300 —

CLASSIFICATION OF FRUITS.—Simple; Aggregate; Multiple; dry
dehiscent; dry indehiscent; fleshy indehiscent. Forms of
Simple Fruits: 1. Capsular (follicle, legume, loment, capsule,
pyxis, regma, siliqua, sillicule). 2. Schizocarpic (carcerulus,
cremocarp, samara). 3. Achenial (akene, utricle, caryop-
sis,nut). 4, Baccate (berry, uva, bacca, pepo, hesperidium). 5.
Drupaceous (drupe, pome). Aggregate Fruits: Etario. Forms
of Multiple Fruits: strobile or cone; sorosis; syconium; galbulus. 300-307

HISTOLOGY OF A CAPSULE, VANILLA.

OM ag scien Ree

CONTENTS XXV
PAGES
CHAPTER XIV
THe SEED
THE SEED.— Definition; The Embryo and its parts; vitality of seeds; seed
structure; functions; appendages including the coma, awn,
caruncle, etc.; aril and arillode; kernel, funiculus; hilum; raphe. 311-314

MODE OF FORMATION OF DIFFERENT TYPES OF ALBUMEN:
Perispermic; endospermic; perispermic and _ endospermic.
Perisperm; Endosperm; exalbuminous and albuminous seeds. 314-315

A MONOCOTYL SEED.—Its gross structure and histology . . . . . 315-318

GERMINATION OF THE SEED.—Germination of Indian Corn and
structure 08 the Sernunie ec ee ee ay RN SED

A DICOTYL SEED.—Its gross structure and histology; germination of
GICOEY) BeCe me Ss Ce apne hy cast ge ee, ts 0 Cie. DI Sae

CHAPTER XV
Tue CLASSIFICATION AND NAMING OF PLANTS
PRINCIPLES OF CLASSIFICATION.—The modern classification of
plants and its basis. Organic evolution. Natural system.
Artificial system. Artificial systems of Theophrastus, Fuchs,
Bock, Czsalpinus, Bauhin, Ray, Tournefort and Linnaeus.
The binomial system. Natural systems of De Jussieu, De
Candolle, Bentham and Hooker, Eichler, Engler and Prantl, and
Hutchinson. The groups of a modern natural system.
Complete classification of a medicinal plant, Aconite. . . . . 323-328

COUT EINE Oe Pa re ee a ae a se ae 328

BOTANICAL NOMENCLATURE.—Practice of Pre-Linnean bota-
nists. The binomial system of Linnaeus. Rules for the naming
i of genera, species, varieties, families, orders, classes, sub-classes,
3 divisions and sub-divisions. Names of Cryptogram groups. . 329-332

.

CHAPTER XVI
THe THALLOPHYTES

GENERAL CHARACTERISTICS OF THALLOPHYTES .... . 333

ALGAE.—Characteristics of group . . ........- pete it SOOO oe

CYANOPHYCEA (BLUE GREEN ALG@).—Their characteristics.
Ghaocaréas Siechiatoria; Dlemtee ke 334-336

CHLOROPHYCE& (GREEN ALG).—Their characteristics. Orders
of green algae and life histories of representative forms ofeach. . 336-348

BACILLARIEZ (DIATOMS).—Characteristics, distribution and

: BODO Rec yas 6k es eh ees hw eae 348-351
- PHAEOPHYCEA (BROWN ALGA:).—Their characteristics and
economic uses. Life history of a filamentous type, Ectocarpus.

The kelps. The rockweeds. Life history of Fucus. . . . . 351-357

Xxvl CONTENTS

PAGES
RHODOPHYCE# (RED ALG@).—Their characteristics. Chondrus,
Pant and Maa iouine ye aes Se i ae rn a. S57 AGL
CHAPTER XVII
Tue Funct
PUNGE-—T heir charatteristics'and classes! 3 32 = a es ay 362

SCHIZOMYCETES (BACTERIA).—Definition and characteristics.
Classification according to the phenomena they produce.
Fermentation, decay and putrefaction. Bacterial cultures.

Forms of bacteria. Motility. Nutrition. Nitrification. Den-
trification. Growth Products. Reproduction. Morphology
due to cleavage. Forms of cell groups after cleavage. Sporula-
tion. Rapidity of growth and multiplication. Chemical com-
position. “Exotoxins and endotoxins. . ... . + . . + 49s) egGe oo0

MORPHOLOGICAL CLASSIFICATION OF BACTERIA.—The
lower bacteria. The higher bacteria. Pathogenic bacteria.
Some bacteria producing disease in man, the lower animals and
1h Piniia ace pee ora fe oo, oF eee eee . . 370-374

MYXOMYCETES (SLIME MOLDS). —Their characteristics. . . . 374-375
EUMYCETES (TRUE FUNGI).—Their characteristics and sub-classes 375-376

PHYCOMYCETES (ALGA-LIKE FUNGI).—Their characteristics.
The orders Zygomycetales and Odmycetales. Cultivation of
Molds. Life history of the Black Bread Mold. Related molds. 376-381

ASCOMYCETES (SAC FUNGI).—Their characteristics. Order
Protoascales. Family Saccharomycetacee. The Yeasts.
Life history of a yeast, Saccharomyces cerevisiae. Top and
bottom yeasts. Wine yeast. Other yeasts. The orders
Pezizales, Plectascales, Tuberales, Helvellales, Pyrenomycetales.
Characteristics of the cup-fungi, blue and green molds, truffles,
saddle-fungi and black-fungi. Production of citric acid by molds.
Pife history of Clavicepe purpurea: cs 3 5 ., 381-397 |

BASIDIOMYCETES (BASIDIA FUNGI).—Their characteristics. The
Lower Basidiomycetes. 1. The Ustilaginales or Smuts: their
characteristics. Life history of corn smut. 2. The Uredinales
or Rusts: Characteristics. Life history of wheat rust. The
Higher Basidiomycetes: Their characteristics and sub-groups,
1. Hymenomycetes: weeping fungi, exobasidiales, coral-fungi,
mushrooms and toadstools including tooth-fungi, pore-fungi,
and gill-fungi. Groups of Agaricacez based on color of their
spores. Spore prints. Life history of the Common Field
Mushroom and of the Fly- and Deadly-. ic. Antidote for
poisoning by the Fly Agaric. 2. Gasteromycetes: puff-balls, ae
earth-stars, nest-fungi and carrion-fungi. . . . .... . . 397-411.

FUNGI IMPERFECTI.—Their characteristics, and forms producing :
discents in Mab OR RO ee a ee pe PS

CONTENTS XXVI11

PAGES
CHAPTER XVIII
Tue LicHENS
LICHENS.—Definition. Asco-lichens. Basidio-lichens. _Foliaceous,
crustaceous and fruticose lichens. Structure of a typical lichen.
Reproduction. Economic uses of various lichens . . . . . . 413-417

CHAPTER XIX

THE BrYOPHYTES
BRYOPHYTES.—Their general characteristics. . .... ...-.. 418
HEPATIC OR LIVERWORTS.—Their characteristics and orders. . 418-419

MARCHANTIA (A TYPICAL LIVERWORT).—Life history and
Structure, So es a ee a ae i ee ee ce 5 AISA Z3

MUSCI OR MOSSES.—Their characteristics. Sphagnales or bog
mosses. Andraeales or rock mosses. Bryales or true mosses.. . 426-430

LIFE HISTORY OF A TYPICAL. TRUE MOSS (POLYTRICUM). 426-430

CHAPTER XX
Tue PTERIDOPHYTES

PTERIDOPHYTES.—Their general characteristics and classes. . . . . 431
THE FERNS.—Their characteristics. Eusporangiate and _ Lepto-

sporangiata. True ferns. Waterferns........ . . 431-437
THE HORSETAILS.—Their characteristics. Life history of Equi-

setum arvenses, = 335 ee eee pote . 437-439
THE CLUB MOSSES.—Their characteristics. Life history of Lycopod-

ium clavatum. The Little Club Mosses and Quillworts. . . . 439-442

CHAPTER XXI
Tue SPERMATOPHYTES

SPERMATOPHYTES.—Characteristics and subdivisions. . .... . 443

THE GYMNOSPERMS.—Their characteristics and sub-groups. Life
history of a Gymnosperm (Pinus Strobus). The Cycads or fern
palms. Life history of Zamia. The Ginkgos. The Conifers.
Classification of Conifers. The Pinacez. Table of official and
unofficial drugs yielded by the Pinacee, including official names,
parts used, botanical origins and habitats. The Taxacee and
representatives. The Gnetales. The Gnetacee. Welwitchia,
Gnetum and Ephedra. The drug Ephedra and its uses. . . . 443-471

THE ANGIOSPERMS.—Their characteristics. Life history of an
Angiosperm (Erythronium). Resemblances between Gymno-
sperms and Angiosperms. Fundamental differences between
Gymnosperms and Angiosperms. Classes of Angiosperms. . . 471-480

XXViil CONTENTS

PAGES
CHAPTER XXII
Tue MonocotyLeDONs
MONOCOTYLEDONS.—Their characteristics. . . . . ... . . . 481-483

DRUG YIELDING MONOCOTYLEDONS.—The orders and families
of monocotyledons yielding drugs with characteristics of the
families. Tables of official and unofficial drugs with names of
drugs, parts used, botanical origins and habitats. . . . .

CHAPTER XXIII
Tue DicorTyLepons
DICOTYLEDONS.— Their characteristics... Ss a

DRUG YIELDING DICOTYLEDONS.—The sub-classes, orders and ©
families of Dicptyledons yielding drugs. Tables of official —
and unofficial drugs under each family description, with the —
drug names, parts used, botanical origins, and habitats. . . . 499-61!

CHAPTER XXIV
EcoLocy
INTRODUCTION.—Ecology defined. Environmental factors.

PLANT COMMUNITIES AND ASSOCIATIONS.—Definitions.
Classification based on relation plant associations have assumed
in regard to water. Characteristics of Hydrophytes, Helophytes,
paca de Xerophytes, Mesophytes and a ese

RELATION OF PLANTS AND ANIMALS.—aaksinypon ae PLANTS:
The Sundews, Venus Fly-trap, Submerged Dionza, the Pitcher
Plants, the Bladderworts, the Butterworts and their biological
relations. Significance of the carnivorous habit. . . . . .—

CHAPTER XXV
GENETICS AND EvoLuTION

SCOPE OF GENETICS.—Genetics and Bai ;
Biereditary eubstance 3 a Ne ee

and his experiments.
Laws. Structure of the Chiveuceaint:
mosomes. Distribution of chromosomes tura
and fertilization. Distribution of genes. -Dominants a
recessives. Genotype and Phenotype. Monohybrids. Re SU
of crossing individuals with one pair of contrasting |

PHARMACEUTICAL BOTANY

CHAPTER I

THE SCOPE AND IMPORTANCE OF BOTANY

Botany is the science which treats of plants. Plants are liv-
ing things or organisms belonging to one of the two great king-
doms of living things, called the VEGETABLE or PLANT Kincpom.
The other kingdom of living creatures is that made up of animals
and called the AnmmAL Kincpom. ‘The science which treats of
animals is called ZodLocy. Botany and Zodlogy collectively
constitute the branch of natural science called BroLocy, the
science which seeks to inquire into all living things.

The term botany is derived from the Greek word Boravyn
meaning pasture, grass, or fodder. Man’s first inquiry into
plants was undoubtedly prompted by his instinct to seek food
and doubtless began with his first appearance on the earth.
History records the first pursuit of botany as a science by the
ancient Chaldeans, Egyptians and Greeks who cultivated it
particularly in relation to sources of food supply.

Later man found that some plants possessed medicinal
properties and interest was next focused in the direction of plants
as sources of medicines. The earliest pioneers in pharmaceutical

botany were the medicine men of primitive peoples including the
Indians of America who were acquainted with the virtues of
many medicinal plants before the coming of the white settlers.
~ They were followed in Europe by the rhzzotomoz or root collectors
who gathered medicinal herbs and prepared them for the
pharmacopolai or drug venders. Probably the greatest of the
early European pioneers in this field was Theophrastus, a Greek
_ philosopher and pupil of Aristotle, who lived between 372 and
- 285 B.C. He wrote ten books on the History of Plants in which

C. - about 500 species of plants used in the treatment of various

I

i. PHARMACEUTICAL BOTANY

diseases are described. Dioscorides, a Greek physician, about
77 A.D. wrote a Materia Medica which included a large number
of descriptions of medicinal plants and all the medicinal sub-
stances then known together with their properties. The early
Romans also contributed to the development of Pharmaceutical
Botany. Outstanding among these were Pliny the Elder
(23-79 A.D.), who in his Historia Naturalis crudely described
about 1000 plants, many of which possessed medicinal properties,
and Galen (131-200 A.D.), a Graco-Roman, who wrote a
Materia Medica.

Attention was next concentrated upon the classification
of plants, and gradually investigations followed on their structure,
functions, habits, relationships, diseases, etc., until to-day the scope
of botanical study embraces every kind of inquiry about plants.

The field of botany has thus developed many specialties or
departments of inquiry.

DEPARTMENTS OF BOTANICAL INQUIRY

1. PLant Morpuovocy treats of the parts, or structure of
plants. It is divided into:

(a2) MACROMORPHOLOGY or Gross ANATOMY which deals
with the external characters of plants and of their parts or organs;
(6) MicroMorPHOLOGY or HisToLocy which considers the minute
or microscopical structure of plants and plant tissues and (c)
Cyto ocy which treats of plant cells and their contents.

——2, Pant Empryovoey treats of the stages of growth, differen-
tiation and development of the individual plant body.

3. PLANT PuystoLocy deals with the study of the life processes
or functions of plants. It explains how the various parts ees

plants perform their work of nutrition, growth, reproduction, —
and the preparation of food for the support of animal life from
substances not adapted to that use.

4. Taxonomy or PLANTs or SysteMATIC BoTANy considers
the classification or arrangement of plants in groups in accord-
ance with their relationships toone another. It also is concerned
with the identification, the accurate description and the naming
of plants. All of the plants found growing in a certain <e
constitute the Flora of that region.

THE SCOPE AND IMPORTANCE OF BOTANY 3

5. Prant Ecoxoey treats of plants and their parts in relation
to their environment, 7.e. to soil, moisture supply, temperature,
light and other plants and animals.

6. PLant GENETICS deals with the coming into being of new
individuals or races of plants. It seeks to account for the resem-
blances and differences shown by plants related by descent.

7. PLrant PATHOLOGY or PHYTOPATHOLOGY treats of diseases
of plants.

8. PLanr GEoGRAPHY or PHYTOGEOGRAPHY treats of the
distribution of plants upon the earth. The center of distribu-
tion 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. When they grow in a locality other than their
original home they are said to be naturalized.

9. GroLocicaL BoTANy or PHYTOPALZONTOLOGY treats of
plants of former ages of the earth’s history traceable in their
fossil remains.

10. Cryptocamic Botany is the botany of the flowerless
plants or cryptogams such as the pond scums, fungi, mosses,
ferns and their allies.

11. PHanERoGaAmic Botany is the botany of seed plants.

12. Atcoocy is the botany of algae.

13. Mycotocy is the botany of fungi.

14. Bryo.ocy is the botany of mosses and liverworts.

15. Economic or APPLIED Borany deals with the science from
a practical standpoint, showing the special adaptation of the
vegetable kingdom to the needs of everyday life. It comprises
a number of subdivisions, viz.: PHARMACEUTICAL BoTANy, or
the botany of drug, spice, dye and poisonous plants; BAcTERI-
-oLocy, which treats of microérganisms called bacteria; PHAR-
MACOGNosy which treats of the history, commerce, selection,
identification, valuation and preservation of crude drugs and
other raw materials of plant and animal origin; AGRICULTURAL
Botany or AcRonomy, which treats of crop plants; Horti-
CULTURE, which treats of the cultivation of garden and orchard
plants; Forestry which treats of the management of forests and
of timber plants; PLANT BREEDING or the application of principles

4 PHARMACEUTICAL BOTANY

of selection to the growing of improved races of plants. All of
these subdivisions interlock closely with departments of pure
botanical inquiry and no training in either of them can be con-
sidered sound without the assimilation of the broad underlying
principles of morphology, taxonomy and physiology.
PHARMACEUTICAL Botany should include a sound foundation
in the morphology, taxonomy and elementary physiology of
plants before taking up its applied aspects, in order to adequately
prepare the student for the more advanced studies of Phar.
macognosy, Industrial Microscopy and Drug Plant Cultivation-

THE IMPORTANCE OF BOTANY TO MAN

It is an established fact that without plants there could be no
human or lower animal life, for green plants supply the food for
all living animals and maintain the oxygen content of the
atmosphere. They supply man with timber to build homes,
ships and furniture, with hairs and fibers with which to manu-
facture clothing, papers and books, with fuel to heat our build-
ings, with dyes to color our garments and ornaments and
preparations, with oils and resins to manufacture our soaps,
paints and varnishes, with drugs to treat our ills, with raw
materials for our explosives, with spices and flavorings to season
our food, and with vitamins without which we could not main-
tain our health, with perfumes, waxes, etc. In fact, for our
very existence, comfort and protection we are dependent upon
plant life.

Again, many of the fundamental principles of biology and
genetics have been first established through observations and
experiments with plants.

We look to the plants for a large number of our drugs and to
the patterns obtained by the examination of many of their
constituents for the chemical synthesis of a large number of
important organic chemicals, medicines and dyes. Coal,
representing the fossil remains of plants which lived millions of
years ago, furnishes the chemist with the raw material from which
he has built up hundreds of organic chemicals which are daily
used in pharmacy, medicine and the industries.

THE SCOPE AND IMPORTANCE OF BOTANY 5

Without some of the microscopic forms of plants, especially
the bacteria, yeasts and molds which represent agencies of decay
and fermentation, the earth would be literally covered with the
remains of the dead bodies of animals and plants accumulated
through the ages. While a number of these produce disease,
most are believed to be harmless, and many are of great eco-
nomic importance to pharmacy, agriculture, and to the dairy,
alcohol and tanning industries.

The study of botany unfolds many of Nature’s secrets, enhances
one’s appreciation of Nature’s beauty, develops one’s powers of
observation, and forms a sound foundation for the understanding
of many problems in pharmacy, chemistry, agriculture, indus-
try, and the biological sciences. .

CHAPTER II
THE GENERAL CHARACTERISTICS OF PLANTS

If we look about in nature we are. confronted at once with
many kinds of living objects called plants, but the more familiar
ones to the beginner in Botany are the higher green plants such
as the common herbs, shrubs and trees of field, meadow or
woodland. Each of these is a living entity or ORGANIsM which
has descended from a parent or parents with an ancestry and
which is capable of carrying out all of the life processes which are
essential to its existence and to the perpetuation of its species or
kind. Each is composed of Orcans whose life processes are
separate but mutually dependent and essential to the healthful
maintenance of the individual, and the continuation of its kind.

These organs are the Roots, Stems, Leaves, F LOWERS,
Fruits and Seeps. The roots, stems and leaves are so con-
structed as to carry out the VEGETATIVE PRocessEs, Be, the
activities which have to do with the maintenance and protection
of the individual, whereas the flowers, fruits and seeds are
structurally adapted to the carrying out of the REPRODUCTIVE
Processes or the activities which have to do with the continuance
of the species of the plant of which the individual is a
representative. :

PARTS OF THE PLant Bopy.—The parts of the body of a higher
green plant are the Axis and AppENDAGEs. The axis consists of
an usually aerial portion or SreM and an underground portion or
Root. The appendages include the Leaves, BRANCHES, EMER-
GENCES such as prickles, and the Hairs.

The Roor grows down into the soil and anchors the plant,
absorbs water with nutrient salts in solution and stores food to
tide the plant over the season when growth activity is suspended.
It represents the descending axis. The Stem grows upward
and so represents the ascending axis of the plant. Its special
processes are those of bearing and supporting leaves, branches

6 ,

THE GENERAL CHARACTERISTICS OF PLANTS 7

_- Leaf Rudiment
----- Growing Point
~~~. Young Leaf

Terminal Bud .....

Za —
Bie EAU EEE TRIG
at ath Line
Sar ACH ae oss oe

Som

.;

“SS LL
RESoe -

Mesophyl / Vein ‘Stoma
Intercellular Space

Pith.... ~
Cortex and... ‘
Pericycle <_ ~~ Petiole
“Axillary Bud

Conducting Tissue ..-}-1

Cotyledonary Bud

~“Y.

aera Cotyledons

Fic. 1.—Diagram of the body of a seed plant, showing its principal parts. (From
Robbins, after Holman and Robbins.)

8

PHARMACEUTICAL BOTANY

flowers and fruits, conducting sap containing nutrient materials
from roots to leaves and other aerial organs and vice versa.
Many stems also store food and green parts of stems manufacture

Fic. 2.—Flower and fruit of Flax.
A, floral diagram—, calyx; co, corolla;
s, stamens; p, pistil. B, median length-
wise section of flower. C, flower with
calyx and corolla removed showing the
essential organs, stamens and _pistil.
_ D, fruit, external view. £, cross sec-
tion of fruit. (Robdbins.)

food. The GREEN LEAVEs,
representing outgrowths of the
stem, are spread out to the sun’s
rays for carrying out the process
of manufaeturing-complex food
from simple inorganic materials
absorbed from the soil and the
air. ‘This process is called photo-
synthesis. ‘The FLOWERS are
equipped with sexual organs
called SraMens (male organs)
and Pistirs (female organs) and
are so constructed as to bring
about fertilization. The Fruits
which follow the flowers function
to form and disseminate the
seeds. ‘The SEEDs serve to carry
a young plant or Empryo which
has separated from its parent in
its dormant stage to the soil and
so function to spread the plant.
In addition to the life processes
noted, all of these plant organs
carry Out activities necessary for
their own maintenance such
as ‘TRANSPORTATION OF Foop
MateriAts, GrowrTu (increase
in size), RESPIRATION (the break-

ing down of carbohydrates accompanied by the taking in of
oxygen and giving off of carbon dioxide), and AsstmiLation (the

changing of food materials into

living matter).

Just as the plant organism is made up of organs which are

visible to the unaided eye, so each of these organs is made up of _

parts called Tissues, but the tissues are visible only with the aid
of a simple or compound’ microscope.

THE GENERAL CHARACTERISTICS OF PLANTS 1

Some of the Tissues in the stem for instance, are the outer
protective tissue or EpmeErmis, the soft region beneath the epi-

dermis and in the pith of the stem or PARENCHYMA, the woody
&

gre i " ’ yng

SS

Be

sie i ll

Fic. 3.—-Cross section of a representative portion of the stem of a clover (Tri-
folium) illustrating the tissues and tissue regions found in a stem of an herbaceous
dicotyledon. X 80. ¢p., epidermis or outer protective tissue; s, stoma, ss, sub-
stomal air-space, co, primary cortex containing parenchyma, f, sclerenchyma
fibers, ph, phloem containing sieve tissue, ca, cambium, and x, xylem of an open
collateral fibrovascular bundle; mr, medullary ray composed of parenchymatous
cells, m, medulla or pith composed of large parenchyma cells and small angular
air-spaces. Note the xylem contains long conducting tubes, here shown in cross
section as large open circular areas arranged radiately, which conduct water
upward through the plant axis.

portion which gives support to the stem or SCLERENCHYMA (stony
tissue), the water tubes which conduct crude sap upward or
TRACHEARY TISSUE and the sieve tubes and phloem cells which
carry elaborated sap downward or Steve Tissue.

10 PHARMACEUTICAL BOTANY

Other plant organs are also differentiated into secondary
parts or tissues which have been set aside to perform some definite
kind of work in the economy of the organs. The healthful
performance of the functions of an organ depends upon the har-
monious relation existing between the tissues of which it is
composed.

Chondriosomes
¢

2 aN past

x en"

sat Te
+

Pon tapas

Cyloplasm--~ 22, shh
Nuclear 5 rhea “oe
Membrane ~~ = ig —Plastids
a s
- x #
Nucleolus- DSc} ——-Linin
eg 7 Chromatin
4 *
Ps
oa sia a
“ai ~-*Chondriosomes

Fic. 4.—Sections of fixed and stained cells from the root tip of the garden pea
(Pisum sativum). A, young cell before appearance of vacuoles. 8B, older cell with
vacuoles. (Mottier.)

Every tissue, when examined microscopically, is seen to be
composed of numerous units of similar form which are in intimate
relation one with another. ‘These appear like small compart-
ments or chambers, separated from each other by walls and have
been termed Cretis. It will be noted that every cell possesses
contents.

While these cell contents vary in character depending upon
the cell examined, every living cell contains a viscid, often gran-
ular, reticular or alveolar, mobile, or jelly-like substance in
which life resides. This is called Prorop.asm or living matter.
It is this living substance which constructs the celis or funda-

THE GENERAL CHARACTERISTICS OF PLANTS 11

mental units of which the plant body is formed, and carries out
all of the life processes of the plant.

The herbs, shrubs and trees discussed in the preceding
paragraphs represent but few of the vast array of forms which

Fic. 5.—Two Blue-green Alge. A, B, C, D, E, Gleocapsa; F, Oscillatoria showing
a dead cell (¢d) which marks a place of separation into segments. (A), Gleocapsa,
parent cell composed of central protoplast containing scattered chromatin granules,
surrounded by cell wall and 3 mucilaginous 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.

make up the Plant Kingdom. Many additional kinds of plants
varying in form, habitat and method of living will be discussed in
the chapters to follow, but in order that the student may
approach the study of these with better understanding, it seems

12 PHARMACEUTICAL BOTANY

necessary, at this point, to make some general introductory com-
ment on the larger groups of plants.

Groups oF PLants.—There are four great divisions in the
Plant Kingdom, the thallophyta, bryophyta, pteridophyta and
spermatophyta. Each of these divisions is further divided into
subdivisions, classes, orders, families, genera and species. ‘The species
are made up of individuals.

ky Nee sy Qh
ye N GS
es wk

Fic, 6.—A mold, Mucor mucedo, showing the branched filaments or hyphe
which collectively constitute the mycelium of this fungus and the erect spore bearing
hyphz or sporangiophores. (Palladin.)

The lowest of these in scale of organization and the most
ancient is the THALLopHyTA or Thallophytes [Gk. thallos, a
young branch; phyton, plant]. ‘They have been so named because
their bodies, each of which is called a thallus, show no differentia-
tion into roots, stems and leaves. To this division belong two
large groups commonly known as the Acar and Funai. The
Algae possess chlorophyll, a complex mixture of green and yellow
pigments, while the Fungi are chlorophyllless. The ALGAE
include the green and blue-green scums which occur in stagnant
pools, ditches and in running streams, the diatoms and the green,
brown and red seaweeds. The Funct include the bacteria or

PT ae ee ae ee ee ee ee

THE GENERAL CHARACTERISTICS OF PLANTS 13

minute germs of decay and disease, the slime molds, the yeasts,
molds which occur on bread, cheese, fleshy drugs, some phar-
maceutical preparations, etc., the water molds, some of which
attack the gills of fishes, the mildews, the mushrooms and toad-
stools, and many forms which cause plant diseases, such as the
smuts, rusts, chestnut tree blight, ergot of rye, etc. There is
another subdivision of Thallophytes
called Licuens which comprises the
leathery-like growths which occur on
the bark of trees, on rocks and the
soil of woodland. They are peculiar
plants in that each represents a living
together of an alga and a fungus.
Next to the Thallophytes in order
of ascending complexity of structure
come the Bryopuyta or Bryophytes
[Gk.bryon, moss; phyton, plant}. ‘This
division includes the liverworts or
Hepatic and the mosses or Musct.
The liverworts are small, leaf-like
plants which occur on moist earth —
and in water. Some of them are
thought to be descendants of forms Fic. 7.—A liverwort (Lanu-
: laria). Below, portions of the
which were the first plants to take up i palloid body, showing the lunar-
a land existence. ‘The mosses in- _ shaped cupules, with brood-buds,
clude pale green forms of swampy 0F S8emma. Above a single
ground and bogs called peat mosses, ‘Gager) a ee
the black mosses of siliceous rock,
and the bright green or common mosses which so conspicuously
carpet portions of our forests and rocks. (See Figs. 315 and 318.)
The next highest division is that known as the PreRIDOPHYTA
or Pteridophytes [Gk. pieris, fern; phyton, plant] which have as
their representatives the CLus Mosses, HorseETaits or Scouring
Rushes and the Ferns. While most of these are low growing
terrestial plants, a few forms are aquatic in habit and some of the
tropical ferns assume tree-like forms. (See Figs. 325, 329 and
330.)
The Thallophytes, Bryophytes and Pteridophytes are col-
lectively termed the CryPTOGAMS [Gk. cryptos, hidden; gamos,

14 PHARMACEUTICAL BOTANY

marriage or union], since it was formerly believed that the sexual
organs of these plants were obscure. They reproduce by spores,
and, for the greater number, also by sexual reproduction.

_ 4

‘ *.
, =

Fic. 8.—A.gymnosperm, Zamia. Note the large cone which when mature bears
naked seeds.

The most highly evolved group of plants is that of the seed
plants or SPERMATOPHYTA [Gk. sperma, seed; phyton, plant] of
which the more familiar herbs, bushes, vines and trees are
examples. The Spermatophytes are also called the PHANERO-
GAs [Gk. planeros, evident; gamos, union], because it was formerly
believed that they alone showed evident sexual organs. They
are characterized by possessing flowers and producing seeds.

Oe oe el ee ee ee oe

THE GENERAL CHARACTERISTICS OF PLANTS 15

The SpERMATOPHYTES include two sub-divisions, namely
the GyMNosPERMS and the ANGIOsPERMS. ‘The Gymnosperms
[Gk. gymnos, naked; sperma, seed] have their seeds uncovered.
They include such plants as the pines, spruces, firs, hemlocks,

ie

Fic. 9.—A dicotyledon. The English Oak (Quercus pedunculata). A, twig with
leaves and flowers. B, staminate flowers. C, two stamens. D, pistillate flower,
enlarged. £, fruit, acorns. F, cup (cupula) with nut removed. G, seed in cross
section. H, seed in longitudinal section. The perianth of the staminate flower
consists of 5~7 segments; stamens 6-12. (After Wossidlo.)

redwoods, larches, yews, junipers, cedars, arbor vita, the cycads,
ginkgos, gnetums and ephedras.

The Ancrosperms [Gk. angeion, vessel; sperma, seed] have their
seeds enclosed within the seed vessel or fruit wall. They include
two sub-groups or classes, namely, the MonocoTyLepons and

16 PHARMACEUTICAL BOTANY

~~~.the DicotyLepons. The MonocotyLepons [Gk. monos, alone;
kotyledon, a cup-shaped hollow] are represented by the cat-tails,
grasses, sedges, lilies, palms, bananas, gingers, orchids, etc.
They have a single seed-leaf or cotyledon, mostly parallel veined
leaves and the parts of their flowers are in threes.

The Dicory.epons [Gk. di, two; kotyledon, a cup-shaped hol-
low] are represented by such forms as the willows, poplars,
maples, oaks, chestnuts, geraniums, violets, cherries, roses, beans,
asters, etc. ‘They have two seed-leaves or cotyledons, netted
veined leaves, and the parts of their flowers are in fours and fives.

Every seed plant normally comes from an ovum (egg cell) which
has been fertilized within the ovule of a flower. The fertilized
egg or zygote divides and redivides to form at first a nearly uniform
mass of cells. As these cells continue to multiply a process
early becomes manifest. This process is called PuystoLOGICAL
Division or Lapor and consists of the apportioning of separate
kinds of work to different parts of the same organism. The result
of the changes occurring in physiological division of labor is called
DIFFERENTIATION. As the cells continue to increase, unequal
growth takes place among them and, in time, they become
differentiated to form a young seed plant or embryo. ‘The
embryo lies within the ovule which by this time has ripened to
form with the embryo the structure called the seed.

Upon reaching suitable soil the seed absorbs water and its
embryo swells and sprouts forth or germinates, forming a young ;
seedling plantlet with roots, green foliage leaves and stem.
Eventually, through further growth and differentiation, a
mature adult plant is developed which bears flowers in which
female sexual cells or eggs and male sexual cells or sperms are
produced. The sperm unites with the egg, fertilizing it, thus.
starting anew cycle. This process is continued from generation
to generation.

CuaptTer III

THE LIVING CELL

THE CELL AS THE FUNDAMENTAL UNIT

Robert Hooke, an Englishman, in 1665, first described and
figured cells. He observed compartments regularly arranged in
rows in thin sections of cork placed under a microscope and,
likening these to the rooms of monasteries, named them cells.
In 1831, Robert Brown, an English botanist, discovered the

_nucleus which he found in a number of the plant cells.
Matthias J. Schleiden, a German botanist, in 1838, showed
the cell to be the unit of plant structure. The following year
_ Theodor Schwann, a German zoologist, showed that the bodies of
animals are also composed of cell units and announced the Cell
Theory, based upon Schleiden’s earlier work and his own observa-
tions. The bodies of all plants and animals are composed of one
or more of these fundamental units of structure and function.
Each cell consists of a unit mass of living matter to which Hugo
Von Mohl gave the name of protoplasm in 1851. But protoplasm
had been observed (though not so called) for the first time in
the cells of some plants by Corti and Treviranus as early as
1772 and in the cells of small animals by Dujardin in 1835. The
latter described it as a jelly-like substance, ‘‘the primary animal
substance,” and called it SAarcopE. The cell protoplasm may or
“may not have a cell wall surrounding it. Most plant cells con-
sist of a nucleated mass of protoplasm and a-cell wall surrounding
it. The term Protop.ast is given to the mass of protoplasm
found within the cavity of the cell. While most plant cells
contain a nucleus and some contain a number of nuclei, the cells
of the blue-green algze and 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.
17

18 PHARMACEUTICAL BOTANY

A Typicat Co.torLess PLANT CELL

If we peel off a portion of the thin colorless skin or epidermis
from the inner concave surface of an onion bulb scale, mount in
water and examine under the microscope, we find it to be com-
posed 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 (see Fig.
10). This wall is not living itself but is formed by the living
matter within the cell. It is sufficiently rigid to give definite
shape to the cell and sufficiently porous to allow water with sub-
stances in solution to pass through it. Somewhere within the
protoplasm will be noted a denser-looking body. ‘This is the
nucleus. The nucleus is called ‘“‘the superintendent | of the cell”
since it is believed to control all the activities of the protoplast.
Its removal results in the death of the cell. It is usually spherical
in shape but in old “epidermal cells of the onion becomes some-
what flattened and pushed up against the outer plasma membrane
and cell wall. It is surrounded by a nuclear membrane and con-
tains a nuclear reticulum, nuclear sap and one or more nucleoli.
The nuclear reticulum can best be seen after the cells are fixed
and stained. It consists of a thread-like net of a colorless sub-
stance called linin adhering to which are granules of a substance
which takes the stain of a basic dye called chromatin. Chromatin
carries the genes or hereditary characters from parent to offspring.
The nuclear-sap is a watery solution of nourishing substances.
Within the nucleus will be seen one or more smaller highly
refractile and definitely circumscribed, dense protoplasmic bodies,
the nucleolus or nucleoli. During cell division the nucleoli disap-
pear and reappear in the two nuclei of the daughter cells when
cell division is completed.

The protoplasm of the cell outside of the nucleus is called the
“cytoplasm.” It will be seen to be clear and granular-looking.
Within the cytoplasm will be observed a number of clear spaces. ,
These are pacoles and, because they are filled with cell sap (water

THE LIVING CELL 19

with nutrient substances in solution), are called ** sap-vacuoles.””
The vacuoles appear as tiny droplets in young cells but grow with
the cell, becoming larger as the cell increases in size, then fusing
with each other until, in a fully mature old cell, one large vacuole
is present in the center of the cell with protoplasm and nucleus
pushed up against the cell wall.

Fic. 10.—Portion of the inner epidermis of an Onion bulb scale, showing cells
at various stages of maturity. Young cell (1); old cell (3); cell intermediate in
age between 1 and 3 (2); cell wall (c); outer plasma membrane (/); middle lamella
(ml); nucleus (n'); nucleolus (n?); nuclear membrane (n*); cytoplasm (cy); vacuole
(v). Note that the young cell (1) shows numerous small vacuoles and a spheroidal
nucleus near center of cytoplasm. In 2 (cell of intermediate age) the cell has
enlarged, larger vacuoles have formed through the bursting of films of cytoplasm
separating smaller ones, and the nucleus has moved toward the cell wall. In 3,
the films have all burst, the cytoplasm and nucleus have been pushed up against
the cell wall, the nucleus is flattened out, and a large vacuole appears in the center
of the cell.

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, homo-
geneous, 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. All of
these membranes are bounding films of protoplasm belonging to
the region in contact with water and are collectively termed
plasma membranes. Upon mounting another portion of epidermis
in iodine solution, removing the excess of stain and adding a drop
of sulfuric acid and then examining under high power, we note

20 PHARMACEUTICAL BOTANY

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 line represents the fused

Fic. 11.—Elodea, an aquatic plant.

primary walls of adjacent cells and is called the middle lamella. It
is composed largely of calcium pectate.

TypicAL GREEN CELLS

Excellent material for the microscopic study of green cells of
plants is to be found in the leaf of Elodea, a fresh water plant,
found in ponds and house aquaria. The leaves of this plant are
small enough to be mounted beneath an ordinary cover glass on
a microscopic slide. They are oblong to oblong-ovate and
occur in whorls of 2 to 4 along a slender, elongated stem.

THE LIVING CELL ' 21

When one of these is detached, mounted in water and
examined under low power magnification, it is seen to be two
layers of cells thick, except along the margin, where only one
layer of cells is to be observed and in the central midrib region,
where there occur several layers of narrow cylindrical cells. The

CSS
ye

= a
aw sae Cs ae Po Som esd o
7 a kr

XS 5 =
_ wos Orga s sa — stevens Ve \ — am,

— aE og, Ae AND | aig,” x ies es nik SN

5 ey Se Te Cn ge oa SD ——- Pa

as ab si IE SMEs abate. ey,

eS SD IN 5 7229: 3S Si eat hte

Sa “ie aie : ee — \
“— EEE, SJ

an

we
BT HS Vea A Ba HUY 4 i
ea ot RSS ) ay!
LAAT ail HNN

rt ‘
Fic. 12.—Portion of the leaf of Elodea, magnified. v, midrib; ¢, pointed cells pro-
jecting at intervals from the margin.

cells are arranged in rows and vary in size, being shorter and
broader toward the midrib and longer and narrower near the
margin. Projecting tooth-like cells occur along the margin.
_ The cells of the upper surface appear about twice as broad as
those of the lower surface. Each cell is separated from its
neighbors by means of transparent ce// wails of cellulose and
pectic substances. Just as in the onion epidermis, what appears
to be a single partition wall between two cells is really two cell
walls separated by a middle lamella.

22 PHARMACEUTICAL BOTANY

If some of the larger cells are now examined under high power
magnification, each is found to be brick-shaped. Beneath the
cell wall will be seen a narrow zone of cytoplasm containing
numerous, green, ovoid, living bodies, the chloroplasts. Each
chloroplast is a dense, porous, mass of protoplasm containing the
green pigment or rather a mixture of four pigments two green
and two yellow and collectively called chlorophyll. Chlorophyll
is a highly complex and important material. By means of it,
energy in the form of light from the sun is absorbed and utilized
by the chloroplast for the manufacture of sugar. In some of the

Fic. 13.—A typica: green cell of Elodea. Cy, cytoplasm; N, nucleus; N,
nucleolus; C, chloroplast; Pm, outer plasma membrane; VM, vacuolar membrane;
V, sap vacuole; W, cell wall; S, intercellular-air-space. Greatly magnified.

cells the protoplasm will be found to be in motion (rotation),
carrying the chloroplast along with it. The nucleus will be
observed as a clear, dense, somewhat hemispherical shaped body
embedded in the cytoplasm and surrounded by the _ nuclear
membrane which separates it from the cytoplasm... A_ nucleolus
will be seen within the nucleus. The center of the cell is occupied
by a large sap vacuole containing a watery solution of salts and
other nourishing substances. A clear vacuolar membrane lines the
inner surface of the cytoplasm, while another clear membrane,
the outer plasma membrane, lines the outer surface of the cytoplasm.

If the leaf be boiled in water so as to kill the plasma mem-
branes and make them permeable and then mounted in dilute
iodine solution and examined under the microscope, bluish to
bluish-black bodies will be observed in the chloroplasts. ‘These

THE LIVING CELL 23

are starch granules. They have been formed in the chloroplast
through the agency of chlorophyll and sunlight by the process
called photosynthesis. ‘This process is discussed more fully in
another part of the book.

THE CELL WALL

Nearly all plant cells possess cell walls. The cell wall is
composed of non-living substances which are secreted by the

PORT ILIS ——es |
XS 5 SODIO = v8,
—s SIQYS | ¢ ]

One: iS ios 2 primary wall-----4
Cy IS tt secondary wall—--- |,
tertiary walk-~--" ,

Ce ener ae aR,”

cay
sie = 4

Peart

oe

p onecasties

vou

Qa

——

er

primary wall...
<" secondary wall, ™.

f tertiary — *

Fic. 14.—Primary, secondary and tertiary walls. A, B, longitudinal and
transverse sections of trachea of Linden (Tilia americana). C, D, longitudinal and
transverse sections of tracheid of a yew (Taxus brevifolia). (After Eames and Mac-
Daniels, Introduction to Plant Anatomy; McGraw-Hill Book Co. publishers.)

protoplasm.. The youngest meristematic cells in the growing
apical regions of roots, stems, and leaves are separated from each
other by delicate primary cell walls made up of carbohydrate
material called pectose. -As these cells grow older each of their
protoplasts secretes an individual secondary cell wall composed of
cellulose and some pectose. So what often appears to be a single
wall separating two adjacent cells consists of the primary wall
between 2 layers of secondary wall. The primary wall has

24 PHARMACEUTICAL BOTANY

undergone some changes due to the infiltration or addition of
other substances and now becomes known as the middle lamella.
At first the secondary walls are thin but become thickened in
many cells owing to the deposition of successive layers of cell wall
substances through the intercalation of new particles of cell wall
materials between those already present. Some cells like tracheae
and tracheids also possess a tertiary wall which is deposited by
the protoplast in the form of spiral or annular bands on the
inner face of the secondary wall.

Thick cell walls are said to be stratified, since they
show lamellz or layers which vary in appearance under the
microscope due to the presence of different quantities of water
or to their chemical composition.. They may also show, when
examined in surface view and especially after special staining,
fine lines running spirally or obliquely in the wall.

Other substances besides cellulose and pectose may be present
in mature cell walls. These include /ignin found in woody walls,
the water-proof substances cuéin and suberin common to walls of
protective tissues, and mucilage found in the walls of the epi-
dermis of some seeds. A further consideration of the cell wall
will be taken up in Chapter V.

THe AMCEBA

Amceba provides excellent material for the study of proto-
plasm. Itisa microscopic organism rarely more than 1p in. in
diameter, classified by most biologists within the phylum,
Protozoa, a group of unicellular animals. It is found in stagnant
water almost universally, where it adheres to weeds, dead leaves,
or other submerged objects.

If a drop of Amoeba culture be placed on the slide, covered
with a cover-glass, and the mount searched under low power, an
Ameeba should be found.

The beginner in microscopy is very likely to overlook an
Amoeba altogether, or to think that every little irregular, clear

spot in the field of view isone. When a clear object is seen which
is never the same in outline in consecutive moments, one is
certain of viewing Amoeba.

THE LIVING CELL 25

Move the slide if necessary so as to bring the organism into
the center of the field of view. Swing on the clips and bring the
high-power objective into position. Focus.

Note that Amceba is a one-celled organism devoid of a cell
wall, hence a naked mass of protoplasm. Protoplasm is a semifluid,
viscid, granular like substance, in which life resides. It is
shown by chemical analysis to consist mainly of certain sub-
stances known as proteids, bodies of extreme complexity in
chemical constitution, examples of which are white of egg or the
albumen of blood serum. Besides proteids, the protoplasm of

Fic. 15.—The Amceba. A, An amoeba moving in the direction of the arrows;
n, nucleus; v, contractile vacuole; f, food within food vacuole. B, the process of
fission in Amoeba, a nucleus in each half. B, after Jordan, Kellogg and Heath.
(From Bergen and Davis Principles of Botany.)

Ameeba contains fats and small proportions of mineral matters
such as salts of potassium, calcium, magnesium and iron. It
also contains a large proportion of water, called water of
organization.

Protoplasm is caused to shink by strong alcohol, which with-
draws the water of organization. It is coagulated by heat and
dissolved by prolonged treatment with weak acids or alkalies.

Note that the protoplasm of the Amceba consists of an outer
non-granular portion called the ectoplasm and an inner portion
called the endoplasm. Within the endoplasm is to be found the
nucleus which in the Ameeba is difficult to discern without the
use of special stains.

26 PHARMACEUTICAL BOTANY

Observe the Amoeba as it moves and find out whether the
direction of movement is in one line or several.

Note that Amoeba moves by extending protoplasmic processes
called pseudopodia, (false feet). At a particular point the ecto-
plasm is pushed out in the form of a small pimple-like elevation;
This increases in size, still consisting of ectoplasm only until, at
last, granules from the endoplasm stream into it and the projec-
tion or pseudopod comes to possess the same structure as the rest
of the Amceba. This method of movement is called ameboid
movement. ‘The same kind of protoplasmic movement is shown by
the slime molds and white blood corpuscles.

Note that when the organism comes in contact with a diatom
or any other particle of food pseudopodia are extruded and the
food is enveloped, so sinking into the cytoplasm. The food is
taken in with a small amount of water, which forms a surround-
ing bubble or food vacuole. This constitutes the first step in the
process of nutrition and is called ingestion. If watched sufficiently
long, the protoplasm of the diatom or substance of the food
particle ingested is slowly dissolved or digested through the
agency of some enzyme or enzymes within the endoplasm. As
it moves on, the Amoeba ejects or expels the cell wall of the
diatom or any other insoluble substance.

An enzyme is a substance of biologic origin which has the
property of causing a change in its surrounding medium without
becoming involved itself. Through enzyme action the pre-
viously water insoluble (colloidal) protoplasm or food of the
Amceba is broken down into water-soluble (crystalloidal)
substances which are capable of diffusion. The process whereby
water-insoluble foodstuffs are made water-soluble through
enzymic action is called digestion.

Mineral mattefs are taken in by the Amceba with its food in
the form of a weak, watery solution. The next step is for the
organism to rearrange the elements of digested food, mineral
salts and water in such a way as to form new particles of living
protoplasm. These are deposited among the pre-existing par-
ticles and so assimilated or converted into the living substance of
the Amoeba. One effect of this formation of new protoplasm is
growth. This process of growth by the interposition of new

THE LIVING CELL ot

particles among the old ones is called intussusception. All living
things, both animals and plants, grow or increase in size by this
method.

Minerals or other organized non-living things grow by a
deposition of successive layers on their original surfaces. This
process of growth is called accretion.

Fic. 16.—Stages in the division of Ameba polypodia. (From Marshall, after Schulze
and Lang.)

‘Look for a large, clear, pulsating structure or space within
the protoplasm of the Ameeba. This is the contractile vacuole.

Alike with all other living organisms, Amoeba is continually
undergoing destructive as well as constructive changes. Every
time it extrudes pseudopodia or performs work of any kind, a
portion of its protoplasm is broken down by oxidation, which is
accompanied by a release of energy. The nitrogenous waste in
solution, resulting from the oxidation of protoplasm, makes its
way to the contractile vacuole and is expelled by its contraction.
This function of getting rid of nitrogenous waste is called excretion.

28 PHARMACEUTICAL BOTANY

In order that the oxidation of protoplasm, referred to in the
preceding paragraph, may take place, a constant supply of
oxygen is essential. The water in which Amoeba lives contains
this gas in solution. Now, the protoplasm of the Amoeba (as
well as other living organisms) is continually forming carbon
dioxide. When two gases are separated from one another by a
porous septum, an interchange takes place between them, each
diffusing into the space occupied by the other. The process of |
gaseous interchange is precisely what takes place between the
H2O in the surrounding water and the CO, in the interior of the —
Ameceba, the protoplasm acting as a porous partition. This
gaseous interchange whereby this organism takes in oxygen and
gives carbon dioxide off is called respiration or breathing. ‘The
oxygen taken in by the Amoeba oxidizes the complex carbon
compounds within its body with an accompanying release of
energy.

In order that the race of Amoeba may not be exterminated,
there must be some provision whereby its numerous casualties
may be balanced. In other words, Amoeba must produce new
individuals like itself. This function is called reproduction.

Ameeba reproduces itself by the simplest method of reproduc-
tion, known as simple division or binary fission. At first the nucleus
divides into two; then the whole organism elongates, the two
nuclei at the same time traveling away from one another; next a
furrow appears across the middle of the elongated body between
the nuclei; the furrow deepens until finally the organism sepa-
rates into two separate Amoeba, which henceforth lead an
independent existence.

- Under unfavorable circumstances, such as the drying up of
the water in pools in which it exists and the approach of cold
weather, the Amoeba draws in its pseudopodia, loses most of its
water content, becomes spherical in shape and develops on its
outer surface a thick resistant wall called a cyst. Inside of the
cyst the nucleus undergoes division to form a large number of
nuclei. Cells walls then appear dividing up the cell contents
into as many little Amoebas called Amoebulz as there are nuclei.
Upon the approach of favorable conditions, these Amoebulz
escape by — of the cyst, cach Alte dome into an active |

THE LIVING CELL 29

Amoeba. This process is called sporulation and represents
another method of reproduction shown by some species of the
Ameeba.

THE RELATION OF PLANTS AND ANIMALS

One can see many striking differences between the highest
forms of plants such as the sunflower and orchid and the highest
forms of animals such as man and the gorilla, but as one passes to
the lower forms of plants and animals these differences gradually
disappear. There are a number of low types of organisms such
as Euglena and Volvox which have characteristics common to
both plants and animals and which might be placed in either

group.

PLANTS

. Most plants possess the
power of constructing com-
plex organic food substances
like sugar and starch from
water and carbon dioxide
by means of chlorophyll and
light.

. The cells of most plants
possess rigid cell walls com-
posed of cellulose and pec-
tose which surround their
protoplasts.

. Most plants are incapable
of moving from place to
place (locomotion).

. In most of the simple plants
growth is not localized but
in the more complex plants
growth in length becomes
localized in the tip regions
of organs and in these con-

\. tinues indefinitely.

. Most plants are constructive
agents.

ANIMALS

1. Most animals cannot con-

struct their own foods but
are dependent upon green
plants for the food essen-
tial to their nutrition and
existence.

. The cells of most animals

are devoid of such walls.

. Most animals are capable

of locomotion.

. In most animals growth in

length is not restricted to
the ends of body or to the
extremities of organs and
ceases when the animal
attains maturity.

. Most animals are destruc-

tive agents.

30 PHARMACEUTICAL BOTANY

Still more primitive and less differentiated are the Amoeba,
the vegetative stages or plasmodia of slime moulds, and the
bacteria which, along with many other unicellular organisms
of simple structure, have been called ‘‘Protista.”’ It is the belief
of most biologists that plants and animals of the present day
have gradually developed from a common origin which, during
an early era in the earth’s history, gave rise to the Protista from
which arose two diverging branches of organisms known as the
plant and animal kingdoms.

It is impossible to draw sharp lines of distinction which will
separate all plants from all animals owing to the similarity of
structures, functions and laws of inheritance found in organisms
belonging to both groups. It is possible, however, to find points
of difference between most plants and most animals. These are
given in the table on page 29.

PROTOPLASM AND ITs PROPERTIES

Protoplasm, or living matter, is the viscous cell substance in
which life resides. Huxley has defined it as the “‘physical basis
of life.”

Protoplasm occurs as a colloidal solution and as an emulsion
of colloidal particles. It has well been called a ‘polyphase
colloidal system.’ Colloids are substances like glue, agar or gelatin
which do not crystallize when solidifying from solution and
which will not pass through a semipermeable membrane like
pig’s bladder or parchment. Substances like salt and sugar
which crystallize from a solution upon evaporation of the solvent
and which pass through a semipermeable membrane are called
- erystalloids. When examined under the microscope, proto-
plasm appears either as a colorless jelly in which are frequently
imbedded globules and granules of various shapes and sizes or as
a streaming liquid. It occurs both as a sol or liquid that flows
readily or as a gel or jelly-like mass. It is extremely complex
and unstaple. It may change from the liquid to the jelly state or
vice versa under certain conditions. The liquid portion of the
protoplasm contains particles of ultramicroscopic size while the
jelly-like, granular portion contains particles, granules or
globules visible under a compound microscope.

THE LIVING CELL 34

Protoplasm cannot be analyzed chemically. Dead proto-
plasm, which is the material examined when protoplasm is
subjected to reagents in the procedure of chemical analysis,
consists of a variety of different chemical compounds including
proteins, carbohydrates, fats, salts of sodium, potassium, mag-
nesium, calcium and sometimes iron, etc., also gases like oxygen
and carbon dioxide and water.

PROPERTIES OF PRoropLasM.—The peculiar properties which
distinguish protoplasm from non-living matter are as follows:

1. SrructuRE.—Protoplasm, as it occurs in cells of plants
and animals, invariably exhibits structure. No region 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 withinit. 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. Inorganic
‘substances, such as crystals on the other hand, grow by accretion,
e.g. through the addition of similar material in layers on their
outer surfaces. A bit of protoplasm may retain 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. It may also be defined as the series
of processes concerned with the building up and breaking down
of living matter. To those processes concerned with the building
up of protoplasm, such as absorption, photosynthesis, digestion
and assimilation, the term anabolism is applied, Another term
for anabolism is nutrition. To those processes which involve the
breaking down of protoplasm, such as respiration, excretion
and secretion, the term katabolism has been given.

3. IrRITABILITY.—This is the property possessed by proto-
plasm of responding to a stimulus. All living matter responds

32 PHARMACEUTICAL BOTANY

to stimulation. When matter fails to be irritable or responsive
to stimuli, we declare it to be dead. The stimuli that excite
reactions in living matter are of two kinds, viz., intrinsic and
extrinsic.

Intrinsic stimuli are inherited stimuli, being carried by the
chromosomes from parent to offspring. They determine that
the plant shall conform to a particular type, carry on certain
activities, pass through a definite life cycle, and detach a portion
of its own substance for the formation of new individuals of its
kind.

Extrinsic stimuli initiate, inhibit, accelerate or modify the
effects of intrinsic stimuli. They comprise agents of the external
world such as cold, heat, chemicals, food, water, light, oxygen,
electricity, gravity, etc.

Tropisms, TAxres AND Nasties.—The irritable reactions
manifested by protoplasm and living things to the effects of these
external agents are termed tropisms, taxies or nasties and will now
be considered briefly. The term tropism is now largely limited
to the response of non-motile plants and is usually applied to
growth movements of these in response to external stimull.
The response made to a stimulus is usually designated by a
prefix name which implies the kind of stimulus, as for example,
the response to light is called phototropism, to gravity, geotropism,
etc. A response in the direction of the source of a stimulus is
called positive while one which is away from the source of the
stimulus is designated as negative. ‘The term taxy (pl. taxies) is
applied to responses of motile plants and plant cells, as gametes
and zodspores, which involve locomotion. The term: nasty
(pl. nasties) is applied to responses of non-motile plants induced
by stimuli in which the movements have no particular direction,
as for example movements brought about in some flowers as a
result of the succession of night and day (nyctinasty).

TuermorropisM is the response of living substance to the
stimulus of temperature. All living substance is influenced by
variations in temperature. Freezing disintegrates it while
excessive heat causes its coagulation. Active vital phenomena
are therefore only evident within these extremes, the limits
differing depending upon the endurance of the organism under

THE LIVING CELL 33

examination. ‘The lowest temperature at which the activity
of an organism becomes evident is known as the minimum, that
at which the activities are at their best, the optimum, and the
highest at which they can be continued, the maximum. Some
plants are able to endure greater extremes of temperature varia-
tion than others because of special adaptations. ‘Thus, certain
bacteria produce spores which resist exposure for an hour to the
temperature of liquid hydrogen (—225°C.) or to that of a
hot air oven at 100°C. Many higher plants can endure moder-
ately low temperatures by the development of a hairy covering;
others which are killed by frost produce seeds which can endure
rigid cold, still others adapt themselves to existence through
periods of cold by passing through a latent stage in the form of
bulbs, like the Squill or the Lily, or rhizomes, as the Blood Root
or the Hellebores.

CuHEMoTROPISM (or chemotaxy) is the response of protoplasm to
chemical stimulation. Any substances that possess the property
of producing a deleterious effect upon protoplasm are termed
poisons. Poisons may effect an immediate destructive combina-
tion with living substance when they are called caustics, or they
may have an exciting or depressing effect which may eventually
prove destructive without visible structural change, when they
are termed toxins. Caustics may liquefy the protoplasm, as the
alkalies, or coagulate it, as the acids or salts of metals.
When well’ diluted, chemicals may occasion no destructive
effects, but may call forth positive or negative responses, known
as positive or negative chemotropism or chemotaxis.

Thus, Pfeffer, working with the motile sperms of ferns, found
that if a capillary tube, containing a solution of malic acid, be
introduced into water containing them, the sperms moved
toward it and entered. It is now generally believed that the
motile male sexual cells of all flowerless plants are attracted to
the appropriate female sexual cells by means of positive chemo-
taxic influences. Among flowering plants, it has been observed
that pollen grains brought by various agencies from anthers to
stigmas of certain plants of different species will not germinate,
but when they are carried from one plant to another of the same
species or variety, they readily send their pollen tubes through

34 PHARMACEUTICAL BOTANY

the stigma and style to the ovule below. In the former instance,
negative chemotropism is illustrated, while in the latter, positive
chemotropism is shown.

SITOTROPISM is the reaction of living matter to the influence of
food. When the response is by motile organisms or motile
cells, it is called setotaxy. Hertwig found that if a fine capillary
tube be filled with a 1 per cent solution of asparagin or beef
extract and held in contact with a drop of water containing
certain bacteria, a mass of these soon plugged the mouth of the
tube. His experiment shows that these organisms moved from a
poorer to a richer nutrient medium in response to a positive
sitotaxic influence.

OxyTRopisM is the response to the stimulating influence of
oxygen. We see evidence of this everywhere in nature. No
living thing can continue to exist without this element. A
mistaken idea is often prevalent regarding obligate anaérobic
bacteria. Like all other bacteria or organisms, these plants
require oxygen but can only assimilate it in its combined form.
The tetanus bacillus is a good example. Aérobic bacteria, on the
other hand, require free (uncombined) oxygen for assimilative
purposes. Thus the tetanus organism (an anaérobe) grows in
the depth of culture media, whereas the tubercle bacterium
(an aérobe) grows only on the surface.

HyprotropisM is the response of protoplasm to the stimulus
of water. This reaction is seen in both positive and negative
phases in the slime molds. ‘The vegetative stage of these lowly
plants is characterized by a naked, many nucleated mass of
protoplasm, confining itself to the moist crevices of rotten logs,
etc. until the surface of the substratum becomes wet, when and
only when it will emerge (positive reaction). As soon, however,
as its fruiting stage begins, the whole protoplasmic mass wells up
from the substratum, away from moisture (negative reaction).
The roots of young seedling plants show positive hydrotropism by
growing toward moisture in the soil.

RHEOTROPISM is the response made by roots and _ slime
moulds to currents of water.

HeioTropiso is the response of living substance to the stim-
ulus of sunlight.

THE LIVING CELL 35

PHOTOTROPISM is the response of living things to light of any
source. ‘The stems of higher plants tend to grow toward the light
and are, therefore, positively phototropic, whereas the roots grow
away from the source of light and are so negatively phototropic.

Fic. 17.—Venus fly-traps, Dionea muscipula, growing in field near Wilmington, N. C.
(Photograph by Steckbeck and Palmer.)

Grorropism is the response of protoplasm to the stimulus
of gravity. Roots of Pteridophytes and seed plants invariably
grow downward toward the center of gravity and so are positively
geotropic. The fruiting organs of the fungi and the main stems
of higher plants tend to grow perpendicular to the earth’s

36 PHARMACEUTICAL BOTANY

surface and so are negatively geotropic. Branches of stems that
assume a relation parallel to the earth’s surface are diageotropic.
The Lima Bean, Sarsaparilla, Poison Ivy, and other plants whose
stems twine about supports exhibit dateral geotropism in their
horizontal curvatures.

GALVANOTROPISM is the reaction of protoplasm to electrical
stimuli. In this connection it may be said that the degree of
response bears a definite relation to the intensity of the stimulus.
No visible external electrotropic reactions have been observed in
higher plants, although when their cells are examined micro-
scopically, the reaction becomes manifest. Kiihne has shown
that when an electric current is passed through the hairs of the
Spiderwort, the cytoplasm becomes gathered into small globular
masses. :

THIGMOTROPISM is the response of living matter to mechanical
stimulation. Examples of this form of irritability appear to be
far less common among plants than among animals. Certain
species of Mimosa (Sensitive Plants); Oxalis, Drosera (Sundew),
Desmodium and Dionea muscipula or the Venus Fly-trap exhibit
this phenomenon to a marked degree. A few instances only
will be considered. When the tendrils of climbing plants come
into contact with the uneven surface of solid bodies they are
induced to coil. When the tentacles on a modified leaf of the
Sundew (Drosera) are stimulated mechanically by an insect or
artificially they are induced to curve over. If a good plant of
the Venus Fly-trap (Dionaa) is selected, it will be seen to possess
leaves, the terminal portions of which are modified as traps for
catching insects (Fig. 17). Hairs will be seen projecting from
the upper surface of each valve of the hinged blade. If one of
these hairs is touched with a pencil no reaction will be evident
but if, after a lapse of twenty seconds, the hair is touched again,
the 2 valves close. If the stamens of Berberis (Barberry) be
touched near the base during their pollen shedding stage they
will be observed to curve toward the stigma.

The most highly specialized form of thigmotropism observed.
in plants appears to be found in Mimosa Spegazzini, a member of
the Bean family. According to Steckbeck “when a mechanical
stimulus, such as a forceps pinch, is applied to one of the terminal

THE LIVING CELL 37

secondary leaflets, after a latent period of less than 14 second, the
leaflet stimulated rises and its partner almost at the same time.

Sif)

i
= . = = fe .

(>
= SY SSP ;
Sy, ise j
a 1 if. ~~ c ,
=| YES.
: ANS |
ar,
Sh

& ae
‘ae. ?
Gh Nig
Xe
Ks
Ke
—
: nwa
—< Ry hay
a BINS
~ ‘Sa i ‘
{ fan
i \ \
if

=»
™
5

- a
———

>

aed

Re SS —
aad?

B

Fic. 18.—Leaves of Mimosa pudica, A, normal position; B, after stimulation.
p, pulvinus or leaf cushion at base of petiole. Note the compound leaf consists of a
long petiole ending in a blade of 4 leaflets, each of which possesses a secondary
petiole with a pulvinus at its base, and a midrib which bears a large number of
small secondary leaflets. A stimulus applied to the large pulvinus at the base of
the main petiole will cause the latter to fall and the secondary leaflets to become
erect with the upper surfaces of each pair of opposite leaflets against each other

(B). (After Pfeffer.)

The stimulus is then carried down the midrib, the pairs of
secondary leaflets closing in order; in 9 seconds all the secondary
leaflets have closed, the midribs converge, followed in 3 seconds

38 PHARMACEUTICAL BOTANY

by a drop of the entire leaf. The stimulus moves up the other
leaflet with the result that the secondary leaflets close in order.
In 20 seconds after the stimulus has been applied all of the
secondary leaflets, are closed. The stimulus is propagated
through the stem to other leaves.”! (Cf. Figs. 18 and 19.)

Fic. 19.—Mimosa Spegazzini. After the application of a stimulus. Compare with
Fig. 18. (After Steckbeck.)

4. REpRoDUcTION.—Protoplasm also shows a very remark-
able 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.
These detached portions may be of the nature of sexual cells
which unite to form a cell called a zygote or odspore from which a
mature individual develops or of the nature of asexual cells
detached singly or enmasse which eventually develop into
mature individuals.

' “The comparative histology and irritability of sensitive plants” by D. W-
Steckbeck in Contributions from The Botanical Laboratory of the U. of Pa.,
vol. IV, No, 2, p. 217, 1919,

THE LIVING CELL

39

Other physiological characteristics, as motility and con-

ductivity, might be added, but
most satisfactory criteria by
which living may be distin-
guished from non-living matter.

PROTOPLASMIC MOVEMENT

Living matter in its active
state of existence exhibits motion.
The several kinds of movement
of protoplasm which may be
observed within the closed cells
or in naked cells are amoeboid
movement, ciliary movement, cyto-
plasmic rotation and cytoplasmic
circulation. Cold retards and
heat accelerates movement with-
in certain ranges. :

1. Amoebotd movement a
creeping or flowing movement
‘such as is typical of the plas-
modium or vegetative body of
slime molds and of the animal
Amoeba.

2. Ciliary movement can be
observed in the motile bacteria
and in the sex cells of the lower
and some of the higher plants
which possess slender proto-
plasmic outgrowths called c7lza
which lash the water.

3. Cytoplasmic rotation is well
illustrated in Elodea and Nitella,
two common aquarium plants.
In rotation, the protoplasm

is

the four just discussed are the

Fic. 20.—A, embryonic cells from
onion root tip; d, outer plasma mem-
brane; c, cytoplasm; a, nuclear mem-
brane enclosing the thread-like nuclear
reticulum 4, nucleolus; ¢, plastids (black

dots scattered about). 8B, older cells
farther back from the root tip. The
cytoplasm is becoming vacuolate; /f,
vacuole. C, a cell from the epidermis
of the mid-rib of Tradescantia zebrina, in
its natural condition 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 magnified.
(Stevens.)

moves down one side of the cell then across the end and returning
along the other side and end. In Elodea the chloroplasts found

ai

40 PHARMACEUTICAL BOTANY

in the inner layer of cytoplasm are carried along by the moving

cytoplasm. In some plants like Nitella the nucleus is carried
along with the current. (See Fig. 21.)

4. Cytoplasmic circulation or streaming may be seen in the cells
of the stamen hairs of the Spiderwort, in some epidermal cells of
the onion and in the epidermal hairs of the squash. Here the
cytoplasm will be seen in several strands traversing the cell
cavity as well as lining the inner face of the cell wall. The

Fic. 21.—Protoplasmic movement. 1 and 2, nuclei from the tapetal plas-
modium of the spore case of Botrychium, 1, in the usual form, and 2, flattened while
entering a crevice between spore-groups; 3, cell from Nitella showing cytoplasmic
rotation as indicated by the arrows; 4, cell from stamen-hair of the Spiderwort
(Tradescantia), showing circulation of the cytoplasm as indicated by the arrows.
(After Stevens.)

cytoplasm moves to and from the nucleus and cell wall and may
after streaming in one direction for awhile, reverse itself.

PROTOPLASMIC CELL CONTENTS

The name “Protop.ast” has been given to the entire living
or protoplasmic matter within a single cell. The protoplast
consists of six well-differentiated portions, viz.:

(a) Tue Cytop.asm, or the foamy, often granular matrix
of protoplasm outside of the nucleus. Embedded in the cyto-
plasm are non-protoplasmic bodies called inclusions. The most
prominent of these are the vacuoles which are spaces within the
cytoplasm containing either a watery solution of nourishing
materials (cell sap) or, in old cells, air.

THE LIVING CELL 41

(0) Tue Nucteus, a denser, definitely circumscribed region
of protoplasm containing nucleoplasm which consists of gels of
varying consistency containing chromatin, a substance staining
heavily with certain basic dyes, adhering in the form of granules
to a network of dimin. NucLear Sap, consisting of a watery
solution of nourishing substance is also found in the nucleus.
The nucleus may be regarded as the superintendent of the cell

Saad ain, ee
ry i Pia rattle
= 2

o 6 ec: 8 6
Fic. 22.—Diagram of Plant Cells: a, cell wall; 4, cytoplasm; ¢, cytoplasmic
strand; d, nucleus; e, plastid; /, plasmodesma, passing through pit in cell wall; g,
sapvacuole. (Drawing by H. McCarthy, from Stanford’s “‘Economic Plants,” D. Appleton-
Century Co. Publishers.)

since it controls the structural changes and physiological activi-
ties that take place within the living cell. It is usually spherical
in shape but in some plant cells is ovate, ellipsoidal, stellate,
discoidal or elongated. The chromatin is the substance which
carries those characters or factors from parent to offspring which
determine that the offspring shall resemble its parent or parents
and past ancestors.

(c) Tur Nucieo.us, a small body of dense protoplasm within
the nucleus. Sometimes two or more nucleoli may occur within
a nucleus. The nucleoli are believed to represent reserve food
which is employed in the metabolic processes of the nucleus.

Od

ae:

42 PHARMACEUTICAL BOTANY

(d) Tue Pxastips, composed of differentiated protoplasm
called plastic plasm. They are small, discoidal, spheroidal,
ellipsoidal, stellate, ribbon-shaped, or variously shaped, dense,
porous, protoplasmic bodies scattered about in the cytoplasm.
The plastids may become modified and specialized for definite
functions depending upon the position of the cells containing
them in the plant and the external conditions to which the cells
are exposed. Sometimes, as in the cells of lower plants
like the Spirogyra, plastids are large and are then called
CHROMATOPHORES.

In the meristem cells of root and stem tips of plants the plastids
are very small, colorless and usually found around the nucleus.
These minute plastids are termed primordia or proplastids. As the
meristematic cells enlarge and undergo differentiation, the
proplastids increase in size and develop into leucoplastids, chromo-
plastids and chloroplastids.

According to the position of the cells in which plastids occur
and the work they perform, they differ in color, viz.: _

Leucop.astips are colorless plastids found in the under-
ground 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 foods like reserve starch and oils from sugar and other
carbohydrates. Leucoplastids may become transformed into
chloroplastids upon exposure of cells containing them to light
through the formation within them of chlorophyll and related
pigments. They are only evident after properly fixing and
staining cells containing them. When leucoplastids build up
and store starch they are sometimes called amyloplastids; when
they build up and store oils they are called elatoplastids.

CHLorop.astips are plastids found in cells exposed to light
and contain the green pigments chlorophyll a and chlorophyll b
together with two yellow pigments known as carotin and xantho-
phyll. All green parts of plants contain these bodies. Chloro-
plastids may develop into chromoplastids, as in the ripening of
many green fruits. The fully developed chloroplastid is capable
of division by constriction.

CHROMOPLASTIDs are plastids found in cells independent of
their relation to light or darkness and contain a yellow, orange

THE LIVING CELL 43

or red pigment called chromophyll. The chromoplastids may be
circular, oval, needle-shaped, angular or fan-shaped and are
averagely smaller than chloroplastids. They are responsible
for the yellow and orange colors of many petals and sepals of
flowers and of ripened fruits. The root of the carrot contains
large numbers of them.

(ec) Ceti Mempranes.—These are of three kinds, viz.: (1)
The outer plasma membrane or ectoplasm, found as a transparent,
colorless, outer bounding film of cvtoplasm enveloping the

Fic. 23.—A, cell from the epidermis of the upper surface of the calyx of the
Nasturtium (Zropeolum majus) with crystalline chromoplasts; B, cells from the
petal of Lupinus luteus, with yellow chromoplasts; C, cell showing numerous chloro-
plasts scattered through the cytoplasm. A, after Strasburger; B, after Frank.
’ (Stevens.)

granular, inner cytoplasm, the latter sometimes termed ‘“‘endo-
plasm’? ; (2) the vacuolar membranes, clear bounding films of cyto-
plasm surrounding sap vacuoles; and (3) the nuclear membrane, a
clear, transparent film of protoplasm forming the outer bounding
zone of the nucleus. All of these membranes are semipermeable
membranes, since, in their living condition, they will permit the
passage through them of water and certain crystalloidal (water-
soluble) substances in solution while they will not admit the
passage of other crystalloidal substances in solution.

(f) CHONDRIOSOMES OF MrrocHonpRria.—These are small
granular or rod-shaped protoplasmic bodies found scattered in

44 PHARMACEUTICAL BOTANY

the cytoplasm of many cells. ‘They take a deep stain with basic
dyes, like chromatin, but their significance is not understood.
PLasMopEsMA.—The protoplasts of adjacent cells in many
plant organs especially those possessing thick cell walls are con-
nected by delicate threads of cytoplasm which pass through tiny
holes (pits) in the walls between the cells. These protoplasmic
connections are called plasmodesma. They are probably more
common between cells with non-lignified walls than is generally
realized owing to their frequent extreme fineness and the need
of a special method of fixing and staining of preparations to
bring them out. They occur in meristem, medullary rays,
storage tissue, phloem, etc. The author has found large and
prominent plasmodesma in the endosperm of species of Plantago
yielding Psyllium seeds. They have been found in the red
algze, in stems of mistletoe and pine, in nearly every part of the
mosses, ferns, and gymnosperms, especially in the young con-
dition of the cells, and in the thick walled endosperm of seeds of
nux vomica, ignatia, date, persimmon, etc. (See Fig. 22:)

CELL FORMATION AND REPRODUCTION ©

The cells of plants have all been derived from preéxisting
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.

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 proto-
plasmic contents takes place the process is called ‘‘sexual repro-
duction’ ; if, however, there is a mere separation of a cell or cells.
from an individual which later form a new organism, the process
is termed “‘asexual or vegetative reproduction.”

THE LIVING CELL 45

There are five chief modes of AsExuaL REPRODUCTION, Viz.:
Fission or Binary Division, Gemmation, Free Cell Formation, Rejuve-
nescence and Vegetative Multiplication.

Fisston.—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 bacteria, growth of many algze and the formation of tissues of
higher plants.

In fission or binary division, as the process is frequently called,
there may occur (1) a division of the cell by means of a cell
plate of the nature of a plasma membrane formed by the shrink-
_ age of the spindle fibers to the equator of the cell during mitosis.
This cell plate splits, forming two plasma membranes, between
which the cell walls separating the daughter cell are laid down,
or (2) a division by constriction. In the latter method, as seen
in some of the algae, the cytoplasm becomes constricted or pushed
away from the wall of the cell at a point midway between the
ends of the cell and at this constriction the cytoplasm lays down
a transverse wall of ring-like form. As the constriction deepens,
the new transverse wall broadens until, when division is com-
pleted, it forms a completely closed circular disk.

Gremmation or Buppinc.—This is the method of reproduction
common among yeasts. (See Fig. 284.) The cell forms a pro-
tuberance 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 Ceti ForMATION.—This is a type of reproduction
which consists of the cutting out of cells within the common
cytoplasm of a cell by the activity of spindle fibers attached to
the nuclei. These curve around the nuclei and form plasma
membranes each of which lays down a cell wall. Seen in the
formation of ascospores within the ascus of Ascomycetes.

REJUVENESCENCE Or ZOOSPORE ForMATION.—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 zodspore. Later it looses its cilia, develops a cell wall and
passes into a resting condition. Under favorable circumstances

46 PHARMACEUTICAL BOTANY

it grows into a new organism. It is found in C£dogonium,
Ectocarpus, and many other algae. (See Fig. 2572.)
VEGETATIVE MULTIPLICATION.—This form of asexual repro-
duction involves the giving off of multicellular portions or vege-
tative outgrowths of the parent plant, such as bulbs, tubers and
runners, which become detached and develop into new plants.
It occurs frequently in nature but is also accomplished artificially
through the removal by man of parts of the parent plant, such
as stem or root cuttings or leaves, or by layering, 7.e. bending
branches over until they touch the soil and subsequently give

rf Fue Geseinncennraaes 7 arr - : 5 : ae |

Fic. 24.—Propagation of Strawberry (Fragaria) by runners (stolons). The
two plants in the small pots developed from the tips of the runners produced by
the older plant in the central pot. (Gager.)

off roots and buds, when they are cut away from the parent
plant. (See Fig. 24.)

There are two kinds of SexuaL REPRODUCTION, Viz.: Conjuga-
tion and Fertilization. In both of these the sexual cells, called
gametes, or their nuclei come together and their protoplasm
blends to form a new cell. Fertilization is the common method
seen in higher plants.

Conyucation.—A union of two gametes, alike in physical
character, the product being a zygote or zygospore. This method
of reproduction is seen in the molds, Spirogyra, Kygnema, etc.

.FertiLization.—A union of two unlike gametes or their
nuclei, the product being an odspore (or zygote). One gamete,
the male sexual cell, is smaller and active, while the other, the

THE LIVING CELL 47

female sexual cell, is larger and passive. This process is seen
among the higher and many of the lower plants.

InprreEcT Nuciear Drvision (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 forma-
tion 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
chromatin 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
chromatin granules together into a thickened, twisted, chromatic
thread called the .spireme. 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-chromosomes. They lie within the nuclear
cavity united by delicate threads. There now begins a phe-
nomenon concerned with the cytoplasm which is primarily a
process of spindle formation. The granular cytoplasm accumu-
lates at the poles of the elongated nucleus, forming the cytoplasmic
caps. Presently it begins. to show a fibrillar structure, the
threads extending outward around the periphery of the nucleus
forming an umbrella-like arrangement of fibers from both
cytoplasmic caps. With the formation of fibers comes a break-
ing 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. Meanwhile 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-chromosome is drawn to one pole and
the remaining half to the other. At the respective poles the

48 PHARMACEUTICAL BOTANY

brag :

Re 4 h

Fic. 25.—Semi-diagrammatic representation of nuclear and cell-division. a,
resting cell ready to begin division; 6, 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; ¢,the nuclear thread has divided longitudinally
through the middle, and the spindle fibers have become more definite; d, the
nuclear 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 meta-
phase, where the longitudinal halves of the chromosomes are being drawn apart
preparatory to their journey toward the opposite poles; f, 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; A, 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; 7, 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.)

THE LIVING CELL 49

daughter chromosomes form a dense compact knot. A cell
membrane (ced/ plate), 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 membranes a substance com-
posed chiefly of pectose (and represent-
ing the primary cell wall) which later, by
union with mineral substances, chiefly cal-
cium, becomes the middle lamella. 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 chrom-
osomes, 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 distributed through the nuclear
cavity. It is soon possible to distinguish 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.

The process of mitosis may
phases and each of these briefly

1. PropHase.—Construction
spireme.

B

eae
Pe

Mm

Cc

Fic. 26.—Enlarged
views of spindle fibers
and cell-plate formation.
A, swellings appearing
on the fibers. 3B, lateral
union of the swellings,
forming a continuous cell
plate. C, splitting of
the cell plate into two
plasma membranes. D,
a cell wall is secreted be-
tween the new plasma
membranes of the
daughter cells. (From
“4 Textbook of General
Botany” by Smith, Overton
et al., Macmillan Co.
Publishers.)

a

be divided into four stages or
summarized, as follows:

of the chromatic thread or
Transverse segmentation of spireme into chromosomes

and their longitudinal splitting into double chromosomes. Dis-

appearance of the nucleolus or nucleoli.
plasmic caps and spindle.

Formation of the cyto-
Disappearance of nuclear membrane.

50 PHARMACEUTICAL BOTANY

2. MerapHase.—Arrangement of the double chromosomes:
at the equatorial plate with their daughter halves drawn apart in
V-shaped fashion. Attachment of spindle fibers from poles of
spindle to the halves of the chromosomes.

3. ANAPHASE.—Separation of the daughter halves of each
double chromosome. Movement of one-half of each double
chromosome to each of the poles of the spindle. Formation of
two dense knots or masses of daughter chromosomes, one at each
pole.

4. TELOPHASE.—Expansion of the masses of daughter chro-
mosomes at the poles, and their division into linin-carrying
chromatin granules, and the complete organization of the daugh-
ter nuclei. Appearance of a nuclear membrane around each ~
daughter nucleus. Appearance of a nucleolus or of nucleoli in
each daughter nucleus. Construction of a cell plate between
the two daughter cells. Splitting of the cell plate to form plasma
membranes. Formation of the middle lamella between the
plasma membranes. Formation of a cell wall by each plasma
membrane.

NuMBER OF CHROMOSOMES.—The number of chromosomes
found in the cell nucleus varies with the kind of cells considered
in each individual and with the species. Every species of plant
or animal has a definite number of chromosomes that appear
when its body cells undergo mitosis. In all sexually-reproducing
plants or animals two kinds of cells occur, the body- or somatic-
cells and the germ cells. Each of the body cells contains twice as
many chromosomes as each of the germ cells. Thus, the Stra-
monium contains 24 chromosomes in each of its body cells and
12 in each egg and sperm cell. The number of chromosomes in
the body cells is constant, is called the diploid or species number,
and is expressed as 2x. The number in the germ cells is called
the haploid number and is expressed as x.

MEIosis oR MATURATION
The process whereby the diploid number becomes haploid is
called meiosis or maturation.
Each of the spore mother cells which give rise to pollen grains
and to eggs in plants are formed by the division of body cells and

THE LIVING CELL 51

like them contain 2x chromosomes, of which x chromosomes are
contributed by the egg nucleus and x chromosomes by the sperm
nucleus. It is apparent that if each uniting egg and sperm had
2x chromosomes, the zygote resulting would contain twice the
number of chromosomes as found in the body cells of the parents.
Moreover, the body cells of the sporophyte resulting from the

Fic. 27.—Diagram showing various stages in the reduction division of a spore-
mother-cell of a plant. A, resting stage of the mother-cell nucleus; B, the nuclear
skein or spirem during synapsis; C, the spirem after synapsis, showing its double
(diploid) nature; the dotted line indicates the segmentation of the spirem into
2 diploid chromosomes, each of which has split longitudinally in DF, the diploid
chromosomes on the equator of the spindle of the first (heterotypic) division; F,
late anaphase; G, metaphase of the second or homotypic division; i; late anaphase
of same, two haploid chromosomes approaching the poles of each spindle; I, the
four daughter cells (spores) of the tetrad. (After Gager.)

division and redivision of the odspore would contain 4x chromo-
somes. This does not occur, for at a certain stage in the life
history of every sexually reproducing plant between each zygote
formation and the next, a reduction division takes place whereby
the 2x chromosomes are reduced to x chromosomes. ‘This
reduction in the chromosome number takes place in the two
nuclear divisions of spore -mother cells to form pollen grains and

embryo sacs with the result that each of the starting points of the

gametophyte generation have their chromosomes reduced to the

x or haploid number.

52 PHARMACEUTICAL BOTANY

In the first division called heterotypic mitosis, the resultant cells
contain, in their two daughter nuclei, only one-half the number
of chromosomes found in the nuclei of the body cells. ‘These
nuclei do not pass into the resting condition but soon undergo a
second division.

In the second division called homotypic mitosis, each of the two
nuclei divide again to form an aggregation of four daughter cells
in all, each with a nucleus containing the x number of chromo-
somes. The x number of chromosomes continues through the
whole gametophyte generation, so that the uniting egg nucleus
and sperm nucleus possess this number. In the act of fertiliza-
tion the x chromosomes from the egg nucleus are combined with
the x chromosomes from the sperm nucleus to form 2x (diploid)
chromosomes in the zygote. When the zygote divides, each of
its daughter cells contains 2x (diploid) chromosomes and their
number is continued throughout the sporophyte generation.

CHAPTER IV

LIFE HISTORY OF THE MALE FERN [AN
INTERMEDIATE TYPE OF PLANT]

The Male Fern (Dryopteris Filix-mas) has long been known to
the pharmaceutical and medical professions as the source of the
drug Aspmrum, a most valuable remedy for the expulsion of
tapeworm. ‘The parts of the plant 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 riims OF

Alike with other ferns, the Male Fern shows two phases or
generations in its life cycle. The first of these is that which is
represented by the more conspicuous plant which we call “‘the
fern.” It is in this phase that the spores or asexual cells are
produced, hence the plant producing them is called the SPORO-
PHYTE (spore-bearing plant). The second phase is that which is
represented in the development of an inconspicuous, tiny heart-
shaped body which occurs on the surface of woodland soil and
which bears the sexual organs in which sexual cells or gametes
are produced. This phase is called the GAMETOPHYTE or sexual

generation.

History OF THE SPOROPHYTE OR ASEXUAL GENERATION

Gross STRUCTURE OF STEM.—The main axis of Dryopteris
Filix-mas is the creeping underground stem or RuIzoMeE which is
oblique or ascending in habit. It gives off numerous Roots from
its lower and posterior portions and Leaves from its upper and
anterior portions. The leaves of a fern are called FRonps.
Behind the fronds of the present year are to be noted the per-
sistent stalk bases of fronds of previous seasons. ‘The roots are
slender, brown and fibrous with semi-transparent tips. 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 ExtTReEmITy and is marked by the presence of an APICAL
53

54 PHARMACEUTICAL BOTANY

Fic. 28.—Dryopteris (Aspidium) Filix-mas. % nat. size. A, Sorus and portion
of pinnule bearing same, in cross section, X 20. (After Kny.) B, Pinnule with
young sori, each covered with an indusium. C, the same, showing a later stage
of the sori, with shrivelled indusia.* Both slightly enlarged. (From Strasburger,
Noll, Schenck and Schimpers Lehrbuch der Botanik.)

LIFE HISTORY OF THE MALE FERN 55

Bup overarched by young fronds. The opposite end is known as
the Posrerior ExTremity and in the living plant is constantly
decaying, as the anterior portion elongates.

Gross STRUCTURE OF THE FronD.—Each frond consists of (1)
a stem like portion called the St1rz or leaf stalk and (2) a green,
expanded, leafy portion called the Lammna or blade. ‘The lamina
is further divided into a Mipris bearing its first divisions or lobes
which are called Pinn&. Each Pinna (sing. for pinnz) is further
divided into secondary lobes which are called Pinnutes. If a
pinnule be examined carefully it will be seen to possess VEINS
which are arranged in forked fashion. ‘The under side of some
of the pinnules will be seen to possess “fruit dots” or Sori.

Hisrotocy or Srem (Ruizome).—Passing from periphery
toward the center, the following structures are to be observed,
when cross sections are examined under a microscope (see Fig.
30):

1. Epmermis, a protective outer covering tissue, composed of
a single layer of brownish cells from which are given off scales.

2. OuTER CorTEX (hypodermis), a zone ‘of several layers of
thick-walled, lignified, fibrous cells separating the epidermis
from the softer underlying tissue. Its main function is to sup-
port the epidermis.

3. INNER Cortex of several layers of more or less isodiametric
cells (cells of nearly the same length, breadth and thickness)
with thin cellulose walls and containing stored starch surrounded
by a protoplasmic investment. Intercellular-air-spaces occur
between many of the cells. This tissue is called parenchyma or
fundamental tissue. Its cells conduct sap by osmosis and store
food. Between the cells are to be noted intercellular-air-spaces,
many of which contain internal glandular hairs with greenish
contents.

4. Prru, of parenchyma, resembling the last in aspect and
function.

5. VASCULAR BunpLEs.—These are elongated strands of
water tubes and other tissues which appear in cross section as
isolated spot-like areas. If the rhizome be cut lengthwise they
will be observed to join each other forming an open network.
Branch bundles come off of this network, some of which course

56 PHARMACEUTICAL BOTANY

into the roots and others into the leaves. Upon reaching the
blade of the leaf, they branch in such a way as to form the char-
acteristic veins of that organ. These are of two kinds, viz.:
stem bundles and leaf-trace bundles. The leaf trace bundles are
the smaller of the two kinds seen. Both are of elliptical outline,
as seen in cross section, and are embedded in the parenchyma
forming the broad central matrix. The stem bundles are
comparatively broad and, as viewed in
longitudinal sections, form a continuous
network with good-sized meshes, each
mesh being opposite the point of insertion
of one. of the leaves (see Fig. 29). In
transverse section these bundles are seen
to be usually ten in number and arranged
in an interrupted circle within the funda-
mental tissue. The leaf-trace bundles are
comparatively narrow and are observed
Fic. 29.—A, Cylindrical t0 Come off of the stem-bundles and pass
network of vascular bundles Out through the cortex into the leaves
in the stem of Dryopteris (fronds). When each “bundle” is ex-
Filix-mas. B, A portion of : ‘ ‘ :
ihe Tanto witwe Highly dnag: amined under a high-power magnification
nified. At L are the inter- it is seen to be composed of: (a) an
stices over which the leaves endodermis or bundle sheath, a single layer
a ae cae ails © of cells with yellowish walls and granular
passing into the leaves from COntents enveloping the bundle; (b) a
the main vascular bundles. pericycle (pericambium) or phloem sheath of
(Sayrs.) 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 w ls and protoplasmic contents,
which convey sugar in solution from \the leaves to the roots, and
broader steve tubes which appear polygonal in transverse section and
whose function is that of conveying soluble proteins in the same
direction; (¢) abroad central zone, the xylem (wood) formed of thin-
walled xylem cells which conduct sap slowly and store food, and
scalariform tubes or tracheids which conduct crude sap (water
with mineral salts in solution) rapidly from the roots to the leaves
(fronds). The scalariform tubes in lengthwise view have ladder-
like thickenings in their walls which alternate with thin areas.

57

LIFE HISTORY OF THE MALE FERN

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58 PHARMACEUTICAL BOTANY

Since the xylem is surrounded by the phloem, the vascular bun-
~ dle is of the xylocentric type. Strictly speaking, the endodermis
and pericambium are accessory regions, surrounding, but not
part of the bundle proper.

HisroLocy or Growinc ApEx.—When the bases of the leaves
of the current year, the young rolled up 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 observed with a hand lens:

1. The ApicaL Cone (punctum vegetationis), a rounded papilla,
which occupies a terminal position in the apical region.

Fic. 31,—Apical cell of a fern rhizome in vertical longitudinal section. 4.¢.,
apical cell; 4, hair; m, meristem, (After Hofmeister.) Sedgwick @ Wilson’s General
Biology, Henry Holt & Co. - :

2. The Younc Fronps, arranged around the apical cone.

Upon removing the extreme apex of the apical cone with a
sharp razor, mouting 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
(Fig. 31). The cells surrounding it have been derived by
segmentation (cell-division) from it, by means of walls parallel
to its three sides; they are termed segment 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.

HisroLocy or Mature Roor.—Transverse sections cut some
distance above the root apex will present the following structures
for examination: (See Fig. 32).

LIFE HISTORY OF THE MALE FERN 59

1, Eprermis, 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 from
the soil.

Fic. 32.—Cross section of a fern root. Photomicrograph, showing the central,
radial diarch bundle and surrounding tissues. Scalariform tracheids of xylem
patches (x! and x); phloem patches (p! and p*); pericambium (fe); endodermis
(en); cortex (co). X 300.

2. Cortex, of many layers of cortical parenchyma 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 endodermis.

3. ENDODERMIS, a single layer of cells tangentially-elongated,
whose radial walls are lenticularly thickened.

60 PHARMACEUTICAL BOTANY

4. Pericycie (Pericambium), usually of one or two layers of
thin-walled cells containing protoplasm and large nuclei. This
region surrounds the single bundle.

5. Rapiat VascuLar BunpDLe, consisting of two phloem
patches of phloem cells and sieve tubes on either side of two
radial xylem arms of scalariform tubes (tracheids) and a few
xylem cells. :

6. LaTerAL Root ets, which take origin in the pericambium.

Hisrotocy or Roor Apex.—Microscopic examination shows
this region to be composed of soft, pale, growing cells ending in
the triangular 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
divide to form three primary meristems. These are (1) the
dermatogen which, by repeated division of its cells, originates the
epidermis (outer protective covering of the root), (2) the periblem,
originating cortex, and (3) the plerome originating the bundle and
related tissue.

Continuity or Crupe Sap FLow.—The crude sap (water with
mineral salts in solution) penetrates 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
the stem xylem into the leaves.

HisToLoGy oF STIPE (PeTIOLE).—This, in transverse section,
passing from periphery toward the center, presents the following
structural characteristics (See Fig. 33):

1. Epmermis, a single layer of epidermal cells with dark
brown outer walls.

2. OUTER CorTEX (hypodermis), a wide band of small cells
with lignified walls.

3. INNER CorTEx, similar to inner cortex of stem but devoid
of leaf-trace bundles.

4. Piru, of fundamental parenchyma, similar to same region
of the rhizome, in the outer region of which are embedded a
number of xylocentric vascular bundles arranged in an interrupted

LIFE HISTORY OF THE MALE FERN 61

circle. Each of these shows a central xylem mass surrounded by
an outer phloem mass. Each bundle is enveloped by a per-
cambium and an endodermis or bundle sheath.

HisroLocy or A PinnuLe.—When transverse and surface
sections of the pinnule are studied microscopically, the following
structural details will be observed (See Fig. 34):

Fic. 33.—Transverse section of stipe of Dryopteris Filix-mas showing epidermis
(e); hypodermis (fh); inner cortex (ic); concentric fibrovascular bundles, one of
which is shown at (+); endodermis (en); pericambium (); xylem (x); and phloem
(p). (Photomicrograph.) X 50.4.

1. Upper Epwwermis, of wavy-walled, slightly chlorophylloid,
flat upper epidermal cells, devoid of stomata (breathing pores),
but with rather thick cuticle.

2. MESOPHYLL, a region between the upper and lower
epidermis consisting largely of chlorenchyma tissue which is
traversed by vascular bundles. The chlorenchyma contains
irregular shaped, chlorophylloid cells, with abundant chloro-
plasts. Intercellular-air-spaces are found between’ various
cells which are larger in the lower than in the upper region.

62 PHARMACEUTICAL BOTANY

Internal glandular-hairs with greenish oleoresin contents are
frequently to be discerned projecting into many of these spaces.

3. XYLOCENTRIC VascuLAR BUNDLES or laminar veins, that
distribute sap to, and carry sap from the mesophyll. These are

Fic. 34.—Structure of the Fern Pinnule. (Dryofteris Fititiivas) A, lower
epidermis in surface section. B, upper epidermis in surface section. C cross
section of mid-vein and adjacent interneural ions of lamina D, cross aii:
of marginal region. 2, epidermal cells below a vein; st, stoma; ge. aco cells; n
nucleus; c, chloroplastid; sv, sap vacuole; chi, chlorenchyma; nal Sis, iia: Ԥ
internal glandular hair; a, intercellular-air-space; St, sieve dieitic ay. .
xylem; fe, pericycle; en, endodermis; uep, upper epi is: ty
se ih Pidermis; mes, mesophyil; lep,

seen to be embedded in the chlorenchyma of the mesophyll
The xylem portion of each te is nearest, in end re <a f
veins, to the upper surface of the leaf cence

and so
approach the collateral type. wet Bie bundles there

LIFE HISTORY OF THE MALE FERN 63

4, ENDODERMIS, a continuous layer of cells whose radial and
inner walls are more thickened than the outer walls and which
surrounds each bundle and binds it in place.

Between the endodermis and the vascular bundle will be noted
one or more layers of thin-walled PERIcycLE cells. In the regions
of larger bundles, collenchyma tissue composed of thick-angled
cells occurs between the endodermis and the two epidermal layers.

5. Lower Epipermis of wavy-walled, flattened, chlorophyl-
loid 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 continuous around the laminar margin.

CoMPARATIVE PuysioLoGy OF Root, STEM AND LEAF
(FronpD).—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 Srorinc Foopsturrs 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 outer
plasma membrane or 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 scalariform tracheids. It
passes from one cortex cell to another by osmosis and, under con-
siderable root pressure, is forced into the 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 parenchyma cells (mesophyll).

Carbon dioxide (COz2), from the air, enters the leaf through
the stomata. From the stomata it moves through the inter-

64 PHARMACEUTICAL BOTANY

cellular-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 renders the energy of the absorbed light effective in break-
ing up the CO: and H:O into their component elements C, H and
O, and in rearranging them in such a way as to ultimately form
sugar. ‘This process is called PHorosynruEsis. The stroma of

Fic. 35.—Demonstration of the production of starch
foliage leaves. A, leaf with piece of cork attached whic
of the sun. 8B, the same leaf decolorized and treated wit
cork 24 hours later to show that starch was removed fro:
replenished. The illuminated portion turned blue on
cation of an abundance of starch. C, the entire leaf
indicated by letters CHO. (Mottier.)

in illuminated parts of
h cut off the direct rays
h iodine after removal of
m the shaded spot and not
addition of iodine, an indi-
was shaded except the part

the chloroplast has the power of manufacturing starch from the
sugar formed within it.

According to von Baeyer, CO, is split into C and Oy, the C
being retained, the Oz given off. The nascent C is linked with
HO to form CH:O (formic-aldehyde); six molecules of this are
then united to form grape sugar or glucose (CsH1.O,). The
Baeyer theory, however, has been criticized by Willstatter
others on the ground that formaldehyde is employed with
difficulty for the synthesis of carbohydrates and so ¢
considered as an intermediate product of carbon dioxid.

and
great
annot be
€ assimila-

ie.

LIFE HISTORY OF THE MALE FERN 65

tion. According to the theory of Willstatter which is not as yet
generally accepted, COs is first united with chlorophyll to form a
compound, the atoms of this compound become rearranged to
form a peroxide which later is decomposed by catalytic enzymes
with the liberation of free oxygen. The formation of starch may
be expressed by the following equation: 6CO, + 5H:O =
CsHi005 + 602. The molecules of glucose and starch contain
stored-up energy. 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 proto-
plasm of the cell. But the greater portion of sugar manufactured
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
removed 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, sulfur and
sometines phosphorus. They are formed from grape sugar
with the addition of nitrogen and the other elements by the living
protoplasm. The source of nitrogen, sulfur and phosphorus 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. The surplus protein material is stored in
various cells for future use.

Not all of the water carried to the green cells of the leaves by
the tracheids is used in the manufacture of sugar and starch.
Some of it is employed in chemical reactions taking place within
the cells and, some by the protoplasm itself of these cells for the
purpose of repair. Whenever there is a surplus of water in the
cells of the leaf, some of it escapes into the intercellular-air-
spaces as water-vapor. When the humidity of the external
atmosphere is less than that of the air-spaces, the water vapor
passes out through the stomata. The passage of water vapor
_ through the stomata is called TRANSPIRATION.

All of the living cells of root, stem, leaf as well as other organs
of the fern require oxygen. This gas enters the plant through the

66 PHARMACEUTICAL BOTANY

stomata and passes into the intercellular-air-spaces of all of the
organs which communicate one with another throughout
the entire plant. It diffuses through the cell walls lining these
spaces into the cells. Here it unites with the food materials
present, oxidizing a portion of them. Oxidation of glucose, etc.
is a breaking down process, similar to the combustion of fuel in a
furnace, but less intensive and not accompanied by light. The
chief products of oxidation are carbon dioxide and water.

The oxidation of food materials within the cells of organisms
is called ResprrATION. ‘The carbon dioxide resulting from this
process diffuses through the walls of the cells into the intercellular-
air-spaces and finally passes out through the stomata. When
food materials such as grape sugar undergo oxidation, their
stored-up energy (potential energy) is set free as energy in motion
(kinetic energy). This energy is utilized by the protoplasm of
the cell in carrying on its various kinds of works.

Respiration may be expressed by the following equation:

CsHiO, + 602 = 6CO;, + 6H.O
Glucose Oxygen Carbon Water
Dioxide

Respiration differs from photosynthesis in the following
particulars:

RESPIRATION PHOTOSYNTHESIS
1. It occurs in all living cells. 1, It occurs only in cells containing
chlorophyll.
2. It takes place during the day and the 2. It takes place only during the day
night. under natural conditions.
3. Oxygen is absorbed and carbon 3. Carbon dioxide is absorbed acd
dioxide is set free. oxygen is set free.

4. Carbohydrates are combined with 4. Carbohydrates are formed.
oxygen and oxidized. :

5. Potential energy is released from the 5. Kinetic energy is stored in the f, om
food molecules as kinetic energy. molecules as potential Sahay.

6. It is a destructive process. 6. It is a constructive Process,

There is another process which goes on in the various cells of
these organs. It will be recalled that the surplus glucose is
stored up in the form of starch by the leucoplasts and that surplus
proteins are also stored. These stored foods are not in the —
whereby ey can be used by the protoplasm when need a c..

LIFE HISTORY OF THE MALE FERN 67

repair and growth. In order to be serviceable for these purposes,
they must be changed into water-soluble products, This is
accomplished through the activity of substances called enzymes
which are manufactured by the cell protoplasm. ‘The process
whereby water-insoluble foods are converted into water-soluble
products is called DiGEsTIon.

Gross StruecTWRE AND HisroLocy OF THE SORI AND Spo-
RANGIA.—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»

Fic. 36.—Sporangia of an undetermined species of fern; /, 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.)

with a membranous expansion of the epidermis called the
indusium. ‘The whole is called a sorus (Fig. 28A) (pl. sori) and
contains many sporangia. Each sporangium is composed of:
(a) the stalk of considerable length and usually comprising three
rows of cells, outgrowths of the epidermis of the pinnule; and (6)
the head; sub-globular and hollow, consisting for the most part of
a covering wall 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. Beneath the wall of an
immature head of a spore case occurs a tapetum of a single layer of
cells and in the center spore mother cells. The spore mother
cells undergo two successive divisions to form tetrads or groups of
four spores which later separate as mature spores. During
the first of these divisions reduction in the number of chromo-
somes takes place from the number characteristic of the species

(diploid number) to the haploid number or half aaa or

68 PHARMACEUTICAL BOTANY

sporophyte number, so that each spore contains only half the
number of chromosomes found in the sporophyte generation and
thus belongs to the gametophyte generation.

Each spore is a single cell composed of an outer brown wall
with band-like markings called an exosporium, an inner thinner
wall or endosportum, and within this a mass of protoplasm con-
taining a nucleus.

The sporophyte generation ends with division of the spore
mother cell to form a tetrad of young spores.

HisToRY OF THE GAMETOPHTYE OR SEXUAL GENERATION

The gametophyte generation begins with the tetrad of young
spores within the sporangium.

RupTurE OF SPORANGIUM AND SporE DissEMINATION.—As was
previously indicated, each sporangial head had 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 then straightens and a wide rent is made in the
sporangium. The annulus then recoils and hurls the spores out
of the sporangium.

GERMINATION.—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 some-
what 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
antheridia, or male sexual organs, archegonia, or female sexual
organs, and rhizoids or hair-like absorptive structures. The
antheridia appear three to five weeks
rid oo PRES in shape and are situated among the
rhizoids toward the terior ‘5 ‘di :
ppaniingastheiore + 5 = sei Each antheridium consists of

pletely surrounds the 5 ermatocytes
or mother-cells of the spermatozoids. Within each ose

after spore germination.

LIFE HISTORY OF THE MALE FERN 69

the protoplasm arranges itself 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 arche-
gonia makes 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 antheridia of the same

prothallium. Each archegonium is. composed of a venter, neck,
neck canal-cells, ventral canal-cell, and ovum or egg-cell (see Fig.
38B). The neck is composed of cells arranged in four rows,
forming a cylinder, one layer of cells thick. This protrudes from
the surface of 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 matura-
tion of the archegonium, the canal cells are transformed into a

70 PHARMACEUTICAL BOTANY
mucilaginous substance which fills a canal extending from the

outside opening (mouth) to the ovum.
wees PE

gas
7 LEE
ae

ror

‘4
NA
isa
Rs

<
sparerss Ne
8 sr taale Wis re
ATTA ERE Be
SET eee SiG
a0y7, saps

J
¥
4

Fic. 38.—4, under surface of a fern prothallium showing archegonia (f),
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 arche-
gonium showing sperms (s..) moving through neck canal (n.c.) toward ovum (e);
and venter (v). All highly magnified.

During wet weather the mature antheridial wall bursts
open and the many ciliated spermatozoids escape into the water.

These, moving in the water, are drawn by the chemotactic

LIFE HISTORY OF THE MALE FERN 71

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 a zygote (odspore) or fertilized egg. This
completes the gametophyte generation.

OriGcIn OF NEw SpoROPHYTE PLANT FROM FERTILIZED Eco

The fertilized egg now divides into hemispheres, then into
quadrants and again to form eight cells arranged in a spherical
mass and called octant cells. The octant cells divide to form a
many celled embryo which shows the following parts: foot rudiment,
root rudiment, stem or rhizome rudiment, hair rudiments, and first leaf.

GROWTH OF EMBRYO INTO MaATuRE SpoROPHYTE.—The foot
rudiment develops into the foot which obtains nourishment from

Fic. 39.—A, median longitudinal section through immature antheridium, and
cell of prothallium showing prothallial cell (#), and antheridial wall surrounding a
number of spermatocytes; B, similar section through mature antheridium and cell
of prothallium showing fully developed spermatozoids (s.p.) enclosed by wall of
antheridium (an). Both highly magnified.

the prothallium, 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 rudiment becomes a rhizome
_ which, with the first leaf, turns upward. In a few weeks the
prothallus decays and the sporophyte is established as an inde-
pendent plant. More leaves (fronds) and roots are developed
and ere long continued growth results in the formation of a
mature sporophyte which presents for examination: (4}.:2
subterranean stem or rhizome, (2) roots; and (3) aerial fronds, each
of which consists of a stipe or petiole and a lamina or blade, the

72 PHARMACEUTICAL BOTANY

latter divisible into pinne or lobes and pinnules, upon which last
sort are developed.

ALTERNATION OF GENERATIONS.—It will be observed that in
the life cycle of the Male Fern there occur two distinct genera-

Fic. 40.—Pteris (The brake fern). A. Early stage in development of embryo
from odspore showing division of octant cells to form: f, foot rudiment; s, stem
rudiment; 4, first leaf rudiment; w, root rudiment. 8. Later stage in the develop-
ment of embryo; f, foot embedded in archegonium (aw) of prothallus (fr); s,
young stem; 4, young first leaf; w, young root. 4, after Kienitz-Gerloff; B, after
Hofmeister. (From Strasburger’s Lehrbuch der Botantk.)

Sporophyte
Stage

eww aie he awe eww es ee

—-_—-——

\5Q90 | F
;' 4۩ 3
G0<*&) -©<—
: Fic. 41.—Diagram of life-cycle of afern. 1, Mature sporophyte; 2, sporangium;
» Spore — cell; 4, tetrad; 5 spores; 6, germinating spore; 7, prothallus; 8,
archegonium; 8a, antheridium; 9, ovum; 9a, antherozoid; 10, fertilization; 11,
zygote. (Gager.) ; :
tions, one, a sporophyte or asexual generation which begins
with the zygote (or odspore) and ends with the division of the
spore mother cells to form tetrads of spores; a second, the

LIFE HISTORY OF THE MALE FERN 73

gametophyte or sexual generation, beginning with the spore and
ending with the fertilization of the egg to form the zygote or
odspore. ‘The sporophyte gives rise to the gametophyte which in
turn gives origin to the sporophyte. This is called “‘alternation of
generations.”

During this alternation of generations, a series of nuclear
changes takes place in which the diploid chromosomes or full
quota of the sporophyte generation are reduced to the haploid
number in the gametophyte generation. These changes occur
in the mitotic divisions of the spore mother cells to form spores
which contain only half the number of chromosomes found in
the spore mother cell and the rest of the sporophyte generation.
At the time of fertilization the two gametes, antherozoid and
ovum, fuse and restore the full quota or diploid number of
chromosomes to the zygote.

CHAPTER V
NON-PROTOPLASMIG CELL CONTENTS

The non-protoplasmic cell contents comprise many kinds of
substance of non-living character sometimes referred to as
inclusions in the protoplast. They include inorganic and organic
substances which are _deposited_in the cell as a result of. its
physiological activities. Many of them, when separated from
plants, constitute valuable drugs and foods. Some of them like
glucose and other sugars, starch and inulin are products of
photosynthesis, others like proteids, fixed oils and fats are formed
from the elements of glucose, still others like enzymes are direct
products of protoplasm, and a number of mineral compounds
such_as calcium oxalate, calcium carbonate and silica bodies
are formed as deposits as a result of. chemical processes within
the cell.

1. 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.. The MonosaccHAROSES are
simple sugars containing two to nine atoms of carbon, which are
known respectively as bioses, trioses, tetroses, pentoses, hexoses,
etc. Of these the hexoses (CgH,2O¢) are the most important and
of wide distribution. Examples of the hexoses found in drug
plants ES: (a) DextTROse (grape sugar), found in the leaves,
&! sproutin ies of flowers of nearly
all plants; (4) Frucrose (levulose or fruit-sugar), commonly
associated with dextrose; (c) D-MANNosk, 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.

Flickiger’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
74

NON-PROTOPLASMIC CELL CONTENTS 75

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 Drisaccuaroses, having the chemical formula of
Ci2H22011, include _ sucrose, maltose, trehalose, melibiose,
touranose and agavose. Of these e SUCROSE i 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; 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, transferred
to a solution of 1 part KOH in 1 part of water, and heated to boiling, the cells
containing the sugar take on a sky-blue color.

Invertase, an enzyme of the yeast, reduces cane_sugar to dex-
trose and levulose and zymase, another enzyme of the same plant,
ferments these forming carbon dioxide and alcohol.

MALTOSE_ is found in the germinating grains c of barley and
other cereals as a product of the action of the ferment diastase on
starch. Itreduces Fehling’s solution, forming cuprous oxide, but |
one-third less with equal weights.

TREHALOSE or mucose is found in ergot, Boletus edulis, the
Oriental Trehala and various other fungi.

Meurpiose is formed with fructose upon hydrolyzing the
trisaccharose melitose which occurs in the molasses of sugar
manufacture and in Australian manna.

Touranose is produced upon hydrolyzing melizitose, a
trisaccharose which occurs in Persian manna.

Acavose is found in the cell sap of the American Century
Plant, Agave americana.

2. Srarcu.—Starch is a carbohydrate having the chemical
formula of (CoH1005)n_¥ which is generally found as the first
visible product of ‘photosynthesis in most green plants. 2

76 PHARMACEUTICAL BOTANY

found in or more rarely adhering to the chloroplasts of green
parts of plants in the form of minute granules. This kind of
starch is known _as AssIMILATION STARCH.

Assimilation starch is dissolved during darkness within the
chloroplasts by the action of a starch-splitting ferment called
amylase and passes into water-solution as a glucose which is
conveyed downward to those parts of the plant requiring food.
In its descent, some of the surplus glucose not used by the cells in

Fic. 42.—Cell of Pellionia Daveauana, showing reserve starch-grains. The
black, crescent-shaped body on the end of each grain is the /eucoplast. Greatly
enlarged. (Gager.)

their nutrition, is stored up by leucoplasts in medullary-ray cells,

and in various parts of the xylem, phloem, pith and cortex in the

form of transitory starch grains. Considerable of it, however, is_
carried to the underground parts, such as rhizomes, tubers,

corms, bulbs or roots, or into aerial storage tissues, as the storage

cells of seeds, where the leucoplasts store it in the form of larger-

sized grains called Reserve Starcu. 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.

NON-PROTOPLASMIC CELL CONTENTS 77

STRUCTURE AND Composition oF StTarcu.—Starch grains
vary in shape from spheroidal to oval to conchoidal to polygonal.
They are composed of layers of soluble carbohydrate material
and probably other substances called ‘“‘/amella,” separated from
each other by a colloidal substance resembling a mucilage in its

behavior toward aniline dyes. They contain a more or less

distinct highly refractile point of origin or growth called the
“‘fulum,” 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.

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.

MetTHopD 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 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
excentric (outside of the center).

5. The shape of the hilum and the degree to which it is often
fissured.

6. The nature of the lamellz, whether distinct or indistinct;
if distinct, whether concentric (surrounding the hilum) or eccen-

TBO a OE Oe re ee ee” eee

SEN

CI: @
See

PHARMACEUTICAL BOT:

ome @ @©O@
~~ 56D
pad 2) ab

5 YOOO
° good on

Fic. 43.—A, wheat starch grains; B, rye starch; C, barley starc ch; ih potato starch;
text

E, Maranta starch; F, Sago starch. eeniaakts

lie | idl

NON-PROTOPLASMIC CELL CONTENTS 79

tric (apparently ending in the margin of the grain and not sur-
rounding the hilum), or both, as in potato starch.

7. The color of the grains when stained with dilute iodine
solutions; whether indigo, blue, purple, red or yellowish-red, etc.

8. The appearance under polarized light.

9. The temperature at which the paste is formed.

10. The consistency of the paste.

11. Whether the grains are unaltered or altered. *

CHARACTERISTICS OF IMPORTANT COMMERCIAL STARCHES

- Poratro STARCH (Solanum tuberosum)

Mostly simple, conchoidal or ellipsoidal,
with occasional spheroidal and two-
to three-compound grains.

size:'6 to 125.

Hilum: circular, at smaller end of grain.

Lamellz: concentric and eccentric.

Polarization cross very distinct; beau-
tiful play of colors with selenite plate.

MARANTA STARCH (Maranta
arundinacea)

Ellipsoidal to ovoid.

Simple.

Size: mostly 10 to 65z.

Hilum: a transverse or crescent-shaped
cleft in center or near broad end of
grain, or circular.

Lamellz: usually faint, concentric and
eccentric. j

Polarization cross very distinct; fine
play of colors with selenite plate.

Corn Srarcu (Zea Mays)

Polygonal to rounded.

Up to 354. Most grains over 15 in
diameter.

Simple.

Hilum: circular or a two- to five-rayed
cleft.

Lamellz: indistinct.

Polarization cross distinct but no
marked play of color with selenite

plate,

* Starch grains may be altered by heat, press
starch is starch altered by boiling. It gives a sky bl

Rice STARCH (Oryza sativa)

Polygonal.

2 to 10u in diameter.

Simple or two- to many-compound.

Hilum: usually indistinct, occasionally
a central cleft.

Lamellz: indistinct.

Polarization cross distinct but no play
of colors with selenite plate.

WHEAT STARCH (Triticum sativum)

Circular grains appearing lenticular
shaped on edge view; simple.

Large grains 28 to 45u in diameter.

Hilum: central, appearing as dot, but
usually indistinct.

Lamellz: generally indistinct, when
present in partially germinated
wheat, concentric.

Polarization cross indistinct; no play
of colors with selenite plate.

Rye Srarcu (Secale cereale)

Grains having a similar shape to those
of wheat starch but many larger;
simple.

Large grains 25 to 64y in diameter.

Hilum: a star-shaped central cleft or
indistinct in most grains.

Lamellz: indistinct; concentric in partly
germinated rye.

Polarization cross distinct.

, acids and enzymes. Soluble
ue color with iodine water. See

also dextrin, p. 82 and corrosion starch, p. 102.

80 PHARMACEUTICAL BOTANY

CROTCH OTS Oo 0 oF o
© ° Oa Oo Oo
0.8 r O® O° bao, 05
OOYS:; tino BE:
. ln © et Bn 0
© Do Ax0 en? oO oo:
@ © be Oe aa
G H

oo a
Fic. 44.—G, corn starch; H, rice starch; I, corn dextrin; J, pea starch; K, cassava
starch; L, bean starch. Explanation in text.

NON-PROTOPLASMIC CELL CONTENTS 81

BARLEY STARCH (Hordeum distichon)

Grains having a similar shape to those

of wheat starch but frequently tending
to bulge on one side and so appear
sub-reniform; large grains smaller;
simple.
Grains appear elliptical to lemon
shaped in edge view. Large grains
usually 18 to 25u, occasionally up
to 30y in length.

Hilum: centric or circular or cleft,
often indistinct.

Lamellz: concentric in partly germi-
nated barley; usually indistinct.

Polarization cross distinct.

BucKWHEAT STARCH (Fagopyrum
esculentum)

Grains simple and compound.
Simple grains polygonal or rounded-
polygonal.

Fic. 45.—Potato starch, polarized as
d starch under polarized light.

BucKWHEAT STARCH

Compound grains more or less rod-
shaped.

2 to 15u in diameter:

Hilum: central.

Lamellz: generally indistinct.

Polarization cross distinct.

CassAvA STARCH (Manihot
utilissima)

Grains rounded, truncated on one
side.

Simple or two- to three- or four- to
eight-compound.

6 to 35u in diameter.

Hilum: central, circular or triangular
with radiating clefts frequently.

Lamellz: indistinct.

Polarization cross prominent.

Arrowroot
Note that

Fic. 46.—Maranta or

viewed by light passing through crosse
nicol prisms of a petrographic microscope.
(Schneider.)

the two dark bands cross at the hilum of
each starch grain. (Schneider.)

82 PHARMACEUTICAL BOTANY

BEAN STARCH (Phaseolus vulgaris)

Ovoid, ellipsoidal or reniform shaped
simple grains, occasionally obscurely
3- or 4-sided.

Up to 60y in length. Generally from
30-35.

Hilum: central, elongated with bran-
ching clefts.

Lamellz: distinct, concentric.
indistinct.

Polarization crosses shaped thus, x.

In some

PEA STARCH (Pisum sativum)

Oval-oblong, elliptical or sub-reniform.

Up to 51y in length. Generally from
20—40u.

Hilum: similar to that of bean starch
but less cleft or simply elongated.

Lamellz: distinct, concentric.

Polarization crosses similar to - bean

CANNA STARCH (Canna edulis and other
species of Canna)

Broadly elliptical, flattened, with beak
or obtuse angle at one end.

Most grains 50 to 135y in length.

Hilum: excentric near narrower end.

Lamellz: concentric and eccentric.

Polarization cross very distinct; fine
play of colors with selenite plate.

Saco Starcu (Metroxylon Sagu et al. spp.,
Cycas spp.)

Ovoid, muller-shaped, or irregularly
3 or 4 sided with rounded angles.
Some more or less gelatinised.

Simple or 2, 3 or 4-compound.

Larger grains 30—60u long.

Hilum: excentric, often altered by
gelatinisation.

Lamellz: eccentric and concentric.

starch. Polarization cross distinct.

3. Dextrin.—Dextrin is a carbohydrate made from starch
(chiefly from corn or potato starch) by the application of heat
(yellow dextrin) or by treatment with both heat and acids (white
dextrin). It forms a paste with water, the yellow variety tending
to swell up and dissolve much more readily than the white.
When examined microscopically in alcohol mounts, the grains,
while conforming in general outline to those of the type of starch
from which the dextrin was prepared, nevertheless show more
conspicuous striations and clefts. Corn dextrin shows distinct
striations, whereas striations in corn starch are absent. The
grains take on a red coloration with iodine solutions,

4. AMYLODEXTRIN.—This is a carbohydrate intermediate in
properties between starch and dextrin. It occurs in the form
of small, irregularly shaped granules, in Mace, that take on a
reddish brown to reddish violet color with iodine solutions.

5. Inutin.—Inulin is a carbohydrate isomeric with starch
which has the_chemical formula of CuBr. 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, sphzro-crystals of inulin will be seen

NON-PROTOPLASMIC CELL CONTENTS 83

applied to the walls of the cells. When these sections are treated
with a 10 per cent. solution of alpha naphthol in alcohol and 2 or 3
drops of strong H,SQOx,, the sphzro-crystals will dissolve with a
violet color. Fehling’s solution is not reduced bv inulin.

6. Grucosmes (Glyco-
sides)—These substances are
very numerous in the plant
kingdom. ‘They arise in the
cell sap of plants containing
them as products of constructive
metabolism (anabolism) and Fic. 47,—Inulin, in the form of
are thought by many to have sphzro-crystals from the tuber of Dahlia.
the function of protecting plants A, precipitated from an aqueous solution;
against the ravages of animals. 2 Precipitated within the cells by long

standing in alcohol. (From Stevens, after
Some are known to serve as Sachs.)

reserve food. _All glucosides

are characterized by the-property of being split up into glucose
and other substances when acted upon by an enzyme, dilute acids
or alkalies.

EXAMPLES OF GLUCOSIDES

HespEeripIN.—Hesperidin is a glucoside having the chemical
formula of C2:,H2sO12. Like inulin it occurs in solution within
the cell sap. It is found in abundance in the Rutacez family but
occurs in many other plants. (See Fig. 85h.)

If sections of alcoholic material containing this substance, such as Buchu leaves
or unripe orange peel, are mounted in alcohol and examined, sphzro-crystals will
be seen. If these are then treated with a drop of alpha naphthol solution and 2
or 3 drops of strong H2SO,, 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 H2SOx,.

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-
centrated H.SOx, the cells containing strophanthin will assume a
bright green color.

_Sazicin.—Salicin is a glucoside occurring in the cell sap of the
bark and leaves of the Willows and Poplars. Sections of these

84 PHARMACEUTICAL BOTANY

mounted in concentrated H,SO, will show a red coloration in the
cells containing this substance. If water be added, a red powder
is thrown down.

Saponins are glucosides which have the property of frothing
with water. They are found in Soap Bark, Senega, Saponaria,
Yucca and other drugs and take a red color with strong H2SOu..

_ConIFERIN is a glucoside, occurring in the cell sap of the
spruce, pine, and other plants of the Conifere. If sections con-
taining it are first treated with a solution of phenol and then with
sulfuric acid, the cells containing it take on a deep blue color.

_Dierroxin, a glucoside found in the leaves of Digitalis pur-
purea, is colored green with hydrochloric acid.

CyANOGENETIC GLUCOSIDES are common in the seeds of the
Rose and Legume families. They yield hydrocyanic acid when
hydrolyzed by enzymes or other agents.

7. PeNTostpEs.—These are substances related chemically
to glucosides, differing from the latter in yielding pentose as one
_of the products, when decomposed (hydrolyzed) by enzymes,
dilute acids, etc. The aloins found in the juices of various
species of Aloe are examples of these principles. One of these,
“barbaloin,” found i in the official Aloe gives a greenish fluores-
cence in aqueous solutions, when the latter are saturated with
borax.

8. ALKALoIps-—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 katabolic 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 in water or “alcohol. They are
composed of carbon, hydrogen and and nitrogen. Some contain
oxygen. They are precipitated from saline solutions by the
addition of alkalies. They are mostly colorless and crystallizable.
They can be precipitated by one or more of the following alka-

NON-PROTOPLASMIC CELL CONTENTS 85

loidal reagents: tannic acid, gold chloride, phospho-molybdic
acid, picric acid and potassio-mercuric iodide (Maver’s reagent).

EXAMPLES OF ALKALOIDS

_ STRYCHNINE—This alkaloid, with a chemical formula of
C2:H»2N2Oz, occurs in the seeds of Strychnos Nux-vomica, Strychnos
~ Ignatii and other species of Strychnos. When sections of strychnine
containing seeds, previously defatted with petroleum ether and
absolute alcohol, are mounted in a solution of 1 Gm. ammonium
vanadate in 100 cc. of sulfuric acid, they take on a violet-red
color which later changes to brown. Strychnine itself produces
with sulfuric acid containing 1 per cent. ammonium vanadate a
deep violet-blue color changing to a deep purple and finally a
cherry red. It crystallizes from alcoholic or aqueous solutions,
forming rhombohedral crystals.

Brucine—This is another alkaloid obtained from Strychnos
seeds having the formula of C23H2¢O4Ne. It crystallizes from
watery solutions in the form of monoclinic prisms. Nitric acid
imparts to it a red color.

Nicotine.—This is a volatile alkaloid having the formula of
CioHi4No 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
reddish-brown precipitate which in time loses its color.

CAFFEINE.—This alkaloid, with a formula of CsHipNsO2 +
H.O, occurs in Tea, Coffee, Maté or Paraguay Tea, Cassine,
leaves of Ilex vomitoria or Yaupon, and seeds of Kola, Cocoa, etc.
If thin sections containing it are placed on a glass slide in 2 or
3 drops of concentrated 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, branching needles of caffeine gold chloride
will be seen to separate.

CocaInE.—This narcotic alkaloid, having the formula
Ci7HeiN' Ou,, 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 substituted for that of gold chloride, large

86 PHARMACEUTICAL BOTANY

feathers or fern frond crystals of cocaine-chloro-platinate will be

seen separating.
_Aconrring (C34H47O11N) is found in various parts of Aconitum
Napellus. It is particularly abundant in the tuberous root of this

Fic. 49.—Morphine. Orthorhombic prisms from alcoholic solution of this alkaloid.
X 120.

plant. If sections of aconite root are treated on a glass slide with
solution of potassium permanganate, a red precipitate of aconi-
tine permanganate will appear in the cells containing this
alkaloid.

Morpnine (C;7H1sO3N), the chief active principle of Opium,
crystallizes from alcoholic solutions in orthohombic prisms and

NON-PROTOPLASMIC CELL CONTENTS 87

acicular crystals. Sulfuric acid containing 5 mg. of selenous acid
in each cc. gives with morphine sulfate a blue color changing to
green and then to brown.

_ CODEINE (CysH2,O;N.H,O), another alkaloid occurring in
Opium, produces with sulfuric acid containing 5 cc. of selenous
acid in each cc. a green color which changes rapidly to blue,
then slowly back to green.

ye .
‘ SA .
>
: &
sy

Fic. 50.—Codeine phosphate crystals from aqueous solution. X 120.

CotcHicinE (C22H2 sO.N).—This alkaloid occurs in the corm
and seeds of Colchicum autumnale. 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
H.SO, and 3 parts of H2O, the cells containing colchicine will be
colored yellow. If a crystal of KNO; then be added, the color
will change to a brownish-violet.

9, GLuCcO-ALKALOIDS (Alkaloidal glucosides).—These are
compounds ‘ntermediate in nature between alkaloids and
glucosides, having characteristics ofeach. To this group belongs
solanine, a poisonous alkaloidal glucoside with a saponin-like
action, which is found in the potato (Solanum tuberosum), in
Solanum nigrum, Solanum Dulcamara, Solanum carolinense and other
species of the Solanaceae. When sections of those plant parts

88 PHARMACEUTICAL BOTANY

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
sulfuric acid with 18 parts of water, the cells containing solanin
take on a yellow color which changes successively to orange,

Fic. 51.—Various forms of calcium oxalate crystals. A, styloids from the
bark of Quillaja saponaria; B, rosette aggregate from rhizome of Rheum officinale;
C, raphides from the bulb of Urginea maritima; D, crystal fiber as seen in longitudinal
section in either the xylem or phloem regions of Glycyrrhiza; E, micro-crystals
(crystal sand) isolated from the parenchyma of Belladonna root; F, monoclinic prisms;
and G, twin-crystals from leaves of Hyoscyamus niger. All highly magnified.

various shades of red, blue-violet, grayish-blue and then
disappears.

10. AspARAGINE (CyHsN, + H,O).—This is an amino com-
pound 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 Althea officinalis and Atropa belladonna, in
young shoots of Asparagus, in the seeds of Castanea dentata, in the

NON-PROTOPLASMIC CELL CONTENTS 89

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.! If thick sections 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 satu-
rated solution of asparagine are added the crystals already formed
will increase in size. To get satisfactory results the saturated
solution must be of the same temperature as the mount.

11. Catcrum OxALATE.—This substance occurs in many
plants always in the form of crystals. It is 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 as a by-product of protein synthesis and other metabolic
processes. 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. (See Fig. 51.)

Crystals belonging to the monoclinic system possess 3 axes of
unequal length, two of which are obliquely inclined to each other,
the other axis forming right angles with these two. ‘They appear
to be more frequently found in plants than those of the tetragonal
system. |

Crystals belonging to the tetragonal system have 3 axes at
right angles to each other; two of the axes are equal in length, the
third being of a different length.

CRYSTALS BELONGING TO THE Mownoc.inic SysTEM AND
ExampLes oF Drucs ConTaAInING THEM:

1. Solitary—Hyoscyamus, Acer Spicatum, Viburnum
Prunifolium. ;

2. Rosette Aggregates (Druses)—Althza, Gossypii Radicis
Cortex, Stramonium, Granatum, Rheum, Foeniculum, Vibur-
num, Eucalyptus.

3. Columnar (Styloids)—Quillaja, Tris.

4. Raphides—Convallaria, Sarsaparilla, Veratrum Viride,
Scilla, Phytolacca, Ipecacuanha, Hydrangea.

1 Stevens’ Plant Anatomy, 3d Edit., p. 189.

90 PHARMACEUTICAL BOTANY

5. Micro-crystals (Crystal sand)—Belladonne Radix, Cin-
chona, Hyoscyamus, Stramonium, Phytolacca, Capsicum. ‘This
type is frequent in the Solanacez family.

6. Crystal Fibers—Cascara Sagrada, Prunus Virginiana, Gly-
cyrrhiza, Aspidosperma, Senna, Viburnum, Unpeeled Calamus.

7. Membrane Crystals—Aurantii Dulcis Cortex, Limonis

Cortex, other Citrus peels, Condurango, Wafer Ash bark.
Solitary crystals, usually in the
form of rhombohedra, occa-
sionally in twin crystals, occur
as sharp angular bodies, each
one often completely filling up
B the lumen of a cell. Twin
Fic. 52.—Cystoliths frorh the leaf of crystals occur in elm bark, hyos-
the ‘Fig (Ficus carica). A, Cystolith cyamus, etc. They are single
within cystolith sac; B, cystolith from : :
which the calcium carbonate has been shee: which have the appear-
removed for use in other parts of the ance of two prisms united by

plant. 8B is from leaf that has fallen their flat surfaces.
off in autumn. (From Stevens after
Haberlandt.)

Rosette aggregates or druses
consist of numerous small prisms
or pyramids, or hemihedral crystals arranged around a central
axis, appearing like a rosette or star. They often contain an
organic center.

Columnar crystals or styloids are elongated prisms.

Raphides or acicular or needle-shaped crystals, which fre-
quently occur in bundles in long, thin-walled cells containing
mucilage called raphide sacs. They are more frequently found in
Monocotyledons than in any other plant group.

Crystal fibers are longitudinal rows of superimposed paren-
chyma cells each of which contains a single monoclinic prism or
rosette aggregate. Crystal fibers are found adjacent to scleren-
chyma 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. (See Fig. 85k.)

NON-PROTOPLASMIC CELL CONTENTS 91

12. Micro-crysrats (sphenoidal micro-crystals or crystal
sand) are minute arrow-shaped or deltoid forms completely
filling the parenchyma cells in which they occur and giving these
a grayish-black appearance. While sometimes classified with
the calcium oxalate crystals, they are thought to represent some
other compound as yet undetermined.

13. Cysro.irus.—Cystoliths are clustered bodies formed by
the thickening of the cell wall at a certain point and subsequent
ingrowth 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, form-
ing an irregular spheroidal or ellipsoidal deposit. ‘These struc-
tures are abundantly found in the plants of the Nettle, Mulberry
and Acanthus families and constitute a leading peculiarity of the
same (see Fig. 52). They also occur in the Combretum and
Borage families, in species of Phlox, Klugia, etc. ‘They are proba-
bly excretory products. Excellent materials for the study of
cystoliths are the leaves of the rubber plant (Ficus elastica) and
the fig (F. carica). Here, the cystoliths appear as stalked
mulberry-like structures suspended from the outer walls of
specialized epidermal cells called “‘cystolith sacs.”

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 sativa.

The calcium carbonate incrustation of a cystolith dissolves
with effervescence on the addition of a mineral or organic acid.

14. Smica.—Silica (SiO,) occurs in a number of plants
either as an incrustation in the cell wall, as in Diatoms, the
Equisetinee and Gramineae or more rarely in the form of “silica
bodies,” such as are found in certain Palms and Orchids. It is
insoluble in» all the acids except hydrofluoric. It may be
obtained in pure form by placing tissue containing it in a drop or
two of concentrated sulfuric acid and after a time treating with
successively stronger solutions of chromic acid (starting with 25
per cent.) and then washing with water and alcohol.

In a number of members of the Palme, Orchidacea,
Scitaminea, etc. the strands of sclerenchyma fibers are accom-
panied by rows of small, tongue-shaped cells called stegmata.

92 PHARMACEUTICAL BOTANY

In these cells the walls adjacent to the fibers are thickened while
the opposite walls bulge outwards. Each stegma possesses a mass
of silica which usually fills the cell lumen.

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 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 bluish-black color, as that which
is found in Rhus, Castanea, Granatum, Galla, etc.; (b) another
tannic acid producing a dark 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 produced in those cells containing tannin.

Braemer’s reagent (Sodium tungstate 1 Gm. and Sodium acetate 2 Gm. dis-
solved in 10 cc. of water) is superior to solutions of iron salts in the detection of

tannin, since some other plant constituents give a dark-green or bluish-black color
with iron. It gives a yellowish-brown precipitate.

16. ProTeins.—Proteins are complex nitrogenous substances
forming the most important of the reserve foods of plants. ‘They
are found in the cytoplasm, nucleus and plastids of all the living
and many of the dead cells of plants, although most abundant in
seeds. ‘They occur as crystalline or amorphous bodies or in
colloidal solution. Protoplasm, itself, is composed largely
of these substances. They all contain carbon, hydrogen, oxygen,
nitrogen and sulfur, and many contain in addition phosphorus.
They are formed by the addition of nitrogen, sulfur and fre-
quently phosphorus to elements of grape sugar. The nitrogen,
sulfur and phosphorus elements are obtained from nitrates,
sulfates and phosphates which are dissolved in the water taken
in through the roots. All living plant cells are capable of syn-
thesizing proteins from carbohydrates. The process of protein

NON-PROTOPLASMIC CELL CONTENTS 93

synthesis appears to involve the following steps: (1) the union
of molecules of dextrose or some other simple sugar to nitrogen
to form amino acids; (2) the combining of amino acids to form
polypeptides; (3) the building up of polypeptides into simple
proteins; (4) the union of phosphorous or sulfur to simple
proteins to form complex proteins. The names of proteins recorded
may be found by the hundreds. These are grouped into
chemical classes, the most important of which from the stand-
point of their occurrence in plants are the globulins, albumens,
glutelins, 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

strong alcohol.
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 (made by dissolving 5 Gm. of iodine and 10 Gm. of potassium
iodide in enough water to make product weigh 100 Gm.) stains proteins yellow or

brown
Concentrated nitric acid stains proteins yellow. This color becomes deeper

upon the addition of ammonia water. ' a
Milon’s reagent (made by dissolving 3 cc. of mercury in 27 cc. of fuming nitric

acid without heat and adding an equal volume of distilled water) stains proteins a

brick-red. :
Concentrated solution of nickel sulfate 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

94 PHARMACEUTICAL BOTANY

of the genus or species. They are produced by the drying up of
protein-containing vacuoles. Each aleurone grain consists of a
ground substance (composed of amorphous proteid matter
soluble in water, dilute alkali or acid), in which are usually
embedded one or more crystalloids or phyto-globulins (insoluble
in cold water, but soluble in less than 1 per cent. solution of an
alkali, in dilute HCl and 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). (Fig. 53B.)

Fic. 53.—To show Aleurone Grains. A, cells from cotyledon of seed of garden
bean; n, aleurone grain; m, starch; B, cell from endosperm of castor bean; a,
aleurone grain; /, ground substance; k, phytoglobulin; i, globoid. (A, After Sachs;
B, After Frank.)

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. MuciLacrs anp Gums are those substances occurring
in plants which are soluble in water or swell in it, and which are
precipitated by alcohol.

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
Leguminose as Fenugreek, etc.

NON-PROTOPLASMIC CELL CONTENTS 95

CELL-CONTENT Muciace has been found in the leaves of Alee,
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 Buchu leaves,
Elm bark, Psyllium seeds, in Althea, Linum, Astragalus, and
Acacia species, in the Blue-green Algae, and many of the Brown
and Red Alge.

Gum.—When mucilage is collected in the form of an exudate
from shrubs and trees it constitutes what is termed a gum. Chem-
ically gums consist of glucosidal acids combined either partly
or completely with calcium, potassium and magnesium. Many
gums are used in pharmacy, medicine, and the arts. The four
most important from a pharmaceutical standpoint are: ACACIA,
yielded by Acacia Senegal and other species of Acacia; ‘T'RAGACANTH,
yielded by Astragalus gummifer and other Asiatic species of Astra-
galus; CHERRY Gum, obtained from Prunus Cerasus and its varie-
ties; and Rep Gum obtained from Eucalyptus rostrata and some
other species of Eucalyptus.

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 min-
utes, then draining off the stain and mounting in glycerin. Those cells containing
mucilage will exhibit bluish contents of deeper aspect than other elements.

18. Frxep Om anp Fats.—These are fatty acid-esters of
glycerin which are found in the vacuoles of cells or formed with
the cell walls from which they may be liberated as globules
upon treating sections with chloral hydrate or sulfuric acid or
heating them. The fats differ from the fixed oils only in that
they tend to solidify and crystallize at ordinary temperatures.
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 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

96 PHARMACEUTICAL BOTANY

in the chromatophores instead of starch as the first visible product
of photosynthesis. In higher plants it is generally found in
storage regions, such as the parenchyma of seeds, fruits and the
medullary-ray cells and the parenchyma of barks, roots and
rhizomes.

19. Waxes.—These are esters of fatty acids which differ
from fixed oils and fats in not forming glycerin when saponified,
but rather certain of the higher monatomic alcohols which are
insoluble in water.

The characteristic bloom on the epidermis of fruits of
junipers, plums and grapes and the glaucous surfaces of many
leaves are due to waxy coatings. Three types of waxy covering
occur on plants. ‘The most common type consists of densely
crowded granules as seen on the leaves and stems of many lilies,
grasses and irises. On the nodes of the sugar cane the covering
assumes the form of a layer of vertical rods, rolled or hooked at
their distal ends [Fig. 68(3)]. On the leaves of the houseleek
(Sempervivum) and the wax palms it occurs as a crust interspersed
with cracks. On the fruits of the wax myrtle and bayberry this
wax crust is secreted by glandular emergences.

Wax serves as an effective control over transpiration by the
epidermis. It also gives rigidity to the epidermis, lessening the
compression and distortion of subjacent parenchyma which is
apt to occur from its contraction during loss of water.

The more important vegetable waxes of commerce are
Carnauba Wax, from the leaf buds and young leaves of Copernicia
cerifera, the Brazilian Wax Palm, Myrtle or Bayberry Wax from
the fruits of Myrica cerifera, M. carolinensis and M. xalapensis,
and Japan Wax from the fruits of Rhus succedana, R. vernicifera
and R, sylvestris.

20. VoLaTILE Ons.—These are volatile odoriferous princi-
ples found in various parts of numerous plants which arise either
as a direct product of the protoplasm or through a decomposition
of a layer of the cell wall which Tschirch designates a “‘resino-
genous layer.” They are readily distilled from plants, together
with watery vapor, are slightly soluble in water, but very soluble
in fixed oils, ether, chloroform, glacial acetic acid, naphtha,
alcohol, benzin and benzol. They leave a spot on paper which,

NON-PROTOPLASMIC CELL CONTENTS 97

however, soon disappears. They respond to osmic acid, alkan-
nin, Sudan ITI, and cyanin stains similar to the fixed oils and fats.

Volatile oils may be grouped into four classes:

A. PrnENEs or TERPENES, containing carbon and hydrogen
and having the formula of CjoH1s. Examples: Oil of Turpentine
and various other volatile oils occurring in coniferous plants.

B. OxyGENATED Orts, 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. SuLFURATED Ors, containing carbon, hydrogen and
sulfur. Example: Volatile oil of mustard.

21. Resins, OLEoREsINS, GuM Resins, AND BALsAms.—These
substances 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 Storax, Benzoin, Balsam of Tolu
and Peru, etc. They occur usually in special cavities such as
secretion cells, glands, or secretion reservoirs. .

Resins are amorphous, solid or semisolid, fusible substances
insoluble in water but mostly soluble in alcohol and other organic
solvents. They combine with alkalies to form soap. Many of
them are oxidized oils of plants. Examples: Guaiac, Rosin,
Mastic.

OLEoRESINS are secretions composed of resins dissolved in
volatile oil. Examples: Turpentine, Copaiba, Balsam of Fir
and Venice Turpentine.

Gum RESINS are natural compounds of resin, gum and oil.
Examples: Asafetida, Myrrh, Gamboge.

BALSAMs are mixtures of resins with cinnamic or benzoic acid or
their esters and frequently both, the esters being generally called
a volatile oil. Examples: Balsam of Tolu, Storax, Balsam of Peru.

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.
22. PiGMENTS.—JT hese are substances which give color to
various plant parts in which they are found. They occur either

98 PHARMACEUTICAL BOTANY

in special protoplasmic structures, as chloroplasts, chromoplasts
or chromatophores, or dissolved in the cell sap. Of the pigments
named, the following will be considered: Chlorophyll, Xantho-
phyll, Chromophyll, Carotin, Etiolin, Anthocyanin, Phyco-
cyanin, Phycophzin, and Phycoerythrin.

Fic. 54.—Absorption spectra of five different concentrations of chlorophyll a.
The strongest is represented by the lowest spectrum. (From Palladin after Willstatter
and Stoll.)

CHLOROPHYLL is a collective name given to the two green
pigments found in the chloroplastids 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 and Magnesium, Carbon, Hydrogen,
Oxygen and Nitrogen are components of it. If green leaves
be boiled in water to kill the plasma membranes, the water
poured off, strong alcohol added and the leaf material allowed
to macerate in the alcohol for some time, an alcoholic extract
of the green leaves can be obtained. Ifa portion of this alcoholic
extract of green leaf be diluted with water to an 85 per cent.
alcoholic solution and xylene or benzine be gradually added
and the mixture shaken, the pigments in solution will break up

NON-PROTOPLASMIC CELL CONTENTS 99

into a yellowish and a greenish portion. ‘The greenish portion
dissolves in the xylene, which rises, forming the upper stratum,
while the yellowish portion dissolves in the alcohol and water
forming the lower stratum. By repeatedly shaking and adding
alcohol to the greenish fraction it can be entirely separated from
the yellowish fraction. The isolated greenish portion of alco-
holic leaf extract has been found to consist of two green pigments
called chlorophyll a and chlorophyll b. When separated and dis-
solved in alcohol, chlorophyll a appears blue-green and chlorophyll

Fic. 55.—Absorption spectrum of five different concentrations of chlorophyll 4.
The strongest is represented by the lowest spectrum. (From Palladin after
Willstatter and Stoll.)

b, yellowish-green in transmitted light. By reflected light, the
former is blood-red, the latter brownish-red. Chlorophyll a
constitutes 72 per cent. by weight of the green pigments and has,
as so far known, the formula of CssH72O;sN4sMg. Chlorophyll 6
represents the remaining 28 per cent. and has, as far as is known,
the formula of CssbinOeltaMg— > The isolated yellowish portion
of the leaf extract consists of two pigments, carotin and xanthophyll.

Alcoholic solution of chlorophyll, when examined spectro-
scopically in several concentrations, shows absorption bands in
the red, orange, yellow and green with a diffuse band in the

100 PHARMACEUTICAL BOTANY

violet-blue zone of the spectrum, the sharpest band being in
the red.

CHROMOPHYLL is a broad term applied to either the yellow
or orange pigments found in chromoplastids. Sulfuric acid
forms a blue color with chromophyll.

CaROTIN or carotene is an orange to orange-red pigment
which has the chemical formula of CyoHs5. It is found accom-
panying the chlorophyll within chloroplasts, but exists separately
in chromoplastids of leaves, flowers, fruits and seeds, as well as in
subterranean parts of higher plants. It is especially abundant
in the roots of carrots. It is readily soluble in chloroform, ether
and carbon bisulfide, slightly soluble in hot alcohol and insolu-
ble in water. When examined with a spectroscope, it shows two
dark bands in the green-blue half of the spectrum. Carotin is
a source of Vitamin A, the growth-promoting factor. It is
changed into Vitamin 4 in the body.

XANTHOPHYLL is a yellow pigment which accompanies the
chlorophyll in the chloroplast. Chemically it represents an
oxidation product of carotin, having the formula of C4oH55O2.

ETIOLIN is a pale yellow pigment which appears when green
plants are kept for some time in darkness. It is probably identi-
cal with xanthophyll.

_ANTHOCYANINS are applied to the blue, purple and red pig-
ments which occur in the cell sap of floral organs, stems, leaves,
roots and fruits. The character of the color is claimed to be
due to the alkalinity or acidity of the cell sap.

Puycocyanin is the blue pigment found in the blue-green
algze, associated with chlorophyll. It is soluble in water and
glycerin but insoluble in ether and alcohol.

PHYCOPH4IN Or FUCOXANTHIN is the brown pigment found in
the brown algz. It is readily soluble in water.

PHYCOERYTHRIN is the red pigment found in many of the red
algze and some of the blue-green alge. It is readily soluble in
water but insoluble in ether and alcohol. )

The last two pigments are always associated with chlorophyll
but frequently conceal it.

23, Latex.—This is an emulsion of varying composition and
color found in special passages, as latex cells and laticiferous

NON-PROTOPLASMIC CELL CONTENTS 101

vessels, of many plants. It may contain starch, sugar, proteid,
oil, enzymes, bitter principles, 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 Gutta and Payena spp. which yields Gutra PERcHa.

Hevea species, Ficus elastica, Landolphia species, Castilloa
élastica, Hancornia speciosa, Forsteronia species, Funtumia elastica
and F. africana, Manthot Glaziovit, Clitandra species and various
species of Urceola and Euphorbia furnish most of the RuBBer of
commerce.

Lactuca virosa and other species of Lactuca yield the drug
LacTuUcARIUM.

Euphorbia resinifera, contains a white resinous latex which is
gathered as the drug EupHoreium.

24. EnzyMEs.—An enzyme or soluble ferment or unorganized
ferment (according to Hepburn) is a soluble organic compound
of biologic origin functioning as a thermolabile (sensitive to
heat) 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
immediate part in the reaction, remains unaltered at the end
of the reaction and can be recovered again from the reaction
product unaltered in quantity and quality. The biologic
catalysts (enzymes) differ from the inorganic catalysts in that
they are sensitive to heat and light. They are destroyed quickly
at 100°C. and most of them cannot be heated safely above 60°C.
The optimum temperature range for this activity usually is
from 40° to 45°C. Enzymes are soluble in water, dilute alcohol,
glycerin or dilute saline solutions and are precipitated by con-
centrated alcohol. They are colloidal and non-diffusible.
They are.stimulated to activity by substances known as “‘activa-
tors’ and their activity is checked by other substances called

102 PHARMACEUTICAL BOTANY

“paralyzers.” Frequently 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 DiIFFUSIBILITY 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: Invertase of Yeast.

B. Accorpinc To Kinp or Supsrances ACTED UPON AND
‘TRANSFORMED.

1. CARBOHYDRATE ENZyYMEs:

Drasrasg, found in the germinating seeds of barley and other
grains and in Aspergillus oryza, etc., converts starch to maltose and
dextrin. As observed under the microscope, starch grains partly
transformed by diastase (amylase) show depressions, later deepening
to form irregular tunnels. Such grains are called corrosion starch.

INVERTASE, secreted by yeasts, and found in younger parts of
higher plants, transforms cane sugar, producing dextrose and
levulose.

Ma tase, found in malt, in green leaves and Saccharomyces
octosporus, transforms maltose to dextrose.

TREHALASE, found in: Polyporus, hydrolyzes trehalose to
dextrose.

Cyrtase, found in Nux Vomica and Ignatia seéds, in barley,
dates, etc., decomposes hemicellulose and cellulose to galactose
and mannose.

Lactase, found in Kephir grains, hydrolyzes lactose to
dextrose and galactose.

PROTOPECTINASE splits protopectin of the primary cell wall
to soluble pectin.

Pecrase hydrolyzes the methyl alcohol from soluble pectin
to produce pectic acid. |

Pectinasg, found in fruits and other plant organs, transforms
the soluble pectin, pectic acid or pectates of the middle lamella
into galactose, arabinose and galacturonic acid.

NON-PROTOPLASMIC CELL CONTENTS 103

INuULASE, found in plants of the Daisy and Bluebell families,
etc., transforms inulin to levulose.

ZyMASE, found in yeast, hydrolyzes glucose (dextrose and
levulose) to alcohol and carbon dioxide according to the follow-
ing equation: CsH»2Os = 2C,H;OH + 2COr.

2. Fat AND O1L ENZYME:

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 ENZYMES:

PEpsIN converts proteids into proteoses and peptones.

Trypsin, found in yeast and other fungi, as Boletus edulis,
Amanita spp., etc., resolves proteins to peptones and amino-acids.

BROMELW, found in the fruit of the Pineapple, and PAPAYIN
(Papain), found in the latex of the fruit of the Papaw, act
similarly to trypsin.

-Nepentuin, found in the pitchers of Nepenthes species, acts
similarly to pepsin.

4, GiucosipE ENZYMES:

Emutsin (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 ]-mandelonitrile glucoside) to hydrocyanic
acid, benzaldehyde and glucose.

Myrostn (myronase), found in the seeds of the black mustard
(Brassica nigra) and other members of the Crucifere, converts
the glucoside, sinigrin, into allyl-iso-sulphocyanide and glucose.

RuamnaseE, found in Alder Buckthorn (Rhamnus Frangula)
and probably other species of Rhamnus, hydrolyzes the anthra-
glucoside frangulin to rhamnose and emodin.

GAULTHERASE, found in Wintergreen (Gaultheria procumbens)
and other Ericaceous plants, resolves the glucoside, gaultherin,
to methyl-salicylate and glucose.

5. Orcanic Actp ENZYMES:

REDUCTASE of yeast decomposes lactic acid to pyrotartaric
acid and hydrogen. It also decomposes formic acid into carbon

dioxide and hydrogen. -

104 PHARMACEUTICAL BOTANY

CARBOXYLASE, found in higher plants, splits pyrotartaric
acid into acetic aldehyde and carbon dioxide.

C. Oxipizinc ENzyYMEs.

These are enzymes which absorb molecular oxygen from the
air and combine it with plant constituents.

PEROXIDASE occurs in many plants including apples, many
fleshy fungi, potatoes, etc.

PEROXIDASES are common in plants and are responsible for
discolorations seen in the tissues of many barks and fruits when
they are bruised or broken and exposed to the air. The juices
and extracts of plants containing oxidases turn brown upon
exposure to air. This is owing to the formation of organic
peroxides by the action of atmospheric oxygen on phenolic
substances present and the presence of a peroxidase which, with
an oxygen acceptor, brings about oxidation with a resulting
colored oxidation product.

25. Hormones.—The term “hormone” means chemical
messenger and is appropriately named because it produces its
effect at a distance from its point of origin. Hormones are
substances which govern the proper development or functioning
of organisms and organs. The hormones of animals, sometimes
termed “internal secretions,” are formed by the endocrine
glands as the pituitary, thyroid, parathyroid, adrenal, pineal,
pancreas, ovaries, testes, and are secreted directly into the blood,
lymph and cerebro-spinal fluid.

The hormones of plants, while anticipated for some time,
remained unknown until 1928, when F. W. Wendt demonstrated
the existence of a plant growth hormone controlling the enlarge-
ment of the coleoptile of the oat seedling. In 1933, F. Kégl
and coworkers isolated 3 plant-growth hormones in crystalline
form, auxin a (CisH3205) which they obtained from malt, maize
germ oil and human urine, auxin b (CisH300,4) and hetero-auxin
from vegetable material. Hetero-auxin has been identified as
indole-3-acetic acid. Their formation and distribution has been
found to be influenced by light. They regulate growth in plants.

When seedlings are illuminated on one side, the growth
hormone moves from the illuminated side to the shaded side.

NON-PROTOPLASMIC CELL CONTENTS 105°

The result is that the cells elongate more rapidly on the shaded
side, bending the plants toward the light. It has been recently
shown that though growth-hormones are produced in the light,
they function better in the dark.

26. ViTaMins.—Vitamins are essential food principles for the
most part manufactured by plants without which animals cannot
maintain their health. They are designated alphabetically
as A, Bi, Bs or G, Bs, Bu, Bs, C, D, E, and K and by names
indicative of the deficiency disease in animals they correct.

Viramin A, also known as Fat-soluble A, Anti-xerophthalmic,
and Antt-infective Vitamin, was discovered by McCollom and
Davis in 1915. It occurs in conspicuous amounts in carrots,
green leafy vegetables, in a number of fruits, as tomatoes,
prunes, pineapple and banana, in sweet potatoes, in wheat
germ, yellow corn meal, whole milk and in cod and halibut liver
oils. In its chemical and physiological properties, Vitamin A
is closely related to carotene which occurs in carrots and many
other plants. Its presence in oils can be detected by the addi-
tion of a few drops of a chloroform solution of antimony tri-
chloride to the oil when a blue color appears which deepens
and then fades. ‘This vitamin protects epithelial tissues against
bacterial infection, especially that of the respiratory mucosa,
and is essential to the proper growth of the bodies of young
animals. Animals deprived of it in their diet lose weight and
develop a hemorrhagic condition in their eyes known as xero-
phthalmia. The U.S.P. unit of vitamin A is the amount of
vitamin contained in 149909 of a grain of the U.S.P. Reference
Oil taken as a standard.

VITAMIN B, or Anti-neuritic Vitamin is a water-soluble, thermo-
labile factor which has been isolated in pure crystalline form
from rice polishings. Its absence in the diet has been found to
be the cause of beri-beri, or polyneuritis, a disease of the nervous
system characterized by muscular incoordination followed by
paralysis of certain sets of muscles. Its presence in the animal’s
diet promotes growth by stimulating the metabolic processes,
protects it from nerve disease and stimulates appetite and
digestion. It is also essential for normal gestation. This
vitamin occurs abundantly in yeast, rice polishings and wheat

106 PHARMACEUTICAL BOTANY

germ and to a lesser extent in whole rice, rye, wheat, barley,
milk, eggs and a number of vegetables and fruits.

Viramin Bs or G, sometimes called the Anti-pellagric factor
of the Vitamin B complex, prevents pellagra, a disease charac-
terized by soreness of the mouth and tongue, diarrhea, skin
eruptions and disturbances of digestion and mentality. It is
water-soluble and little affected by heat. It occurs in yeast, in
spinach, kale and other green vegetables, in potato, banana,
fresh carrots, cabbage, beets, tomatoes, lettuce, wheat bran,
meat, etc.

Viramins Bs, Bu, and B; are all growth-promoting principles
which have been found to occur in yeast. They are water-
soluble and thermolabile.

Viramin C or Antiscorbutic Vitamin is the factor which prevents
scurvy. It is water-soluble and destroyed by heat. It has been
found to be ascorbic (cevitamic) acid, CsHsO¢. and has been
made synthetically. This vitamin is required for the proper
metabolism of bones and the formation and nutrition of teeth.
It occurs abundantly in the juice of oranges, lemons and limes,
in lettuce, spinach, cabbage and tomatoes and to a less extent
in pineapple, raspberry, strawberry, peach, beans, corn, and
some other fruits and vegetables.

Vitamin D or Anti-rhachitic Vitamin is essential for the preven-
tion of rickets, since it regulates the absorption, retention and
deposition of calcium and phosphorous by the bones and teeth.
It has been found to be irradiated ergosterol. It is formed in

animals by the action of ultra-violet light upon cholesterol —

present in the skin. This vitamin has not been found in plants.
It is soluble in oil and relatively stable to heat. It occurs
abundantly in cod-liver and halibut liver oils and in smaller
amounts in salmon, egg-yolk, oysters, clams, etc. The U.S.P.
unit employed in the biological standardization for vitamin D
is that amount of vitamin contained in 145 of a gram of the
Reference Oil taken as a standard.

Vitamin E or Anti-sterility Vitamin is a fat-soluble principle
which has been found essential for the normal maturation of
the germ cells of the male animal and for the maintenance of
the placenta in the female. Its most abundant source is wheat

NON-PROTOPLASMIC CELL CONTENTS 107

germ oil. It also occurs in lettuce, water cress, whole cereals,
nuts, in cocoanut, corn and cottonseed oils, meat, etc.

VITAMIN K or Anti-hemorrhagic Vitamin has been found to
represent the substance necessary in the diet of chickens (and
possibly other animals) for the prevention of multiple hemor-
rhages. It has been found in the fat of hog liver, in tomatoes,
kale and hemp seed.

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 (primary cell wall) consists of pectose or a related car-
bohydrate. Later the daughter protoplasts secrete two layers of
a mixture of cellulose, (CeH10O5)n, and protopectin, one on either
side of a middle lamella, the latter composed of calcium pectate.
Each of these layers on either side of the middle lamella represents
a secondary cell wall. A tertiary wall may be laid down by the
protoplast beneath the secondary wall. (See Fig. 14.)

The cell walls may remain of such composition or become
modified to meet certain functions required of them. ‘Thus, in the
case of outer covering cells as epidermis and cork, whose function
is that of protecting the underlying plant units, the walls become
infiltrated with cutin (in the case of epidermal cells), suberin
(in the case of cork cells), 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 sclerenchyma
fibers whose function is that of giving strength and support to the
regions wherein found, the walls become infiltrated with dignin
which increases their strength, hardness, and in the case of
sclerenchyma fibers, their elasticity also. Moreover, in the case
of the cells comprising the testa or outer seed coat of the pumpkin,
squash, mustard, flax, psyllium, etc., whose function is that of
imbibing quantities of water, the walls undergo a mucilaginous
modification. These, upon the imbibition of water, swell up and
form layers of mucilage within the cell cavities.

The sub-epidermal cells beneath the upper epidermis of
Buchu leaves exhibit a striking example of mucilaginous modi-

108 PHARMACEUTICAL BOTANY

fication of cellulose as to their walls. If transverse sections of
either fresh or dried Buchu leaf-blades are first mounted in
strong alcohol and observed under the microscope, the walls of
the sub-epidermal cells appear no different from ordinary
cellulose walls; if, however, the alcohol be gradually withdrawn
from beneath the cover slip and gradually replaced with water,
these walls will be seen to swell and form layers of mucilage in
the cell lumina. The mucilage formed absorbs water so rapidly
that, in a relatively short time, the pressure (turgor) within the
cells becomes so great as to cause a rupture of the less elastic
vertical walls of the sub-epidermis with a consequent separation
of the upper epidermal cells from those of the palisade layer
beneath and the appearance of a broad rent in the section.

In pathological conditions, as a result of natural or induced
injury to the tissues, the walls of cellulose and pectose may be
completely transformed to mucilage. This condition is called
gummosis. ‘The gum drugs Acacia and Tragacanth are produced
in this way.

The walls of some bacteria, other fungi and some algae are
infiltrated with chitin. Those of the outer. layers of stems of
grasses, sedges and horsetails contain silica deposits, while
many plant walls contain calcium oxalate and calcium carbonate.

GROWTH IN AREA AND THICKNESS.—The cell wall when first
formed is limited in both extent and thickness. As the proto-
plast 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.

Pores (Prrs).—Pores or pits are small thin spots or holes in
the cell wall. The following terms are used to describe the
character of these structures:

Beaded. Circular pores arranged close together in a single
row.

Bordered. Pores partially covered on either side of the middle
lamella by a thickened, overhanging projection of the cell wall.

Branching. Pore canals which divide and sometimes sub-
divide.

CELL WALLS

109

Various Kinps or CELL WALLS AND BEHAVIOR OF EACH To MICROCHEMICG REAGENTS

Nature of wall

Where found

Reagent and behavior toward same

Lignocellulose
(Lignified wall).

Reserve cellulose.

Mucilaginous
modification of
cellulose.

Suberized walls. .

Cutinized walls. .

Callus of sieve
plates.

Silicified walls...

Parenchyma cells, tri-
chomes such as cotton,
etc,

Woody parts of plants,
such as stem cells, bast
fibers, wood fibers, etc.

Found in certain seeds
such as nux vomica,

ignatia, ivory nut,
date, coffee, etc.
In various parts of

plants.

In cork, wounded areas
of plants, endodermis.

Forming outer walls of
many epidermal cells.

Plates of sieve tubes.

Epidermis of Horsetails,
Grasses, etc.; Diatoms.

Schweitzer’s reagent dissolves it.
Chlorzinciodine solution imparts a
blue or violet color. Iodine solu-
tion followed by sulfuric acid colors
it blue.

Phloroglucin with HCl imparts a red
color except to bast fibers of flax,
mezereon, etc, Corallin-soda solu-
tion imparts a pink color. Aniline
sulphate with H.SO, colors it a
golden-yellow. Chlorzinc-iodine
imparts a yellow color.

As for cellulose,

Alcoholic or glycerin solution of
methylene-blue imparts a_ blue
color.

Alcoholic extract of chlorophyll, in
the dark, imparts a green color.
Alcannin and Sudan III impart a
red coloration. Converted into
yellowish droplets and granular
masses upon heating with a strong
solution of KOH. Sulfuric acid is
resisted.

As for suberized walls.
Corallin-soda solution imparts pink
color.

Soluble in hydrofluoric acid.

110 PHARMACEUTICAL BOTANY

Pores (continued)

Elliptical. Somewhat elongated pores with rounded ends.

Oblique. Slit-like pores in a slanting position in the cell wall.

Simple. Unbranched or non-bordered pores.

Slit-like. Very narrow, elongated pores.

Transverse. Slit-like pores extending in a direction perpen-
dicular to the lateral cell walls.

CHAPTER VI

PLANT TISSUES

A TissuE is an aggregation of cells of common source,
structure and function in intimate union.

Tue Tissues oF SPERMATOPHYTES AND PTERIDOPHYTES

The tissues of seed plants and pteridophytes are all derived
from a fertilized egg (zygote) which has undergone repeated
divisions. At first either an apical cell arises (pteridophytes)
or a mass of cells is formed (spermatophytes) at the tips of grow-
ing regions which are essentially alike and constitute the pri-
mordial meristem; 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 com-
pressed, elongated, and highly modified in respect to their
contents and walls.

CLAssIFICATION OF ‘TissuEs.—Tissues may be classified
according to method of development, form, structure, function,
and whether simple or complex.

According to their method of development, tissues are either
primary or secondary. Primary ‘Tissues are those which
develop at the growing points of different organs by cell division.
They arise from apical and intercalary meristems. They
include parenchyma, epidermis, endodermis, pericycle, primary phloem,

primary xylem, and cambium.
111

112 PHARMACEUTICAL BOTANY

SECONDARY TissuEs are those which take their origin in a
cambium or cork cambium. Examples of secondary tissues

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ylem from the Procambium 4 Xylem from the procambiumn
Xylem.from the cambium Xylemfrom the cambium

Phloem from the procambium and cambium Phloem from the procambium and cambium

Fic. 56.—Diagram showing the evolution of tissues in a stem from the pri-
mordial meristem down to the beginning of cambial activity. In the longitudinal
diagram, at the bottom, the initial C of the word cambium stands directly beneath
this tissue which is radially but one cell in thickness. (After Stevens.)

oe

are secondary xylem and secondary phioem which arise from a primary
cambium, and cork and secondary cortex which arise from cork
cambium. ‘These will be discussed more fully in a later part of

this chapter.

PLANT TISSUES 113

Srmp.e Tissues are those which are composed of one kind of
cell, as ordinary parenchyma, collenchyma and sclerenchyma, whereas
CompLex Tissues are made up of two or more kinds of cells as
xylem and phloem.

The adult or permanent tissues represent those which have
been laid down by the cell-division of meristems.

Primary MEeErIsTeMs.—As was shown by Hanstein,' 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 arrangement 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” the Calyptrogen
layer. These primary layers or groups of cells are called
primary meristems or generative tissues. ‘They are composed of
more or less 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 primary meristems or generative tissues are found in the
growing apices of plant organs, such as root, stem and leaf
apex. By the division and redivisions of their cells they give
rise to the mature or adult tissues of plants.

1. DeERMATOGEN or PROTODERM Originates epidermal tissue
and derivative structures such as stomata, non-glandular and
glandular hairs, glands, and sometimes cork cambium.

2. PeRIBLEM or GROUND MERISTEM originates primary cortex
tissue, chlorophylloid cells (chlorenchyma), colloid cells (collen-
chyma), strengthening cells (sclerenchyma), crystal cells (raphi-
derchyma), latex cells (lacterchyma), pericycle, endodermis,
sometimes cork cambium and pith.

3. PLEROME or PRocamsriuM originates the primary or first
fibrovascular bundles, the cambium and sometimes the pith.

According to the structure of the component cells and their
location, the following tissues are found in various forms of higher

plants:

. Hanstein, “Die Scheitelzellgruppe im Vegetationspunkt der Phanerogamen,”

Bonn, 1868.
2 Am. Sci. Nat. 5 série, tom. xx.

114 PHARMACEUTICAL BOTANY

1. MERISTEM 8. Cork
2. PARENCHYMA 9. LATICIFEROUS TISSUE
3. COLLENCHYMA 10. SIEVE OR CRIBIFORM TISSUE
4. SCLERENCHYMA 11. TRACHEARY TISSUE
5. EprmpermIis 12. MEDULLARY RAYS
6. ENDODERMIS 13. GLANDS AND SECRETION RESERVOIRS
7. PERICYCLE 14. NECTARIES
MERISTEM

MERISTEM, frequently called embryonic tissue, is undifferenti-
ated tissue composed of cells in the state of rapid division. The
first kind of meristem to appear arises from the division of apical
cells at or near the apices of roots, stems and leaves and is in
these regions called PrimorpiAL MeristeM. The primordial
meristem gives rise through cell divisions and slight differentia-
tion to the Primary Meristems (flerome, periblem and dermatogen)
which in turn develop the primary permanent tissues of plants.
Primary meristems retain the power of independent growth and
capacity for division as long as the plant part survives which
contains them. Meristem is also found in other regions of
plant organs such as the cambium and cork cambium, and is
there called SeconpARY MeristeM. Secondary meristem is a
derivative of primary meristem. It gives rise to the secondary
permanent tissues. It loses with its transition to the permanent
tissues the power of division and independent growth.

CampiuM is a secondary meristem lying between the phloem
and xylem in certain collateral bundles (intrafascicular cam-
bium) or between the phloem and xylem portions of the medul-
lary rays (interfascicular cambium). It is cylindrical and
extends longitudinally through the plant body. The cambium
is found in roots and stems of gymnosperms and dicotyledons
where it divides to form secondary xylem and secondary phloem.
These formations are responsible for the increase in diameter
of these organs. (Cf. Fig. 76.)

Cork CAMBIUM or phellogen originates in the pericycle of roots
and in the epidermis or outermost layer of the cortex of stems.
Through division of its cells it cuts off cork on its outer face and
frequently secondary cortex on its inner face. (See Fig. 70.)

PLANT TISSUES 115

According to the position of the meristems in the plant, they
may be classified as apical, intercalary and lateral.

ApIcAL MERISTEMs consist of growth cells which occur at the
tips of roots and stems. They produce growth in length by giving

Thy
So it, ee

Fic. 57.—The cambium and medullary rays in tangential-longitudinal section.
x 50. The section has been cut through the cambium zone adjacent to the
phloem or inner bark of the stem of the tulip poptar, Liriodendron tulipifera, and shows
besides the elongated taper-ended cambial cells, phloem parenchyma which has been
cut off by the transverse divisions of cambial cells, also medullary rays. ca, cambial
cell, in the upper portion of which two phloem cells have been cut off; ca’, cambial
cell at left of which is a row of phloem parenchyma; r, medullary ray.

rise to primary tissues. In the bryophytes and pteridophytes the
apical cell represents apical meristem, while in spermatophytes
a group of apical cells makes up this tissue.

INTERCALARY MERISTEMS represent parts of apical meristems
which have been separated*from the rest of apical meristem by

116 PHARMACEUTICAL BOTANY

permanent tissues and left with the latter as the apical meristem
advances in growth. They are found just above or below the
nodes of many stems and at the bases of some leaves, especially
those of monocotyls.

LaTERAL MERIsTEMS are those which occur lateral in an
organ, as the cambium and cork cambium.

PARENCHYMA

PARENCHYMA or FUNDAMENTAL TIssuE is the soft, primitive
tissue of plants, consisting of cells nearly. equal in length, breadth
and thickness (isodiametric) with usually thin, cellulose cell
walls enclosing cytoplasm and a nucleus and frequently sub-
stances of a non-protoplasmic nature. It is capable of cell
division after maturity. There are four. generally recognized
types of parenchyma, viz.:

OrDINARY PaRENCHYMA (Sorr GrounpD ‘TissuE, FUNDA-
MENTAL TissuE).—Next to the meristem this is the least modified
of all plant tissues. It is generally composed of thin-walled
cells, commonly polyhedral or spheroidal in form and often of
approximately the same length, breadth, and thickness (iso-
diametric); the cell walls are composed of cellulose which is
usually unmodified. Intercellular air-spaces occur in the angles
of adjoining cells. Occasionally the outline of the cells is star-
shaped or stellate, as in the stems of Wood Rush or Pickerel
Weed or the cells may be several times as long as wide, as in the
stems of Geranium, etc. Moreover, markings may occur on
the walls. These may be of the nature of pores, as in the
parenchyma cells of the pith of the Elder or Sassafras, annular
thickenings, as in the Mistletoe, or spiral thickenings, as in
certain Orchids. Protoplasm and a nucleus are always present
as well as colorless plastids, but in old cells are only seen as a
thin layer pushed up against the cell wall. Ordinary Paren-
chyma may be seen composing the soft tissues of roots, stems,
and barks. (See Fig. 69.)

ASSIMILATION PARENCHYMA (CHLOROPHYLL PARENCHYMA,
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

PLANT TISSUES 117

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. Good examples of
assimilation tissue are seen in the green column-shaped cells
below the upper epidermis of foliage leaves which, because of
their shape, are termed “‘falisade parenchyma cells’ and in the
green rounded to irregular-shaped cells beneath the palisade
parenchyma of foliage leaves which, because they are more
loosely arranged around air-spaces, ¢

are termed “‘spongy parenchyma cells.” “\~——
(See Fig. 166.)

ConpucTING PARENCHYMA.—
This type of parenchyma functions
in the rapid translocation of food
materials to distant regions in the
plant. It includes the wooed paren-
chyma 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

: : Fic. 58.—Transverse section of
(carbohydrate and proteid material part of leaf-stalk of a begonia.

in solution). Conducting paren- ¢, Epidermis; c, cuticle; B, collen-

iia ‘differ fram those of chyma, with walls thickened at
chyma ce. the angles, v;_ chl, chloroplasts.

ordinary parenchyma in being (¢4),¢ after Vines.)
usually more elongated and in con-
ducting soluble food materials with greater celerity.

RESERVE PARENCHYMA.—This resembles ordinary paren-
chyma 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 under-
ground stems such as tubers, corms, and bulbs.

CoLLENCHYMA.—This form of tissue is characterized by its
cells being prismatic, more elongated than ordinary parenchyma,
with soft, plastic walls of cellulose thickened in areas, usually
in their angles, but sometimes on the tangential walls or on
the sides abutting air-spaces, with a colloidal substance. The
thickened areas are in the form of longitudinal strips. The

118 PHARMACEUTICAL BOTANY

cells, like those of parenchyma tissue, contain cytoplasm and a
nucleus, and frequently chloroplasts. It remains alive after
maturity (Fig. 58). Collenchyma is generally found under-
neath the epidermis in the elongating parts of many stems, less
often in the midribs and petioles of leaves, and gives support
to these regions. It is the first strengthening tissue to develop
in stems. It is frequently observed forming the “ribs’’ of stems
and fruits of the Parsley Family and ‘‘ribs”’ of stems of the Mint

Fic. 59.—Stone cells from different sources. 1, 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.)

Family. 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 support-
ing elements having thickened cell walls usually composed of
lignocellulose. When first formed these cells resemble those of
ordinary parenchyma in having walls of pure cellulose, but later
lignin, a hard, woody substance, becomes deposited on the inner
surface of the walls in one or more layers. (Occasionally, as
in the bark region of stems of Flax and Mezereon, no lignin is

“ PLANT TISSUES 119

deposited on the walls of the sclerenchyma fibers.) When
sclerenchyma is composed of cells which are more or less iso-
diametric or moderately elongated, with thickened, lignified

| ae? Si (PERE pp

Fic. 60.—Stone cells from various sources. 1, From olive pit; 2, from cocoa-
nut endocarp; 3, from flesh of pear; 4, from aconite root; 5, from capsicum; 6,
from hazelnut; 7, from allspice. (Drawing by Hoffstein.)
walls and conspicuous pores, its elements are called STONE
Cets. Stone cells are distributed in fruits, seeds and barks
of many plants, rarely in woods. They occur singly or in small
groups or large masses. ‘They have been found forming the

120 PHARMACEUTICAL BOTANY

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
examination a cell wall of cellulose with one or several layers
of lignin on its inner surface which surround a central lumen.

Fic. 61,—Sclerenchyma fibers from different sources. 1, From powdered
cinnamon bark; 2, end of bast fiber of flax stem showing transverse markings
(4); 3, middle portion of flax fiber showing characteristic cross markings at }; 4,
bast fiber from cinchona bark; 5, branched bast fiber from choke cherry bark; 6,
above, end, and below, medium portion of bast fiber of jute. All highly magnified.

The latter is in communication with radial pore canals leading
outward to the middle lamella. Longitudinal pore canals
are also evident.

Fisers.—When sclerenchyma is composed of cells which are
greatly elongated and more or less obtusely or taper ended, its
component elements are termed SCLERENCHYMA FIBERS or
Firsers. These fibers are frequently spindle-shaped. When
young they contain protoplasts but when mature their lumina
usually contain air. They exhibit pits of the nature of oblique

sai ih

121

PLANT TISSUES

They are either polygonal,

slits or rounded pores in their walls.

They

rectangular or somewhat rounded in transverse section.

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Fic. 62.—Transverse section of a portion of the wood of the Black Ash.

A,

bers; mr, medullary

wf, wood fi

170. Note the polygonal to rounded-polygonal shape of the

summer wood; B, autumn wood; C, spring wood.
wood fibers in cross section.

ray; tr, tracheae.

occur in various par
supporting elements.

ts of roots, stems, leaves, fruits and seeds as

When sclerenchyma fibers occur in the

, they are termed Xylem

xylem region of fibro-vascular bundles

122 PHARMACEUTICAL BOTANY

Fibers, or Wood Fibers; when they appear in the phloem region,
Phloem Fibers, Bast Fibers or Liber Fibers; when in the pericycle,
Pericyclic Fibers; when in the cortex, Cortical Fibers. Wood fibers
are usually cut off by the cambium only and so are generally

Ti
, Te
Fic. 63.—Tangential-longitudinal section of wood of the Black Ash. 170.
wf, wood fibers; mr, medullary rays. Note how the-tapered ends of the wood fibers
are spliced over each other.

seen in the secondary xylem. They differ from the fibers of
the phloem, pericycle and cortex in that they possess bordered
pits although in many instances these are so reduced by the
thickening of the wall as to be simple. Phloem, pericyclic and
cortical fibers possess simple pits.

EPIDERMIS

EpierMis is the outer covering tissue of a plant and is pro-
tective in function. It provides against mechanical injury and

PLANT TISSUES 123

loss of water. In old stems and roots of secondary growth it is
ultimately replaced by cork. Its cells are living, containing a
thin marginal layer of cytoplasm, a nucleus and a large central
vacuole and frequently leucoplastids. Non-protoplasmic inclu-
sions as crystals, mucilage, etc., may occur in the lumen. They
may be brick-shaped, tabular or polygonal, the vertical wall being
equilateral or wavy in outline. Their outer walls are frequently
cutinized (infiltrated with a waxy-like, waterproof substance called
cutin). The radial walls also are sometimes cutinized.

Fic. 64.—Upper epidermis of Sweet Fern (Comptonia asplenifolia) leaf (surface view)
showing epidermal cells and two non-glandular trichomes.

Stomata.—Among the epidermal 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 guard cells contain chloroplasts and are capable of
undergoing changes in size and shape dependent upon changes
in their turgor. Increased turgor causes them to open. A
decrease in turgor causes them to close. The stomata are in
direct communication with the sub-stomal air chambers beneath
them which in turn are in communication with intercellular
space systems of the tissues of the entire plant. Through the
stomata an exchange of gases between the tissues of the plant
and the external air takes place. The function of the stomata is.

-

124 PHARMACEUTICAL BOTANY

to give off watery vapor (transpiration) and take in or give off
carbon dioxide, and oxygen. In addition to stomata some
leaves possess groups of water stomata which differ from the
ordinary or transpiration stomata in that they always remain open,
are circular in outline, give off water in droplets directly, and
lie over an area of small, secreting, glandular cells (epithem)
which is in connection with a bundle of tracheids that springs
from the end of one or more fibro-
vascular bundles. An air chamber usu-
ally occurs directly beneath the water
pore. Haberlandt gave the name of hyda-
thode to this and related types of structures
which secrete water in the form of drop-
lets. Examples: Leaves of Primula,
Corn, various grasses, Potato, Elm, Lark-
spur, Crassula, Saxifraga and Ficus.
EpmERMAL AppEeNDAGES.—The epi-
Fic. 65,—Radial-longi- Germis of leaves, stems, fruits, and seeds
tudinal section through a Of many plants frequently gives rise to

hydathode from the leaf oyt-growths in the form of papilla, hairs
margin of Primula sinensis. d ]

i, upper, and j, lower epi- sspeae -nabinge

dermis; h, palisade cell; ¢, EPIDERMAL Papi_L@ are short pro-

thin walled parenchyma tuberances of epidermal cells. “They may
eset reusnd be seen to advantage on the upper epi-
water stoma; &, tracheids, Germis of the ligulate corolla of various
(From Stevens, after Haber- species of Chrysanthemum, on the lower epi-
londt.) dermis of the foliage leaves of species of
Erythroxylon and upon the upper epidermis of the petals of the
Pansy (Viola tricolor). EpmpERMAL Hairs or TRICHOMES are more
elongated outgrowths of one or more epidermal cells. “They may
be unicellular, as those of the seed of 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 Hyoscy-
amus, Belladonna and Digitalis leaves; hooked, as on the stems
of the Scarlet Runner or Hops; barbed, as on the stems of Loasa
species; or tufted, as found on the leaves of Horehound (Marru-

PLANT TISSUES 125

Fic. 66.—Trichomes from different sources. 1, Unicellular non-glandular
trichomes as seen growing out of epidermal cells of Senna; 2, uniseriate nonglandular
trichomes of Digitalis; 3, unicellular stellate trichomes from Deutzia scabra; 4,
unicellular twisted trichomes from lower epidermis of Eriodictyon; 5, clavate non-
glandular trichomes from scraping of epidermis of the fruits of Rhus glabra; 6, 2-
branched trichomes of Hygoscyamus muticus, a substitute for Henbane; 7, branched
multicellular trichome of Marrubium; 8, glandular trichomes from strobile of
Humulus (Lupulin) 9, glandular trichomes from leaves of Digitalis purpurea; 10,
aggregate, non-glandular trichomes of Kamala; 11, lateral view (to left) and vertical
view (to right) of glandular trichomes of Kamala; 12, vertical view (above) and pro-
file view (below) of 8-celled glandular hair from Mentha piperita. All highly
magnified.

126 PHARMACEUTICAL BOTANY

bium vulgare). They may be simple as in Cotton, etc., or
branched as in Hyoscyamus muticus.

Nonglandular hairs are said to be uniseriate when they consist
of a single row of cells, as those of Digitalis or Stramonium, etc.,

wd, YgX>

Fic. 67. Fic. 68.

Fic. 67.—1, Epidermis of oak leaf; 2, epidermis of Iris leaf, both viewed from
the surface; 3, group of cells from petal of Viola tricolor, showing conical papille;
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; 5 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.)

Fic. 68.—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 remainder from Haberlandt.)

multiseriate, when they consist of two or more rows of cells, as
those of the corolla of Calendula.
The glandular hairs comprise those whose terminal cell or
cells are modified into a more or less globular gland for gummy,
resinous or 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, uniseriate (consisting
of a single series of superimposed cells) or multiseriate (consisting

PLANT TISSUES 127

of two or more rows of superimposed cells). The head varies
from a one- to many-celled structure. The drug Lupulin consists
of the glandular hairs separated from the strobiles of the ioe
(Humulus lupulus).

ScALEs are flat outgrowths of the epidermis composed of
one or several layers of cells. They occur attached to the stipes
of Aspidium, Osmunda and other ferns, where they are called

“chaff scales.””» They are also found on a number of higher
plants.

Functions or Harrs.—Plant hairs are adapted to many dif-
ferent purposes. They may absorb nourishment in the form of
moisture and mineral matter 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 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 layer of cells, constituting the innermost
layer of the cortex. It forms a cylindrical sheath around the
stele in roots and stems of pteridophytes and spermatophytes.
It also occurs around the vascular bundles of leaves. In
Angiospermous stems it usually resembles the other parenchyma
layers of cortex as to structural characteristics, save that it
frequently contains more starch. In fern stems, roots of Mono-
cotyledons and of Dicotyledons of primary growth, however, its
cells are clearly distinguished from the other cells of the primary
cortex by their vertically elongated form and cutinized or sub-
erized (occasionally lignified) radial and end walls.

_ The most outstanding characteristics of typical endodermal
cells are to. be found in their walls. These may be thin or thick

128 PHARMACEUTICAL BOTANY

but the radial and end walls are usually thickened with deposits
of a waxy, waterproof material representing cutin or suberin.
The thin-walled endodermal cells possess strips or bands of
cutin which extend around the cells on the inner surface of the
radial and end walls. These strips or bands are called Casparian

1D Epidermis
ap Seseestoe
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Fic. 69.—Cross section of the stele and a portion of the epidermis and cortex of
the monocotyledonous root of the corn. The outgrowths of the epidermis are root
hairs. (From “A Textbook of General Botany” by Smith, Overton et al., Macmillan Co.,

publishers.)

strips. In cross sections the Casparian strips often appear like
dots or lenticular shaped areas in the radial walls and are
frequently termed ‘‘Casparian dots or spots.’ 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

PLANT TISSUES 129

nucleus; starch grains are frequently but not always present
and sometimes mucilage, tannin and crystals occur. Its
functions seem to be to give protection to the stele (tissues within
it), to reduce permeability between primary cortex and stele

Py.

Fic. 70.—Transverse section of the outer regions of the stem of the elder (Sam-
bucus canadensis), showing cork, cork cambium, etc. (From Strasburger, Macmillan
Co.)

and to serve as an air-dam, preventing the clogging with air of
the water-conducting elements.

Cork

Cork or suberous tissue is a protective tissue composed of
cells of tabular shape, whose walls possess a layer of water-proof
substance called suberin. It is formed by the division of the
cells of the phellogen or cork cambium which cuts off cells out-
wardly. Its cells, when first formed, contain cytoplasm and
nucleus but, as the cork cambium forms new layers of cork on
its exterior, the older cork cells lose their protoplasmic contents,
die, become filled with air or a yellowish or brownish substance,
and gradually sluff off. Under the microscope, dead cork cells
usually appear black due to air filling their cavities. Thin
sections of cork, when mounted in suitable reagents, will show

in. PHARMACEUTICAL BOTANY

that the walls of the cork cells are composed of three layers or
lamella. The lamella next to the cell cavity is cellulose.
Outside of this is a middle lamella which may be lignified and
a lamella of suberin. Cork tissue is usually devoid of inter-
cellular-air-spaces. It forms a protective covering to the roots
of secondary growth, perennial stems (after the first season)

i paces

wt ln

cs i £ Naat
»“ a @ as oe we 4
a we —, & 3

roseum. X50. ep, epidermis; co, primary cortex; fer, pericycle filled with peri-
cyclic fibers; ph, phloem; ca, cambium (intrafascicular); x, xylem; mr, medullary
ray; m, pith, composed of ordinary parenchyma.

of Dicotyledons and Gymnosperms, and wounds of stems and
branches. It protects the delicate underlying tissues from
invasion by fungal parasites, from sudden changes in tempera-
ture, and from loss of water.

The suberized walls of cork cells resist the action of concen-
trated sulfuric acid. They are colored green, when in contact
with alcoholic extract of chlorophyll for several days in the dark.

RHYTIDOME sometimes called “scale bark” or “shell bark”
represents the outer crust of alternating cork and cortical or
pericyclic and phloem tissues resulting when cork cambia arise

PLANT TISSUES 131

in deeper regions of the bark and form cork or periderm. (See
Fig. 131.)

BorkE is a term which has been applied by German botanists
to designate the dead tissues which sluff off when cork cambium
arises and forms layers of cork in deep regions of the cortex,
pericycle and phloem. ‘This occurs in the oak, birch, grape
vine, elm, viburnum, hickory, cherries, etc.

PERICYCLE

The PericycLe or PERICAMBIUM, as it was formerly called,
is a derivative of the ground meristem. It is a narrow cylinder of:
tissue which. lies between the endodermis and the outer margin
of the primary phloem of the fibrovascular bundles of stems or
between the endodermis and the radial fibrovascular bundle of
roots. In many stems it is composed entirely of thin-walled
parenchyma cells but in others, whether woody or herbaceous
and notably climbing or twining stems, it is composed of two
kinds of tissues, thin-walled parenchyma and sclerenchyma.
Frequently stone cells as well as sclerenchyma fibers occur in
this region, as in stems of Cinnamon, etc. The sclerenchyma
fibers occurring in the pericycle are called “‘pericyclic fibers.”
Normally these fibers are lignified but may consist of cellulose,
as in Flax and Mezereon stems. The parenchyma cells
sometimes contain chloroplasts and function in the slow con-
duction of sap and the storage of starch. The sclerenchyma
fibers give strength. In roots, the pericycle consists of one or
two rows of thin-walled cells which undergo division in places
to form meristems which give rise to lateral roots. In roots of
primary growth of Exogens the pericycle divides into two layers,
the outer layer becoming a phellogen or cork cambium. The
pericycle also may give rise to meristems forming adventitious

roots and stems.

LATICIFEROUS ‘TISSUE

This form of tissue comprises either /atex cells, latictferous
vessels, or coenocytic latex cells, differing from each other in
origin and method of development. (1) Latex Cexts (latex
sacs) are uninucleate cells which take their origin from minute

y
i

132 PHARMACEUTICAL BOTANY

meristematic cells of the embryo. They are arranged in longi-
tudinal rows and often occur in all of the organs. Each latex
cell is a single unit and does not anastomose with other latex

: bem PVH)

Fic. 72.—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.)

cells. Such cells are abundant in the following families:
Apocynaaceea, Papaveracea, Urticacee, Sapotacee and Convolvulacea.

(2) Latictrerous Vessets are long 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

PLANT TISSUES 133

roots, stems, and leaves. The branches connect with those of
other tubes forming an anastomosing network. ‘These vessels
occur in the following families: Composite, Papaveracee, Campanu-
laceea, Convolvulacea, Euphorbiaceae, Aracee, Oleaceea, Geraniacea,
and Musacee.

(3) Ca@NocYTIG LATEX CELLS are elongated, multinucleate,
laticiferous cells which occur in the milkweed family (Asclepiada-
cee) and in Euphorbia species. ‘They arise, in each instance, from
a single cell in the embryo which elongates at an equal rate with
the growth of the plant, extending itself between other cells like a
filamentous hypha of a mold, branching freely, but unlike
laticiferous vessels the branches do not anastomose.

All of the latex elements are living, possessing thin layers of
protoplasm just within their walls and containing one or more
nuclei depending upon the type, also at times, /eucoplastids,
elaioplastids and proteinoplastids. The leucoplastids form starch
grains, the elaioplastids build up oil globules and the proteino-
plastids organize protein granules.

The exact role of latex elements remains undiscovered.
Some botanists believe them to be food reservoirs, others excre-
tory reservoirs, while still others regard them as representing a
conducting system for the movement of starch and proteins.

All lacticiferous 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 ‘“‘/atex.”’ For further details see under ‘‘Latex”’

in Chapter V.

Sreve (LEPTOME OR CRIBIFORM) Tissur

This tissue, found in the phloem region of fibro-vascular
bundles, is a product first of the procambium and later of the
cambium. It consists of superimposed, elongated, tubular
cells whose longitudinal walls are thin and composed of cellulose
and whose oblique or transverse walls, called “sieve plates,” are
thickened and perforated, permitting of the passage of proteids,
amino-acids and amides from one cell to another. Frequently
sieve plates are formed on the longitudinal walls, as in Cascara
Sagrada. The mature sieve tube has a thin layer of cytoplasm

*

134 PHARMACEUTICAL BOTANY

lining its walls and a large central vacuole but no nucleus.
The tubes are arranged in longitudinal rows and the protoplasts
of each series are connected by means of protoplasmic strands
which extend from one sieve. tube to

another through perforations in the sieve #
plates. The cell sap found in the vacuole sees
of sieve tubes contains protein substances o|

often of slimy character. When fixed assed

990

939

°
3

9°

t)

Fic. 73. Fic. 74.

Fic. 73.—Stages in the development of sieve tubes, companion cells, and
phloem parenchyma. A. a and 6, Two rows of plerome cells; in c and d, a has
divided longitudinally and ¢ is to become companion cells; d, a sieve tube, and
b, phloem parenchyma. B. c, Companion cells, and d, a beginning sieve tube
from ¢ and d, respectively in A. The cross-walls in d are pitted; 6, phloem paren-
chyma grown larger than in A. C. The same as B with the pits in the cross-walls
of the sieve tubes become perforations, and the nuclei gone from the cells composing
the tube. (From Stevens.)

Fic. 74.—Vascular elements. A, annular tracheal tube; B, spiral tracheal
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
Strasburger.)

and stained sections are examined, coagulated masses of
protein will often be seen filling the pores of the sieve plates.
These are called slime blugs.

_ Sieve tubes are usually accompanied by companion cells
excepting in Pteridophytes and many Gymnosperms. Both

PLANT TISSUES 135

companion cells and sieve tubes arise by the division of the same
mother-cell of the procambium. ‘The companion cells occur in the
phloem of Angiosperms only. In cross sections they appear as
small, rounded or angular cells at the corner or lateral to a larger
sieve tube. In longitudinal section they are seen to be elongated,
slender cells. They may be distinguished from the mature sieve
tubes by their abundant granular, cytoplasmic contents, their
small vacuoles, and also by the fact that they retain their nuclei
after complete maturation. Their walls are pitted where they
are in contact with sieve tubes with which they are associated,
phloem parenchyma and medullary rays. ‘Their work is to take
over food materials from the sieve tubes and pass them on to
other tissues needing them for growth, repair or storage.

Besides sieve tubes and companion cells, phloem fibers also
called bast fibers, and parenchyma cells called “‘phloem-parenchyma”’
or “‘steve parenchyma’’ are often found in thephloem. The phloem-
parenchyma cells transmit soluble carbohydrates, amino-acids
and amides slowly downward and, through their communication
with medullary rays, deliver food materials to them for radial

distribution and storage.

TRACHEARY ‘TISSUE

The tracheary tissue of plants comprises two kinds of elements,
the trachee (ducts or vessels) and tracheids. They are formed first
by the procambium or plerome and later by the cambium.
Both of these conduct crude sap (water with mineral salts in solu-
tion) upward through the roots and stems into the leaves. The
trachee are very long tubes of a cylindrical or prismatic shape
which are formed by the breaking down of the transverse walls
between vertical groups of superimposed cells, during the growth
of the plant. Thousands of these trachea are arranged one
above the other from the smallest branches of the root to the
highest part of the stem and out into the farthest reaches of the
leaves. Cross or end walls which are perforated with openings
separate each trachea from its neighbors. In this way each
line of trachez serves as a continuous tube for the conduction
of water rapidly from root to leaf. Most trachee have two
perforations, one in either end but as many as four have been

136 PHARMACEUTICAL BOTANY

found, each opening into a different cell. ‘The cross or end wall
separating one trachea from another usually possesses a single,
large, rounded or elliptical perforation called a simple perforation,
but some end walls possess two or more smaller perforations
forming an opening divided by transverse bars and called a
scalariform perforation. ‘The longitudinal or vertical walls of these
tubes are of varying thickness, usually, however, thinner than
those of woody fibers. ‘The thickness is due to an infiltration of
lignin upon the original cellulose wall. The walls show char-
acteristic thickenings on their inner surfaces. These thickenings
are for the purpose of giving strength to the tube while the thin
parts of the wall are for the passage into and out of the tube of
water with nutrient materials in solution.

TRACHE/# 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.

Bordered pored, with rows of pores or slits, each being sur-
rounded by a thickening of variable shape (circular, ellipsoidal
- and prismatic are the usual shapes).

Annulo-spiral, with both ring and spiral thickenings.

Scalariform, with ladder-like thickenings.

TRACHEIDS are primitive conducting xylem cells which have
thickenings similar to trachea but are usually, though not always,
smaller in size. Their ends are usually tapered, sometimes
chisel-like, but not sharply pointed. They may be distinguished
from trachez: by the absence of perforations in their end walls.
Their end walls have, however, thin places or pits which permit
the more rapid passage of sap from cell to cell. Like trachee
their walls usually give the characteristic lignin reaction with
phloroglucin and HCl. The tracheids of conifers (see Fig. 75E,
D.) such as the pine, spruce, cedar, etc., usually show one row
of good-sized bordered pores (bordered pits) on their longitudinal
walls. Each bordered pore exhibits a wall surrounding the pore
which forms a dome-shaped protrusion into the cell. Like
trachez, also, tracheids convey water with mineral salts in solu-
tion upward through the plant axis. Tracheids with bordered

PLANT TISSUES 137

>

Fic. 75.—Types of tracheae and tracheids. A, annulo-spiral trachea; B, spiral
trachea; C, tracheid with several rows of bordered pores; H, pitted vessel; £,
portion of a coniferous tracheid with a single row of bordered pits; D, tangential-
longitudinal section of a bordered pit and portion of adjacent wall shown in £,
enlarged; p, pore or orifice pit; 4, eeteenns border; cm, limiting membrane which

t);

bears at its center a thickened pod or torus ¢, cavity of pit; F, reticulate trachea in
longitudinal section; G, longitudinal section of spiral trachea.

138 PHARMACEUTICAL BOTANY

ae
fe)
ro)
SI
a oO
oO
°o
2bned 310
A
Z ah “ 4 j 5 an

Fic. 76.—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 overhanging 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 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

PLANT TISSUES 139

pits and medullary-rays make up the wood of Conifers and most
of the other orders of Gymnosperms.

The functions of tracheids are longitudinal conduction of
crude sap and support to the part containing them.

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 (sec-
ondary medullary rays or “‘vascular
rays”). Secondary medullary rays ©
occur only as a result of the activity
of the cambium during secondary
growth of the plant axis. The xylem
portion of a medullary ray is termed
a xylem ray or wood ray while the
phloem portion is termed a phloem ray. oy ove a
In _ tangential-longitudinal section 6 © Molle@
medullary rays usually appear

: : a Fic. 77.—Portion of radial-

- : e in

epacie shapes (Fig aP icy longitudinal section through
radial-longitudinal sections they are cq of White Pine stem. {,
seen crossing the other elements. tracheid; », bordered pore of
Their primary function is to supply ee medullary —_ ray.
the cambium and wood with elabo- ~*8"™""
rated sap formed in the leaves and conveyed away by the sieve
tubes and phloem parenchyma and to supply the cambium
and phloem with crude sap which passes up mainly through the
tracheze and tracheids from the absorptive regions of the roots.
They distribute food materials radially. They furthermore
with cells longer and more pointed and walls becoming thickened and pitted; 4,
complete wood fibers with walls more thickened than in the previous stage and
lignified, as shown by the stippling. The protoplasts in this last stage have disap-
peared 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.)

00)|\OCOOO

O||

140 PHARMACEUTICAL BOTANY

serve as storage places for starch, alkaloids, resins, and other
substances. (Cf. Figs. 62, 127, 139.)

VASCULAR AND FIBROVASCULAR BUNDLES

VASCULAR BuNDLEs are strands of conducting elements con-
sisting of elongated cells specialized for the conduction of sap.
The conducting strands are of two kinds: the PHtozm which
carries elaborated food materials (e/aborated sap) from the leaves
and green parts of stems to all parts of the plant, and the XyLem
which carries soil water with dissolved mineral salts (crude sap)
from the roots to the leaves.

Tracheal tube carrying food upward Sieve tube carrying food
downwards from
Kylem parenchyma recéiving 3] Chk eaves
P food from medullary rays }
> i “i\ and storing it if
2° p Lea? 4 2 09 \* el *%e?
id Ay *? 99 0\0% "| «
es yout \\e 47% Cambium { m5 oe.
oho (9¢ Sy "OO | ile@s oA
a Soe A ae hie Hie ee
Saare- em ae rarer — agaee oo oo = > ~ 3 é <
[40 9 eS Sloeee Va \\ ly de o,° ee
vo
f Parenchyma
Medullary ray cells carrying food inward and outward storing food

from the sieve tubes The ray cells also store food
Fic. 78.—Diagram showing the transport of food through the sieve tubes,
medullary-rays and tracheal tubes (trachez), and its storage in the parenchyma
cells of the wood and bark. The black bodies in the cells indicate stored food.
(Stevens.)

FIBROVASCULAR BUNDLES are groups of fibers, vessels and cells
coursing through the various organs of a plant and serving for
conduction and support. The term “Vascular Bundle’ is
employed to indicate bundles whose xylem and phloem contain
no fibers and which are devoid of enveloping fibrous sheaths.
According to the relative structural arrangement of their xylem
and phloem masses, they may be classed as follows:

I. Collateral, consisting of a xylem and a phloem strand
extending side by side.

II. Closed collateral, consisting of a strand of xylem lying
alongside of a strand of phloem, in intimate contact, with no
cambium zone between them. Stems and leaves of most Mono-
cotyledons and Horsetails.

III. Open collateral, consisting of a strand of xylem and a
strand of phloem separated from each other by a cambium.

PLANT TISSUES 141

Stems and leaves of Dicotyledons and roots of Dicotyls and
Gymnosperms of secondary growth. In stems and roots the
phloem faces toward the exterior and the xylem toward the
center of the axis.

IV. Bicollateral, characterized by a xylem strand befng
between an inner and an outer phloem mass. Cambium may

; VG :
Fic. 79.—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; S, companion cells; CP, crushed primary
sieve tubes; F, thin-walled parenchyma of the ground or fundamental tissue.
(From Sayre after Strasburger.)

be present or absent on either outer or inner face of the xylem
or on both faces. Seen chiefly in stems and leaves of the Cucurbz-
tacee, Combretacee, Loganiacea, Apocynacee, Asclepiadacee, Oleacea,
Convolvulacee, Gentianacee, Myrtaceae and Solanaceae.

V. Concentric, characterized by a central xylem mass sur-
rounded by a phloem mass or vice versa. No cambium present.

(a) Concentric, with xylem central in bundle. (Xylocentric
bundle.) Seen in stems and leaves of nearly all ferns and the
Lycopodiacea.

(b) Concentric, with phloem central in bundle. (Phlocentric
bnudle.) Seen in stems and leaves of some Monocotyledons. Exam-
ples: Calamus and Convallaria rhizomes.

142 PHARMACEUTICAL BOTANY

VI. Radial, characterized by two or more xylem and phloem
masses arranged in alternate manner usually around a central
pith. The terms diarch, triarch, tetrarch, pentarch, hexarch,
heptarch and polyarch are respectively employed to designate
radial bundles having 2, 3, 4, 5, 6, 7 or many xylem strands and

ss a

ay

Fic. 80.—Vascular bundles. A, the concentric-monocotyl or phlocentric
type, with xylem, k, surrounding the phloem, h. B, the collateral type, with
phloem, h, standing in front of the xylem,k. C,a portion of the radial type (under-
going transition) shown complete in D, where the part outlined at a corresponds
toC. c, xylem, b, phloem, f, cambium ring; e, pericycle; d, endodermis. Pericycle
and endodermis are not parts of the bundle. (From Stevens after Haberlandt.)

a like number of phloem strands. Seen in the primary roots
of all Spermatophytes and Pteridophytes.

XYLEM! is that part of a conducting or vascular bundle that
usually contains wood parenchyma cells, tracheze and tracheids.
It frequently contains, in addition, wood fibers. The tracheid
is the fundamental cell type in the xylem. The xylem of most
Gymunosperms is devoid of trachez.

* The conducting elements of the xylem comprising trachez, tracheids and
wood parenchyma constitute the hadrome. :

PLANT TISSUES 143

PRIMARY XYLEM (PROTOXYLEM) is that xylem in all conduct-
ing bundles which directly develops from the plerome. In
exogenous stems of secondary growth, the primary xylem is in
small strands adjacent to the pith, and in roots of secondary
growth it appears as a small, several rayed strand in the center.

:

a a
we ae 4

a a ek dite as pees |
Fic. 81.—Bicollateral bundle of the stem of Squash (Cucurbita). X70. P’,
external phloem; P?, internal or intraxylary phloem; x, xylem; ca, cambium; si, a

sieve tube.

SECONDARY XyLeM (METAXYLEM) is xylem layed down
by the cambium.

PHtoem! is that part of a vascular bundle that contains sieve
tubes, phloem parenchyma cells, and often bast fibers. The
sieve tubes are the most important structural and functional
elements of the phloem. In Pteridophytes and many Gymno-
sperms these.and phloem parenchyma are alone present in the

1 The conducting elements of the phloem which comprise the sieve tubes,
companion cells and phloem parenchyma constitute the leptome.

144 PHARMACEUTICAL BOTANY

phloem while in Angiosperms sieve tubes, companion cells,
phloem parenchyma, bast fibers, secretion cells and stone cells
may occur in this region.

PriMARY PHLOEM (PROTOPHLOEM) is the phloem formed by
the plerome and occurs directly beneath the pericycle.

Fic. 82.—A Polyarch Radial Bundle. Cross-section through a portion of a
root of Acorus calamus, a monocotyledon. A, Cortical parenchyma; B, endodermis;
C, pericycle; E, phloem; F, xylem. At Y, 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 polyarch, radial vascular bundle,
occur thin-walled, parenchymatous pith cells. (From Sayre after Frank.)

SECONDARY PHLOEM (METAPHLOEM) is phloem formed by
the cambium.

LeaF AND BrancH TRaces.—These are prolongations of the
vascular bundles from the stele of the stem which extend into
leaves and branches. (See Fig. 84.)

THE STELE

The STELE represents the central cylinder of a stem or root
and includes all of the tissues interior to the cortex.

PLANT TISSUES 145

A PRosTELE is a primitive form of stele in which the vascular
tissues form a solid cylinder, the phloem surrounding the xylem.
It occurs in young parts of stems of club-mosses, in the stems of
some ferns and in the roots of secondary growth of most plants.
(See Fig. 83A.)

A SIPHONOSTELE resembles the prostele except that a pith
occurs in its centre. There are two kinds of siphonosteles, viz.,
the ectophloic in which the phloem is external to the xylem, as

(a) ; 3 »

|

A B Cc
Fic. 83.—Diagrams showing the types of arrangement of vascular tissues in

steles. A, protostele. B, siphonostele. C, dictyostele. (From “An Introduction to
Plant Anatomy” by Eames and Mac Daniels, McGraw-Hill Book Co., Inc. Publishers.)

in the stems of gymnosperms and angiosperms, and (2) the
amphiphloic in which there is an external phloem outside the
xylem and an internal phloem on the inner face of the xylem,
as in the stems of some ferns and certain families of dicotyledons.
(See Fig. 83B.)

A DicryosTELe is a siphonostele whose vascular mass is
broken up into a number of longitudinal strands or vascular
bundles. This type is common to most monocotyl stems, the
bundles being sometimes called meristeles. (See Fig. 83C.)

BraNcH TRACES.—These are strands of vascular tissue which
arise from the vascular bundles of the stele of the stem and run
out into the lateral branches. Usually two traces or bundles
arise for each branch, but sometimes only one arises. When

146 PHARMACEUTICAL BOTANY

y
:
:
/
/
|
:
]
]
/
]
:
'

ee ee

CG
Yf ry)

Fic. 84. Diagrams illustrating leaf traces, branch traces and gaps. A, longi-
tudinal section of node through leaf trace and gap. __B, similar to A, but with branch
trace and gap also present. C, view of vascular cylinder showing departure of leaf
and branch traces, and the gaps associated with each. D, E, F, cross sections of
stem illustrated in A at levels a-a, 6-b, and c-c, respectively. G, face view of outside of
cylinder shown in C, the leaf and branch traces cut away at the surface of the cylin-
der. H, transverse section of G at a-a. (In diagrams A-H the vascular tissue is not
differentiated as xylem and phloem; the traces are doubly cross-hatched. In /,
more detailed structure is shown, protoxylem, metaxylem (secondary xylem) and
pith being indicated. (From “An Introduction to Plant Anatomy” by Eames and Mac
Daniels, McGraw-Hill Book Co., Inc., publishers.)

PLANT TISSUES 147

the branch supply consists of two bundles, these soon unite on
their way into the branch to form a cylindrical stele which
appears circular as viewed in cross section. In instances where
only one trace or bundle strand courses into the branch, this
appears in cross section as a horseshoe or crescentic structure
with an opening. The opening is closed as the trace passes
into the branch so that in the branch a complete cylindrical
stele is formed.

LEAF AND Brancu Gaps.—These are breaks in the continuity
of the vascular cylinder of stems around and above the places
where the leaf or branch traces leave the cylinder. The break
or opening is designated as a leaf gap or branch gap according
as to whether a leaf trace or branch trace is involved in its
formation. It contains parenchyma and through it the cortex
and pith are confluent. (See Fig. 84.)

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 (Schizogenous 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 (xerophytes) the intercellular-air-spaces are
avéragely small and more or less angular. In plants of swamp

148 PHARMACEUTICAL BOTANY

or marsh habit they are medium-sized, while in those which
live entirely in the water (Aydrophytes) they are of large size and
more or less rounded.

INTERNAL SECRETION GLANDS AND RESERVOIRS

These structures are either found as globular cavities as in
Orange and Lemon Peel and Eucalyptus leaves, containing oil

Seness

Y}
NW),

Fic. 85.—Cross section Fic. 86.—Lysigenous gland
through a portion of Orange in the leaf of Dictamnus fraxinella,
Peel showing the cavity of an B, young glands, with cells
internal, schizogenous glob- beginning to secrete oil; C,
ular gland at g; crystals of mature gland where the secreting
hesperidin at A; membrane cells have broken down and left
crystals of calcium oxalate their secretion in the cavity thus
at k. (After Tschisch and ' formed; o, large drop of secreted
Oesterle.) oil. (Stevens after Sachs.)

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 or reservoirs. In the Parsley family these canals
are found in the fruits, where they are called witte. Occasion-
ally they are named according to the nature of their contents—
resin or oil canal, reservoir, etc. The internal glands usually
arise by the division of a cell or group of cells of the ground

PLANT TISSUES 149

meristem but sometimes the dermatogen gives rise to them, as
in Clove, Myrtle, Asarum, etc., or both the ground meristem
and dermatogen may contribute to the formation of the same
gland. Internal glands and secretion reservoirs are termed
schizogenous when the secreting cells split apart at the center of
the group and draw away from the line of separation, as in the

Fic. 87.—Schizogenous secretion reservoir (resin duct) as seen in cross section
in the young stem of English Ivy. A, early and £, later stage in the formation of
the duct. g, the mature duct; c, cambium; wh, phloem; 4, bast fibers, rp,
parenchyma. (Stevens, after Sachs.)

stems and leaves of the pines; when they break down completely,
leaving their secretion in the resulting cavity they are termed
lysigenous, as in the rinds of the orange, lemon and other Citrus
fruits. The secretion reservoirs, also called internal tubular glands,
are formed by a long vertical series of cells forming a space
schizogenously or lysigenously. The schizogenous type of
gland and reservoir is 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 “secretory epithelium.”

150 PHARMACEUTICAL BOTANY

NECTARIES

Nectaries are nectar-secreting tissues, consisting either of
groups of modified cells or of specialized glands, tubes or cavities
found on insect-pollinated plants. ‘The nectar secreting tissue
occurs usually on parts of the flower but may occur on petioles,
stipules, bracts or other structures. Nectar is a sugary secretion
which is exuded through the outer walls of the cells of the

Fic. 88.—Resin duct (secretion reservoir) in leaf of Pinus sylvestris, in cross-
section at A, and in longitudinal section at B; h, cavity surrounded by the secret-
ing cells; f, f, sclerenchyma fibers surrounding and protecting the duct. (Stevens
after Haberlandt.)

nectary. When the nectar secreting tissue is superficial the
modified epidermal cells are often palisade-like or papillose in
form, but may consist of dense groups of club-shaped hairs.
In numerous plants as more elaborate structures such as sunken
pouches, as in the buttercups, or elevated glandular structures
lined with nectar secreting structures, as on the petioles of
Viburnum Opulus, etc.

CLASSIFICATION OF ‘TIssuES ACCORDING TO FUNCTION.—
According to their particular function, tissues may be classified
as follows:

II.

HT.

EN;

VI.

. ConpucTING TIssuES

. STORAGE TISSUES

PLANT TISSUES 151

Medullary rays

Xylem cells (wood parenchyma)
Trachez (vessels) and tracheids
Phloem cells (phloem parenchyma)
Sieve tubes

Companion cells

Epidermis (outer cell walls cutinized)

= (fundamental tissue)
lees (suberized tissue)

PROTECTIVE TissuES

Phloem fibers
Wood fibers
MECHANICAL TIssuES se
Stone cells
Collenchyma
Internal glands
SECRETING TISSUES gana ag aoads ’
External glands (glandular hairs)

ectaries

Medullary-rays

Wood parenchyma

Phloem parenchyma
Laticiferous tissue
Primordial meristem
Dermatogen (protoderm)
Periblem (ground meristem)
Plerome (procambium)
Cork cambium

Cambium

Parenchyma
Reserve parenchyma
MERISTEMATIC TISSUES }

CHAPTER VII

THE ROOT

The Root is that part of the plant that grows into or toward
the soil, that never develops leaves, rather rarely produces buds,
and whose growing apex is covered by a cap.

The chief functions of a root are absorption, storage and
support. Its principal function is the absorption of nutriment
and to this end it generally has branches called Rooters that
are covered with Root-nairs which largely increase the absorb-

Fic. 89.—Cross-section of rootlet in the region of the root-hairs. (From Stevens.)

ing surface. These Roor-Harrs are of minute and simple
structure, being merely elongations of the epidermis of the root,
for a short distance back of the root cap, into slender tubes with
thin walls. Like other living cells, each shows cytoplasm, sap
vacuoles containing cell sap, a nucleus with nuclear membrane,
outer plasma membrane against the covering cell wall of
cellulose and vacuolar membranes around sap vacuoles. That
portion of the epidermis which bears root-hairs is called the
PitiFEROus LAYER.

Som. AND WaTER Retation.—Under conditions favorable
to the growth of land plants, the soil is open and porous, the
particles of soil being separated by spaces containing air, while
every soil particle is enveloped by a film of water. The root-

hairs are in close contact with these soil particles (F ig. 92) and
152

THE ROOT 153

soil water from the films surrounding them soaks through their
cell walls into the root-hairs. This soil water is a solution of
various mineral salts. In order to understand how this solution
passes into the root-hairs, it is essential to first understand the
processes of diffusion and osmosis.

Dirrusion.—This is the process whereby the particles of sub-
stances making up a solution tend to be uniformly intermingled.
It may be observed by introducing a ‘
crystal of copper sulfate into a tumbler :
of water. The crystal slowly dissolves
and the particles of which it is com-
posed in time diffuse themselves
equally throughout the water, so that
the water is colored uniformly blue.
The movement of the particles of the
dissolved copper sulfate from the region
of greater density or concentration to
that of less implies a pressure which is
termed diffusion tension.

Osmosis.—If the denser copper sul-
fate solution in the bottom of the tum-
bler were separated from the less dense
water above by a semipermeable mem-
brane such as parchment or bladder,
diffusion would take place through the oe cede ote
membrane and the water above would .o.. rapid inflow of liquid
become colored as proof of this diffu- from A to B has elevated the
sion. Moreover, the water above the liquid surface within from 64 to
membrane would pass through it in the Re rad ha och apiatcl amtiaad
opposite direction and more rapidly
than the solution of the copper salt. It would continue to do
so until the solution was of the same density on both sides of the
membrane. Diffusion through a porous membrane is known as osmosts.
It is a well-known law of physics that when two liquids or gases of
different densities are separated by a porous (osmotic) membrane,
diffusion through the membrane will take place until the density of
the fluids or gases becomes the same on each side. The diffusion
will be more rapid from the less dense to the more dense region.

154 PHARMACEUTICAL BOTANY

Osmosis AppLIED TO Root-Hatrs.—Normally the cell sap of
the root-hairs is a denser liquid than the soil water outside of

1 2

Fic. 91.—Successive stages of plasmolysis. JV, nucleus; V, vacuole; p, proto-

plasm; e, area formerly occupied by protoplast now replaced by plasmolyzing
solution. (Palladin after de Vries.)

the hairs. The outer plasma membrane and the vacuolar
membranes represent porous osmotic membranes separating the
denser solution within the hair
from the less dense soil water
without. The soil water
imbibed by the cell wall passes
by osmosis through the outer
plasma membrane, _ diffuses
through the cytoplasm to the
vacuolar membranes through
which it passes by osmosis into
the sap vacuoles and_ there
becomes a portion of the cell
sap. '

_ The kind of osmosis just indi-
cated is known as endosmosis or
osmosis from without in. A

Fic. 92.—Root-hairs, with soil-particles
adheae: (Case, after Sachs) reverse process takes place
when small traces of COs, acid

and other substances are excreted, ¢.g., passed out of the root-
hair. This is called exosmosis.

THE ROOT 259

Turcor.—If the cell sap of the root-hair or any other plant
cell becomes concentrated above that of the soil solution, the cell
is caused to bulge. ‘This bulging is a manifestation of osmotic
pressure within and the condition resulting is termed ¢urgor.
Cells of plants exhibiting this phenomenon are said to be in the
state of turgescence or rigidity. Since turgor is coexistent with
growth, conditions affecting turgescence affect at the same time
all growth processes.

PLAsMOLysIs.—If root-hairs or other plant cells are placed in
solutions of a greater density than that of their cell sap content, as
for instance 5 to 10 per cent. salt or sugar solution, exosmosis
becomes more rapid than endosmosis. The cell sap is extracted
more rapidly than the fluid enters from without, with a result that
the protoplasm is loosened from the cell wall and caused to
collapse. This condition is called plasmolysts and the cell is said
to be plasmolyzed. Plasmolysis throughout a tissue or organ
results in wilting. Many plants wilt on account of too high a
concentration of soil solutes.

ABSORPTION OF NUTRIENT SALTs.—The plasma membranes of
all living plant cells are permeable to certain solutes (substances
in solution) and impermeable to others. ‘These membranes in
the root-hairs absorb each solute particle separately and accord-
ing to the need or attraction for that substance; this is called
selective absorption.

The higher green plants vary greatly in respect to their
mineral requirements. Of thirty-one elements found in the ash
of plants, only eleven occur regularly, ¢.g.—sulfur, chlorine,
phosphorus, silicon, potassium, sodium, calcium, magnesium,
iron, aluminum (a_trace) and manganese (a trace), Ten
elements are absolutely required in the form of water-soluble salts
for the normal growth of higher green plants. These ten are
termed essential elements and comprise the following: nitrogen,
sulfur, phosphorus, calcium, potassium, magnesium, tron, manganese,
boron and zinc. Traces of copper have also been found essential
for the growth of some plants.

IMPORTANCE OF EssENTIAL ELEMENTS.— Nitrogen, sulfur and
phosphorus are constituents of proteids and so of protoplasm and
are essential to their formation. Without sufficient nitrogen,

156 PHARMACEUTICAL BOTANY

growth is retarded. Without sufficient sulfur, cell-division and
fruit formation are checked. Deficiency in phosphorus retards
the maturation of the plant.

Fic. 93.—Median longitudinal section through the root tip of the barley. f,
calyptrogen; d, dermatogen, c, the thick outer wall of the same; pr, periblem; fl,
plerome; en, endodermis; /, intercellular-air-spaces; a, cell rows which will develop
the central vessels; 7, detached cells of root cap; s, starch grains in cells of root
cap. X 180. (After Strasburger and Koernicke in Lehrbuch der Botanik.)

Calcium not only neutralizes many harmful substances but
facilitates the absorption of other salts and is necessary for normal
leaf development. It is the chief element in the ash of leaves.

Potassium accompanies carbohydrates during their transloca-
tion and is thought to play a part in their formation.

THE ROOT 157

Magnesium is a constituent of chlorophyll and accompanies
proteins. It is necessary for the formation of fats. Deficiency in
this element results in chlorosis and poor fruit formation.

Iron is essential to the formation of chlorophyll although it is
not a component of that substance. When deprived of iron,
green plants become pale and chlorotic, even though they are
grown in light.

While not absorbed as mineral salts, carbon, hydrogen and oxygen
are likewise indispensable elements. The very life of all plants
depends upon their availability. All three enter into the forma-
tion of proteins, carbohydrates and protoplasm and the first two
are essential to the formation of fats and oils. Water, which
makes up the greatest part of protoplasm and forms the solvent
for soil salts, consists of a combination of two parts of hydrogen to
one part of oxygen.

In determining which elements are essential for the normal
growth and health of plants, it is customary to grow them in
water solutions of various salts or in sterile sand or ground
pumice containing a water solution of the salts.

NON-ESSENTIAL ELEMENTS ABSORBED BY Root-Hairs.—These
elements include those which are also absorbed as water-soluble
salts but which are not required for the normal growth of all
plants. In many specific instances, however, they may be
beneficial.

Silicon is an abundant element in many plants, notably the
grasses, sedges and scouring rushes. It occurs in the ash of
these plants as silicon dioxide (SiO:) and in most soils as silicic
acid (H2SiO;). It is deposited mainly in cell walls of the periph-
eral parts of stems, leaves and seeds and affords protection from
penetration by the hyphz and parasitic fungi as well as from
animal attack such as plant lice and scale insects.

Chlorine, another inessential element, probably has an influ-
ence on the translocation of carbohydrates from leaves to other
organs under natural conditions. Most plants can attain com-
plete development without it. The question of its rdle still
remains unsettled.

Aluminum is only occasionally found in plants. It is known to
influence the color of flowers in Hydrangea hortensis. When

158 PHARMACEUTICAL BOTANY

grown in forest or moor soil, this species has reddish flowers but
when grown in soil containing pent aluminum compounds, its
flowers are blue.

REGIONS OF THE Root. vee are five regions of a root
beginning with the extreme tip and passing backward. These
are the root cap, the embryonic region, the region of elongation or growth
in length, the region of maturation or cell differentiation and the
mature region (see Fig. 94). These phases of growth can be seen
in fixed and stained longitudinal sections of root tips.

Root Cap.—The tip of each root or rootlet is protected by a
smooth, sheath- or scale-like covering known as the root cap or
calyptra. ‘This is a mass of cells which not only protects the
delicate cells of the embryonic region but serves as a mechanical
aid as the root pushes its way through the soil.

Empryonic Recion.—This region is composed of small,
densely arranged, somewhat polyhedral or quadrangular cells
which undergo repeated division. ‘This region is about a milli-
meter in length.

Growinc Point.—The growing point of the root of a seed
plant consists of a group of actively dividing cells directly beneath
the rootcap. These cells make up the tissue called the primordial
meristem.

Primary MErIsTEMS.—Just in back of the primordial meris-
tem, the cells have begun to show differentiation into three
groups of tissues called primary meristems or generative tissues. In
the posterior part of this region the cells gradually cease their
rapid division and become larger due to the absorption of soil
water.

GENERATIVE TissuEs.—The generative tissues directly back of
the growing point are, passing from the center toward the out-
side: plerome or procambium, producing fibro-vascular tissue (xylem
and phloem) and pith; pertblem or ground meristem, producing
cortex including endodermis, and pericycle; and dermatogen or
protoderm, producing epidermis. The name ca/yptrogen has been
given to those cells of the primordial meristem which form the
root cap. (Cf. Fig. 93.)

RecIon oF ELoncAtion.—This region occurs just behind the
embryonic region where the cells enlarge chiefly inlength. Root

THE ROOT ‘ 159

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a Yio. 94,—Lengthwise section of a root tip (diagrammatic), showing its various
“regions. (From “A Textbook of General Botany’? by Smith, Overton et al., Macmillan
Co., Publishers.) Rie!

160 PHARMACEUTICAL BOTANY

hairs begin to make their appearance in this region. Differences
in size and shape of the cells also begin to occur here.

REGION oF MaTurATION.—In this region more differentiation
takes place, the cells become larger and frequently their walls
become thicker. Root hairs are usually evident throughout this
region.

Mature Recion.—This region is the part of the root between
the region of maturation and the base of the stem. In its lowest
part the primary tissues have been laid down by the primary
meristems. These, in order of their arrangement from the
periphery to the center, are the epidermis, primary cortex (composed
of parenchyma), endodermis, pericycle, ycle, radial i vascular bundle composed
of alternating strands of primary xylem and primary phloem, and
pith (composed of parenchyma). Root hairs usually occur in this
lower region as outgrowths of epidermal cells but gradually
disappear in most roots as we pass further upward,

DIFFERENCES BETWEEN Root AND STEM

THe Roor Tue Stem
1. Descending axis of plant. 1. Ascending axis of plant.
2. Growing point sub-apical, being 2. Growing point apical, but covered
covered by the root cap. by protective bud scales or young
foliage leaves.
3. Contains no chlorophyll. 3. Chlorophyll sometimes present.

4. Does not bear leaves or leafrudiments. 4. Bears leaves and modifications.
5. Branches arranged irregularly, and 5. Branches with mathematical regu-
arising from deeper tissue. larity, the branches arising at
surface of stem, in axils of leaves
and from nodes.
6. No division into nodes and internodes. 6. Division into nodes and internodes.
7. Internal structure different. 7. Internal structure different.

CLAassIFICATION OF Roots AS TO ORDER.—1. Primary or first
root, a direct downward growth of the radicle (from embryo
within seed), which, if greatly in excess of the lateral roots, is
called the main or tap root.

2. Secondary roots are. lateral branches of the primary root.
They may, in turn, give rise to finer branches, or rootlets. Both
primary and secondary roots may be either fibrous or fleshy.

3. Adventitious roots are such as arise from any part of the
plant other than the seed embryo or normal root system. Exam-
ples: Roots developing on Bryophyllum and Begonia leaves when

sy

ia aeerttntaiindihiiallinietin alain emeee

THE ROOT 161

placed in moist sand; roots arising from stems of climbing plants
or from stem cuttings.

4. Prop roots or brace roots are such as grow out of the stem a
short distance above the soil
and extend diagonally into the
ground, serving as supports to
the stem. Examples: Indian
Corn and Pandanus.

5. Epiphytic roots are the
roots of epiphytes or air-plants,
many of which are common
to tropical forests. In some
instances, as in Vanilla and
other epiphytic Orchids, these
roots, arising from aerial stems,
hang down free in the air and
absorb water from rain by
means of their several layered
epidermis called a velamen. In
others, as in the English Ivy,
several short roots grow out of
the stem at various intervals
and adhere by their tips to
walls, thus serving as supports
for climbing.

The Banyan (Ficus religiosa)
of India can extend itself over
large areas by means of its aerial ‘
roots. ‘These extend down ver- Fic. 95.—Vanilla planifolia, an epi-
tically from horizontal branches phyte, growing on a tree in Jamaica,
aiid formtee nks, upon anchor- 2 ; (Gager, after Maxon, U. S. Dept.
ing themselves in the soil.

6. ‘The roots of parasitic plants are known as Haustoria. These
penetrate the bark of plants upon which they find lodgment,
known as hosts, and absorb nutritious juices from them. The
Mistletoe, Dodder and Gerardia are typical parasitic higher plants.

CLASSIFICATION OF Roots As TO ForM.—Fusiform, or Spindle-
shaped, like that of the radish or parsnip.

a +

ieee atalino tae ena

162 PHARMACEUTICAL BOTANY

Napiform, or turnip-shaped, somewhat globular and becoming
abruptly slender then terminating in a conical tip, as the roots
of the turnip and some radishes.

Conical, having the largest diameter at the base then tapering,
as in the Maple and Aconite.

Fibrous, when very slender or fiber-like, as in the grasses and
cereals.

Tortuous, when irregularly
bent or twisted, as Pareira.

Nodose, when swollen at
intervals in its course and
fibrous otherwise, as the roots
of the Dropwort.

Moniliform, when the swell-
ings occur at very close intervals
giving the root the appearance
of a close chain of beads.

Tuberous, when fleshy,
swollen, especially in the central
portion, and resembling tubers
in shape, as in Jalap and the
Sweet Potato.

Fascicled, when a number of
thickened and similarly looking
roots occur in a cluster, as in

Fic. 96.—Dodder (Cuscuta sp.) in the Dahlia.
flower. Parasitic on a _ golden rod Roor Consistency or TEx-
(Solidago ulmifolia). (Gager, photo by Elsie’ TURE.—lIn this particular, roots
A, Retireige.) are termed woody, when the
hard, lignified tissue predominates and fleshy, when the soft cellu-
lar tissues predominate. Fleshy roots may be (1) simple or single
or (2) multiple, when a number arise from the base of the stem in
a cluster, as in the Dahlia.

Duration oF Roor.—Plants are classified according to the
duration of the root, as follows:

1. Annual plants are herbs with roots containing no nourish-
ment for future use. They complete their growth, producing

THE ROOT 163

flowers, fruit and seed in a single season, then die. Examples:
Stramonium, Lobelia, etc.

2. Biennial plants develop but one set of aerial organs the
first year, e.g., the leaves, and, as Digitalis, Conium, etc., a large
amount of reserve food material is stored in the roots for the

Nd

ed

2
poms

Fic. 97.—Photomicrograph of a transverse section of the stem of a dicot-
yledonous host-plant infested with the parasite, Dodder. Note the haustoria extend-
ing from the Dodder (D, D’) into the cortex of the host (H). Greatly enlarged.
(Gager.)

support of the plant the following season, when it flowers, fruits
and dies.
3. Perennial plants are those whose roots or underground stems
and roots live indefinitely, as trees, shrubs and perennial herbs.
Root Hisrotocy.—The histology of roots varies, depending
upon character of the surrounding water relations or soil in which

164 PHARMACEUTICAL BOTANY

the plants are accustomed to live which may be aquatic, meso-
phytic, halophytic or xerophytic, etc.

Aquatic plants or hydrophytes are those which live in water or
wet soil as the water-lilies, calamus, cat-tails, etc.

Fic. 98.—Root systems. Fleshy tap root of Carrot (1); napiform root of a
Radish (2); multiple, tuberous roots of a Dahlia (3); nodose roots of the Dropwort
(4); fibrous roots of a Grass (5); fibrous-tuberous root system of an Orchid (6).

Mesophytic plants or mesophytes are those which thrive with a
moderate supply of moisture, as most of the land plants of our
fields and meadows.

Halophytic plants or halophytes are those which thrive in salt
marshes, on saline flats near the sea coast and on the alkali flats of
the interior, as the salt marsh samphire, the mangroves, cheno-
podium, etc.

Xerophytic plants or xerophytes are those which thrive in very
dry soil, as the cactus, century plant, hemlock, spruce, etc.

ee

THE ROOT 165

HistoLocy oF Monocotyt Roor

In this connection we will discuss only the type of greatest
pharmacognic importance, #.e., the mesophytic type as seen in its
most typical form in the transverse section of a Smilax root.

Examining such a section from periphery toward the center,
one notes the following:

1. Epidermis (derived from dermatogen) of a single layer of
cells many of which give rise to root-hairs.

2. Hypodermis (derived from periblem) of one, two or three
layers of cells whose walls are extremely thickened.

3. Cortex (derived from periblem) consisting of a broad zone
of parenchyma cells many of which contain starch grains.

4. Endodermis (derived from periblem) of one layer of endo-
dermal cells whose walls are extremely thickened through the
infiltration of suberin and lignin.

5. Pericycle, also called pericambium (derived from periblem),
of one or more layers of meristematic cells whose walls are
extremely thin.

6. A radial fibro-vascular bundle (derived from plerome) of many
alternating xylem and phloem strands and hence polyarch. The
phloem tissue consists of phloem cells and sieve tubes arranged in
small ovate areas beneath the pericycle and between the outer
reaches of the xylem strands. The xylem is composed of xylem
cells, tracheze and wood fibers.

7. Medulla or pith (derived from plerome) composed of paren-
chyma cells containing starch and often showing xylem patches
cut off and enclosed within it.

The tissues within the endodermis collectively constitute the
STELE.

The Smilax root resembles most monocotyl roots in show-
ing no secondary increase in thickness which is sometimes
called secondary growth. Secondary increase in thickness is the rule
in roots of most dicotyledons and gymnosperms and is due to the
formation in those roots of acambium. No such tissue arises in
most monocotyl roots. However, some difference in diameter
will often be noted between sections cut through the same root at
different levels above the root cap, those cut nearer the stem end

Fic. 99,—Cross section through a representative portion of a monocotyl root
of a species of Smilax. (X 70.) ep, epidermis; hy, hypodermis with lignified radial
and outer walls; co, cortex; par, parenchyma; en, endodermis whose radiatand inner
walls are thicker and more lignified than the outer walls and which show a V-shaped
lumen; fc, pericycle; ph, strand of phloem, the end of the leader being on a sieve
tube; x, xylem; wf, wood fibers in xylem; ¢r, trachea in xylem; m, pith. Note the
numerous xylem and phloem strands collectively constituting a polyarch radial
fibrovascular bundle.

THE ROOT

being broader.

167

The explanation is that the cells in the broader

sections have become larger through the process of growth,
rather than having been augmented in number.

Histotocy oF Dicotyt Roots

The typical dicotyl root is a tetrarch one, which is a root

whose radial bundle exhibits four
xylem patches alternating with four
phloem patches. These roots usually
have an unlimited power of growth.
The early growth is called primary
growth which means first growth and is
represented in sections cut a little above
the root cap in the region of the root
hairs. Above the region of primary
growth is the region of transition where
changes are taking place in the root
which lead to the appearance of
secondary structure.

A transverse section of a dicotyl
root in its YOUNG GRowTH shows the
following structure from periphery
toward center:

1. Epidermis (from dermatogen)
with cutinized outer walls, the cells
often elongating to form root-hairs.

2. Primary cortex (from periblem)
of parenchyma cells with usually small
intercellular spaces. Whenever the
outermost layer or layers of cells of
this region have more thickened walls
than those beneath, the term hypodermis

Fic. 100.—Cross-section of
a young root of Phaseolus multi-
florus. A, pr, cortex; m, pith; x,
stele or central cylinder—all

tissue within the _pericycle,
inclusive; g, primary xylem
bundles; 6, primary phloem
bundles. 8B, cross-section of
older portion of root; lettered
as in A; 6’, secondary phloem;
k, cork. (Stevens, after Vines.)

is employed for this outer portion of the cortex.

3. Endodermis (from periblem) or innermost layer of cells of
the cortex, with lenticularly thickened radial walls.

4. Pericycle (Pericambium) (from periblem) of one to two layers
of actively growing cells which may produce side rootlets.

5. Radial fibro-vascular bundle (from plerome) of four, less fre-
quently of two, three, five or six primary phloem (protophloem)

Late Mee ee et ae ere

1 68 PHARMACEUTICAL BOTANY

patches alternating with as many primary xylem (protoxylem)

arms. It is not uncommon to find bast fibers along the outer

face of each phloem patch. Xylem has spiral trachez, internal
Cork Cambium : = PP ie

1 3 -E-ndodermis
‘

Root Hair

Root Cap—--

Fic. 101.—Diagram of a longitudinal section of a root, showing secondary
thickening and the disappearance of the primary cortex. Endodermis then
becomes surface layer beneath which is formed the cork by the pericycle. p', p”, p*,
phloem formed during first, second and third years; xy!, xy’, xy°, xylem formed dur-
ing first, second and third years. (Mottier.)
to these a few pitted vessels, then, as the root ages, more pitted
vessels, also xylem cells and wood fibers make their appearance.

6. Pith (from plerome), a small zone of parenchyma cells

(not present in all dicotyl roots of primary growth).

ON GS > oe te Se ee Se ees Sees seed

THE ROOT 169

HistoLoGy oF TRANSITIONAL GROWTH AND DEVELOPMENT OF
SECONDARY STRUCTURE.

At about six weeks of growth, one notes cells dividing by
tangential walls in the inner curve of each phloem patch. This is
intrafasctcular cambium. Its cells in each location start to cut off on
their inner side a quantity of secondary xylem and add a little
secondary phloem on their outer side. The protophloem tracts
become pushed out and the protoxylem tracts in. The pith (if
originally present) disappears as the protoxylem encroaches upon
it. Secondary xylem finally fills up the patches between the
arms. ‘The pericambium has a tendency to start division into an
outer and an inner layer. The outer layer becomes a cork
cambium (phellogen) surrounding the bundle inside of the endo-
dermis. It cuts off cork tissue on its outer face, hence all liquid
material is prevented from filtering through and cortex including
endodermis, as well as the epidermis, shrivel, dry up and separate
off at the age of two to three months. The cork cambium
(phellogen) lays 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 intrafasctc-
ular cambium to form a continuous cambium ring which then cuts
off additional secondary xylem on its inner face and additional
secondary phloem on its outer face pushing inward the first-
formed or protoxylem and outward the first-formed or proto-
phloem. Medullary-rays are formed by the cambium as it cuts
off secondary xylem and secondary phloem elements.

Thus, in a transverse section made through a portion of a
Dicotyl root showing secondary growth, the following regions are
noted passing from periphery to center:

1. Cork
2. Cork cambium (phellogen)
3. Secondary cortex

= 4, Protophloem

% «\5. Secondary phloem

z 2 6. Cambium

5 = (7. Secondary xylem

& 3\8. Protoxylem

170 PHARMACEUTICAL BOTANY

9. 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.

The xylem or wood portion of each fibrovascular bundle
appears as a wedge placed between the xylem portions of two
medullary-rays and so is sometimes called a Woop Wepcr. The
xylem portions of the medullary-rays between the wood wedges
are termed Woop-rays.

ees oe : : ‘
; . oy : ee : eae Z

Fic. 102.—Photomicrograph of a transverse section of a California Privet root
of primary growth showing epidermis (ep); hypodermis (h); primary cortex (fc);
endodermis (en); pericambium (ca); a xylem arm of the radial bundle (x); a phloem
arm (ph); and pith (m). x 40.

HistoLoGy AND DEVELOPMENT oF A TypicAL Dicotyit Root
(CALIFORNIA PriveT).—A. Make a permanent mount of a cross
section of the root of the California Privet (Ligustrum Californicum)
cut just above the root cap, and note the following structures,
passing from periphery toward the center (see Fig. 102):

1. Epidermis, composed of a layer of epidermal cells whose
outer walls have been infiltrated with a substance called cutin.

2. Hypodermis, a layer of somewhat thick-walled cells just
beneath the epidermis.

3. Cortex, composed of cortical parenchyma cells with small,
angular, intercellular-air-spaces.

SPY RTARTA SAA tae AA RINE: NS ae a

THE ROOT 171

4, Endodermis, or innermost layer of cells of the cortex, whose
radial walls are lenticularly thickened.

5. Pericambium (Pericycle), of a layer of actively growing meri-
stematic cells, which has the power of producing lateral rootlets.

ep

Fic. 103.—Photomicrograph of a transverse section of a California Privet root
made about 114 inches above the root tip and showing transition structure. The
epidermis (ef), primary cortex (pc) and endodermis are in the process of sluffing
off, since cork (ck) has been laid down by the cork cambium (pn) directly beneath
the endodermis. The cork cambium has also formed several layers of secondary
cortex (sc) on its inner face. The protophloem ( pp) represented largely by hard bast
has been pushed out, while a small amount of secondary phloem (sp) represented
by soft bast has been deposited beneath it by the cambium (c) which now is
nearly circular in aspect. The protoxylem (px) has been pushed into the center
by the encroaching secondary xylem (sx) which has been laid down by the cambium
on its inner face. mr, medullary ray. Highly magnified.

6. Radial fibro-vascular bundle of five or six primary xylem arms
alternating with as many primary phloem patches. Note the
narrow spiral trachez in the xylem patches.

7. Pith, a small central zone of parenchyma.

172 PHARMACEUTICAL BOTANY

The section you have just studied illustrated in general the
appearance of any Dicotyl root of primary growth.

B. Mount permanently another cross section 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

pe

development. More secondary xylem has been laid down on the inner face of
the cambium and there has been a slight increase in the secondary phloem. ¢f,
epidermis; h, hypodermis; fc, primary cortex; ck, cork; pn, phellogen or cork cam-
bium; sc, secondary cortex; pp, protophloem; sf, secondary phloem; ¢c, cambium; sx,
secondary xylem; px, protoxylem.

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 is composed of a mass of
protophieom (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 (interfascicular cambium) which becomes joined to the
intrafascicular cambium to develop secondary phloem on the outer
surface and secondary xylem on the inner face.

ee ee ee

THE ROOT 173

C. Mount permanently a third T. S. cut through the same
root a short distance above the second. Note that this is still
larger in diameter than the second. The pertcambium has already
divided into an inner and an outer layer. The outer layer has
become the cork cambium, cutting off the cork on its outer face

2 x"

x

Fic. 105.—Transverse section of California Privet root made about an inch and
a half above the section shown in Fig. 104 and showing early secondary structure.
Note that epidermis, primary cortex and endodermis have completely disap-
peared. Cork (ck); phellogen (pA); secondary cortex (sc); protophloem (p’);
secondary phloem (#?); cambium (c); secondary xylem (x?) and protoxylem (x):
(Photomicrograph, X 40.)

beneath the endodermis. Cork, being an impermeable barrier
to water, has prevented the nourishing sap from percolating
through to the endodermis, cortex and epidermis. ‘These
regions have consequently begun to sluff off. Note that the
cambium has begun to spread out into the form ofaring. More
secondary xylem has been formed on its inner face and additional
secondary phloem has appeared on its outer face (Fig. 104).

174 PHARMACEUTICAL BOTANY

D. MA@ke a permanent mount of a fourth cross section cut
through the same root some distance above the third. Note that
the epiderr#s, brimary 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

on gemma neem RS — —

ccc

Fic. 106.—Photomicrograph of a transverse section of an older portion of Cali-
fornia Privet *0ot, showing further secondary development. XX 35. Note the
prominent medullary rays (mr); cork (ck); phellogen (ph); secondary cortex
(between ph and ’); protophloem (p’); secondary phloem (2); cambium (c);
secondary xylem (x?), the leader pointing to a trachea; wood fibers between
trachez; and Pprotoxylem (x’).

cortex on its inner face. The cambium has assumed a circular
aspect. Just beneath the secondary cortex will be found flat-
tened patches of protophloem, and beneath these secondary phloem
masses have 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. The pith has disappeared and its place is
taken by protoxylem (Figs. 105 and 106).

THE ROOT 175

ABNORMAL STRUCTURE OF DicotyL Roots.—In certain
Dicotyl roots as the Beet, Jalap, Pareira, and Poke, after the
normal bundle system has been formed, there then develop suc-
cessive circles of cambiums in the cortex outside of these bundles.

Fic. 107.—Cross section of the daughter tuberous root of Aconitum Napellus,
Spark’s variety, cut slightly below mid-region. Photomicrograph X 25. ¢p,
epidermis; fc, primary cortex; en, endodermis; si, sieve strands in inner bark; c,
cambium; wp, wood parenchyma; xy; tracheary tissue of xylem; m, pith.

Each of these cuts off phloem on its outer face and xylem on its
inner face, thus producing concentric series of open collateral

bundles.
Hisrotocy or A Dicotyt Tuserous Roor (AconITUM).—

Transverse sections made through the tuberous roots of Aconitum
Napellus vary in appearance depending upon the level at which

176 PHARMACEUTICAL BOTANY

they are cut and upon whether cut from the daughter or parent
tuberous root. In general, the commercial root shows the fol-
lowing internal structure: An outer layer consisting either of an
epidermis or, in older parts, of one or more rows of subertzed cells
with blackish-brown walls, beneath which in younger portions of
the root occur a primary cortex of starch containing parenchyma in
which occur scattered stone cells of numerous shapes and a modified
endodermis, all or a portion of these regions being suberized in older
parts, forming a brown tissue called metaderm or entirely exfoliated;
an inner bark of numerous rows of starch parenchyma and scattered
islets of sieve tissue. Stone cells may occur in this zone directly
under the endodermis. Next, a five- to eight-angled cambium,
in each angle of which is a 2- to 4-rayed collateral vascular
bundle. In some sections of Aconite roots smaller bundles may
be found between the angles and along the cambium line. In
the center is found a broad, five- to eight-rayed pzth composed of
large parenchyma cells. The parenchyma cells of the cortical
regions and pith contain single or two- to five-compound starch
grains.

Root Brancutnc.—The branches of roots are formed by the
pericycle. The cells of this region adjacent to the protoxylem
undergo division by laying down tangential walls forming a mass
of meristem. This meristem develops a root tip which pushes its
way out laterally through the tissues of the primary root (Fig. 108).
In its passage outward, enzymes are secreted by its cells which
dissolve the tissues in its path, the root growing into the cavity
thereby formed. The first branches ofa root are termed secondary
roots. ‘These may give rise to branch roots and those to further
branches by the same method.

Root TuBERCLES

The roots of plants of the Bean and Bayberry families as well
as some species of the Birthwort family and of the Alders, Cycads,
Spruces, etc. are characterized by the appearance upon them of
nodule-like swellings called root tubercles.

Lecuminous TuBERCLES.—In the case of the members of the
Bean family (Leguminose), the causative factor is a species of
bacteria named Pseudomonas radicicola. This is a motile rod-

177

Pericycle

THE ROOT

Endodermis

es
EQ
ss
2 Qs
s =
5 yo
x 3 ee
oC g s& 3
ws O CS -]
SS x) sae mS
A - t ! /
\ H / /
se) | /
Acct

(From Curtis.)

f ee Sr oeig, i ‘ ‘ _
ae an
i . } 4 a

e

a

aS

Se

aa Ss

ENG

Oe:'

Cross section of root of lupine, showing a secondary root breaking
through the cortex.

108.

Fig;

(Marshall.)

Fic, 109.—Root system of a legume showing tubercles.

178 PHARMACEUTICAL BOTANY

shaped organism which appears widely distributed in soils. It is
apparently attracted to the root-hairs of leguminous plants by a
chemotaxic influence probably due to the secretions poured out
by these structures. A number of these organisms penetrate the
walls of the root-hairs by enzymic action. Upon entering the
hairs they form bacterial strands which branch and rebranch and

extend into the middle cortex

% ® ( cells. Within the cortex cells
yK >. the organisms multiply rapidly
a a ® SI . producing nest-like aggrega-

4 4 1 tions. Their presence here
stimulates the rapid multipli-

with the consequent forma-

en ve a # cation of the parenchyma cells
y

me ° tion of nodules or tubercles.
Ww o a ;

te R 5 Under oil-immersion magni-

S 3 ° eo fication these bacteria are often

Fic. 110.—Pseudomonas radicicola. 1, fouaid 0 aes : VariGnsly
From Melilotus alba; 2 and 3, from Shaped forms called involution
Medicago sativa; 4 and 5, from Vicia villosa. forms. They remain within
(Marshall, after Harrison and Barlow from the cells of the middle cortex
Lipman.) 3 S

region breaking down the
starch and sugar present into simpler compounds. In doing
this the Pseudomonas obtains the energy required to ‘‘fix’’ the
free atmospheric nitrogen. This process is called “fixation of
atmospheric nitrogen” and consists of uniting the free nitrogen
of the air with some other elements to form a nitrogenous com- .
pound. This is assimilated into its own body. ;

The bacteria gradually swell up into zodglwa masses, until
finally their bodies break down into soluble nitrogenous sub-
stances which are partly absorbed and assimilated and partly
stored as reserve nitrogenous food by the green leguminous
plant.

Rotation or Crops.—Upon the decay of the leguminous
roots and their bacteria in the soil, the nitrogenous compounds
are set free and are ready to be decomposed by various soil

organisms into the various decomposition products, including
amino-acids and ammonia,

THE ROOT 179

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, such as cereals,
cabbage, potatoes, etc.

Many strains of Pseudomonas radicola exist, one for every kind
of leguminous crop. Cultures of many of these, such as “‘Nitro-
Germ,” etc. are now obtainable from the drug trade.

Saeee

Fic. 111.—Tubercular clusters on underground stem and roots of a Bayberry;
Myrica Macfarlanci, observed by the author at North Wildwood, N. J., Jan. 31,
1915.

Myrica TuBeRCLES.—The writer has found tubercles on
Myrica cerifera, Myrica Caroliniensis 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 certfera, 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.

180 PHARMACEUTICAL BOTANY

The tubercles occur either singly, as is frequently the case on
subterranean branches, in small groups the size of a pea, or in

&

oe
cea ES

(MITROGEN OF Al

— \——___y-

C

oe * : : s NY : fg -
eis Ate €
i : @ 4. :
| Ys uteier
Ee OO ATES |
s Bee ioyy: eoNI R

Fic. 112,—Nitrogen cycle in the life of alfalfa, a leguminous plant. (From Brownell?s
General Science.)

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 trichot-

THE ROOT 181

Kate

tAST
3

&
xn

Fic. 114.—Ectotrophic mycorrhiza of the beech (Fagus); a, humus particles; 4,
strands of fungus hyphz penetrating the soil. (Palladin.)

182 PHARMACEUTICAL BOTANY

omously after attaining a certain length. The branches fre-
quently 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.

Tue Nirrocen Cycie

Living plants make use of nitrogen in the form of nitrates
found in the soil and water. The nitrates are absorbed by the
plant and split up within its cells into the
component elements, the nitrogen of which
is linked with some other simple elements
there present including carbon, hydrogen,
oxygen, sulfur and sometimes phosphorus
to form proteins. Animals eat the plants
and part of the proteins are digested by
them and excreted as urea, uric acid and

hippuric acid. These products are later
Fic. 115. Endo- decomposed by bacteria (micrococcus ure@,
trophic mycorrhiza in etc.) to form ammonia. The remainder of
epidermal cells of the : . ;
root of a member of the the proteins are assimilated and stored in
heath family, Andromeda the growing animal cells. When the ani-
polfolia, the root shown mals die their bodies decay and the pro-
in cross section. (Palla- |. : ‘
din.) tein or nitrogenous compounds are split
into decomposition products (peptones,
amino-acids, ammonia) of which ammonia is the final end
product.

The soil contains Nirriryinc Bacteria. Of these the Nitro-
somonas and WNitrosococcus secrete enzymes which oxidize ammonia
to nitrous acid. Some of the nitrous acid is neutralized by
alkaline substances found in the soil to form nitrites. Another
nitrifying organism, the Nitrobacter secretes enzymes which oxi-
dize nitrous acid and nitrites forming nitric acid; free nitric acid
then combines with other substances (bases) in the soil forming
nitrates, thus completing the cycle.

THE ROOT 183

MycorRHIZA

The roots of many plants, especially those growing in humus,
are associated with fungal hyphz which form a weft over their
surface, as in the case of the beech, etc., when this weft is called
an ectotrophic mycorrhiza, or the surrounding hyphz may penetrate
into the cells of the roots, as in many plants of the Heath and
Orchid families, when the association is termed an endotrophic
mycorrhiza.

There are a number of plants, particularly non-green seed
plants, which cannot thrive without the mycorrhiza with which
their roots are normally associated and die when transplanted to
soil not containing them. Mycorrhizz assist the roots to absorb
nutrient materials from the soil.

A number of mushrooms form ectotrophic mycorrhizas on
the roots of trees.

CuapPtTer VIII

THE BUD

Bups are short, young, undeveloped stems or shoots with
rudimentary leaves 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 trans-
formed into scales; these 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. The
official Poplar Bud is an example.

Naked buds are those which are devoid of protective scales.
They are common to herbaceous plants. The rose buds, head of
cabbage or brussels sprouts are good examples.

CLASSIFICATION OF Bups AccoRDING TO DEVELOPMENT.—1. A
leaf bud is a young shortened shoot bearing a number of small
rudiments of leaves. It is capable of elongating into a branch
which bears leaves. The leaf buds of Populus balsamifera and
P. candicans are official in the National Formulary.

2. A flower bud is a rudimentary shoot bearing one or more
concealed and unexpanded young flowers. The drug Clove is
an example of a flower bud which, if allowed to expand, would
form a single flower.

3. A mixed bud is an undeveloped stem or branch bearing
concealed, unexpanded leaves and flowers. The buds which
give rise to flowers and leaves on the apple trees are mixed buds.

CLASSIFICATION OF Bups 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-
184

THE BUD 185

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 plant other than at the
apex of the stem or in the axil of
aleaf. Such buds may be seen
developing along the veins of a
Begonia leaf or in the notches
along the margin of a Bryo-
phyllum leaf after these have
been planted in moist soil for
several days. The roots of the
poplars, willows, cherries and
many other trees develop
adventitious buds underground
which develop suckers which
may eventually become mature
plants that lead an independent
existence.

4, An accessory bud is an extra
bud which forms in or near the
leaf axil. These may be seen
on the young stems of the Red
Maple.

5. A sub-petiolar bud is one
arising beneath the basal part of
the petiole or leaf stalk, so that
when the leaf falls off the bud
becomes exposed, as in the
Buttonwoods.

CLASSIFICATION OF BupDs
AccoRDING TO THEIR
ARRANGEMENT ON THE STEM.

1. When a single bud is

» ferminal leaf-bud

AS
a

} flower- buds.
x \stipule-sear

J ..- J lower-bud-scars

Fic. 116.—Cottonwood twig, two years
old. (From Robbins after Longyear.)

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.

CHAPTER IX

THE STEM

_ The Stem is that part of the plant axis which bears leaves or
modifications of leaves and its branches are usually arranged with
mathematical regularity.

Stems usually grow toward the light and so are positively
heliotropic.

The principal functions of a stem are to bear and support
leaves, branches and reproductive organs, connect roots with
leaves, conduct water with minerals in solution from roots to
leaves, to transport elaborated plant foods from the leaves to the
roots and to store reserve foods.

When the stem rises above ground and is apparent, the plant
is said to be caulescent, as the various trees, shrubs, geraniums, etc.

When no stem is visible, but only flower or leaf stalks, the
plant is said to be acaulescent, as the Bloodroot and Violet.

Stems vary in size from scarcely !¢5 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 and has a
trunk diameter at its base of between 25 and 30 feet. Some of
the Eucalyptus trees of Australia and Tasmania are reported to
attain the height of 500 feet.

Nopes AND INTERNODES.—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 GrowrH.—Generally the growth of the
stem is upright or erect. Very frequently it may be:

Ascending, or rising obliquely upward. Example: Saw
Palmetto.

Reclining, ox at first erect but afterward bending over and
trailing upon the gound. Examples: Raspberry and Blackberry.

Procumbent, lying wholly upon the ground. Example:
Pipsissewa.

186

THE STEM 187

Decumbent, when the stem trails and the apex curves upward.
Examples: Vines of the Cucurbitacee or
Gourd family.

Repent, creeping upon the ground
and rooting at the nodes, as the
Strawberry. NOUS Terminal Bud

STEM ELONGATION.—At the tip of }
the stem there is found a group of
very actively dividing cells (primordial
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 or protoderm forming
epidermis;

2. Periblem or ground meristem form-
ing the cortex, pericycle, and pith.

3. Plerome or procambium forming
the fibro-vascular elements.

DurRaTION OF STEMS.

Annual, the stem of an herb whose
life terminates with the season.
Example: Corn.

Perennial, when the stem lives for
many years. Example: Oak.

Forms oF Stems.—The following
adjectives appear in pharmacognostic
and botanical literature in relation
to the form of stems and in some |
instances other plant organs: Cylin- }f
draceous or cylinder-shaped; subcylin-
drical or somewhat cylinder-shaped; rye. 117.—End portion of a
terete or circular in cross section; com- branch of the horse-chestnut
pressed or pressed together above and “Speeg amare te : ee) the
below or laterally; triquetrous or three- SS
angled, quadrangular, or four-angled; alate or winged; exfoliating or

=.

patee--- Auxillary Bud
Ce a Leaf Scar

a Leaf Scar

ae----- Scars of Terminal
j Bud Scales

wy ~~ ~~

188 PHARMACEUTICAL BOTANY

shedding cork in layers; obconical, or having the shape of an
inverted cone; flexuous or bent; tortuous or twisted; truncate, or end-
ing abruptly, as if cut across at the summit.

Stem MobpirIcaTIONs.

Twining stems are those which coil spirally around a support,
as the stems of the Hop, Bittersweet, Dodder, etc.

Tendriliform stems are those which undergo thread-like modi-
fication and become sensitive to contact of a side branch or other

Cette Prins object, coiling around it, as in

es Passion flower, Grape, Squash,
Cle.

Spines are formed by the

oF Leaves
a

checking and hardening of a
(f ))) : branch that may then become
wo, }f| + sity Buds defensive, as in hawthorn,
ie honey locust, etc.

Aerial tuberous stems are those
in which one or more inter-
eee & nodes enlarge above ground
— and store reserve food, as in

Fic. 118.—Diagram of a longitudinal pseudobulbs of orchids.
section through the tip of astem. 1, 2, Subterranean tuberous stems

3, 4, successively older beginnings (pri-

are those in which an under-
mordia) of leaves. (Mottier.)

ground 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).

Phylloid stems or phylloclades are green, flat, leaf-like stems in
which flattening branch expansion has occurred, as in Asparagus,
Ruscus, etc.

Cactoid stems are those in which reduced, condensed branches
or stems become swollen for water (and food) storage, as in Cacti,
some Euphorbia species, etc.

ABOVE-GROUND OR AERIAL 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 STEM 189

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, leaflet or branch, capable of coiling spirally and used by a
plant in climbing. The tendrils
of the Grape Vine are modified
inflorescence branches, those of
Sarsaparilla are modified stipules
and those of the Pea are modified
leaflets. The ends of the tendrils
of the Japan Ivy become swollen
and flattened, forming adhesive
discs which cling to objects with
which they come into contact.

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
on the stems of the roses, green-
briers, etc.

A stolon or runner is a prostrate
or reclining branch, the end of
which, on coming in contact with
the soil, takes root, so giving rise
to a new plant. Examples:
Currant and Raspberry. Short
stolons like those of the Houseleek Fs is ted ex Hevea (Bice
are called offsets. dioica). (Palladin.)

A bulbel or bulbil is a small,
bulb-shaped, young aerial shoot which serves as an organ of veg-
etative multiplication by falling off the stem and developing into
a new plant. The Garlic and Tiger Lily are good examples of
plants which produce bulbels.

An herbaceous stem is one which is soft in texture and readily
broken. Example: Lily-of-the-Valley.

An undershrub or suffruticose stem is a stem of small size and
woody only at the base. Examples: Bittersweet, Thyme, etc.

*

190 PHARMACEUTICAL BOTANY

A shrubby or fruticose stem is a woody stem larger than the pre-
ceding and freely branching near the ground. Example: Lilac,
rc;

A trunk is the woody main stem of a tree.

A twig is that portion of a woody branch which has developed
during the last growing season.

HERB, SHRUB AND TREE

An Hers is a plant whose stem does not become woody and
permanent, but usually dies, at least down to the ground, after

|

Fic. 120.—An elm tree in summer and winter. To illustrate alike the deliques-
cent habit of branching and the deciduous nature of this tree (dropping its leaves
in autumn and developing new ones in the spring). (Atwood.)

flowering. Herbs are either annual, biennial or perennial (see
pp. 162-163). When a perennial herb possesses an aerial stem
which develops leaves that persist on the stem over winter, as
the Partridge Berry, it is called an evergreen herb.

A Tree is a perennial woody plant of considerable size,
attaining a height of 15 or more feet, and having as the above-
ground parts a trunk and a crown of leafy branches.

THE STEM 191

There are two plans of branching in trees. When the trunk,
or main stem, extends vertically upward to the tip, as it does in
the pines, junipers, spruces and other conical trees, the type of
branching is called excurrent; when it divides into several more or
less equal divisions as in the elm and other spreading trees, it is
said to be deliquescent. ‘The deliquescent plan is the more com-
mon one among our deciduous trees.

Fic. 121.—A, tunicate bulb of Onion (Allium Cepa), in longitudinal section;
B, Scaly bulb of Canada Lily (Lilium canadense); C, Same as B, but in longitudinal
section; D, Bulb of Garlic (Allium sativum) with portion of the scales cut away so
as to enclose the young bulbs within; E, Portion of stem of Tiger Lily (Lilium
tigrinum), showing bulbils in the axils of the leaves; G, Corm of Crocus showing a
terminal and one lateral bud. The horizontal lines represent the scars of the few
scales that have been removed. At the base is the remnant of last year’s corm.
The various buds will develop into new corms. F, Vertical section of a familiar
corm. H, flat corm of Cyclamen; I, Bulblets at the apex of a flower stalk of the
Onion. (After Gager.)

A Surus is a perennial woody plant of smaller stature than a
tree whose stem is relatively short and usually branches near the
ground. Examples: Viburnum, Elder, Buchu, Krameria, etc.

Fatt or Leaves.—The fall of leaves in autumn from our
deciduous woody dicotyledons and gymnosperms is caused by the
formation beneath the leaf stalks of an abscission or cut-off layer
or band of delicate cells. ‘The cellulose walls of these break down
into pectic acid and pectin which forms a mucilaginous substance,
the middle lamellae between the cells disintegrates, the veins
become ruptured and the leaf falls of its own weight. A protec-

192 PHARMACEUTICAL BOTANY

tive layer of cork is developed beneath the abscission layer either
before or after the fall of the leaf which seals the wound and forms
the leaf scar. The shape of many of the leaf scars are diagnostic
for the stems bearing them.

UNDERGROUND OR SUBTERRANEAN STEMS.—A RHIZOME is a
creeping underground stem, more or less scaly, sending off roots
from its lower surface and stems or leaves from its upper. The
rhizome grows horizontally, vertically or obliquely, bearing a

Nit WI
Fic. 122.—Rhizome of Solomon’s Seal (Polygonatum biflorum). 1, scar of last
year’s overground stem; 2, base of this years aerial stem; 3, terminal bud of next
year’s aerial stem. sc, scale leaves; s, scars (annulations) of old scales; r, rootlets.
5¢ natural size. (Gager, after Gapp.)
terminal bud at its tip. Its upper surface is marked with the
scars of the bases of aerial stems or leaves of previous years.
Examples: Triticum, Rhubarb, etc.

The Tuser is a short and excessively thickened underground
stem, borne usually at the end of a slender, creeping branch, and
having numerous “eyes” or axillary buds. Examples: Tubers of
the Potato and Jerusalem Artichoke.

The Cor 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. Exam-
ples: Colchicum, Jack-in-the-Pulpit, Crocus, etc. (See Fig. 127.)

A BuLB is a very short underground stem invested by fleshy
scales which represent storage leaves. It produces rootlets from
its lower face and leaves and flower from its 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
enclosing the inner. Example: Lily. (See Fig. 127.)

THE STEM 193

A Sucker is a slender underground branch that arises either
on aroot orastem. Root suckers are produced by roses, poplars,
elms, etc. and arise as adventitious buds on the roots of these
plants. Stem suckers are produced by peppermint and other
mints, etc., arising from the axils of scaly leaves on the rhizome,

— decurrent
leaf base

_ gerial stem

Fic. 123.—Portion of a sprouting potato tuber. (After Robbins.)

or they may arise from wound tissue which has developed on
sawed-oft branches or from old wood near the base of old trees.

ExoGENoUsS AND ENDOGENOUS STEMS.—Exogenous stems are
typical of Gymnosperms and Dicotyledons and can increase
materially in thickness due to the presence of acambium. Such
stems show differentiation into an outer bark cylinder and an
inner or wood cylinder. Between the bark and the wood cylin-
ders is a cambium.

194 PHARMACEUTICAL BOTANY

Endogenous stems are typical of most Monocotyledons and can-
not increase materially in thickness due to the absence of cam-
bium. The limited increase in diameter that does take place is
due to the enlargement of the cells of the primary tissues. Such
stems show no differentiation into bark and wood regions.

ee ie t
0
spt ie halk

/

Fic. 124.—Sunflower stem. Representative portion of a transverse section
cut through upper, young part of stem. ep, epidermis bearing trichomes (2);
hy, hypodermis; co, primary cortex; Pf, pericyclic fibers; p, protophloem of primary
fibrovascular bundle; c, intrafascicular cambium of bundle; x, protoxylem of bundle;
m, pith or medulla. All of the tissue between each two bundles is a primary
medullary ray. Greatly enlarged.

Hisrotocy or AnnuaL Dicory. STEM.—Excellent stem
material of an annual dicotyledon for the study of the stems of
this type is to be found in the sunflower. Transverse sections
cut through the upper, thin, younger portion of this stem,
present the following structures: (In both annual and perennial
dicotyledonous stems, endodermis and pericycle are rarely dis-

THE STEM 195

tinguishable from the cortex, except tor the knowledge of their
relative position, since each has become so similar to the cortex
through passage of food, etc.)

1. EpIweRMis, a single layer of somewhat flattened cells whose
outer walls are cutinized. Some of the cells show outgrowths as
hairs. Stomata also occur.

mr mr

Fic. 125.—Diagram of a representative portion of a transverse section cut
through an old part of the Sunflower stem. ep, epidermis; col, collenchyma;
co, cortex; p.f., pericyclic fibers in pericycle; ph, protophloem; ph’, secondary
phloem; cam, intrafascicular cambium; cam’, interfascicular cambium; x, proto-
xylem; x’, secondary xylem; mr, primary medullary ray; mr’, secondary medullary
ray; m, pith. The three longest bundles represent primary bundles which have
been broadened by the cambium adding secondary phloem outwardly and second-
ary xylem inwardly. The smaller bundles represent secondary bundles, each
consisting of secondary phloem, cambium and secondary xylem.

2. Primary CorTEX, composed of two zones: an outer region
of several layers of narrow, thick-walled cells (hypodermis) and a
broad zone of larger, thin-walled cortical parenchyma cells, many

of which contain chloroplasts.

3. The innermost layer of cells of the cortex is called the
Enpopermis. (Not generally distinguishable.)

4. Pertcycie (Pericambium). Not generally distinguish-
able in most dicotyl stems but showing early formation of groups
of fibers in sunflower stems.

196 PHARMACEUTICAL BOTANY

5. PRIMARY FIBRO-VASCULAR BuNDLEs of open collateral type
arranged in a circle with primary medullary rays between the
bundles. Each bundle shows an outer protophloem, a cambium
called intrafascicular cambium, because it is within the bundle, and
an inner protoxylem. .

6. Piru, composed of parenchyma cells.

If a series of additional cross sections are now cut at different
descending levels along the Sunflower stem until the old part of

Mechanical Tissue, Phloem
Parench | mbium

Xylem .

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Fic. 126.—Portion of a longitudinal section of a sunflower (Helianthus) stem.
(From “A Text Book of General Botany’? by Smith, Overton et al. The Macmillan Co.

Publishers.)

the stem near the ground is reached, and these be examined
microscopically, it will be noted that some secondary changes
have taken place which explain the gradual increase in thickness
of this stem as the base is approached. One of the first of these
changes incident to secondary growth is that observed in the
cells of the primary medullary rays between the intrafascicular
camlia of the various bundles. These lay down tangential walls
to form interfascicular cambia or cambiums between the bundles.

When this process is completed, a continuous cambium ring is
formed.

THE STEM , 197

The cambium ring adds more xylem elements called secondary
xylem on its inner face and more phloem elements called secondary

hyma of Cortes

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Fic. 127.—A diagram to show the character of the tissues and their disposition
in a young stem of the typical dicotyledon type. (From Stevens.)

_ phloem on its outer face. As it cuts off sieve tubes and companion

cells outwardly and tracheze and wood fibers inwardly all along

the line it also cuts off here and there a medullary ray cell on

198 PHARMACEUTICAL BOTANY

each of its faces. It will be noted that more secondary xylem is
added than secondary phloem. The result of the activity of the
cambium ring is the deepening of the primary bundles and the

x
5 *lloge, pce
Brey Cambium
Ylem fro
S iM the Camby
a
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gente 2s
Fic. 128.—Diagram similar to the preceding but representing a later stage and
showing the tissues formed by the cambium. (From Stevens.)

primary medullary rays, the appearance of new or secondary
bundles (which are smaller than the primary bundles) in the
primary medullary rays and the appearance of narrow secondary
medullary rays between the secondary bundles as well as inside

THE STEM 199

of the deepened primary bundles. The secondary medullary
rays within the bundles are also called “‘vascular rays.”’

As secondary increase progresses, it will be noted that certain
cells in widely separated regions of the pericycle begin to divide
rapidly producing groups of thick-walled pericyclic fibers here and
there within the pericycle but chiefly outside of the protophloem
of the larger or primary bundles.

Fic. 129.—Photomicrograph of cross-section of stem of Aristolochia sipho, where
cambial activity is just beginning. a, Epidermis; 4, 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 parenchyma of
the pericycle; f, primary medullary-ray; g, ‘phloem; ’, xylem; 7, interfascicular
cambium; j, medulla or pith. > 20. (From Stevens.)

It will be noted that cross sections through the young terminal
portion of the sunflower stem show an interrupted cylinder of
separate bundles whereas similar sections cut through the older
lower portion of the stem show a continuous cylinder of bundles,
the continuity being due to the laying down by the interfascicular
cambia of secondary bundles between the primary bundles during
secondary growth. The stele or central cylinder of the younger
part of this stem is known as a dictyostele or dissected siphonostele
whereas that of the older portion is a s¢phonostele (see Fig. 83).

GROWTH OF PERENNIAL DicoTyL STEM AND Its HisroLocy.—
A perennial dicotyl stem, in the first year of its growth does not

200 PHARMACEUTICAL BOTANY

differ in structure from an annual. By the close of the year a
cork cambium (phellogen) has originated beside the epidermis.
In the origin of the cork cambium—one of two methods: (a) either
the epidermis may divide into an outer layer of cells that remains

irae SGT LS:
Po

Fic. 130.—Portion of cross-section of four-year-old stem of Aristolochia sipho, as
shown by the rings of growth in the wood. The letters are the same as in Fig. 129
but new tissues have been added by the activity of the cambium; and a cork cam-
bium has arisen from the outermost collenchyma ceils and given rise to cork.
The new tissues are: /, cork cambium; f, cork; g, secondary phloem from the
cambium, and just outside this is older crushed phloem; n, secondary xylem pro-
duced 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. 129 and 130, 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.)

epidermis and an inner layer of cells that becomes cork cambium,
or, (b) the outermost layer of cortex cells underneath the epi-
dermis becomes active after being passive for one year, and lays
down walls, the inner layer becoming cork cambium, the outer
becoming a layer of cork. The cork cuts off water and food
supplies from the epidermis outside, and so the epidermis sepa-

THE STEM 201

rates 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

\periderim

epidermis

Fic. 131.—Diagrams, based on the red oak, Quercus rubra, showing the position
and extent of successively formed periderm or corky layers in a typically woody
stem. A, a one-year-old twig, the first periderm layer, a complete cylinder, formed
beneath the epidermis. B, a two-year-old twig, the epidermis and first periderm
ruptured; new, shell-shaped layers formed deeper in the cortex. C, a three-year-
old stem, the outer tissues weathered away and more periderm layers formed still
more deeply in the stem, invading the secondary phloem. D, a four-year-old stem,
the cortex and outer secondary phloem with their periderm layers weathered away,
the new cork layers invading the younger phloem. £, the outer tissues of an old
tree-trunk, showing the narrow band of young, living secondary phloem, and the
thick, deeply fissured layer of older, dead phloem with its many shell-shaped peri-
derm layers; a considerable amount of similar tissue has exfoliated. (Krom Eames
and MacDaniels “Introduction to Plant Anatomy,” McGraw-Hill Book Co., Inc., Publishers.)

joins the cambium of the first-formed bundles (intrafascicular
cambium) to form a complete cambium ring. By the rapid multi-
plication 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-

202 PHARMACEUTICAL BOTANY

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, Hickories,
Cherries, Viburnums, and Oaks, etc., instead of cork cambium
arising only from the epidermis or the outer layer of cortex
cells, it may arise at any point in the cortex or even as far inward
as the pericycle or outer layers of phloem, and frequently several
cork cambia appear successively. As each new cork cambium
arises and cuts off cork cells on its outer face, all the living tissues
outside of the cork are deprived of nutrient sap and die. It is the
origin of cork cambia at varying depths that causes extensive
sheets of tissue to separate off. This dead tissue has been termed
borke by the German botanists (see Fig. 131). That is what gives
the stringy appearance to the stems of climbers.

At close of first year in a Perennial Dicotyl Stem we note:

1. Epidermis—development of dermatogen—in process of peeling off, later on
entirely absent.

2. Cork tissue.

3. Cork cambium or phellogen.

4. Sometimes a zone of thin-walled cells containing chloroplasts cut off by
cork cambium on inner face and known as phelloderm or secondary cortex.

5. Primary cortex—in perennials, stem cells of cortex may undergo modifica-
tion into mucilage cells, into tannin receptacles, crystal cells, spiral cells,
ete,

6. Endodermis, or innermost layer of cells of cortex, usually similar to other
cortex layers in character of cells.

7. Pericycle, a parenchymatous region usually appearing similar to the cortex,
but sometimes containing sclerenchyma fibers or stone cells or both.

8. Vascular bundles of open collateral type which are now arranged in a
compact circle, and between which are found primary- and often secondary
medullary rays. The longest bundles represent Primary Bundles which
have been deepened by the cutting off of secondary xylem and phloem by
the cambium.

From without inward the following tissues make up the bundles:

Protophloem Hard Bast—long tenacious bast fibers (sometimes
absent). ;

Secondary Phloem _) Soft Bast—phloem cells, companion 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 trachez.
9. Pith.

THE STEM 203

EXCEPTIONAL Types or DicotyL STEMS

In a number of Dicotyledons and Gymnosperms, the second-
ary 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 stems of Phytolacca, etc., there first arises a ring of primary
bundles with broad, loose medullary-rays. Then the stem

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Fic. 132.—Transverse section of the herbaceous dicotyledonous stem of Solanum
tuberosum (Fam. Solanacez) showing bicollateral bundles with internal phloem.
X 75. ep, epidermis; co, cortex; ph, external phloem; ca, cambium; x, xylem; 7,
internal phloem; si, sieve tissue; m, medulla. Both phloem regions contain primary
phloem.

cambium ceases its activity and, outside the bast of the bundles
already formed, in the pericycle 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, the cambium closes up. Still
another cambium ring arises without this which lays down
another circle of bundles and, in a single season quite a number of
these are found successively arranged in concentric fashion.

204 PHARMACEUTICAL BOTANY

In Gelsemium, species of Solanacea, Combretacea, Cucurbitacee,
etc., there arises a cambium on the inner face of the xylem which
forms tnternal phloem (or intraxylary
| phloem), thus giving rise to bicol-
| lateral bundles. (See Fig. 132.)

In Strychnos Nux Vomica,
internal phloem, exactly as in
Gelsemium, 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 FoRMA-
TION.—Lenticels are openings in
the periderm which provide a

Fic. 133.—White birch (Betula Means of communication between
populifolia). Portion of a branch the external air and the living cells
soil: the prominent _lenticels. of the bark beneath. The epi-

oy dermis 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.

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Fic. 134.—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 some of the
stomata loosely arranged cells called “complimentary cells which

THE STEM 205

are cut off by the cork cambium in certain places instead of cork.
These swell up during rain and pressing outward, form fissures
in the epidermis, called lenticels.

The function of lenticels is similar to that of stomata, namely,
to permit of aération of delicate cells of the cortex beneath.

Fic. 135.—Representative: portion of a transverse section of a Linden stem, 3
years old.—1x, 2x, 3x, Successive annual rings of wood. ph, Phloem, with bast
fibers showing as lighter parts and sieve-tubes as darker parts. c, Cortex containing
chloroplasts and rosette crystals of CaO. m.r., spreading end of primary medullary
ray. pe, periderm. (From Small, after Kny.)

While common to many plants, they are very conspicuously
prominent in the birches and cherries.

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 growth is lessened and small ducts and much
mechanical woody fiber are formed. Thus the open, loosely

206 PHARMACEUTICAL BOTANY

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 “Annuat Rinc.” By counting the number of these rings
it is possible to estimate the age of the tree or branch.

Primary Medullary Ray

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= SERN
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Spring | ¥ (55 0 nae aoe ee a
Wood ita e Y cigee
See
ee
Summer q e
Wood

Fic. 136.—Stem of Linden, showing detailed structure of a small square area
of Fig. 135, magnified 200 times. S, sieve tubes; the dark spaces at one corner of
the sieve-tubes are companion cells. V, pitted trachee. The summer wood
consists of wood fibers interspersed with parenchyma. (From F. O. Bowers? ‘Plants
and Man.” By permission of the Macmillan Co., Publishers.)

Barx.—Bark 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 region (sometimes with epidermis).
_ 2. Middle Bark or Cortex region.
3. Inner Bark or Phloem region (sometimes with pericycle attached).

THE STEM 207

Corx.—Cork or phellem is all that tissue produced by the cork
cambium on its outer face. Its walls become impregnated with
a waterproof substance of a waxy character called suberin.

og, KE)
“
CO . SE
eves S:85e ee

f eornsh

Fic. 137.—Part of a cross-section through branch of Cytisus laburnum. (The
branch was cut from the tree at the end of October.) From A to £ the last annual
ring of wood; from A to B the spring growth with large tracheal tubes (7, 7, 7);
between B and C and D and D are wood-fibers; between C and D and D and £,
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 paren-
chyma formed the previous year. (From Sayre after Haberlandt.)

PERIDERM.—Periderm is a name applied to all the tissue
produced by the cork cambium (Phellogen) externally and inter-

208 PHARMACEUTICAL BOTANY

nally and includes the cork cambium itself. ‘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. Its
cells frequently contain chloroplasts.

Fic. 138.—Cascara Sagrada Bark. Transverse section. A, cork; kk, cork
cambium; fh, phelloderm; st, group of stone cells; C, primary cortex; rer, rosette
aggregate of calcium oxalate; P, phloem; df, bast fibers; crf, crystal fibers more or
less surrounding a group of bast fibers; mr, medullary ray; mp, monoclinic prism

of CaO.

Histrotocy or A TypicaL Bark, CascaraA SacRaDa.—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 rec-
tangular 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, brownish red or
purple contents.

v

THE STEM 209

2. Cork Campium (phellogen), a layer of delicate cells in the
process of division with protoplasmic contents.

2 rer
(=3
ei ay a
CS mr
ee ; —bf
= ni J =
p
LF = ee ee
Po is
=iS-4 eae, 9 285%

—crf

A

te .

Fic. 139.—Cascara Sagrada Bark. Radial-longitudinal section. &, cork, kk,
cork cambium; ph, phelloderm of collenchymatous cells; c, primary cortex; st,
group of stone cells; mp, monoclinic prism; rcr, rosette crystal of calcium oxalate;
mr, medullary ray; bf, bast fibers; s, sieve tube; , phloem; crf, crystal fibers adhering
to bast fibers.

3. CorTEx, or middle bark, consisting of two regions, viz.: an
outer zone of two or three rows of brownish collenchyma cells,
and an inner broader zone of tangentially elongated, cortical
parenchyma cells. Imbedded within this zone will be noted
numerous groups of stone cells. Rosette crystals and monoclinic

210 PHARMACEUTICAL BOTANY

prisms of calcium oxalate occur in parenchyma cells of this
region.

4. PHLoeM, or inner bark, a very broad zone composed of
irregular-shaped, elongated phloem masses separated from each
other by medullary-rays which converge in the outer phloem
region. Each phloem mass consists of numerous sieve tubes,

rcr

a) 2 $i

mp

fi

Fic. 140.—Cascara Sagrada Bark. Tangential-longitudinal section through
outer phloem region. , phloem parenchyma; sf, group of stone cells; rcr, rosette
aggregates of calcium oxalate; mr, medullary ray; si, sieve tubes; crf, crystal fibers
containing monoclinic prisms of calcium oxalate; mp, monoclinic prism.

companion cells and phloem cells, some of which latter contain
spheroidal starch grains while others contain either monoclinic
prisms or rosette aggregates of calcium oxalate. Embedded
within the phloem masses, in tier-like fashion, will be noted
groups of bast or phloem fibers, each group of which is sur-
rounded by a row of crystal fibers, individual cells of which can
only be made out in this kind of a section. Each of these con-
tains a monoclinic prism of calcium oxalate. The medullary-

THE STEM 211

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

f

co

ST

ph

xT

td?

td‘

Fic. 141.—Transverse section of representative portion of the stem of the second
year’s growth of spruce (Picea sp.), a conifer. (X 80.) ep, epidermis; ck, cork;
pn, phellogen; co, cortex; sr, secretion reservoir; ph, phloem; ca, cambium; xr,
xylem ray; td', tracheids of autumn wood of first year; id?, tracheids of wood of
second year.

fibers here will be seen to be composed of vertical rows of super-
imposed thin-walled cells each of which contains a monoclinic
‘ prism of calcium oxalate. The phloem (bast) fibers appear
elongated and taper-ended and are associated with crystal fibers.
The sieve tubes show sieve plates on their longitudinal as well as
their transverse walls in this bark.

212 PHARMACEUTICAL BOTANY

In a tangential longitudinal section which has been cut through
the phloem, the exact range in width of the medullary-rays may

on
LSS»

<n

‘Medullary ray

Fic. 142.—Diagrammatic representation of a block of pine wood, highly mag-
nified. a, Early growth; 4, late growth; c, intercellular space; d, bordered pit in
tangential wall of late growth; m, f and e, borderéd 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; /, thin place in radial wall of ray
cells that carry food. (From Stevens.)

be ascertained. In this bark the medullary-rays are spindle-
shaped in tangential view and one to four cells in width.
Woop.—From a practical pharmacognic standpoint as well
as that of the lumber trade, wood is all that portion of woody
exogenous plant axis inside of the cambium line. In Dicotyl

THE STEM 213

and Gymnosperm stems it therefore includes the xylem regions
of the bundles, the 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. From the pure botanical viewpoint, wood is
synonomous with xylem and so represents that portion of a
vascular bundle that contains trachez or tracheids or both.

As the cambium year after year adds new layers of wood to
that already present on its inner face, the conveying of sap and
storing of starch, etc. is gradually relegated to the outer wood
layers, since the inner layers, step by step, lose their protoplasmic
contents and power of conducting sap and become filled with
extractive, resinous and coloring matters. The outer whitish
layers of wood which contain living cells, functioning in the
vegetative processes of the plant, constitute the ALBURNUM or
Sap-woop. The drug Quassia is a good example of this kind of
wood. ‘The inner dead colored layers constitute the DURAMEN
or HEART-woop. Important examples of this kind of wood used
in pharmacy are Lignum Guaiaci, Haematoxylon and Santalum
Album.

Ty.tosrs.—These are bladder-like enlargements of cell walls
usually of wood parenchyma or medullary ray cells which extend
into the lumina of adjacent trachez or tracheids through pit
cavities in the walls of the latter. ‘They are formed by the surface
growth of the pit membranes of the one-sided bordered pits which
push their way into the lumina of the vessels or tracheids. In its
living state the tylosis contains cytoplasm, cell sap, and often a
nucleus, the latter having passed into it from the parent cell. In
its fully developed state it may contain starch, gum, resin and
crystals, and its wall may be wrinkled. Sometimes a single
parenchyma cell may produce several tyloses. Tyloses occur
both in angiospermous and coniferous woods although are
probably more common in the former. They are more abundant
in heartwood than in sapwood and are of constant occurrence in
some species. Excellent examples of tyloses can be seen in the
rhizomes of Asarum and Serpentaria and in the stems of Sassafras,
Juglans, the locust (Robinia), Catalpa, Poplar, Sumachs, Oaks

and White Pine.

214 PHARMACEUTICAL BOTANY

Microscopic CHARACTERISTICS OF ANGIOSPERMOUS AND
Conrrerous Woops.—The wood of Angiosperms is character-
ized by the presence of tracheze (vessels) with various markings
on their walls, particularly by small pits in the walls of some of

rte gt

sk a
rs oyee 9
a eee

B

Fic. 143.—A, diagram of the course of vascular bundles in a palm stem (mono-
cotyledon); 1m, 2m, 3m, bundles from the median portions of the leaves. B,
diagram of vascular bundles in external view and in cross section of the dicotyledon-
ous stem of Cerastium. The leaves are shown cut off at 1, 2, 2’ and 3. (After
Vines.)

the tracheze, together with wood fibers, fiber tracheids,' wood
parenchyma cells and medullary-rays.

The wood of Conifers is made up for the larger part of
tracheids with bordered pits which latter are characterized 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 wood parenchyma cells (in larches and Douglas Fir, etc.),
are also found.

1 Forms intermediate between typical tracheids and wood fibers.

EE eS eS Pe. Se

THE STEM 215

Hisrotocy or TypicaL Hrersactous Monocoryt STEMs
(ENDoGENous).—Passing from exterior toward the center the
following structures are seen:

1. Epidermis whose cells are cutinized in their outer walls.

2. Hypodermis, generally col- a
lenchymatic in young, terminal
portions, becoming later scler-
enchymatous.

Fic. 144. Fic. 145.

Fic. 144.—Photomicrograph of cross-section of very young cornstalk (a herbace-
‘ous monocotyl stem), where certain plerome strands have just gone over into
vascular bundles. For comparison with Fig. 145. (Stevens.)

Fic. 145.—Photomicrograph of cross-section of older cornstalk stem; a, epi-
dermis; 6, cortex and c, ground tissue of pith. (After Stevens.)

3. Cortex, of cortical parenchyma cells. - Sometimes vascular
bundles of the closed collateral or the concentric type occur in
this region.

4. Endodermis or innermost layer of cortex and Pericycle
(frequently appearing like and often indistinguishable from corti-
cal parenchyma cells).

5. Stele or Pith, a large central zone of parenchyma matrix in
which are found scattered fibro-vascular bundles of the closed
collateral or more rarely of the concentric monocotyl (phlocentric)
type. In this latter type, which is typical of old monocotyl stems
and also seen in Calamus, Convallaria, etc., the xylem grows
completely around the phloem so that phloem is found in the
center and xylem without and surrounding it.

HisroLocy oF A TypicaL MonocotyL Ruizome.—The
rhizome of Convallaria majalis (Lily-of-the-Valley) shows typical
herbaceous monocotyl rhizome structure. A transverse section

216 PHARMACEUTICAL BOTANY

(see Fig. 147) presents the following structural details, passing
from periphery toward the center:

1. Epidermis of a layer of epidermal cells whose outer walls are
covered with a waxy cutin deposit.

2. Cortex of about 20 rows of ordinary parenchyma cells, some
of which contain more or less spheroidal starch grains, others
raphides of calcium oxalate.

. ‘ o.
eo *e@

et
‘6 .

Ss
AR
t*

q { ke .

» y OF. Aes:
=4 a a ar
\ A, i ies
@%a a og
“a LA

}

leeds

ane,

AS

§ 8.
Sy

“eres
=e aM
e,*

Sand
x

Fic. 146.—Transverse section through a representative portion of the mono-
cotyledonous stem of the corn (Zea Mays) (X 50). ep, epidermis; A, hypodermis
containing lignified fibers; co, cortex (poorly defined in corn); ph, primary phloem
and x, primary xylem of a closed collateral bundle surrounded by a sclerenchyma
sheath of lignified fibers (scl); par, parenchyma of pith in which the bundles are
scattered; sz, sieve tissue; tr, tracheae; a, a ring of an annular trachea bordering a
large air-space; scl, sclerenchyma fibers. The two large lateral trachez in each
bundle are pitted tracheae.

3. Endodermis of usually 2 layers of endodermal cells, the radial
and inner walls of which are strongly indurated with lignin.

4. Stele, a broad central region consisting of a matrix of
starch- and crystal-bearing parenchyma and through which
region course closed collateral and concentric bundles.

The closed collateral bundles are arranged in an interrupted
circle just beneath the endodermis. The concentric phlocentric
bundles are scattered through the center of the stele.

THE STEM 217

Longitudinal-radial sections show that the trachez are of the
spiral, scalariform, and pitted types.

HistoLocy or A TypicAL Woopy Monocoty.L StemM.—The
stem of the Greenbrier, a woody monocotyl, will here be con-
sidered. In transverse section passing from periphery toward the
center the following structural details will be noted:

ep

nt ;

eR

Fic. 147.—Transverse section of the rhizome of Convallaria majalis, a mono-
cotyledon. X 40. ep, epidermis; co, cortex; end, endodermis; p, phloem and x,
xylem of a closed collateral fibrovascular bundle; p’, phloem and x’, xylem of a
concentric bundle; m, pith, in which the concentric bundles are imbedded.

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 evapora-
tion of water, the ingress of destructive parasites, and injury
from insects.

2. A cortex, composed of about ten or twelve layers of thick-
walled parenchyma cells, the outer two or three layers of which
are termed hypodermis.

218 PHARMACEUTICAL BOTANY

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 sclerenchyma (pericyclic) fibers and unde-
veloped fibro-vascular bundles of the closed collateral type. —

Fic. 148.—Photomicrograph of a representative portion of Greenbrier stem
showing epidermis (¢.p.), cortex (c), endodermis (e.n.d.), cylinder sheath (c.s.),
sclerenchyma fibers of closed collateral bundle (6), fundamental parenchyma
(f.p.), trachea (t). X 22.

5. A pith or central matrix of strongly thickened, more or less
lignified, parenchyma 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 trachez and 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

THE STEM 219

bundle will be seen an area of soft, small-celled sieve tubes,
companion cells and phloem parenchyma. The entire bundle is
enclosed by a several layered ring of sclerenchyma fibers.

HistoLocy or Stems oF Grasses (MoNOcOTYLEDONOUS).—
The stems of grasses are solid in their terminal and nodal regions;

6a
Sau:

tec

Pee, .
36

~

Fic. 149.—Structure of a Grass Stem. ‘Transverse section of the underground
stem (rhizome) of the couch grass, Agropyron repens, X 35. ep, epidermis; h, hypo-
dermis; co, cortex; 6, concentric bundle; en, endodermis; scl, sclerenchymatous ring
of several rows of sclerenchyma fibers in which are imbedded an irregular, inter-
rupted circle of undeveloped vascular bundles; ér, trachea and ph, a phloem strand
of a closed collateral bundle; cb, closed collateral bundle; m, pith. Note the hollow
space in the center due to the disintegration of the pith in centrifugal fashion.

but as we pass backward from the terminal internode into the
older internodes the central pith is seen to disintegrate in cen-
trifugal fashion, leaving a hollow cavity in the central region
bordered by parenchyma cells. A distinctly differentiated
endodermis may or may not be present. The vascular bundles
may be entirely closed collateral, as in corn, but are frequently of
two kinds, namely, small, phlocentric bundles in the cortex and

220 PHARMACEUTICAL BOTANY

larger closed collateral bundles in the pith. The pith bundles
may be more or less uniformly scattered, as in the stems of corn
and bermuda grass, or arranged in one or more interrupted
circles, as in Triticum species, etc. The epidermal cells usually
_ possess cutinized outer walls and frequently all of their walls are
lignified. ‘The hypodermis contains lignified fibers, and there is
usually present a sclerenchyma ring directly beneath the endo-
dermis or the cortex in which bundles may be imbedded or to
which they may be attached. A sclerenchyma sheath usually
surrounds each bundle.

SECONDARY GROWTH IN MonocotyL StTEems.—Secondary
thickening in stems of monocotyledons is exceedingly rare and
the exception rather than the rule. It is seen, however, in some
of the more woody and tree-like genera of the Lily family
as Aloé, Yucca, Cordyline and Dracaena and in the tuberous Diros-

-coreacee. In these forms a cambium arises in the inner region of
the cortex or the pericycle and lays down procambial strands and
secondary parenchyma on its inner face. As it retreats outward,
the procambial strands, by repeated division and differentiation,
develop into secondary collateral (rarely phlocentric) bundles
which lie imbedded in secondary parenchyma.

CHAPTER X

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.

Fic. 150.—Complete leaf of Slippery Elm (Ulmus fulva). 1, lamina; p, petiole;
5, stipules; mr, midrib; 2, secondary vein; st, branch to which leaf is attached. The
margin of this leaf is doubly serrate.

Leaves seldom develop buds over their surface or along their
margin and in connection therewith roots. The capacity for bud
development by the leaf is restricted to three families, viz.: Cras-
sulacee, Begoniacee and Gesneracea.

Tue Comptete Lear.—The leaf when complete consists of

three parts, Jamina, petiole, and stipules. The Lamtna or BLapE
221

222 PHARMACEUTICAL BOTANY

is the largest, most prominent and essential part. Other parts
may be wanting. It consists of upper and lower surfaces
provided with a layer of protective epidermal cells which may give
rise to outgrowths in the form of hairs or papilla. Either or
both of these surfaces show openings called stomata (sing. stoma)
which are protected by guard cells that regulate their opening
and closing. ‘The stomata serve for the exchange of carbon

: U
Palisade parench. tia
> y F) 3 _

Fic. 151.—Stereogram of leaf structure. Part of a veinlet is shown on the right.
Intercellular spaces are shaded. (From Stevens.)

dioxide and oxygen and the escape of water vapor. Between
the epidermal layers, in the region called the mesophyll, is a soft
green tissue called chlorenchyma which is made up of leaf paren-
chyma cells containing chloroplasts and intercellular-air-spaces.
Coursing through the mesophyll are the veins or fibro-vascular
bundles which are in continuity with the fibro-vascular bundles
of the stem, These veins branch and rebranch and so make up
the frame-work of the lamina. The outer walls of the epidermal
cells are provided with a cuticle which is generally thicker on
the upper than on the lower surface. In most leaves the meso-
phyll is differentiated into upper palisade parenchyma, lower

i
|

:
4
4
4
:
4
3

THE LEAF Zio

spongy-parenchyma and border parenchyma regions, and stomata are
only found in the lower epidermis.

The Petio.e is the leaf stalk which supports the lamina in a
position where it will receive the most light for photosynthesis.
The Sriputes are leaf-like appendages appearing at. the base
of the petiole.

The leaf of the Slippery Elm (Ulmus fulva) affords a good
example of a Complete Leaf (see Fig. 150).

7

oY es
&
9 sew $02
Ww wi.

5 @
rg 0
oO 6 @ ; (Kirt >
: WALI ZA
% 8 \\\SR8 xo
o : oe f 86 2 =s

Fic. 152.—Camera-lucida drawing of a tangential section of a leaf (section
parallel to leaf surface) showing /f, intercellular-air-spaces between the palisade par-
enchyma cells (e); g, border parenchyma; h, tracheal elements of a vein. (Stevens.)

Sometimes the lamina or blade is attached directly to the
stem by its base and the leaf is then said to be sessile. If the
petiole is present, the leaf is petzolate.

When leaf stipules are absent, the leaf is said to be exstipulate;
when present, stzpulate.

The petiole is seldom cylindrical in form, but usually chan-
nelled on the upper side, flattened, or compressed. The stipules
are always in pairs and closely resemble the leaf in structure.

Lear Functions.—The most essential function of green plants
is the conversion 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.

224 PHARMACEUTICAL BOTANY

The chief functions of a leaf are photosynthesis, assimilation,
- respiration and transpiration.

PHOTOSYNTHESIS is the process possessed by the cells of all
green leaves or other green parts of plants of building up sugar,
starch or other complex organic (carbon containing) substances
by means of chlorophyll and sunlight. This process takes place
in nature, only during sunlight but will also occur in artificial
light. In this process CO, is taken into the leaf through the

Fic. 153.—The end of a very small veinlet as seen in a section of the spongy
parenchyma cut parallel to the leaf surface. The walls of the conducting cells
of the veinlet have spiral thickenings. The border parenchyma cells surrounding
the veinlets serve to conduct the sugar from the mesophyll to the phloem cells of
the vein. (From Mottier, after Strasburger, Macmillan Co.)

stomata and passes into the intercellular-air-spaces. between
the chlorenchyma cells. It diffuses through the cell walls of
these into the protoplasts where it passes into solution and enters
the chloroplastids along with H:O from the soil. The chloro-
plasts contain chlorophyll which absorbs the light energy. This
light energy is then used either by the chlorophyll or, as seems
most likely, by the whole chloroplast to unite the CO: with the
H,O to form carbon compounds. These contain stored or poten-
tial energy. ‘The kinetic energy of the sun’s rays thus becomes
stored as potential energy and O; is given off. The plant gains

THE LEAF 225

in weight and, so the process is constructive in character. In
photosynthesis, sunlight is the power, the chloroplasts the work-
ing machinery or energy transformers, the chlorenchyma cells
the factories, CO. and H,O the raw materials and glucose,
starch or oil the finished products, while oxygen is a waste

OXYGEN
CARBON DIOXIDE

~ MINERAL SALTS
WATER

Fic. 154.—Diagram of photosynthesis. (After Atwood.)

a is

<— GLUCOSE

ipsam

product. The production of glucose may be represented by
the equation:

6H2O + 6CO, + light = C.H,.0, + 602

AssIMILATION is the process of converting food material into
protoplasm. The chlorenchyma cells of the leaf synthesize
proteins from the carbohydrates. In this process molecules of
glucose derivatives containing CHO are united with nitrogen
absorbed from the soil as nitrates to form amino acids. The
amino acids combine to form polypeptides which are later built
into simple proteins. The simple proteins are joined with sulfur
or phosphorous or both to form complex proteins. ‘The latter
enter into the formation of protoplasm.

REsPIRATION is the process which takes place in all living
plant and animal cells whereby complex carbon compounds
are oxidized with an accompanying release of energy. In
higher green plants oxygen is taken in through the stomata of
leaves, lenticels of stems, and root hairs. It passes into the
intercellular-air-spaces which communicate with each other

226

PHARMACEUTICAL BOTANY

throughout the entire plant and diffuses through the various
cell-walls of cells, in contact with these, and passes into the

interior of the cells.

Here it oxidizes some of the carbohydrates

breaking these down into carbon dioxide and watery vapor.

Wid

rp yrtay
I

py eo

Fic. 155.—Demonstration of the
liberation_of oxygen during photo-
synthesis. A, release of bubbles of
oxygen gas from the freshly cut end of
the stem of Elodea in sunlight. B,
collection of the gas, mainly oxygen,
from Elodea in sunlight during photo-
synthesis. If an aquatic plant be
placed in a glass vessel of water under
an inverted funnel over whose stem
is suspended an inverted test tube
filled with water, the gas given off
by the plant will collect in the upper
end of the tube and displace the water.
Upon removing the tube and thrust-
ing into it a glowing splinter, the
latter will increase in brightness, indi-
cating the presence of oxygen. (After
Mottier.)

These diffuse out of the cells into
the intercellular-air-spaces and
pass out of the plant through the
same channels as oxygen entered.
Respiration goes on both in light
and darkness, organic compounds
are broken down, O is absorbed
and CO, set free, and potential
energy is transformed into kinetic
energy. The plant loses weight
and, accordingly, this process is
destructive in character.
TRANSPIRATION is the action of
giving off watery vapor. ‘The
greater portion of the crude sap
consisting very largely of water is
conveyed upward as a transpiration
stream through the trachee and
tracheids of the roots and stems
into those of the leaves. The
latter pervade the soft, green leaf
parenchyma and end in proximity
to air spaces between the green
cells. A portion of the crude
sap diffuses into the leaf paren-
chyma cells and is utilized in the
nutritive processes such as photo-
synthesis, digestion and assim-
ilation occurring there. The

remainder, which normally constitutes the larger part, passes
through the intercellular-air-spaces and out of the leaves as

watery vapor.

About 90 per cent. of the water thus transpired

escapes through the stomata, the remainder through the epi-
dermal surface. It has been estimated that a large oak tree

THE LEAF 227

transpires about 180 gallons of water per day and that during a
growing season of five months it would give off water enough to
cover the ground shaded by it to the depth of 20 feet.

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.

: “ By

ay Th JATT ®) eo
iY, ‘ ,
ras rt,
iimenan’:

\

Fic. 156.—Semi-diagrammatic cross section of a DorstvENTRAL Lear of a meso-
phyte showing by arrows how the water passes from the tracheal elements of
a vein into the border parenchyma cells, and thence into the palisade and spongy
parenchyma, from which it evaporates into the intercellular spaces and passes
from the leaf through the stomata. a, upper epidermis; 4, lower epidermis; ¢,
palisade parenchyma; g, spongy parenchyma; d, border parenchyma; e, tracheal
elements; and the stippled cells below ¢, the phloem cells. (After Stevens.)

4. Bract leaves; (a2) primary at base of inflorescence; (5)
bracteolar leaves (bracteoles) at the base of individual flowers.

5. Sepals.

6. Petals.

7. Microsporophylls (stamens).

8. Megasporophylls (carpels).

CotyLepons.—Cotyledons are the first leaves to appear upon
the ascending axis and are single in Monocotyledons, double in
Dicotyledons. 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

228 PHARMACEUTICAL BOTANY

Gymnosperms. In Monocotyledons 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.

Scate Leaves.—Scale leaves are reduced foliage leaves.
They are found on certain rhizomes, above ground stems, such
as Dodder, Indian Pipe, etc., on bulbs, and forming the pro-
tective scales of scaly buds.

Surface View Section

Fic. 157.—The growing end of the stem of the aquatic plant, Elodea canadensis,
showing the origin of leaves. The larger over-lapping leaves are omitted. A, sur-
face view; B, in section. (Courtesy of Standard Scientific Corp., formerly Scientific
Equipment Dept. of Kny-Scheerer Corp.)

Fouace Leaves.—These are the common green leaves so
familiar to all.

Bracr 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.

Oricin AND DeveLopMEnt or 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

THE LEAF 229

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 or leaf parenchyma 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 two motions, the circular and the longitudinal, and its
most common modification is the circle.

_ Foliage leaves are usually arranged in separated fashion along
the stem, one occurring at each node. This is the form of spiral
arrangement called the alternate arrangement, as shown by most
plants.

The alternate 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 (counting the leaf from which the start is
made), two turns being made around the stem to reach it.
Examples: Cherry, Apple, Peach, Oak and Willow, etc. As the
distance between any two leaves in this arrangement is two-
fifths of the circumference of the stem or 144°, the five-ranked
arrangement is expressed by the fraction 74. In the eight-
ranked arrangement, the ninth leaf stands over the first, and three
turns are required to reach it, hence the fraction 3g expresses 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, count-
ing along the spiral, from any one to the one directly above it.

230 PHARMACEUTICAL BOTANY

The one which is chosen as the starting point is marked and taken
as 0.

Other forms of leaf arrangement are:

Opposite, when a pair of leaves is developed at each node, on
opposite sides of the stem. Examples: Mints, Lilac.

Decussate, when the leaves are arranged in pairs successively
along the stem, at right angles to each other. Example:
Thoroughwort.

Whorled or Verticillate, when three or more form a circle about
the stem. Examples: Canada Lily and Culver’s Root.

Fascicled or Tufted, when a cluster of leaves is borne from a
single node, as in the Larch and Pine.

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
a we Tulip Tree, it is said to be

es inflexed or reclinate vernation; if
convolute plcate conduplicaté Goybled on the midrib so that
Fic. 158.—Three principal types of the two halves are brought
vernation. (Robdbins.) ‘
together, as in the Oak or
Peach, 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 toward the midrib on
the upper side, as the leaves of the Apple or Violet, involute;
when rolled outward from each margin as Dock or Willow leaves,
_ revolute. The inner surface is always that which will form the
upper surface when expanded.

Lear Venation.—This refers to the arrangement of the veins
within the leaf. It is dependent upon the course pursued by the
fibrovascular bundles after they enter the leaf blade.

The central vein in the leaf is termed the midrib or primary.
The branches coming off of the midrib are called secondary veins or
secondaries; those arising from secondary veins are called tertiares.

THE LEAF 231

Strong and thick veins are called bs and less prominent ones,
especially if lateral to the midrib, nerves.

FuRCATE- or FoRKED-VENATION is characteristic of leaves of
the Ferns.

3
6.

2. Parallel (Convallaria),
(Drawing by Canis.)

5. Palmate-reticulate (Acer).

1. Furcate (Botrychium, a fern).

4. Pinnate-reticulate (Castanea).

Pinnate-reticulate with marginal veins anastomosing (Eucalyptus).

Toa

Fic. 159.—Leaf venation.

Pinnately-parallel (Musa).

PARALLEL VENATION is typical of the leaves of Monocotyledons,
as Palms, Lilies, Grasses, etc. In this type of veining the veins
visible to the unaided eye extend in somewhat parallel fashion
through the leaf, converging near the summit. In pinnately-
parallel veined leaves, as the Banana and Calla Lily, parallel
veins extend from midrib to margin.

232 PHARMACEUTICAL BOTANY

RETIcCULATE or Netrep Verns characterize the leaves of
Dicotyledons, as Matico and the Poplar or Oak. The primary
veins in these are generally pinnate or palmate while the second-
ary ones and their branches are arranged in netted fashion.

PINNATELY-VEINED or FEATHERED-VEINED leaves consist of a
midrib with lateral veins, extending from’ midrib to margin at
frequent intervals and in a regular manner. Examples: Beech
and Chestnut leaves.

PALMATELY VEINED leaves consist of a number of veins of
nearly the same size, radiating from petiole to margin. Exam-
ples: Maple and Castor Oil leaves.

In palmately-veined leaves, the central vein is termed the
midrib or middle primary and those veins lateral to it, the dateral
primaries.

Most pinnately-veined leaves and palmately-veined leaves
show net-veins combined with the basic pinnate or palmate type
and so are said to exhibit pinnate-reticulate or palmate-reticulate
venation respectively. )

Veins are said to be ANAsTomosinc when they subdivide and
join each other, as the veins near the margin of Eucalyptus leaves;
TESsELLATED, when they form square meshes, as in Matico
leaves; and ImpresseD when sunken below the surface.

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 or a root.

Cauline, when upon an aérial 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).

When the bases of two opposite leaves are so united as to form
one piece, the leaf is called connate-perfoliate, as Eupatorium per-
foliatum or Boneset.

Leaves are called eguitant when they are all radical and succes-
sively folded on each other toward their bases, as in Jris spp.

THE LEAF 233

Tur Forms or LEAvVES.—SIMPLE LEAveEs are those having a
single blade, either sessile or petiolate.

Compounp LeAvEs are those whose blades are divided into
two or more distinct subdivisions called /eaflets. The leaflets may

(Drawing by

lanceolate (3); ovate (4); elliptical (5);
(8); spatulate (9); falcate (10); ensiform (11).

Fic. 160.—Leaf outline: Linear (1); linear-lanceolate (2);

orbicular (6); oblanceolate (7); cuneate

Canis.)

be either sessile or petiolate. If the compound leaf has leaflets
arising from the midrib, it is called Pinnately Compound.
The midrib of a compound leaf is sometimes called the rachis.
If the leaflets of a compound leaf come off of the summit of the

(234 PHARMACEUTICAL BOTANY

petiole, the compound leaf is termed PALMATELY- or DiciraTELy-
CoMPOUND. The leaf stalk of a leaflet is called the petiolule.

A compound leaf develops from a single leaf bud as does a
simple leaf. Even though a group of small leaves may appear as

Cordate (12);
Acuminate

paripinnate (28); bi-pinnate (29); decompound (30);

=

ss

Fic. 161.—Leaf bases (12-17); leaf apices (18-26); compound leaves (27—31).

auriculate (13); connate-perfoliate (14); sagittate (15); hastate (16); peltate (17).

(18); acute (19); obtuse (20); truncate (21); retuse (22); emarginate (23); cuspidate (24); mucronate
Imparipinnate (27);

(25); aristate (26).
palmately 5-foliate (31).

a compound leaf, they can be distinguished as simple leaves by
the presence of a bud in each of their axils.

Forms or Lamina.—The lamina of simple leaves and the
leaflets of compound leaves are described as to general outline,
apex, base, marginal indentations, surface and texture.

THE LEAF 235

(a) GENERAL OurT.ine (form viewed as a whole without
regard to indentations of margin).

Linear, long, narrow, ribbon-like of nearly equal breadth
throughout, as Linaria.

Filiform, thread-like, as the Asparagus and one of the Sundews.

Acerose or acicular, needle-like, as in the Pines.

Subulate, awl-shaped, as in the Arbor Vitz.

Ovate, egg-shaped, with its length not more than two or three
times its breadth, as Boldo.

Obovate, reversely ovate as some leaves of Short Buchu.

Rhomboidal, angularly-ovate, as the Salt Bush.

Lanceotate, 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. |

Equilateral, margin of the same length on both sides.

Inequilateral, margin longer on one side than the other, as the
Hamamelis, Elm and Linden.

Orbicular, the blade circular in shape. Example: Nasturtium.

Reniform, or kidney-shaped, as the leaves of Canada Snakeroot.

Oblanceolate, reversely lanceolate. Example: Chimaphila.

Spatulate, like a spatula, with narrow base and broad rounded
apex. Example: Some leaves of Uva Ursi.

Ensiform, when shaped like a sword. Example: Calamus. _

Falcate, scythe- or sickle-shaped, as leaves on older branches of
the Eucalyptus.

Deltoid, when the shape of the Greek letter A, as Chenopodium.

(b) Apex or Lear.—Acute, when the margins form an acute
angle at the tip of the leaf giving the tip the appearance of a sharp
point. Examples: Eriodictyon, Digitalis.

Acuminate, when the point is longer and more tapering than -

the acute. Examples: Pellitory, Coffee.

Tapering, if the summit of the leaf be narrowed very grad-
ually to a point, as in leaves of many grasses.

Attenuate, if the summit is still more drawn out than the taper-

ing type.

236 PHARMACEUTICAL BOTANY

Obtuse, blunt or rounded. Example: Long Buchu.

Rounded, when extremely obtuse, as in some leaves of F oxglove.
Truncate, abruptly obtuse, as if cut square off. Example:

Melilotus leaflets.

Mucronate, terminating in a short, soft point. Example:
some Senna leaflets.

: Cuspidate, like the last, except that the point is long and rigid.

Anstate, with the apex terminating in a bristle.

Emarginate, notched. Example: Pilocarpus.

Retuse, with a broad, shallow sinus at the apex. Example:
Petal of Rose gallica.

Obcordate, inversely heart-shaped. Example: Oxalis.

(c) Base or Lear.—Cordate, heart-shaped. Examples: Cat-
nep, Violet and Coltsfoot leaves.

Reniform, kidney-shaped. Examples: Ground Ivy, Asarum.

Hastate, or halberd-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. Examples: Bindweed and Sagittaria.
Cuneate, wedge-shaped. Examples: Short Buchu and Uva
Ursi.

Peltate, or shield-shaped, having the petiole inserted at the
center of the lower surface of the lamina. Example:
Podophyllum.

Oblique, when the base of the lamina is attached lower on one
e of the midrib than the other, as in the Stramonium.

The bases of many leaves extend downward along the stem
beyond the main point of insertion when they are called decurrent.
If the base of the lamina is prolonged downward along the sides
of the petiole to which it is joined, the leaf is said to have a
winged petiole, as in Digitalis,

The terms acute, tapering, attenuate, truncate, obtuse and rounded
which have been defined under “Apex of Leaf’? are also applied
to bases of like character.

(2) Marc or Lear.—Entire, when the margin is an even
line. Example: Belladonna.

Revolute, margin rolled backward, as in Thyme.

sid

os

es a st ae

THE LEAF 237

Involute or incurved, margin rolled inward.

Serrate, with sharp teeth which incline forward like the teeth
of a hand-saw. Examples: Peppermint, Yerba Santa, Buchu.

Serrulate, when finely saw-toothed, as in Rose leaflets.

Dentate, or toothed, with outwardly projecting teeth.
Example: Damiana.

Denticulate, when finely toothed.

Crenate, or Scalloped, similar to the preceding forms, but with
the teeth much rounded. Examples: Digitalis, Catnip.

Crenulate, when finely scalloped, as in Sage.

In the serrate, dentate and crenate forms, when the teeth
show divisions into smaller or secondary teeth, the word “doubly”
is used as a prefix to the term expressing the character of the
primary tooth, thus, Doubly-serrate, etc.

Spinose, when the teeth are extended as spines, as in the
American Holly. :

Ciliate, when the margin is fringed with hairs, as in Drosera.

The sinuses are the indentations of the margin. _ If sufficiently
deep, they separate the leaf into. segments or lobes.

Repand, or Undulate, margin—a wavy line. Example:
Hamamelis.

Sinuate, when the margin is more distinctly sinuous than the
last. (Stramonium.)

Incised, cut by sharp, irregular incisions. Example:
Hawthorn.

Runcinate, the peculiar form of pinnately-incised leaf observed
in the Dandelion and some other Composite in which the teeth are
recurved.

A Lobed leaf is one in which the indentations extend toward
the mid-rib, or the apex of the petiole, 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 or the petiole. Example: Geranium maculatum.

In the Divided leaf the incisions extend to the mid-rib, or the
petiole, but the segments are not stalked nor are they in the form
of leaflets. Example: Watercress.

238 PHARMACEUTICAL BOTANY

If the venation is pinnate, the preceding forms may be
described as pinnately-cleft (incised), -lobed, -parted, or -divided.
If the venation is (radiate) palmate, then the terms palmately-
lobed, -incised, etc., are employed.

pinnately-parted (34);

-lobed (36); palmately tri-cleft (37); palmately 3-parted (38);

pinnately-cleft (33);
serrate (41); dentate (42); repand or undulate (43); sinu-

: Pinnately-lobed (32);

162.—Leaf margins
pinnately-divided (35); palmately tri

Fic.
palmately 3-divided (39); crenate (40);

ate-dentate (44).

Pinnatifid is employed by some authors for describing 4
pinnately-cleft leaf, pinnatipartite for a pinnately-parted one, and
pinnatisect for one that is pinnately-divided. Likewise palmatifid,
palmatipartite and palmatisect are sometimes employed in place

THE LEAF 239

of palmately-cleft, palmately-parted and palmately-divided
respectively. |

We

Fic. 163.—Forms of Leaves. Runcinate leaf of Dandelion (1); lyrate leaf of
Turnip (2); interruptedly-pinnate leaf of Potato (3); palmate leaf of Horsechestnut
(4); inequilateral leaf of Witch Hazel (5); pedate leaf of Hellebore (6). Modified
leaves of insectivorous plants 7-9. Leaf of a Sundew bearing tentacles (7); leaf
of Venus Trap (8); pitchered leaf of Sarracenia purpurea (9); leaf of Heartsease
showing free lateral stipules (10).

A Pedate \eaf is one which is palmately-parted or divided but
which has its lateral lobes in turn divided in more or less linear

fashion.

240 PHARMACEUTICAL BOTANY

When the lobes or divisions of a pinnately-divided, -parted
or cleft leaf are alternately large: and small, the leaf is inier-
ruptedly pinnate, as the Potato leaf. When the terminal lobe is
the largest, and the remaining ones diminish in size toward the
base the form is known as /yrate, illustrated in the leaf of the
Turnip. |

Compounp LEAvEs.—The transition from Simple to Com-
pound 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.

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, ér2folzate, or ternate, as in
Menyanthes; when four in number, quadrifoliate, as in four-leafed
clover, etc. If each of the leaflets of a palmately 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 the Honey Locust
(Gleditschia); if these become again divided, as in many Acacia
species, the leaf is termed tripinnate. Beyond this point the leaf
is known as decompound. Examples of pinnately-decompound
leaves are seen in Cimicifuga and Parsley.

Lear Moprrications.—The leaves of a number of plants have
become modified in one or more parts for the purpose of carrying
out special functions coincident with habits acquired by the
plants possessing them. Thus, in the common Garden Pea, the

THE LEAF 241

upper leaflets are transformed into tendrils for climbing purposes;
in the Barberry some of the leaves have become transformed into
spines for the defense of the plant from browsing animals; in
Squill and Garlic the leaves of the bulbs have become succulent
scales for food storage; in the Bladderworts certain submerged
leaves have become modified as bladder-like traps for capturing
crustaceans; in Sundews, Venus Fly Trap and the Pitcher Plants
(Heliamphora, Darlingtonia, Sarracenia, Nepenthes and Cephalotus)
the leaves are variously modified as traps for alluring, capturing
and devouring insects. (See Carnivorous Plants in Chapter on
Ecology.)

Lear TEXTURE.—Leaves are described as:

Membranous, when thin and pliable, as Coca.

Succulent, when thick and fleshy, as Aloe, and Live Forever.

Coriaceous, when thick and leathery, as Eucalyptus, Uva Ursi
and Magnolia.

Lear 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.

Lear SurFACE.—Any plant surface is:

Glabrous, when perfectly smooth and free from hairs or pro-
tuberances. Example: Tulip.

Glaucous, when covered with bloom, as the Cabbage leaf.

Punctate, when dotted as if by punctures, as in the leaf of the
Wax Myrtle. ;

Pellucid-punctate, when dotted with projections formed by
subjacent oil glands, as the leaves of the many members of the
Orange family.

Papillose, when covered with minute, teat-shaped projections,
as the lower surface of Coca leaves.

Scabrous leaves have a rough surface with minute, eeccied,
hard papillae or warts, as those of Slippery-elm and many

composites.

Pubescent, covered with short, soft hairs. Example:
Digitalis.

Appressed-hairy, when the hairs are curved so as to appear
somewhat pressed against the surface, as in Senna leaflets.

242 PHARMACEUTICAL BOTANY

Villose, covered with long and shaggy hairs. Example:
Forget-me-not.

Sericious, when covered with hairs which give a silky appear-
ance. Example: Silverleaf.

Hispid, when covered with short, stiff hairs. Example:
Borage.

Strigose, when covered with appressed, stiff and sharp hairs.

Tomentose, densely pubescent and felt-like, as the Mullein leaf.

Spinose, beset with spines, as in the Thistle.

Rugose, when roughened with closely set wrinkles. Example:
Sage.

Verrucose, covered with protuberances or warts, as the calyx of
Chenopodium.

Tuberculate, when covered with minute wart-like projections.

Bullate, when blister-like projections appear between the
veins.

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 Lear.—The parts of a typical leaf are
petiole or leaf stalk, /amina or blade, and stipules.

Tue PETIOLE

The PretioLE iv MonocotyLepons is usually a broadened,
sheathing basal structure which connects the lamina to the stem.
In the Grasses and Bananas it forms a long tubular sheath which
surrounds the stem. At the point of union of the sheathing
petiole and the lamina of a grass is a membranous or leathery
ring or flap called the Licute. The ligule is not a part of the
petiole but is a formation due to the upgrowth and union of a pair
of stipules. Into the Grass petiole a set of closed collateral
vascular bundles of the stem extend, these showing xylem upper-
most and phloembeneath. Inthe Palms, Aroids, Yams, Bananas,
the petiole in part or throughout may be much thickened,
strengthened and developed as a semi-cylindric or cylindric
structure frequently showing, as in Palms, generally, two sets of

THE LEAF 243

bundles. In all of these the petiole shows distinct closed col-
lateral to concentric bundles embedded in parenchyma which in
turn is surrounded by epidermis. In the Monocotyl genus
Maranta of the Arrowroot family, a special swelling is found at
the apex of the petiole which is termed a Pulvinus.

In DicoryLepons, the PETIOLE attains its most perfect devel-
opment and here usually shows differentiation into a short basal
part called the Putvinus or leaf
cushion and a longer part, the
STALK portion. (See Fig. 18.)

The pulvinus is sensitive to
environal stimuli and in some
groups, as Oxalidacee and Legu- end \
minosé, 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. Fic. 164.—Barley, (a member of

The stalk portion of the petiole the Grass Family). 4, portion of
: . : leaf at juncture of blade and decurrent
In Dicotyledons 1S usually plano- sheath; B, stem cut in median longi-
convex or nearly to quite Cir- tudinal section. 214. (Robbins.)

cular in outline; rarely in certain
families does it resemble 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 Parsley family (Umbellifere) where the flattened
sheathing leaf stalk is known as the PericLaprum. Such a
structure is not peculiar to the Umbellifere, for in many members
of the Crowfoot family (Ranunculacee), etc., a similar sheathing
development is observed. The stalk may bear the laminar tissue
on its extremity. This is most commonly the rule, but when the

Sas i a

244 PHARMACEUTICAL BOTANY

plant is exposed to xerophytic conditions, as the Acacias of
Africa and 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. Such a
structure is known as a PHyLLope or PHyLiopium. (Fig. 165.)

Stoma

Fic. 165.—A Phyllode, or Blade-like Petiole, from Acacia melanoxylon, and its
Cross-Section. Drawing by H. McCarthy. (After Stanford, General and Economic
Botany, D. Appleton-Century Co. publishers.)

The INTERNAL STRUCTURE OF THE PETIOLE in primitive types
of Dicotyledons resembles that seen in Monocotyledons except
that the bundles are more condensed side by side. In these the
petiole is somewhat flattened and shows an external epidermis, a
flattened cortex and a single row of vascular bundles, each with
xylem uppermost and phloem below. From this we can pass to
another group of these plants in which the bundles form three-
fourths of a circle and in which the upper bundles show incurving

THE LEAF 245

orientation, to still another in which, as in Nepenthes, all of the
bundles form nearly a cylinder. Finally, in Ficus, Geranium,
Podophyllum and other plants showing completely formed cylin-
drical petioles, the bundles form a continuous ring enclosing
pith and surrounded by cortex and epidermis, as in Dicotyl stems.

STIPULES

Stipues 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 Leguminosae, 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 membranous to leathery, scale-like and cadu-
cous. Caducous scaly stipules only function as bud scales, forming
a protective covering to the bud through the winter and fall in
spring as the buds expand. Cf. Fig. 163 (10).

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 Roses, in Clovers, etc.

LATERAL CONNATE STIPULES are such as join and run up with
the petiole to form a structure which is called a Licute. This
structure is common to the Gramineae or Grass family. (See Fig.
164.)

LATERAL INTERPETIOLAR STIPULES are common to many
species of the Rubiaceae. ‘This family has opposite leaves and in
some of the genera, as Cinchona, one stipule from each leaf on
either side of the stem unites by its margin with the margin of
the other stipule to form an interpetiolar stipule. In this way a
stipule is formed on either side of the stem between the opposite
leaves.

The AxILLAry Group represents stipules which stand in the
axil of the leaf with the stem. Such may be Free AxILLary
STIPULES, arising as distinct processes, or CONNATE, when the two

246 PHARMACEUTICAL BOTANY

stipules unite at their margins, grow up and sheathe the stem,
forming a legging-like sheath, as in many species of the Poly-
gonacee such as Buckwheat, Rhubarb, Yellow Dock, Knot Weeds,
etc. The sheath formed is called an OcHREA.

Mobptrriep StreuLEs.—In some plants such as the Locust and
several other trees and shrubs of the Legume family, the stipules
become modified for defensive purposes as spines or prickles.
In the Sarsaparilla-yielding plants and other species of the genus
Smilax they undergo modification into tendrils which are e useful i in
climbing. (See Fig. 366.)

THe LAMINA

This, as was previously indicated, represents an expansion of
the tissues of the petiole, but in sessile leaves is directly attached ~
to the stem and so is a direct stem outgrowth.

Mope or DEVELOPMENT OF THE LAMINA OF LEAvEs.—The
lamina of leaves develops in one of six ways.

1. Normal or Dorsiventral.

Convergent.
Centric.
Bifacial.
Reversed.

. Ob-dorsi-ventral.

The first four will only be considered.

A. DorsIvENTRAL (the commonest).

(a) DorstvENTRAL UmsBropuytic.—Flattened from above
downward. Plants with such leaf blades tend to grow in the
shade.

(6) DorstvENTRAL Mesopuytic.—Similar to the former, but
plants usually grow directly in the pporhe and exposed to sunlight
and winds.

(c) DorstvenTRAL XeEROpHyTIC.—Similar to former, but —
plants not only grow exposed, but exposed to hot desert conditions
or to cold vigorous conditions.

(7) DorstvENTRAL Hypropuytic.—All transitions occur
between typical mesophytic forms to those of marshy places, to
swamps and borders of streams and finally with leaves wholly
emersed, the last a completely hydrophytic type.

Auk WN

THE LEAF 247

Gross Structure AND Hisrorocy or DirreERENT TYPES OF
DorsIVENTRAL LEAF BLApEs.—1. DorsIVENTRAL 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.

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- 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 chlorenchyma is fairly
spongy, the spongy parenchyma having decided intercellular
spaces. Examples: Hydrastis, Asarum, Maranta.

2. DorsIVENTRAL Mesopuytic.—In this type of dorsi-
ventral leaf there is a tendency to subdivision, either to slight or

OS ee ee ge ee ee Te EAE SE

248 PHARMACEUTICAL BOTANY

moderate lobing, seldom to complete subdivision in pinnate or
tripinnate fashion. Examples: Belladonna, Hyoscyamus,
Dandelion, Grasses, etc. In microscopic structure, they consist
of an upper and lower epidermis, the upper epidermis being the
thicker of the two, the lower epidermis having wavy vertical
walls. The stomata are wholly or are mainly on the lower
epidermis. In mesophytic trees and shrubs they are restricted
generally to the lower surface while in herbs of mesophytes they
may occur on both surfaces although mostly on the lower surface.
In mesophytic dicotyledons, the palisade parenchyma is toward
the upper epidermis and consists of one to three layers of cells.
The spongy parenchyma is toward the lower and consists of
irregular, loosely arranged cells with prominent intercellular-
air-spaces. In a number of mesophytic monocotyledons like
the grasses and sedges the chlorenchyma is devoid of ore
cells, being compact and uniform.

3. DoORSIVENTRAL XEROPHYTIC.—Leaves characterized by a
thick upper and lower cuticle and by having their numerous,
small stomata restricted to the lower surface or present more or
less equally on both surfaces, where they are sunken in depres-
sions or pits. They may be either firm, leathery, tough, or
fibrous, or may become swollen up in their mesophyll, chiefly in
their spongy parenchyma cells, and store considerable mucilage.
The. object of mucilage is to retain water. Very frequently they
develop a water-storage tissue above the mesophyll. Their
palisade tissue is usually well developed, occurring in the cacti
in many rows. Examples: Boldo, Eriodictyon, Ficus.

4. DorstvENTRAL Hypropnytic.—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 on the upper surface. In Ranunculus,
the lower leaves are cut up into filiform segments. These are
devoid of stomata. Their mesophyll is soft, open, and spongy
showing large air-spaces. 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 sub-
merged leaves are modified into bladders which trap insect
larvee and smaller Crustacez (see Fig. 481).

THE LEAF 249

B. CoNvERGENT.—This type is mostly seen in monocotyle-
dons. In Phormium tenax or New Zealand Flax 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 sides of the
upper part being fused and functioning as a lamina.

é.
Af
a e\ sé rs
asserere

Fic. 167.—Transverse section through portion of dorsiventral leaf blade of
horehound (Marrubium vulgare). Upper epidermis devoid of stomata (up. ¢p.);
lower epidermis which possesses stomata (/. ef.); palisade parenchyma (pal.);
spongy parenchyma (sp. p.); xylem (x) and phloem (fh) regions of fibrovascular
’ tissue of stronger vein; long-pointed non-glandular trichome (f); branched tri-
chomes (ft!, ft2, ft); several types of glandular trichomes (gt', gt?, gt*, gt*).

C. Centric.—This type of lamina is semi-centric to centric
in shape. It is nearly always associated with Xerophytes.
Centric laminz are produced gradually by an encroachment of
the under on the upper surface, and the swelling of the whole.
There are two kinds of centric laminz, the succulent-centric and

the xerophytic centric.

Fl

ya ge PHARMACEUTICAL BOTANY

SuccuLENT.—In a completely centric leaf of the succulent
kind, like that of Sedum murale the difference between the upper
and lower surface is lost. Stomata are found scattered over the
entire epidermis. The bundles are arranged in a circle, the mid-
rib being in the center. A great deal of mucilage is found stored
in the central cells.

XEROPHYTIC.—In a typical Xerophytic Centric leaf, like that
of the Pine (see Figs. 168 and 169) or Sansevierta cylindrica, the

Fic. 168.—Transverse section of the xerophytic-centric leaf of the Austrian
Pine (Pinus nigra), magnified. ep, epidermis; s, deeply sunken stomata; Af, hypo-
dermal fibers; 7p, infolded parenchyma; 0.r., schizogenous oil reservoirs; f, fibers;
n, bundle sheath (endodermis) surrounding the stele in which will be noted two
vascular bundles with xylem (x) directed toward the flat face of the leaf; ph, phloem
of vascular bundle; mr, medullary ray: scl, sclerenchymatous cells; p, small celled
parenchyma of pith.

epidermis shows a thick cuticle; the stomata are sunken in
cavities of the epidermis; the epidermis and leaf tissue are
strengthened by scleroid bands in the outer portion of the
centric mesophyll.

D. BrractaL.—Leaves with laminz which stand edge on in
relation to the sun’s rays. The best illustrations are seen among
dicotyledons, such as Eucalyptus, Callistemnon, and other genera of

en ae ee a a

Pe ee ee

THE LEAF 251

the Myrtaceae. The leaflets of Cassia Senna and Cassia angustifolia
which constitute the drug Senna, show a marked bifacial struc-
ture. Both surfaces are similar, having stomata about equal in
number. ‘The mesophyll (chlorenchyma) is differentiated into a
central spongy parenchyma containing bundles, and a zone of
palisade cells on either side facing the epidermises (see F ig. 170).

STRUCTURE AND DEVELOPMENT OF STOMATA.—Stomata or
stomates 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

Fic. 169.—Portion of a transverse section of the White Pine leaf showing
xerophytic-centric structure. ¢p, Epidermis; s, stoma; gc, guard cells; s’, sub-
stomal air-space; h, hypodermis, consisting of strengthening fibers which form a
scleroid band beneath epidermis; f, infolded parenchyma cell of mesophyll.
Greatly enlarged.

communication between the intercellular-air-space (respiratory
cavity) beneath them and the exterior. The slit-like openings,
taken with the guard cells, constitutes what is known as the
stomatal apparatus. The stomatal apparatus is frequently called
in practice, ‘‘stoma” or “‘stomate.”

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, Peonia, Ferns, etc.,
are very similar to the other epidermal cells, but in a large num-
ber 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 mostly two in number, one larger
than the other and arranged parallel to the guard cells of the

252 PHARMACEUTICAL BOTANY

stoma; in Coca a similar arrangement occurs but the cells are more
even in size, nevertheless they lack the characteristic papille
found on the other epidermal cells; in Pilocarpus 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.

ep pal
Fic. 170.—Photomicrograph of a transverse section of a bifacial leaf of Eucalyp-
tus globulus showing epidermis (ep), palisade parenchyma (pal), toward both
surfaces, spongy parenchyma (sp), vein (v), and oil reservoir (o0.r.) lined with
secretory epithelium. (Highly magnified.)

On all dorsiventral leaves, the stomata arise more abundantly 4
on the lower epidermis, less abundantly on the upper. Excep-
tions to this rule are due to the peculiar readaptation of the leaf to
its surroundings. 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 phys-
iological lower surface.

THE LEAF 253

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 the 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 numerous; if they dwell in dry soil in hot sur-
roundings and dense light they are very small and frequently
sunken. If the plants are Xerophytic and the leaves dorsiven-
tral, the stomata are quite abundant, small, with narrow slit,
and depressed below the level of the epidermis.

There are five types of stomatal development, viz.:

First Typr.—Each primitive epidermal cell (or the majority,
or only certain ones of the epidermis) at the close of the dermato-
gen 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, quad-
rangular, 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 length-
wise 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 Sguzll, Hyacinth,
Daffodil, Sambucus, Silene, etc.

SEconp Type.—After the cutting off of the stomal mother-
cell, there are ¢ut off, on either side, portions of neighboring
epidermal cells which form subsidiary cells to the stoma. This
condition is seen in Gramineae, Cyperacea, Juncacee, in various
species of Aloe, Musa and Proteacea.

Turp Type.—Instead of two parallel subsidiary cells, four
are cut off, as in Heliconia, in species of Tradescantia, Araucaria, or
four to five, as in Ficus elastica, or four to five or more, as in the
Conifere and Cycads.

Fourtu Type.—Instead of only four subsidiary cells, each of
these again subdivides by parallel walls, more rarely by radial

254 PHARMACEUTICAL BOTANY

walls, into eight radiating subsidiary cells, as in Maranta bicolor,
Pothos argyrea, some of Proteacea, etc.

FrrtH 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 cells. This mode of development is seen in the

Fic. 171.—Types of stomatal apparatuses and neighboring cells from different
sources. In A, a portion of the lower epidermis of Easter Lily leaf. The stomatal
apparatus is surrounded by neighboring cells that are similar to other epidermal
cells adjacent to them; in B, lower epidermis of Senna leaflet, note the two neighbor-
ing cells parallel to the guard cells, one being larger than the other; C, lower
epidermis of Coca leaf showing two neighboring cells, parallel to the guard cells
but nearly equal in size as well as papillated regular epidermal cells; D, lower
epidermis of Pilocarpus showing rounded stomatal apparat@s and four crescent-
shaped neighboring cells; E, lower epidermis of Uva Ursi, showing eight neighbor-
ing cells arranged radiately around stomatal apparatus; F, lower epidermis of
Stramonium,

Labiate, Papilionacee, Crucifere, Solanacea, Crassulacea, Cuactacea,
and Begoniacee, also in a number of ferns.

HisTo.ocic DirFERENCES BETWEEN LEAVES OF DicOTYLEDONS
anD Monocotytepons.—The following may be cited as broad
comparative histologic differences between Dicotyl and Mono-
cotyl leaves:

THE LEAF 255

Dicotyi LEAVEs

. Epidermal! cells usually iso-diametric
or sinuous in surface sections.

. The stomata are on the whole more
numerous but smaller.

. Non-glandular and glandular hairs
frequent on upper but more fre-
quent on lower surface, or both.

. Leaf glands which excrete varied
products are rather abundant.

. Water stomata over the upper sur-
face, more rarely over the lower
surface, are frequent, especially
along margins of leaves.

. Palisade and spongy parenchyma
in dicotyledons are more distinct
and palisade parenchyma is
denser.

. The vascular bundles, in their in-
trinsic elements, are more indur-
ated but the accessory fibrous
sheath is feebly developed.

. A greater variety of accessory prod-
ucts of assimilation are devel-

oped.

Monocotyt LEAVES

1. Epidermal cells usually elongate

with rectilinear vertical walls, in
surface sections.

. Stomata larger.

. Hairs rare in Monocotyls.

. Leaf glands rare and only seen as a

rule on the sepals.

. Water stomata absent or very rare.

Present in some Araceae.

. Palisade and spongy parenchyma

are less distinct and dense.

. The vascular bundles, in their in-

trinsic elements, are less indu-
rated. The fibrous sheath is
strongly developed.

. A comparatively small variety of

accessory products of assimilation
are developed.

CHAPTER XI

THE INFLORESCENCE

INFLORESCENCE OF ANTHOTAXY.—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.

Indeterminate, racemose, ascending, or centripetal inflorescence is
that form in which the terminal bud of the flower cluster con-
tinues to develop and increase the length of the stem indefinitely.
As the stem elongates new flower buds arise behind the terminal
bud in the axils of reduced foliage leaves called bracts, so that the
oldest bud is at the base of the flower cluster and the youngest one
is nearest the apex. Example: Digitalis.

Determinate, cymose, descending, or centrifugal inflorescence is
that form in which the opening of the terminal bud as a flower
determines or completes the growth of the stem of the flower
cluster. Example: Ricinus communis.

Mixed inflorescence is a combination of the other two forms.
Example: Horse Chestnut.

The flower stalk is known as the peduncle. Its prolongation
beyond the point where flowers or branches begin to appear is
called 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 the peduncle or its extension,
the rachis, are termed bracts. These vary much the same as leaf
forms, are described in a similar manner, 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 generally green,
but sometimes petaloid, as in the Dogwoods. The modified

leaves found on pedicels are called bracteolar leaves or bracteoles.
256

a a oo eee ee

THE INFLORESCENCE 257

The Spathe is a large bract more or less enveloping the
inflorescence and often colored, as in the Calla, or membranous,
as in the Daffodil.

INDETERMINATE INFLORESCENCES.—In the indeterminate or
centripetal anthotaxy, the flowers are either produced from base

(Drawing by Canis.)

J AESCULUS HiePocASTAN UY

1, indeterminate; 2, determinate; 3, mixed.

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rachis or the flowers open from margin tocenter. Indeterminate
inflorescences thus fall into two groups, viz., the elongated or
ascending group and the flat-topped or centripetal group. The
principal forms of indeterminate inflorescence are:

258 PHARMACEUTICAL BOTANY

SOLITARY INDETERMINATE.—The simplest form of inflores-
cence 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 a lengthened
rachis, the first flower to open being at the base of the rachis and
the last to open at the summit. Examples: Convallaria, Cimici-
fuga, and Wild Cherry.

ft WY

Fic. 173.—Types of indeterminate or centripetal inflorescence. a, raceme:
6, spike; c, panicle: d, corymb; e, umbel; f, head (capitulum) of Compositz type; g,
head of clover type. (Gager.)

Corymp, 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: Hawthorn.

UmseL, which resembles the raceme, but has a very short
axis, and the nearly equal pedicels radiate from it like the rays
of an umbrella. When the pedicels of an umbel put forth
branches, these branches are termed secondary pedicels. Such,
then, bear flowers at their tips and make up with other parts of
the inflorescence a Compounp UmBet. Many examples of this
mode of inflorescence are seen in the family Umbellifera, as
indicated by the name, including Anise, Caraway, Fennel and
other drug-yielding official plants.

THE INFLORESCENCE 259

A Spike is a cluster of flowers, sessile or nearly so, borne in the
axils of bracts on an elongated rachis. The Mullein and com-
mon Plantain afford good illustrations. If branches arise from
the rachis of a spike and bear flowers, a COMPOUND SPIKE results.
In this type of inflorescence which is seen in Wheat, Rye, Barley
and many other members of the [iG =a aaa Se
Grass Family, the branchiscalled | — /¢@@t/
the rachilla and the secondary
inflorescences are termed sprkelets.

The CaTKIn or AMENT resem-
bles the Spike, but differs in that it
is deciduous and usually bears
only one kind of flowers, stami-
nate or pistillate, as in the Oak,
Hazel, Willow, Comptonia, etc.

The Heap 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
spike of unisexual flowers with
overlapping scales concealing the
flowers, as the inflorescence of the
Hop.

‘ ° Fic. 174.—Photomicrograph of
The Spaprx is a thick, fleshy longitudinal section through a stami-

rachis with flowers closely sessile pate catkin of Comptonia asplenifolia
or embedded on it. It is usually x10, showing catkin axis (ax),

rhore he Wed elon oy a sensi: St, ore ©) and bract ©).

or sheathing bract. Examples: Calla, Calamus, Arum triphyllum.
The raceme, corymb, and umbel, are frequently com-
pounded through the branching of their pedicels. The com-
pound raceme, or raceme with branched pedicels, is called a
Panicte. Examples: Brayera, Yucca and the Oat.
DETERMINATE INFLORESCENCES.—Determinate Anthotaxy is one
in which the first flower that opens is the terminal one on the axis,
the others appearing in succession from apex to base or from

260 PHARMACEUTICAL BOTANY

center to margin. If the floral axis is elongated, the order of
flower development is descending, or from apex to base; if more or
less ‘‘flat topped,” the order is centrifugal, or from center to
margin. ‘The principal varieties are:

The Soxirary DETERMINATE, in which there is a single flower
borne on the scape, as in the Anemone, or Windflower, the Tulip,
and Hydrastis.

The Cyme, a flower cluster resembling a corymb, except that
the flower buds develop and open from center to circumference.

Oo Pp © ,

WY Wy

es

VY
be ef

F G H
Fic, 175.—Cymose inflorescences. F, A terminal flower; G, a simple cyme; H,
acompound cyme. (From Hamaker.)

The simplest kind of cyme is seen in the flower cluster of the
Linden tree, where a peduncle (arising from a large bract) is
elongated into a rachis which ends in a flower bud. At the
junction of the rachis with the peduncle, 2 bracteoles arise. In
the axil of each of these bracts a bud is formed which elongates in
each instance into a pedicel bearing a flower bud at its end.
This type of cyme is called a SimptE Cyme. If each of the
pedicels give rise to a pair of bracts from the axils of which
secondary pedicels arise bearing flowers, a type of compound cyme
arises called a DicuastaL Cymer.

A Scorpiomw Cymer imitates a raceme, having the flowers
pedicelled and arranged along alternate sides of a lengthened
axis. These open in right-left hand fashion from summit to
base. Example: Sundew.

THE INFLORESCENCE 261

A GLOMERULE is a cymose inflorescence of any sort which is
condensed into a head, as the so-called head of the Dogwoods
(Cornus).

A VERTICILLASTER Is a compact, cymose flower cluster which
resembles a whorl, but really consists of two glomerules situated

Fic. 176.—Flowering-branch of a Dogwood showing glomerules. Note the
four notched petaloid bracts subtending each glomerule or determinate head of
flowers.

in the axils of opposite leaves. Clusters of this kind are seen in
Catnip, Horehound, Peppermint and other plants of the Labvate.

Mrxep INFLORESCENCES are those in which the determinate
and indeterminate plans are combined, and illustrations of this
are of frequent occurrence.

A Tuyrsus is a mixed inflorescence more or less pyramidal or
oblong in shape consisting of a panicle of cymes, as in the Privet,
Lilac, Grape or Horsechestnut. Racemes of cymes and corymbs
of cymes are other examples of mixed inflorescences.

CHAPTER XII

THE FLOWER

The FLower is a shoot which has undergone a series of
changes so as to serve as a means for sexual reproduction. It
represents a highly modified stem or branch which bears
appendages.

A TypicAL OR CoMPLETE FLOWER possesses four circles or
whorls of floral leaves arranged upon a more or less shortened
and frequently swollen upper end of a stem axis called a RECEP-
TACLE, Torus or THALAMus. These whorls passing from periph-
ery toward the center are: CALYx, composed of parts called
sepals; CoROLLA, composed of parts termed petals; ANDRECIUM,
composed of parts called stamens or microsporophylls; and
GyNncciuM, composed of one or more parts termed carpels, pis-
tils, or megasporophylls. In a complete flower the calyx and
corolla constitute the perianth or floral envelope.

The stem bearing the flower is called the flower stem or
PEDUNCLE.

The stamens and carpels constitute the EssENTIAL ORGANS,
and a flower is said to be Perrect when these are present and
functional. An ImpEerFecT FLower shows one set of essential
organs wanting. The calyx, corolla and receptacle constitute
the Accessory Orcans of the flower. 3

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 RecuULAR FLOWER possesses parts of each whorl of the
same shape and size, as the flower of the buttercup. This kind of
flower possesses radial symmetry and is also called for that
reason ACTINOMORPHIC. :

An IRREGULAR FLoweEr is one in which the parts of one or
more whorls differ in shape and size. If it possesses bilateral

symmetry, as the flower of the violet or sweet pea, it is also called
262

THE FLOWER 263

ZYGOMORPHIC. _Irregularity may result either from variation in
form or size of the parts of a whorl, from appendaging, as in the
calyx of the Larkspur where one of the 5 sepals bears a long spur,
or from abortion.

A SYMMETRICAL FLOWER is one in which the parts of each
whorl are of the same number, or multiples of the same number,
as the lily or rose.

Fic. 177.—Diagrams of floral structures. A, Shows the relations of the floral
parts in a hypogynous flower; B, the same in a perigynous flower; C, the same in
an epigynous flower; D, a stamen; F, a simple pistil in longitudinal section; F,
the same in cross-section; G, transitional forms between true petals (left) and true
stamens (right); H, slight union of two carpels to form a compound pistil; J and 7,
union of carpels more complete; K and L, cross-sections of compound pistils, of
three carpels. In B: a, stamen; 6, petal; c, sepal; d, pistil; e, receptacle; f, pedicel.
In D: a, anther cell; 4, connective; ¢, filament. In £: a, stigma; 4, style; c, ovules;
d, ovary. (From Hamaker.)

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 trans-
formation of stamens (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 one or more pistils are present and stamens wanting, the
flower is called PistrLLATE, or female; if it possesses one or more
stamens but no pistil, it is described as SraMINATE, or male; if
both are absent, NEUTRAL, as marginal flowers of Viburnum

264 PHARMACEUTICAL BOTANY

and Hydrangea. When staminate and pistillate flowers are
borne separately they are called diclinous flowers. Flowers con-
taining stamens and pistils are monoclinous. Some plants, as the
Begonias, Corn, and Castor oil, bear both staminate and pistillate
flowers on the same individual, and are called Monecious.
When the staminate and pistillate flowers are borne on different
plants of the same species, the plants are termed Diecious, as the
Sassafras and Willow. When staminate, pistillate and hermaph-
rodite flowers are all borne on one plant, as on certain species of
Maple and Buckwheat, they are polygamous.

A Hypocyrnous FLow:r is one in which the sepals, petals and
stamens are attached near the base of the receptacle while the
pistil or the carpels are arranged over the upper portion of
the receptacle, as in the magnolia and strawberry. ‘The recep-
tacle of this kind of flower is generally conical, dome-shaped or
elongated like a spire. The ovaries of this type of a flower are
superior.

A PericyNnous FLower is one in which the sepals, petals and
stamens are inserted on the edge or summit of a saucer or cup-
shaped receptacle while the pistil or carpels are inserted at the
bottom of the receptacle, as in the flowers of the cherry, rose and
almond. The ovary of this type of flower is superzor.

An Epicynous FLowsr is one like that of the apple or quince
where the receptacle is greatly hollowed out in vase-shaped
manner, its summit overarching the top of the ovary and bearing
the sepals, petals and stamens above the ovary. The ovary of
such a flower is inferior.

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 under vernation, the fol-
lowing are important.

Valvate Prefloration, in which the margins of the parts meet but
do not overlap. 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 Althza.

THE FLOWER 265

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 one.

Contorted, where one margin is invariably exterior and the
other interior, giving the bud a twisted appearance, as in the
Oleander and Phlox.

CoNNATION AND ADNATION.—In the development of the
flowers of primitive species of flowering plants, the parts of each
whorl are disjoined or separate from each other. In many
higher types, however, the parts of the same whorl frequently

Fic. 178. Fic. 179.
Fic. 178.—Flower of the Ash (Fraxinus); both calyx and corolla absent (achlam-

ydeous).
Fic. 179.—Flower of the Goosefoot (Chenopodium) with a calyx but no corolla
(monochlamydeous). (After Small.)

become partly or completely united laterally. This condition is
termed connation, coalescence, cohesion or syngenests. Illustrations of
this may be seen in Belladonna, Stramonium, and Uva Ursi
flowers, where the petals have joined laterally to form gamo-
petalous corollas. When the one or more parts of different
whorls are united, as of stamens with petals (Rhamnus) or stamens
with carpels (Apocynum), the union is called adnation or adhesion.
Tue Receptacite.—The Receptacle, Torus, or Thalamus is a
more or less shortened axis or reduced branch which bears the
floral leaves. It is usually flat or convex, but may be conical
and fleshy as in the Strawberry, concave as in the Rose and Fig,
or show a disc-like modification, as in the Orange. Like other
stems it has nodes and internodes. The internodes of the
receptacle in many species lengthen and separate various whorls.
When the lengthening of the internode occurs between calyx and

266 PHARMACEUTICAL BOTANY

corolla, as in Lychnis, the structure resulting is called an ANTHO-
pPHORE; if between corolla and androecium, as in Passiflora, a
Gonorpuorg; if between androecium and gyncecium, as in Geum,
a GyNnopuorE. In the flowers of the Umbellifere, the receptacle
elongates between the carpels producing the structure called a
CARPOPHORE.

Tue Pericone.—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. :

Fic. 180. Fic. 181.
Fic. 180,—Chorisepalous and hypogynous calyx of the Pimpernel (Anagallis).
Fic. 181.—Gamosepalous and bilabiate calyx of the Deadnettle (Lamium).
Fic. 182.—Urceolate calyx of the Henbane (Hyoscyamus). (After Small.)

Tur Caryx.—The Calyx is the outer whorl of modified
leaves. Its parts are called Sepats. These may be distinct or
separated from one another when the calyx is called CHo-
RISEPALOUS, APOSEPALOUS or PoLysEpALous. If the sepals are
more or less united at their margins, the calyx is called GAmMo-
SEPALOUS or SYNSEPALOUS.

The sepals are usually green—foliaceous or leaf-like—but
may be brilliantly colored, hence the term petaloid (like the
petals) is applied. Examples of flowers having a petaloid
calyx: Tulip, Larkspur, Columbine and Aconite.

In the Daisy Family, etc., the calyx is modified in the form of
teeth, bristles, hairs or awns and is then called the PAPPus.

_ Ina gamosepalous calyx, when the union of sepals is incomplete,
the united portion is called the tube, the free portion, the /imb, the
orifice of the tube, the throat.

THE FLOWER 267

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

Fic. 183. “Fic. 184. Fic. 185.
Fic. 183.—Campanulate calyx, surrounded by an epicalyx, of the Rose of
Sharon (Hibiscus).
Fic. 184.—Perigynous and chorisepalous calyx (cf) of Rose with concave
receptacular tube (ct) with stamens (e) inserted perigynously on its edge.
Fic. 185.—Persistente pigynous calyx (c) of the Quince (Cydonia), showing the
receptacular tube completely enveloping the matured carpels. (After Small.)

Fic. 186. Fic. 187. Fic. 188.

Fic. 186.—Caducous hairy calyx of the Poppy (Papaver).
Fic. 187.—Globose persistent calyx of Winter Cherry (Physalis).
Fic. 188.—Perigynous chorisepalous calyx of Indian Cress (Tropa@olum) with

spurred receptacular cup. (After Small.)

or urn-shape; hypocrateriform, or salver-shape; bilabiate, or two-
lipped; corresponding to the different forms of corolla, under
which examples illustrating each will be given.

The calyx usually remains after the corolla and stamens have
fallen, sometimes even until the fruit matures, as in the Night-

268 PHARMACEUTICAL BOTANY

shade family; in either case it is said to be persistent. fit falls with
the corolla and stamens, it is deciduous, and if when the flower
expands, caducous, as in the calyx of Poppy and May-apple. If
the calyx falls very early, it is called fugactous.

SEPALINE NECTARIES AND 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 the Crucifere or Mustard Family, we occasionally see a

\\
“
15
\\\‘nim of receptacle

“fleshy receptacle

{7
stamens’

|

Fic, 189.—Strawberry (Fragaria chiloensis). Median lengthwise section of flower.
(After Robbins.)

slight pouching of the two lateral sepals. These act as nectar
pouches for the nectar secreted by nectar glands on 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 the Larkspurs, the posterior sepal
forms an elongated spur into which pass the two spurred nectar-
iferous petals. In Aconitum the same sepal, instead of being
spurred, forms an enlarged hood-like body called the ga/ea or
hood which arches over the flower like a helmet; into this pass the

THE FLOWER 269

two hammer-shaped nectariferous petals. In the Wild Colum-
bine (Aquilegia) each of the 5 sepals develops a spur.

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 supple-
mentary whorl of green sepal-like lobes. This is outside of the
normal calyx, and is called the Epicatyx. 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.

SEPALINE Posirion.—The most simple and primitive position
for the sepals in relation to the floral parts is Hypocynous, 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, Leguminosae, etc., the
floral axis forms a saucer-like, transverse expansion which pushes
out the sepals, petals and stamens on its edge. ‘Thus originates
the Pericynous insertion of the sepals. In not a few higher
Rosacea, Saxifragacea, 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.

Tue Coroiia.—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 Perats,

and when the calyx closely resembles the corolla in structure and

270 PHARMACEUTICAL BOTANY

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 saccharine secretions serve to attract pollen-carry-
ing insects.

When the petals are not united with each other, the corolla is
said to be CHORIPETALOUS, APOPETALOUS or POLYPETALOUS.

ca

Fic. 190.—Sour Cherry (Prunus cerasus). Median longitudinal section of flower.

Note the perigynous insertion of the sepals, petals and stamens. The ovary is
superior. (After Robbins.)

When more or less united into a tubular structure, it is Gamo- me
PETALOUS, often called SYNPETALOUS.

ForMS OF THE COROLLA AND PERIANTH.—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 e
plants belonging to the family Cruciferz. :

The Papilionaceous corolla is so called because of a taseied
resemblance to a butterfly. The irregularity in this form i is —

THE FLOWER 271

striking, and the petals bear special names: the largest one is the
vexillum, or standard; the two beneath it the alg, or wings; the
two anterior, the carina or keel. Examples: Locust, Pea, and
Clover. [Fig. 193 (9).]

Orchidaceous flowers are of peculiar irregularity, combining
calyxand corolla. The petal in frontofstamen and stigma, which
differs from the others in form and secretes nectar, is called the
Labellum (Fig. 192). Examples: Cypripedium and other Orchids.

Fic. 191.—Apple (Malus sylvestris). Median lengthwise section of flower.
Note the epigynous insertion of sepals, petals and stamens. The ovary is inferior.
(After Robbins.)

When calyx and corolla each consist of three parts closely
resembling 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 Composite. 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 Labiate, while in the family Leguminosa this arrange-

272

ment is sometimes reversed.

PHARMACEUTICAL BOTANY

The corolla may be either ringent,

or gaping, as in Rosemary and Sage, or personate, when the throat
is nearly closed by a projection of the lower lip, as in Snapdragon

Fic. 192.—Flower of an orchid

(Orchis militaris). A, a, bract; 4,
ovary; c, the outer, and d, the two
anterior perianth leaves; ¢, thé label-
lum or lip with spur, f, (nectary);
g, the gynostemium the union of
stamens and pistil. B, flower after
removal of all the perianth leaves
with the exception of the upper part
of the labellum; A, stigma; /, rostel-
lum, a beak-like extension from the
upper edge of the stigma; k, tooth-
like prolongation of rostellum; m,
anther; n, connective; 0, pollinium;
q, Viscid disk; p, staminodium, C,a
pollintum removed and enlarged;
r, its stalk (caudicle); s, pollen mass.
D, ovary in cross section. (After
Strasburger, Macmillan Co., publishers.)

apex.

and Linaria.

Rotate or 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.

Cratertform or Saucer-shaped, like
the last except that the margin is
turned upward or cupped. Exam-
ple: Kalmia latifolia (Mt. Laurel).

Hypocrateriform or Salver-shaped
(more correctly, hypocraterimor-
phous), when the tube is long and
slender, as in Phlox or Trailing
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

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,

Chimaphila and Gaultheria.

THE FLOWER 273

Caryophyllaceous, when the corolla consists of five petals, each
with a long slender claw expanding abruptly at its summit into a

Fic. 193.—Illustrating various forms of the corolla. 1, Personate bilabiate
corolla of Linaria; 2, cruciform corolla of Rocket; 3, companulate corolla of
Harebell; 4, infundibuliform corolla of Bindweed; 5, ringent bilabiate corolla of
Rosemary; 6, spurred calyx of Larkspur; 7, ligulate corolla of Chrysanthemum;
8, rotate corolla of Pimpernel; 9, papilionaceous corolla of Irish Broom; 10, urceo-
late corolla of Heath.

broad limb. Examples: Carnation and other members of the
Pink family.

274 PHARMACEUTICAL BOTANY

APPENDAGES OF THE PERIGONE.—In addition to the spurs
already discussed, the following outgrowths of the perianth may
also be encountered:

The AuricLe, an ear-shaped appendage occurring on the
calyx of Lobelia. This term is also applied to similarly shaped
lobes of foliage leaves.

The Awn is a sharp, rigid outgrowth from the tip or dorsal
surface of a perigone lobe.

The Horn or Cornua is a hollow, horn-like appendage.

The Caupa is a long, drawn out, tail-like appendage, as
seen in the petals of the Cacao plant.

The Paxate is a sac-like upgrowth of the lower lip of a
personate, bilabiate corolla.

The Keet is a ridge resembling the keel of a boat.

‘The Corona or Crown isa circular upgrowth from the throat
of the perianth, as seen in Narcissus. It may represent the up-
growth and coalescence of stipules of perianth segments or stamens.

Tue ANDRGCIUM OR STAMEN SystEM.—The andrecium is the
single or double whorl of male organs situated within or above
the corolla. It is composed of SraMENS or MICROSPOROPHYLLS.

A complete stamen (Fig. 177D) consists of a more or less
slender stalk portion called a FILAMENT and a terminal appendage
called the ANTHER or Microsorus. 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 pass-
ing from these to those of more evolved families there occurs a

2
f
s
,

THE FLOWER 275

gradual reduction from many to ten, as in Caryophyllacea,
Leguminose and some Aceracee, these being in two circles. In
Malvaceae, Umbellifere and other Apopetalous families as well as
many Synpetalz, the number five is typical. But in Scroph-
ulariacee, while five are developed and fertile in Verbascum, four
with a fifth staminode (sterile stamen) are found in the allied
genus Ceélsia. 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 Scrophulariacee 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 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:

Hypocynous, when inserted upon the receptacle below the
base of the pistil (see Fig. 177A).

PEericyNnous, when inserted on the calyx or corolla above the
base of and lateral to the pistil (see Fig. 1778).

Epicynous, when inserted above the ovary (see Fig. 177C).

GyNANbRows, when inserted upon the pistil, as in Orchids and
Aristolochia (see Fig. 194 No. 8).

PROPORTIONS OF THE STAMENS.—The stamens may be of
equal length; unequal, or of different length.

Diwynamous, when there are two pairs, one longer than the
other. Examples: Snapdragons, Digitalis, Mints. [Fig. 194
4).
geen three pairs, two of the same length, the
third shorter. Example: Mustard. [Fig. 194 (6).]

CoNNATION OF STAMENS.—Terms denoting connection
between stamens are:

MonapeE.pHous (in one brotherhood), coalescence of the
filaments into a tube. Examples: Lobelia and Malvaceae [Fig. 194
(5)].

DiapELpHous (in two brotherhoods), coalescence into two
sets. Example: Glycyrrhiza. (Fig. 194 (2).]

276 PHARMACEUTICAL BOTANY

12 13 a

Fic. 194.—The Stamen System. 1, Stamen of Hepatica showing adnate
anther, the 2 lobes separated by connective; 2, diadelphous stamens of a leguminous
plant; 3, polyadelphous stamens of Orange with calyx at base; 4, flower of Teucrium
showing didynamous stamens; 5, monadelphous stamens of Malva; 6, tetradyna-
mous stamens of a Crucifer; 7, flower of Hypericum cut vertically to show triadelphous
stamens; 8, androecium and gyneecium of Aristolochia showing gynandrous stamens;
9, stamen of a Labiate showing branched connective with one anther-sac aborted
and absent; 10, adnate anther of Campanula; 11, stamen of Malva with confluent
anther, which is 1-celled; 12, stamen of Lily with versatile anther; 13, stamen of
False Rue Anemone with innate anther; 14, stamens of Sunflower showing syngene-
sious anthers.

: TRIADELPHOUS, with filaments united into three sets. Exam-
ple: St. John’s Wort. [Fig. 194 (7).]

THE FLOWER 277

PoLYADELPHOUS, when there are several sets or branched
bundles. Example: Orange.!

SYNGENEsIOUS, when the anthers cohere. Example:
Composite.

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 Sassafras, Cucumber and Golden Club; greenish-
yellow, yellow to red, in Maples; yellow-pink to pink and pink-
ish-red, in some Mallows; in Azalea amena the filaments are
crimson-purple and the anthers, purple-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 a dilated base as in the Harebell, three-toothed as in
Garlic, appendiculate, when it bears an appendage as in
Chetostoma, Alyssum, etc. The filament is covered with a protec-
tive 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 (M1cROSPOROPHYLL) bears a special development or
appendage, as a rule, on its extremity which is the ANTHER or
Microsorus. This consists, fundamentaily, of a median pro-
longation 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, con-
sists 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

1In the Orange flower each set of stamens consists of a filament bearing in its
distal region phalanges of stamens or staminal leaflets.

278 PHARMACEUTICAL BOTANY

time of dehiscence. Again, in various species of Laurace@, 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.

. - ex
Connective

a ed
5.

ms —_
Stomium —

Pollen Mother ae
Cell a Gate, he Ae

Tapetum= ~

Fic. 195.—Cross section of immature anther of Johnson grass (Holcus halepensis
L.). The pollenm other cells are contained within four microsporangia. x,
exothecium. (Mottier.)

Externally the mature anther is bounded by an ExoTHECIUM
or epidermis, 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
EnpoTHECIUM. 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 ExorHecrum, 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

THE FLOWER 279

by developing PoLLEN or microspore cells. Filling the cavities
of the four sporangia are the mature pollen grains. ‘The con-
nective shows in or near its center a vascular bundle with xylem
uppermost and phloem downward, surrounded by thin-walled
cellular tissue, from which the indusial and sporangial substance
has matured by extension.

ANTHER DeEHISCENCE.—This is the breaking open of the
anther to discharge the pollen.

When fully ripe the dividing partition between each pair of
sporangia usually becomes thinned, flattened and ultimately
breaks down, 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 /ongitudinal anther dehiscence on either side of the
anther sacs. In the division Solanee of the family Solanacee which
includes Belladonna, in some of the Evicacee 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 Lauracee and Berberidace@, the anthers
dehisce by recurved valves (see Fig. 202). This is called valvular
dehiscence.

Moreover, in Malvacea, 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 par-
tition becomes absorbed, thus becoming a one-celled anther
with transverse dehiscence in its mature state [see Fig. 194 (11)].

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 filament develops later. When such a young sorus
or anther is cut across and examined microscopically, it shows a
mass of nearly similar cellular tissue in which the first observable
changes are the following:

The surface dermatogen cells become somewhat flattened and
regular 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

280 PHARMACEUTICAL BOTANY

vascular bundle in the connective. Then, along four longitudinal
tracts, rows of cells remain undivided or only divided slowly as
they increase in size and around them cells divide and redivide
to form the future endothecial 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 protoplasmic 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 chromatin substance takes place in

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Fic. 196.—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. Copy-
right by the American Book Company, Publishers.)

these cells. Thus originate fetrads (groups of four) of spore
daughter-cells inside the 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 irreg-
ular layer around the maturing pollen cells. When the anther
is finally ripe the partition between each pair of microsporangia
becomes narrowed, flattened and ruptured and thus numerous

THE FLOWER

microspores or pollen grains fill two cavities, one on either side
of the connective. The microspores or pollen grains at first show

m Typha latifolia (A),

Fic. 197.—Various forms of pollen grains. Pollen fro :
Zea mays (B), Ambrosia elatior (C), Lilium philadelphicum (D), Pinus (E), Ranunculus

bulbosus (F), Carpinus caroliniana (G), Althea rosea (H), Ocnothera biennis (I). All
highly magnified. Drawing by Hogstad. some

only a thin, clear, cellulose layer, but from this, by a differentia-
tion of the exterior film, the exospore layer or extne becomes cut off.

282 PHARMACEUTICAL BOTANY

This becomes cuticular. The cellulose inner layer (endospore or
intine), 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 pollen
tube. These are called germ pores.

ATTACHMENT OF ANTHER.—The attachment of the anther to
the filament 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.

Fic. 198. Fic. 199.
Fic. 198.—Orchid Pollinia (~), composed of masses of pollen grains, with
caudicle (c), and disc or retinaculum (r). (After Small.)
Fic. 199.—Upper part of Orchid flower, showing the retinacula (a) in place
within the vostellum or sterile stigma. (After Small.)

Versatile, when the anther is attached near its center to the top
of the filament, so that it swings freely. The adnate and versatile
types are introrse when they face inward, extrorse when they face
outward.

Po._en.—The pollen grains or microspores vary in form for
different species and varieties and while they are averagely
constant for these, nevertheless, many exceptions have been
recorded. The following 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 Evening Primrose (CEnothera) and Clove.

Echinate, as in the Malvaceae.

Spherical, as in Geranium, Cinnamon and Sassafras.

THE FLOWER 283

Lens-shaped, as in the Lily.

Spinose, as in the Composita.

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,”’ and an 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.

Po.unta.—These are agglutinated pollen
masses which are common to the Orchidaceae and
Asclepiadacea.

Hay Fever Potiens.—The pollen of many
plants, notably certain species of Composite,
Chenopodiaceae, Graminee, Amaranthacee, Fagacee,
Ulmacea, Salicacee and Juglandacee, has been
shown to be responsible for “Rose Cold” and
“Hay Fever.” The chief hay-fever pollens are
those of the Ragweeds (Ambrosta spp.), Pigweeds 5, 200—Pol-
(Amaranthus spp.), Docks (Rumex), Wormwoods jinia of a Milkweed
(Artemisia), Russian Thistle (Salsola), various fapiecinde ¥ svi
grasses and cereals including timothy, corn, fe = ar vie .
wheat, oats and Buffalo grass, and a number of ( After Small.)
trees including the oaks, elms, poplars, black
walnut, and red cedar. From the investigations of Walker and
others, it appears that the symptoms of “Rose Cold” and “‘Hay
Fever” which are seasonable forms of bronchial asthma are
induced by the sensitivity of individuals to the proteins of certain
pollens. At the present time pollen extracts, also called pollen
allergens, pollen antigens and pollen atophans, are manufactured
from various pollens to be used in the diagnosis, prevention and
relief of this disease.

Tue Gynacrum or Pst 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 megasporo-
phyll is a female organ of reproduction. In the Spruce, Pine,
etc., it consists of an open leaf or scale which bears but does not

j
a
‘

3

284 PHARMACEUTICAL BOTANY

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

Fic. 201.

Fic. 204. Fic. 205.

Fic. 203.

Forms or Gyncac1uM

Fic. 201.—Monocarpellary pistil of Broom (Scoparius). 0, ovary; s, style;
t, stigma. Fic. 202.—Barberry (Berberis). 0, one-celled or unilocular ovary;
ov, ovules attached to marginal placenta (f); st, sessile stigma. One stamen dehisc-
ing by recurved valves is shown. Fic. 203.—Bicarpellary, syncarpous gyncecium
of Tobacco (Nicotiana Tabacum). 0, ovary; s, style; g, capitate stigma. Fic. 204.—
Syncarpous gynoecium of Primrose (Primula) with simple style and capitate stigma.
Fic. 205.—Median lengthwise section through flower of Pzony (Paonia). ds,
dorsal suture of carpel; vs, ventral suture. (After Small.)

unites these two. The last is nonessential and when wanting
the stigma is called sessile, as in the Poppy. 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

THE FLOWER 285

portion, the style. A leaf thus transformed into an ovule-bearing
organ is called a carpel or pistil. The pistils 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 corresponds 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 may consist of a number of separate carpels, as
in the buttercup or Water Lily flowers, when it is said to be apo-
carpous, 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 mono-
carpellary; if two carpels enter into its formation, it is said to be
bicarpellary; if three, tricarpellary; if many, polycarpellary.

Compound Pistils are composed of carpels which have united to
form them, and therefore their ovaries will usually have just as
many cells (Jocules) 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 placentz as there
are carpels, but all more or less consolidated into one. The
partitions are called dissepiments and form parts 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.

Ovutes.—The Ovutes or MEGAsori are transformed buds,
destined to become seeds in the mature fruit. Their number
varies from one to hundreds. In position, they are erect, growing
upward from the base of the ovary, as in the Composite;
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 perpen-
dicularly from the top of the ovary.

In Gymnosperms the ovules are naked; in Angiosperms they
are enclosed in a seed vessel.

286 PHARMACEUTICAL BOTANY

Fic. 206. Fic. 208.

Fic. 211. Fic. 212,

O GW

Fic. 214. Fig. 215:
Forms OF GyNC&CIUM

Fic. 206.—Cone of apocarpous carpels (c) of Tulip-tree (Liriodendron tulipifera).
a, receptacle; e, stamens. Fic. 207.—Apocarpous carpels of Strawberry, raised
upon a swollen, hemispherical carpophore._ Fic. 208.—Ripened apocarpous
carpels of Raspberry on conical receptacle (/). Fic. 209.—Apocarpous carpels
of Pheasant’s Eye (Adonis). Fic. 210.—anthoxylum with gynophore (g) and apo-
carpous ovary (0) free styles but united stigmas (s). Fic. 211.—Two bi-lobed
carpels and gynobasic style (s) of Horehound (Marrubium vulgare). Fic. 212.—
Similar but more distinctly divided gyncecium of Forget-me-not (Myosotis). Fic.
213.—Flower of Rue (Ruta graveolens) to show carpels (ov) united only at base.
Fic. 214.—a, Diagram of three apocarpous carpels; }, cross-section of the ovaries
of the same. Fic. 215.—a, Diagram compound pistil of three carpels with free
styles and united ovaries; 5, cross-section of the ovaries of the same. (After Small.)

THE FLOWER 287

A complete angiospermous seed ovule which has not under-
gone maturation consists of a NucELLus or body; two coats, the
outer and inner InTEGUMENTs; and a Funicutus, or stalk.
Within the nucellus is found the Empryo Sac or MEGASPORE
containing protoplasm and a nucleus. (See Fig. 221.)

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
CyuaLaza. The Hirum 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 of the funuculus is called the
RAPHE.

Fic. 216.—Forms of Ovules. A, orthotropous. 8B, campylotropous. C, amphi-
tropous. D, anatropous. (From Gager, redrawn from Gray.)

Forms or Ovu.es.—The shape of the ovule may be ORTHO-
TROPous, or straight; CAMPYLOTROPOUS, bent or curved;
Amputrropous, 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, as the ovule of mustard.

Tue PLACENTA.—The placenta is the nutritive tissue con-
necting the ovules with the wall of the ovary. The various types
of placenta 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 Polygonacee
(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.

288 PHARMACEUTICAL BOTANY

SUTURAL PLACENTATION is seen in the Leguminose (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 PLac-
ENTATION is seen in Campanu-
lacee (Lobelia, etc.), Solanaceae,
etc., where the two, three, or
more carpels have folded in-
ward until they meet in the
center and in the process have
carried the originally parietal
placenta with them. This then
Fic, 217.—Heterostyly, a means of may form a central swelling

insuring cross pollination. Two forms of :
flowers of bluets (Houstonia) opened out bearing the ovules over the

and seen from the inside. A, short surface.
stamens and long style. 2B, long stamens FREE CENTRAL PLACENTA-

and short style. An insect visiting A, will TION occurs perfectly in the
receive pollen upon the front part of its

body; upon visiting B this pollen will rub Prémulacea, Plantaginacee and a
against the stigmas, and the pollen from few other families. In this the
this flower will be received by the under carpels simply cover over or
part of the body; upon visiting another : n
flower like A, the pollen from B will be FOOfin a central placental pillar
brushed against the stigma of A. (From around which the ovules are
Darwin’s “Cross Pollination.”? Courtesy of scattered.

my cote & Ce.) 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 considerably thickened.
It frequently divides into branches in its upper part. These are
called Styte Arms. As many style arms as carpels may be
present. In the one-carpelled pistil of some Leguminosae, the
usually bent-up style is the tapered prolongation of a single
flower. Again, in the apocarpous carpels of many flowers of the
Ranunculacee, each carpel bears a short to long stylar prolonga-
tion, When the carpels, however, are syncarpous, the common

THE FLOWER 289

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 gyncecium.

In some plants remarkable variations from the typical stylar
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 Campanulacea,
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 stigmas
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 prolon-
gation, viz.: (a) collecting stylar hairs,
functioning for pollen collection and
distribution; and (b) stigmatic hairs for
pollen reception from another flower.
In Vinca, the style swells near its
extremity into a broad circular stigma
and then is prolonged intoa short column
bearing a tuft of hairs that prevents the

Fic. 218.—Lilium Martagon.
Longitudinal section of the
stigma and upper part of the
style. The pollen grains,
caught on the papillze of the
stigma, have germinated, and
the pollen-tubes are growing
down along the walls of the
style canal. (From Gager after
Dodel-Port.)

entrance of insect thieves into the flower. In the genus Jris, the

common style breaks up at the insertion of the perianth into three,
wide, petaloid, stylearms. Each of these bifurcates at its extrem-
ity. On the lower or outer face of this is a transverse flap that
bears the stigmatic papilla. 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 por-
tion with five radiating ribs. At the extremity of each bifid end
of each rib is a minute, peg-like, stigmatic surface.

ae ea ee ee

290 PHARMACEUTICAL BOTANY

Tue SticMA.—This is usually a viscid papillose surface of
greater or less expanse functioning for pollen reception. If no
style intervenes between the ovary and stigma, the stigma is said
to be SEssILE, as in the poppy.

In wind-pollinated flowers
such as the grasses, the stigmas
are the numerous feathery hairs
which cover the ends of the styles
and are intended to catch flying
pollen grains. Such stigmas are
called plumose. In animal-pol-
linated flowers, the stigmas are
usually small restricted knobs,
lines or depressions. The stig-
matic papille 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 stig-
matic papillae are elongated,

Fic. 219.—Trimorphism and hetero- columnar, hair-like structures,
styly, a means of insuring cross pollina- Whereas in the short styles of
tion. The three forms of flowers in the short-styled flowers the papillae

loosestrife (Lythrum salicaria). The are small knob-like cellular
corollas and calyces are omitted in order :
swellings.

to show the different lengths of stamens
and styles. The functional crosses are Various terms are ascribed to

indicated by the arrows. (Fram Dar- stigmas to denote their shape,
win’s ‘Cross Pollination,’ Courtesy of ‘ :
D. Appleton & Co.) lobing, markings, appendages,
etc. Among the more common
of these are capitate, referring to a spheroidal stigma, bifid, or two-
lobed, truncate, when cut square off at the tip, annular, or ringed
below the summit, penicillate or brush-like, barbellate, having
minute barbs, etc.

POLLINATION

PouinaTIoN is the transfer of pollen from anther to stigma
and the consequent germination thereon. It is a necessary step
to fertilization.

THE FLOWER 291

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 sta-
mens and pistils standing in extraordinary relation to each other,
(6) by the anthers and stigmas maturing at different times, (¢) by
the pollen in many cases germinating better on the stigma of
another flower than its own and (d) by staminate and pistillate
flowers being borne on different plants of the same species.

In flowers like the geranium, clerodendron and chamomile,
the stamens mature and shed their pollen before the stigmas are
receptive. Such flowers are called protandrous. In the plantains,
figworts, etc., the stigma matures before the stamens. Such
flowers are protogynous.

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, Oak, Birch, Beech,
Wormwood, Hop, etc.

Insect-pollinated plants are called Enromopuitous. These,
being dependent upon the visits of insects for fertilization, possess
brilliantly 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 appa-
ratus. Examples: Orchids, Irises, F oxglove, Aconites, etc.

Plants pollinated through the agency of water currents are known as
Hypropuitous. To this class belong such plants as live under
water and which produce flowers at or near the surface of the
same. Some of these as Ruppia disperse floating pollen grains,
others like Zostera produce pollen of the same specific gravity as
water while Vallisneria, the common eelgrass, liberates floating
male flowers.

Bird-pollinated plants are called OrnirHopuiLous. Some
plants like the Trumpet Creeper, Nasturtium and Honeysuckle
are pollinated by humming birds. Other plants, as Aspidistra,

FO Te oe ON eee

eP PHARMACEUTICAL BOTANY

are pollinated by snails and slugs when they are called Mat-
ACOPHILous. These have dark colored flowers.

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 AND FERTILIZATION
(In ANGIOSPERMS)

MATURATION OF THE POLLEN GRAIN AND FORMATION OF THE
MA e GAMETOPHYTE.—The substance of the microspore (pollen
grain) divides into two cells, the mother and tube cells of the future
male 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 parti-
tion disappears and the thin endospore (¢ntine), carrying within it
the protoplasm in which are embedded the tube nucleus and two
sperm nuclei, penetrates through a deficiency of the exospore
(extine). 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 undergoes division until eight daughter-nuclei are produced
which are separated into the following groups:

(a) 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.

(b) 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 constitutes 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

THE FLOWER 293

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 nucleus to form the odspore.
The second sperm nucleus is absorbed as food. ‘The oéspore
develops at once into an embryo or plantlet which lies passive

Fic. 220.—Stages in the germination of the pollen grain and development of the
pollen tube. (From Robbins, after Holman and Robbins, after Bonnier and Sablon.)

‘until the seed undergoes germination. In a number of plants
including corn, some lilies, etc., the second sperm nucleus unites
with the previously fused polar nuclei to form the endosperm
nucleus. The union of one sperm nucleus with the egg nucleus
and the other sperm nucleus with the two polar nuclei is called
double fertilization.

FoRMATION OF ENDOSPERM.—The endosperm nucleus within
the embryo sac soon undergoes rapid division into alarge number
of nuclei that become scattered about through the protoplasm
of the embryo sac. These accumulate protoplasm about them,
cell walls are laid down, ENDosPERM resulting.

294 PHARMACEUTICAL BOTANY

Fic. 221.—Development of the female gametophyte in an angiosperm. A,
lengthwise section through a pistil showing a single inverted ovule within the
locule of the ovary. The embryo sac is in the center of the ovule surrounded by
the nucellus () and the two integuments (1) better shown in C. Note how the
pollen tube reaches and enters the micropyle of the ovule. _B shows the develop-
ment of the eight nuclei in the embryo sac. C shows the mature female gameto-
phyte and the tip of the male gametophyte; the former containing eight nuclei
and the latter containing only three. AP, antipodal nuclei; E, egg nucleus; EN,
two polar nuclei fusing to form endosperm nucleus; J, integuments; WV, nucellus;
S, sperm nuclei. (After Atwood.)

Fic. 222.—Development of the embryo. This figure is a continuation of the
preceding. D shows the antipodal cells (AP) disappearing, and the fusion of the
two sperm nuclei with the endosperm nucleus (ENV) and the egg nucleus (£).
The remains of the pollen tube (P) are seen in the micropyle (M). £ shows the
developing embryo with its suspensor (5) and the endosperm (END). F shows
the young embryo and its parts. C, cotyledons; PL, plumule; R, radicle. Note
that the embryo is surrounded by four coats. END, endosperm; .V, nucellus;
1 and OF, inner and outer integuments. His the hilum. (After Atwood.)

aac

j

THE FLOWER 295

DEVELOPMENT OF THE Empryo (DicoTyLepons).—In the
development of the embryo of a typical Dicotyledonous type, as
seen in Brassica (the mustard genus), the odspore, lying within
the embryo sac, undergoes division and redivision to form a sev-
eral celled filament. At the end of the filament farthest away
from the micropyle of the ovule, unequal growth and differentia-

Fic. 223.—Successive stages, 1, 2, 3, 4, 6, 8, in the development of the embryo
of Brassica. 1. Proembryo. 2. Octant stage. x, Octant. 7, i, Suspensor.
3. Formation of dermatogen. 4. Differentiation of periblem and plerome. 6. Last
cell or hypophysis, (v), of suspensor completes the dermatogen by division.
8. Inner part of v. d!, d?, Completes periblem. 2, Region of stem apex. A, A,
Bases of cotyledons. D, Dermatogen. Pe, Periblem. Pl, Plerome. (From

Small, after Kny.)

tion takes place which eventually produces the body or main part
of the embryo. Just below this region, the cells of the filament
develop into a suspensor whose basal cell is larger than the rest
and directed toward the micropylar end of the embryo sac.
The body of the embryo is formed as follows: The upper end
cell of the axis of the young developing embryo divides to form
2 rows of four cells each. The upper row of 4 cells by unequal
growth and repeated division form the terminal epicotyl and 2

296 | PHARMACEUTICAL BOTANY

lateral cotyledons. The 4 cells of the lower row similarly form
the hypocotyl.

MonocotyrLepons.—In the average Monocotyledon, the
odspore divides to form a short row of 3 cells. The basal cell
which is the largest of the three does not divide. The middle
cell of the row by repeated division and unequal growth of the
cells, develops into a suspensor, a hypocotyl] and a lateral epicotyl.
The terminal cell, by repeated divisions and unequal growth
forms the single cotyledon which is terminal on the axis of the
embryo. — :

CHAPTER XIII

THE FRUIT

The Fruir 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 Com-
posite, the modified calyx or pappus adheres to the ovary during
its maturation into the fruit and renders the fruit buoyant. In
Wintergreen, the calyx becomes fleshy, surrounds the ovary,
reddens, and forms the edible part of the fruit. In Ground
Cherry (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 Composite,
leathery and prickly as in the Chestnuts, scaly woody cups
(cupules) as in the Oaks, or foliaceous cups as in the Filberts.
Occasionally, as in the Fig, Osage Orange, Mulberry, etc., the
fruit may consist of the ripened flower cluster or inflorescence.

Fruir STRUCTURE

Tue Pericarp.—The Pericarp, or seed vessel, is the ripened
wall of the ovary. In general the structure of the fruit wall
resembles that of the ovary, but undergoes numerous modifica-
tions in the course of development.

The number of cells of the ovary may increase or decrease,
the external surface may change from 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
297

Pe Ne ee ee SES eee See

PL ee Me ae ne Ee ee ee ee

DE NS Pe ee ae Fy ET, ee ee ee ed

298 PHARMACEUTICAL BOTANY

the rind of the Orange, or assume the forms seen in the Gourd,
Peach, Grape, etc.

Frequently, as in the Apple, Pear, Quince, Wintergreen etc.,
the pericarp consists for the most part of other elements than the
ripened ovarian wall and is then termed a pseudocarp or anthocarp.

The pericarp consists of three layers of different texture, viz.:
epicarp, mesocarp and endocarp.
The epicarp is the outer layer.
The mesocarp the middle, and the
endocarp the inner 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

PS te ae union between adjacent septa or

1G. 4.—Frit o e a :
op on me ge 0 avalos,
section. (After Robbins.) : areca

into which the mature fruit sepa-
rates to permit the escape of the seeds. Depending upon the
number of these the fruit is said to be univalved, bivalved, trivalved, etc.

DeEHIscENCcE.—This is the opening of the pericarp to allow the
seeds to escape.

Fruits are either Dehiscent or Indehiscent according as to
whether 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 (INDEHIS-
CENT). Dehiscent fruits open regularly, or normally, when the

THE FRUIT 299

pericarp splits vertically through the whole or a part of its length,
along sutures or lines of coalescence of adjacent carpels.
Legumes usually dehisce by both ventral and dorsal sutures.
Irregular or abnormal dehiscence has no reference to normal
sutures, as where it is TRANSVERSE or CIRCUMSCISSILE, extending

--- posh of
Q recep ahh

medulla if
Y on ge

\ {

rh
\ |
Neal lobe J

Fic. 225,—A. The aggregate fruit of the Strawberry as seen in median longi-
tudinal section, X 24. B,Asingle achene. The greater bulk of this fruit consists
of a fleshy receptacle. Partially imbedded in the surface of this receptacle are
numerous seed-like bodies which represent true fruits or achenes. (After Robbins.)

around the capsule either entirely or forming a hinged lid, as in
Hyoscyamus, or detached.

Deniscence is called Porous or Apicat when the seeds
escape through pores at the apex, as in the Poppy; VaLvu.ar,
when valve-like orifices form in the wall of the capsule.

VALVULAR DEHISCENCE.—There are three kinds of valvular
dehiscence, viz.: septicidal, loculicidal and septifragal. Srp-

300 PHARMACEUTICAL BOTANY

TICIDAL DEHISCENCE is that form in which the constituent carpels
of a fruit become disjoined, and then open along their ventral
suture, as in Colchicum and Marsh St. John’s Wort capsules.

LocuticiDAL DEHISCENCE, or dehiscence into loculi or cells, is
the form which is seen when each component carpel splits down
its dorsal suture, and the dissepiments or partition walls separat-
ing the ripened carpels remain intact, as in Cardamomand Iris.
SEPTIFRAGAL DEHISCENCE is that form in which there occurs a
breaking away of the valves from the septa or partitions, as in
Stramonium and Morning Glory fruits.

CLASSIFICATION OF Fruits (according to structure).—SIMPLE
Fruits result from the ripening of a single pistil in a flower, as
the Tomato or Banana.

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,
Dewberry and Strawberry.

Muttipte. Fruits are those which are the product of the
ripening of a flower cluster instead of a single flower, as the Fig
and Pineapple.

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 < II. Schizocarpic (splitting).
III. Achenial (indehiscent).

S os Baccate (berries).

ucculent ;
V. Drupaceous (stone fruits).

The Capsuxar fruits include all of those, whether formed of
one or more carpels, which burst open to let their seeds escape.

Scuizocarpic or splitting fruits are those in which each carpel
or each half carpel (in Labiate) splits asunder from its neighbor
and then falls te the ground. The split portion is one-seeded.

ACHENIAL fruits are dry, one-celled, one-seeded and indehis-
cent 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

THE FRUIT 301

pulp formed by the endocarp or endocarp and mesocarp
combined.

Drupaceots 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 parenchyma 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 asin Peaches, Cherries, etc.

I. CapsuLAR Fruirs.—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 Fo.uicte or pod [Fig. 226 (1)] is a dry, simple, capsular
fruit formed of a single carpel which dehisces by one suture.
This is usually the ventral suture, as in Aconite, Staphisagria, Lark-
spur and some other Ranunculacee, but may be the dorsal suture as
in Magnolia and Illictum (Star anise).

A LecumE is a dry, simple, capsular fruit formed of a single
carpel and dehiscent by both ventral and dorsal sutures. Exam-
ples: Peas, Beans, etc. The legume is typical of most Leguminosae,
Fig. 226 (8).

A LomentvuM or Loment is a modified, usually jointed legume
that is separable transversely into one-seeded portions. Seen in
Acacia, Peanut, Tamarind, Entada scandens, Cathartocarpus Fistula,
Desmodium, etc. of Leguminosae, Fig. 227 (5).

A CapsuLe is a fruit formed of two or more carpels composing
a compound pistil which dehisce longitudinally or by apical teeth
or valves. Examples: Cardamom, Poppy, Irs, etc., Fig. 226 (2
and 3).

A Pyxis or Pyxipium is a capsular fruit formed of two or more
united carpels that dehisce transversely. Examples: Hyoscyamus,
Plantago, Portulaca. ‘The upper portion forms a lid which fits
upon the lower pot-like portion, Fig. 226 (4).

A Recma is a capsular fruit of two or more united carpels that
first splits into separate carpel parts and then each of these
dehisces. This type of fruit is typical of Hura crepitans (Sandbox),
Pelargonium (House Geranium) and Geranium, Fig. 226 (5).

302 PHARMACEUTICAL BOTANY

A S1uigua is a long, slender, one or two-celled capsule, often
with a spurious membranous septum (when two-celled) and two
persistent parietal placentz, the valves opening from below
upward. Examples: Mustard, Chelidonium and Wallflower, Fig.
226 (6).

Fic. 226.—Types of capsular fruits. 1, Pod of Aconite; 2, Capsule of Colchi-
cum showing septicidal dehiscence; 3, capsule of poppy, having porous dehiscence;
4, pyxis of Henbane; 5, dehiscing regma of Geranium; 6, siliqua of Celandine show-
ing valves opening from below upward; 7, silicule of Cochlearia; 8; Legume of Pea.

A Smicutz is a short siliqua in which the length is never
much greater than the breadth. Examples: Cochlearia, Fig.
226 (7); Shepherd’s Purse, etc.

II. Scuizocarpic Frurrs.—A Carceruus or NuTLET is the
typical fruit of the Labiata, but is also seen in the Boraginacea,
The ovary that has become four-celled at the time of flowering
matures into four little pieces which split asunder lengthwise.

THE FRUIT 303

Each split part is composed of one-half of a ripened carpel,
Fig. 227 (2).

A Cremocarp is the characteristic splitting fruit of the
Umbellifere family. It consists of two inferior akenes or meri-
carps separated from each other by a forked stalk called a carpo-
phore. These mericarps usually cling to the forks of the

W)
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4
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re
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3 Ao aS

<.

Fic. 227.—Schizocarpic fruits, and loment. 1, Cremocarp of Fennel, composed
of two mericarps (m) and a split carpophore (c); 2, carcerulus of the Bugle; 3, one-
carpelled: samara of the Ash; 4, double samara of the maple; 5, loment of purging
cassia, a portion of the pericarp being removed to show chambers, each containing

a Sirigle seed.

carpophore for a time after the cremocarp splits, but sooner or
later fall, Fig. 227 (1).

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. See Fig. 227 (3 and 4).

III. Acuentat Fruits (all indehiscent)—The AKENE is a
dry, one-chambered, indehiscent fruit, in which the pericarp is
firm and not united with the seed, the style remaining in many
cases as an agent of dissemination, Fig. 228 (1). The latter
may be long and feathery as in Clematis or be hooked. Exam-

304 PHARMACEUTICAL BOTANY

ples of akenes: Fruits of the Composite, Sedges, Buttercup,
Anemone, etc. ‘The Hip of the Roses consists of a number of
akenes in a ripened concave receptacle.

The Urricte is like the akene, except that the enveloping
calyx is loose and bladder-like. Examples: Chenopodium, Fig.
228 (3).
A Caryopsis or GRAIN is a dry, one-celled indehiscent fruit
whose pericarp is always fused with the seed coat. This fruit is
more likely than any other to be mistaken for a seed. Examples:

Fic. 228.—Achenial fruits. 1, Akene of Pulsatilla cut vertically, showing
adherent feathery style (st), pericarp (p), testa (t), endosperm (e), hypocotyl (A)
and cotyledons (cot) the last two structures making up the embryo; 2, caryopsis
of wheat showing beard of h4irs. above and position of embryo of seed below;
3, utricle of Chenopodium cut vertically to show calyx (c), pericarp (p) and seed
(s) regions; 4, nut of an oak consisting of a glans (g) and cupule (cu).

Wheat, Corn, Barley, Oats and other members of the Graminee
or Grass Family. Fig. 228 (2).

A Nut or GLans is a one-celled, one-seeded fruit with a
leathery or stony pericarp. Examples: Oaks, Beeches, Chestnuts,
Alders and Palms, Fig. 228 (4).

IV. Baccate Frurrts (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 suc-
culent 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, Fig. 229 (1).

THE FRUIT 305

A Bacca is a berry from an inferior ovary. Examples:
Gooseberry, Honeysuckle, Currant.

The Pepo or Gourp 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 Cucurbitacee and the Banana, Fig. 229 (2). ;

The Hesperipium 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 the membranous cover-

Fic. 229.—Baccate fruits. 1, Berry (uva) of Belladonna with adherent calyx;
2, Pumpkin, cut transversely illustrating a pepo fruit; (A), a locule; 3, hesperidium”
fruit of the Orange cut transversely showing epicarp (e), mesocarp (m), endocarp
(en), pulp (p), and seed (s).

ings 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, Fig. 229 (3).

V. Drupaceous Fruits (succulent fruits in which the meso-
carp is more or less succulent, but the endocarp leathery or
stony).—A Drupe is a one-celled, generally one-seeded, dru-
paceous fruit such as the fruit of the Plum, Peach, Prune, Sabal,
Rhus, Piper, Cherry, etc., whose endocarp or putamen is com-
posed wholly of stone cells or stone cells and sclerenchyma fibers,
Fig. 230 (1).

The Pome is a fleshy, drupaceous fruit, usually 5-carpelled,
with a tough epicarp, succulent or fleshy mesocarp, and leathery
or stony endocarp, the chief bulk of which consists of the adherent
torus. Quince, Apple and Pear are examples. The carpels

306 PHARMACEUTICAL BOTANY

constitute the core, and the fleshy part is developed from the
torus, Fig. 230 (2) and 231.

if

2

Fic. 230.—1, Drupe of cocoanut cut vertically, showing epicarp (¢), mesocarp
(m), stony endocarp (d), seed coat (s), endosperm (end), and embryo sac cavity
(e.s.) which in the mature seed contains a nutritive fluid. 2, Pome of an apple
cut vertically to show core composed of 5 ripened carpels and flesh of matured
torus. 3, Etzrio of raspberry. 4, Same, cut vertically to show arrangement of
the little drupes on fleshy receptacle.

AGGREGATE Fruirs.—An Era€rio consists of a collection of
little drupes or of achenes on a ripened torus of a single flower.

awe
——

a

A

Fic. 231.—Pome of apple. A, median lengthwise section; B, cross section of fruit.
(Robbins.)

Examples: Raspberry, Blackberry, Strawberry, etc., Fig. 230
(3 and 4).

a ea

THE FRUIT 307

Mu tiple Fruits.—The Syconrum is a multiple fruit con-
sisting of a succulent, hollow torus enclosed within which are
akene-like bodies, products of many flowers. Example: Fig.
Zam41},

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
Pineapple all the parts are blended into a fleshy, juicy, seedless
mass, and the plant is propagated by cuttings.

Fic, 232.—Miultiple fruits. 1, Syconium of Fig cut vertically to show hollowed
out receptacle (r) of ripened flower cluster; 2, strobile of the Hop; 3, galbulus of

Juniper.

The STROBILE or Cone is a scaly, multiple fruit consisting of a
scale-bearing axis, each scale enclosing one or more seeds. The
name is applied to the fruit of the Hop, Fig. 232 (2), and also to
the fruit of the Conzfere in which the naked seeds are borne on the
upper surface of the woody scales.

A GaLsBuLus is a more or less globular multiple fruit formed of
three fleshy connate scales each bearing a seed on its upper or
inner surface, as in Juniper, Fig. 232 (3).

HistoLocy 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.—Pass-
ing from periphery toward the center, the following structures
present themselves:

308 PHARMACEUTICAL BOTANY

1. Epicarp or epidermis consisting 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.

See

i Set antes

seresreuel. Roane
Vaczghepricasse se: ?
RR : mee gs

Pay"

ae
ge
(Alea “

eaae

PRC ONT ESS

NM
ay
an
Bh
ae
CS
4
AS

H,
KS
:
‘

Fic. 233.—Vanilla. Structure of the capsular fruit. ¢, cuticle; ¢p, epidermis;
hy, hypodermis; mes, mesocarp of parenchyma in which region fibrovascular bundles
(4) course; ir, tracheae, si, sieve tissue and J, sclerenchyma fibérs of a bundle; end,
endocarp; f/, placenta; h, glandular hairs; sd, seed; s, suture; cr’, prismatic crystals
and cr?, raphides of calcium oxalate.

2. MesocarpP, a broad region divisible into a narrow, outer
hypodermis and broad inner parenchyma zone. The hypo-
dermis 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. The inner
parenchyma zone is composed of somewhat loosely arranged,
large, thin-walled parenchyma cells becoming smaller in the
inner part 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

THE FRUIT ee

a drop of strong sulfuric acid is added, a carmine-red color will
be observed showing the presence of vanillin in this region. A
number of closed collateral bundles will be seen coursing through
the mesocarp.

3. ENDocarpP, 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
contain an abundance of balsam; the placental region covers the
six bifid placente which extend into the cavity of the capsule. _ Its
(inner) epidermis is composed of mucilaginous cells.

Fic. 234.—Photomicrograph of a transverse section of a mericarp of Feniculum
vulgare, showing epicarp (A), mesocarp (B), endocarp (F), vitta (C), endosperm
of seed (D), carpophore (G) and fibro-vascular bundle in primary rib (Z). (Highly
magnified.)

4. SeEps.—These are minute blackish bodies attached to the
placental twigs of the placentae. Some of them may have been
torn off in cutting the section. ;

HistoLocy oF A TypicAL MERICARP, Fa@nicuLuM.—This
five-angled fruit, in transverse section, shows a concave com-
missural and convex dorsal surface. Passing from the surface
toward the center we note:

1. Epicarp, or outer covering tissue, composed of colorless
epidermal 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,
isodiametric cells, beneath which are two to several additional

310 PHARMACEUTICAL BOTANY

layers of thicker-walled cells with brownish walls. Through the
angles or rib portions of the mesocarp extend the /ibro-vascular
bundles. Each of these consists of a few spiral, annular and
simple pored .tracheids and tracheae and many sclerenchyma
fibers. 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 vittze contain the official oil of fennel. Two vitte gen-
erally occur in the mesocarp of the commissural side although
four are reported to have been found in this region of some
varieties of fennel.

3. Endocarp, a narrow layer of oe transversely elongated,
except over the regions of bundles where they may be seen elon-
gated in several directions.

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, somewhat reniform mass of more or
less polygonal cells containing aleurone grains and fixed oil
globules. Each aleurone grain generally contains a rosette
aggregate of calcium oxalate and one or two globoids.

6. Embryo, embedded in the endosperm of the upper region of
the seed, with its hypocotyl directed upward.

CHAPTER XIV
THE SEED

A Seep is a fertilized and matured ovule containing an
embryo.

The Empryo is a young plant in a dormant state of existence.
It is capable of remaining alive for a long period under a broad
range of conditions of temperature and moisture. When
fully developed within the seed it consists of four principal
parts: the caulicle or embryonic stem, the plumule or embryonic
bud, the radicle or embryonic root and one, two or more seed
leaves called cotyledons.

The plumule is at the summit of the caulicle and consists of
the young growing apex of the caulicle with delicate rudimentary
leaves enveloping it. The radicle is a continuation of the lower
end of the caulicle. It develops, upon the sprouting of the
seed, the primary or first root of the seedling. The cotyledons
frequently are carried above the soil, upon germination, and
there function to build up food for the seedling plantlet until the
latter has developed foliage leaves. In some seeds the cotyledons
contain stored food which is used by the embryo as it develops
into the seedling plantlet.

All that portion of an Empryo PLANTLET above the cotyledons
is termed the eficotyl and all that part below the cotyledons, the
hypocotyl.

The purpose of the seed is to insure the continuation and
distribution of the species. It is therefore an organ of reproduc-
tion. Like all other organs of the plant, seeds exhibit irritability,
nutrition, respiration and growth. These, in the dry seed, are
in the state of reduced activity. When properly stored, seeds
may live for one year, as those of Aconite and Larkspur, or one to
three years, as those of the Grass family, or two to five years, as
those of the Rose and Mallow families, up to 20 years, as those of

Tobacco or rarely 5 to 87 yearsas those of the Bean and Pea family.
311

312 PHARMACEUTICAL BOTANY

SEED SrRuCTURE.—The average seed shows one or two SEED
Coats that enclose an Empryo which fills the seed cavity; the
seed is then termed exalbuminous, as in beans, peas and other
leguminous seeds; or between the seed coat and embryo reserve
food may be stored, and the seed is then termed albuminous, as in
the cereals, castor bean, etc. The stored food in or around the
embryo may consist of starch and protein (pea, bean), or starch,

Chalaza

4 -~-Endosperm

a = Cotyledons

Raphe---~ :

x _ Plumule
£. -~ Hypocotyl

ee Eig - <<< Caruncle

i

1

; B

J

i

!

Hilum

Fic. 235.—Castor-oil seed Ricinus communis), an albuminous type. A, view of
seed halved flatwise, showing surface of cotyledon and hypocotyl. B, longitudinal
section at right angles to the surface of the papery cotyledons. (Mottier.)

protein and oil (peanut), or oil and protein and active medicinal
principle (castor and croton), or of hard reserve cellulose and
protein (date, nux vomica, coffee, ivory-nut, palm-nuts). 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 SPERMO-
DERM. 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

THE SEED 313

crustaceous; if smooth and glossy, it is polished, if roughened, it
may be pitted, furrowed, hairy, reticulate, etc.

The testa may often present outgrowths or seed appendages
whose functions are to make the seeds buoyant, whereby they
may be disseminated by wind currents. Examples of these are
seen in the Milkweed, which has a tuft of hairs at one end of the
seed called a Coma, in STROoPHANTHUSs, which has a long bristle-
like appendage attached to one end of the seed and called an

Fic. 236.—Appendages of seeds. A, coma (c) on milkweed (Asclepias); B,
strophiole (st) on bean (Phaseolus) h, hilum, m, micropyle; C, strophiole on Corydalis
aurea; D, aril on white water-lily (Castalia); E, projecting raphe (r) on pitcher plant
(Sarracenia purpurea); F, wing on trumpet creeper (Tecoma radicans); G, arillode
(“mace”) on nutmeg. A, B, and G from nature; C-F, redrawn from Gray. (After

Gager.)

Awy, and in the seeds of the Trumpet Creeper and Pines which
have wings. ‘The excrescence or slightly developed aril at or
near the hilum of some seeds, as in Castor and Colchicum Seed, is
called the CARUNCLE or STROPHIOLE.

The tegmen or inner coat surrounds the nucellus closely and
is generally soft and delicate.

A third integument, or accessory seed coat covering the out-
side of the testa, is occasionally present and is called the Arm.
or arillus. Examples: Evonymus (succulent) and Cardamom

(dry).

314 PHARMACEUTICAL BOTANY

When such an integument arises from the dilatation of the
micropyle of the seed, as in the Nutmeg, it is known as an
ARILLODE. :

The KERNEL consists of all the seed substance within the seed
coats. ALBUMEN is the name of the nutritive matter stored in the
seed. The Funicutus or seed stalk is absent in most of the
official seeds. The scar left by its separation is called the Hitu.
When the funiculus is continued along the outer seed coat, it is
called the RAPHE.

MopbeE oF ForRMATION OF DIFFERENT TYPES OF ALBUMEN

If the fertilized 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 nutritive material is called Perispermic albumen.
The tissue of the seed containing it is called the PerisPERM and
represents the ripened nucellus of the ovule. Seen in the Poly-
gonacee or Buckwheat family, etc.

In by far the greater number of Angiosperms, the endosperm
nucleus, divides and redivides, giving rise to numerous nuclei
imbedded in the protoplasm of the embryo sac, outside of the
developing embryo. Gathering protoplasm about themselves
and laying down cell walls they form the ENDosPERM tissue out-
side of the embryo. Into this tissue food is passed constituting
the Endospermic albumen.

In the Arrowroot family (Marantacee), the Pepper family
(Piperacee), 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 nourish-
ing substance, thus resulting in the formation of abundant
PERISPERM and a small reduced ENDosPERM.

ExaLpuminous SEEps 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: Puysosticma, AMYGDALUS, etc.

ALBUMINOUS SEEDs are those in which the nourishment is not
stored in the embryo until germination takes place. Such

THE SEED 315

seeds show a larger nourishing tissue region and a smaller embryo
region, Examples: Nux Vomica, Myristica, Coxcuicum,
Castor, etc.

Gross STRUCTURE OF A Monocotyt SEED (wiTH Frurr 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 erroneously 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 base or “tip” region marks the part of the grain
which was inserted into the cob. Upon it may be found papery
chaff, representing parts of the pistillate spikelet. The groove
noted on the broader surface indicates the position of the
embryo.

HisToLoGy OF THE INDIAN Corn SEED (wiTH FRuir WALL
ATTACHED).—If a longitudinal section be cut through the lesser
diameter of a soaked grain, the following histologic character-
istics 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 SPpERMODERM or seed coat, a single layer of delicate
elongated cells. .

3. The PerRispeERM, another layer directly underneath the
Spermoderm, difficult to distinguish without special treatment,
and representing the ripened nucellar tissue of the ovule.

4. The ENposPERM 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. (6) Starch Parenchyma, consisting of two
regions: an outer horny zone composed of cells containing for the

316 ; PHARMACEUTICAL BOTANY

5S ~ 69) fo )

—

Fic. 237.—Indian Corn grain and seedling. 1, Median longitudinal section
through lesser diameter of grain of corn showing fruit wall (pericarp) and adher-
ent seed coat (spermoderm) p + 5; horny endosperm (end!); starchy endosperm
(end”); palisade layer (pal) scutellum (s) with vascular tracts (v); cotyledonary
sheath (cot); radicle (r) and plumule (pl). 2. Transverse section of a representa-
tive portion of outer region of corn grain showing pericarp (#) and spermoderm
(s) with tube cells (¢.c.); endosperm (end) with an outer layer of aleurone cells
containing aleurone grains (a/) and an inner starch parenchyma zone (st). 3.
Young seedling showing first leaves (/); primary root (r) bearing secondary roots
(sr) with root hairs (A); cotyledonary sheath (cs) and seed coat (s).

THE SEED 317

most part polygonal starch grains and oil droplets; and an inner
mealy zone of cells with mostly rounded starch grains.

Fic. 238.—Median longitudinal section through the lesser diameter of a grain of
corn (Zea Mays). p, pericarp which is attached to subjacent seed coat; al, aleurone
layer of endosperm; end, horny endosperm; fe, floury endosperm; e, epithelium of
scutellum; sc, scutellum; cr, coleorhiza; r, radicle; pl, plumule. X 8.

5. The Empryo, 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 rudi-
mentary root, with its tip covered by a root cap. The cotyledon

318 PHARMACEUTICAL BOTANY

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 surrounds and protects the rest of the embryo.
That portion of the cotyledonary sheath which surrounds the
radicle is called the coleorhtza; that portion surrounding the
plumule, the coleoptile.

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
colored 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 contents of its cells are not baled out to the embryo until
after germination begins. Indian Corn is therefore an albuminous
seed.

GERMINATION OF THE SEED

When any viable seed is planted in suitable soil, and furnished
with oxygen and water and a certain degree of heat, germination
takes place. The first step in germination is the zmbzbition or
taking in of water. In the presence of moisture, etc., the seed
imbibes water and the embryo and endosperm (when present)
swells. ‘The ferments present within the cells of the embryo and
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 oxygen and gives off carbon
dioxide. The oxygen oxidizes the food materials within the
tissues with an accompanying release of energy, which is neces-
sary to life and growth.

The combined pericarp and spermoderm bursts opposite the
tip of the radicle, and the radicle, piercing through the cotyle-
donary 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,

THE SEED 319

grows downward into the soil forming the primary root, and devel-
ops upon itself secondary or lateral roots, all of which giverise to
root-hairs just above their root caps. Additional lateral roots
emerge above the scutellar region which ere long attain the size
of the first or primary root. The caulicle, 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 surrounding both
the root and the stem for a portion of their length. By this time

Fic, 239.—Different ways in which seedlings emerge from the soil. A, garden
pea; B, corn; C, Lima bean; g, ground level. (Motter.)

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 DocoTyL SEED, PHAsEoLus LuNATUS
(Lima Bean).—The Lima Bean Seed shows a flattened-ovate to
somewhat reniform outline. Externally it exhibits a polished
seed coat which is perforated on its basal side by a minute pore

320 PHARMACEUTICAL BOTANY

Fic. 240.—Lima Bean. A, seed showing scar (hilum); B, embryo after removal
of seed coat showing one cotyledon (cot) and caulicle (c); C, embryo after removal
of one of the cotyledons showing the second cotyledon (cot), plumule (p), caulicle
(c), and radicle (r); D, young seedling showing epicotyl (ep) bearing first pair of

green leaves (/) and a terminal stem bud (4); cotyledons (c), hypocotyl (A) with first
root (r) and rootlets (rt); E, cross section of representative portion of seed showing
cuticle (c), palisade cells (f), column cells (4) and spongy parenchyma (sf) of
spermoderm; epidermis (ep) and parenchyma (fa) of mesophyll of cotyledon
(magnified).

THE SEED o21

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

Fic. 241.—Stages in the germination of the common bean (Phaseolus vulgaris).
Showing nutation and hyponasty. (After Atkinson.)

and called the plumule, the caulicle or rudimentary 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 structure will be evident:

1. SPERMODERM of three regions, viz.; Palisade cells, Column
cells, and Spongy Parenchyma. ‘The palisade cell layer is com-
posed of vertically elongated, thick-walled cells which are covered
on their outer faces by a clear glistening cuticle. The column
cells, found forming a layer directly beneath the palisade zone,
are hour-glass-shaped.

322 PHARM ACEUTICAL BOTANY

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.

2. Empryo, the two cotyledons of which make up the greatest
bulk. These are composed of an epidermis covering over a
region of mesophyll. ‘The mesophyll is constituted of moderately
thick-walled cells which contain ellipsoidal and kidney-shaped
starch grains up to 65u 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.

GERMINATION OF Dicotyt SEEDs.—Following the imbibition
of water and swelling of the seed contents, in dicotyledonous
plants the hypocotyl is generally the first part of the embryo to
elongate and emerge from the seed coat. Responding positively
to the stimulus of gravity, it grows directly toward the center of
the earth and enters the soil, becoming the primary root. The
primary root continues to grow in the soil and ere long develops
secondary roots. In the seeds of the Lima and other beans, the
cotyledons and epicotyl are pulled above the soil by the elonga-
tion of the hypocotyl which makes a crook. In seeds of the pea
and corn, etc., in which the cotyledons or cotyledon remain
within the seed coat under the ground, the elongation of the
epicotyl brings the aerial organs above the soil surface.

CHAPTER XV E
‘THE CLASSIFICATION AND NAMING OF PLANTS ;

The classification and naming of plants belongs to that depart-
ment of Botany called Taxonomy, a name which was derived
from two Greek words, taxts, arrangement, and nomos, law.

PRINCIPLES OF CLASSIFICATION

The modern classification of plants is an attempt to arrange
plants in groups based upon their natural relationship or genetic
affinities. The basis of all modern classification alike of plants
and animals is the theory of organic evolution.

Organic evolution presumes that all organisms living to-day
have descended from preexisting organisms either by direct
descent or by common. ancestry. Sufficient evidence has been
brought forth during the past century to substantiate plant
evolution. The sources of this evidence have been found in part
through the study of comparative embryology, anatomy and
histology of plants, the study of fossil plants found in the rocks of
past ages of the world’s history in comparison with living forms of
the present age, the comparison of life histories of present day
plants, and through the more recent sero-diagnostic tests of Carl
Mez and his pupils. 7

A system of classification based upon natural resemblances of
plants related by descent is called a NATURAL System. Any
system of classification which does not take into account the
natural kinships and evolutionary tendencies existing between
plants is called an ARTIFICIAL SYSTEM. ©

A number of artificial systems were derived and used by
botanists of the past. Most of them were based upon some
arbitrarily chosen character such as woody stems, petals,
stamens, etc.

Theophrastus (370-285 B.C.), a pupil of Aristotle, and prob-
ably the earliest of pioneers in attempting a scientific grouping of

plants, classified them structurally into trees, shrubs, undershrubs,
323

324 PHARMACEUTICAL BOTANY

and herbs. He was the first to distinguish between certain
natural families.

Leonhard Fuchs (1501-1566), a German physician and
botanist, in his Strpium Historia (1542), an Herbal, chose to
arrange plants alphabetically by their Greek names.

Hieronymus Bock (1498-1554) published an Herbal in 1546
called Kreuter Buch in which he carefully described from nature
many plants which he grouped under herbs, shrubs and trees.

Fic. 242.—Linnaeus, the father of modern systematic botany. (After Gager.)

Caesalpinus (1519-1603) whose real name was Andrea
Cesalpino, an Italian physician and botanist, was dissatisfied
with the alphabetical arrangement and the narrowness of the
earlier artificial systems which were largely restricted to medicinal
plants. He attempted in his De Plantis (1583) a classification of
all plants without regard to their economic importance and
based it upon their fruit and seed characters.

Gaspard Bauhin (1560-1624), a Swiss botanist, grouped
plants into seaweeds, grasses, lilies, trees and shrubs.

‘yee wee

KK .
Baie
THE CLASSIFICATION AND NAMING OF PLANTS 325 *

John Ray (1627-1705), an English botanist, in 1682 intro-
duced a method of classification which distinguished between
flowerless and flowering plants. He arranged plants into. two
chief divisions, viz.; woody plants and herbs, and made cotyle-
dons a basis for subdividing flowering herbs into monocotyledons
and dicotyledons.

Joseph P. De Tournefort (1656-1708), an eminent French
botanist, chose petals as his basis of grouping and subdivided the
flowering plants into the Apetal: or the plants without petals and
the Petalodes or those possessing petals, distinguishing herbs and
undershrubs, trees and herbs. He was the first botanist to
describe genera.

The greatest of these artificial systems was that of the Sexual
System of Carl von Linné, popularly called Linnaeus (1707—
1778), a Swedish naturalist, who through his great works,
Systema Nature, published in 1735 and Species Plantarum, published
in 1753, did more to establish Botany on a scientific foundation
than any previous investigator. Linnaeus divided plants into
sexual and asexual, the former of which he designated as Phanero-
gamia or flowering plants and the latter as Cryptogamia or flower-
less plants. His classification of the flowering or seed plants was
based primarily on the number and position of the stamens, each
of the larger groups being subdivided according to the number
and character of the carpels. In his Species Plantarum, he
systematically described a great list of plants giving to each species
a binomial name consisting of two Latin words. ‘This system of
naming plants, called the “Binom1AL System,” has been followed
by all scientific botanists since Linnaeus and is still in use at the
present day.

The first natural system of classification was that devised by
Antoine Laurent de Jussieu, a French botanist, who in 1789,
issued his Genera Plantarum. ‘This work formed the basis of all
future natural systems. Jussieu was the first to group the genera
into families, and many of his concepts on families are used with
modifications to-day.

In 1813, Augustin P. De Candolle, a follower of the teachings
of Jussieu, issued his Theorie elementaire de la botanique in which
he demonstrated that the affinities of plants are to be found

326 ‘ PHARMACEUTICAL BOTANY

through the study of the comparative embryology and structure
of plant organs. He later published his famous Prodromus which
was continued by his son Alphonse de Candolle with the aid of
eminent collaborators. This work gives descriptions of genera
and species of Dicotyledons and represents a modification and
elaboration on the system of Jussieu.

From 1862 to 1883, George Bentham and Joseph D. Hooker,
two eminent British botanists, issued their Genera Plantarum.
This work contains a systematic arrangement of the Phanerogams
with a description of all of the genera then known.

This system was followed by the systems of A. W. Eichler
(Berlin 1883) and of Engler and Prantl. The latter, called ‘‘Die
naturlichen Pflanzenfamilien,” has since its appearance become the
most popular guide of systematists. It is based on the belief that
the absence of a perianth is a primitive condition and that
Monocotyledons precede Dicotyledons.

In 1926, J. Hutchinson, of the Royal Botanical Gardens at
Kew, issued the first volume of his work, ‘‘ The Families of Flower-
ing Plants.” ‘This is a new natural system on the seed plants and
is based on the assumption that plants with sepals and petals
associated with other floral and anatomical characters regarded
as primitive are more ancient than plants without sepals or
petals. The Monocotyledons are placed after the Dicotyledons
from which they are claimed to have been derived.

In every modern natural system, every individual belongs to
a species or unit of classification.

A SpPEctEs is a mental concept of a group of individual plants
having their more stable characters in common. While the
individual plants making up the group called a species may differ
in the size and number of their various organs and even show small
differences in the shape of their leaves or fruits, the parts of their
flowers will usually be quite similar in form, number and arrange-
‘ment. These more stable characters enable one to recognize
any members of a given species and to distinguish it from mem-
bers of other species. When an individual of a species repro-
duces, its offspring will show the same essential or more stable
characters as its parent or parents. Thus, all the Oak trees
which we commonly recognize as white oak constitute the species
Quercus alba,

THE CLASSIFICATION AND NAMING OF PLANTS 327

Moreover, every species belongs to a genus, every genus to a
family, every family to an order or cohort, every order to a class,
every class to a subdivision and every subdivision to a division
or phylum of the Plant Kingdom.

A GENUs is a group of related species. ‘Thus, the plants we
recognize as red oak, scarlet oak, pin oak and white oak belong to
the genus Quercus. The chief structure upon which the concept
of a genus is based is the flower, but other organs also play a part.

A FamIty is a group of similar and related genera. ‘Thus, the
oak genus (Quercus), the chestnut genus (Castanea) and the beech
genus (Fagus) constitute the family called Fagacee.

An Orpber is a group of related families. ‘Thus, the Birch
Family (Betulacee), the Hazel Family (Corylacee) and the Oak
Family (Fagacee) constitute the order known as Fagales.

It will be noted that the common family ending is ACEz&,
more rarely 2 and the order ending ALEs.

A Crass is a still higher group composed of related orders.
Thus, the Fagales, Ranales and a large number of other orders are
placed in the Class Dicotyledonez, because all of them have
several outstanding characteristics in common, such as netted-
veined leaves, two cotyledons in embryo, parts of their flowers in
fours and fives, and open collateral bundles.

A SuppIvision is a group of related classes. ‘Thus, the Class

Dicotyledonez and the Class Monocotyledonez, while differing
in several characters, resemble each other in possessing covered
ovules and seeds and so are placed in the subdivision Angiospermae.
A Division or PoyLum is composed of related subdivisions. ‘Thus,
the Subdivision Angiosperme or related plants having covered
seeds and the Subdivision Gymnosperme or plants with naked
seeds agree in possessing seeds and so are placed in the Division
Spermatophyta.
- The Divisions Spermatophyta (seed plants), Pteridophyta
(ferns and their allies), Bryophyta (liverworts and mosses) and
Thallophyta (seaweeds, fungi, etc.) make up the Kincpom oF
PLANTS.

In some instances species are subdivided into smaller groups
called Varieties and these in turn into SuB-VARIETIES, Forms and
Strains. Classes may be subdivided into Sus-cLasses. Thus,
the Class Dicotyledonee contains two subclasses, the ARCHI- —

328

PHARMACEUTICAL BOTANY

Division-Spermatophyta (seed plants).
Subdivision-Angiospermz: (seeds enclosed).
Class-Dicotyledonez (seeds with two cotyledons).
Sub-class-Archichlamydez (petals disjoined).
Order-Ranales (carpels disjoined, etc.).
Family-Ranunculacez (herbs with many hypogy-

nous stamens).

Sub-family-Helleborez (petaloid sepals hooded).
Species-Aconitum Napellus (carpels, 3 or 4;

Oivispor't
PLA
THALLOPHYTA
BRYOPHYTA }
PTERIDOPHYTA !
SPERMATOPHYTA |

f\

—) eae

Angiospermz 1

ete.)

OuTLINE OF PLANT Groups

Cyanophycez
Chlorophycez
Bacillarieze

Phaeophycez
Rhodophyceze

Schizomycetes
Myxomycetes

Algze

Phycomycetes
Ascomycetes
Basidiomycetes
Deuteromycetes
Ascolichens
Basidiolichens
Ricciales
Marchantiales
Jungermanniales
Anthocerotales

Fungi

Lichens

Hepatic

Andrezales
Bryales
Sphagnales
Lycopodiales
Selaginellales
Tsoetales
Equisetales

|
|
oa
|

Musci

Lycopodineze
Equisetineze
Filicinee Filicales
Hydropterales
Cycadales
Ginkgoales
Coniferales
Gnetales
Monocotyledonez
Dicotyledonez

Gymnospermz

Blue-green alge
Green alga
Diatoms
Brown alge
Red alge
Bacteria
Slime molds
Alga-like fungi
Sac-fungi
Club fungi
Fungi imperfecti
Lichens
1 Lichens
Riccia, etc.
Marchantia, etc.
Leafy liverworts
Anthoceros, etc,
Black mosses
True mosses
Peat mosses
Club mosses
Little club mosses
Quillworts
Horsetails

ead s tongue ferns

Water ferns

Sago palms

Ginkgos

Conifers

Gnetums and ephedras

Grasses, lilies, etc.
yaan. cherries, etc.

THE CLASSIFICATION AND NAMING OF PLANTS 329

CHLAMYDE# or that group in which the petals are either distinct
and separate or entirely wanting and the METACHLAMYDE# or
SYNPETAL@ in which the petals are more or less joined to form a
tubular corolla. Large families may be subdivided into Sus-
FAMILIES or tribes which in some works are given full family rank.

A Hyprw is a cross breed of two varieties or species, rarely of
two genera.

A complete classification of the medicinal plant, Aconite, will
show the principal subordinate groups of one of the divisions.
(See p. 328.)

BoTANIGAL NOMENCLATURE

Before Carl von Linné (Linnaeus), the great Swedish natural-

ist, brought forth the binomial system of nomenclature, no
uniformity existed in the assignment of plant names. Among the
pre-Linnean botanists there were some who designated plants
by single names, others who employed sentences in naming
‘them, some of which were quite lengthy, and a number who
adhered to the practice of naming them in their own modern
tongue. The result was quite obvious, a number of systems
were employed and confusion prevailed among students.

Specific Name.—According to the binomial system which
has been universally adopted, every plant belongs to a species
which is given two Latin names which together represent its
Sprciric NAme. The first name is the name of the genus or
Generic Name, the second, the name of the species or SPECIFIC
Eprruet. The generic name corresponds, in the naming of
persons, to the surname or family name, while the specific
epithet is analagous to the given name. Thus, the Wild Cherry
Species is named Prunus serotina, Prunus representing the name
of the genus, serotina the specific epithet or kind of Prunus.

The application of the specific name is determined by the
type specimen, i.e., the specimen or more than one with which
the author of the species worked.

Generic Name.—The name of the genus (pl. genera) is
always a noun in the singular number and must not be applied
to more than one genus. Its spelling should begin with a
capital letter. Genera names may be taken from any source

a
‘a
a
4
a

330 PHARMACEUTICAL BOTANY

whatever. Some, like Fagus for the Beech genus, and Acer for
the Maple, are of Latin origin. Others have been latinized
from other languages. Some have been named after some
therapeutic property, their roots, leaves, flowers or seeds were
thought to possess; for example, Jateorhiza, a latinized compound
of two greek words, idrecpa, healing, + pifa, root, because of the
healing virtues of the root. A number have had names ascribed
to them because of some peculiarity of structure, color, taste, odor,
behavior, habit or appearance of the plant or portion thereof.

Thus, Eriodictyon (from Gr. €pov, wool + dixrvor, net) alludes
to its wooly, netted veined leaves; Melaleuca (from Gr. pé\as,
black, + Aevxds, white) alludes to the black bark of the trunk and
white bark of the branches; Marrubium (from Hebrew marrob,
bitter) refers to its bitter sap; Barosma (from Gr. Baptis, heavy
+ ooyuy, odor) in allusion to its strong smell; Epifagus (from Gr.
éri, upon + Lat. Fagus, the beech) alludes to its growth on the
roots of that tree; Impatiens (from Lat. in, not and patiens, endur-
ing) refers to the sudden bursting of the capsules of this genus
when touched; Lycopodium (from Gr. \bcos, a wolf, + zods, a foot)
pertains to the appearance of the shoots of this genus. Many
have been named in honor of eminent naturalists or ‘friends of
these, or other noted persons. For example, Collinsonia was
named in honor of Peter Collinson, an English botanist of the
18th century; Dvzoscorea in honor of Dioscorides, the Greek
naturalist; Paullinia after Paullini, a German botanist of the 17th
century; Cinchona in honor of the countess of Chinchon, who
brought the bark to Europe in 1640, and Jeffersonia, in honor of
Thomas Jefferson.

Speciric EpirHet.—The specific epithets are for the most
part adjectives which agree with the names of genera to which
they belong in case, gender, etc. They may, however, be nouns
and in a few instances consist of two nouns or a noun and an
adjective. If an adjective, it should begin with a small letter,
as in Rhus glabra and Evonymus atropurpureus. This holds if it is
drawn from a geographical name, ¢.g. Rubus allegheniensis.
When the specific epithet is a noun, it may either be a proper
noun in the genitive case when it should begin with a capital,
as Garcinia Hanburyi; or it may be a common noun in. the geni-
tive, when it should begin with a small letter, as Grindelia cam-

THE CLASSIFICATION AND NAMING OF PLANTS 331

porum; or the noun may be in apposition to the generic name
and so in the same case, as Cytisus scoparius. Names that had
formerly been used for genera but since reduced to species are
usually capitalized, whether originally proper nouns or not, as
Aristolochia Serpentaria and Anacyclus Pyrethrum. In cases where
two nouns make up the specific epithet, the first of these is in
the nominative case, the second in the genitive, the two names
being connected by a hyphen, as Capsella bursa-pastoris. The
botanical name of the species yielding the drug, Aspidosperma,
(Aspidosperma Quebracho-blanco) will serve as an example of the
specific portion of the names being composed of a noun and an
adjective. Specific epithets taken from names of persons begin
with a capital.’

Variety EpirHeT.—Names of varieties are applied in two
different ways. Either the name of the species is given and
followed by the prefix var. before the varietal epithet, as Cheno-
podium ambrosivides var. anthelminticum; or the varietal epithet
may be appended to the name of the species, as Chenopodium
ambrostotdes anthelminticum.

It frequently happens that a botanist is careless in naming a
species and, without ascertaining whether the same name has
been assigned to another species, applies it to his, thus causing
duplication. For example, there have been two distinct species
of plants named Eupatorium laevigatum, one a form of Joe Pye
Weed, the other a species of the American tropics. In this case
the name Eupatorium laevigatum does not tell us which species
the writer or speaker refers to; it might be the tropical plant
called Eupatorium laevigatum by Lamarck or the Joe Pye Weed
named Eupatorium laevigatum by Torrey at a later date. Accord-
ingly, it is necessary to add to the name of the species the author’s
name. In this connection it is customary to abbreviate the
name of the author thus, Lam. for Lamarck, Torr. for Torrey,
L. for Linné, Mill. for Miller, Ait. for Aiton, Loisel. for Loiseleur-
Deslongchamps, or Tourn. for Tournefort.

Whenever a plant is transferred from one genus to another, it
must retain its original specific epithet, unless the genus to which
it is transferred already possesses a species with that epithet, in

-1 There is a tendency amongst American botanists to use the small letter in
beginning the spelling of all specific names irrespective of their source.

332 PHARMACEUTICAL BOTANY

which case a new specific name must be given it. Moreover,
the name of the botanist who assigned the original specific
epithet but placed it under a different genus must be placed in
parenthesis between the specific epithet and the name of the
botanist who later connects it with another genus. For example,
we read as the definition for Purging Cassia in the National
Formulary V—‘‘The dried fruit of Cathartocarpus Fistula (Linné)
Persoon.” ‘The significance of the name Linné in parenthesis
is that he had previously given the specific epithet Fistula to
the plant indicated but placed it under a different genus, which
genus happened to be Cassia.

Famity NAme.—The names of families are designated by the
name of one of their principal genera or ancient generic names
with the ending acez, e.g., Ranunculacee from Ranunuculus,
Malvaceae from Malva, etc. The following names, because of
long usage, are exceptions to the rule: Palma, Gramineae, Guttifera,
Umbellifera, Labiate, and Composite.

OrpDER NAmE.—Orders are generally designated by the name
of one of their principal families, with the ending ales, ¢.g.,
Rhamnales from Rhamnacea, Rosales from Rosacee. Suborders are
likewise designated, but with the ending—ineae, e.g., Malvinee
from Malvacee. Other older endings may, however, be retained
for these names providing they do not lead to confusion or error.

Crass NAME.—Names of classes, subclasses, divisions and sub-
divisions are designated from one of their characters by words of
Latin or Greek origin, some similarity of form and ending being
given to those that designate groups of the same nature, as Mono-
cotyledonee, Dicotyledonee; Archichlamydeew, Metachla-
mydex; Thallophyta, Spermatophyta; Gymnosperme,
Angiospermz. ©

In the case of Cryptogams, the use of old family names as
Algz, Fungi, Lichens, Musci, etc. is permissible for designating
groups above the rank of family.

For further details on botanical nomenclature, the student is referred to the
following articles: :

International rules of nomenclature revised by the International Botanical
Congress of Cambridge, 1930, 3 ed., Gustav Fisher, Jena, 1935.

An abridged version of The International rules, inaccurate in a few details, was
published as a supplement to volume 72 of the [English] Journal of Botany (1934).

ow, Mighie-S

Carter XVI

THE THALLOPHYTES

The Thallophyta or thallophytes constitute Division I of the
Vegetable Kingdom. As a group they represent the most
simple and most primitive plants. Many of their living repre-
sentatives like the bacteria, flagellates and slime molds possess
characteristics of both plant ane animal so that they are believed
to be near the first forms of life that existed on the earth. The
greater number of the members of this division have bodies
consisting of a thallus, a structure undifferentiated into roots,
stems or leayes. Under the microscope the thallus shows neither
tracheids nor tracheze. Some of the higher forms possess struc-
tures superficially resembling and functioning like the roots,
stems and leaves of higher groups. They reproduce asexually
or asexually and sexually.

They include the Schizophyta or fission plants represented by
the bacteria and blue-green algae, the Flagellata or flagellates, the
Myxomycetes or slime moulds, the Lygophycee represented by the
diatoms, pend silks and desmids, the Chlorophycea or green alge,
the Charophyta or stoneworts, the Phaeophycee or brown alge, the
Rhodophyceae or red algae, the Hyphomycetes or fungi and the
Lichens.

The evolutionary or phylogenetic relationship of the above
groups is so incompletely understood that the forms now to be
considered will be discussed under the long established physio-
logical groups of Algae, Fungi and Lichens.

THE ALG
The algze (sing. alga) are forms of thallophytes found on land
and in fre kish or salt water. The lower forms consist 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, while appearing like stems and leaves of higher
333

334 PHARMACEUTICAL BOTANY

plants, are merely sheets or cylinders of many celled filaments,
closely arranged and connected by protoplasmic connections.
They contain chlorophyH, and so can use the CO; and H.O in
the same manner as higher plants, ¢.g., in building up food and
providing for their own nutrition. In addition to chlorophyll,
some contain one or more additional pigments. Asexual repro-

Fic. 243.—Blue-Green Algae. A-C. Gloeocapsa polydermatica. A, single cell in
which division has begun; B, two daughter cells resulting from fission; the cell on
the left is dividing; C, colony of four granddaughter cells; D, fixed and stained cell
of Rivularia pisum. The net-like central body is the nucleus. X 3000. fy Pe,
Oscillatoria princeps. E, terminal portion of a filament; P, portion of a filament
showing a dead cell. X 540. G, as maller species of Oscillatoria. H-K, Nostoc.
H, portion of filament; A, heterocyst; I, portion of a filament with spores (sp);
J; spore; K, short filament produced by germinating spore. X 650. (A to C,
after Strasburger; D, after Haupt; H, I, F, K, after Bornet.)

duction is accomplished by motile or non-motile spores. Sexual
reproduction occurs in all but the simpler forms. The gametes
may be motile or non-motile. The female gamete is lodged
within an od6gonium or carpogonium both of which are unicel-

lular female sexual organs. Archegonia are absent in this group.
Crass I.—CyANopuyce#, THE BLUE-GREEN ALG

Plants which are sometimes termed blue-green alge. They
contain chlorophyll, a green \ pigment, and phycocyanin, a blue 2
pigment, a combination giving a blue-green aspect to most of the

¢

THE THALLOPHYTES 335

plants of this group. Some of them also possess phycoerythrin, a
red pigment. Their bodies are usually covered with a gelatinous"
envelope which may be in layers. In many of them the proto-
plast is differentiated into a peripheral pigment containing region
and a central colorless region. The unicellular forms reproduce
by binary fission while many of the filamentous forms reproduce
by the separation of one or more segments of the filament from
the rest of the body which move away and develop into new
individuals. These segments are called hormogonia. ‘They are
found everywhere in fresh and salt water and also on damp logs,
rocks, bark of trees, stone walls, in the roots of Cycads, etc.
Examples: Oscillatoria, Gleocapsa, Nostoc, Anabena, Rivularia, and
Spirulina.

GiaocapsA.—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 microscope
a mount of Gloeocapsa will be seen to consist of isolated proto-
plasts and groups of protoplasts, surrounded by concentric
_gelatinous 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 cell wall which undergoes mucilaginous modi-
fication producing thus the soft gelatinous envelopes which
encircle parent-, daughter-, grand-daughter- and even
great-grand-daughter-cells.

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 and composed of compactly arranged disc-
shaped cells which are all alike, excepting the terminal ones
which appear rounded off. Each filament is surrounded by a
gelatinous sheath. The filaments tend to be agglomerated in
thick felts or gelatinous masses and each possesses peculiar
oscillating and forward creeping movements. At the time of
reproduction the filament breaks up transversely into short
segments or hormogonia each of which, by fission occurring
among its cells, grows into a new filament.

336 PHARMACEUTICAL BOTANY

Nosroc.—Nostoc occurs on the damp ground bordering
streams or in slow bodies of water as greenish or brownish, tough,
gelatinous balls 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, imbedded in a gelatinous matrix, numerous
serpentine filaments, composed of spherical or elliptical cells
loosely attached to each other in chain-like fashion. Most of

Fic. 244.—Blue-Green Algae. A, Anabena azolle; B, Anabaena inequalis; C,
Cylindrospermum catenatum; h, heterocyst; s, spore; D-G, Rivularia bullata: D, Single
filament; FE, F, hormogonia; G, formation of hormogonia; H, Merismopedia. (A,
C, from Campbell’s “University Text-Book of Botany.” By permission of The
Macmillan Co., Publishers; B, from “A Treatise on The British Freshwater Algae’’
by the late G. S. West; revised by F. E. Fritsch, Cambridge Univ. Press, 1927;
D-G, after Bornet, from Engler and Prantl.)

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 filaments break apart on
either side of the heterocyst, setting free segments of cells (hor-
mogonia) which grow into new filaments.

Crass II.—CuHLoropHyce#, THE GREEN ALGAE

Green alge are unicellular (sometimes motile), filamentous,
colonial, or sheet-like water plants characterized by the presence
of solitary, or numerous chloroplasts in the cells which compose

THE THALLOPHYTES a5T

the thallus. These chloroplasts vary considerably in form, being
_in some cases spiral bands, in others star-shaped, in others like a
napkin ring, and in others granular. In the chloroplasts of most
“green algz are pyrenoids, which consist of a central crystalline por-
tion of protein (aleurone-like) surrounded by a starchy envelope
of variable magnitude. These are called starch centers and the
starch is frequently in the form of rounded, or angular grains.
The nutrition of these algz is autotrophic. ‘There is a defi-
nite nucleus present, but in the ccenocytic forms the nuclei
may be many within the confines of the cell wall. The motile
cells have two (except in Oedogoniacee and by fusion in the
Vauchertacee) to many cilia, as likewise some of the reproductive

cells. No cilia occur in the conjugate alge. Reproduction is by_

Cell division, the formation of zodspores (motile cells), by zygo-
spores produced by conjugation, by egg cell and sperm cell union
(odspores) odgamous reproduction. Green algz live mostly in
fresh water. Some live in sea water and a few in brackish water.
Some are associated with fungi to form lichens. .

1. ORDER PROTOCOCALES OR ONE-CELLED GREEN ALGA.—
This order contains nearly all of the one-celled green algze
excepting the diatoms and desmids. It also contains a variety
of many celled forms, the cells of which are devoid of cilia.

FamiLty PLEuRococcACEa.—Pleurococcus vulgaris [Fig. 245 (2)]
is a one-celled green alga, millions of which, living together in
colonial fashion, constitute the so-called “green stain’ that is
common on the north sides of tree trunks, stone walls and fences.
Each organism consists of a protoplast surrounded by a cell wall
of cellulose. ‘The protoplast contains a chromatophore (capable of
division into 2 or more chromatophores), 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 granddaugh-
ter-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 VotvocaLes.—This order comprises free-swim-
ming aquatic forms whose vegetative cells are bi-ciliated, green,
more or less spherical or compressed. Some of the organisms

oo

338 PHARMACEUTICAL BOTANY

like Spherella [Fig. 245 (3)] and Chlamydomonas consist of single
cells bearing a pair of cilia, while others like Pandorina, Eudorina

Fic. 245.—Various types of Green Algee. Ulothrix (1); Pleurococcus (2); Spharella
(3); Cosmarium (4); Rhaphidium (5); Vaucheria (6); Rhizoclonium (7); Hydrodictyon (8);
Oedogonium (9); Botrydium (10); pgnema (11); Draparnaldia (12); Zygnema during
conjugation (13).

and Volvox show varying degrees of colony formation. Repro-
duction sexual or asexual.

ae Meare San

THE THALLOPHYTES D330

The organisms of this order are sometimes classified
with the Flagellates, a group having characteristics partly like »
plants and partly like animals and represented by Euglena and
many other unicellular, aquatic forms possessing flagella or
cilia.

_ Volvox globator, a typical representative of this order, is found
in fresh water pools as a tiny, hollow, spherical, green colony
about 19 to 19 of aninchin diameter. When examined under
the microscope (Fig. 246), it is found to consist of hundreds of
green, more or less spherical cells, united by fine strands of
cytoplasm (protoplasmic bridges), the whole being enveloped
by a gelatinous sheath. The peripheral cells are provided with
cilia, in order that the colony may rotate and roll through the
water. In a young colony, all of the cells are alike, each con-
sisting of a mucilaginous-like cell-wall enclosing cytoplasm, a
nucleus, a chloroplastid and often a red pigment spot. Ina
mature colony, however, throughout the greater part of its
existence, two kinds of cells may be discerned: small, sterile,
vegetative cells that do not divide and from 10 to 12 larger
vegetative ones that divide to form new colonies. The latter
slip inward below the level of the smaller cells and through
repeated divisions form a number of ciliated cells joined by cyto-
plasmic threads, which in reality is a miniature colony. This
then escapes to the exterior through the rupturing of the gelati-
nous wall of the old colony.

During autumn of the year, certain of the ordinary cells
undergo differentiation, some to form sperm cells, others, egg-
cells. When about three times the size of the ordinary sterile
cells, the sperm cells divide repeatedly to form a cluster of
elongated secondary cells [Fig. 246 (1a, a® and 5)] each of which
contains an orange pigment, a red corpuscle, has an elongated
beak and bears a pair of flagella (lash-like processes). The
cluster in time separates into motile antherozoids [Fig. 246
(6, 7)] which finally escape into the cavity of the volvox sphere
through rupture of the investing wall. The flask-shaped egg cells
(16, 6) increase greatly in size without dividing. Each shows
vacuoles, then becomes filled with a dark green pigment, becomes
spherical and acquires a gelatinous envelope. It then passes

340

Fic. 246.—Volvox globator. Mature colony in center (1); sexual cells (2a);
endochrome of primary cell has resolved itself into a cluster of secondary cells
(1a, a* and 5); antherozoids (6, 7); bundle separated into component antherozoids
in cavity of primary cell (1a*); breaking of wall of primary cell showing escape
of antherozoids into cavity of volvox sphere (1a*); egg cells (14, 6); flask shaped
germ (egg) cells with large vacuoles in protoplasm (147, 57); globular egg cells :
prepared to pass into cavity of volvox sphere (5%); antherozoids collected about
egg cell (3); odspore (4). (Carpenter.)

THE THALLOPHYTES 341

into the cavity of the sphere where it is surrounded by numerous
antherozoids (3) and is finally fertilized.

The product of this fertilization is an odspore or zygote (4) which
ere long becomes covered with an internal smooth membrane and
a thicker external spinose coat. The chlorophyll within then dis-
appears and starch and a reddish- or orange-colored oil make
their appearance. Up to 40 of these odspores have been
observed in a single volvox sphere. Not long after the formation
of these odspores the whole parent colony breaks up and the
odspores fall to the bottom of the pool to pass the winter season.
As early as February each odspore germinates to form another
volvox colony, which repeats the life cycle described.

Fic. 247.—Vaucheria terrestris, a siphon alga. anth, antheridium (empty); 0, odgonia.
(Gager.)

3. ORDER CONFERVALES.—In this order are included a
variety of green filamentous and membranous forms some of
which show sexual reproduction.

Famity ULoTHRIcACE&.—Ulothrix zonata, [Fig. 245 (1)] 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 is elongated and serves
as an attachment structure. Each cell consists 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 (chloroplast) lies next to the cell wall and
contains pyrenoids or starch-forming centers. The filament

342 PHARMACEUTICAL BOTANY

grows in length by the fission of its various component cells.
After attaining a certain size it reproduces either asexually
or sexually. Asexual reproduction takes place by certain cells
becoming altered in their protoplasmic contents, through divi-
sion, to form rounded or pear-shaped zodspores. Each zodspore
contains a red pigment spot and bears four cilia (protoplasmic
outgrowths). The zodspores 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 produc-

tion of many gametes, in cells of the filament. These resemble
the zodspores 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 fuse with one another. The product of the fusion of
each pair of these like gametes is termed a zygospore. The zygo-
spore 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 zodspores which,
escaping from the cell, swim about for a while and finally,
attaching themselves to objects, grow into filamentous Ulothrix
organisms.

Closely related to Ulothrix is the sea-lettuce, Ulva lactuca (Fam.
Ulvacee), a membranous, leaf-like form found attached to rocks
along the shores exposed to the action of the waves.

4. ORDER ConjuGALEs.—To this order belong the desmids
and pond scums which are distinguished from other green alge
by presenting no motile stages in their life histories. No zoé-
spores are, therefore, produced by members of this order. T hey
are all of fresh-water habit and reproduce by conjugation.

Famity Desmipace#.—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 [see
Figs. 245 (4, 5)] is characterized by being composed of two like
halves frequently separated from each other by a constriction
called the isthmus. In each half there is a chromatophore contain-
ing pyrenotds. The nucleus is found in the isthmus. Reproduc-

THE THALLOPHYT ES 343

tion is accomplished either asexually by fission or sexually by
‘ conjugation.

FamiLy ZYGNEMACE&.—This is a family of pond scums includ-
ing the well-known genera, Spirogyra and Zygnema [Fig. 245 (11,
. 13)].

Fic. 248.—Spirogyra sp. A, terminal portion of vegetative filament; B, stages
of scalariform conjugation; C, preparation for lateral conjugation; D, zygospores
formed by lateral conjugation. (Gager.)

Spirogyra or Brooksilk is a filamentous organism found sus-
pended 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 ter-
minal cells having rounded extremities. Each cell has a cell wall

344 PHARMACEUTICAL BOTANY

of cellulose within which is to be found a thin film of ectoplasm.
One or more spirally shaped chromatophores 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 Sperogyra 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

Fic. 249.—Spirogyra longata. Portion of a monoecious filament showing two
adjacent cells in the early stages of conjugation. This type of conjugation is
called lateral. Ladder-like or scalariform conjugation of dioecious filaments is shown

in figure 248. Note the pulsating or contractile vacuoles. (From Gager, after
Lloyd.)

occur every night. In this way the filaments are rapidly made
longer. Sooner or later they break and in this way the plant
multiplies. |

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 in the vessel in which it is growing to slowly evaporate.
Two filaments arrange themselves side by side, and the cells lying
opposite each other undergo internal changes so as to form gametes
or sexual cells. Each gamete-containing cell is called a game-
tangium. Each gametangium protrudes a process or lateral
branch. The tips of these fuse upon meeting each other and
form a conjugation tube or connecting tube. ‘Then the protoplast
(gamete) of one cell passes over and coalesces with that in the cell

THE THALLOPHYTES 345

opposite. ‘The result of the process is a new cell called a zygospore
or zygote. ‘This method is called scalariform conjugation. The
zygospore forms a thick cell wall about itself. It is set free by
decay of the walls of the old cell and falls to the bottom of the

Fic. 250.—Vaucheria sessilis. A, Terminal portion of a filament showing the
appearance of a partition wall separating the newly formed sporangium from the
rest of the filament. B, Same, after escape of zodspore (<). C, A portion of
the margin of the zodspore. D, Young plant developed from a zoéspore with
rhizoids (r). (From Strasburger, Noll, Schenk and Schimper; A, B, &, after Gotz; D,
after Sachs from Altmann’s Alga; C, after Strasburger.)

water, there to undergo a resting stage until favorable conditions

for growth arise.
Professor F. E. Lloyd has shown that in Spirogyra longata, con-
jugation takes place between adjacent cells of the same filament.

346 PHARMACEUTICAL BOTANY

This second type of conjugation is called Jdateral conjugation.
In this case a protrusion of the cell wall of the filament occurs
opposite the partition wall of adjacent gametes forming a
channel of communication between the gametes. Pulsating or
contractile vacuoles, such as have been described in Amoeba
appear in the protoplast and upon contraction propel the gamete
in one cell cavity (supplying gamete) over into the cell cavity of the
other gamete (receiving gamete). ‘The fusion of these gametes
results in the formation of a zygospore.

5. ORDER SIPHONALES (SIPHON ALG&).—This group is char-
acterized by the peculiarity that the organisms constituting it
possess protoplasm containing myriads of nuclei within a common
filament or cell cavity not segmented by cell walls. The term
cenocyte has been given to such structures which consist of a many-
nucleated mass of protoplasm surrounded by a cell wall. Some
of the siphon alge reproduce by zoéspore formation, others by
conjugation as well as zodspore formation while Vaucheria, the
‘green felt,” stands out alone in reproducing both by the forma-
tion of a single compound zoéspore and by the production also of
oégonia and antheridia with resultant fertilization (see Figs. 245
(6) and 247).

The various species of Vaucheria occur in water or moist soil
where they form dense mats of tubular filaments. Each filament
is unicellular until the time of reproduction, containing cytoplasm
in which are imbedded thousands of nuclei as well as numerous
chloroplasts devoid of pyrenoids, and small drops of oil. At the
time of asexual reproduction a transverse wall is laid down near
the tip of the filament cutting off a terminal cell or zodsporangium
in which a many ciliated compound zoéspore is formed. This
escapes by a break in the end of the sporangium, swims about for
a brief time and then germinates, sending out one or more
Vaucheria filaments. In some species a colorless simple or
branched rhizoidal cell or organ of attachment is developed by
the filament where it contacts a solid substance.

6. ORDER CHARALES (THE STONEWORTS).—FAMILY CHAR-
ACE&.—The highest group of green algze, possessing forms which
are differentiated into stem-like and leaf-like structures and
rhizoids. Examples: Chara and Nitella.

THE THALLOPHYTES 347

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. _ Its
plant body consists of a many noded (jointed) axis or so-called
“stem”? which bears whorls of slender, green, dwarf branches,

- Apical Cell

Dwarf Branch-~4

8
E
~~~ Internodal Cell

Fic. 251.—Chara fragilis. Wabit and structure of certain vegetative parts.
A, portion of plant showing long branches (a), dwarf branches (6), and rhizoids.
B, longitudinal section through growing tip. CC, cross section of internode, showing
internodal cell and surrounding cortex. (From Mottier, adapted from Campbell .)

sometimes called leaves, at its nodes. The dwarf branches, like
the main axis, possesses nodes and internodes. The nodes of
these produce short abortive “leaves.” Filamentous rhizoids
arise from the lower node. Long branches are also found issuing
from the axils of some of the dwarf branches at the nodes which
repeat the structure of the main axis. Reproduction is either

348 PHARMACEUTICAL BOTANY

asexual or sexual. Asexual reproduction 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 reproduc-
tion is effected through the formation of odgonia (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 of dwarf
branches. The oval oégonium is
borne above the spherical anther-
idium. The oégonium develops
within itself a large ovum or egg.
The antheridium produces within
its wall numerous motile sperms.
Upon the maturation of the an-
theridium the sperms are liberated
into the water, and, propelled by
iS their cilia, find their way to the
Fic. 252.—A, Portion of the axis of °0§0nia which they enter, the one
Chara fragilis, a stonewort. s. Oogoni- best adapted fusing with the egg
um (one-celled female sexual organ). in each case and’ fertilizing it.
Abortive “leans” Br ie oom The resultant cell is called the
“leaves.” c. Crown ofodgonium, B. 00Spore or zygote. The zygote
Young stage of the same. r. Corti- undergoes a resting stage within
Se sts 6. Antberiiien, sh Oligo 1h. odgonium. The odgonium
nium. (From Small, after Sachs.) z

with the enclosed zygote falls from
the plant and sinks to the bottom of the water. The zygote later
germinates as a proembryo or juvenile form of Chara plant. The
proembryo consists of a small shoot of two nodes and a long
rhizoid. From the upper node of this proembryo, the adult form
of Chara arises as a side shoot, and from the lower node rhizoids
are formed.

Crass ITI.—BacitLariE& or DIAToms

This class comprises 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

THE THALLOPHYTES 349

unicellular, they frequently are united in colonies. They all
possess chromatophores containing chlorophyll 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

Nodule — fe:

ia

mia
ett

Nodule -

Orn

Et =" Nucleus

Fic. 253.—Two views of the diatom Pinnularia. A, valve view; B, girdle view,
showing cell contents. (After Strasburger’s Text Book of Botany, Macmillan Co.
publishers.)

a pill box. The valves, which are beautifully sculptured with
intricate patterns of lacework or of beading, 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, stellate, wedge-shaped, 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

350 PHARMACEUTICAL BOTANY

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.
One of 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 24 to 40 feet in depth and extends
for many miles. Another deposit at Lompoo, California, is
claimed to cover an area of 12.
square miles with an average
depth of over 100 feet. Many
of the diatomaceous earths are
useful as absorbent and polish-
ing powders.

| Terra Silicea Purificata (Puri-
fied Siliceous Earth) is a powder
consisting of the frustules and
fragments of diatoms which has
been purified by boiling with
diluted hydrochloric acid,
washed and calcined.

Many diatoms exhibit a

Fic. 254.—Two mag Diatoms. gilding or jerky motion. This
To left, Diatoma vulgare, a, side view of 15 stated to be caused by the.

frustule; 4, frustule undergoing division. streaming movement of their
To right, Grammatophora serpentina: a, cytoplasm.

front and side views of single frustule; : bs
b, 6, front and end views of divided Diatoms exhibit two modes

frustule; c, frustule about to undergo Of reproduction, viz., fission
division; d, frustule completely divided. and formation of an auxospore.
Bet CORR The more common method is
that of fission but this: is peculiar for these plants. Thecell-contents
within 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

THE THALLOPHYTES 351

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 forma-
tion of succeedingly smaller and weaker individuals. This
process, however, goes on only for a certain number of genera-
tions 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
contributed to itsformation and fees ><
is endowed with renewed vigor =
for growth and division.

Cxiass [V.—PHAOPHYCE,
‘THE Brown ALG

Mostly marine forms show- hn \
ing great diversity in the form Fig. 255.—Licmophora flabellata, a dia-
of their vegetative bodies. pa nga ee ee
They occur for the most part eA (4 sehag ohne a
in salt water between the high
and low tide marks. Some of them are always submerged.
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 a hundred feet or more in length
as, for example, Macrocystis, a giant kelp, which grows in the
Pacific Ocean off the coast of California. They all contain the
brown pigment called phycophein or fucoxanthin and the green pig-
ments, chlorophyll, both of which are present in their chromato-
phores. A yellowish pigment called phycoxanthin has also been
isolated from some of the species. Many of the kelps and rock-

352 PHARMACEUTICAL BOTANY

weeds belonging to this class have long been sources of iodine,
potash and sodium.

There are four orders of brown algae, viz., the Pheosporales,
Laminariales, Dictyotales and the Fucales. Types of three of these
will be considered. :

A FrtaMentous Brown ALGA, Ecrocarpus SILICULOSUS.—
Ectocarpus, a representative of the Order Phaosporales, occurs as

Fic. 256.—Fossil diatoms: a, a, a, Gaillonella procera and G. granulata; 6, 6,
Surirella plicata; c, Surirella craticula; d, d, d, Gaillonella (Melosira) biseriata (side view);
e, Gomphonema gracile: f, Cocconema fusidium; g, Tabellaria vulgaris; h, Pinnularia dacty-
lus; i, Pinnularia nobilis; k, Surirella caledonica; 1, Synedra ulna. (After Carpenter.)

tufts of branching filaments, each of whichis many celled. ‘These
tufts are found on eelgrass or other algz 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 reproduction 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 irreg-
ular chromatophore containing a brown pigment called phyico-
phein. From certain cells of the filament spherical to elliptical

THE THALLOPHYTES 353

sporangia (spore cases) arise, which are unicellular. They con-
tain numerous biciliate zodspores, 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 chambers have resulted, which are
called plurilocular gametangia. Each cell of a_ plurilocular

Fic. 257.—Ectocarpus siliculosus, a filamentous brown alga. A. Unicellular
sporangia, one containing zodspores, the other empty; z, a zodspore. 8B. Pluri-
locular gameangia, the larger mature, the smaller still showing the outlines of the
original cells in the branch from which it arose. C, Union of two gametes to form
the zygospore.—C, after Oltmanns. (From Bergen and Davis Principles of Botany.)

gametangium contains a gamete or sexual cell, which resembles
in many details a zoéspore. When the gametangium matures
these gametes are discharged into the salt water. They fuse
together in pairs and form zpgospores. Each zygospore under-
goes a resting stage and upon the advent of favorable conditions
develops into a new Ectocarpus filament.

Tue Keips.—To this group (Order Laminariales) belong the
largest living representatives of the Alga. _ They are represented

Oe ee ee ee ee ee. ee a es ee SA ee ee ee

354 PHARMACEUTICAL BOTANY

by the Devil’s Aprons or Laminaria and the Sea Palms (Postelsia)
which grow on rocks along the North Atlantic coast and by the
giant kelps Macrocystis (Fig. 258) and Nereocystis (Sea Otter’s
Cabbage) found along the Pacific coast of the United States.
Laminaria digitata is an abundant kelp off the coast of New
England where it grows attached to rocks by a holdfast consisting
of a whorl of rootlet-like structures. From this arises a solid
cylindrical stipe bearing on its end an oval or lance-shaped

Fic. 258.—A giant Kelp (Macrocystis), one of the Brown Algz. (From U.S.D.A.
Bur, Soils.)

blade, the latter becoming split into finger-like segments at its

summit. Postelsia or the Sea Palm has a holdfast of several —

whorls of stubby rootlet-like structures and a rigid stipe about a
foot in length, the latter terminating in a number of lanceolate
blades.

Because of their rich content of potassium salts, the kelps are
used extensively as fertilizer. They also contain a fair amount of
iodine and represent one of the chief sources of this drug.

Tue Rockweeps.—To this group (Order Fucales) belong the
brown seaweeds which cover many of the rocks along our North

tial ita

%
.

THE THALLOPHYTES 355

Atlantic shores and mussel beds of our inlets. They all possess
floats or bladder-like structures con-
taining carbon dioxide and oxygen
which aid in buoying up the plants
in the salt water when submerged.
Common forms occurring along the
eastern coast of North America are
Fucus, Ascophyllum and Sargassum.
The last named is called the Gulf
Weed because it is often found float-
ing freely in the gulf stream. Large
numbers of these plants often ac-
cumulate in quieter areas of the
ocean where they form the so called
**Sargasso Sea.”

Fucus Vesicutosus (THE Biap- ¥ pity
DER WRACK).—This form, a brown ea) voy
alga, occurs as a flat thallus, which Ky
forks repeatedly, a type of branching
called dichotomous. It grows near
the surface of sea water, attached to
rocks or to mussels along banks 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 ES Se ok sae oe
with minute warts, each having an North Atlantic. A, The simple
aperture at its summit which leads type of Laminaria, some of whose
into: aH eneieee eeuis acthads the SPete Ere: te be thirty oF more

3 : : feet ‘long. B, The digitate type
receptacle. This cavity is called (jominaria digitata) which is never
the conceptacle. Within the concep- very long, but is broad at the base.
tacles of the male plants of thespecies (from Bergen and Davis Principles
the antheridia, or male sexual organs, —
are formed while in the conceptacles of female plants the odgonia or
_ female sexual organs, are produced. The conceptacles also con-

356 PHARMACEUTICAL BOTANY

tain numerous, branching, protective filaments called paraphyses,
which arise from the cells lining the cavities. ‘The antheridia are
found as outgrowths of some of these paraphyses and produce

Vegetative Branch

‘> r cH 3
Sgt —- Fruiting Branch

:

SS
We,
Hy.

aga

lal ae Ree 38
ACS ESA)
On Fanty

By
eae
nese

ay
13
aNd
vt

)

2

ee

is
OR
P aw,
uty.
(Le

1)
ui
UL

Fic. 260.—Fucus vesiculosus. A, portion of plant showing habit. B, diagram
of cross section of thallus; r, outer rind; m, loose inner pith or medulla. C, cross
section of a fruiting receptacle. D, longitudinal section of a growing point taken
at right angles to the flat surface, showing the pit at the bottom of which is the
apical cell. (A, B, C, after Mottier; D, from King’s chart.)

sperms or male sexual cells. The odgonium is a large, globular,
stalked cell and produces eight eggs, each of which is a female
sexual cell. The eggs and sperms 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

THE THALLOPHYTES

that of the egg to form an odspore. The odspore at once develops
into a new Fucus plant.

The drug Fucus consists of the dried ‘thallus of Fucus vesiculosus,
Fucus serratus, Ascophyllum nodosum or Halidrys siliquosa (Fam.
Fucacez). Fucus has been used in the treat-
ment of obesity and goiter.

T

Cass V.—RHODOPHYCEA, THE RED ALG

A greatly diversified group comprising the
majority of marine algz but represented also ¢~-%
by some fresh-water forms of which Batracho- Seat.
spermum and Lemaneaareexamples. Themarine Fic. 261.—Fucus
red alge are gencrally found at or just beyond en
the low water mark. Their vegetative bodies , conceptacle, T,
vary from simple branching filaments through cellular _ thickness.
all gradations to forms differentiated into hold- ange aa (After
fasts and branching stem- and leaf-like struc- van
tures. 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
superficially resemble the tissues of plants of higher domain.
Their color may be red, purple, violet, or reddish-brown or even
green. The red color of most of them is due to the presence of
phycoerythrin, a red pigment, which is found in the chromatophores
with, but frequently masking, the chlorophyll. Phycocyanin, a
blue pigment occurs in the chromatophores of some red algae.
In most of them several chromatophores occur in each cell.
These are usually small and oval in shape.. Starch is not formed
but rather another carbohydrate staining red with iodine.

Reproduction is asexual by spores or sexual by spermatia
functioning like sperms and by eggs. The spores and gametes
are devoid of cilia.

SuB-cLAss FLoRIDE#.—To this group belong plants which
consist of branching filaments or filaments closely bound by an
intercellular substance and plasmodesma into dichotomously
branched, cylindrical or ribbon like thalli. Some of them have
their bodies composed of cells all alike while others show differ-
entiation into an outer chlorenchymatous and an inner conduc-

358 PHARMACEUTICAL BOTANY

tive tissue. Several rounded or flattened chromatophores occur
in each cell.

Reproduction may be accomplished vegetatively by frag-
mentation and by budding. Asexual reproduction is commonly
by means of ¢etraspores which are formed in tetrasporangia that
occur on the ends of lateral filaments or in subepidermal cells.

Pra :

a4

par

an

Fic. 262.—Fucus vesiculosus. ‘Transverse section through outer region of a portion
of the receptacle of male plant showing a male or antheridial conceptacle and
adjacent tissues. X 50. an, antheridium; os, ostiam or opening of conceptacle
through which paraphyses are seen protruding; far, paraphyses protruding from
another conceptacle, the latter concealed by surrounding tissue; b, compact pseudo-
parenchyma; fs, loose pseudoparenchyma with mucilaginous cells:

Sexual reproduction is by union of the nuclei of Spermatia and ova.
The female sexual organ is called a carpogonium, the male sexual
organ an anthertdium. The carpogonium consists of a basal cell
which contains the ovum and a terminal filament which ends
in a hair-like portion called the trichogyne. The contents of each
mature antheridium as it emerges from the organ is termed a
spermatium. It is a spherical, non-motile male cell. The sper-
matia cling to the trichogyne and the nucleus of a spermatium

agp aie ieel ae ae

THE THALLOPHYTES 359

enters it and passes to the ovum in the basal cell where fertiliza-
tion is effected. The zygote forms filaments called gonimoblasts
which give rise to carpospores or fruits called cystocarps which con-
tain carpospores.

Chondrus crispus and Gigartina mamillosa (Fam. Gigartinacee)
when dried and bleached, constitute the official drug CHoNDRUS,

& ™
a \ SAT "

cna TE at
Se NR

Fic. 263.—Fucus vesiculosus. Transverse section through the outer region of a
portion of the receptacle of a female plant showing an odgonial conceptacle and
adjacent tissues. 50. 0, odgonium within a conceptacle; p, paraphyses; ¢f,
epidermis; co, cortex; ps, pseudoparenchyma of irregular cells having mucilaginous.
walls.

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

apy ia Si RO a, ee eal a im ies "Pk a oa AN ae aia ie al

360 PHARMACEUTICAL BOTANY

elliptical and embedded in the thallus near its surface, whereas in
Gigartina they are ovate and borne on minute stalks which

Fic. 264.—Fucus vesiculosus. Fertilization. A, egg approached by biciliated
sperms; B, sperms attached to egg and ee it prior to fertilization (highly
magnified). (After Thuret.)

project outward from the surface of the segments. Upon the
ripening of these structures the spores are discharged into the sea

Fic. 265.—Chondrus crispus. (After Strasburger.)

water. These sooner or later germinate into new Chondrus or
Gigartina organisms.

:
;
:

THE THALLOPHYTES 361

The dried mucilaginous substance extracted from Gelidium
corneum or other species of Gelidium (Fam. Gelidiacee) and closely
related red algz growing in the sea along the eastern coast of
Asia and along the coast of California constitutes the drug, AGAR.

Fic. 266.—Gigartina mamillosa. Ss, fruiting body consisting of short projection
bearing tiny sporangia. (After Strasburger.)

Agar is a most valuable ingredient in culture media as well as a
laxative.

Rhodymenia palmata or Ir1sH Dutse 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.

CHAPTER XVII
THE FUNGI

The fungi or hyphomycetes constitute a great assemblage of
thallophytes which are characterized by the total absence of
chlorophyll. The members of this subdivision, “therefore,

possess no independent power of manufacturing food materials
such as starches, sugars, etc. from CO, and H2O. Conse-
quently they are either parasites, depending 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 sapro-
phytic habit. Some of the fungi like the bacteria and the yeasts
are unicellular while others like the molds and mildews, etc.
have bodies in the form of thread-like filaments called hyphe
(sing. hypha) which are frequently branched. A mass of inter-
tangled hyphz is called a mycelium. The hypha may be uni-
cellular or multicellular. Fungal hyphe cither ramify through
decaying matter or invade the tissues of living organisms and
derive nourishment therefrom. In the cases of parasitic fungi
like the rusts, etc., absorbing connections are present as hyphal
outgrowths which penetrate into the cells of the host plant.
These are called haustoria. In the higher forms the hyphz
become consolidated into false tissues, and assume definite
shapes according to the species. Of this character are the
fruiting organs which constitute the above ground or exposed
parts of Puff Balls, Cup Fungi, Bract F ungi, Mushrooms,
etc.

There are three classes of fungi, viz.:

Class I Schizomycetes (bacteria)

Class IIT Myxomycetes (slime moulds)

Class ITI Eumycetes (true fungi)

362

THE FUNGI i SD

Crass I.—ScuizomyceTes (BACTERIA)

The bacteria, also called “schizomycetes”’ or “fission fungi,”
are now regarded by leading systematists as the lowest types of
plant life extant. Among them are to be found the smallest
living organisms known to science.

BACTERIA are minute, unicellular, colorless, rarely weakly red
or green colored, vegetable organisms destitute of chlorophyll.
They serve as agents of decay and fermentation and are fre-
quently employed in industrial processes. The curing of certain

Animals
Non-
chlorophyll-
bearing

plants

Chilorophyll- Disintegration
went by saprophytes
plants & parasites

Inorganic
matter
(Chemical
elements and
compounds)

Fic. 267.—The organic cycle. (After Gager.)

drugs and tobacco, the tanning of leather, the manufacture of
vinegar, buttermilk, cheese and indigo, the making of ensilage
and sauerkraut, and the retting of flax and some other textile
fibers are all dependent upon fermentative processes induced by
bacteria. Their form varies from spherical, rod-shaped, spiral
to filamentous, the filamentous types being frequently surrounded
by a capsule or sheath.

"The bodies of most bacteria possess nitrogenous cell walls
representing outer plasma membranes, although hemicellulose
has been found in the walls of many and mucilage occurs as a

364 PHARMACEUTICAL BOTANY

gelatinous sheath surrounding the primary wall in some forms.
Cytoplasm occurs within the cell as well as chromatin, but the
latter is usually diffused in fine particles through the cell proto-
plasm and never has been demonstrated to be condensed into a
definitely organized body 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 becomes a
separate and independent organism. Sexual reproduction has
not been conclusively proven, although suspected by some
investigators. Bacteria occur everywhere but are relatively few
on high mountain peaks and beneath 3 meters of natural soil.
They have even been found in the hot geyser springs where no
other form of life can exist (Thermophilic bacteria).

According to the various phenomena they produce, they may
be classified as follows: (a) Zymogens producing fermentation; (b)
Aerogens producing gas; (c) Photogens producing light; (¢) Chromo-
gens producing color; (¢) Saprogens, producing putrefaction; (/)
Pathogens, producing disease; (g) Thermogens, producing heat.

FERMENTATION, DECAY AND PuTREFACTION.—Fermentation is
the process which consists of the breaking down of organic sub-
stances through the action of enzymes produced by living organ-
isms. ‘The organic substances fermented are various, including
carbohydrates, proteins and fats. Decay is the fermentation of
proteins in the presence of oxygen. PurTrEractTIon, although
frequently used in the sense of the decomposition of proteins, is
more properly applied to the anzrobic fermentation of proteins,
many of the products of which possess an offensive odor. When
carbohydrates undergo fermentation, the products are useful.
Bacteria cause considerable spoilage of foods and drugs owing to
their power of producing enzymes.

BacTERIAL CuLTuREs.—Because of their minute size (a
space the size of a pinhead may 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 differentiated. These cultures are obtained
by growing the microorganisms on sterile artificial culture media
usually in an incubator, the temperature of which is carefully
regulated to suit the requirements of the organisms grown. The

ee ge eS Oe ee ee

Pee eee ar NS) ici

THE FUNGI 365

culture media employed are both solid and liquid. Beef broth
or nutrient bouillon forms the basis of most of the culture media.
To this either agar agar or gelatin are added to solidify the
medium (solid media) and various chemical substances and
sugars for the purpose of studying the physiological character-
istics of the various forms. Formulas for the preparation of
various culture media and directions for the isolation of bacteria
will be found in the National Formulary VI and in works of
bacteriology and mycology.

Forms oF BAcTERIA.—Bacteria occur as spheres, straight
rods or cork-screw like (bent rods) forms (lower bacteria) or as
filaments (higher bacteria). The spherical shaped form is

Fic. 268. Fic. 269.
Fic. 268.—Monotrichous bacteria. Microspira comma. (After Migula from
Schmidt and Weiss and Marshall.)
Fic. 269.—Lophotrichous bacteria. Pseudomonas syncyanea. (After Migula
from Marshall.)

termed a micrococcus or coccus (plural-micrococct or cocci). The
straight rod form is called a baczllus (plural-bacill:). The cork-
screw shaped form is known as a spirillum (plural-spirila).
Mortury.—A large number of bacteria possess the power of
independent movement when observed in a fluid medium which
will permit of their locomotion. This true movement should be
distinguished from the so-called “Brownian movement,” a quivering
or oscillating phenomenon shown by minute particles or bodies
when freely suspended in a suitable fluid. True movement is:
due to the activity of the living cell, whereas Brownian movement
is a purely physical phenomenon due to the bombardment of
particles, dead bacteria or non-motile bacteria, by the molecules
of the menstruum. In true movement the individual form moves
from one part of the microscopic field to another, whereas in

366 PHARMACEUTICAL BOTANY

Brownian movement the individual forms do not change their
relative positions to any extent.

The power of locomotion which certain bacteria possess is due
to the possession by these forms of fine protoplasmic extensions
from their bodies which are thought to pass from the cytoplasm
through the cell wall into the surrounding liquid medium.
These are called flagella or cilia. ‘They are endowed with the
power of contractility, and through their vibratory motion the
organism is enabled to propel itself through the fluid medium.
The great majority of bacteria possessing flagella are the bacilli
and spirilla. These flagella are rarely seen without special
methods of treatment and staining. :

Fic. 270. Fic. 271.

Fic. 270.—Lophotrichous bacteria. Spirillum rubrum. (After Migula from
Schmidt and Weiss and Marshall.)

Fic. 271.—Peritrichous bacteria. Eberthella typhi. (Bacillus typhosus.) (After
Migula from Schmidt and Weiss and Marshall.)

The distribution of flagella varies in different forms of
bacteria. Some possess a single flagellum at one pole and are
called monotrichous. When a flagellum occurs at each pole, they
are termed amphitrichous; when a tuft occurs at one pole, lopho-
trichous; and when they occur over the whole cell, as in the case of
the typhoid bacillus, peritrichous.

The motility or non-motility of bacteria is best determined
through the examination of a hanging drop.

Nurrition.—Most bacteria derive their sustenance from
organic substances. Those which obtain their nourishment from
dead organic substances are termed saprophytic bacteria and those
which obtain their nourishment from living organic material,
parasitic bacteria. A number of the saprophytic forms will
thrive both on living and dead organic matter and so are called

THE FUNGI 367

faculative parasites, while many others, like the majority of the
common soil and water bacteria, are restricted to dead organic
matter and so are obligate saprophytes. Most of the parasitic
bacteria are faculative saprophytes, since they can be grown upon

artificial culture media. The leprosy bacillus is one of the few

examples of parasitic bacteria which has resisted attempts to
grow it upon artificial culture media and so is an obligate parasite.

NitTRIFICATION.—There occur a group of soil bacteria called
nitrifying bacteria which thrive in the presence of simple inorganic
salts and in the entire absence of organic matter. Some of these
(Nitrosomonas and Nitrosococcus) oxidize the ammonia of the soil
to nitrous acid while others (Nitrobacter) convert the nitrous acid
to nitric acid. The nitric acids reacts with alkaline soil con-
stituents to form nitrates which pass into solution in the soil
water and are utilized by higher plants in their growth. This
process is called nitrification.

‘DENITRIFICATION.—Denitrification is a process carried on by
denitrifying bacteria including Pseudomonas fluorescens, Ps. pyocyanea,
and the colon and typhoid bacilli. Many of the denitrifying
organisms flourish in “‘sour”’ soil, in the absence of air, and obtain
oxygen for their respiration by reducing the nitrates and nitrites
to gaseous oxides of nitrogen or to free nitrogen, liberating the
gas into the atmosphere.

GrowtH Propucts.—The products resulting from bacterial
growth include pigments, enzymes, acid and alkali, putrefactive
substances, ptomaines and toxins.

REPRODUCTION, —Bacteria multiply and reproduce them-
selves by ¢ 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.

MorpHo.ocy DuE To CLEAVAGE.—According to limitations
imposed by cleavage directors, the spherical forms called micro-
cocci or cocci assume a chain appearance, or a grape cluster-
like appearance, or an arrangement in packets or cubes having
three diameters. These aggregations are the basis for the follow-
ing genera of spherical-shaped bacteria:

Diplococcus (plural, diplococci), occurring in twos, as organisms
producing meningitis, pneumonia and gonorrhea.

368 PHARMACEUTICAL BOTANY

Staphylococcus (plural, staphylococci), from a Greek word refer-
ring to the shape of a bunch of grapes, as the organism producing
golden pus.

Streptococcus (plural, streptococci) from a Greek word meaning
chain-shaped, as the scarlet fever organism.

Sarcina, package-shaped or cubical, as the Sarcina lutea, a
common saprophyte.

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 morpho-

2. ee > Be
s oe

Fic, 272.—Division forms of micrococci. a, Diplococcus, perfect form with
flattened opposed surfaces (gonococcus), lanceolate form (pneumococcus); b, strepto-
coccus; ¢, consecutive fission yielding a tetrad; d, sarcina form resulting from division
of a tetrad; e, staphylococcus. (From Marshall after Novy.) —

logical characteristics. Similarly, among the bacilli, character-
istic forms result as single individuals and others which form
chains of various lengths.

SPoRULATION.—A large number of bacteria possess the power
of developing into a resting stage by a process known as sporula-
tion or spore formation. Sporulation is regarded as a method of
resisting unfavorable environment. This is illustrated by the 4
anthrax bacilli which are readily killed in twenty minutes by a 4
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. While the vegetative forms of
anthrax 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

THE FUNGI | 369

return to the vegetative form and then are capable of multiplica-
tion by fission in the ordinary way.

Two kinds of spores occur amongst bacteria, arthrospores and
endospores, Arthrospores represent entire bacterial — vegetative
cells which lose some of their water and thicken their walls and
thus become resting cells. Endospores or true spores of bacteria
are formed within the cell. They have the power of resisting a
greater heat than the arthrospores.

Rapipity oF GRowTH AND MULTIPLIcATION.—The rapidity
with which bacteria grow and multiply is dependent upon species
and environment. The rapidity of the growth is surprising.
Under favorable conditions they may elongate and divide every

Fic. 273.—Spores and their location in bacterial cells. (Marshall after Frost and
McCampbell.) :

twenty or thirty minutes. If they should continue to reproduce
at this rate for twenty-four hours, a single individual would have
17 million descendants. 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. However, 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 for bacteria.

CHEMICAL CoMpPosITION OF BAcTERIA.—The quantitative
chemical composition of bacteria is subject to wide variations,
dependent upon the nutritive material furnished them. About

eee:

370 PHARMACEUTICAL BOTANY

80 to 85 per cent. of the bacterial body is water; proteid sub-
stances 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.

00000 8 :

Fic. 274.—Spore germination. a, direct conversion of a spore into a bacillus
without the shedding of a spore wall (B. leptosporus); b, polar germination of Bact.
anthracis; c, equatorial germination of B. subtilis (hay bacillus); d, same of B.
megaterium; e, same with “horse-shoe” presentation. (From Marshall after Novy.)

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
the exotoxins or bacterial poisons secreted by certain bacteria like

fh tt Be (Ul

Fic. 275.—Types of bacilli. (After Williams.)

the tetanus and diphtheria organisms during the process of
growth, also occur.

MorRPHOLOGICAL CLASSIFICATION OF BACTERIA *
THE Lower BAcTERIA

The lower bacteria comprise most of the forms which produce
disease and the majority of the forms which are used in the arts

* No strictly morphological system of classification for the bacteria will subserve
the purpose of properly identifying species of this group owing to a deficiency of
structural characteristics for the separation of so many species. For the main
divisions and some of the smaller details, the application of a morphological key
will be sufficient, but cultural tests for their physiological phenomena and staining
reactions are essential in working out the identity of a specific bacterial organism.

THE FUNGI aTt

and industries. They are grouped under the first 3 families, viz.,
Coccacea, Bacteriacee and Spirillacea.

ef swV

%
ee
ne

oore

Fic. 276.—Types of spirilla. (After Williams.)

Famity I.—Coccace#.—Cells in their free condition
globular, becoming but slightly elongated before division.

Cell-division in one, two or three
directions of space. |

A. Cells without Flagella.

1. Division only in one direc-
tion of space forming an aggrega-
tion 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 direc-
tions of space forming a package-
shaped or cubical aggregation—
Sarcina. 3

B. Cells with Flagella.

1. Division in two directions
of space—Planococcus.

2. Division in three direc-
tions of space—Planosarcina.

Famity II.—BACTERIACE.
Cells longer than broad, gen-
erally two to six times, straight
or only with an angular bend,
never curved or spiral, division

only at right angles to axis or rod;

endospores.

Fic. 277.—Crenothrix polyspora, an
iron bacterium, which oxidizes iron
causing a deposit of iron hydroxide to
accumulate in water pipes where it may
cause trouble. (After Migula from
Schmidt and Weiss and Marshall.)

with or without flagella and

ate PHARMACEUTICAL BOTANY —

1. Flagella and endospores absent—Bacterium.
2. Flagella and endospores present.
(a) Flagella over entire surface—Bacillus.
(b) Flagella at one end—Pseudomonas.
Famity III.—Spirmttace#.—Cells curved or spirally bent,
generally motile through polar flagella.
1. Cells stiff, not flexile.
(a) Cells without flagella—Spirosoma.
(4) Cells with one, very rarely with two polar flagella—
Microspira (Vibrio).
(c) Cells with a bundle of polar flagella—Spirillum.

‘Tue HicHer BAcTERIA
ate" ;

Famity IV.—Mycopacrertace@.—Cells short or long,
cylindrical or clavate-cuneate in form, without a sheath surround-
ing the chains of individuals, without endos-
pores, with true dichotomous branching.

A. In cultures possessing the characters of
4 true bacteria. Growth on solid media
smooth, flat, spreading. Rod with swol-
3} len ends, or cuneate or clavate forms—
1 Corynebacterium.

B. Cultures on solid media raised, folded
or warty. Generally short slender rods,
rarely short branched. Take the tubercle
stain—M ycobacterium.

Suctente fas ak Famity V.—CHLAMYDOBACTERIACEZ.—
alba). (After Wino. Lhread-like, composed of individual cells, sur-
gradsky fron Schmidt rounded by a sheath. Simple or with true
~ Mes iss and Mar- branching. Ordinary vegetative growth by

: division in only one direction of space, 7.¢.,
at right angles to the longer axis.

A. Cell contents without sulfur granules.

1. Filaments unbranched.
(a) Cell-division only in one direction of space.
(6) Cell-division in gonidial formation in three direc-
tions of space—Actinomyces (Streptothrix).
*Marine forms with cells surrounded by a very
delicate hardly discernible sheath—Phragmidiothrix.

Fic. 278.—Sulfur

THE FUNGI 373 :

**Fresh-water forms with easily discernible sheath—
Crenothrix. (Iron bacteria.)
2. Filaments branched.
B. Cell contents with sulfur granules— Thiothrix.

Some BAcTERIA PropucinG DisEASE IN MAN OR THE LOWER ANIMALS

ORGANISM _. DISEASE :
Alcaligenes melitensis 24,5 3:0 eie.g We ae nee Ss oe Malta fever 3
Actinomyces bovis 6:2 0)5.. i Saha een sv deem eee ee Actinomycosis in cattle a
Bacillus anthracis. 3.23035 itn tess eae ste Anthrax
Clostridium chauvels, 23 (see Ss See “Blackleg’’ of cattle
Clostridium :tetani. 450. 4. eee POA ees Tetanus or Lock-jaw
Clostridium: welthil 355 63. Ses Oo ee res Gas-gangrene :
Corynebacterium diphtheriz................--: -... Diphtheria 4
Diplococcus pneumoniae... .. 6... 666 ee eee ere eee Lobar pneumonia }
Eberthella typhi. 6500265 5. AG ea se eines ons Typhoid fever q
Hemophilus pertuisis.2. .-5.. s4ie8 ci ene hae Whooping cough 3
Hemophilus influenzz..........--++++eeer eee eee Influenza a
Mycobacterium leprae... .....--- +. sete eee eee Leprosy
Mycobacterium tuberculosis. ........---+++++e+005 Tuberculosis
Neisseria catarrhalis. .... 2.2... 26-0 e ee ee eee eee Catarrh
Neisseria intracellularis...........- Nae er eee = tere Cerebrospinal meningitis
Neisseria gonorrhoez......----+esereee erect Gonorrhoea
Pasteurella pestis... 0... 200s eee cece e etter terete tes Bubonic plague
Pfeifferella mallet 52 0. es ee Glanders’

Rickettsia prowazeki.........-- ee Os ene Typhus :
Staphylococcus aureus. ......---+2++seeeeerereeee Boils, abscesses, carbuncles a
Streptococcus scarlatin®........--+++ sree eee ... Scarlet fever “4
Streptococcus erysipelatis.......---++++sere seers Erysipelas

Salmonella paratyphi.....-.------+-++seeee eres Paratyphoid fever (A)

Salmonella schottmulleri.........---+-+-+e+ereeee Paratyphoid fever (B)

Vibrio ComMA (scsi = ee ee oe aes te Be Cholera

(Spirochaetales, intermediate forms between bacteria and protozoa)
Borrelia recurrentis.......--+----ercttrc cts Relapsing fever (European)

Borrelia duttont(. (25022505 eo ee ies Relapsing fever (African)
Vincent’s angina

Borrelia vincentie 44 v4 oso e token he se eee ine en
Treponema pallidum. .....-.+--+-++seetererrreee Syphilis or lues
Some Bacteria Producing Disease in Plants
. ORGANISM DISEASE
Actinomyces Myricarum....---+-+----- pete Aaee Tubercles upon and _ lesions
within Myrica and Comptonia
’ Bacterium tumefaciens. ....--- +--+ -es+r terre Crown gall
Bacterium savastanoi......------+:ssertrtrtee Olive knot
Bacillus amylovorus.....-.--+-++ssrrtcrrtttt Pear blight
Bacillus Solanacearum.....-----++ee sere rrr Wilt of Solanaceae
Bacillus tracheiphilus.......-----++sersr tcc Wilt of Cucurbits
Pseudomonas juglandis.....-------++++++r-877+ Walnut blight

Pseudomonas Stewarti....-----errrcr terre Wilt of Sweet Corn.

374 PHARMACEUTICAL BOTANY

The iron bacteria utilize iron salts forming iron oxide which is
deposited in a sheath around their walls. They are responsible
for the rusty appearance of some water.

Famiry VI.—Beccratoace& (Sulfur Bacteria).—Thread-
like, without a capsule, but with an undulating membrane. Cell
contents show sulfur granules.

Threads apparently not separated, septa only faintly visible
with iodine staining. Colorless or faintly rose-colored—
Beggiatoa.

The sulfur bacteria oxidize the hydrogen sulfide of sulfur
compounds, liberating sulfur in the form of granules within the
cell.

ParuocEnic BacTEertiA.—Bacteria which produce diseases in
living organisms are called pathogenic bacteria or pathogens. A list
of some of the more important of these and the names of the
disease caused are shown on p. 373.

Crass II.—MyxomyceTEs, or SLIME MOoLps

Terrestrial or aquatic organisms, frequently classified as
belonging to the animal kingdom and found commonly on decay-
ing 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 of its body called
pseudopodia. ‘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, 7.e., turning away from the sun’s rays. 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 (ethallia)
to more elevated bodies in which the net-like structure of the
plasmodium is preserved (plasmodiocarps) to stalked sporangia

THE FUNGI Be | be

(spore cases). All of the fructifications, however, produce Spores.
During wet weather amoeboid protoplasts (swarm spores) escape
from the spores, each developing a single celium and moving
actively about. In time the cilia disappear and _ these
swarm spores coalesce in smaller then larger groups to form a
plasmodium.

Aes

LES

ah TC

Fic, 279.—Slime molds. A, B, Comatricha nigra. A, Sporangium, natural
size; B, capillitium, 20/1; C, E, Stemonitis fusca; C, sporangium, natural Bize;-D),
and £, capillitia, 5/1, 20/1; F, H, Enerthema papillatum, F, unripe; G, mature
sporangium, 10/1; H, capillitium, 20/1. (C, D, after nature. A, F, G, H, after
Rostafinski; B, E, after de Bary in Die natiirlichen Pflanzenfamilien 1. 1, p. 26.)

Crass III.—Eumycetes (True Funct)

The True Fungirepresent a large class of chlorophylless plants
in which are included plants ordinary called fungi, such as the
molds, water-molds, yeasts, blights, mildews, rusts, smuts, toad-
stools, puffballs, etc. In nutrition, they are either saprophytes
or parasites or may be adapted to both modes of existence. Most
of them have a vegetative body called a mycelium consisting of
filamentous hyphe. This class is subdivided into the following
sub-classes:

Sub-Class I. Phycomycetes or Alga-like Fungi—molds,
water-molds, blights, mildews, rots.

Sub-Class II. Ascomycetes or Sac-Fungi—yeasts, blue and
green molds, powdery mildews, cup-fungi, morels, ergot, etc.

Sub-Class III. Basidiomycetes or Club-Fungi—smuts, rusts,
mushrooms, puff-balls, earth stars, etc.

376 PHARMACEUTICAL BOTANY

Sub-Class IV. Deuteromycetes or Imperfect Fungi—fungi
whose life histories are incompletely known, as celery-blight,
trichophyton, monilia, oidium, etc.

THE PHYCOMYCETES, OR ALGA-LIKE FUNGI

The Phycomycetes represent a small group of fungi showing
close affinity with the green algee. ‘Their mycelium is composed
of hyphz which are usually unicellular up until the time of
reproduction and many-nucleated. This suggests a close rela-
tion with the Spihonales group of green alge. ‘Their sexual
organs are likewise similar in structure. ‘Transverse septa
appear in the hyphe upon the formation of reproductive organs
which separate these structures from the vegetative hyphz.
They reproduce by spores as well as by sexual reproduction.

Those Phycomycetes in which sexual reproduction is brought
about through the union of like gametes belong to the order
called <ygomycetales which includes the black bread molds and
their kin; those in which sexual reproduction takes place through
the union of unlike gametes belong to the Odmycetales or order of
the watermolds and their allies.

CuttivaTion or Motps.—Molds and other fungi will grow
on most bacteriological culture media but do better if some sugar
is added. ‘They may also be grown on moistened slices of bread,
on slices of potato or carrot, or on decoctions of prunes, potatoes,
manures, peas, beans, etc. Beerwort is also a good substratum
for their growth. The liquid media may be made solid by the
addition of agar. An excellent medium for the growth of molds
and most other fungi in the laboratory is a modification of
Czapek’s medium, the formula for which follows:

BREN Se 1G iathes be ieee Pie eee a. ee 30.0 Gm
Sethu Nitrates sent ia t e e ee 20
Dibasic Potassium Phosphate....-................ 1.0
Magnesium Sulfate Crystals. os cc. ccc. 0.5
pumennin CRORGG oe ee 0.5

Ferrous Sulfate

ya gs aay baie s Gen te ee 0.01
WER ak ei Pc oe eek ee 1000.0 cc
For solid medium, add Ager... 60 15.0 Gm

Slide cultures are of great value in the study of molds since
they permit the study of the aerial and submerged hyphae

THE FUNGI 377

which are often tangled in mounts prepared by transferring the
mycelium with a forceps or needle. These may be prepared by
the Henrici method as follows: Clean large rectangular cover
glasses and deposit a drop of de Khotinsky cement or melted
sealing wax on each end; spreading it out with a small hot iron
file to form a layer 5 mm. wide. Heat a slide in a flame and
place the cover glass on it with the cement side down. The slide
should be just hot enough to soften the cement rather than to
liquefy it. The culture chamber thus prepared has a space
between the cover slip and slide about 1 mm. deep. Melt a tube
of Czapek’s agar, cool and inoculate with spores of the mold to be
examined. With asterile pipette transfer some of the inoculated
agar to the edge of the cover glass and allow it to run under until
part of the space has been filled. ‘The slides are then placed in a
clean glass staining jar containing moistened filter paper in the
bottom and the lid sealed with vaseline, ‘and the slides incubated
until growth has occurred.

OrRDER ZYGOMYCETALES

THe Biack Breap Mo.p.—Ruizopus Nicricans (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 isincontact. Each
of these, upon germinating, sprouts out and forms a vegetative
mycelium consisting of three kinds of hyphe, viz.: rhizoidal or
submerged hyphae, sporangiophores or aerial hyphe and stolontferous
hyphe. The branching rhizoidal hyphe penetrate the substratum
and secrete a diastatic ferment that changes the water insoluble
carbohydrate materials into a soluble sugar which passes into
solution and is absorbed by their walls. ‘This, upon entering the
hyphez, is converted into protoplasm, and so the mold increases in
size. Sporangiophores or aerial hyphz arise vertically or
obliquely from a bulged-out common base of the rhizoidal hyphz.
Each of these when mature bears upon its summit a spheroidal
sporangium containing numerous small, brownish, multinucleate

378 PHARMACEUTICAL BOTANY

spores called endospores. The wall of the sporangium is beset
with asperites of calcium oxalate. Springing from the base of
the sporangiophores or aerial hypha, one or more stoloniferous
hyphze traverse a portion of the surface of the substratum, and
their tips, coming in contact with the substratum, swell up form-
ing an adhesive organ or appressorium which branches out below
into a cluster of spreading submerged hyphz and above into
several aerial hyphz bearing sporangia. This method of growth
proceeds until the entire surface of the nutritive medium is |
covered with a dense fluffy mycelium.

Fic. 280.—Black mold (Rhizopus nigricans). A, older plant; myc, mycelia;
sph, sporangiophore; sp, sporangium; st, stoloniferous hypha produced by A, and
giving rise at its tip to a new plant, B. Greatly enlarged. (Gager.)

Rhizopus reproduces by two methods. The most common
one is that of division by cleavage furrows. Inthis ASExUAL METHOD,
a transverse wall is laid down in the sporangiophore near its tip.
The terminal cell thus formed swells up, becoming globular in
shape and its multinucleate protoplasmic contents, by progressive
cleavage, become divided 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 sporan-
gium 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.

402 PHARMACEUTICAL BOTANY

The Cedar and Apple Rust produces aecia on the leaves of the
apple and hawthorn and telia on the red cedar (Juniperus virgimi-
ana). The mycelium of this fungus growing in the leaves of the
juniper produces the large brown galls known as “cedar apples.”

OrpeR 3.—AURICULARIALES.—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 earlike fruiting body is known as the sporophore.
Its internal surface is lined with a hymenium or fruiting membrane
consisting of numerous four-celled basidia, each of which cuts
oft at its tip a basidiospore. A common example of this group
is the Jew’s ear fungus.

OrpeR 4.—TREMELLALES.—Saprophytes which live on
decaying wood as moist, soft, quivering gelatinous growths
becoming later dry and horny.

Tue HicHer BAsIDIOMYCETES

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.—HyYMENOMYCETES
(Hymenium or spore-bearing surface exposed)

This division of higher basidiomycete fungi comprises the
following orders: Dacromycetales, Exobasidiales, Thelephorales,
Clavariales and Agaricales.

OrvER 1.—DacromyceraALes.—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 the mycelium may break
up into oidiospores, if the wet period is prolonged. In consisting
of slimy gelatinous masses the ‘“‘weeping fungi” approach the
Tremellacee but are distinguished from them in the basidium
being undivided in the former and divided in the latter.

THE FUNGI 403

OrvER 2,—ExopasIDIALEs.—This group is found growing
parasitically on shrubs especially those of the heath family.
The mycelium lives in the tissues of the stems, leaves, sepals and
petals and produces spongy, fleshy, yellowish or brownish galls
which are popularly called ‘Azalea apples.” The galls are
edible. ‘They are covered with a hymenium.

ORDER 3.—THELEPHORALES. ‘These possess fruit-bodies
appearing on tree trunks as flattened leathery or crust-like

Fic, 298.—Coral-like fruit-bodies of Clavaria flava. (Harshberger, from Photo by
W. H. Walmsley.)

or bracket-like outgrowths often overlapping each other.
Example: Corticium.

ORDER 4.—CLAVARIALES, the coral or fairy club fungi.
Fleshy coral or club-shaped forms, all of which are saprophytes
found in woods growing in bunches out of leaf mold. They
are all edible and of a white, yellow, lavender or some other
brilliant color. (See Fig. 298.)

OrbDER 5.—AGARICALES, the mushroom or toadstool alliance.
Alike with the other members of the Basidiomycetes, the plant
body consists of the mycelium, ramifying through the sub-

404 PHARMACEUTICAL BOTANY

stratum, but the part which rises above the surface (the sporo-
phore or fruiting body) 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.

Famity 1.—HypNaAcE#, or tooth fungi. This group is
characterized by the hymenium being placed over purplish,
spiny or long finger-like projections of the pileus. Many of
the species of the genus Hydnum are edible.

[i ee a. a ree ‘i
Fic. 299.—Boletus felleus in three stages of development. (After Patterson, Flora W.
and Charles, Vera K., Bull. 175, U. S. Dept. Agric., pl. xxxi, Apr. 29, 1915.)

Famity II.—Potyporace#, or pore fungi. ‘The sporophores
or fruiting bodies of these fungi are various. They may be
entirely supinate with pores or shallow depressions on their
upper surfaces (Merulius), or mushroom-like (Boletus), or of the
nature of woody or corky (Fomes) or fleshy (Fistulina) brackets.
In all cases the hymenium or basidial layer lines the inner
surface of tubes which open on the exterior as pores.

The sporophore or fruit body of Polyporus officinalis, when
deprived of its outer rind and dried, constitutes the drug AcarI-

THE FUNGI 405

cus. This species grows abundantly on various species of pines,
spruces and larches.

Famity III.—Acaricace&, 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. Accord-

Fic. 300.—Meadow mushroom (Agaricus campestris L.). A, view showing
under side of pileus; g, gills; 2, annulus, or remains of the veil attached to the stipe,
B, side view; s, stipe; a, annulus; p, margin of pileus, showing at intervals the
remains of the veil. (Gager, after W. A. Murrill.)

ing to the color of their spores the members of this family are
divided into 5 groups:

1. Leucospore or white-spored, as Amanita.

2. Rhodospore or red-spores, as Pluteus.

3. Ochrosporz or brown-spored, as Pho/iota.

4. Porphyrosporee or purple-brown spored, as Agaricus.

5. Melanospore or black-spored as Coprinus.

The color of the spores is best determined by cutting off the
pileus, placing it gill-surface downward on a piece of smooth
paper and covering it with a bell jar. Within a day or less the
spores, falling off the gills, leave an imprint of the gill arrange-
ment which is called the spore print.

Acaricus CAMPESTRIs (CommMon MusHroom).—This plant.
also referred to as Psalliota campestris (L.) Fr. by some authors
is a common edible gill fungus which grows in open, grassy

THE FUNGI 383

Yeast plants grow in dilute saccharine solutions containing
dissolved 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 nitrog-

Fic. 283.—Yeast, Saccharomyces cerevisiae, the variety known as brewers’ bot-
tom yeast; a, spore formation; 6, elongated cells. (After Schneider, Pharmaceutical
Bacteriology.)

enous and other matters. 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

384 PHARMACEUTICAL BOTANY

effect on the medium in which they live. Yeast causes a whole-
sale destruction 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. .

The protoplasm of the yeast plant manufactures two kinds of
enzymes, viz., an extracellular enzyme called invertase which
passes out of the yeast cell and hydrolyzes cane sugar to dextrose
(invert sugar) and an endocellular enzyme called zymase which

Fic. 284.—Saccharomvees cerevisie. The form or variety known as brewers’ top
yeast. (Oberhefe.)

remains within the yeast cell and breaks down dextrose into
alcohol and carbon dioxide.

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 conditions by eliminating most of their water, suspend-
ing physiological processes, and becoming dormant. Some-
times they enter the resting 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

THE FUNGI 385

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

Fic. 285.—Saccharomyces ellipsoideus. A common yeast found on grapes, jams,
jellies, etc. Budding process is shown in many of the cells as also the vacuoles.
(Schneider, Pharmaceutical Bacteriology.)

later perish. But some will 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 commences 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-granddaughter-buds before separation
from the parent-cell takes place.

386 PHARMACEUTICAL BOTANY

There are two varieties of brewer’s yeast, viz.: Tor YEAsts
and Bottom Yeasts. ‘Top yeasts grow on or near the surface
of the liquid and produce rapid fermentation at summer tem-
peratures causing great quantities of carbon dioxide to arise to
the surface and thus forming the froth which is characteristic of
ale, stout and porter. They tend to remain attached in clusters.
Most of the yeasts used for compressed yeast and bread making
are top yeasts.

a 3 =) : at . aa
E Vai ce af t Gacy <p

Fic. 286.—Saucer-shaped fruit-bodies of Peziza repanda. (Harshberger, from Photo
by W. H, Walmsley.)

A

Bottom yeasts grow at about 4°C. at or near the bottom of
the vat. They are used in the manufacture of lager beers. |

Compressed yeast (Cerevisiae Fermentum Compressum) or
“yeast cake” consists of the moist, living cells of Saccharomyces
cerevisie or of other species of Saccharomyces, combined with a
starchy or absorbent base.

Yeast contains the antineuritic vitamin B, and the pellagra
preventive vitamin B, or G.

OTHER YEASTS.—Saccharomyces ellipsoideus (Fig. 285) 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 from other wild species
found in grape juice, like S. apiculatus and S. membranifaciens

THE FUNGI 387

which exert a deleterious eftect in wine production. Its cells
are ellipsoidal, 64 long, occurring singly or in rows of several
generations, which are rather loosely joined.

ED

aN

Fic. 287.—Cup Fungi. A, B, Lachnea scutelata. A, Habit; B, ascus with para-
physis; C, D, Lachnea hemispherica; C, habit; D, ascus with paraphysis; E, Sarco-
sphera arenosa habit; F. G, Sarcosphera coronaria; F, ascus; G, habit; H, Sarcosphera
arencola ascus with paraphysis. (See Die natiirlichen Pflanzenfamilien 1, 1, p. 181.)
(Harshberger.)

S. pastorianus is a common contaminant in brewer’s yeast
and in yeast cakes. It forms large sausage-shaped cells. It is a
cause of disagreeable flavors in beer.

OrDER 2.—PEzIZALES or cup fungi. Examples: Peziza,
Lachnea and Ascobolus.

Parasitic or saprophytic plants, whose vegetative bodies con-
sist of a mycelium ramifying through the substratum and whose
above ground fruiting bodies are sessile or stalked, cup or saucer-

. 388 PHARMACEUTICAL BOTANY

shaped structures termed apothecia (sing. apothecium), in which -
a fruiting membrane (ascigeral layer) lines the concave upper
surface. The asci are usually eight-spored and separated from
each other by filamentous structures called paraphyses. (Figs.
286 and 287.)

sees?

Y ZS

Za
sweesss
wette

|

Fic. 288.—Three aerial hyphz showing the characteristic brush-like branching
and spore formation of Penicillium glaucum. This fungus is a true saprophyte and
is never found on living fruits or vegetables. a, Conidiophore branching above into
secondary conidiophores; 6, sterigmata; c, conidiospores. (Schneider.)

OrDER 3.—PLecTASCALES, the blue and green molds.
Examples: Aspergillus and Penicillium. |

PenicrLtium GLaucum (GREEN Mop or MILpEw), a type
of mildew, belonging to the Ascomycetes class of Fungi, forms
sage-green crusts on bread, jellies, old boots, gloves, and various
pharmaceutical preparations. It consists of a felt-like mass of
interlaced tubular hyphz called a mycelium. - From the mycelium

THE FUNGI 389

numerous hyphz project into the air and bear a green powder,
the spores. These hyphz are called aerial hyphe. Other
hyphz grow down into the substratum and are called sub-
merged hyphae.

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 that each hypha has a transparent wall and proto-
plasmic contents and is divided by transverse septa into a number

—

mt 3

Fic. 289.—Penicillium Roqueforti. a, Part of a conidiophore; 6, c, other types
of branching; d, young conidiophore, just branching, ¢, /, conidiiferous cells; g,
j, diagrams of types of fructification. k, /, m, n, germinating spores. (After Thom.)

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 hyphe bear brush-like branches, which become
constricted on their ends into a moniliform aggregation of
rounded spores appearing like a row of beads. Each aerial

390 PHARMACEUTICAL BOTANY

hypha is composed of a vertical septate branch of the mycelium
called the conidiophore, branches of this, which are called secondary
conidiophores, and chains of spores at the tips of sterzgmata (cells

al

Fic. 290.—Penicillium Camemberti. a, Conidiophore with common type of
branching with conidiospores; (6) a common less-branched form; ¢, d, f, diagrams
of large fructifications; g, i, j, germinating conidiospores. (From Bull. 82, Bureau
of Animal Industry, also after Thom.)

bearing conidia) which are called conidia or conidiospores. ‘The
conidia form the loose green powder characteristic of Penicillium.

A number of species of Penicillium are useful in the arts.
Penicillium roqueforti is the principal ripening agent of Roquefort,
Gorgonzola and Stilton cheeses. It possesses blue-green globu-
lar conidia 4 to 5u in diameter.

THE FUNGI 391

Penicillium camemberti is the principal agent in the ripening of
Camembert cheese. It possesses ellipsoidal bluish-green conidia
4.5 to 5.5u in diameter.

Penicillium brevicaule grows on old moist paper and has been

used to detect the presence of arsenic, for when grown in media -

containing this element, it develops the compound, diethylarsine.
It is yellowish-brown in color and its conidia are rough and spiny.

Penicillium expansum is often found on decaying apples where
it produces brownish coremia.

Cirric Acip is produced by fermenting solutions of carbo-
hydrates (mono- or poly-saccharides) by means of molds of the
genera Penicillium, Citromyces, Aspergillus. and Sterigmatocystis.
The unfermented carbohydrate is removed by an alcoholic
fermentation using yeast.

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, tobacco, cotton-seed meal, old leather, stale
black bread, etc. Like Penicillium its vegetative body consists
of a mycelium consisting of aerial and submerged hyphe.
It differs from Penicillium, however, mainly in not possessing
septated conidiophores and by the upper portion of the conidio-
phores being globular. Upon the globular extremity of the
conidiophores are placed numerous elongated sterigmata which
bear chains of grayish-green conidia. These are spherical and
prickly and range from 7 to 30u in diameter. Under certain
conditions closed brownish fruit bodies called perithecia are
produced. These arise on the surface of the substratum from
spirally coiled hyphae and when mature possess numerous asci,
each of which contains five to eight ellipsoidal ascospores.

Aspergillus oryze is a yellowish-green to brown mold which
secretes diastase, a valuable digestive ferment, having the power
of converting starch into sugar and dextrin. For centuries the
Japanese have employed this species in the preparation of rice
mash for Saké, as well as in manufacture of Miso and Soja
sauce. The spherical conidiospores are 6 to 7 in diameter
and of a yellowish-green color.

392 PHARMACEUTICAL BOTANY

Aspergillus fumigatus is a pathogenic species which produces a
disease in birds, horses, cattle and even, though rarely, in man
that is called aspergillosis. ‘The organ most prone to infection

O) QAA?ADO
Q ” on O S BG
0) Cy & CO OOO
QQ aa _ WW oA
CORRWIEE :
B ay?
(). a
5 8 4
oar 8
Ns
O

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Fic. 291.—Aspergillus oryze associated with yeasts in the making of the Japanese
beverage Saké. Vegetative hyphae (a) and spore-forming hyphz (4, c, d) are
shown. (Schneider, Pharmaceutical Bacteriology.)

by this organism is the lung, although the skin, cornea, ears and
other parts are also subject to its parasitic influence. It pro-
duces short conidiophores with sterigmata bearing long chains
of rounded, colorless conidia 2.5 to 3u in diameter. Harsh-

- pa Na Tol aii toi a.) oot Le, sae Ne eer ead RMD Oo sn aaa aE oe ahs Ci the |
Geta ae ~ lege cert ga : . A
ie ial

THE FUNGI 393

berger’ cites the presence of perithecia in this organism which
are nut-brown, globular, 250 to 350u in diameter, and inclose

Fic. 292.—Aspergillus niger showing conidiophores and conidiospores formation
with stages in germination of spores. (Harshberger, after Henri Coupin.)

oval, thin-skinned asci with eight, red, lenticular ascospores each

of which has a diameter of 4 to 5yu.
Aspergillus niger develops dark brown mycelial masses in which

are to be noted tall conidiophores terminating in large, black,

1 “Mycology and Plant Pathology,” p. 147.

ee a ee ee ee Tey

ee ee ee ee ee ee ee ee eS ee ee Mee eee ee ee ee ee, ee oe ee ee

394 PHARMACEUTICAL BOTANY

globular spore heads. The globular vesicle of the spore head
bears radially arranged, branched sterigmata that cut off from
their tips chains of rounded black to dark brown conidia 3.5 to
5u in diameter. This fungus has been found to produce sup-
purative inflammation of the external and middle portion of
the human ear. It is also a cause of cork disease, so often impart-
ing a disagreeable taste to bottled beverages.

This mold, however, is of considerable economic importance.
While a number of molds produce citric and oxalic acids from

Fic. 293.—The morel, Morchella esculenta. (Gager, from photo by W. A. Murrill.)

sugars, Aspergillus niger is especially active in this direction and
selected strains of it are used commercially in the production of
citric acid on a large scale.

Orper 4.—TuBERALES, the truffles. Fungi whose septate
mycelium is often connected with the roots of trees forming the
structure known as mycorrhiza. Several species of the genus
Tuber growing in woods of France, Germany and Italy produce
tuberous subterranean bodies called Truffles, which are highly
prized as a table delicacy by the inhabitants of these countries.

OrpER 5.—HELVELLALES, the saddle fungi. Fleshy fungi
entirely saprophytic, living attached to leaf mold or growing in
humus soil or, in a few cases, on decaying wood. The fleshy
fruiting bodies (ascocarps) are divided into stalk (stipe) and cap

}
!
i
‘
s
t
'
'

THE FUNGI 395

(pileus) portions. 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 depressions are covered with an ascigeral layer composed
of asci and paraphyses. This species is edible. (See F ig.
293.)

ORDER 6.—PYRENOMYCETALES, the mildews and black fungi,
common as superficial parasites on various parts of plants. To
the Family Hypocreacee of the black fungi division of this order
the Ergot fungus, Claviceps purpurea, belongs.

Lire History or Caviceps 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 superficially, secrete a ferment, and cause decom-
position of its tissue and the resultant formation of a yellow,
mucous 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 penetrate 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, some-
what 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 numerous flask-shaped invaginations called peri-
theca, from the bases of which several sacs or asci develop from
the ends of hyphae. Within each ascus are developed eight fili-
form spores (ascospores) which, when the ascus ruptures, are

396 PHARMACEUTICAL BOTANY

Ee

ert
Fy

Fic. 294.—A, Balansia claviceps on ear of Paspalum; B-L, Claviceps purpurea; E,
sclerotium; C, sclerotium with Sphacelia; D, cross-section of sphacelial layer; E,
sprouting sclerotium; F, head of stroma from sclerotium; G, section of same; H,
section of perithecjum; 7, ascus; K, «germinating ascospore; C, conidiospores
produced on mycelium. (See Die natiirlichen Pflanzenfamilien 1, i, p. 372,)

THE FUNGI ed

dichascd and are carried by the wind to other fields of grain,
there to begin over a new life cycle.

The drug ergot contains a number of physiologically active
constituents chief among which are the alkaloids ergonovine,
ergotoxine and ergotamine and the amines /istamine and tyramine.
It is used in medicine chiefly because of its power to contract the
pregnant uterus and to check post partum hemorrhage.

Tue BAsIDIOMYCETES OR BAsIDIA FUNGI

This large class of fungi, including the smuts, rusts, mush-
rooms, gill and tooth fungi, etc., is characterized by the occur-
rence of a basidium in the life eee, A basidium is the swollen
end of a hypha consisting of one or more 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 one or more kinds of other types of
spores. All of the members have a mycelium composed of

septated hyphe.
THE Lower BASIDIOMYCETES

This group comprises the smuts and rusts which are parasites
_ living within the bodies of seed plants.

~ Orper 1.—UstiacInaLes, the smuts. Destructive parasites
which attack the flowers of various cereals, occasionally other
parts of these plants. Examples: Ustilago Maydis, the corn smut.
The basidiospores in this group are borne on a three- to five-
celled basidium called a promycelium.

Ustritaco Mayopis (Ustitaco ZE#) (Corn Smut).—Corn
smut is a destructive parasite which for a long time was supposed
to be confined to the Indian Corn, but which now is known to
occur on Mexican Grass. It is the only smut useful in medicine.
The mycelium of the fungus extends through all parts of the
infected organ of the host through the intercellular-air-spaces,
sending short absorbing branches into its cells. Im due course
of time, it produces large tumor-like swellings on the ears, tassels,
husk, leaves, root and stem. Each swelling is at first made up
of masses of hyphz, the whole being covered with a tightly
appressed membrane which has a whitish appearance like

ay
si

398 PHARMACEUTICAL BOTANY

German silver. Later the cells of the hyphae lay down walls
becoming divided into smaller cells, each of which becomes
rounded off as a spore. This spore is known as a chlamydospore
or brand spore. ‘The spores are at first a dark olive-green, but on

Fic. 295.—Smut boil of Ustilago zee on ear of corn, developed from one infected
kernel. (After Jackson, F. S., Bull. 83, Del. Coll. Agric. Exper. Stat., December, 1908.)

maturity are dark brown. They are sub-spherical and show
prominent but tiny spines. ‘These spores fall to the ground
and pass the winter. In the spring each germinates into a three-
to five-celled filament called a promycelium, from the cells of which

THE FUNGI 399

basidiospores arise. (Fig. 296.) The basidiospore, upon coming
in contact with young growing tissue or a wound upon the host
plant, starts a new infection and develops a mycelium which
penetrates the infected organ.

ORDER 2.—UREDINALES, the rusts. Obligate parasites
possessing a septated, branched mycelium which ramifies

a
Ai 4
N a

Fic. 296.—Germination of the chlamydospores of corn smut (Ustilago zee); 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
140 per cent. acetic acid, formation of infection threads. a, Spores; 6, promycelia;
c, basidiospores; d, infection threads; e, detached pieces of mycelia. (After Bull.
57, Univ. Ill. Agric. Exper. Stat., March, 1900.)

through the intercellular-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 peculiarities of the group.

Tue Wueat Rust (PucctntA GRAMINIs).—If we examine the
wheat plant just before harvest we will find on the stems and

400 PHARMACEUTICAL BOTANY

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 disturbed 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 |

Fic. 297.—Spore forms of wheat rust, Puccinia graminis. A, Section through
barberry leaf showing pycnia (spermagonia) on upper surface and zcia on lower;
B, two urediniospores; 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.)

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 uredospores (uriediniospores) are thrust out. These
uredospores are orange-brown in color and covered with minute
spines. ‘The mass of them has been called a uredinium. These
spores are.detached from the pedicels and blown by the wind
to healthy plants. After summer is over and dry weather

THE FUNGI 401

comes on, an examination of stubble in the field (blades of grass
and stems of wheat left carelessly), shows that 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.

The teleutospores are two-celled and have thick walls and
persistent 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 ~
appearance of a number of small depressions on the upper side
of the leaf. These in section are a rich chocolate brown and
known as spermagonia. In the center of a spermagonium are
produced hyphz, which project out to its orifice and obstrict
off minute spores called spermacia. On the opposite side of the
leaf, cup-shaped depressions are formed, each with a limiting
membrane (feridium). Within the cup-shaped depression thou-
sands of spores are formed in chains closely packed together.
These are the acidiospores (aciospores). The cluster cup is called
an Atcidium (Aicium). These zciospores are conveyed to wheat
and cause infection, thus completing the life cycle. It has been
observed that in America the uredospores or summer spores may
_ winter over and infect healthy plants, so that the Barberry
phase is completely eliminated from the life cycle.

Some OTHER Rusts.—The White Pine Blister Rust,
Cronartium ribicola, causes great damage yearly to the Pinus
strobus and other 5-needle pines. It has its aecial and spermatial
stages on these pines and the uredospore and teleutospore stages
on currants (Ribes spp.). The Hollyhock Rust, Puccinia mal-
vacearum occurs on the hollyhock (Malva rosea) and other species
of Malva, notably the drug-yielding Malva rotundifolia and M.
sylvestris. It passes through two stages, the teleutospore and
basidiospore, on the same host, only a single host being involved.

402 PHARMACEUTICAL BOTANY

The Cedar and Apple Rust produces aecia on the leaves of the
apple and hawthorn and telia on the red cedar (Juniperus virgini-
ana). ‘The mycelium of this fungus growing in the leaves of the
juniper produces the large brown galls known as “‘cedar apples.”

ORDER 3.—AURICULARIALES.—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 earlike fruiting body is known as the sporophore.
Its internal surface is lined with a hymenium or fruiting membrane
consisting of numerous four-celled basidia, each of which cuts
off at its tip a basidiospore. A common example of this group
is the Jew’s ear fungus.

ORDER 4.—TREMELLALES.—Saprophytes which live on
decaying wood as moist, soft, quivering gelatinous growths
becoming later dry and horny.

THE HIGHER BASIDIOMYCETES

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 higher basidiomycete fungi comprises the
following orders: Dacromycetales, Exobasidiales, Thelephorales,
Clavariales and Agaricales.

Orper 1.—Dacromycetares.—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 the mycelium may break
up into oidiospores, if the wet period is prolonged. In consisting
of slimy gelatinous masses the “weeping fungi” approach the
Tremellacee but are distinguished from them in the basidium
being undivided in the former and divided in the latter.

THE FUNGI 403

OrDER 2.—ExosasIDIALES.—This group is found growing
parasitically on shrubs especially those of the heath family.
The mycelium lives in the tissues of the stems, leaves, sepals and
petals and produces spongy, fleshy, yellowish or brownish galls
which are popularly called ‘Azalea apples.” The galls are
edible. ‘They are covered with a hymenium.

ORDER 3.—THELEPHORALES. These possess fruit-bodies
appearing on tree trunks as flattened leathery or crust-like

Fic. 298.—Coral-like fruit-bodies of Clavaria flava. (Harshberger, from Photo by
W. H. Walmsley.)

or bracket-like outgrowths often overlapping each other.
Example: Corticium.

OrDER 4.—CLAVARIALES, the coral or fairy club fungi.
Fleshy coral or club-shaped forms, all of which are saprophytes
found in woods growing in bunches out of leaf mold. They
are all edible and of a white, yellow, lavender or some other
brilliant color. (See Fig. 298.)

OrbDER 5.—AGARICALES, the mushroom or toadstool alliance.
Alike with the other members of the Basidiomycetes, the plant
body consists of the mycelium, ramifying through the sub-

ee
gait
P rT al va

404 PHARMACEUTICAL BOTANY

stratum, but the part which rises above the surface (the sporo-
phore or fruiting body) 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.

Famiry 1.—Hypnace#, or tooth fungi. This group is
characterized by the hymenium being placed over purplish,
spiny or long finger-like projections of the pileus. Many of
the species of the genus Hydnum are edible.

Fic. 299.—Boletus felleus in three stages of development. (After Patterson, Flora W.
and Charles, Vera K., Bull. 175, U. S. Dept. Agric., pl. xxxi, Apr. 29, 1915.)

Famity II.—PotyporacE&, or pore fungi. The sporophores
or fruiting bodies of these fungi are various. They may be
entirely supinate with pores or shallow depressions on their
upper surfaces (Merulius), or mushroom-like (Boletus), or of the
nature of woody or corky (Fomes) or fleshy (Fistulina) brackets.
In all cases the hymenium or basidial layer lines the inner
surface of tubes which open on the exterior as pores.

The sporophore or fruit body of Polyporus officinalis, when
deprived of its outer rind and dried, constitutes the drug AGARI-

Lentved 5/3 /o

THE FUNGI 405

cus. This species grows abundantly on various species of pines,
spruces and larches.

Famity III.—Acaricace&, 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. Accord-

Fic. 300.—Meadow mushroom (Agaricus campestris L.). A, view showing
under side of pileus; g, gills; 2, annulus, or remains of the veil attached to the stipe.
B, side view; s, stipe; a, annulus; p, margin of pileus, showing at intervals the
remains of the veil. (Gager, after W. A. Murrill.)

ing to the color of their spores the members of this family are
divided into 5 groups:

1. Leucosporee or white-spored, as Amanita.

Rhodosporz or red-spores, as Pluteus.

Ochrosporee or brown-spored, as Pholiota.
Porphyrospore or purple-brown spored, as Agaricus.
. Melanosporze or black-spored as Coprinus.

The color of the spores is best determined by cutting off the
pileus, placing it gill-surface downward on a piece of smooth
paper and covering it with a bell jar. Within a day or less the
spores, falling off the gills, leave an imprint of the gill arrange-
ment which is called the spore print.

Acaricus CAmpEsTRIs (Common MusHroom).—This plant.
also referred to as Psalliota campestris (L.) Fr. by some authors
is a common edible gill fungus which grows in open, grassy

mR ON

406 PHARMACEUTICAL BOTANY

fields during late summer 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 mush-
rooms tons of horse manure are used. This is covered with
loamy soil 114 inches thick. The whole mass is compacted
together. Into the resultant beds is introduced English-grown
spawn, which comes in flat brick-shaped masses (horse manure
through which mycelium has grown). Pieces of these ‘‘bricks”’

ai
Diseaies =
-- ey 3

Fic. 301.—Fruiting surface (hymenium) of a mushroom (Agaricus). m, hyphe
of the trama and sub-hymenium; 4, basidium; st, sterigma; sp, spores. Diagram-
matic. (Gager.)

are put in the horse manure bed only after the heat has first
disappeared. The beds are then watered well and in a short
time the sporophores or fruiting bodies of the fungus spring up.
The mycelium or vegetative body of Agaricus which develops
in the soil from spores (basidiospores) is white and thready.
On this mycelium develop little white buttons, first about the
size of a pin head, becoming later pea size and then assuming a
pear-shaped form. At this stage the sporophore consists of a
cylindrical solid stipe or stalk and a pileus or cap. The border
of the pileus is joined to the stipe by means of a “partial veil.’
Within this veil is found a circular cavity, into which the gills |
grow. At first the stipe grows faster than the rest of the fruiting

THE FUNGI 407

body. ‘The pileus expands transversely and the gills keep pace.
After a while the veil ruptures, leaving a portion attached to the
stipe. This constitutes the annulus or ring (true annulus). The
hyphe in the pileus form the Tela contexta. If we make a section
through a gill, the hyphz are seen to run longitudinally. The
central part is called the ‘rama; next and outside trama is the
sub-hymenium; next, hymenium, consisting of basidia (hence a
basidial layer). Each basidium bears two or four little points

Fic. 302.—Deadly amanita (Amanita muscaria) showing volva at base of stipe
and frill-like stipe ring. (After Chestnut, V. K., Bull. 175, U. S. Dept. Agric., pl.
i, Apr. 29, 1915.)

known as sterigmata. Each sterigma bears a purplish-brown
basidiospore. The basidiospores falling to the ground germinate
into hyphz and these become interlaced to form a mycelium.

In the wild mushroom the gills are at first white in the button
stage, later pink; in the cultivated, fawn-colored. Ultimately in
the wild form the gills turn blackish brown. The spores are
purplish-brown. The color of the stipe is white; that of the
upper surface of the pileus varies from satiny white to pallid
brown.

Tue AMANITAS (Polsonous FuNGI).—Amanita muscaria and
Amanita phalloides, commonly known as the “‘fly agaric’? and

408 PHARMACEUTICAL BOTANY

the ‘“‘deadly agaric,” respectively, are very poisonous forms.
Amanita muscaria is common in coniferous forests, although it
may be found in cultivated soil along roadsides, in open woods
and wood margins. It occurs singly or in patches. Its chief
poisonous constituent is muscarine for which atropine is an
antidote. An emetic should first be given. Amanita phalloides

Fic. 303.—The deadly amanita, Amanita phalloides. Note the cup at the base of
the stipe. (Gager, from photo by E. M. Kittredge.)

is found in woods and borders of fields and occurs singly and not
in groups.

Each of these have fruiting bodies (sporophores), which
begin at the surface of the ground as a button similar to that of
the edible mushroom. This enlarges and assumes a dumbbell
shape. The whole button is covered by an outer veil, known
as the velum universale, which encloses the pileus, gills and stipe.
As the stipe lengthens more rapidly than the pileus, the upper
part of the veil is stretched and finally breaks in its middle

THE FUNGI “409

portion. ‘The lower part remains as a cup, out of which the
stipe grows. ‘The upper part is carried up as shreds adhering
to the margin of the pileus. The lower part is called the volva
or death cup. The annulus present is a false annulus, for it
represents a peeling down of the upper part of the stipe. Both
have chalk-white gills, an usually white stipe with a bulbous base,
and white spores.

The pileus of Amanita muscaria varies from orange-red or
blood-red to orange or yellow. ‘The surface is covered with

portion of a rotten log. (Photograph by author.)

prominent warty scales, the latter being remnants of the veil

which are readily removed. Its stipe is white or pale sulfur
colored, at first stuffed, later hollow. The annulus is large,
white and ragged.

The pileus of Amanita phalloides varies from pure white to
dull yellow to olive or umber. It does not possess the warty
scales found in the Amanita muscaria, but occasionally has a few
membranous patches. Its stipe is generally smooth, usually
white, solid and bulbous below but hollow upwards.

Amanita phalloides is the most poisonous of all species of toad-
stools. Its chief poisonous constituents are the glucoside
amanita-hemolysin and the toxin called amanita-toxin. Amani-
tahemolysin is a blood laking principle. No antidote is known
which will neutralize the deadly effects of this fungus.

Fic. 304.—A colony of Puff Balls, Lycoperdon, growing saprophytically upon a

PREP Ge ne oP Peas, Se Bw a ee ME EEL

410 PHARMACEUTICAL BOTANY

Division B.—GASTEROMYCETES
(Hymenium inclosed)

OrpDER 1.—LycopEeRDALESs, or puff ball alliance. This
order includes a number of interesting parasites and saprophytes,
the most common of which are the earth stars belonging to the
genus Geaster and the puft balls, the most common form being
Lycoperdon. In these, the fruiting sporophore consists for the
most part of a shell-like covering called the peridium, composed

Fic. 305.—Bird’s Nest Fungus (Crucibulum). Note the open sporophores appear-
ing like nests with eggs. The “eggs”’ are the peridiola or special bodies in which
the basidiospores are produced. (Mottier.)

of an outer layer or exoperidium and an inner layer of endoperidium.
The peridium in the unripe condition of the sporophore covers
a mass of soft cellular tissue called the gleba. Upon the ripening
of this mass, the interior is seen to be divided into many-branched
compartments that are separated from each other by walls
made up of branched hyphez. These walls are lined with a
hymenium composed of many basidia, each of which constricts
off usually four basidiospores. The earth stars differ from the
puff balls in possessing an outer wall or exoperidium which
splits in star-shaped fashion.

OrpDER 2.—Niwu aries, the nest fungi. A group of
Gasteromycetes whose sporophores are crucible- or crater-like.

SE ae a.

THE FUNGI 411

These arise from a subterranean mycelium and show an outer
and inner peridial layer. The outer peridium is roughened at
its base. The inner peridium is leathery and may or may not be
continued over the top. When mature the crucible-like body
shows black seed-like bodies inside which
resemble eggs in a bird’s nest. Each
one of these is connected with the inner
peridium by a cord which resembles the
umbilical cord of an animal. These
inner bodies are called peridiola (sing.
peridiolum). Each peridiolum consists of
a hard glistening outer layer and a
spongy inner layer surrounding a cavity
into which basidia and _ basidiospores
project. These fungi are found in stiff
clayey soil.

OrvER 3.—PHALLALES, the carrion
or stink-horn fungi. This group con-
sists of highly and characteristically
colored forms which, when mature, emit
most vile and penetrating odors. ‘The
fruiting body, in each instance, begins as
an egg-shaped structure which starts its
growth from a widely spread under-
ground mycelium of chalky-white color.
As the “eggs”? enlarge they push above
the surface of the ground. The central Fic. 306.—Mature stink-
portion, elongating, then breaks through horn, Dictyophora duplicata.

the outer or peridial portion, which (arshberger, from photo. by
W. H. Walmsley.)

remains as a cup or volva at the base of
the mature fruit body. Upon the summit of the central stalk
rests the cup-like, many-chambered gleba. ‘The basidiospores
are imbedded in a greenish fetid slime formed by a mucilaginous
disintegration of the substance of the hymenium. ‘This fetid
green material is attractive to carrion flies which visit the plants
and remove the material with its embedded spores. ‘The latter
will not germinate until after passing through the alimentary
canal of these flies.

41 2 PHARMACEUTICAL BOTANY

Tue Func IMPERFECTI

An assemblage of varied forms, the life histories of most of
which are imperfectly understood. Only one kind of spore, the
conidiospore, is known to occur in this group. In this group
are included numerous parasites which produce diseases in crop
plants and some in man and the lower animals.

Among the fungi of this group producing diseases in man are
Trichophyton species which produce a skin affection called “ring-
worm,” and Monilia albicans which produces an infection of the
mucous membrane of the mouth known as “thrush.”

CHAPTER XVIII

THE LICHENS

Lichens are variously colored, usually dry and leathery
plants, consisting of symbioses of algae and fungi. In each case
the fungus derives its food from materials manufactured by the
alga and in return extracts water from the substratum and shares
it with the alga. The association is therefore mutually benefi-

Fic. 307.—A foliaceous lichen, Physica stellaris (L.) Nyb., growing on a rock.
The cup-shaped structures are the fruiting bodies (apothecia). At the left are
seen two very young plants. (Gager.)

cial. While Blue-green and Protococcus forms of Green Algz
and Ascomycete Fungi are for the most part concerned in lichen
formation, in some instances Basidiomycete Fungi are also
involved with the algze. ‘Thus we have two distinct classes of
Lichens depending upon the type of fungus associated with the
alga, viz.: Asco-lichens and Basidiolichens.

Lichens are found on the bark of trees, on rocks, logs, old

fences, etc.
413

414 PHARMACEUTICAL BOTANY

According to the manner of growth of the thallus and nature
of attachment to the substratum, three different sub-groups of
lichens may be distinguished, viz.: (1) Foliaceous, where the
thallus is flat, leathery and leaf-like and attached to the sub-
stratum at different points. To this group belong Physica,
Sticta and Parmelia. (2) Crustaceous, where the thallus closely _
adheres to rocks and bark of trees. To this group belong
Graphis and Pertusaria. (3) Fructicose, where the thallus is
upright and branching. To the last group belong Cetraria
islandica, species of Cladonia, and Usnea.

Fic. 308.—Cetraria islandica, or Iceland Moss, a medicinal lichen. (Sayre.)

STRUCTURE.—The body of a typical lichen shows a differ-
entiation into the following regions:

(1) A tegumentary layer of several rows of hyphal cells.

(2) An upper cortex of densely arranged hyphal cells.

(3) An algal layer containing green algal cells, often called
gondia.

(4) A medullary layer of loosely interwoven fungal hyphe,
with large air-spaces. This is usually the broadest layer and
sometimes also contains algal cells.

(5) A lower cortex of hyphal cells, less dense than the upper
cortex. From the surface cells of this layer rhizoids are

developed which absorb water and nutrient salts from the
substratum.

THE LICHENS 415

rat

ZRAOS SESS
Ss

Ot

390 090° a

ooo ree Oat eee
.

Fic. 309.—A lichen, Parmelia perlata (L.) Ach. 1. Plant, slightly reduced in
size; a, apothecia; 6, lobe of thallus; c, soredial patches. The soredia are vegeta-
tive reproductive bodies composed of both algal and fungal elements, and there-
fore able to reproduce the lichen; the ascospores, alone, cannot do this. 2. Longi-
tudinal section of apothecium; a, thecium; 5 and c, the two layers of the hypo-
thecium; d, upper algal layer; e, colonies of algae distributed through the medullary
layer; f, lower algal layer; g, lower cortical layer. 3. Cross-section of vegetative
portion of thallus. 4. Paraphyses (sterile fungal filaments), and spore-sac (ascus),
containing ascospores. 5. Ascospores. 6. Algal cells, surrounded by fungal
hyphae with haustoria (absorbing branches). (Gager.)

416 PHARMACEUTICAL BOTANY

Repropuction.—Lichens reproduce either by producing
spores or by detaching scale-like portions of their bodies. In
the Asco-lichens (which make up the majority) either saucer-
shaped structures called apothecia (sing. apothecium) occur on the
upper surface or dark-colored, spherical structures called peri-
theca somewhat embedded in the upper surface, arise. In either
type of structure asci and sterile hyphz (paraphyses) arise. Each
ascus usually contains eight spores. The. spores_upon_maturity _
are liberated, each germinating in the presence of moisture into a

Fic. 310.—Roccella tinctoria, a dye-yielding lichen. (From Strasburger after Wiesner.)

hypha. The hypha, upon coming into contact with an alga cell,
grows around it to form a lichen. ~~ >

Seales or soredia aré found on many lichens. These consist
of a network of hyphze enclosing algal cells. By becoming
detached from the parent plant, they develop new lichens and so
constitute a means of vegetative propagation.

Usrs.—To the pharmacist and chemist lichens are chiefly of
interest because of the coloring principles which they contain.
Species of Lecanora and Roccella tinctoria yield, when subjected to
fermentation, the dyes orcein and litmus. Litmus is one of the
best indicators in volumetric analysis. Cudbear, a purplish-red
powder, used extensively for coloring pharmaceutical prepara-
tions in the form of tincture, is prepared by treating species of
Roccella, Lecanora or other lichens with ammonia water. Other

THE LICHENS 417

lichens, such as Cetraria islandica, various species of Parmelia,
Usnea and Alectoria, have been used in medicine because of
demulcent principles which they contain.

One of the most conspicuous lichens in northern latitudes is
the Reindeer Moss (Cladonia rangiferina). It is a slender,
branched, fruticose form which grows to the height of about a
foot and bears its brownish apothecia at the ends of branches.
In Arctic regions it occurs in dense patches covering large areas
and forms the only kind of fodder available in many sections for
reindeer and caribou.

Another lichen, Lecanora esculenta, is thought to represent the
source of Scriptural Manna. It has been employed as a food in
periods of famine by the natives of southwestern Asia.

CHAPTER XIX

THE BRYOPHYTES

This interesting group, comprising the second division of the
Vegetable Kingdom, is represented by the liverworts and the
mosses. These plants show a beginning of definite alternation of
generations, 7.¢., gametophyte (sexual_phase) alternating with _
sporophyte (asexual phase of development) i in their life history, the
two phases being combined in one plant. The sporo _Is
attached to the gametophyte throughout its existence, being
parasitic upon rit. Of the two Pp hases, the. gametophyte i iS always
the larger. r and more conspicuous structure. ‘The female sexual
cell is always lodged in an archegonium (a multicellular female
sexual organ).

There are two classes of Bryophytes: Crass I, Hepatic
or liverworts and Cass II, Musci or mosses.

HEPATIC2 OR LIVERWORTS

The liverworts are a class of aquatic or terrestrial plants
occurring in tropical and temperate regions. While most of
them live in_ moist land situations, some are found submerged or
floating in water, some are epiphytic, and others occur in localities
where long dry seasons prevail. The body of a liverwort either
consists of (1) a small, flat, dichotomously branching thalloid
body showing differentiation into a green upper or dorsal region
and a nearly colorless under or ventral part, as in Riccia or
Marchantia, or (2) of a branching stem which bears two rows of
dorsal leaves containing chloroplasts and one row of pale ventral
leaves, as in Porella and other higher liverworts. Reproduction
is by fertilization or by vegetative multiplication. Some species
develop both kinds of sexual organs (antheridia and archegonia)
on the same gametophyte and so are monecious, while others
develop antheridia on one gametophyte and archegonia on

another gametophyte and are therefore diwcious.
418 gees

THE BRYOPHYTES.

There are three orders of liverworts, viz.:

ORDER 1.—MArCHANTIALES, including Marchantia and Riccia.

ORDER 2.—JUNGERMANNIALES, the leafy liverworts, including
Porella.

(3
[A
Pike

Fic. 311.—Marchantia polymorpha. Male gametophyte. Lower figure: Thalloid
body with cupules containing gemmz and male receptacles (antheridiophores) in
various stages of development, a, 6, c, d. Upper figure: antheridia in depressions
of upper surface of disk of male receptacle, in successive stages of development,
hukeew sé depression with the antheridium taken out. (From Small after Kny.)

ORDER 3.—ANTHOCEROTALES, having the most complex
sporophyte generations among liverworts, including Anthoceros
and Megaceros.

MarcuantTiA (A TypicaAL Liverwort)

One of the most common and widely distributed liverworts is
| Marchantia polymorpha. It occurs in dense patches on the moist

426. PHARMACEUTICAL BOTANY

surface of soil in flower pots of greenhouses and on the damp
rocks and humous soil of woodlands.

GAMETOPHYTE GENERATION.—The gametophyte or sexual
plant consists of a dichotomously branched thalloid body with a
distinct midrib. With a hand lens it will be noted that its green
upper surface exhibits numerous pale green lines which divide it
into small rhomboidal-shaped areas. In the center of each of
these is a pore which leads into an air-chamber within the body.
Its lower surface is nearly colorless and exhibits rhizoids and
delicate scales.

Guard Cell _ mina Guard Cell Upper Epidermis

Air Chambers

Partition sey rs 2 ce cae ae
between Air--> a & g

Chambers

Lower
Epidermis” ~

Rhizoid

Fic. 312.—Marchantia polymorpha. Microscopical structure of the thallus which

shows dorsioventral differentiation. A, vertical section showing one large air-

cavity beneath the epidermis. B, C, D, portions of each of three kinds of rhizoids,
the last two with inner thickenings of the wall. (Moittier.)

A cross section of the thalloid body, when examined micro-
scopically, will disclose an upper epidermis of epidermal cells
containing chloroplasts and showing stomata, each stoma sur-
rounded by four vertical rows of cells. Directly beneath this is a
row of air-chambers separated by partitions. In each air-cham-
ber branched rows of densely chlorophylloid cells grow upward from
the cells which constitute its floor. ‘These do most of the work
of photosynthesis. Below the air-chambers are several rows of
clear parenchyma cells containing only a few chloroplasts.
From the lower surface of these are to be noted outgrowths as
slender rhizoids and scales.

Some of the gametophytes will show cup-like structures on
their upper surface. These are called cupules. They contain

THE BRYOPHYTES 421

special buds-called gemma which, upon being detached, may give
rise directly to a new gametophyte. This is the vegetative
method of reproduction.

Marchantia is diecious for it produces two kinds of gameto-
phytes, ope bearing antheridia (male) and another archegonia

TK t-4 x :
Li. AL Secepe
pete,

, F
een
a
a]

&
t
:

i
i
|

Fic. 313.—Marchantia polymorpha. 1, Vertical section of one lobe of the disk
of the archegonial receptacle, showing archegonia on lower surface, the group
being surrounded by a laciniate perichaetium. 2a, 2a2, 3, 4, 5, 9, 10, Successive
stages in the development of the archegonium. 6, 7, 8, Transverse sections of the
same. 11, Mature archegonium with antherozoids attracted to mouth. 12,
Transverse section of venter of mature archegonium. 13, Transverse section of
neck of same, showing six cells. 14. Tuberculate rhizoid. 15, Transverse section
of same. (From Small after Kny.)

(female). There is no way of distinguishing one from the other
until stalked structures or receptacles arise from the upper surface
of the gametophyte.

The antheridial receptacle consists of an erect stalk surmounted
by a disk with an eight lobed edge. Embedded in numerous
separate cavities of the upper region of the disk are the antheridia.

~

422 PHARMACEUTICAL BOTANY .

The archegonial receptacles consist of a stalk surmounted by an
umbrella-shaped disk with about nine somewhat pendant rays.
On the lower surface of the disk the archegonia arise as flask-
shaped structures which hang downward.

The anthertdia are somewhat oval and consist of a wall of sterile
cells surrounding a mass of sperm mother cells. The latter give rise
to the sperms or antherozoids which are biciliate.

The archegonia are flask-shaped structures, each of which
possesses a basal bulbous part called the venter and an elongated
tubular part termed the neck. The flask has an outer layer of
stertle cells and a central axial row of cells which from the base of
the venter upward are the egg cell, ventral canal cell and neck canal
cells. As the archegonium matures, the horizontal walls of the
canal cells are dissolved by an enzyme and the contents of the -
cells are transformed into a mucilaginous mass. During wet
weather the upper portion of the wall of the mature antheridium
bursts open and the biciliate sperms are liberated in a mucilagi-
nous mass. Attracted by the substances then issuing from the
mouths of the archegonia, they swim toward them and usually a
number pass down the neck of the canal of each archegonium
and gather around the egg. One of them unites with the egg
fertilizing it and forming an oospore. The act of fertilization
concludes the gametophyte generation.

form a spherical mass of cells which in time become differentiated
to form a sporophyte or asexual plant which consists of (1) a basal
foot which attaches the sporophyte to the base of the archegonium
and absorbs nutrient substances in solution from it, (2) a short
stalk which gradually lengthens and (3) a capsule composed of an
outer layer of sterile cells enclosing spores and slender, elongated
cells called elaters. :

Upon the rupture of the wall of the capsule the spores are
discharged, being aided in their dispersal by the elaters found
among the spores which, being hygroscopic and with spirally
thickened walls, bend and twist, giving force to the process of
spore distribution. Each spore, upon absorbing water, may
develop into a gametophyte.

THE BRYOPHYTES 423

ALTERATION OF GENERATIONS.—In the life history of Mar-
chantia there are two distinct generations. The gametophytes
bear the sex organs in which gametes (sperm and egg) are pro-
duced. The sperm unites with the egg forming an oospore.

SSS ue

Fic. 314.—Marchantia polymorpha. 1. Thalloid body of Female Gametophyte -
with archegonial receptacles (archegoniophores) in successive stages of develop-
ment, a1, a2, b,c, d,e,f. 2,3, Upper and under views of archegoniophore when the
sporophytes are ripe. 4, 5, Stages in the development of the sporophyte. 6,
Young sporophyte enclosed by a perigynium. 7, Mature sporophyte shedding
spores. 8, Elater and spores. 9, Surface view of cells from wall of capsule.
(From Small after Kny.)

The oospore develops into a sporophyte which forms spores.
Each spore may develop into a gametophyte.

male gametophyte—sperm raha sporophyte—
female gametophyte—egg eee
male gametophyte

steed female gametophyte

424 PHARMACEUTICAL BOTANY

Musci or MosseEs

Plants found on the ground, on rocks, trees and in running
water. Their life histories consist of two generations,

pF e

Fic. 315.—Sphagnum acutifolium, Ehrb. A, prothallus (pr), with a young leafy
branch just developing from it; B, portion of a leafy plant; a, male cones; ch, female
branches; C, male branch or cone, enlarged, with a portion of the vegetative
branch adhering to its base; D, the same, with a portion of the leaves removed so
as to disclose the antheridia; FE, antheridium discharging spores; F, a single sperm;
G, longitudinal section of a female branch, showing the archegonia (ar); H, longi-
tudinal section through a sporogonium; sg', the foot; ps, pseudopodium; ¢, calyptra;
sg, sporogonium, with dome of sporogenous tissue; ar, old neck of the archegonium;
F. Sphagnum squarrosum Pers.; d, operculum; c, remains of calyptra; gs, mature
pseudopodium; ch, perichatium. (Gager, from Schimper.)

gametophyte and sporophyte similar to the liverworts but differ
from liverworts, generally, by the ever-present differentiation of

the gametophyte body into distinct stem and simple leaves, and

THE BRYOPHYTES 425

the formation of the sexual organs at the end of an axis of a
shoot. They are either monecious, when both kinds of sexual
organs are borne on the same plant, or diwcious, in which case
the antheridia and archegonia arise on different plants.

OrpvER 1.—SPHAGNALES, or Bog Mosses, including one
family, Sphagnacee with the single genus, Sphagnum which has
many species. Pale mosses of swampy
habit whose gametophytes grow
indefinitely at the apices of the stems
repeating their growth periodically
while their lower portions die away
gradually and form peat, hence their
frequent name of Peat Mosses. Peat
in a dry state is used as fuel. The
peat mosses possess a protonema
which differs from that of other mosses
in being thallose and bearing a resem-
blance to the prothallus of a fern.
The leafy stem ends in a three-angled
apical cell and bears a branch at every
fourth leaf. The branches divide to fy
form secondary branches. The an- }
theridia arise on small lateral branches
in the leaf axils, each one consisting of
a slender stalk and a globular capsule.
The archegonia arise on the leafy tips of
stems. The zygote gives rise to the

Fic. 316.—Surface view of
portion of a leaf of Sphagnum
palustre showing reservoir cell (r);
Sporophyte which possesses a large foot, transverse bands that prevent

a small stalk and a spherical sporan- cells from collapsing (h); pores

in reservoir cells (p); and green

gium the end of which splits offasa 4. Sauuiaiae chiceopayil ()-

lid to allow the spores to escape.

A number of species of Sphagnum have been employed in
surgery as absorbents in place of gauze. For this purpose they
must be thoroughly cleaned and sterilized.

The power which bog mosses have to absorb water depends
on the presence of large chlorophyll-less cells the walls of
which have pores and are supported by spiral or ring-like
thickenings. These cells absorb water by capillarity. Between

426 PHARMACEUTICAL BOTANY

these empty cells are elongated narrow cells containing
chloroplasts.

OrDER 2.—ANDREAALES Or Rock Mosses consisting of one
family, Andreeacee with the single genus Andreea, of xerophytic
habit, occurring on siliceous rock. ;

OrpDER 3.—BRYALES, or true mosses, comprising the most
highly evolved types of bryophytes. Examples: Polytrichum,
Funaria, Hypnum, and Mnium.

LirE History oF PoLyTricHuUM CoMMUNE (A TypicaAL TRUE
Moss).—Polytrichum commune is quite common in woods, forming

Fic. 317.—Protonema of a Moss. A, Early stages of germinating spores: v,
vacuole; w, rhizoid; s, spore wall or exosporium. B, Developed protonema
about three weeks after germination; 4, primary filament; k, bud and w, rhizoid.
The bud develops into a gametophytic plant. (After Sachs from Small.)

a carpet-like covering on the ground beneath tall tree canopies.
It is dicecious, the plants being of two kinds, male and female.
Beginning with a spore which has fallen to the damp soil, we
note its beginning of growth (germination) as a green filamentous
body composed of chloroplast-containing cells arranged end to
end. This is called a protonema. ‘This protonema soon becomes
branched, giving rise to hair-like outgrowths from its lower por-
tion called rhizoids and lateral buds above these which grow into
leafy stems bearing rhizoids at their bases and which are com-
monly known as ‘‘moss plants.” At the tips of some of these
leafy stems antheridia (male sexual organs) are formed while on
others archegonia (female sexual organs) are developed. The leafy

THE BRYOPHYTES 427

Fic. 318.—Hair-cap moss (Polytrichum commune). A, male plant; B, same,
proliferating; C, female plant, bearing sporogonium; D, same; g, gametophyte; s,
seta; c, capsule; 0, operculum; a, calpytra; E, top view of male plant. (Gager.)
organs are surrounded at the tips by delicate hairy processes
called paraphyses as well as leaves for protection. The antheridia
bear the antherozoids, the archegonia, the eggs or ova, as in
the liverworts. When an abundance of moisture is present, the

428 PHARMACEUTICAL BOTANY

antherozoids are liberated from the antheridia, swim through the
water to an archegonium and descend the neck canal, one fertiliz-
ing the egg by uniting with it. This completes the sexual or
gametophyte generation. The fertilized egg now undergoes

ed

\ Ws
Qi icici
AN Y
: ee ie i
SHA j | y
Yo NY Nest i} f yt lH t
AS WALA

Fic. 319.

Fic. 320. Fig. 321.
Fic. 319.—Vertical section through summit of male gametophyte of a Moss.
Mnium hornum (X 40) an, Antheridia; par, Paraphyses.
Fic. 320.—A, Antherozoid of Moss (X 1,200); B, Antherozoid of fern (X 700).
Fic. 321.—Funaria.—A, Apex of female gametophyte with archegonia. B
Neck of archegonium, showing mode of separation of the ca
archegonium. (From Small after Sachs.)

BI

P cells. C, Immature

division until an elongated stalk, bearing upon its summit a
capsule, is finally produced, this entire structure being known as
the sporogonium. The base-of the stalk remains imbedded
in the basal portion of the archegonium, at the tip of the leafy
stalk, and forms a foot or absorbing process. In growing upward
the sporogonium ruptures the neck of the archegonium and

THE BRYOPHYTES 429

carries it upward as the covering of the capsule, or calypira.
The calyptra is thrown off before the spores are matured within

Fig. 322. Fic. 323.

Fic. 322.—Funaria hygrometrica. A, Young sporophyte (c) attached to gameto-
phyte (g). B, Adult sporophyte with seta (s) and capsule (f); gametophyte (g),
part of which forms the calyptra (0). C, Vertical section of capsule (X 80).
Operculum (d); Peristome (p); Annulus (2); Air-space with trabeculz or filaments
(t); Sporogenous layer (s). (From Small, after Prantl.)

Fic. 323.—Surface view of peristome of Funaria, after removal of operculum.
pf, peristome tooth. (After Small.)

the capsule. The upper part of the capsule becomes converted
into a lid or operculum at the margin of which an annulus or ring of
thick-walled cells forms. Attached to the annulus by their bases

Male Votant Sperm
. Nantheria ee Fertil ized—Sporophyte —Spore
Female plant—Archegonium —Egg egg

Fic. 324.—Diagram of the life history of a moss. (After Atwood.)

Spore Protonema ~<e

are a number of long triangular teeth called collectively the
peristome. The cells of the annulus are hygroscopic and expand

430 PHARMACEUTICAL BOTANY

at maturity, throwing off the lid. The peristome teeth are very
hygroscopic. During rainy weather or when the atmosphere is
very humid they bend inward preventing the escape of the spores.
During dry weather they straighten out and in this process lift
_ out some of the spores and allow others to sift out from the cavity
of the capsule. This completes the asexual or sporophyte
generation. The spores falling to the damp soil germinate into
protonemata, thus completing the life cycle in which is seen an
alteration of generations, the two phases, gametophyte alternat-.
ing with sporophyte.

CHAPTER XX
THE PTERIDOPHYTES

The Pteridophytes or Vascular Cryptogams constitute the
third great division of the Plant Kingdom. ‘They include the
plants known as ferns, horsetails or scouring rushes, club mosses
and quillworts. These are the most highly developed crypto-
gams, showing not only a distinct alternation of generations in
their life histories, but a considerable structural advance over
the Thallophytes and Bryophytes. They differ from the Bryo-
phytes (1) by the sporophyte becoming an independent plant
which shows differentiation into true root or roots, stem, and
leaf or leaves each of which possesses a well organized vascular
bundle ‘system, (2) by the gametophyte body or prothallus
nourishing the young ~ sporophyte only until the latter has
produced its own root and leaf after which the gametophyte
perishes, and (3) by presenting large, conspicuous sporophytes
and very small gametophytes.

Pteridophytes formed the chief vegetation of the earth during
the Carboniferous period.

The Pteridophytes are grouped into three classes, viz.:

Crass I. FiiciInE# or FERNS.

Cuass II. EguiseTinE2# or Horseralts.

Crass III. LycopopInE#, THE CLuB MossEs_ AND
QUILLWORTSs.

THe FERNS

The ferns, constituting the class Filicinea, represent the largest
group of the vascular cryptogams.

Ferns occur in a variety of situations in tropical and temperate
countries. Most of them prefer damp, shady places. Some
grow in forests and fields, others in the crevices of rocks, still
others in water. In the tropics, some ferns become tree-like in
form, possessing long erect trunks bearing at their summit a

crown of compound leaves called fronds. In temperate regions,
431

432 PHARMACEUTICAL BOTANY

the land ferns have underground stems or rhizomes from which
fronds arise.

Fic. 325.—Dryopteris marginalis, a fern whose rhizome and stipes constitute the drug,
American Aspidium.

The main axis of the majority of the typical land-growing
ferns is a creeping underground stem or rhizome which at its
various nodes bears rootlets below and fronds above. These
fronds are highly developed, each being provided with a petiole-

THE PTERIDOPHYTES 433

like portion called a stipe which is extended into a lamina usually
showing a forked venation. Some ferns possess laminae which
are lobed, each lobe being called a pinna. If a pinna be further
divided, its divisions are called pinnules. In some ferns the
pinnules are further subdivided into secondary pinnules, as in the
“brake? or “bracken,” Pterts aquilina. The unfolding of a
frond is circinate and it increases in length by apical growth.
On the under surface or on the margin of the lamina, pinnz, or
pinnules may be seen small brown patches each of which is called
a SOrUs, and usually covered by a protective membrane called the
indusium. Each sorus consists of a number of sporangia (spore
cases) developed from epidermal cells. In some ferns the entire
Ieaf becomes a_spore-bearing organ (sporophyll). Many
sporangia have a row of cells around the margin, the whole being
called the annulus. Each cell of the annulus has a U-shaped,
thickened cell wall. Water is present within these cells and when
‘it evaporates it pulls the cell walls together, straightening the
ring and tearing open the weak side. The annulus then recoils
and hurls the spores out of the sporangium. Upon coming into
contact with damp earth, each spore germinates, producing a
green septate filament called a protonema. This later becomes a
green heart-shaped body called a prothallus. It develops on its
under surface antheridia or male organs and archegonia or female
organs as well as numerous rhizoids. Within the antheridia are
developed motile, many ciliated sperms, while ova are produced
within the archegonia. The many ciliate sperms escape from
the antheridia of one prothallus during a wet season and, moving
through the water, are drawn by a chemotaxic influence to the
archegonia of another prothallus, pass down the neck canals of
these and fuse with the ova, fertilizing them. The fertilized
egg or odspore divides and redivides and soon becomes differ-
entiated into a stem, first leaf, root, and foot. The foot obtains
nourishment from the prothallus until the root grows into the soil,
when it atrophies, and the sporophyte becomes independent.
Unequal growth and division of labor continue until a highly
differentiated sporophyte results, the mature “fern plant.”

In some species of ferns as the Cinnamon Fern (Osmunda
cinnamomea) and the Sensitive Fern (Onoclea sensibilis), two kinds

434 PHARMACEUTICAL BOTANY

of fronds are produced, sterile fronds which represent green foliage
leaves and fertile fronds which contain little or no chlorophyll and
bear clusters Of sporangia.

There are two sub-classes of Filicineze, viz.:

SuB-cLass EusPORANGIAT&, including ferns like Ophioglossum
or adder’s tongue and Botrychium or moonwort in which the

sporangium originates from a group of epidermal cells and

2h
ite

ee

Fic. 326.—Fern leaves, showing various degrees of subdivision or branching of the
blade. A, Phyllitis; B, Polypodium; C, Pteris; D, Adiantum. (From Gager.)

contains similar spores, the latter giving rise to prothallia which
bear both antheridia and archegonia.

Susp-cLass LEPTOSPORANGIAT& which include most of the
common ferns and the water ferns. In these the sporangium
develops from a single epidermal cell. Most of them have
sporangia containing similar spores but the water ferns are
heterosporous.

Orver 1.—FivicaLes or TRuE Ferns (Homosporous)

Famity CyATHEACE®.—Mostly tree-like ferns of the tropics
and subtropics possessing sporangia with a complete annulus.

THE PTERIDOPHYTES 435

The long, silky, yellow or brownish hairs obtained from the bases
of Cibotium glaucum, C. Barometz, Alsophila lurida, and other
members of this family growing in Java and Sumatra have been
successfully used in checking external hemorrhages under the
names of “‘Pengawar Djambi’’ or “Golden Moss.”

Famity PotypopiAce&#.—In this family the capsule of the
sporangium possesses a vertical, incomplete annulus which splits

ph end
Fic. 327.—Cross section of the main axis or rhizome of the common brake,
Pteris aquilina. X8. ep, epidermis; scl, sclerenchyma fibers in hypodermis; 4,
xylocentric vascular bundles; sc/b, sclerenchyma band composed of lignified fibers;
r, lateral ridges; pa, parenchyma; x, xylem; s?, sieve tissue (phloem) ; end, endodermis;
ph, phloem of a vascular bundle.

the mature sporangium transversely. It includes most of the
more common land ferns of temperate regions. Outstanding
genera of this family are Pteris or brake, Polypodium or polypody,
Adiantum or maiden hair, Asplenium, or spleenwort, Dryopteris
or wood fern, Camptosorus or walking leaf, Polystichum or christmas
fern, Pellaea or cliff brake and Phegopteris or beech fern.

The rhizomes and stipes of Dryopteris Filix-mas and Dryopteris
marginalis are valuable tape worm remedies. , Those of the former
species are official under “Aspidi wey the U.S.P. The
vascular bundles of the stems and leaves of these and other

«

436 PHARMACEUTICAL BOTANY

ferns are concentric in type but differ from the concentric vascular
bundles of the stems and leaves of some monocotyledons in that
xylem is innermost and phloem surrounds the xylem. This kind
of concentric bundle is called a xylocentric bundle. For further
details on fern structure see Chapter IV. The dried fronds of
the native Adiantum pedatum and of the European A. Capillus-
Veneris are used in domestic medicine as a demulcent expectorant
in the form of syrup or infusion.

Famity OsmMuNDACE&.—This family is characterized by
having short sporangia, the annulus of which consists of a group of

is 5

Fic, 328.—Cyrtomium falcatum. Under (dorsal) surface of a portion of a sporophyll,
showing the numerous sori on the pinnae. (Gager.)

thick walled cells near the apex on either side of the wall of the
sporangium, the rest of the wall becoming gradually thinner
walled. The mature sporangium splits open longitudinally.
- It is represented in North America by the genus Osmunda, three
species of which, O. regalis, the royal fern, O. cinnamomea, the
cinnamon fern and O. claytoniana, the interrupted fern, occur
locally. The rhizomes and stipe bases of the last named have
frequently been found as substitutes and adulterants of Aspidium.

ORDER 2.—HybRopTERALES OR WATER FERNS (HETEROSPOROUS)

FamiLty Satvintace&.—Floating ferns with broad floating
leaves and submerged dissected leaves which bear fruit bodies
called sporocarps containing microsporangia and megasporangia.
The microsporangia contain microspores which give rise to male
prothallia. The megasporangia contain megaspores which

THE PTERIDOPHYTES 437

develop female prothallia. Examples: Saluinia. Marsilea or the
Water Shamrock, and Azolla.

THe HorseEtTAILs

The Equisetinee, commonly known as the Horsetails or Scour-
ing Rushes, are perennial herbs with hollow, cylindrical, jointed
and fluted stems, sheath-like whorls of united leaves and terminal
cone-like fructifications. Their bodies contain large amounts of
silicon, hence the name scouring rushes. Only 25 species exist
today and all of these belong to the genus Equisetum. ‘These
are believed to represent degenerate descendants of tree-like
Horsetails, fossils of which have been found in coal-bearing rocks
of the Carboniferous period. :

In some species of Equisetum the fruiting cone is borne on the
ordinary aerial stem, in others on a special’ stem of slightly
different form. The spores are provided with ribbon-like
appendages called elaters, which, being hygroscopic, coil and
uncoil with increase or decrease in the amount of moisture
present, thus aiding in the entanglement of the spores into small
masses, and so preventing isolation, an advantage on account of
the « dioecious character of the prothallia which they originate.

Lire History or EquisETUM ARVENSE, A HorseTAIL

Equisetum arvense, a frequently occurring species along railroad
embankments but also in fields, may be taken as a type form of
the group. Like other horsetails its life history involves an
alternation of sporophyte and gametophyte generations of which
the sporophyte is the more conspicuous.

The spompkge consists of a subterranean, horizontally-
branched, food-storing rhizome (main axis) bearing on its lower
surface water- and mineral nutrient absorbing-roots and on its
upper surface 2 kinds of jointed, hollow shoots—sterile and fertile.
The sterile shoots are green and bear whorls of shortened, green
branches from their upper nodes and scale-like leaves from their_
lower nodes, Theirs is the work of manufacturing food for the
plant. The fertile shoots are nearly or completely devoid of
chlorophyll and bear whorls of scale-like leaves at their nodes and
terminate in a cone. The central stalk of this cone is a prolonga-

438 PHARMACEUTICAL BOTANY

tion of the shoot axis and bears numerous whorls of sporophylls.
Each sporophyll consists of a short stalk and a disk; the latter is
arranged in peltate fashion on the end of the stalk and bears a
number of sporangia on its lower surface. Within these sporangia
are produced numerous spores which are liberated upon the

jn epyens
Tt

a

Goat AS
\ V3
YSAY if iy

é

Fic. 329.—Equisetum arvense. P, sterile branch; P', fertile branch with strobilus,
or cone; R, rhizome (underground); 7, cross-section of cone, showing insertion of
sporophylls in a whorl; V, V1, sporophylls with pendant sporangia; S, S, S?, spores
with coiled elaters (el). (Gager.)

ripening of the sporangia. While alike as to size, some of them
give rise upon germination to male prothallia, others to female
prothallia. These prothallia represent the sexual or gametophyte
plants. The male prothallia produce antheridia along the margins
or tips of their lobes in which, when mature, numerous many-
ciliated antherozoids are found. ‘The female prothallia produce

ho Benin ia a
reek

THE PTERIDOPHYTES 439

flask-shaped archegonia on the prothallial cushion each of which
contains anegg. Both kinds of prothallia bear absorptive rhizoids
on their lower surace. The antherozoids, liberated from the
antheridia of male prothallia, swim to the archegonia of neigh-
boring female prothallia, enter their mouths, pass down their
neck canals and gather about the eggs. In each instance the best
adapted one fertilizes the egg. The fertilized egg (odspore)
develops into an embryo which through continued growth becomes
a mature sporophyte plant.

THE Cius MossEs

The Class Lycopodinee or club mosses consists of small peren-
nial, vascular, dichotomously-branched, trailing, evergreen
herbs with stems thickly covered with awl-shaped leaves. They
are represented, today, by over 900 extant species. ‘These are
relatives of giant tree-like species of club mosses such as Lepido-
dendron and Sigillaria which with the giant horsetails and ferns
formed the chief flora during the Coal Age. They differ from
ferns and horsetails in possessing a suspensor, like seed plants, in
their early sporophyte stage and in having biciliate sperms.
Except in a few instances the sporangia are borne on sporophylls
crowded together and forming cones or spikes (strobili) at the
ends of erect branches or as lateral branches. The spores found
within the sporangia are either alike or unlike in size, hence the
plants are either homosporous or heterosporous. In heterospory the
small spores give rise to male gametophytes and the large spores
to female gametophytes.

Famity I.—Lycopopiace#, including the single genus
Lycopodium with widely distributed species. The group is
homosporous. The spores of Lycopodium clavatum are official.

Lire History or Lycopoprum CLAvAtuM.—This species of
club-moss thrives in temperate and sub-tropical forests of both
hemispheres and yields a light-yellow, mobile powder consisting
of spores. This powder is of value in pharmacy to prevent the
adhering of suppositories, pills, etc., and in medicine as a protec-
tive to tender surfaces. When examined under the microscope
the spores appear tetrahedral with one convex and three plane
surfaces.. The outer wall (exosporium) of each is reticulated,

440 PHARMACEUTICAL BOTANY

Yaar,

SJ i Y
f WI fae

a

Fic. 330.—Lycopodium clavatum L. A, portion of sporophyte plant showing
habit (natural size); a, foliage leaf; 5, sporophyll, with kidney-shaped sporangium
at base (magnified); c, d, ¢, spores viewed from different aspects (X 580); B
prothallus of another species. When the sporophylls are mature, the spikes are

cut and dried, and the spores separated by shaking. They are freed from debris by
sieving through flour-sieves. (After Luerssen.)

>

THE PTERIDOPHYTES 441

- showing meshes that are surrounded by polygonal ridges with —
straight sides. In nature, these spores, falling out of the ripe
spore cases, germinate on moist soil, each producing a fleshy
prothallus which is submerged in its lower portion and exposed
above ground in its upper part. This structure has antheridia

. Fic. 331.—Selaginella Martensii. a, vegetative branch; 6, portion of the stem,
bearing cones (x); c, longitudinal section of a cone, showing microsporangia (mic.
sp.) in the axils of microsporophylls, and megasporangia in the axils of megasporo-
phylls; ¢, microsporangium with microsporophyll; ¢, microspores; f, portion of wall
of sporangium, greatly magnified; g, megaspore; h, microsporangium opened, and
most of the microspores scattered; i, megasporangium, with megasporophyll; ,
same, opened, showing the four megaspores. (Gager.) :

and archegonia imbedded in its upper end and shows elongations
of its epidermal cells as rhizoids in its tuberous, sunken, lower part
which absorb nutriment from the soil. The mature antheridia
burst open during wet weather and liberate biciliate antherozoids.
Each mature archegonium produces an ovum in its basal cavity.
Antherozoids (sperms) swim to the mouth of the. archegonium,
pass down its canal and gather about the ovum.

442 PHARMACEUTICAL BOTANY

One of these fertilizes the ovum forming an odspore (zygote).
The odspore begins to divide and ere long develops into an
embryo which soon becomes differentiated into foot and shoot
regions. ‘The foot absorbs nourishment from the prothallus, for
a short time, until the first root commences to function. The
shoot elongates carrying ‘the first leaves above the soil and giving
rise to the first root or radicle at its base. Through continued
growth and differentiation a mature sporophyte is developed which
consists of a creeping, prostrate, dichotomously branching stem
covered with crowded, light green, linear, awl-shaped, and
_bristle-tipped leaves and bearing several ascending fertile.
branches which terminate in one or two spikes. Each of these
spikes consists of a slender axis bearing numerous more or less
deltoid, bristle-tipped scales, each one having a kidney-shaped
sporangium at the base of its upper surface containing numerous
Spores.
FamiLy II.—SE.LaGINELLACEZ# (Little Club Mosses), includ-
ing the single genus Se/aginella with species for the greater part
tropical. Plants similar in habit to the Lycopodiacee but produc-
ing microspores and megaspores, hence showing heterospory.
The microspores are formed in microsporangia which are borne

all

for ornament. One of them, Selaginella lepidophylla, a native of
Mexico, is a “resurrection plant,” curling up if allowed to dry
and, when apparently dead, capable of being revived in water.
Famity III.—Isorracra (Quillworts) including the single
genus Isoetes whose species are plants with short and tuberous
stems giving rise to a tuft of branching roots below and a thick
rosette of long, stiff, awl-shaped leaves above. They produce
microspores and megaspores and so show heterospory.

CHAPTER XXI
THE SPERMATOPHYTES

This is the highest evolved division of the vegetable kingdom.
It includes all plants which produce real flowers and seeds. The
older name for the group is Phanerogamia. In this group the
sporophyte attains its greatest development while the gameto-
phytes are reduced to minute structures parasitic on the sporo-
phyte. The Spermatophytes are subdivided into two subdivi-
sions based primarily upon whether their seeds are borne naked
or enclosed within a seed vessel or pericarp. —

Subdivision I. Gymnospermz (Gymnosperms)—plants with
naked seeds. _

Subdivision II. Angiospermae (Angiosperms)—plants with
covered seeds. ao

THE GYMNOSPERMS

The Gymnosperms comprise an ancient and historic group of
seed plants which were more numerous in the Triassic and Car-
boniferous periods than now. They differ from the Angiosperms
in several respects, viz.: they bear naked ovules on the edges or
flat surfaces of leaves called carpels, while Angiosperms bear
covered ones; each ich_megaspore_ produces within itself a bulky
prothallus, in the upper portion of which originate one or more
archegonia, while in Angiosperms no recognizable prothallus has
been proven to exist; the stored food. tissue within their seeds is
prothallial tissue loaded with starch, etc., while that in Angio-
sperm seeds (endosperm) is developed from the endosperm
nucleus; the mode of growth of their stems is always indefinite
while that of Angiosperms is either indefinite or definite.

The groups still extant are the Cycads or Fern Palms, the
Gnetums, the Ephedras, the Ginkgos and the Conifers. Of
these the Conifers comprising over 300 species are the most
numerous. Many of them yield valuable products to pharmacy

and the arts.
443

444 PHARMACEUTICAL BOTANY

The Conifers include the pines, spruces, hemlocks, cedars, firs,
redwoods, yews, arbor vitae, chamzecyparis, and larches. All of
their number are evergreen except the European larch, which
drops its foliage upon the advent of winter.

i

Fic. 332.—The White Pine (Pinus Strobus).

Lire History oF A GyMNosPERM (Pinus Stropus)

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 expectorant properties and is official in the National
Formulary.

THE SPERMATOPHYTES 445

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 ina terminal bud. The trunk rarely exceeds
3 feet in diameter and 125 feet in height and is averagely 11% to

ae * ee sith 4 : Siidier ae
: ‘i : : a : ae persis i

Fic. 333.—Transverse section of white pine stem of four years’ growth, showing
cork (a), cortex (6), phloem (c), cambium (d), xylem (e), secretion reservoir ( i,
pith (g) and medullary-ray (4). (Photomicrograph.) % 16.

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 emanating 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, grad-
ually shorter until the summit is reached, giving to the crown
or upper part of the tree the appearance of a pyramid. These

446 PHARMACEUTICAL BOTANY

branches give rise to other branches which agree with the lateral
branches in ending in terminal buds. Another kind of branch,
however, is found which is always shorter than the former
branches. This type of branch is called a ‘spur shoot” and arises
from the former branches. The spur shoots bear the needles or

See ace ewe ; oa

Fic, 334.—Transverse section of the needle (leaf) of the white pine. Highly
magnified. ¢p, epidermis; s, stomata; m, infolded parenchyma cells of mesophyll;
c, cuticle; r, oil reservoir; f, colorless central parenchyma corresponding to the pith
of the stem; 7, endodermis; x, xylem and fh, phloem of collateral vascular bundles.
The xylem is composed of tracheids with bordered pores. The leaf shows xero-
phytic-centric structure.

foliage leaves which are light-green, when young, and bluish-green,

soft, flexible, 21¢ 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 sheath. These foliage leaves persist until
the end of their second year, when they are shed with the spur
shoot which bears them.

THE SPERMATOPHYTES 447

The white pine, like most of its allies among the Conifer,
bears cones. ‘These structures are of two kinds, viz.: staminate
and carpellate. Both kinds are produced on the same tree.

STAMINATE ConeEs.—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 called microspor-
ophylls or stamens. On the under surface of each scale are the

Fic. 335.—Staminate cones of the Austrian pine (Pinus austriaca). Below, before -
shedding pollen; above, after shedding. (Gager.)

spore-cases (microsporangia), which develop the microspores (pollen
grains). A section of the microsporangium shows it to consist of
a wall beneath which is a layer of cells called the tapetum and in
the interior a large number of spheroidal spore mother-cells.
Each of the spore mother cells divides into four spores. During
this process, reduction division takes place and the chromosomes
are reduced to one-half the number, so that the spores are haploid
as those of the fern. Each pollen grain when mature consists of
a central fertile cell and a pair of air-sacs 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.

448 PHARMACEUTICAL BOTANY

CARPELLATE Congs.—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 a lateral axillary branch. <A carpellate cone is
composed of a main axis which bears spirals of scales, termed

Fic. 336.—Scotch pine (Pinus sylvestris). A-D, stages in the development of
the carpellate cone, and its carpotropic movements. £, very young carpellate
cone much enlarged; F, ventral, G, dorsal views of a scale from £; 1, ovuliferous
scale, 2, ovule (in longitudinal section); 3, pollen chamber and micropyle leading
to the apex of the nucleus (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; J, longitudinal section of a mature cone; 6, ovule; 7, 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; ¢, embryo; f, remains of nucellus; L, embryo; c, cotyle-
dons; ¢, hypocotyl; r, root-end. (Gager.)

megasporophylls or carpels. Each megasporophyll is composed of an
ovuliferous scale bearing two ovules or megasori and a bract. Each
megasorus (ovule) is attached to the surface of the megasporo-
phyll. It contains a nucellus or megasporangium which is sur-
rounded by an integument, except at the apex where an opening,
the micropyle, is evident. The micropyle is the gateway to the pol-
len chamber which lies below it. Within the nucellus a single spore

THE SPERMATOPHYTES 449

mother cell is formed. This divides to form four spores but only
one persists. This is called the megaspore or embryo sac. It
contains haploid chromosomes.

DESCRIPTION OF THE GAMETOPHYTE GENERATION

The Gametophyte generation of the White Pine begins with
the development of the male and female gametophytes and ter-
minates with the fertilization of the egg.

Fic. 337.—Mature carpellate cones of white pine showing separated scales.

Tue Mate GametopHyTe.—The male gametophyte com-
mences 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 prothallial 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. (See Fig. 338.) At this
stage the pollen is shed and some of it is carried by air currents
to the carpellate cones where it sifts in between the ovule-bearing
scales and accumulates at the scale bases. 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

450 PHARMACEUTICAL BOTANY

fluid they germinate forming pollen tubes. The transfer of
pollen grains from the pollen sac to the pollen chamber and
consequent germination therein is called pollination. The con-
tents of a mature pollen-grain constitutes the male gametophyte.

Fic. 338.—Pine Pollen and development of pollen tube. In A the pollen grain.
is shown Containing prothallial cells (6). They have been cut off before the grain
was shed. The tube nucleus (7) has been divided off from the generative nucleus
(G).

In B the tube has been put forth and the tube nucleus has moved down into it.
Between the stages represented in B and C the generative nucleus has divided
to form a stalk nucleus (ST) and a body nucleus (B).

In C four nuclei are present in the pollen tube,—the tube nucleus (TJ), the stalk
nucleus (S7), and the two sperm nuclei (S). The sperm nuclei have been pro-
duced by a division of the body cell, the latter having disappeared. (After Atwood.)

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 large number of nuclei which are scattered throughout
the cytoplasm. Each nucleus accumulates around itself a por-

eR eRe

THE SPERMATOPHYTES 451

tion 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 (frothallus)
originate several archegonia.

_Antheridial
ae Cell

ik Spong
He Tice

Fic. 339.—Longitudinal section of one-year old ovule of Austrian Pine showing
- jmternal structure. (Mottier.)

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 contents of the mature embryo-sac constitutes the female gametophyte.

452 PHARMACEUTICAL BOTANY

After pollination the carpellate cone gradually bends on its
axis so as to ultimately reverse its position and hang downward.
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
Pe 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
t | egg nucleus and fertilizes it form-
| ing an odspore or zygote. The
aii 4} odspore contains the diploid
Fic. 340.—White pine (Pinus Strobus), NUMber of chromosomes.
Vertical section through the upper part = SEED FORMATION AND Dis-
peepee a Faia nha) a TRIBUTION.—The odspore under- -
tube-nucleus; arch, archegonium; en, Z0€S repeated divisions and
egg-nucleus. (Gager, after Margaret C. forms the embryo or young sporo-
Ferguson.) phyte 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 integument
becomes modified to form the hard protective seed coat. A por-
tion 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.

THE SPERMATOPHYTES 453

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.

Neck of

7 °
7 Archegonium

| Wall of
Megaspore

Fic. 341.—Austrian Pine (Pinus nigra). Longitudinal section of the upper end
of the endosperm showing two archegonia. The one at the right is mature and
ready for fertilization; that at the left is younger; the ventral canal cell is being cut
off. (Mottier.) :

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

454 PHARMACEUTICAL BOTANY

Outuine oF Lire History or Pinus Strosus
Oéspore

Embryo (Young Sporophyte)—(see 1)
7 Seedling Sporophyte (Composed of hypocotyl, cotyledons,

tap root, first leaf and bud)
Mature Sporophyte............. (see 2)

Staminate Cones hides ay Carpellate Cones
Microsporophylls (stamens) Megasporophylls (carpels)
Megasorus (Ovule) (see 3)
Microsporangium (Spore case) Megasporangium (nucellus)

Megaspore mother cell

Microspore-mother-cell Megaspore (Embryo sac)
Microspore (Pollen grain) Prothallus (Endosperm tissue)
Male Gametophyte (see 4) Female Gametophyte (see 5)
Pollination (transfer of pollen) Archegonium
sy , oan
One of Sperm Nuclei Ovum—egg nucleus
(Fertilization)
— Oéspore—

Embryo and “‘suspensor” in seed

1. Embryo is in a seed structure composed of a seed coat (developed from the
integument of the ovule, “‘megasorus’’) enclosing perisperm (developed from mega-
sporangial tissue, the nucellus). The perisperm surrounds endosperm (prothallial
tissue developed in embryo sac) and within the endosperm is the embryo which was
developed from a fertilized ovule within one of the archegonia. The seed wings
are developed from a portion of the scale leaf (ovuliferous scale or megasporophyll).

‘2. Mature Sporophyte consists of a root system, trunk, lateral branches, spur
shoots bearing foliage leaves in fascicles. Staminate cones are borne at base of
new growth. Carpellate cones are borne at end of a lateral axillary branch.

3. Megasorus or ovule is composed of an integument surrounding the mega-
sporangium (nucellus) except at apex where the micropyle (opening) leads to
pollen chamber.

4. Male Gametophyte—consists of the mature pollen grain containing two pro-
thallial cells, a tube nucleus and a generative cell.

5. Female Gametophyte consists of prothallial tissue (endosperm) with eee
archegonia, each containing an ovum. It is the mature “embryo sac.”

THE SPERMATOPHYTES

455

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

Gametophyte § t
Stage yeti

‘
\
\
‘
\
\
'
' 3

' tage
‘

i)

‘

'

Ly

\

Rex —O.— O- 6 ‘s
‘e— —@+g-6 4-0-4.

Fic. 342.—Diagram of life-cycle of a pine. (From Gager after Schaffner.)

CLASSIFICATION OF THE (GYMNOSPERMS

The gymnosperms are grouped into 4 orders, as follows
Order 1. Cycadales, the Cycads.

Order 2. Ginkgoales, the Ginkgos.

Order 3. Coniferales, the Conifers.

Order 4. Gnetales, the Gnetums and Ephedras.

ORDER CYCADALES

The Cycads or fern palms, the most primitive of the seed
plants, are represented by about 100 species of fern-like plants
that are believed to have been descended from the Cycadofilices, a

group of fossil plants which possessed the leaf characters of ferns
and ovules and seeds like the pines

Most of them have plant
bodies consisting of a thick cylindrical stem covered by the bases

of spirally arranged leaves and bearing upon its summit leathery,
pinnately-compound leaves and scale leaves. The male and
female cones are borne on different sporophytes. They are
represented by. the genera Cycas, Lamia, and Dioon.

456 PHARMACEUTICAL BOTANY

Lire Hisrory or ZAMIA

Aamia, a representation of the Cycads, possesses a plant body
or sporophyte which is short with a cylindrical erect stem con-

Fic. 343.—Cycas revoluta. The trunk of this cycad stores an enormous amount
of starch, It is one of the sources of Sago starch of commerce. Note the crown of
pinnately-compound leaves.

taining a large central pith which is surrounded by a circle of
open collateral bundles and by a broad cortex in whose cells
great amounts of starch are stored. The summit of the stem

THE SPERMATOPHYTES 457

is crowned by a cluster of pinnately-compound leaves of a
leathery texture.

STAMINATE ConE.—The staminate cone of Zamia arises on the
ends of short branches emanating from the end of the stem of

& Se ee

x 4.

Fic. 344,—Mature sporophytes of Zamia bearing staminate strobili (cones).
(Photograph by F. F. Bacon.)

the male plant (sporophyte). Each consists of a central axis bear-
ing compactly arranged microsporophylls. Each microsporophyll
bears on its lower surface a large number of microsporangia.
Within each microsporangium spore mother cells are developed,
each of which by division form 4 microspores (pollen grains).

458 PHARMACEUTICAL BOTANY

CARPELLATE CoONE.—The carpellate cone arises on a short
branch arising slightly from the end of the main stem of the female
plant. It consists of a fleshy central axis bearing peltate mega-
sporophylls. Each megasporophyll bears on the inner surface of
its expanded outer portion two megasort or ovules. Each mega-
sorus when mature consists of a megasporangium or nucellus sur-
rounded by a single integument, the latter being perforated by a
narrow tubular opening or micro-
pyle leading to a small cavity or
pollen chamber at the end of the
nucellus. (See Fig. 348.)

Within the nucellus is devel-
oped one megaspore mother cell
_ which by division gives rise to 4
megaspores or embryo sacs. Only
one of these persists, the other
three perish.

GAMETOPHYTES.—The male
gametophyte begins to develop
from the young pollen grain
(microspore) while the latter is still
within the microsporangium. The
pollen grain divides to form 2

Ais, RAO aging alae ce cells, a smaller one or prothallial
Zamia. B, microsporophylls bearing cell and a larger cell. "The larger
on their lower surfaces many micro- cell then divides to form a genera-

sporangia. (From “‘A Textbook of Gen- ti
@ C
dali Buswr® bp nite Dadi 6 ell and a tube cell. There

Moacidlles Co. Publiches results a 3-celled, undeveloped

male gametophyte or mature pollen
grain. ‘These are liberated by the bursting of the wall of the
microsporangium of the now ripened male cone with its separated
microsporophylls, and are carried by winds. Some are carried to
the ripened carpellate cones of female Zamias. Some of the
pollen grains sift between the megasporophylls and come to rest
about the ovules. Some of them become caught in a mucilagi-
nous fluid at the outer end of the micropyle and are sucked into
the pollen chamber. Upon reaching the pollen chamber they
resume their development. Each pollen grain germinates

THE SPERMATOPHYTES 459

size. The starch obtained from the stem of this plant is known as Florida arrow-
root. (Photograph by F. 7. Bacon.)

460 PHARMACEUTICAL BOTANY

through the elongation of its tube cell to form a pollen tube which
grows into the nucellus (megasporangium). ‘Then, the genera-
tive cell divides into a stalk cell and a body cell. The body cell
then divides to form 2 top-shaped antherozoids, each of which
bears many cilia along a spiral grove. The mature pollen tube and
its contents represents the male gametophyte of Zamia.

.

bas sbi eee a Dae ee
4 D
Fic. 347.—A, mature carpellate strobilus (cone) of Zamia. B, megasporophyll
bearing two megasporangia. C, lengthwise section of a young megasporophyll,
showing an early stage in the development of female sexual plants or megagameto-
phytes. D, similar section of a later stage; megagametophytes fully matured.
(From “A Textbook of General Botany” by Smith, Overton @ Co. by permission from
Macmillan Co., publishers.)

The female gametophyte develops within the megaspore (embryo
sac) of the nucellus of the ovule by a series of nuclear divisions
of the remaining nucleus within the embryo sac, resulting in
many scattered nuclei within the cytoplasm. Cell walls are laid
down and many cells result, and the embryo sac increases in
size forming a prothallus. The nucellus is encroached upon by
the enlarged developing gametophyte or prothallus in whose cells
nourishment is stored. An archegonial chamber appears between
the end region of the prothallus and nucellus, in line with the

THE SPERMATOPHYTES 461

pollen chamber and micropyle. Two to six archegonia are
developed from the prothallial tissue just beneath the archegonial
chamber. Each archegonium consists of two neck cells protruded
into the archegonial chamber and a large ovum embedded in
the prothallus. The contents of. the mature embryo sac, therefore,
represents the female gametophyte.

FERTILIZATION.—It will be recalled that at the time pollina-
tion was effected, the male gametophyte’ had assumed the char-

Pollen Chamber _Integument Nucellus. -

Zz ee Integument {

Microgametophyte Macrogametophyte ;

Fic. 348.—Zamia. Stages in the development of a megasporangium and a
macrogametophyte (megagametophyte). A, at the time when the macrospores
(megaspores) are formed. B, after the germination of one macrospore to form a
macrogametophyte. C, at the time of gametic union. (From ‘A Textbook of Gen-
eral Botany” by Smith, Overton et al., the Macmillan Co., Publishers.)

acter of a pollen tube (see Fig. 348) containing a prothallial
cell, stalk cell, and two antherozoids which lay in the pollen
chamber. Several of these are usually present. Now, when the
eggs are mature, the tube grows through the nucellus to the arche-
gonial chamber and the end of the tube bursts. The antherozoids
of the several tubes move about in the fluid of the chamber and
some of these pass through the necks of the archegonia, one
uniting with each egg. The sperm nucleus then unites with the
egg nucleus forming a zygote.

SEED DEVELOPMENT.—The zygote, through repeated division,
forms a young sporophyte or embryo Kamia in the upper end of the
female gametophyte (prothallus) of the ovule. This, by the
elongation of several cells, becomes differentiated into a slender

1 Microgametophyte is another name for male gametophyte, and macrogametophyte
a synonym for female gametophyte. :

462 PHARMACEUTICAL BOTANY

suspensor attached to the anlage of the embryo which has been
pushed through the basal membrane of the egg deep into the pro-
thallus by the elongating cells of the suspensor. The embryo
lives as a parasite on the tissue of the female gametophyte. After
differentiation has proceeded in the body of the embryo to a
certain point, z.e. where two cotyledons or seed leaves and a central
axis consisting of a hypocotyl (part below the attachment of the
cotyledons) and an epicotyl (part above the cotyledons) have been
formed, growth of the embryo becomes suspended.

Nourishment is stored in the cells of the prothallus around the
embryo forming the endosperm. The integument undergoes
changes in its cells to form a firm seed coat consisting of a strong
and a fleshy layer surrounding the nucellus. The micropyle
persists as a tubular opening in the seed coat.

GERMINATION OF THE SEED.—The mature seeds drop from the
megasporophyll of the mature cone to the ground and some of
them germinate. In this process the seed absorbs water. ‘The
enzymes within its cells change the reserve or stored food within
the endosperm into water soluble substances which are converted
into new substances or protoplasm, as a result of which the

Out ine or Lire Hisrory or ZAMIA

Zygote | Zygote

Embryo Embryo

Mature male sporophyte Mature female sporophyte
Microsporophyll Megasporophyli
Microsporangium Megasporangium
Microspore mother cell Megaspore mother cell
Microspore Megaspore

Male gametophyte Female gametophyte

Antherozoids

-_—~ Fertilization

|
a

Zygotes (destined to develop into male and
female sporophytes)

THE SPERMATOPHYTES 463

embryo grows. ‘The hypocotyl, elongating, forces its way out
through the micropyle and arches downward into the soil, most
of it becoming the primary or main root. The upper part
becomes the stem. The cotyledons remain within the seed.
The epicotyl pushes its way upward and develops the rest of the
stem, also the green leaves of the seedling sporophyte. As

Rew ik *

Fic. 349.—Ginkgo biloba, the Maiden-hair tree.

soon as the green leaves and the root with its absorbing root hairs
are developed, the seedling becomes an independent organism
developing either into a mature male or female sporophyte.

ORDER GINKGOALES

The Ginkgoales is represented by the single species, Ginkgo
biloba, the maidenhair tree, a native of western China and grown
extensively in China and Japan and the United States as an
ornamental tree. It possesses a straight trunk from which

464 PHARMACEUTICAL BOTANY

spring long shoots bearing scattered leaves and short shoots
with crowded leaves. Its leaves are fan-shaped, more or less
2-lobed, with furcately-branching veins and deciduous. It
bears two kinds of cones (strobéli) each kind on a different tree.
The staminate strobili occur in clusters on the short shoots, each
strobilus consisting of a central axis bearing a loosely arranged
series of stamens (microsporophylls). Each stamen consists of a
stalk ending in a knob and bearing on its underside two or more

Fic, 350.—Inflorescences of the pine. 1, Terminal twig; 2, ovulate cone; 3,
staminate cone; 4, two-year-old cone. (Hamaker.)

microsporangia. "The ovulate (female) strobili arise from the axils
of leaves on the dwarf shoots of female trees. Each consists of an
elongated stalk bearing near its end two ovules each of which has a
small, cup-like megasporophyll at its base. The mature, naked
seed resembles a drupe, the seed coat having an outer fleshy and
inner stony layer.

OrDER CONIFERALES

Trees with a single upright stem which develops side branches
that spread out horizontally and taper to a point at the summit,
giving the crown of the tree the appearance of a huge cone, rarely

THE SPERMATOPHYTES 465

shrubs. This order includes two families, the Pinacee or Pine
Family and the Taxacee or Yew Family.

Pinaceae (Conifere) or Pine Family.—Trees or shrubs with resin-
ous juice whose wood is characterized by being composed
largely of tracheids with bordered pits. Leaves entire, awl- or
needle-shaped, frequently fascicled, exstipulate, usually ever-
green. Flowers, monoecious (Pines, etc.) or rarely dicecious
(Juniper), achlamydeous, in cones. Staminate cone of a large
number of microsporophylls (stamens) closely packed together
and arranged spirally around a central axis, each stamen bearing
usually two pollen sacs. Carpellate cone composed of spirally
arranged scales, each of which bears a pair of naked ovules
(megasori) near the base of its upper face. Fruit a cone with
woody or fleshy scales (Pinus, Thuja, Abies, Picea, etc.) or a
galbulus (Juniperus). Seeds albuminous. Embryo with two or.
more cotyledons.

The following genera of the pine family are of especial
pharmaceutic and general economic interest:

GENUS Common NAME

Pinus Pine

Picea Spruce

Abies Fir

Larix Larch

Tsuga Hemlock

Pseudotsuga Douglass Fir

Thuja Arbor vitae

Juniperus Juniper

Sequoia Redwood

Cedrus Cedar

OrrictAL Druc Part Usep BorAnIcAL ORIGIN HABITAT
Terebinthina Concrete oleoresin Pinus palustris and Southern United -
(Turpentine) other species of States
Pinus :
Resina (Rosin) Resin Pinus palustris and Southern United
- other species of Pinus States
Oleum Tere- Volatile oil Pinus palustris and Southern United
binthinge other species of Pinus States
Pix Pini Product of destruc- Pinus palustris and Southern United
(Pine Tar) tive distillation of other species of Pinus States
wood

Oleum Picis Volatile oil from pine tar
Rectificatum :
Pinus Alba Inner bark Pinus Strobus United States

and Canada

s

466

OFFIcIAL Druc
Oleum Pini Pumi-
lionis
Juniperus

Oleum Juniperi

Pix Juniperi

UnorriciAL Drua

Sabina

Pix Burgundica

Terebinthina
Canadensis

Terebinthina
Laricis
Thuja

Oleum Abietis
(Oil of Siberian
Fir)

’ Tamarack Bark

Sandaraca

Dammar
Kauri Gum
Succinum
(Amber)
Bordeaux
Turpentine
Pix Canadensis
Oregon Balsam

Spruce Gum
Oil of Cedarwood

Hemlock

PHARMACEUTICAL BOTANY

Part Usep
Volatile oil

Fruit

Volatile oil
Empyreumatic oil
Tops

Resinous exudate
Liquid oleoresin
Oleoresin

Leafy young twigs

Vol. oil from fresh
leaves

Stem bark
Resinous exudate
Resinous exudate
Resin

Fossil resin

Concrete oleoresin

Oleoresin
Oleoresin

Gum
Oil from wood

Bark

HABITAT
Tyrolese Alps

BOTANICAL ORIGIN
Pinus montana

Juniperus
communis North America,
Juniperus Europe and Asia
communis
Juniperus Oxycedrus So. Europe
Juniperus Sabina Europe
Picea excelsa Europe
Abies balsamea Northern United
States and
Canada
Larix europa Alps and
Carpathians

United States
and Canada
Russia and
Siberia

Thuja occidentalis

Abies sibirica

Larix laricina Cold swamps of

North America

Callitris Africa
quadrivalvis —

Agathis loranthifolia EE. India
Agathis australis New Zealand

Pinites succinifer Basin of Baltic

Pinus maritima France

North America
Western United
States and British
Columbia

Tsuga canadensis
Pseudotsuga
taxifolia

Picea canadensis

z i Canada and New
Picea mariana

Picea rubra zaaone
Juniperus North America
Virginiana

Tsuga canadensis North America

TAxXACE& oR YEW Famity.—Trees or shrubs with dark green,
evergreen, exstipulate, linear leaves without resin ducts and with

hard, close-grained wood of a reddish-color.

The flowers are

usually dioecious; the staminate globular with 4 to 8 stamens each
having their anthers under a shield-shaped connective: the

THE SPERMATOPHYTES 467 .

pistillate consisting of a single terminal ovule with several
bracts at its base. The fruit consists of an erect bony seed
situated in a fleshy, reddish, cup-shaped disk, representing an aril.

c D

Fic. 351.—Ephedra a member of the order Gnetales. A, branches with ovulate

strobili. B, branches with staminate strobili. C, ovulate strobilus. D, an

ovulate flower. £, staminate strobilus with staminate flowers in the axils of bracts.
(From Watson.)

The leaves, seeds and bark of Taxus canadensis (American
Yew) and T. baccata (European Yew) contain the alkaloid,
taxin, which is poisonous to horses, cattle, sheep and pigs. The
fleshy disk (aril) surrounding the seed is not poisonous.

468 “PHARMACEUTICAL BOTANY

Fic. 352.—Ephedra sinica. A, entire plant. B, portion of a green aerial stem
from A, enlarged; n, node bearing two opposite scale-leaves (/); i, internode. on
fruiting branch showing terminal cone (f) and leaves (i). This species yields the
commercial variety of Ephedra known in commerce as Tsaopen Mahuang and is
gathered in Northern China.

THE SPERMATOPHYTES - 469

OrpDER GNETALES

The Gnetales comprise the single family, Gnetacea, which has
three genera which in certain respects show a closer resemblance
to the angiosperms than do other gymnosperms.

Fic. 353.—Cross section of a lower internode of the green stem of Ephedra sinica,

a source of the drug Mahuang. 17, ridge; note many of the ridges are papillated;
c, cuticle of epidermis; co, cortex; mf, mesocortical fibers; Af, hypodermal fibers; x,
xylem of an older (primary) bundle which has undergone secondary thickening;
Pp; pericyclic fibers just beneath which is phloem, ca, cambium; sx, secondary xylem;
px, end of protoxylem; ph, phloem of a secondary bundle; /d, tracheids with bordered
pits; ér, trachea; m, pith. Note the narrow medullary rays separating the open
collateral bundles.

Family Gnetacea. The Gnetum family includes the genera
Ephedra, Welwitschia and Gnetum, each genus representing a differ-
ent type of plant. The Ephedras include about 30 species of
straggling shrubs or low bushy plants of warm dry regions of the
northern hemisphere, a number of species of which are used
medicinally because of their ephedrine content. Welwitschia

470 PHARMACEUTICAL BOTANY

contains the single species Welwitschia mirabilis, a low growing
plant of arid regions in southwest Africa. The genus Gnetum

contains about 15 species of woody climbers native to the tropics
of both hemispheres. The principal features of the Gnetum
family are, as follows:

Stems either unbranched and tuberous (Welwitschia) or
branched and shrubby (Epfedra) or often climbing (Gnetum) and

st

td

Fic. 354.—Transverse section through the outer region of a representative
portion of the stem of Ephedra sinica. 400. c, cuticle; ep, epidermis; h, hypo-
dermal fibers; ch, chlorenchyma of cortex, the outer layers of which are palisade-like;
J, mesocortical fibers; td, tracheids and tr, trachea of secondary xylem; st, stoma.

possessing open-collateral fibrovascular bundles. Resin canals
are absent while true vessels (tracheae) are present in the second-
ary xylem (distinction from conifers). Tracheids also occur in
the xylem. Internodes and nodes are conspicuous in Ephedra
and Gnetum. Leaves usually small, often reduced to scale-like
structures and arranged in opposite or verticillate fashion.
Flowers nearly always dioecious and exhibiting a membranous
perianth of 2- to 4-bracts. Perianth of male flowers surrounding
2 to 8 stamens. Female flowers showing an erect ovule with 1 to

THE SPERMATOPHYTES 471

2 integuments and a bract-like perianth. Fruit in Ephedra a
reddish, somewhat globular, galbulus-like structure (syncarp).
Seed albuminous, the embryo with two cotyledons imbedded in
endosperm.

The drug EpHepra consists of the green stems of Ephedra
sinica, Ephedra equisetina, E. distachya, E. Gerardiana or other
species of Ephedra containing the alkaloid ephedrine. These
species are largely Asiatic in distribution. The United States
Pharmacopoeia recognizes the alkaloid ephedrine and two of its
salts, ephedrine sulfate and ephedrine hydrochloride. Prepara-
tions of these are official in the National Formulary. Ephedrine
and its salts are used locally to shrink mucous membranes in
colds and asamydriatic. They are given internally for LeSieeenses
asthma, and certain circulatory conditions.

THe ANGIOSPERMS

The angiosperms comprise a large assemblage of seed plants
which are more highly developed than those of the preceding
group. Like the gymnosperms they possess flowers with sporo-
phylls (stamens and carpels), but the floral axis has been
shortened so that the sporophylls and the accessory floral leaves
(sepals and petals), when present, are brought closer together on
the axis, instead of being separated at strikingly different levels
along the axis. The ovules, instead of being borne naked on a
scale, are enclosed within the ovary and the seeds are covered
by the ripened wall of the ovary which is called the pericarp or
seed vessel.

Lire History oF AN ANGIOSPERM (ERYTHRONIUM)

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 /eaves of pale green color mottled with purple and
white, and later, a flower stalk (scape), which bears upon its sum-

472 PHARMACEUTICAL BOTANY

mit 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, andrecium and gynecium. The calyx is composed
of three lance-shaped and recurved, yellow parts called sepals;
the corolla of three similarly looking parts called petals which

Fic. 355.—Dog’s tooth violet (Erythronium americanum). Stages of development
from the seed. 1-5 show the stage of development in each of five successive years.
Full explanation in the text. 6, Bulb showing a surface bud (the sprout has been
destroyed). (Gager after F. H. Blodgett.)

alternate in position with the sepals. Both of these whorls are
collectively called the fperianth 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) consists of
an awl-shaped stalk or filament bearing upon its summit an
oblong-linear body called an anther. The anther consists of two

THE SPERMATOPHYTES 473

lobes called microsori. Each lobe or microsorus contains two
anther sacs or microsporangia in which, when mature, are to be
found microspores or pollen grains. . In the center of the flower will
be noted the gynoecium or female system of organs. This, upon
dissection, will be found to consist of three fused carpellary leaves
termed megasporophylls (carpels) forming a somewhat flask-
shaped structure called a pistil. "The swollen basal portion of the

Fic. 356.—Cross section of the ovary of a lily formed by the coalescence of the
lower portions of three carpels. X70. 5, dorsal sutures; ep, epidermis; c, cuticle of
epidermis; / bundle in mesophyll; /, locule; ov, ovule (anatropous); p/, infolded edges
of adjacent carpellary leaves forming the axile placentas from which the ovules arise.

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.

Microscopical examination of sections of the ovary will reveal
it to be composed of three chambers called Jocules, within each of
which are to be noted several inverted ovules. Each of these
ovules is developed upon a nourishing tissue termed “placenta”
which connects the ovules to the inner angle of the wall of the
locule. The ovule is composed of a central prominent mega-

474 PHARMACEUTICAL BOTANY

sporangium 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 micropyle (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 sections examined have been properly
fixed, will be found a megaspore or embryo sac.

The megasporangium or nucellus is formed from a mass of
meristem on the placenta. While in the state of differentiation,
one of its cells is set apart as a megaspore mother cell. Its nucleus
(containing 2x chromosomes) undergoes two successive divisions
during the first of. which a reduction in the number of chromo-
- somes from the diploid (2x) to the haploid (x) number occurs. The
x number persists in the ovum or egg.

DEVELOPMENT OF THE FEMALE GAMETOPHYTE THROUGH THE
MATURATION OF THE EmBryo Sac.—In its immature condition
the embryo sac (megaspore) contains a mass of protoplasm surround-
ing 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 opposite end of the sac three
nuclei known as antipodals take their position; 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 gametophyte. [See Fig. 358 (1-8).]

MATURATION OF THE POLLEN GRAIN AND FORMATION OF THE
Mae 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.
The pollen mother cells like the rest of the sporophyte generation
contain the diploid or 2x number of chromosomes. During
their division to form tetrads a reduction division occurs which
reduces the chromosomes to the haploid or x number. This

THE SPERMATOPHYTES 475

number continues into the sperm nuclei. Each pollen grain,
after the tetrads have separated into their components, consists of
an outer firm wall or exosporium, an inner wall or endosporium,
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 develop-
ment of the male gametophyte. The nucleus and protoplasm
enveloping it divides to form two cells, one a generative-cell

A

Stomiumy |
Mechanical Cells,
Endothectum ~

Broken Down Partition ~
between Miucrosporangia

Generative Cell-~ es

Fh
Tube Cell-~

Fic. 357.—A, cross section of dehisced anther of a lily (Butomus). The layer of
cells beneath the epidermis (exothecium) acts as mechanical tissue to open the
pollen sacs. (After Sachs.) B, Section of a pollen grain, diagrammatic. (Mottier.)

containing a generative 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. (See Fig. 220.)

POLLINATION AND FERTILIZATION

The mature pollen grains are discharged from the ripened
anther through the splitting open of its wall. ‘They are trans-
ferred to the stigma of the pistil of another Erythronium flower
through the agency of insects. Here they germinate, each put-

476 PHARMACEUTICAL BOTANY

ting forth a tube (pollen tube). The pollen tubes, each 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

YL

Z LZ
LL

S$
s
Y

aN

Fic. 358.—At the left, diagram of the anatomy of an angiospermous flower
shortly after pollination; anth., anther; fil., filament; s¢., stamen; stig., stigma;
p.g., pollen grains germinating; sty., style; pt., pollen tube; 0.w., ovary wall; o.
ovule, containing embryo-sac; fet., petal; sep., sepal. 1-8, Stages in the develop-
ment of the female gametophyte (embryo-sac); meg.sp., megaspore-mother-cell;
i.2., inner integument; 0.7., outer integument; fun, funiculus; chal., chalaza; nu.,
nucellus (megasporangium); emb., embryo-sac. All diagrammatic, (Gager.)

of these sperm nuclei (containing x chromosomes) passes between
the synergids and fuses with the nucleus of the egg (containing x
chromosomes) to form an oéspore or zygote in which the 2x number
of chromosomes found in the sporophyte of the species is restored.
By this time the tube nucleus has disintegrated. The odspore 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

foe aes

a eae

THE SPERMATOPHYTES — 477

undergoes rapid division into a large number of nuclei scattered
about through the protoplasm of the embryo sac. Later cell
walls are laid down and endosperm is formed. ‘The endosperm
cells soon become filled with abundant starch which is later to be
utilized by the embryo during germination. |

RIPENING OF THE OVULE TO ForRM THE SEED AND OF THE OVARY
To Form THE FRuitT

When 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 cap-
sule or fruit splits open to discharge them. ‘They fall to the
ground and lie dormant until the following spring when they
germinate or commence 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 protoplasm changes it into additional
protoplasm and so the embryo increases in size, therefore, grows.
The pressure of the swollen endosperm and growing embryo
becomes so great that the seed coat bursts; the hypocotyl emerges
first, dragging the cylindrical cotyledon out of the seed coat and
epicotyl with it. The hypocotyl elongates and extends itself
- into the soil where it develops a root near its tip. ‘The tip
enlarges through the storage of starch, manufactured 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.

478 PHARMACEUTICAL BOTANY

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 fifth year
after formation and shedding of seed) one of these bulbs develops
_ into a mature sporophyte plant, bearing a single flower at the
summit of its elongated scape. (See Fig. 355.)

RESEMBLANCES BETWEEN GYMNOSPERMS AND ANGIOSPERMS

1. In both are developed flowers.

2. In both the flowers develop at least two sets of leaves
(either on one or two plants of the same species) called sporophyll
leaves, the stamens and carpels. ‘The stamens or staminal leaves
are also termed microsporophylls. The carpels or carpellate
leaves are also known as megasporophylls.

3. Both groups produce microspores or pollen grains and
megaspores or embryo sacs.

4. In both are developed on the evident generation, the
plant or sporophyte and the gametophyte, the latter concealed
within and parasitic upon 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 pericycle.

8. Both produce collateral vascular bundles. Occasion-
ally we meet with phlocentric bundles in the stem or leaf of
Angiosperms.

“FUNDAMENTAL DIFFERENCES BETWEEN GYMNOSPERMS AND
ANGIOSPERMS

1. The flowers of Gymnosperms are often moncecious or
dicecious but very rarely hermaphrodite (as in Welwitchia),
whereas those of Angiosperms are usually hermaphrodite, rather
rarely moncecious, 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

THE SPERMATOPHYTES 479

whorls or spirals set around a shortened axis, the floral axis or
receptacle, 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 Angiosperms, 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 gameto-
phyte plant that later becomes the so-called “‘endosperm’’ of
the gymnosperm, whereas in Angiosperms no recognizable
prothallus has been proven to exist.

5. The stored food tissue in Gymnosperm seeds is prothallial
tissue loaded with starch, etc., whereas in Angiosperm seeds the
stored food tissue fendldeperat} is a — formation after
fertilization.

6. Gymnosperms bear naked ovules and seeds while Angio-
sperms bear covered ones.

7. In Gymnosperms there are distinct recognizable arche-
gonia 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.

CLASSES OF ANGIOSPERMS

The Angiosperms are subdivided into two classes, the Mono-
cotyledonez or Monocotyledons, and the Dicotyledoneze or
Dicotyledons.

The Monocotyledons include such plants as the cat-tails,
grasses, sedges, palms, lilies, daffodils, irises, bananas, gingers
and orchids. The Dicotyledons are represented by a vast

480 —S*SW PHARMACEUTICAL BOTANY

array of plants which include nearly all the familiar trees and

shrubs except the conifers and the majority of weeds, garden

plants, and ornamental herbs which have netted veined leaves.

The monocotyledons appear from studies in comparative
morphology and serum-diagnosis tests to have originated from
the Archichlamydez of the Dicotyledons.

3
:
‘
3

CHAPTER XXII

THE MONOCOTYLEDONS

This is a class of Angiosperms characterized by the following
peculiarities:

Fic. 359.—Morphology of the typical monocotyledonous plant. 4A, leaf,
parallel-veined; B, portion of stem, showing irregular distribution of vascular
bundles; C, ground plan of flower (the parts in 3’s); D, top view of flower; E, seed
showing monocotyledonous embryo. (Gager.)

(1) One cotyledon or seed leaf in the embryo, terminal in
position.
(2) Stems usually endogenous with closed collateral or

concentric fibro-vascular bundles, which are scattered. ,
481

482 PHARMACEUTICAL BOTANY

(3) Leaves alternate in arrangement, with an entire margin
and generally parallel-veined.

Fic. 360.—Wheat plant showing the general habit of grasses. (Robbins.) _

(4) Flowers usually pentacyclic and trimerous (having the 2
parts of each whorl in threes or a multiple thereof). :
(5) Secondary growth in roots generally absent, the olyarch a
radial bundle persisting from its formation.

THE MONOCOTYLEDONS 483

(6) Medullary-rays generally absent.
(7) Cambium generally absent.

Fic. 361.—Pistillate and staminate inflorescences of corn (ea mays). Pistillate
on left, staminate on right. (Roddins.)

Druc YIELDING Groups OF MONOCOTYLEDONS

The Monocotyledons comprise 45 families which contain
about 25,000 species of described plants. Only those groups
which yield drugs are considered in this chapter.

OrDER GRAMINALES

This is the most important economic order of Monocoty-
ledons. It contains two families, the Graminee or family of
grasses and cereals and the Cyperacee or family of sedges. The

484 PHARMACEUTICAL BOTANY

flowers of both of these families are devoid of sepals and petals
but possess scale-like structures called glumes and palets which
enclose one or more flowers. The grasses and cereals have
stems with solid nodes and hollow internodes whereas the sedges
have stems which are solid throughout. The above ground
stem of a sedge is usually 3-sided, unlike any grass stem. The
fruit of a grass is a caryopsis whereas that of a sedge is an akene.

GRAMINE# OR Grass Famity.—Mostly herbaceous (Triticum,
Sorghum, Saccharum, cereals, etc.), rarely
5) , shrubby (Arundinaria, etc.), more rarely
arborescent (Bamboo, etc.) plants, vary-
ing in height from 2 in. to 100 ft.

Stems, leaves and even parts of flowers
“often abounding in silica. The leaves are
alternate, with long split sheaths usually
open on the side opposite the lamina, and
a ligule. Flowers small, generally her-
maphroditic, occasionaly moncecious
(Corn and Rice Grass) and usually borne

Fic. 362.—Diagram of a IN sptkelets arranged along a spike-axis
typical spikelet. n¥ lower called a rachis, but sometimes in panicles

glume; @Y upper glume; Oat) 8h : . :
nl outer palet; ®J, inner ( ) eS ikelet is the unit of a gr ass

palet; #, lodicule; st, sta- Mflorescence. A typical spikelet consists
mens; J-/, rachilla; 2, lateral Of a shortened zigzag axis (rachilla) bear-

axes. (Somewhat modified ing floral leaves. The two lowest floral
after Warming.) l : h ikel

eaves in each spikelet are barren and
termed glumes. The succeeding floral leaves are of bract-like
character, each supporting one flower in its axil, and are called
outer palets or lemmas. Each flower has a bracteole which is
placed on the inside, opposite and with its back to the rachilla
and known as the inner palet. Immediately following the inner
palet are 2 delicate scales called Jodicules, 3 stamens (rarely
1, 2 or 6) with versatile anthers that are usually notched at each
end and a single pisti/, having a unilocular, superior ovary with
one ovule and usually 2 styles, with simple or spirally branched
(plumose) stigmas. Fruit, a caryopsis or grain. Seeds albu-
minous with starchy endosperm and a small embryo at the base
and opposite the hilum side. Seed coat fused with fruit coat to

Paes eee

THE MONOCOTYLEDONS 485

form one layer. Embryo with a scutellum. The scutellum
represents a portion of the single cotyledon of these plants,
which bales out the nourishment from the endosperm to the
rest of the embryo, during the germination of the grain.

OrriciAL DRuG Part Usep BOTANICAL ORIGIN HABITAT
Triticum Rhizome and roots Agropyron repens Europe and Asia
Sucrosum Refined sugar Saccharum offici- Tropics

narum and other
sources
Glucosum Product obtained by incomplete hydrolysis of starch
Dextrosum Sugar obtained by hydrolysis of starch

Amylum Starch Zea Mays Mexico
Zea Styles and stigmas Zea Mays Mexico
Avena Grain Avena sativa Eurasia
Oleum Maydis Fixed oil Zea Mays Mexico

UnorriciAL DrucG

Cymbopogon Nardus
Oleum Citronella Volatile oil Cymbopogon Asia
Winterianus
Job’s Tears Fruits Coix Lacryma-Jobi _ E, Indies
Fructus Fruits Phalaris Canary Islands
Canariense canariensis
Diastasum Enzyme Hordeum spp. Asia
Vetiver Rhizome and Andropogon E. Indies
roots © squarrosus
Maltum Grain partially Var. of Hordeum Asia
germinated and vulgare
dried

OrDER PRINCIPES

PALM& or PatmM Famity.—Tropical or subtropical shrubs,
rarely trees, having unbranched trunks which are terminated
by a crown of leaves, in the axils of which the flowers are pro-
duced. The leaves are well developed with pinnate or palmate
blades and fibrous, sheathed, clasping petiole. The flowers are
small, of one or two sexes, and crowded on a spike or spadix,
which is subtended by a large bract, or spathe which may
become woody, as in the Cocoanut Palm. The perianth con-
sists of 6 parts in 2 whorls (3 sepals and 3 petals) or it may be
inconspicuous or absent. The stamens are 6 in number, rarely
3, inserted below the ovary. The ovary is superior, of 3 cells,
with central placenta. The fruit is either a nut, with leathery
epicarp, fibrous or cellular mesocarp and thin membranous

486 PHARMACEUTICAL BOTANY

endocarp, or a drupe (Cocoanut) with leathery epicarp, broadly
fibrous mesocarp and stony endocarp, or a berry as in the Date

ba

Fic. 363.—Cocoanut Palm (Cocos nucifera). Note pinnate leaf-blades. (From
Bull. Agricole du Congo Belge. 1910.)

Palm, Phoenix, with membranous epicarp, succulent mesocarp
and soft succulent endocarp. The seeds are albuminous with

THE MONOCOTYLEDONS 487

the reserve food frequently in the form of hard (reserve) cellulose
(ivory-nut-palm) or ruminate albumen (Areca).

OrrFicIAL Druc Part Usep BotANnicAL NAME HABITAT
Sabal Fruit (drupe) Serenoa serrulata South Carolina to
(Saw Palmetto) ; Florida and Texas
Areca Seed

Arecolinz Hydrobromide of Asia and Malayan
Hydrobromi- _aikaloid from ee Islands.

dum seed

UnorrFictiAL Druc

Dragon’s Blood Inspissated juice Damonorops Draco East Indies
Cocoanut oil Fixed oil Cocos nucifera Tropics

Carnauba wax Wax from leaves Copernicia cerifera Brazil

Palm oil Fixed oil Elzis guineensis West Africa

OrDER ARALES

ARACE& OR ARUM Famiry.—Perennial herbs with fleshy
rhizomes or corms, long petioled leaves, and acrid or pungent
juice. Leaves mostly radical; when cauline, alternate, entire to
divided, often hastate or sagittate, with a membranous sheath
at the base. Flowers small, usually imperfect, rarely hermaph-
roditic, usually monoecious, the pistillate usually below, the
staminate above, crowded on a spadix, which springs from the
axil of a spathe. Fruit an indehiscent, 1 to several celled, 1 to
many seeded berry occurring in a cluster on the fleshy fruiting
spadix. Seeds filled with large, curved, fleshy embryo or mostly
with copious endosperm with embryo in center.

This family includes many ornamental plants among which
are the calla lily (Richardia ethiopica), the ceriman (Monstera ©
deliciosa) whose large leaves have many holes, the Anthuriums with
colored showy bracts and the Elephant’s ear (Colocasia esculenta).

OrrictAL DruG Part Usep BoTtanicAL NAME HAsiTrAtT
Calamus Peeled rhizome Acorus Calamus Europe, Asia, North
America
UNOFFICIAL
Druc

Skunk cabbage Rhizome Symplocarpus North America
foetidus

Indian turnip Corm : Ariseema North America
triphyllum ;

Tonga _ Rhizome Epipremnum pin- Fiji Islands

natum (in part)

*

;
:
‘
:

488 PHARMACEUTICAL BOTANY

Fic. 364.—Acorus Calamus, the Sweet flag, a member of the Aracee. (Sayre.)

Fic. 365.—Diagram of A, lily flower, and B, grass flower, showing homologous
structures. A, f, bract; ax, axis; op, outer perianth; i, inner perianth; s, stamens;
(c) tricarpellary ovary. B, shaded structures are aborted; /], lemma (bract); ax,
axis; and p’, palet (outer perianth); / and /’, lodicules (inner perianth); s and s,
two whorls of stamens; ¢, tricarpellary ovary. (A, Robbins. B, after Shuster.)

THE MONOCOTYLEDONS 489

ORDER LILIALES

LILiAcE# oR Lity Famity.—Herbs (Lilium), shrubs (Yucca),
or trees (Dracena Draco or Dragon’s Blood), with regular and

Fic. 366.—Smilax officinalis—Portion of vine and root system. Note the 3-nerved
leaves with stipules modified as tendrils and the berry fruits. (Sayre.)

symmetrical almost always six-androus flowers. Stem either
short, creeping under ground (Convallaria), or, swelling up and
forming bulbs (Hyacinth), or corms (Colchicum), or, stem may

2
a
hee
a
14
iz
“a

eh wee a a

490 PHARMACEUTICAL BOTANY

elongate above ground and become wiry and herbaceous or
semi-shrubby as Smilax, or the stem may remain short, giving
rise to thick, fleshy and sap-storing leaves, as in Aloe. Leaves
linear to lanceolate, ovate rarely wider, divisible into sheating
base, narrow petiole and expanded blade. Venation, parallel,
becoming in some ovate leaves parallel with oblique connec-
tions, reticulate or highly reticulate, as in the Sarsaparilla plants
and other species of Smilax. The perianth is parted into six
segments, the calyx and corolla being alike in color. Anthers
introrse. Ovary three-locular with a single style. Fruit a
three-locular, loculicidally dehiscent capsule (Lilium, Colchicum,
etc.) or rarely a berry (Asparagus, Lily-of-the-valley, etc.).

Seeds usually numerous with fleshy endosperm.

OrrictAL Dru Part Usrp BoTranicAL NAME HasitaT
Smilax medica Mexico
Smilax ornata Costa Rica
Sarsaparilla Root Smilax officinalis or Guatemala,
an undetermined Honduras and
species Nicaragua
Veratrum Viride Rhizome and roots Veratrum viride United States
Colchici Cormus Corm Colchicum Mediterranean
autumnale Basin
Colchici Semen _—_ Seed Colchicum Mediterranean
autumnale Basin
Aloe vera Dutch West
‘ : Indies
Aloe am juice of Aloe Pecryi ORES
Aloin Pentoside East Africa
Scilla Inner scales of bulb Urginea maritima Mediterranean
(Scilla maritima) Basin
Convallariae Rhizome and roots Convallaria majalis Europe, Asia,
Radix United States
Trillium Rhizome and roots Trillium erectum North America
Aletris Rhizome and roots Aletris farinosa Eastern United
States
Helonias Rhizome and roots Chamelirium luteum Eastern United
States
UnorriciAL DruG
Cape Aloe Inspissated juice Aloe ferox S. Africa
jez and
Sabadilla Seed Asagrza officinalis Central
America
Red Squill Bulb Urginea maritima Mediterranean
(red variety) Basin

THE MONOCOTYLEDONS 491

HABITAT

UnorrFiciAL DruGc PArT UsepD BoTANICAL ORIGIN
Allium Bulb (fresh) Allium sativum Southern Europe
Convallarize Inflorescence Convallaria majalis Europe, Asia,
Flores United States
Veratrina Mixture of Asagrzea officinalis Mexico and
alkaloids Central America

DIoscOREACE OR YAM FamILy.—T wining perennial herbs or
undershrubs arising from large tuberous roots or knotted root-
stocks. Leaves alternate, peti-
oled, simple, palmately-ribbed
and netted-veined. Flowers
small, dioecious, regular, having a
six-cleft, calyx-like perianth, six
stamens and a three-celled inferior
ovary. Fruit usually a membra-
nous, three-angled or winged,
three-celled capsule.

OFFICIAL DRUG Part UsEep
Dioscorea Rhizome
BoTANICAL NAME HABITAT

Dioscorea villosa United States

IRIDACEZ OR IRIs FAMILY.—
Perennial herbs with underground
rhizomes (Iris) corms (Crocus) or

bow

Fic. 367.—Iris_ versicolor (Blue

bulbs (Gladiolus) sword shaped
or linear, equitant, two-ranked
leaves and usually showy, perfect,
regular or irregular flowers which
emerge from a spathe of 2 or more
bracts. Perianth of 3 sepals (often
petaloid), and 3 petals arising

Flag). Note the fleshy rhizome bear-
ing equitant, sword-shaped leaves and
long stalked flower clusters. The
flowers are violet-blue, the outer
segments of the perianth limb being
spatulate, variegated with yellow,
green or white on the claw and marked
with purple veins. (Reproduced from
U. S. Dept. Agric. Miscellaneous Publ.
77.)

from the summit of a tubular re-

ceptacle above the ovary; stamens 3; ovary inferior, 3-celled, style
single, stigmas 3, sometimes expanded and colored, as in Jris.
Fruit a three-celled, loculicidal, many-seeded capsule. Seeds
with fleshy or horny endosperm. Rootstocks, tubers, or corms
mostly acrid. Fibrovascular bundles in stems and _ leaves
phlocentric.

492 PHARMACEUTICAL BOTANY

OrriciAL DrucG Parr Usep BoraAnicAL NAME HasitaAt
wslinsb ines Mediterranean
Iris (Orris) Peeled rhizome Iris pallida :
; : Region
Iris germanica
Iris Versicolor Rhizome ee versicolor Eastern North
(Blue Flag) Iris caroliniana America
Crocus (Saffron) Stigma Crocus sativus Southern Europe
and Asia

ORDER SCITAMINALES

This order includes herbs with asymmetrical flowers and
pinnately-veined leaves. Some of the members, notably the
bananas, are gigantic in size. It contains the Musacee or
Banana Family, Cannacee or Canna Family, Zingiberacee or
Ginger Family and Marantacee or Arrowroot Family.

ZINGIBERACE# OR GINGER Fami_y.—Tropical plants, peren-
nial herbs, usually with fleshy rhizomes and large, elliptical,
pinnately-veined leaves. The leaf sheaths are folded tightly
around each other so as to give the appearance of a stem.
Flowers, very irregular, trimerous; sepals three, short, often
green; petals three, elongate, often fused below; stamens three
to four abortive, petaloid, one often absent, a sixth alone fertile
and stamen-like. The petaloid stamens constitute the most
attractive parts of the flower. Pistil, inferior, tricarpellary.
Fruit, a loculicidally dehiscent, usually 3-celled capsule. Seeds
albuminous with both perispermic and endospermic albumen.

OrrFiciAL Drue Part Usep BoraAnicAL NAME HABITAT
Zingiber Rhizome Zingiber officinale §. Asia
Cardamomi Semen Seeds Elettaria Indo-China

Cardamomum
Oleum Volatile oil from —_Elettaria Indo-China
Cardamomi seeds Cardamomum
UnorriciAL Druc
Galanga Rhizome Alpinia China
officinarum
Zedoaria Rhizome Curcuma Zedoaria India
Curcuma Prepared rhizome Curcuma longa S. Asia and E.
Indies
Grains of Paradise Seeds Amomum Asia
melegueta

MARANTACE2 OR ARROWROOT Famity.—Perennial herbs
chiefly of tropical climes with fleshy rhizomes or tubers rich in

- THE MONOCOTYLEDONS 493

starch content. The leaves are radical and cauline, usually
long petioled, with a sheathing base and a pulvinus at the sum-
mit of the petiole. The inflorescence is a panicle or raceme

Fic. 368.—<ingiber officinale, the Ginger plant. (Sayre.)

bearing irregular trimerous flowers with a one-celled ovary (by
abortion).

UnorFIcIAL DruG PART UsEpD BoTANICAL ORIGIN HAsirar
Amylum ' Starch Maranta West Indies
Marantz arundinacea Northern S.

America

494 PHARMACEUTICAL BOTANY

OrDER ORCHIDALES

ORCHIDACE2 OR OrcHIp Famity.—Perennial herbs of
terrestrial or terrestrial saprophytic or epiphytic growth, having
grotesque flowers. Roots fibrous or tuberous, often saprophytic

tf
J

Fic. 369.—Floral organs of an orchid (Catleya sp.). A, the entire flower; sep,
sepal; pet, petal; B, column, showing s, stigma and r, the rostellum (beak), with
the small glands at the tip; to the glands are attached the four strap-shaped caudicles
of the pollinia; C, pollinia, with the four caudicles; below, the gland; D, longitudinal
section of the column; /, pollinium; E, the same, enlarged. (Gager.)

in relation, or aerial and with velamen. Stems and branches
upright, in epiphytic types, often forming pseudobulbs. Leaves
alternate, entire, parallel-veined, sheathing at base, rarely
reduced to yellowish or pale scales in saprophytes. Flowers
irregular, usually attractive, entomophilous, but sometimes

THE: MONOCOTYLEDONS 495

small, greenish, and inconspicuous, arranged in elongated spikes
or racemes, trimerous. Sepals 3, usually similar; petals 3 of
which two often resemble sepals, third is variously, often greatly
modified and fused with two outer petaloid stamens as a labellum
or lighting-board for insects. Third stamen of outer whorl

Fic. 370.—Vanilla planifolia—Branch showing leaves and flowers. (Sayre.)

fertile (Orchidee@) or a barren knob (Cypripedia); pollen of fertile
anther agglutinate and forming a pollinium. Three stamens of
inner circle barren and petaloid or one absent (Cypripedia).
Stamens all epigynous and often three are fused with the style
as a gynandrium. Carpels three, syncarpous, with inferior, 3-,
rarely 4-, 1 (usually)-celled ovary. Fruit a capsule, 3-valved
and 1-celled. Seeds minute, abundant and wind disseminated.

J
a
fi

iS) aa PHARMACEUTICAL BOTANY

OrriciAL DruG ' Part Usrep BoranicAL NAME HABITAT
Vanilla Fruit (cured, full- Vanilla planifolia © Mexico
grown, unripe)

UnorFiciAL Druc

- (Orchis mascula
Salep Tubers kere ee Europe

-Corallorhiza Rhizome Corallorhiza Eastern U. S.

odontorhiza
Cypripedium
_ \ bulbosum
Cypripedium Rhizome and roots lat oo ee United States
Cypripedium
pubescens

CHAPTER XXIII

THE DICOTYLEDONS

The Dicotyledons are a class of angiosperms having the
following characteristics:

(1). Two-seed leaves (cotyledons) in embryo, these being
lateral on the embryonic axis.

Z¥
eas
SS
Sy
Ja
ih,
LL

ay,
Pe,

- eS: a oe ~.
PRY

L>

ne,

Fic. 371.—Morphology of a typical dicotyledonous plant. 4, leaf, pinnately-
netted veined; B, portion of stem, showing concentric layers of wood; C, ground-
plan of flower (the parts in 5’s); D, perspective of flower; E, longitudinal section
of seed, showing dicotyledonous embryo. (Gager.)

(2) Foliage leaves with pinnate-reticulate or palmate-
reticulate veins.

(3) Leaf blades varying from simple to lobed to compound
-in pinnate or palmate fashion depending upon character of

venation.
497

498 PHARMACEUTICAL BOTANY

(4) Stems, leaves, and roots of secondary growth with open
collateral or (excepting roots) bi-collateral fibro-vascular bundles;
in stems radially arranged about a central pith.

(5) Exogenous stems.

(6) Medullary-rays present in stems and in roots of secondary

growth.
(7) Cambium present in the plant axis.

Fic. 372.—At left, fruiting branch of Black Pepper (Piper nigrum). Portion
of shoot with inflorescences. Terminal portion of an inflorescence (enlarged).
(After Wossildo.)

At right, the Cubeb (Piper Cubeba). a, cluster of fruit; b, staminate flower;
¢, pistillate flower in longitudinal section; d, fruit in longitudinal section. (After
Berg and Schmidt.) Both from Gager after Strasburger, Noll, Schenck and Schimper.

(8) Roots developing secondary structure.

(9) Flowers usually pentacyclic and tetramerous or pentam-
erous (parts of each whorl, four or five or multiple thereof),
with usually a reduced number of parts in the gyncecium.

This is the largest class of angiosperms containing more than
100,000 species of described plants which have been grouped
into 240 families. We shall consider in this chapter only those
groups of dicotyledons which yield drugs.

THE DICOTYLEDONS 499

The Dicotyledons are subdivided into two sub-classes; viz.:
Sub-class 1. Archichlamydee.
Sub-class 2. Sympetale or Metachlamydez.

Druc YreELpiInGc DicoTyLEDONS

Sus-CLAss ARCHICHLAMYDE

Those dicotyledonous plants in which the petals are distinct
and separate from one another or are entirely wanting. That
group of the Archichlamydez whose flowers show the absence
of petals and frequently of sepals is called the Apetale. The
group whose plants have flowers showing the parts of their
corolla (petals) separate and distinct is called the Choripetale or
Polypetale.

ORDER PIPERALES

PIPERACEZ OR PEPPER Famity.—A family of apetalous aro-
matic herbs and shrubs with jointed stems, opposite, verticillate,
or sometimes alternate leaves without stipules, and spiked,
inconspicuous, wind-pollinated flowers. The characteristic fruit
is a drupe enclosing a single, upright, albuminous seed with
abundant perisperm (from megasporangial tissue) and reduced
endosperm (from matured embryo sac).

OrriciAL Druc Part UsEpD BorAnicAL NAME HAsitat

Cubeba Unripe fruit Piper Cubeba Borneo, Java
Sumatra

UnorrFiciAL DrucG

Piper (Pepper) . Unripe fruit Piper nigrum Cochin-China,
India

Piper Longum Spike of imma- fae longum as

_ ture. fruit Piper officinarum
Kava Rhizome and Piper methysticum ‘Sandwich Islands
: roots :
Matico Leaves Piper angusti- Peru, Bolivia
folium

OrDER SALICALES

SALICACE2 OR WILLOW FamiLy.—Shrubs or trees of tem-
perate or cold regions, with upright woody stems, rarely herbs
(Salix retusa). Bark often containing bitter principles (Salicin,
etc.).

500 PHARMACEUTICAL BOTANY

Leaves alternate, simple, entire, stipulate; stipules rarely
green, persistent, usually functioning as winter bud-scales and
falling in spring.

Inflorescences dicecious spikes, so on separate plants. Stami-
nate spikes forming deciduous catkins of yellowish flowers,
pistillate as persistent spikes of green flowers, at length maturing
fruit.

Flowers of catkins numerous, each of two to five (Willow) or
six to fifteen (Poplar); stamens in axil ofa small bract leaf,
sometimes with small nectar knob or girdle at base; pollen
abundant, hence plants anemophilous, rarely entomophilous.
Pistillate flowers green, each of a bicarpellate pistil in axil of
bract, ovary one-celled with parietal placentation, style simple,
stigma bilobed.

Fruit a capsule dehiscing longitudinally. Seeds small, exal-
buminous, surrounded by a tuft of hairs for dissemination.

OrrFiciAL DruG Part UsEep BoTANnicAL NAME HasitTAtT

Salicinum _ Glucoside Several species of Europe, North
Salix and Populus America
Populus

Populi Gemma Closed winter leaf rat North America

bud Populus candicans-

UnorriciAL Druc

Salix Bark Salix alba Europe

Salix Nigra Bark Salix nigra North America

Poplar bark Bark Populus tremuloides North America

ORDER MyRICALES

Myricack& OR BAYBERRY FamiLy.—Dicecious or sometimes
moncecious, aromatic shrubs or trees with watery juice and
possessing underground branches which arch downward then
upward producing many suckers. Roots fibrous and bearing
many short rootlets upon which are frequently found coralloid
clusters of tubercles containing the Actinomyces Myricarum
Youngken. Leaves alternate, revolute in vernation, serrate,
irregularly dentate, lobed or entire, rarely pinnatifid, pinnately
and reticulately veined, pellucid-punctate, evergreen or decidu-
ous, generally exstipulate, rarely stipulate. Flowers naked,
unisexual, monoecious or dicecious, in the axils of unisexual or

THE DICOTYLEDONS

501

androgynous aments from scaly buds formed in the summer in
the axils of the leaves of the year, remaining covered during the
winter and opening in March or April before or with the unfold-

ing of the leaves of the year.
Staminate flowers in elongated
catkins, each consisting of two to eight
stamens inserted on the torus-like base
of the oval or oval-lanceolate bracts of
the catkin, usually subtended by two
or four or rarely by numerous bracte-
oles; filaments short or elongated,
filiform, free or connate at the base
into a short stipe; anthers ovoid, erect,
two-celled, extrorse, showing longitu-
dinal dehiscence. Pistillate flowers in
ovoid or ovoid-globular catkins.
Gynoecium of two united carpels on
a bract. Ovary sessile, unicellular,
subtended by two lateral bracteoles
which persist under the fruit, or by
eight linear-subulate bracteoles, accres-
cent, and forming a laciniate involucre
inclosing the fruit; styles short and
dividing into two elongated style arms
which bear stigmatic surfaces on their
inner face; ovule orthotropous, solitary,
with a basilar placenta and superior
micropyle. Fruit an akene or cerifer-
ous nut. Pericarp covered with glan-
dular emergences which secrete wax or
fleshy emergences, smooth and lustrous
or smooth, glandular. Seed erect, ex-
albuminous, covered with a thin testa.
Embryo straight, cotyledons thick,
plano-convex; radicle short, superior.

Fic. 373.—F lowers and fruit
of willow (Salix viminalis). <A,
twig with staminate catkins.
B, staminate flower with sub-
tending bract, enlarged. C,
pistillate catkin. D, E. pistil-
late flowers, enlarged. F, fruit,
natural size. G, the same en-
larged. H, seed _ enlarged.
(From Strasburger, Macmillan Co.)

There are two distinct genera of this family, e.g., Myrica and

Comptonia.

502 PHARMACEUTICAL BOTANY

UnorriciAL Druc Parr Usep BoraAnicAL NAME HABITAT
Myrica Bark of root Myrica cerifera veiw North
Ma carolinensis America
Comptonia Leaf Comptonia aspleni- Eastern North
folia America
Bayberry Wax Wax from fruits Myrica cerifera me North
Myrica carolinensis America

Fic. 374.—Fructiferous branches of Myrica caroliniensis (to left) and Myrica
cerifera (to right) as they appear in early winter. The former species is decidu-
ous while the latter is evergreen. Collected by author at Wildwood, N. J., in
mid December,

ORDER JUGLANDALES

JUGLANDACEZ OR WALNUT Famity.—Tall, branching,
monececious trees or shrubs with branches and leaf-stalks usually
beset with glandular-resinous hairs. Leaves alternate, exstipu-

THE DICOTYLEDONS 503

late and odd-pinnate which, upon falling, leave usually horse-
shoe shaped scars on branches, and in whose axils develop
frequently one axillary bud and several ascending accessory
buds. Inflorescence of staminate catkins and pistillate spikes.

Fic. 375.—Fructiferous branch of Myrica Gale, showing mature pistillate catkins
and nature of leaves. Collected at the south end of Peak’s Island in Casco Bay,

Maine, September, 1913.

The staminate catkins occur on the old wood of the previous
year. The pistillate spikes are borne at the extremity or on the
sides of the new wood. The staminate flower is composed of
an elongated floral disk bearing a terminal bract, a perianth of
6 to rarely 4 parts and 40 to 4 stamens. The pistillate flower is
composed of an upgrown, hollowed-out receptacle bearing on

504 PHARMACEUTICAL BOTANY

its margin or extremity 5 to 4 perianth segments and containing
a pistil consisting of an inferior ovary with a single ovule and 2
styles having radiating penicillate stigmas. Fruit an aromatic
drupe composed of fused recep-
tacular tube and ovarian wall
and exhibiting a leathery to
rarely membranous epicarp,
more or less succulent mesocarp,
and stony endocarp enclosing a
seed with a double layered seed
coat surrounding an enlarged
embryo with irregular, semi-
ruminate cotyledons, into and
between which the fruit and seed
layers grow.

| The family embraces six
genera, of which Carya (Hicko-
ries) and Juglans (Walnuts) are
represented in the United States.
The seeds of Carya Pecan (Pecan)

sees

. Fic. 376.—The sweet-or black birch,

Betula lenta. Above, branch with stami-
nate catkins. Below, branch with

pistillate catkins. The tree occurs in
rich woodlands from Newfoundland to
Florida. (Reproduced from U. S. Dept.

and C. ovata (Shagbark Hickory)
are valued as food. The wood
of the walnuts is used for furni-

Agric. Miscell. Publ. 77.
gric. Miscell. Pu ) ture and the seeds of Juglans

cinerea (Butternut), 7. nigra (Black Walnut) and 7. regia (English
Walnut) are prized as foods. Black walnut hulls are used as a
coloring agent.

Part Usep
Inner root bark
Leaves, bark,
hulls, kernels

UNoFFICIAL DruG
Juglans

Black Walnut 1

‘Hasirrar
United States

BorAnicAL NAME
Juglans cinerea

Juglans nigra United States

ORDER FAGALES

BETULACE2 OR BircH Famiry.—A family of aromatic trees
or shrubs distinguished by monaecious flowers with scaly bracts
and astringent resinous bark. Differs from Fagacee by superior
ovary and absence of cupule. To this family belong the hazel-
nuts (Corylus), birches (Betula), alders (Alnus), the hop hornbeam,
ironwood (Ostraya) and the American hornbeam (Carpinus).

THE DICOTYLEDONS 505

The seeds of the hazelnuts or filberts are prized as food. The |
bark and oil of the sweet- or black-birch are employed as |
flavoring agents and antirheumatics.

OrriciAL Druc Part Usep BoTANICAL HABITAT
NAME
(Oleum Betulz) Volatile oil Betula lenta North America
Methylis Salicylas
Oleum Betula Empyreu- Pyroligneous oil Betula alba Eurasia

maticum Rectificatum
UNoFFICIAL Druc
Birch Bark Bark of stem Betula lenta North America

Fic. 377.—Quercus infectoria—Branch and nutgall. (Sayre.)

506 PHARMACEUTICAL BOTANY

FAGACE# OR BEECH FAMILY (CUPULIFER&).—Beeches, Chest-
nuts, Oaks, the trees of this family, are found in the temperate
forests of the eastern and western hemispheres and comprise
about 368 species. North America has over 50 species of oaks;
2 species of Chestnuts; 1 species of beech and 1 species of golden-
leaved chestnut. The most important American oaks used for
building, for furniture, for cooperage, for wagons, for tanning
_. leather etc. are white oak, Quercus
alba; chestnut oak, Q. prinus; black
oak, Q. velutina; live oak, Q. vir-
_ gimana; swamp white oak, Q.
platanoides; cow oak, Q. Michauxii,
and the two Pacific coast oaks, Q.
__ chrysolepis and post oak, Q. garryana.

_ The uses of the fast disappearing
American Chestnut, Castanea
dentata, are well known. The
_ wood of the beech, Fagus grandi-
@® folia, is used for chairs, tool
-%® handles, plane stocks, shoe lasts

and for fuel. The nuts (mast)
_ _ § fatten hogs and feed wild animals
Fic. 378.—Ulmus fulva, the Slippery and birds. The cork of commerce

Elm. Flowering branch bearing bell- i :
shaped flowers, leaf and leaf and fruit- Ss obtained from the bark of

ing branch. The fruits are circular, Quercus Suber and Quer cus occiden-
. single samaras. ‘The inner bark of this talts, plants indigenous to Spain
tree is rich in mucilage. (Reproduced and France.

from U.S. Dept. Agric. Miscell. Publ. 77.)

‘

The above trees are all monce-
cious, that is the staminate (male) and pistillate (female) flowers
are distinct from each other, but borne on the same tree. Most
of the species are trees, a few oaks are shrubs. The leaves are
simple, netted-veined and alternate. A pair of deciduous stipules
are found at the base of the leaf-stalk (petiole). The margins of
the chestnut and beech leaves are sharply cut with large teeth.
The leaves of the oaks divide the genus into two groups, viz.—the
white oaks with rounded lobed leaves and annual acorn production,
and the black oaks with sharp, bristle-tipped lobes and biennial
acorn production. The male flowers are in dangling heads

THE DICOTYLEDONS 507

(beech), or in catkins (chestnut and oaks). The male flowers have
a united perianth, which is 4-6 parted and encloses an indefinite
number of undivided stamens. The female flowers have a
superior 6-parted perianth; the pistil consisting of 3 carpels
with a corresponding number of stigmas. The ovary is 3-6
celled and each cell has 2 pendulous ovules. The fruit is a one-
seeded nut. The cup, or cupule, in the beech is 4-sided and
covered externally with weak spines and encloses two 3-sided
seeds. The cupule in the chestnut forms the spiny bur, which
splits into 4 valves at maturity, enclosing 3 nuts. The cupule
in the oak is saucer-, or cup-shaped, and encloses a single rounded
nut, or acorn. The seeds are exalbuminous and the cotyledons
are thick and fleshy, edible in the beech, chestnut and a few of
the oaks.

OrrictAL Druc Part Usep BorANICAL NAME HABITAT
Galla Excrescence Quercus infectoria Europe and W. Asia
Castanea Leaf Castanea dentata North America

Acidum Tannicum Tannin from gall Quercus infectoria Europe
Acidum Gallicum Product made from tannin

Pyrogallol Product made from gallic acid or extract of galls
UnorriciAL Druc
Quercus Inner bark. Quercus alba North America

‘ORDER URTICALES

Utmacem or Exim Famity.—Forest trees indigenous to the
temperate and tropical zones, characterized by being woody
plants, with simple, often inequilateral, pinnately-veined leaves
having caducous stipules and without milky juice. Their
flowers are unisexual or hermaphroditic with six or four parts
to the calyx. Fruit a single winged samara (Elms) or a drupe

(Hackberry).

OrrictAL DruGc Part UsEeD BorANICAL NAME HasiratT
Ulmus Inner bark Ulmus fulva United States
and Canada

MorAce# on Mutserry Famity.—Mostly apetalous shrubs
or trees, as the mulberries (MM orus), figs (Ficus), osage orange
(Maclura), etc., rarely herbs as hop and hemp, many of them

‘
:
;
.
:

508 PHARMACEUTICAL BOTANY

containing a milky juice in latex tubes with small, axillary,
clustered or solitary, unisexual flowers with sepaloid perianth,
variously colored; leaves either simple, ovate with serrate margin

Fic. 379,.—Hop (Humulus lupulus), a member of the Moracee. A, portion of
plant showing pistillate inflorescences; B, staminate inflorescence; C, rachis of
pistillate inflorescence (“hop”). (Robbins.)

or palmately lobed (fig) to palmately compound (hemp) and
having caducous stipules; ovules mostly amphitropous; fruit an
akene enclosed by the persistent perianth, as in hop and can-

THE DICOTYLEDONS 509

Scr

Fic. 380.—Cannabis sativa or Indian Hemp (Fam. Moracez). At left, staminate
plant, bearing axillary panicles of staminate flowers. At right, young pistillate
plant whose panicles constitute the drug Cannabis. Note the palmately compound
leaves, the leaflets of which are lanceolate with serrate margins.

nabis, a sorosis, as in the mulberry and osage orange or a
syconium, as in the fig; embryo usually curved.

OrriciAL Druc Part Usep BoTrANnicAL NAME HAsItraAT
Cannabis Flowering tops of 1 ee ativan ‘i
pistillate plant

510 PHARMACEUTICAL BOTANY

OrriciAL Druc Part Usep BotANicAL NAME Hasirat
Humulus Strobile Humulus lupulus ee Asia
Lupulinum Glandular trichome Humulus lupulus (North America
UnorrFiciAL Druc

Fructus Cannabis Fruit Cannabis sativa Asia

(Hemp seed)

Ficus : Fruit Ficus Carica Persia

ORDER SANTALALES

SANTALACEZ OR SANDALWOOD FamILy.—Herbs, shrubs or
trees having entire exstipulate leaves, greenish flowers, nut-like
or drupaceous fruits and oily seeds. Many are parasitic on the
roots of other plants or on trees. The woods and volatile oils
of a number of species are of economic importance.

OrrFiciAL Druc ParT UsED BoTANICAL NAME HABITAT
Oleum Santali Volatile oil
Santalum Album Heartwood

UnorrictaL Druce
Oleum Santali Volatile oil Eucarya spicata W. Australia
Australiensis

t Santalum album India and Malaya

LoORANTHACEA OR MistLeTor Famity.—Most of the mem-
bers of this family are woody shrubs, parasitic on trees; leaves
opposite or whorled, entire, simple, exstipulate, sometimes
reduced to scales; flowers hermaphroditic or dioecious, the calyx
annular or cup-like, adnate to the ovary; petals disjoined or
united into a tube, often split down one side, stamens equal in
number to the petals and inserted on their base, the anthers
dehiscent by pores or terminal slits: nectar disk present or absent;
ovary inferior with simple style or sessile stigma; fruit a berry or
drupe with a solitary seed devoid of testa and containing copious
endosperm and a straight embryo.

UnorrictAL Druc Part Usep BoTANIcAL NAME HABITAT
Phoradendron Entire plant Phoradendron flavescens - United States
(American Mistletoe)

Viscum Entire plant Viscum album _ Europe and Asia

(European Mistletoe)

ORDER ARISTOLOCHIALES

ARISTOLOCHIACEE OR BirtHwort Famity.—Herbs or twin-
ing semi-woody or woody plants, having more or less swollen

THE DICOTYLEDONS 511

nodes from which spring alternate, cordate, reniform or ovate
leaves. Flowers regular (Asarum, etc.) or irregular (Aristolochia),
often offensively smelling. Sepals varying from 6 to rarely 5
to more commonly 4 or 3; in Asarum, etc. forming a regular’
symmetrical cup with 3 teeth or lobes, in Arzstolochia divisible
_ into a bowl, a tube and a limb. Stamens from 24 to 3, the fila-

Fic. 381.—Aristolochia Serpentaria. (Sayre.)

ments distinct or slightly fused with styles, in Asarum, or com-
pletely fused, in Aristolochia, to form a gynostemium, the anthers
separate. Pistil of as many carpels as stamens, the stylar por-
tions more or less distinct from stamens in Asarum, fused in
Aristolochia, the ovary semi-inferior to inferior in Asarum, com-
pletely inferior in Aristolochia and as many celled as there are
carpels. Fruit a capsule. Seeds with copious albumen and
minute embryo.

512 PHARMACEUTICAL BOTANY

OrrictAL Druc Part Usep BoTANICAL ORIGIN HABITAT
Aristolochia

Serpentaria Rhizome and roots Bers ania United States
Aristolochia
reticulata

Asarum Rhizome and roots Asarum canadensis United States

OrDER POLYGONALES

PoLyGonace® Famrty.—Usually herbs [Knot-grasses (Poly-
gonum), rhubarb (Rheum), docks (Rumex), buckwheat (Fagopy-

Fic. 382.—Serpentaria—Cross-section of rhizome. (25 diam.) A, parenchyma
of cortex; B, medullary ray; C, xylem; D, phloem; FE, medulla. (Sapre.)

rum), etc.| rarely trees (Coccoloba uvifera or Sea-Grape) or shrubs
(Muehlenbeckia or Ribbon Bush) having strong vertical tap roots
and ‘Spreading secondary roots more or less provided with
tannin compounds. Stems elongate, green, to woody, rarely
flattened, leathery, phylloidal (Muehlenbeckia platyclada), still
more rarely tendriliform (Antigonon leptopus or Mountain Rose
of Tropical America). Leaves alternate, rarely opposite or
_whorled (Eriogonum), entire, rarely lobed (Rheum palmatum,
Rumex acetosella), petiolate, rarely sessile, and stipulate. Stipules
fused and forming a greenish membranous upgrowth (ocrea)
_ which sheaths the stem. Inflorescence racemose with many
dense scorpioid or helicoid cymes, which in some forms condense
into single flowers. Flowers regular, pentamerous, with simple
calyx, becoming trimerous with two whorls of three sepals each.
Stamens varying from fifteen or twelve to nine or six, more

THE DICOTYLEDONS 513

rarely to five, four, three to one (Kenigia), hypogynous, more
rarely by enlargement of receptacle and slight fusion of sepals,
perigynous, Pistil tri- to bi-carpellate, often three- to two-
sided; ovary one-celled with one ovule. Styles three, rarely
two, radiating penicillate in wind-pollinated, inconspicuous
flowers, becoming condensed, knob-like in conspicuous insect-
pollinated flowers. Fruit a triangular or biconvex akene often
crowned by persistent styles and surrounded by persistent closely

Nectar -&

glands X

Fic. 383,—Rhubarb (Rheum) flower, external view, median lengthwise section,
and with perianth and stamens removed. (Robbins, after Luerssen.)

applied sepals. Seeds solitary, albuminous, with straight
embryo, or in Rumex, curved embryo.

OrrIciAr Drug PART UsED _ Boranicar Origin Hasirat
Rheum officinale
Rheum palmatum and

i other species (except-

Rheum gran — ing R. rhaponticum) or China and Thibet
ee : hybrids of Rheum,

*{ grown in China and
Thibet

Unorricra, Drue
Rumex crispus

Rumex Root Rumex Europe
obtusifolius
Bistorta Rhizome Polygonum Bistorta Europe and Asia

OrDER CHENOPODIALES OR CENTROSPERMZ

This order contains seven families, all of which have seeds
with a curved or coiled embryo. These families are the Goose-

514 PHARMACEUTICAL BOTANY

foot Family (Chenopodiaceae), the Pokeweed Family (Phytolaccacee);
the Amaranth Family (Amaranthacee) which includes the orna-
mental Cock’s Comb; the Four-o’clock Family (Nyctaginacee) to
which belong the Marvel of Peru or Four-o’clock and the
beautiful Bougainvilleas; the Azzoacee containing the Carpet
Weed (Mollugo verticillata) and the Sea Purslane (Sesuvium
maritimum), etc.; the Portulaca Family (Portulacacee) which is
represented by the Spring Beauty (Claytonia virginica) and the
Purslane (Portulaca); the Pink Family (Caryophyllacee) which
includes the Carnations, Corn Cockle, Bouncing Bet, etc.
CHENOPODIACE2 OR GoosEFooT Famity.—Usually herba-
ceous halophytes or shore growers, growing also in any alkaline
soil, more rarely shrubs (Atriplex) or low trees (Haloxylon).
Among them are several garden vegetables (Spinach, Beets,
Mangels) and a number of weeds. Leaves alternate to opposite
sometimes reduced to teeth, entire or lobed. Inflorescence
spikes or short.racemes of condensed cymes. Flowers regular,
usually small and greenish. They are either perfect (Beta),
monoecious (Chenopodium), dicecious (Aériplex sp.), or poly-
gamous (Kochia or Summer Cypress). Fruit a utricle. Seed
albuminous. Embryo curved, enclosing central albumen.

OrrictAL Druc PART Usep BoTranicAL NAME HABITAT
Oleum Chenopodii Volatile oil Chenopodium ambrosioi- United States
des var. anthelminticum

Sucrosum Refined sugar ee wang Prope
Beta maritima
UnorFiciAL Druc

Chenopodium Fruit Chenopodium ambrosioi- United States
des var. anthelminticum

PHYTOLACCACE2 OR PoKE Famivy.—A family of apetalous
trees, shrubs, and herbs with alternate, entire leaves, and flowers
resembling those of the goosefoot family (Chenopodiacee), but
differing in having the several-celled ovary composed of carpels
united in a ring, and forming a berry in fruit. It embraces 21
genera, and 55 species, mostly tropical and sub-tropical... Only
the genus Phytolacca occurs in the United States.

OrFIcIAL DruG PART Usep BoTANIcAL NAME HasrratT

Phytolacca Root Phytolacca North America
(Poke Root) americana

THE DICOTYLEDONS 515

ORDER RANALES

This order includes the following families: Nympheacee or
Water Lily Family, Magnoliacee or Magnolia Family, Ranuncula-
cee or Crowfoot Family, Berberidacee or Barberry Family,
Menispermacee or Moonseed Family, Anonacee or Custard Apple

Fic. 384.—Phytolacca americana (in foreground), growing in damp woodland.

Family, Mpristicacee or Nutmeg Family, Monomiacee or Boldo
Family, and Lauracee or Laurel Family.

MAGNOLIACEH OR Macnouia Famity.—Trees and shrubs
having alternate leaves with large deciduous stipules that
enclose the young buds and single large flowers with several
series of sepals and petals which are often colored alike. Sepals
and petals deciduous. Carpels and stamens numerous, spirally

516 PHARMACEUTICAL BOTANY

arranged and free. Bark aromatic and bitter. Fruit a collec-
tion of follicles dehiscing dorsally. Seeds large, albuminous and
oily.

Fic. 385.—Mpristica fragrans—Branch and fruit. (Sayre.)

To this family belong the stately Tulip Poplar, Liriodendron

Tulipifera, prized for its wood and beautiful flowers, and the
showy Magnolias of our gardens.

OFFIciAL Druc Part Usep BoTANICcAL NAME HABITAT
Oleum Anisi Volatile oil Illictum verum South China
UNoFFICIAL DruG
Winter’s Bark Bark,.::; Drimys Winteri South America

Illicium Fruit Illictum verum S. Chin
(Star Anise) er

THE DICOTYLEDONS - 517

MyristicAcE& or Nurmec Famity.—A group of apetalous
trees comprising the genus Myristica, of about 80 species.

Myristica.—A large tropical genus of fragrant, apetalous
trees—the nutmegs—coextensive with the nutmeg family, having.

aggregate fruit. (After Strasburger, Noll, Schenck and Schimper.)

alternate, entire, often pellucid punctate leaves, small dicecious,
regular flowers, and a succulent, two-valved, one-celled fruit
with a solitary seed containing ruminate albumen and covered
by a fleshy arillode.

518 PHARMACEUTICAL BOTANY

M. fragrans, a handsome tree, 20 to 30 feet high, indigenous
to the Molucca Islands in the East Indian archipelago but
now cultivated extensively in the West Indies, supplies the nut-
megs and mace of commerce.

OrFicIAL Druc Part Usep BoTANICAL NAME HABITAT
Myristica Kernel of seed Myristica.fragrans
Oleum Myristice Volatile oil Myristica fragrans a
UnorriciAL Druc
Macis _ Arillode Myristica fragrans

RANUNCULACEZ OR BuTTERCUP FAmity.—Herbs, rarely
shrubs (Clematis) with acrid, poisonous,
watery juices and with alternate, rarely
opposite, simple, rarely compound,
exstipulate leaves. Flowers pentam-
erous, regular to irregular, incomplete
to complete, aposepalous and apopet-
alous. Sepals five—rarely more or less
—green to petaloid, regular, passing to
irregular (Larkspur, Monkshood).
Petals none or five, regular to rarely
irregular, often nectariferous and with
nectariferous petals often variously
transformed. Stamens __ indefinite,
hypogynous. Pistil of many to few
apocarpous carpels, each carpel with
one to several ovules. Fruit a collec-
tion of achenes as in Buttercup (Ranun-
culus), Pasque Flower (Pulsatilla), or a

\ collection of follicles (Columbine,

Larkspur, Peony, Aconite) or rarely a
Fic. 387.—Above ground berry as in Baneberry (Actea) or head
portion of Aconitum Napellus of crimson berries as in Golden Seal

Odonkshoad) caring”: Bae (Hpdrastis). Seeds albuminous with q

mately-divided leaves and

hooded flowers. large oil-containing endosperm and
small embryo.

OrFIcIAL Druc Parts Usep Botanica, NAME Hasrrat

Hydrastis Rhizome and roots Hydrastis canadensis Eastern United

(Golden Seal) Stites and

Canada

THE DICOTYLEDONS 519

OrFIciIAL DruG PARts Usep BoTANICAL ORIGIN HABITAT
Aconitum Tuberous root Aconitum Napellus —__ Europe, Asia
Cimicifuga Rhizome and roots Cimicifuga racemosa E. North America

Pulsatilla vulgaris E :
Pulsatilla Entire herb Pulsatilla pratensis oe ae ee
Pulsatilla patens W. North America
Adonis Overground portion Adonis vernalis Europe and Asia
Delphinium Seed : Delphinium Ajacis Europe
UnorFIciAL DruG
Aconiti Folia Leaves and flower Aconitum Napellus Europe and Asia
tops
Coptis Entire herb Coptis trifolia United States
(Goldthread) and Canada
Helleborus Rhizome and roots Helleborus niger Alps
Nigella Seed jee sativa E
(Black Caraway) Nigella damascena gies i
: : Delphinium South Europe,
bie: nats, alee i: Staphisagria Ta hthice
Western Coptis Entire herb Coptis occidentalis Western United
States
Bear’s Foot Root Rhizome and roots Helleborus foetidus | Europe
Xanthorhiza Rhizome Zanthorhiza apiifolia United States

BERBERIDACEZ OR BARBERRY FAmiILy.—Herbs and woody
plants with watery juices and alternate or radical, simple or
pinnately-compound, mostly exstipulate leaves, either stems
or leaves of Berberts (Barberry) species often bearing spines or
barbs, which give them a barbed appearance. Rhizomes and
roots often of yellow color internally, due to berberine content.
Flowers either in racemes of cymes, cymes, racemes or solitary.
Sepals and petals usually in 2 series of 3 each, similar, hypog-
ynous; stamens 9 to 4, hypogynous; pistil single. Fruit a
berry or capsule. Seeds exalbuminous with a straight large
embryo in axis of albumen.

OrrictAL Druc Parts UseD BOTANICAL ORIGIN HAsitat
Berberis Rhizome and roots Berberis species of Western North
(Barberry) * the Sect. Mahonia = America
Podophyllum Rhizome and roots Podophyllum Eastern North
(May Apple) peltatum America
Caulophyllum Rhizome and roots Caulophyllum Eastern North
thalictroides America .

UnorFIciAL DruG —
Podophyllum Rhizome and roots Podophyllum emodi Tibet and

Indicum B.P. : Afghanistan
Barberry Bark Bark of root and Berberis vulgaris Europe
stem

520 PHARMACEUTICAL BOTANY

Fic. 388.—Cimicifuga racemosa—Plant and rhizome,
pound leaf, the wand-like racemes which bear white
rhizome. This perennial herb grows in rich woods, It is
Snakeroot and Black Cohosh and belongs to the Ranunc.

Note the pinnately decom-

flowers, and the knotted
commonly known as Black
ulacee. (Sayre.)

THE DICOTYLEDONS 521

MENISPERMACEZ, OR MoonsgEED Famity.—Choripetalous,
woody, climbing, mostly tropical plants with alternate, exstipu-
late, simple often peltate leaves. Flowers green to white. Fruit

Fic. 389.—Podophyllum peltatum. Entire plant, above ground portion, and
fruit. Note the flowering stems bearing in each instance two one-sided leaves
and a nodding flower from the forks. This plant also sends up from its rhizome
flowerless stems each of which terminates in a 7—9 lobed peltate leaf.

a one-seeded succulent drupe. Seeds albuminous. They usu-
ally contain tonic, narcotic or poisonous bitter principles.

OrrFiciAL Drue Parts UseD BoTANIcAL NAME Hasitat

Calumba Root Jateorrhiza East Africa
palmata

Cocculus Fruit Anamirta Cocculus Asia

UnorrFictAL Druc

Menispermum Rhizome and roots Menispermum United States

(Yellow Parilla) canadense and Canada

Pareira Root Chondodendron Brazil and

tomentosum Peru

522 PHARMACEUTICAL BOTANY

MoNnoMIACE# OR Botpo FamiLy.—Tropical trees and shrubs
with opposite or rarely alternate, entire or serrate, pellucid-
punctate leaves devoid of stipules, but containing cystoliths.

Flowers hermaphrodite or unisexual in cymes or racemes;
calyx of many to 4 lobes or teeth in 2 or more series, stamens
numerous to few, hypogynous, carpels several to one, each with

Fic. 390.—Blue Cohosh (Caulophyllum thalictroides). Top of plant. Note the
triternately compound leaf bearing 2- to 4-lobed leaflets and the panicle of yellowish-

green flowers. The fruits which ripen in August are dark blue berries. (Reproduced
from U.S. Dept. Agric. Misc. Publ. 77.)

a single ovule. Fruit a drupe (Peumus), nut or achene. Seeds
with membranous testa and fleshy endosperm.

OrrFiciAL Drug Part Usep

BoTanicAL NAME HasiraAT
Boldus Leaf

Peumus Boldus Chile

Laurace® or Spicepusy F AMILY.—Shrubs or trees of sub-
tropical or tropical, rarely of temperate regions. Bark, wood and
leaves rich in spicy, aromatic, hydrocarbon oils. Leaves alter-
nate, simple, entire, often leathery and shining and exstipulate.
Inflorescenes usually cymose clusters. Flowers small, green,
yellow or rarely whitish, hermaphrodite or more or less dicecious;
regular calyx alone present as a floral whorl, of three to six

THE DICOTYLEDONS 523

small sepals. Stamens four to twelve in several rows of three to
four each; anthers continuous with the filaments, each anther
opening by recurved valves, the valves opening from the base
upward. Pistil monocarpellary, ovary superior, one-celled with

Fic. 391.—Jateorhiza palmata, the source of Calumba—Portion of vine. Note

tendril for winding about a support as this plant climbs. (Sapre.)

solitary pendulous ovule, style’ simple with usually rounded
stigma. Fruit a succulent berry (Spicebush and “Bacca
laurea’’) or a succulent drupe (Sassafras). Seeds usually

richly albuminous.

OrriciAL DruGc Part Usep BoTraANicAL NAME HABITAT
Camphora Ketone Cinnamomum Eastern Asia
Camphora
Sassafras Bark of root Sassafras North America

variifolium

524 PHARMACEUTICAL BOTANY

OrrFiciAL DruGc ParT Usep BotaAnicAL NAME HABITAT
Oleum Sassafras Volatile oil from Sassafras varii- © North America
root folium
Cinnamomum Bark Cinnamomum Cochin China
Loureirii
Oleum Cinnamomi Volatile oil Cinnamomum China
Cassia
UnorriciAL Drucs
Cinnamomum Bark Cinnamomum Ceylon
Zeylanicum Zeylanicum
Coto Bark Nectandra Coto Bolivia
Laurus Leaves Laurus nobilis Europe
North America
Fagot cassia Bark Cinnamomum Asia
Burmanii
Clove Bark Bark Dicypellium » Beez
caryophyllatum
Cassia Buds Immature fruit Cinnamomum Cochin China
, Loureirii
Cassia Bark Bark Cinnamomum China
Cassia
Sassafras Medulla _ Pith Sassafras North America
variifolium

Orver Rua@apALes (PAPAVERALES)

PAPAVERACEZ OR Poppy Famity.—Herbs or low shrubs.
Root only, in some cases root and stem, and in still others the
entire plant, traversed by anastomosing latex tubes that contain a
milky juice, varying from white (Papaver or Poppy) or pale-
yellow, as in Celandine (Chelidonium), to red (Bloodroot). Latex
tubes in the axis wholly confined to cortex or phloem but occa-
sionally sending branches into the medullary rays, rarely into
the pith of the plant. Leaves alternate, varying from simple
(Platystemon) to pinnatifid, pinnatipartite or even pinnately-
compound. Inflorescence varying from a loose raceme or
panicle of cymes to a raceme of cymes or condensing until, as in
Papaver, only one large terminal flower is left. Flowers regular
tetramerous or dimerous. Sepals typically two, more rarely
three. Petals typically four, more rarely six. Stamens indef-
inite, in most forms hypogynous, except in California Poppy
(Eschscholtzia) where hypogyny is modified into perigyny. Pistil
of sixteen to four, rarely two carpels generally fused together.
Placenta parietal. Ovules numerous, anatropous. Fruit a

THE DICOTYLEDONS 525

S e ®
AWS .
OR
Nie eae
\

Sreava N

SSS

S

~ SS
SS
a FF

\
\

hos

Uy)
Vr,

Wf

Fic. 392.—Cinnamomum Zeylanicum, the source of Ceylon cinnamon bark.

A member of the Lauracee. (Sayre.)

Fic. 393.—Flower of Ceylon Cinnamon (Cinnamomum Zeylanicum). Note
the uplifted lids of the anther locules in dehiscence, the calyx of six sepals (in a), the
perigynous stamens and 1-celled ovary with pendulous ovule, in 6. (Agter Baillon.)

526 PHARMACEUTICAL BOTANY

Fic. 394.—Bloodroot (Sanguinaria
open woods of eastern North America.

canadensis), a perennial herb native to rich
The waxy white flowers and grayish green
5-9-lobed, kidney-shaped leaves make their appearance only in spring. The
rhizome containing a red latex constitutes the drug. It should be gathered in early
summer, deprived of its rootlets and carefully dried. The drug possesses expectorant
and emetic properties. (Reproduced from U. S. Dept. Agric. Misc: Pub. 77.)

THE DICOTYLEDONS

527

Fic. 395.—Papaver somniferum—F lowering branch and fruit. (Sayre.)

capsule (Papaver) or 2-valved pod (Bloodroot, etc.). Seeds
richly albuminous.

OrriciAL DruG
Opium

Morphine Sulfas
Morphinz
Hydrochloridum
Codeina

Codeine Phosphas

Codeine Sulfas
Papaverinz
Hydrochloridum

Part UsepD
Air-dried milky
exudate from
unripe capsules
Sulfate of alkaloid
Hydrochloride of
alkaloid
Alkaloid
Phosphate of alka-
loid
Sulfate of alkaloid
Hydrochloride of
alkaloid

BorTANICAL ORIGIN HABITAT

Eastern Mediter-
ranean countries

Papaver somni-
ferum and its
var. album

528 PHARMACEUTICAL BOTANY

OrricAL Druc Part Usep BoTANICAL ORIGIN HABITAT
Sanguinaria Rhizome Sanguinaria United States and
canadensis Canada
UnorriciAL Druc
Chelidonium Entire flowering Chelidonium United States and
plant majus Canada ;
Maw seed Seeds Papaver somni- Eastern Mediter-
ferum var. nigrum ranean countries
Papaveris Fructus —_ Unripe capsules Papaver somni- Eastern Mediter-
ferum ranean countries
Papaver Rhoeas Fresh petals Papaver Rhoeas Europe

Fic. 396. Fic. 397. Fic. 398.
Fic. 396.—Transverse section of flower of Poppy. (Sayre.)
Fic. 397.—Gyneecium of Poppy, with one stamen remaining. (Sayre.)
Fic. 398.—Transverse section of ovary of Poppy. (Sayre.)

FUMARIACEA OR Fumirory Famity.—Delicate herbs, rarely
shrubs containing milky-watery to watery latex containing
alkaloids. Leaves radical, alternate and more or less compound.
Inflorescence a raceme or spike. Flowers irregular, hermaph-
rodite, zygomorphic; sepals 2, petals 4, the two outer often
saccate or with a spur at the base; stamens 4 or 6; ovary superior,
1-celled with 2 parietal placentas. Fruit a one-chambered
capsule. Seeds albuminous. Idioblasts common.

OrrFicIAL DruG Part Usep BoraAnicAL NAME HaAsitratT
Corydalis Tubers Dicentra canadensis Ethie States and
Dicentra Cucullaria ) Canada

CrucIFER& OR Musrarp Famity.—Herbs, rarely shrubs,
mostly of temperate regions. Many species of this family con-
tain glucosides which on hydrolysis yield pungent volatile oils.
Stem and branches upright or diffuse spreading (Arabis).
Leaves alternate, simple, rarely compound, exstipulate, entire
or toothed, often more or less hairy. Inflorescence at first

THE DICOTYLEDONS 529

a corymb or shortened raceme, later elongating into a loose
raceme. Bracts at base rarely reduced, usually absent. Flowers
regular, rarely irregular (Candytuft), tetramerous. Sepals four,
green, equal, or two laterals at times pouched as nectar recep-
tacles. Petals four, yellow, white, pink or purple, cruciform,
often divisible into claw and blade. Stamens six, four being
long and anteroposterior in position, two being short and lateral,

Fic. 399.—The Black Mustard, Brassica nigra—Branch. (‘Sayre.)

hence termed tetradynamous, insertion hypogynous. Pistil
syncarpous, bicarpellate, superior, carpels lateral. Ovary one-
celled but falsely two-celled by a placental replum (a spurious
membranous partition dividing the ovary into 2 locules); style
simple; stigma rounded or bifid or bilobed. Ovules several,
rarely few, attached to marginal placenta. F ruit a siliqua, as in
the mustards or silicule, as in the shepherd’s purse (Capsella)
rarely indehiscent, bursting lengthwise by two valves. Seeds
exalbuminous with folded cotyledons. (See Fig. 400.)

530 PHARMACEUTICAL BOTANY

OrriciAL Druc Part Usep BotanicAL NAME HABITAT

Oleum Sinapis Volatile oil Flas nigra Europe and Asia
Volatile Brassica juncea Asia

Sinapis Nigra Seed (Brassica nigra Europe and Asia

Brassica juncea Asia

UnorriciAL Druc

Semen Rapz Seed Brassica napus Europe and Asia

Armoracia Root - Cochlearia Armoracia Europe

Sinapis Alba Seed Sinapis alba Europe and Asia

Capsella Aerial plant Capsella bursa-pastoris Europe

OrDER SARRACENIALES

This order includes several families of plants whose leaves are
modified for the purpose of catching and digesting insects.
The principal families are the Sarra-
P crn cemacee and Nepenthacee both of which
have their leaves modified as pitchers,
and the Droseracee@ which includes the
Fly-Traps and Sundews. (See pp.
622-638.)

DROSERACEZ OR SUNDEW FamILy.
Herbs (Drosera, Dionaea, etc.) rarely
shrubs (Roridula of South Africa),
growing in bogs or swamps or purely
aquatic in habit (Aldrovanda). Leaves,
either rosettes or more or less scattered
in alternate fashion over stem, usually
glandular-hairy or sensitive-hairy and
insectivorous. Inflorescence a loose
raceme or cymose umbel. Flowers
regular, pentamerous; sepals five,

_ Fic. 400.—Crucifer, Floral aposepalous, green; petals apope-
siagiendiagl aa pre talous, varying from white to whit-
(Robbins. ish-pink, pink, scarlet, purple to

bluish-purple; stamens varying from
fifteen to five, hypogynous; pistil syncarpous of five to three,
rarely fewer carpels. Fruit a capsule. Seeds albuminous.

‘

UnorricitaL Druc Parr Usep BoTANICcAL ORIGIN HApiraT

Drosera : Drosera rotundifolia E

(Sundew) Entire plant Drosera anglica astern a
Drosera longifolia ern Hemispheres

.

THE DICOTYLEDONS 531

ORDER ROSALES

SAXIFRAGACEZ OR SAXIFRAGE FamiLy.—Herbs (Mitrewort
(Mitella), Saxifrage (Saxifraga), Alum-root (Heuchera), etc.) or
shrubs (Hydrangea, Philadelphus or Mock-orange, etc.) rich in
tannin content, with opposite or alternate leaves usually devoid
of stipules. Flowers regular, hermophrodite; sepals 5 usually,

——~

s-

|

a

Fic. 401.—Drosera rotundifolia, Production of a young plant from the leaf of an

older plant. (Gager.)

petals 5 to 10, sometimes absent, petals and stamens perigynous to
epigynous; ovary 3 to 1 celled, free or adnate to upgrown recep-
tacle. Fruit a follicle, capsule, or berry. Seeds small with
copious albumen and tiny embryo.

OrriciAL Druc Parts Usep BoTANICAL NAME HABITAT

Hydrangea Rhizome and roots Hydrangea United States
arborescens
UnorFIciAL DruG ;
Heuchera Root Heuchera United States
americana

Hines or Wircu Haze. Famity.—Shrubs or small
or large trees. Leaves simple, alternate and usually pinnately
veined; stipules deciduous to caducous, paired and slightly fused

Bie ait:
to eee

532 PHARMACEUTICAL BOTANY

at the bases of petioles. Flowers frequently yellow to yellowish-
white, in axillary clusters or heads or spikes, hermaphrodite or
moneecious; sepals and petals five to four, rarely indefinite,
superior (petals absent in Fothergilla); stamens twice as many as
the petals, but the outer row alone fertile, the inner row being
more or less barren, scale-like; gyncecium of two carpels united

Fic. 402.—Liquidambar orientalis, the Sweet i eck.

(Sapre.)

below. Fruit a two-beaked, two-celled, woody capsule dehiscing

at the summit, with a bony seed in each cell, or several, only one
or two of them ripening.

OrrFiciAL DruG Part Usep BoranicAL NAME eae
Styrax Balsam from wood — orientalis The Levant
(Storax) and inner bark

Liquidambar Styracifilua Central America
Hamamelidis Leaf Hamamelis United States

Folium virginiana and Canada
UnorFiciAL DruG
Hamamelidis Bark Hamamelis United States
Cortex virginiana

and Canada

THE DICOTYLEDONS 533

Fic. 403.—Hamamelis virginiana. Upper figure shows this shrub as it appears
in autumn after the leaves have fallen. Note that the plant is in blossom. Lower
figure shows a flowering branch from the same plant. The bright yellow flowers
occur in axillary clusters appearing at the same time as the ripening of fruits —
blossoms of the previous year. A leaf of the plant, commonly known as Witch
Hazel, is shown on p. 239. The recently cut and partially dried twigs and leaves
are used in the preparation of Witch-hazel water or Aqua Hamamelidis.

534 PHARMACEUTICAL BOTANY

RosAce# OR Rosz Famity.—Herbs, shrubs, or trees mostly of
temperate regions. Stem and branches upright or creeping
(Strawberry, Cinquefoil), herbaceous to woody. Leaves alter-
nate, stipulate (stipules green, persistent to scaly, deciduous),
compound, condensing to “simple.” Flowers regular, pen-

Fic. 404.—Branch of Quillaja Saponaria, a Chilean tree from whose trunk and
branches Soap Tree Bark is obtained. (Sayre.)

tamerous, either solitary, as in the Peach (Prunus persica) or
Apricot (Prunus armeniaca), in racemes, as in the Wild Black
Cherry (Prunus serotina), in cymes, as in Apple (Malus), or in
umbels as in Bird Cherry (Prunus Avium) and Sour Cherry
(Prunus Cerasus), sepals and petals 5, rarely 4, inferior to ovary,
becoming by stages superior to it. Sepals green, at times with

THE DICOTYLEDONS 535

epicalyx (Strawberry, Cinquefoil, Kousso, etc.), persistent round
fruit. Petals usually yellow to white or pink, crimson, rarely
purple, rosaceous, deciduous. Stamens indefinite, perigynous
(Strawberry, etc.), to semi-epigynous (Rose, Peach, etc.),
and epigynous (Apple, Pear, Quince). Pistil apocarpous with
many (Strawberry, Rose) carpels or fewer to five (Apple),
or two to one (Plum, Cherry), becoming falsely fused by union
with upgrowing receptacle (Hawthorn, Quince, Pear, Apple).
Fruit a collection of achenes on dry (Cinquefoil) or succulent
receptacle (Strawberry), or dry follicles (Bridal Wreath), or
drupels (Blackberry), or a drupe (Peach, Plum, Cherry), or a
pome (Apple, Quince, Pear). Seeds exalbuminous, embryo
filling seed cavity. .

Some botanists divide this family into three sub-families, each
of which, in the former American System, had been given the
rank of afamily. These according to this system of classification
are: Rosacea, Malacee and Amygdalacee. Rosacez include the
genera Geum, Agrimonia, Duchesnia, Fragaria, Potentilla, Wald-
steinia, Cercocarpus, Spirea, Rubus, Dalibarda and Rosa.

Malacee include the genera Sorbus (Mt. Ash), Malus (Crab
Apple and Apple), Amelanchier (Service Berry and Shad Bush),
Pyrus (Pear), Aronia (Choke Berry), Crategus (Hawthorn) and
Cotoneaster (Christ’s Thorn).

Amygdalacee comprise the genera Prunus (Plum group), Padus
(Wild Cherry group) and Amygdalus (Almond and Peach group).

OrriciAL Druc Part Usep BoraAnicAL ORIGIN HAsitat
Prunus Virginiana Bark Prunus serotina United States and
Canada
Quillaja Inner bark Quillaja Saponaria Chile and Peru
Rosa Petals Rosa gallica W. Asia and S. Europe
Oleum Rosz Vol. oil Rosa gallica id Asia and 8. Europe
Rosa damascena Balkans
Rubus Idzeus Fresh ripe Rubus Idzus and —_— Europe and Asia
fruit Rubus strigosus _
Prunus Cerasus Fresh ripe Prunus Cerasus Asia Minor
fruit
Oleum Amygdale Vol. oil from Amygdalus commu- W. Asia
Amare kernels nis, Prunus persica,
etc.
Oleum Amygdalze Fixed oil Amygdalus commu- W. Asia

Expressum nis (var’s.)

536 PHARMACEUTICAL BOTANY

UnorriciAL Druc Part Usep BoTANICAL ORIGIN HAsirAT
Amygdala Dulcis Seed Amygdalus commu- W. Asia
nis var. dulcis
Amygdala Amara Seed Amygdalus commu- W. Asia
nis
Folia Laurocerasi Leaves Prunus Laurocerasus Asia Minor, Persia
Cydonium Seed Cydonia vulgaris Cultivated widely
Rosa Centifolia Petals Rosa centifolia Western Asia
Rosa Canina Spurious Fruit Rosa canina Europe -
Tormentilla Rhizome Potentilla silvestris Europe and Asia ;
Rubus Bark of rhizome en Eubatus of United States :
and roots genus Rubus
Prunum Fruit Prunus domestica Caucasus region .
Brayera Panicles of Hagenia abyssinica Abyssinia ,
pistillate flowers
Succus Pomorum Fresh juice of Malus (cult. var’s.) | Europe ;
fruit : |
4

Fic. 405.—Prunus domestica, or Damson plum, the source of prunes. Fruiting branch
: and flowering branch. (Sayre.)

Lecuminos# or Pea Famiiy (FABACE®).—Herbs, shrubs or
trees of all regions, with tubercled roots. Stem usually erect,
rarely creeping (Trifolium repens). Leaves alternate, compound,
rarely simple, stipulate, sometimes tendriliform, as in Peas, or
reduced to phylloid petioles (some Acacia spp.). Inflorescence a

;
2
i
4
:
.

537

THE DICOTYLEDONS

;
i!
14M
ty)

'
N’

\\
\

=,"
—S
SW

iy

y
i,

Yi,

\ Waa

Fic. 406.—Glycprrhiza glabra—Branch. (Sayre.) The source of Licorice. The
varieties yielding the drug are perennial herbs with alternate, imparipinnate

leaves and axillary spikes of violet-colored papilionaceous flowers.

\ +}-—standard~__ =
Ze) }
wing~ {
a eat
B

Fic. 407.—Flower of Papilionacez. A, floral diagram of Vicia faba; B, sweet
(From Robbins, A after Eichler, B after Bergen

pea flower, dissected, diagrammatic.
and Caldwell.)

538 PHARMACEUTICAL BOTANY

raceme, at times, condensed almost to a head or capitulum (Spp.
of clover, Mimosa, etc.). Flowers pentamerous (rarely tetram-
erous as in Krameria), regular (Mimosacee), to irregular (Ces-
alpinacea) to very irregular with corolla butterfly-shaped
(Papilionacee). Sepals five united, green; petals five (rarely
four) variously related; in Papilionacee the one superior, external

-
SY)
WY,
ALD
Kor

Fic. 408.—Cassia Senna—Branch showing flower and fruit. (Sayre.)

and ‘posterior is called the standard or vexillum, the two lateral
form the wings or ale, the two inferior, internal and anterior,
slightly adherent, form the keel. Stamens ten to four, free, or in
Papilionacea united by filaments into a monadelphous (ten) or a
diadelphous (nine to one) ‘tube, inserted perigynously. Pistil
typically monocarpellary, ovary with sutural placentation, style
simple. Fruit a legume, more rarely becoming transformed into

loment by transverse constrictions (Entada scandens). Seeds
exalbuminous.

THE DICOTYLEDONS 539

There are three natural subdivisions of the Leguminosae: (1)
The Mimosacee or Mimosa alliance is characterized by possessing
regular flowers, often in capitula, the petals of which do not
overlap or are they overlapped, and by often furnishing mucilage.
To this group belong the plants yielding Acacia and Catechu.
(2) The Cesalpinacee which have slightly irregular flowers, the
petals of which are imbricate in aestivation, the odd posterior
petal in the flower being overlapped by the 2 side petals. To this
group belong the plants yielding Copaiba, Heematoxylon, Senna,
Cassia Fistula, Brazil Wood, Peach Wood, Sappan and Tam-
arindus. (3) The Papilionacee which have very irregular flowers,
the petals of which are imbricate in aestivation, but the odd petal
here called the standard, overlaps the 2 side petals and the two
side petals overlap the keel petals. To this group belong the
plants yielding Tolu, Balsamum Peruvianum, Baptisia, Glycyr-
rhiza, Goa Powder, Foenugreek, Galega, Melilotus, Trifolium,
Tragacantha, Tonka, Santalum Rubrum, Scoparius, Physo-
stigma, etc. _ }

OrrFiciAL Druc PART UsepD BoTANICAL ORIGIN HABITAT
Acacia Gummy exudation Acacia Senegal and Africa
some other African
species of Acacia
Cascara Bark Sweetia S. Mexico to
Amarga panamensis Panama
Tragacantha Gummy exudation Astragalus gummifer Asia
and other Asiatic
species
Balsamum Balsam Toluifera Pereirze Central America
Peruvianum (Myroxylon Pereirz)
Balsamum Balsam Toluifera Northern South
Tolutanum Balsamum America
Stantalum Heartwood Pterocarpus Indo-China and
Rubrum santalinus S. India
Glycyrrhiza glabra
var. typica
Glycyrrhiza glabra S. Europe, W. and
Glycyrrhiza Rhizome and var. glandulifera C. Asia
roots and other var’s. of
G. glabra yielding
a yellow and sweet wood
Cassia Senna pane te
Senna Leaflets Soudan

Cassia angustifolia

Arabia and Africa

540

OrrFiciAL Druc
Copaiba
Chrysarobinum
Kino

Trifolium
Krameria

Physostigminz
Salicylas

PHARMACEUTICAL BOTANY

Part Usep
Oleoresin

Neutral principle

Inspissated juice

Inflorescence

Root

Salicylate of alka-

loid from seed

UnorrFiciAL Druc

Baptisia
Hzmatoxylon

‘Tamarindus
Cassia Fistula

Scoparius
Galega

Sappan
Brazil Wood

Peachwood
Derris

(Tuba Root)
Cube Root

Physostigma
Melilotus

Mesquite Gum
Fenugreek

Piscidia
Indigo
Soy Bean
Abrus
(Jequirity)
Catechu
Mucuna
(Cowhage)

Root

Heartwood

Pulp of fruit
Fruit

Tops
Flowering herb

Heartwood
Heartwood

Heartwood

Rhizome and root

Root
Seed

Leaf and flower-_

ing top
Gum
Seed

Bark
Coloring matter
Seed
Seed

Extract
Hairs from fruits

BoTrANICAL ORIGIN HABITAT

Copaiba species South America
Vouacapoua Araroba Brazil
Pterocarpus India
Marsupium
Trifolium pratense Europe
Krameria triandra Peru and Bolivia
Krameria argentea Brazil
Physostigma veneno- Africa

sum

Eastern North
America

Baptisia tinctoria

Hzmatoxylon Central America
campechianum

Tamarindus indica ‘Tropical Africa

Cathartocarpus India
Fistula

Cytisus Scoparius Europe

- Galega officinalis S. Europe and W.
Asia

Caesalpinia Sappan India, Malaya

Caesalpinia South America
brasiliense
Caesalpinia echinata South America
Derris elliptica Malaya
Derris malaccensis -. Indies
Lonchocarpus Spp. South America
Physostigma Africa
venenosum
Melilotus officinalis Europe
Prosopis juliflora Mexico
Trigonella Mediterranean
Foenum-greecum region
Piscidia Erythrina West Indies
Indigofera tinctoria India
Soja max China and Japan
Abrus precatorius Tropics and sub-
tropics
Acacia Catechu India
Mucuna pruriens East and West
Indies

ORDER GERANIALES OR GRUINALES

GERANIACEZ OR GERANIUM F AMILY.—Herbaceous, rarely
semi-succulent, sub-shrubby plants. Stems cylindrical, often
hairy or glandular hairy. Leaves alternate to opposite, stipulate;

THE DICOTYLEDONS 541

venation from pinnate to palmate, so leaf shape from ovate to
pinnatifid to pinnatipartite to sub-palmatifid to palmatipartite to
compound palmate. Inflorescence either a dichasial or scorpioid
cyme. Flowers regular, pentamerous (Geranium) to irregular
pentamerous (Pelargonium); sepals five, aposepalous; petals five,

AR

Mi}
i

!

Fic. 409.—Physostigma venenosum—Portion of plant and fruit. (Sayre.)

apopetalous, varying in color from greenish-white or pink-red to
scarlet, scarlet-crimson to crimson-purple; stamens with anthers
ten or five, hypogynous or inserted into slightly developed
hypogynous disk; pistil pentacarpellary, ovary five-celled with
two, rarely one ovule in each cell, styles elongate, fused round a
stylar column of receptacle then continued as a stylar tip which
splits into five stigmatic surfaces. Fruit a regma, rarely a simple
capsule. Regma splits into five recurved carpels, each then
dehiscing to set free two or one seeds. Seeds exalbuminous.

542 PHARMACEUTICAL BO TANY

UnorrFiciAL DruG Part Usep BorAnicAL NAME HAsirAtT

Geranium Rhizome Geranium maculatum North America
Pelargonium
odoratissimum
Oil of Rose i z Pelargonium Radula Mediterranean
: Volatile oil z a
Geranium Pelargonium regions
capitatum

LinACEZ OR FLAx Famity.—Herbs with slender stems and
alternate, simple, narrow leaves.
Inflorescence cymose with regu- €3 74 Te!
lar pentamerous flowers; pistil “& Yi"
five-carpelled with a five-celled »
ovary containing two ovules in
each cavity and having a single

meee I

Fic. 410. Fic. 411.
Fic. 410.—Cranesbill (Geranium maculatum), a perennial herb with palmately-
parted leaves, pale purple or rose-purple flowers, a tuberculated rhizome and regma
fruits. The latter, appearing like the bill of a crane, are seen to arise from slender

stalks forming a fork near the center of the picture. (Reproduced from U.S. Dept.
Agriculture Misc. Publ. 77.)

Fic. 411.—ZLinum usitatissimum, an ainvittnk herb with linear-lanceolate leaves
pale blue flowers and capsular fruits, the source of Flaxseed, Linseed oil and Linen

fiber. (Sapre.)

style with a knob-like stigma. While the flower is still in bud
condition or soon after, there commences an ingrowth of the
mid-rib of each carpel which proceeds until, when plant is in

THE DICOTYLEDONS 543

fruit, there are formed 10 cavities each enclosing aseed. Seeds,
anatropous, mucilaginous, flattened, containing a large embryo
and small enveloping endosperm.

OrrFiciAL DruGc PART Usep BotANnIcAL NAME Hasirat
Linum Seeds Linum usitatissimum pane Cult. in
Oleum Lini Fixed oil Linum usitatissimum )'Temperate regions

ERYTHROXYLACEZ OR Coca Famity.—Shrubs (Erythroxylon)
or trees with alternate, simple, entire, glabrous and pinnately

Fic. 41 2.—Guajacum sanctum—F lowering branch. (Sayre.)

veined leaves, the cells of the lower epidermis of which show
papilla. Flowers regular, hermaphroditic, each with five
sepals, five hypogynous petals, ten stamens and a two- to three-
celled ovary subtending three styles, each with a capitate
stigma. Fruit an ovoid, angular, one celled drupe containing
a single seed.

Pd rae eR Pe A

Kee Pee

544 PHARMACEUTICAL BOTANY

OrriciAL DruG Part Usep BoTANICAL ORIGIN HABITAT
Cocaina Alkaloid Erythroxylon Coca }/ Peru and Bolivia,
Cocainze Hydrochloride of js other species of South America
Hydrochloridum alkaloid Erythroxylon

UnorFictiAL DruG
Coca Leaves Erythroxylon Coca Peru and Bolivia
and E. truxillense

ZYGOPHYLLACEZ OR CALTRoP Famity.—Herbs, shrubs, or
trees (Guajacum or Guaiac) having jointed, often divaricate

Fic. 413.—Citrus Aurantium, the bitter or Curacao orange, a small tree bearing
evergreen leaves with winged petioles, white flowers and a spherical fruit with

bitter, acid pulp. The source of bitter orange peel and volatile oils of the peel and
flowers. (Sayre.)

branches. Leaves usually opposite, stipulate and compound.

Flowers regular or irregular, pentamerous, white, yellow, red or
blue (G. officinale). Fruit a capsule.

OrrictAL Druc PART Usrep

BoTANICAL ORIGIN HABITAT
Guaiacum Resin of wood aie officinale oe Indies and S.
America

Guajacum sanctum

THE DICOTYLEDONS 545

UnoFFICIAL DruG Part Usep BOTANICAL ORIGIN HABITAT
Guaiaci Lignum —_ Heartwood G. officinale and G. ie Indies and S.
sanctum America

RutTAceE& oR Rue Famity.—Herbs (Rue), shrubs (Zaborandi
and Buchu) or trees (Orange, Lemon, Grape Fruit, etc.). Stems
upright, often wiry and xerophytic in the Rue sub-family or
Rutee, elongated and spiny in the
Prickly Ash sub-family or Zanthoxy-
lee, woody and green in the Citrus
sub-family or Aurantiee. Leaves
alternate or opposite, simple (Buchu),
rarely whorled (Pilocarpus) or pin-
natifid, as in the Rue plant (Ruta
graveolens), or pinnate, as lower parts
of Ruta graveolens, becoming reduced °
pinnate in the Orange (Citrus
Aurantium). Leaves exstipulate or
with spiny stipules (anthoxylum).
Stems and leaves abound in more
or less sunken glands. Flowers
pentamerous, varying in color from
yellow in Ruta to white in Citrus to
pink (Barosma betulina) or pink crim-
son, as in some Barosma and Diosma
species, rarely to pinkish-purple
(Pilocarpus); sepals five, aposepalous,
becoming in Cztrus more or less Fic. 414.—Barosma betulina, the
synsepalous; petals five, apopetalous, yoni ie agape . Ls
becoming more or less synpetalous (¢,,,,.)
and tubular (Correa grandiflora);
stamens five, simple or with expanded bases, lobed, or lobes
developed as staminal stipules and more or less split (Citrus);
pistil of ten, five, three or two carpels, ovary as many-celled,
with frequently 2 to 1 ovules in each cell. Fruit a capsule
(Dittany, Zanthoxylum), a hesperidium (Citrus) or rarely a samara,
as in the Wafer Ash (Ptelea). Seeds albuminous or exalbuminous.
Many of the plants contain volatile oils in their secretory

cavities.

546
OrriciAL DruG
Aurantii Dulcis

Cortex
Oleum Aurantii

Aurantii Amari
Cortex

Limonis Cortex
Oleum Limonis
Pilocarpine Nitras
Pilocarpine
Hydrochloridum
Buchu

Xanthoxylum
(Prickly Ash)

Oleum Bergamottz

Oleum Aurantii
Florum

Oleum Aurantii
Amari

Xanthoxyli Fructus

UnorriciAL Druc
Ruta (Rue)

Ptelea (Wafer Ash)
Belz Fructus

Pilocarpus

Cusparia Bark

PHARMACEUTICAL BOTANY

Part Usep
Fresh outer rind
of ripe fruit
Vol. oil from
fresh peel of
ripe fruit
Rind of fruit

Outer rind of
ripe fruit

Vol. oil from
fresh fruit peel
Nitrate of alka-
loid from leaflets
Hydrochloride of
alkaloid from
leaflets

Leaves
Bark

Volatile oil from
fresh fruit rind
Volatile oil from
fresh flowers
Volatile oil from
fresh peel

Capsules

Leaves
Bark of root
Unripe fruit

Leaflets

Bark

BOTANICAL ORIGIN
Citrus Aurantium var.

sinensis

Citrus Aurantium var.

sinensis

Citrus Aurantium

Citrus medica var.

Limon

Citrus medica var.

Limon

Pilocarpus Jaborandi

Pilocarpus
microphyllus

Barosma betulina,

B. crenulata and
B. serratifolia
Zanthoxylum
americanum
Zanthoxylum
Clava-Herculis
Citrus Aurantium
var. Bergamia
Citrus Aurantium

Citrus Aurantium

Zanthoxylum
americanum
Zanthoxylum
Clava-Herculis

Ruta graveolens
Peltea trifoliata
Egle Marmelos

Pilocarpus Jaborandi

Pilocarpus
microphyllus
Galipea officinalis

HABITAT
Northern India

Northern India

Northern India
Northern India

Northern India

Brazil

S. Africa

Northern United
States

Southern United
States

France, Italy

Northern India

Northern India

Northern United
States
Southern United
States

Southern Europe
North America
Malabar, India

Brazil

Venezuela

SIMARUBACEZ or AILANTHUS Famity.—A family of chiefly

tropical shrubs or trees containing bitter
are mostly alternate and pinnate.

polygamous and arranged in axilla
or in terminal racemes (Quassia a

principles.
The flowers are unisexual or
ry panicles (Picrasma excelsa)
mara). -The fruit is a drupe

The leaves

THE DICOTYLEDONS 547

~
—- -_.

\\ SS = SN
SF <=
Sd ONN\\\ Wir
at SA : —
=~ oi Pa \

Fic. 415.—Branch of a Quassia yielding plant, Picrasma excelsa. Note the imparipi-
mate leaves with winged petioles and axillary flower clusters. (After Sayre.)

(Picrasma) or pod (Quassia). The plants are distinguished from
those of the Rutacee by the absence of secretory cavities.

OFFICIAL DruGc Part Usep BOTANICAL ORIGIN HAsIrat
: Picrasma excelsa West Indies
Quassia Wood 1 ; :
Quassia amara Surinam

UnorriciaL Druc

Si Bark of Simaruba officinalis South America
weiner cn vce -Ssseatne amara Central America
Bahamas and

Florida

BuRSERACE2 OR Myrru Famity.—Shrubs and trees of
tropical climes having secretion reservoirs in their bark which

PO RE ee ey ee ae eae Sara pesee

548 PHARMACEUTICAL BOTANY

contain gum-resin or oil. Leaves alternate, exstipulate and
compound (trifoliate in Commiphora Myrrha). Flowers small,

regular and hermaphrodite,
arranged in racemes or pan-
icles. Fruit a drupe.

OrrFicIAL DruG Part Usep
Myrrha Gum resin
BorAnicAL ORIGIN HABITAT

Commiphora N. E. Africa and

abyssinica et al.spp. Arabia
UnorriciAL Druc Part UseED
Olibanum Gum resin
BoTANICAL ORIGIN HABITAT
Boswellia carterii East Africa and
Arabia

MELIACEZ OR MAHOGANY
Famity.—Iropical trees or
shrubs with wood often hard,
colored and odoriferous.
Leaves alternate, exstipulate,
pinnately-compound, rarely
simple and entire. Inflores-
cence a terminal or axillary
raceme. Flowers hermaphro-
dite or rarely polygamo-dice-
cious, regular; sepals five to
four, small; petals usually five
to four, hypogynous; stamens
generally ten to eight, rarely
five, very rarely twenty to sixteen, inserted outside the base of the
hypogynous disc; filaments united into a tube; carpels usually
five hate three; style simple; ovary free, usually five- to three-celled.
Fruit a drupe (Melia), berry (Vavaea), or capsule (Cedrela). Seeds
exalbuminous or with fleshy albumen.

Fic. 416.—Commiphora myrrha—Branch.
(Sayre.)

UnorFIcIAL Druc Part Usep BoTANICAL ORIGIN HABITAT
Aen Bark of root Melia Azedarach Asia
illana Bark Guarea Rusbyi Bolivia

PoLYGALACE® or Mirkwort Famity.—Herbs or shrubs
with upright, herbaceous to woody stems often branching pro-

THE DICOTYLEDONS 549

fusely, the branches occasionally becoming geotropic or sub-
terranean and bearing cleistogamous flowers. Leaves simple,
often lanceolate or linear, exstipulate, alternate. Inflorescence
a raceme, spike (Polygala Senega) or head (P. lutea). Flowers

‘TD

Fic. 417.—Polygala senega—Plant and rhizome. (Sapre.)

irregular, hermaphroditic with 5 distinct sepals, the 2 lateral
ones being large and petaloid, 5 petals of which the two lateral -
are wanting or rudimentary and the anterior large and boat-
shaped, eight stamens, and a bicarpellate pistil. Fruit a two-
celled capsule (P. Senega), rarely a drupe or samara. Pollen
grains barrel-shaped.

550 PHARMACEUTICAL BOTANY

OrriciAL DruG PART UsEep BOTANICAL ORIGIN HABITAT
Senega Root Polygala Senega United States and
Canada

EupHorBIACEZ OR SpurGE Famity.—Often herbaceous,
more rarely shrubby, rather seldom arborescent plants. Stem,
leaves and other parts in several genera traversed by latex canals

=.

a

end

Fic. 418.—The Castor Oil plant (Ricinius communis). (From Strasburger after Baillon.)

that are either ramifying cells (Euphorbia) or laticiferous vessels
(Manihot, Hevea, etc.) or rows of laticiferous sacs (Micrandra)
and contain a white latex with acrid often poisonous contents
or alkaloid or hydrocarbon, at times, rubber contents. Leaves
alternate, exstipulate to stipulate, simple to pinnate or palmate.
Inflorescence cymose. Flowers usually, as in Ricinus, etc.,

THE DICOTYLEDONS 551

pentamerous, moncecious; sepals five, green, aposepalous,
becoming rudimentary or absent in Euphorbia. Petals none or
five more or less petaloid; stamens numerous to ten to five or one
(Euphorbia); pistil in pistillate flowers rarely of twenty to ten
apocarpous or loosely syncarpous carpels (Sandbox tree), com-
monly of three syncarpous carpels with distinct radiate styles;
ovary as many-celled as carpels with two to one ovule in each
cell. Fruit a tricoccoid regma or capsule, rarely winged,
indehiscent, nutlike. Seeds with oily endosperm. Flowers at
times surrounded and subtended by more or less petaloid and
expanded bracts and bracteoles.

OrrFictAL Drauc Part Usep BOTANICAL ORIGIN HABITAT

Euphorbia Herb Euphorbia pilulifera India
‘ Pilulifera

Stillingia Root Stillingia sylvatica Southern United

: States

Kamala Hairs of capsule = Mallotus philip- Asia

pinensis
Oleum Ricini Fixed oil Ricinus communis _ India
Oleum Tiglii Fixed oil Croton Tiglium Asia
UNOFFICIAL
Druc

Tapioca Starch Manihot utilissima South America
Elastica Prepared latex Hevea braziliensis —_ Brazil
(Rubber) and other species
Cascarilla Bark Croton Eluteria West Indies
Euphorbium Resinous latex Euphorbia resinifera Morocco

Tung Oil Fixed oil Aleurites cordata China

OrDER SAPINDALES

ANACARDIACEZ OR SUMAC Famity.—Shrubs or trees pro-
ducing in stems and leaves secretion contents that are either
acrid-watery or acrid-opalescent or white-viscid, viscid-acrid
and poisonous. Leaves alternate, rarely opposite, simple (Rhus
Cotinus), three foliate (Rhus toxicodendron) or pinnate (Rhus
glabra, R. venenata, etc.). Inflorescence frequently terminal and
composed of racemes of cymes, often reduced to a simple raceme.
Flowers small, clustered, green, greenish-white to greenish-
yellow; sepals five, rarely six or four green, small; petals five,
smaller than sepals; stamens equal in number to the petals and
alternate, rarely fewer, sometimes double in number, rarely

552 _ PHARMACEUTICAL BOTANY

indefinite, inserted hypogynously or upon an enlarged disc that
surrounds or swells up between stamens and pistil; pistil mono-
carpellary (Rhus), more rarely bicarpellary, very rarely, as in
Spondiea, of ten to five carpels, ovary one-celled with single
ovule. Fruit a drupe. Seeds exalbuminous with large embryo
filling seed cavity.

OrrFictAL DruG PART Usep BoTANICAL ORIGIN HABITAT

Mastiche Concrete resinous Pistacia Lentiscus Grecian Archi-
exudate pelago
UnorriciAL Druc
Chinese Galls Excrescences Rhus semialata China
Japanese Galls Excrescences Rhus japonica Japan
Rhus Glabra Ripe fruits Rhus glabra - Canada and United
States
Rhus Typhina Ripe fruits Rhus typhina United States
Acajou Gum Gum Anacardium West Indies
occidentale
Pistachio Seeds Pistacia vera Western Asia
Anacardium Fruit Anacardium West Indies
occidentale
Rhus Fresh leaflets Rhus toxicodendron United States
Toxicodendron

AguIroLiace# (ILicacE#) or Horry Famity.—Trees or
shrubs, chiefly evergreen; leaves alternate, petiolate, simple with
entire to spiny-toothed margin (J/ex opaca or American Holly).
Flowers regular, small, greenish to white, hermaphrodite or
polygamous or dioecious. Fruit drupaceous and containing 4
to 6 nutlets. Caffeine has been obtained from Maté and
Cassena.

UnorriciAL Druc Part Usep BoraAnicAL NAME HABITAT
Maté Leaves Ilex paraguariensis South America
Cassine (Cassena) Leaves Ilex vomitoria Southeastern

United States
Black Alder (Prinos) Bark Ilex verticillata North America

CELASTRACE@ OR Starr TREE Famity.—Shrubs (Evonymus
or Burning Bush), or shrubby climbers (Céelastrus or Climbing
Bittersweet). Leaves rarely alternate (Celastrus), usually oppo-
site (Evonymus, Pachystima), simple, entire or toothed. Inflores-
cence of axillary cymes of terminal racemes. Flowers perfect
(Evonymus, Pachystima) or polygamo-dicecious (Celastrus), greenish
(Celastrus), greenish or yellowish-white (Evonymus Europeus),

THE DICOTYLEDONS 553

egreenish-purple (Evonymus americanus) to dark purple (Evonymus
atropurpureus); calyx four- to five-lobed; corolla of four to five
petals; stamens four to five, perigynous, on a disc, which fills the
base of the calyx and sometimes covers the ovary; ovary three- to
five-celled. Fruit a two- to five-celled capsule. Seeds albumi-

Fic. 419.—The Burning Bush or Wahoo, Evonymus atropurpureus. Flowering
branch to left; fruiting branch to right.

nous with fleshy reddish aril (Evonymus, Celastrus) or white mem-
branous aril (Pachystima).

OrriciAL Druc Part Usep BoTANIcAL ORIGIN HABITAT
Euonymus Bark of root Evonymus United States
atropurpureus

SAPINDACE OR SOAPWORT FamiLy.—Trees (Sapindus), shrubs
(Paullinia), undershrubs or rarely herbaceous vines of tropical

554 PHARMACEUTICAL BOTANY

climes (Cardiospermum) containing the glucoside saponin. Stem
erect or climbing (Paullinia) often provided with tendrils.
Leaves commonly alternate and compound. Flowers in racemes
or panicles (Paullinia) or corymbs (Cardiospermum), perfect or
polygamodicecious, yellowish in Paullinia Cupana. Fruit a
poe: | capsule (P. Cupana), samara,
7 os + drupe or berry (Sapindus). Seeds
exalbuminous.
OrriciAL Druc PART UsEpD
Guarana Dried Paste, made
_ from crushed seed

BoTANICAL ORIGIN Hapsirat
Paullinia Cupana Brazil

ACERACE& OR MAPLE
Famity.—Chiefly trees, occa-
sionally shrubs, of temperate
regions with watery sap. Leaves
opposite, simple and palmately
lobed or cleft (Acer) or pinnate
: (Negundo). Inflorescence a

Fic. sone Te da raceme, condensing in some
(Rhamnus purshiana). Leaf and fruiting Species to a capitulum of cymes.

branch. The cured bark constitutes Flowers small, regular, poly-
a valuable tonic-laxative. (Reproduced amous or diccci : if
from U. S. Dept. Agric. Misc. Publ. 77.) ® cious; sepals five

to four, green; petals none or five,
variously colored; stamens usually eight, hypogynous_ or
perigynous; nectar disc around stamens or between them and
pistil; pistil bicarpellary with two-celled ovary. Fruit a samara.
Seeds green, exalbuminous with coiled or folded embryo and long
thin cotyledons.

UnorrictAL DruG PART Usep BoranicAL Oricin

: HABITAT
Acer Spicatum Bark Acer spicatum

United States

ORDER RHAMNALES

RHAMNACEA OR BUCKTHORN Fami_y.—Shrubs or low trees
usually of branching or spreading habit. Branches either
cylindric, long, green or hardened, checked back and spines-
cent, occasionally, especially flowering branches, developing

THE DICOTYLEDONS 555

tendrils for support. Leaves simple, usually alternate and
stipulate. Inflorescence cymose. Flowers hermaphrodite or
more or less diclinous, pentamerous to tetramerous, greenish to
greenish-yellow to yellowish-white; sepals five to four; petals
five to four alternating with sepals; stamens five opposite the

Fic. 421.—Alder Buckthorn or Rhamnus Frangula—Branch. (Sayre.)

petals, perigynous; pistil either free in center of receptacular
cup or more or less fused with it and so semi-inferior, ovary
typically three-celled, becoming rarely four-celled with two to
one atropous ovules in each cavity. Fruit of three indehiscent
cocci, each enclosing a single albuminous seed with straight
embryo imbedded in albumen.

556 PHARMACEUTICAL BOTANY

OrriciAL DruGc Part Usep __ BorANICAL ORIGIN HABITAT
Cascara Sagrada Bark Rhamnus Purshiana Northern California,
Oregon, Washington,
and southwestern Brit-
ish America

Frangula Bark Rhamnus Frangula Europe

Rhamnus Cathartica Fruit Rhamnus cathartica Asia and Africa
UnorricIAL Drue ,

Ceanothus Root Ceanothus americanus North America

Zizyphus Fruit Zizyphus sativa Mediterranean region

Vitace& orn Grape Famiry.—Rarely tall, herbaceous,
usually shrubby and climbing, more rarely shrubby, upright

Fic, 422.—Northern Fox Grape (Vitis Labrusca). 1, portion of twig, with leaf,
inflorescence and tendril, about natural size; 2, median longitudinal section of a ae
fect flower, enlarged; 3, perfect flower (enlarged) with petals removed; 4, longi-
tudinal section of fruit (berry); 5, seeds, natural size and enlarged. (7-4 afi Gray;
5, after Engelmann, from C. E. Bessey, Nebraska Hort. Soc. Ann. Rept. 1895.) (Gager.) ;

THE DICOTYLEDONS 557

plants. Stems rarely short, more usually elongate, feeble,
rather brittle, climbing by tendrils which represent modified
inflorescence shoots. Leaves alternate, simple to lobed (either
pinnately or more often palmately) to compound-pinnate or
palmate. Perfect graded series of lanceolate leaves with pinnate
veining to palmate veining; from pinnately veined to compound-
pinnate; from palmately veined to compound-palmate. Stipules
greenish to membranous or none. Flowers in racemes of com-
pressed cymes, hermaphrodite or diclinous, nearly always small,
clustered, green to greenish-yellow or greenish-white, rarely
otherwise; sepals five, rarely four, small to minute (mere rim
of receptacle) more or less persistent. Petals five, deciduous to
caducous, typically distinct (in Vitis), united by their tips into
calyptroform corolla, so in June, as Grape Vine flowers expand,
corolla splits at base into five lobes that separate below, being
attached at tips, while whole becomes tumbled off by lengthen-
ing stamens. Pistil bicarpellate. Ovary two-celled, superior
or at most sub-inferior. Ovules two to one, erect. Style short,
often more or less thickened, with terminal, capitate, slightly
two-lobed stigmas. Stamens equal to petals or sepals and
opposite petals. Receptacle internal to stamens, often expanded
into nectariferous girdle or, in Vitis, into receptacular knobs
alternating with stamens. Fruit a berry rarely six- to three-
celled, typically two-celled and with two to one seeds in each
cavity. Seeds like ovules, erect with bony testa. Embryo
small, imbedded at base of cartilaginous albumen.

Brandy or Spiritus Vini Vitis is an alcoholic liquid obtained by the distillation

of the fermented juice of sound, ripe grapes and containing from 48 to 54 per cent.
of ethyl alcohol.

OrpDER MALVALES

STERCULIACEZ OR CoLa Famity.—Rarely herbs, usually
shrubs or tall, often heavy trees with soft wood and broad annual
rings. The cambium, in developing bast, produces one, two,
three, four, or five alternating layers of hard and soft bast which
in some species of this as well as the Tiliacee or Linden family
form long finger-like processes pushing out into the cortex.
Leaves alternate, sometimes simple and pinnately-veined or
passing to palmately-veined or palmately-compound. Flowers

558 PHARMACEUTICAL BOTANY

hermaphrodite; sepals five, sometimes surrounded by bracteoles
forming an epicalyx; petals usually five; stamens typically five,
hypogynous, opposite petals, distinct or slightly fused in mon-

“A \W

Fic. 423.—Theobroma Cacas, the Chocolate plant. 1. Trunk and branches
bearing fruit. 2. Branch bearing flowers. 3. Single flower. 4. Staminal ring.
5. Stamen. (After Strasburger, Noll, Schenck and Schimper.)

adelphous fashion (Melochia, Waltheria) or, stamens subdivided
above into few or numerous staminal leaflets, anthers two-celled;
pistil many- to ten- to five- or four-carpelled; carpels apocarpous
or more usually partially or completely united. Fruit either

ee ee ge eee ae ee ee ae 7

THE DICOTYLEDONS 559

follicles, or fused to form a capsule of ten or more, frequently five
dehiscent carpels or, carpels splitting asunder into cocci or,
becoming a woody capsular nut (Theobroma) or, rarely the fruit
may become succulent. Seeds globose or subglobose and often
provided with wings, arils or similar appendages; embryo
straight, large and surrounded by scanty albumen.

OrrictAt Druc Part Usep _ BorAnicAt ORIGIN HAsiraT
Oleum Fixed oil Theobroma Cacao ‘Tropical America
Theobromatis
Cacao Prepara- Prepared powder from Theobroma Cacao Tropical America
tum roasted, cured kernels and other species

of seeds
Kola Cotyledons Cola nitida and Africa

other species

TILIACE OR LinDEN FAmiLy.—Shrubs or trees, rarely herbs,
having stellate hairs on both _
stems and leaves. Leaves alter-
nate, pinnately- more rarely
palmately-veined, stipulate.
Inflorescence cymose. Flowers
hermaphrodite, more rarely, by
absorption, more or less dicli-
nous; sepals and petals five each,
more rarely four, sepals decidu-
ous; stamens five opposite the
petals or, as in Sterculiacea, five
phalanges of stamens represent-
ing subdivided stamens (Tz/ia),
pistil of ten to five or two syn-
carpous carpels; ovary superior.

Fruit either a nut-like drupe or Fic. 424.—Tilia americana, The
American Linden or Basswood. Leaf
and flowering branch. The leafy,

drupe, rarely baccate.

UnorrictaL Druc Part Usep tongue-shaped bract and simple cyme
Tilia (Linden Inflorescence springing from near the center of its
flowers) ; midrib constitutes the drug ‘‘Linden

BorAnicAL ORIGIN HABITAT Flowers.”’ (Reproduced from U. S. Dept.
Tilia species United Statesand 44, Misc. Pub. 77.)

Europe

Ma.tvace& or Martiow Famity.—Herbs in temperate
regions (as the Mallow, Malva rotundifolia, and Marshmallow,

560 PHARMACEUTICAL BOTANY

Althea officinalis, etc.), occasionally shrubs in temperate regions
(Hibiscus Syriacus or Rose of Sharon, etc.), frequently shrubs or
tall trees in the tropics. Stems, as in Sterculiacee and Tiliacea,
sometimes forming numerous layers of hard and soft bast.
Leaves alternate and stipulate, ovate, ovate-cordate, orbicular
or palmately-compound; venation pinnate or palmate. Stems,
roots and leaves contain mucilage cells. Inflorescence a raceme

locyles7~
with ovules

Fic. 425.—Median lengthwise section through flower of American Upland Cotton
(Gossypium hirsutum) X 2. (After Robbins.)

or fascicle of cymes. Flowers regular, pentamerous; calyx
green, of five aposepalous sepals but frequently surrounded
outside by an epicalyx. Both calyx and epicalyx are persistent.
Corolla of five petals, varying in color, which are more or less
fused with stamens at their bases, stamens monadelphous and
forming an upright column (staminal column) enclosing the styles;
anthers one-celled, dehiscing transversely; pollen grains echinate;
pistil loosely or strongly syncarpous, rarely sub-apocarpous of
thirty to five carpels. Fruit either a circular set of cocci (Malva),

THE DICOTYLEDONS

561

Fic. 426.—The Asiatic- or Levant Upland Cotton, Gossypium herbaceam—Branch.

(Sayre.)

follicles, or a capsule (Gossypium or Cotton). Seeds albuminous
with oily and mucilaginous albumen.

OrriciAL Druc Part Usep

Althza Root (peeled)

Gossypii Radicis Bark of root
Cortex (recently)
gathered)

Gossypium
Purificatum Hairs of seed

BoTANICAL ORIGIN
Althzea officinalis
Cultivated varieties of:
Gossypium
herbaceum
Gossypium
Barbadense
Gossypium
hirsutum
Cultivated varieties
of Gossypium
herbaceum

HABITAT
Europe and Asia

Arabia, United
States, East Indies
West Indies, Fgypt,
and India

Arabia, East
Indies, United
States, and India

.

562 - PHARMACEUTICAL BOTANY
OrriciAL Druc —_—s—~PArT:-~Usep BOTANICAL ORIGIN HABITAT
Cultivated varieties of aed Stic
Oleum Gossypii Fixed oil from G. herbaceum and re ae
yee ia, Africa and
Seminis Seeds other Gossypium We Eadie
: species
UnorriciAL Druc
Malve Folia wires Wet sylvestris | Burope
‘ Malva rotundifolia
Althza Folia Leaves ' Althzea officinalis Europe and Asia
Althza Flores Flowers” i Althza rosea Europe
(larger mallow (dark colored)
flowers) <o
Malvz Flores Wovens ae sylvestris ae
(mallow flowers) Malva rotundifolia 1

Ambrette Seed Seed Hibiscus Abelmoschus India

4

Fic. 427.—Upland cotton (Gossypium hirsutum). A, mature boll opened out; B,
Cross-section of young boll; C, single seed with hairs; D, young boll. (Rodbdins.)

ORDER PARIETALES

This order includes a large variety of plants most of which
have parietal placentation, separate sepals and numerous
stamens. Belonging to this order are the Violets, Begonias,

St. John’s Worts, Damiana, Passion Flowers, Rock Roses, Tea
and Camellias, Canella, etc.

THE DICOTYLEDONS 563

THEACEH OR TEA Famiry (TERNSTR@MIACE#, CAMMELI-
AcEa:).—Evergreen shrubs or low branching, or tall, often heavy
trees with watery juice. Leaves for the most part alternate,
evergreen, often leathery, sometimes membranous; stipules
either bud scales and caducous or often absent; leaf margins
sinuate or serrate (Thea). Inflorescence a raceme becoming by
condensation terminal, one-flowered. Flowers regular, perfect,
pentamerous; sepals five, rarely four to three, deciduous,
occasionally subtended by bracteolar scales; petals five, brittle
and succulent, varying from greenish-white or greenish-yellow
through yellow to white or, whitish pink to pink, scarlet, crimson,

very rarely a tendency toward purple; stamens typically five
but, as they grow, they subdivide into staminal leaflets, so that
in their mature condition they are apparently indefinite and
mono- to polyadelphous; stamens inserted hypogynously or
perigynously and opposite the petals; pistil of typically five
syncarpous carpels but reduced in some species to four to three
or two. Fruit usually a capsule (Thea), five- to three-celled,
dehiscing longitudinally, more rarely a fleshy, semi-baccate,
semi-drupaceous, indehiscent fruit. Seeds with scanty or no
albumen and often attached to the inner angle of cells by project-
_ Ing spongy placentz.

OrriciAL Druc Part Usep BoTANICAL ORIGIN HABITAT
Caffeina Feebly basic Thea sinensis Eastern Asia
substance varieties, etc.
UnorriciAL Druc
Thea (Tea) Leaves Thea sinensis var. _ Eastern Asia

Bohea and viridis

GUTTIFERA OR GAMBOGE Famity.—Tropical trees (Gamboge
and Mangosteen), rarely shrubs, containing resinous principles
in schizogenous resin canals found in cortex and pith. Leaves
Opposite, coriaceous. Flowers dioecious, generally pentamerous
_ or tetramerous with usually five stamens which are subdivided.
_ Fruit a berry (Garcinia Hanburyi), drupe or capsule. Seeds
generally large; embryo large to huge, often with enlarged
radicle and reduced or absorbed cotyledons.

OrrictaL Druc Part Usep BoTANICAL ORIGIN HABirat
aie Cochin

Cambogia Gum resin Garcinia Hanburyi China and Annam

564 PHARMACEUTICAL BOTANY

HyprrIcAce& or St. Joun’s Wort Fami_y.—Herbs or shrubs
of temperate climes with opposite (Hypericum perforatum), rarely
whorled branches and balsamic, resinous juices, which, in the
herbaceous species, are secreted by black or pellucid glands
found in the leaf parenchyma. Leaves entire, opposite, usually

Fic. 428.—Garcinia Hanburyi—Branch of Gamboge plant. (Sayre.)

sessile, exstipulate, and dotted. Flowers, regular, hypogynous
and arranged in panicles or forked cymes. Petals usually 5.
Stamens usually indefinite, rarely definite, often in 3-5 sets,
more rarely monadelphous or free. _Pistil of usually 3-5 carpels
with 3-5 celled, compound ovary and as many filiform styles as
carpels. Fruit a capsule with usually septicidal dehiscence

THE DICOTYLEDONS 565

(Hypericum) or a berry. Seeds small, numerous, anatropous and:
exalbuminous.

UnorriciAL Druc Parr Usrep BoranicaAL OriGIN Hasirat
Hypericum Entire plant Hypericum perforatum Europe

CANELLACE# FamiLy.—Trees the bark of which contains aro-
matic 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.

OrriciAL Druc Part Usep BoTANICAL ORIGIN Hapsirat
Canella Inner bark Canella Winterana era and West
Indies

Brxace& Famity.—Tropical shrubs or trees. Leaves alter-
nate, simple with minute or no stipules. Flowers hermaphrodite
or unisexual, regular; stamens hypogynous, mostly indefinite
with anthers opening by slits, rarely by one or two apical pores
(Bixa). Fruit fleshy or dry. Seeds with fleshy albumen and
sometimes covered with a fleshy arilus (Bixa Orellana).

_UnorrFictAL Druca Part Usep BoTANICAL ORIGIN HABITAT
Annatto Coloring matter __ Bixa Orellana oe America
and Madagascar

FLACOURTIACEZ OR FLacourtiA Famity.—A family of
chiefly tropical trees and shrubs with simple, alternate leaves,
staminate, pistillate and hermaphrodite flowers, frequently
dicecious or polygamous, with a one-celled ovary. The fruits
are either berries (Taraktogenous Kurzti or Leprosy plant), drupes
or rarely capsules. The seeds are albuminous.

Orric1AL Druc Part Usep BoTANICAL ORIGIN Hasirat
Oleum Fixed oil from Taraktogenos Kurzii Burma
Chaulmoogre _ seeds Hydnocarpus

Wightiana India
Hydnocarpus
anthelmintica Cochin China

VIOLACEH OR VIOLET Famity.—Herbs or shrubs. Stems
upright, rarely creeping, spreading or acaulescent. Leaves
either cauline or radical, stipulate, alternate, simple to pinnatifid
or palmate.

566 PHARMACEUTICAL BOTANY

Flowers pentamerous, regular or irregular (Viola) and some-
times cleistogamous; sepals 5; petals 5, unequal, the lowermost
often spurred at the base; stamens 5, hypogynous, with adnate,
introrse anthers and usually closely arranged or coherent
around the ovary, in Violeis the two lower stamens bearing spurs
which extend into the spur of the corolla; ovary, sessile, one-
celled with 3 to 5 parietal placentas; style usually club-shaped.
Fruit a loculicidally dehiscent capsule (Viola), rarely baccate.
“Seeds albuminous.

UnorrictaL Druc Part Usep BotTANIcAL OriGIN HABITAT

Pansy Entire herb Viola tricolor Temperate regions
Viola Odorata Fresh flowers and Viola odorata Europe
root

TURNACE OR Damiana Famity.—Tropical herbs, shrubs or
trees. Leaves alternate, simple, petioled, exstipulate. Flowers
perfect, regular, axillary, pentamerous with one-celled ovary.
Fruit a capsule with three valves. Seeds strophiolate with
albuminous embryo.

OrriciAL DruG PART UsEep BOTANICAL OrIGIN HABITAT

| diffusa ct Lower

Damiana Leaves Turnera California, Mexico, S.

aphrodisjaca America, West Indies

PASSIFLORACEZ OR Passion FLower Famity.—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 crowned by one or more
series of subulate filaments which are frequently colored; gyno-
phore elongating, supporting the stamens and pistil. Fruit a
one-celled berry (Passiflora) or a three- to five-valved, dehiscent
capsule containing numerous seeds.

UnorriciAL Druc Part Usep Boranicay Oricin HaAsirat
Passiflora Flowering and fruiting top Passiflora incarnata’ United States

CaRICACE2 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

THE DICOTYLEDONS 567

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
dicecious flowers. The fruit is a berry, the size of one’s head and
contains an acrid milky juice
from which papain can be pre- ~
cipitated by the addition of —
alcohol.

CisTACE2 OR Rock Rose
Famity.—Herbs or shrubs whose
stem and branches are often
glandular, pubescent or tomen-
tose, with simple or stellate tri-
chomes. 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; Fic. 429.—Passion Flower or May-
petals five, as in Helianthemum or PoP (Passiflora incarnata), a climbing,
Rock R PER ial 1 perennial herb used as a motor de-
se a re pvc bet cediolans & pressant in neuralgia, etc. Note the
three or none, as in Lechea or palmately 3-cleft leaves and axillary
Pinweeds; stamens hypogynous, tendrils. The flowers possess a lavender
: : ink and 1 .
indefinite; carpels three to five, “ P™ pacts tan soiree pe
ovary free, one-celled. Fruit a one-celled, three- to five-valved

capsule.

2
ee |

UnorriciAL Druc Part UsEep BoTANICAL ORIGIN HABITAT
Helianthemum Herb Helianthemum canadense Eastern United States

OrDER OPUNTIALES.—CACTACEZ OR Cactus FAMILY.—
Herbaceous, rarely arborescent (Cereus giganteus or Giant
Cactus), more or less succulent plants living in warm, dry
(Pereskia), usually desert situations, rarely becoming epiphytic
and correspondingly modified. Stems, accordingly, varying
from elongate, slightly enlarged, green (Pereskia), to flattened
and ribbed (Selenicereus), to flattened and shortened as in the
Prickly Pear (Opuntia), to condensed (Echinocactus, Echinocereus,

568 PHARMACEUTICAL BOTANY

etc.), to greatly condensed (Mamillaria). Leaves alternate, stip-
ulate or exstipulate, enlarged and more or less fleshy (Pereskia),
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 crim-
son; stamens indefinite, inserted at varying levels in the throat

Fic. 430.—The peyote cactus, Lophophora Williamsii, a perennial herb whose
dried tops yield the drug known as peyote or mescal buttons. This drug has been
used by the Aztec and Kiowa Indians as an intoxicant in their religious ceremonies.
A, side view; B, view from above; #, /?, tubercle; r rib; ¢’, scar of tubercle of older
growth. xX 4.

of a greatly expanded upgrown receptacle; _pistil generally
tricarpellary; ovary inferior, often deeply sunk in upgrown
receptacular part; style thread-like, divided above into as many
stigmas as carpels. Fruit a receptacular berry enclosing numer-
ous small seeds. Seeds exalbuminous.
OrriciAL Druc Parr Usep BOTANICAL ORIGIN
Cactus Fresh succulent
Grandiflorus stem
UnorrictAL Druc
Peyote Crown of plant Lophophora Williamsii Mexico

HaAsitrat

Selenicereus grandiflorust Jamaica

ORDER MyrTALES OR MyrTIFLORA

This order includes trees, shrubs and herbs, most of which
possess tetramerous flowers with an inferior ovary. It contains

THE DICOTYLEDONS 569

in addition to the families here considered the Rhizophoracea or
Mangrove Family, the Onagracee or Evening-Primrose F amily
which is represented by the Oenotheras or Evening Primroses,
the Willow Herbs (Epilobium) and the cultivated Fuchsia, the
Elaagnacea, represented by the Buffalo-Berry, the Trapacee, con-

!
r Hf

Fic. 431.—Daphne Mezereum (Fam. Thymelaacea)—Fruiting branch and flowers.
(Sayre.)

taining the Water-Chestnut or Trapa natans and the Lecythidacee
or Monkey-Pot Family which contains the Brazil-Nut whose
seeds are edible.

THYMELZACEZ OR MEzEREUM Famity.—Shrubs (Daphne
Mezereum) or low trees, usually of branching habit, the stems
developing long tenacious bast fibers. Leaves alternate, rarely

*

Bitch i Sea eee i i nS xe ae ace

570 | PHARMACEUTICAL BOTANY

opposite, exstipulate, coriaceous, simple, varying from lanceolate
to ovate. Inflorescence a condensed raceme or spike. Flowers
perfect, polygamous or dicecious, 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).
OrFIcIAL DruG Part Usep BoTANICAL ORIGIN HABITAT
Joe Mezereum

Daphne Gnidium Europe and Asia
Daphne Laureola

Mezereum Bark

LYTHRACEZ OR LoosestrirE Famity.—Herbs, as Swamp
Loosestrife (Decodon), Cigar plant (Cuphea) etc., shrubs (Law-
sonia) or low trees containing astringent principles. Leaves
either alternate, opposite or whorled, simple, exstipulate, usually
lanceolate to ovate, entire, often glandular, viscous and contain-
ing idioblasts. Stems and leaves possess bicollateral bundles.
Inflorescence a raceme, spike, or condensed cyme. Flowers
perfect, often dimorphic or trimorphic, as in the spiked or purple
loosestrife (Lythrum Salicaria), usually regular, but pass more or
less to irregular, sometimes very irregular as in genus Cuphea;
Calyx tubular, 3 to many toothed, the tube ribbed; sepals seven to
four, petals equal in number and alternate with the inner teeth
of the calyx, inserted on the mouth of the calyx tube, rarely
absent; stamens fifteen, ten, or five in alternate rows of five each,
inserted hypogynously or perigynously; pistil six-, five-, four-,
two-, to one-carpeled with as many cavities in the ovary and
numerous small ovules; style elongate with knobbed or 2-lobed
stigma. Fruit a capsule. Seeds anatropous, exalbuminous.
Calcium oxalate occurs in rosette aggregates and monoclinic
prisms.

UnorriciAL Druc Part Usep . BoTANICAL ORIGIN

Hasrrar
Lawsonia Leaves Lawsonia inermis N. Africa to
(Henna) as

GRANATACEZ (PUNICACEZ) OR POMEGRANATE Famity.—A
small family of shrubs and trees all of which are included in the

THE DICOTYLEDONS 571

single genus Punica. Leaves opposite to whorled, entire, glabrous
and extipulate. Flowers hermaphrodite, scarlet in the Pome-
granate (Punica Granatum) solitary or aggregate, terminal, calyx
petaloid; corolla of 7 to 5 petals inserted to calyx throat; stamens
numerous; in many series, inserted below the petals; ovary
inferior and imbedded in upgrown receptacle, its upper portion
7 to 5 celled with parietal placentation, its lower portion 3 celled
with central placentation; style filiform; stigma capitate; ovules

Fic. 432.—The Pomegranate, Punica Granatum—Branch with flowers. (Sayre.)

numerous, anatropous. Fruit a spherical, baccate capsule
crowned by the persistent calyx, its locules separated by mem-
branous septa. Seeds numerous, exalbuminous.

OrriciAL Druc Part Usep BorTANnicAL ORIGIN HABITAT
Pelletierinz Tannas Mixture of tannates Punica Granatum India

of the alkaloids
Unorrictat. Druc

Granati Fructus Rind of fruit
Cortex Punica Granatum India
Granatum Bark of stem and

root

Myrtace&® or Myrtie Famiry.—Rarely herbs (Careya)
mostly shrubs or trees, some being the tallest trees known (Euca-
lyptus). Stems often tend to develop cork in flakes which separate

572 PHARMACEUTICAL BOTANY

much as in the Buttonwoods. Leaves rarely alternate, nearly
always opposite, entire, often glistening, subcoriaceous tocoria-

Hi dys
Gp
to

Fic. 433.— The Blue Gum Tree, Eucalyptus Globulus,—Branch. (Sayre.)

ceous (Eucalyptus, Pimenta, etc.), frequently edge-on in position
upon branches. Inflorescence cymose, at times forming scor-

THE DICOTYLEDONS

poid 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, typ-
ically pentamerous, not infrequently reduced to tetramerous

6) F
woul} Ve

i.
ay ER IM

\

SS OT Tp
—

Xs ws
SS

Fic. 434.—Caryophyllus aromaticus. (Sayre.)

(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 (Caryophyllus) or from

574 PHARMACEUTICAL BOTANY

whitish to pink, scarlet, crimson, purple and blue, petals some-
times synpetalous and cup-like, detaching as the flower opens;
stamens usually indefinite and epigynous, varying in the color
of their filaments as do the petals; pistil rarely of ten to six carpels
usually of five, not infrequently, as in Clove, of four carpels;
ovary inferior or semi-inferior, as many-celled as there are carpels
and with central placentation; style elongate; stigma undivided.
Fruit either a hard, woody indehiscent nut (Brazil Nut), a
capsule dehiscing at apex (Eucalyptus) or berry (Caryophyllus).
Seeds exalbuminous.

OrrictAL Druae PAarT Usep BoTANICAL ORIGIN HABITAT

Oleum Eucalypti Volatile oil Eucalyptus Globulus :

and bi other docpabeeany

and

Eucalyptus sp. T ;
Eucalyptol Organic oxide Eucalyptus Globulus ) ~°°"#™*
Caryophyllus Flower buds Caryophyllus aro-

maticus
Oleum Caryophylli Volatile oil Caryophyllus aro- ies Vtandla

maticus
Eugenol Aromatic phenol Caryophyllus aro-

maticus

West Indies
Oleum Pimentze Vol. oil Pimenta officinalis fc America
: Mexico
Oleum Myrciz Volatile oil Pimenta acris West Indies
UnorriciAL Druc
Bay Leaves Leaves Pimenta acris West Indies
EucalyptiGummi Dried gummy Eucalyptus rostrata Australia
exudate and other sp. of

Eucalyptus
Eucalyptus Sythe-shaped leaf Eucalyptus Globulus Australia
Oleum Cajuputi Volatile oil from (Var’s. of Melaleuca :

leaves and twigs crit dara dove or iae

Pimenta Fruit Pimenta officinalis West Indies

CoMBRETACEZ OR MyrosBALANs Famity.—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.

Unorricta, Drug —_—s— Part Usep BorAaNnIcAL OriGINn HAsitAT

Combretum Leaves Combretum Sumatra
sundaicum

Pe ante eee ere ne fi ae

THE DICOTYLEDONS 575

ORDER UMBELLALES OR UMBELLIFLORZ&

This order includes the Ginseng Family (Araliacee) repre-
sented by Ginseng, Spikenard, and Virginian Sarsaparilla, the
Umbellifere or Parsley Family which contains Fennel, Anise,
Coriander, Caraway, Poison Hemlock, Water Hemlock and
other medicinal plants, and the Dogwood Family (Cornacea)
which includes the Dogwoods and Sour Gum. All of these
families agree in usually possessing inflorescences consisting of
umbels and flowers with an inferior ovary and a single whorl of
stamens.

ARALIACE OR GINSENG FamiLy.—Herbs, as Ginseng (Panax
quinquefolium), English Ivy (Hedera Helix), Wild or Virginian
Sarsaparilla (Aralia nudicaulis), etc., undershrubs (Aralia hispida,
etc.), shrubs, as Bristly Sarsaparilla (Fatsia horrida), or trees, as
Hercules Club (Aralia spinosa), with stems which are more or less
hollow along internodes and solid at nodes. Leaves alternate,
varying 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 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, with a
single seed coat.

OrrFiciAL DruG PART UsEpD BOTANICAL ORIGIN HABITAT
Aralia Rhizome and Aralia racemosa Eastern United
(Spikenard) — roots States and

Canada

576 PHARMACEUTICAL BOTANY

UnorFiciAL Druc | Part UsEep BorANnICAL ORIGIN HABITAT
Aralia Nudicaulis Rhizome Aralia nudicaulis Eastern United
: States and

Canada

Aralia Spinosa - Bark Aralia spinosa Eastern United
States

American Ginseng Root Panax quinquefolium North America

Chinese Ginseng Root Panax Ginseng E. Asia

Panax Repens Rhizome Panax repens Japan

UMBELLIFER& OR ParsLEY Famity.—Herbs (Anise, Fennel,
Sweet Cicely, Poison Hemlock, etc.), rarely shrubs, often of
rapid growth, and with upright, fistular (hollow at internodes,
solid at nodes), often grooved and ridged stems. Leaves alter-
nate, 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 surrounded 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 inflexed 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, each with a single seed coat.

OrriciAL DruG Part Usep BorANICAL ORIGIN HAsirat
Anisum Ripe fruit Pimpinella Anisum ) Asia Minor, Egypt
Oleum Anisi Volatile oil Pimpinella Anisum t Ade Greece

Methyl ether of | Pimpinella Anisum
Ancthol |p jest vulgare
penyl phenol
Foeniculum Ripe Fruit Feeniculum vulgare ? Mediterranean
Oleum Feeniculi —- Volatile oil Foeniculum vulgare a
Rhizome and Ferula Sumbul or Asia
Sumbul ba other closely
related spp. of
Ferula having a

musk-like odor

THE DICOTYLEDONS

577

OrriciAL Druc Part Usep BoTANICAL ORIGIN HABITAT
Carum Fruit Carum Carvi Europe, Asia
Oleum Cari Vol. oil Carum Carvi
Asafoetida Gum resin Ferula foetida, F. Persia and

Assa-foetida, F. Afghanistan
rubricaulis, etc.
Coriandrum Ripe fruit Coriandrum isiiecrheaa,
: sativum :
Oleum Coriandri ‘Volatile oil Coriandrum — Snsiconiug
: regions
sativum
Apii Fructus Ripe fruit Apium graveolens Southern Europe
UnorFic1AL Druc
Angelic Fructus Ripe Fruit Angelica Archangel- Northern Europe
ica and other and Siberia
species of Angelica
Angelic Radix Rhizome and Angelica Arch- Northern Europe
roots angelica and other and Siberia
species of Angelica
Conium Unripe fruit Conium maculatum Europe
Ammoniacum Gum-resin. Dorema Persia
Ammoniacum
Galbanum Gum-resin Ferula galbaniflua Persia and
Afghanistan
Imperatoria Root Imperatoria Europe
(Masterwort) Ostruthium
Levisticum Root Levisticum officinale Europe
(Lovage) ee ces
Pimpinella Roots Hew ae Sone Central Europe
Petroselinum Ripe fruit
Petroselini Radix Root Petroselinum Southern Europe
Apiol Oleoresin from sativum and Asia Minor

CoRNACEA

root

or Docwoop F amiLy.—Herbs, as Bunchberry

(Cornus canadensis), etc., shrubs, as Blood-Twig Dogwood (Cornus
sanguinea), etc. or trees, as Flowering Dogwood (Cornus florida)
and Sour Gum (Nyssa sylvatica), etc. Leaves simple, alternate
(Sour Gum), or opposite (Dogwoods). Inflorescence an umbel
or determinate head (glomerule), the whole being 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

578 PHARMACEUTICAL BOTANY

between the nectar disc; pistil syncarpous, bicarpellate, nie:
ericab ellie: ovary as many celled with one pendulous ovule in
>

a

Fic. 435.—Conium maculatum L., the poison hemlock. Note spotted stems,
decompound leaves and compound umbels of white flowers. The poisonous
juice of this plant was employed as a potion by the ancient Greeks which was
given to criminals as a means of capital punishment.
each cavity; style usually simple, ending in rounded or slightly

bilobed stigma. Fruit a two-seeded drupe. Seeds albuminous
with single seed coat.

THE DICOTYLEDONS 579

UnorFIcIAL DruG PART UsEp BoTANICAL ORIGIN HABITAT
Cornus Bark of root Cornus florida Eastern United States
and Canada

SUB-CLASS—SYMPETALA (GAMOPETALZ OR METACHLAMYDEZ)
A division of dicotyledonous plants in which the flowers
possess both calyx and corolla, the latter with petals more or less
united into one piece.
ORDER ERICALES

EricAcE# OR HEATH Famity.—Sub-herbaceous, as Pipsis-.
sewa (Chimaphila), suffruticose, as the Heather (Calluna),

eeeeen = — - “sy
t ‘

Fic. 436.

Fic. 436.—Wintergreen (Gaultheria procumbens), an aromatic, evergreen, peren-
nial with ovate leaves, nodding, white, urn-shaped, axillary flowers and red, berry-
like fruits, each of the latter representing a 5-celled capsule enveloped by a fleshy
calyx. Native to sandy soils in cool, damp woods of Eastern North America.
(Reproduced from U. S. Dept. Agric. Miscellaneous Publication 77.)

Fic. 437.—Pipsissewa (Chimaphila umbellata) on left. The two plants on right
are Chimaphila maculata, the spotted pipsissewa”’ or “‘spotted wintergreen.” (Repro-
duced from U. S. Dept. Agric. Misc. Publ. 77.)

fruticose (Azaleas, Mountain Laurel, 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 creeping, more or less

580 PHARMACEUTICAL BOTANY

woody, rarely through a saprophytic or parasitic connection
becoming soft, annual and white above ground, as in Indian
Pipe (Monotropa uniflora). Leaves alternate, simple, entire,
exstipulate, rarely soft, delicate, herbaceous (Azaleas), usually
leathery to wiry and evergreen, more rarely (Pterospora (Pine
Drops), Monotropa, etc.) becoming greenish-blue, bluish-yellow,
yellowish-white to white and correspondingly saprophytic.
Inflorescence typically a raceme (Pyrola, Andromeda, Gaylussacia,
Erica, Arctostaphylos Uva Urst, 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 (Mono-

Fic. 438.—Flower of a Blueberry (Vaccinium). A, mediam lengthwise section;

. B, external view. (Robbins.)

tropa uniflora). Flowers regular, passing to irregular (Rhodo-
dendron), pentamerous or tetramerous ; sepals five to four, rarely
fewer, apo- to synsepalous, usually green, sometimes brightly
petaloid; petals five, more rarely four, slightly to deeply syn-
petalous, cup-shaped, as in Mountain Laurel (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, becoming by absorp-
tion of inner circle, five to four only, hypogynous, epipetalous or
epigynous; anthers two-celled, dehiscing by apical pores, as in
Bearberry (Arctostaphylos) or apical slits; pollen sometimes
agglutinated into long viscous threads; pistil five- to four-,

THE DICOTYLEDONS 581

_rarely six- to eight-carpeled, superior, rarely semi-inferior to
inferior, as in Blueberries (Vaccinee); ovary as many celled as
there are carpels; style elongated, filiform, usually five- to four-
lobed. Fruit a capsule, as in Epigea repens or Trailing Arbutus,
a berry, as in Blueberries and Cranberries (Vaccinium) and

Fic. 439.—Arctostaphylos Uva ursi. 1. Twig bearing spatulate leaves and urn-
shaped flowers. 2. Flower in longitudinal section. 3. Pollen tetrad. 4. Fruit.
5. Cross section of fruit. > 2-5. (From Strasburger, Noll, Schenck and Schimper
after Berg and Schmidt.)

Huckleberries (Gaylussacia) or false drupe, as in the Wintergreen
(Gaultheria procumbens). Seeds small, anatropous.

OfFFicIAL DruG Part Usep BOTANICAL ORIGIN HABITAT
United States,
Chimaphila Leaf Chimaphila umbellata + Canada, Northern

Europe and Asia.
en Northern United

Uva Ursi Leaf Arctostaphylos Uva Ursi¢States and Canada,
Europe and Asia

Methylis Salicylas Volatile oil Gaultheria procumbens United States and

(in part) Canada
UnorriciAL Druc
Gaultheria Leaf Gaultheria procumbens United States and
Canada
tera moe Macointam parr avonee United States
Neomyrtillin Gncwe Co

OrpDER EBENALES

SAPOTACEZ OR STAR APPLE Famity.—Tropical shrubs or
trees (Palaquium) characterized by the presence of laticiferous sacs

582 PHARMACEUTICAL BOTANY

in the pith and cortex of the stems and adjoining the veins of the
leaves. Leaves alternate, exstipulate, evergreen and coriaceous.

Fic. 440.—The manzanita (Arctostaphylos manzanita), an ericaceous shrub growing
in western United States and common on hillsides near the Pacific coast. (After
Stanford, General and Economic Botany, D, Appleton-Century Co., publishers.)

Flowers perfect, large and axillary. Fruit a berry (Palaquium)
rarely a capsule (Ponteria).

UnorriciaL Druc Part Usep BorANIcAL ORIGIN HABITAT
Gutta Percha Purified coagulated Various species of Malay Archipelago
milky exudate Palaquium and
Payena
Chicle Dried latex Achras Sapota Mexico and Central
America
Balata Dried latex Mimusops West Indies and S.
globosa America

STYRACEZ OR Benzoin Famity.—Shrubs or low trees.
Leaves alternate to opposite, entire, often acuminate and
exstipulate. Flowers perfect, 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 to two, perigynous or sub-
hypogynous; pistil bicarpellary or four to five carpellate. Fruit

either drupe-like or dry, often winged and rarely as many-celled
as there are carpels.

THE DICOTYLEDONS 583

-OrFiciAL Druc PART Usep BoTANICAL ORIGIN HABITAT
Styrax Benzoin East Indies
Styrax tonkinensis Siam

or other species of

Styrax

Benzoinum Balsamic resin

ORDER GENTIANALES

OLEACEa OR Olive Famity.—Shrubs, as Lilacs (Springa),
Golden Bell (Forsythia) or trees (as the Ash (Fraxinus), Olive
(Olea), etc.) with stems possessing close white wood, and slightly
swollen or enlarged nodes. Leaves opposite, decussate, simple,
rarely pinnately-compound (Ash). Inflorescence dichasial 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 syn-
petalous, elongated into a narrow tube, expanding 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 inde-
hiscent akene (Ash). Seeds with moderate to scanty albumen,
becoming occasionally exalbuminous.

OrriciAL Druc Part Usep . BoTANICAL ORIGIN HABITAT
Manna Dried exudation Fraxinus Ornus Southern Europe
Oleum Olivze Fixed oil Olea europza Southern Europe,

s Algeria, Asia
Chionanthus Bark of root Chionanthus virginica Southern United
States

UnorrictaL DruaG
Fraxinus Bark Fraxinus americana Northern United
States and Canada

LoGANIACE OR LoGANIA Famity.—Herbs (Spigelia or Pink
Root, etc.), woody vines. (Gelsemium or Yellow Jasmine, etc.) or
trees (Strychnos Nux Vomica, etc.) with a bitter juice usually con-
taining 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 cymose (Spigelia) (scor-

584 PHARMACEUTICAL BOTANY

pioid cyme in Siérychnos), sometimes condensed into solitary,
axillary flowers. Flowers perfect, usually regular; calyx gamo-
sepalous; corolla gamopetalous, hypogynous, rotate, campanulate

Fic. 441.—Strychnos Nux-vorica—F lowering branch and seeds. (Sayre.)

or infundibuliform; stamens 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 marilandica), sometimes a berry

THE DICOTYLEDONS 585

as

Fic. 442.—Pinkroot (Spigelia marilandica) a perennial herb of the Loganiacee.
Note the opposite leaves and one-sided spike of tubular flowers. The corollas of
these are red outside and yellow within. The rhizome and roots are used as a worm
remedy or anthelmintic. (Reproduced from U. S. Dept. Agric. Misc. Pub. 77.)

(Strychnos Nux-Vomica) or drupe. Seeds numerous or solitary,
sometimes winged.

OrriciAL DruG Part Usep BoTANICAL ORIGIN HABITAT
Nux Vomica Seeds Strychnos Nux- India
Vomica
Brucinz Sulfas Sulfate of alkaloid from
seed
Strychnina Alkaloid from seed also Strychnos Nux- Pe tee,
sulfate, nitrate and Vomica and other here
phosphate of strychnine (Loganiacee
Gelsemium Rhizome and roots Gelsemium Southern United
sempervirens States
UnorriciAL Druc
Curare Extract from bark Strychnos toxifera S. America
et al. spp.
Ignatia Seed Strychnos Ignatii Philippine
Islands
Spigelia Rhizome and roots Spigelia marilandica Southern United
States

GENTIANACE OR GENTIAN FamiLy.—Herbs, often low-grow-
ing. Roots and short stems sometimes more or less thickened
(Gentiana lutea). Leaves opposite, decussate, entire, exstipulate.
Inflorescence cymose (Gentiana lutea) or condensing to a single

586 PHARMACEUTICAL BOTANY

solitary, terminal flower (Gentiana verna, G. acaulis, etc.). Flowers
regular, perfect, pentamerous or tetramerous, sepals five to four,
green, more or less synsepalous, not infrequently everted or
reflexed, corolla of five, rarely four petals, more or less

Nit

Uy
ie

Fic. 443.—The yellow gentian or bitter root, Gentiana lutea,

—Flowering h
dissected flower. (Sayre.) oe aed

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 ‘ss
elongated with bilobed to divided stigma. Fruit a ca

: ans sule.
Seeds albuminous. Plants contain bitter glucosides, 5

THE: DICOTYLEDONS 587

OrriciAL Druc Part Usep BorANICAL ORIGIN HABITAT
Gentiana Rhizome and Gentiana lutea Europe
root
UNOFFICIAL
Druc
Chirata Entire plant Swertia Chirayita Northern India
Centaurium Flowering plant Erythraa Centaurium Europe
Menyanthes Leaves Menyanthes trifoliata | Europe and Asia
Sabbatia Herb Sabbatia angularis Eastern United States

and Canada

Fic. 444.

Fic. 444.—Menyanthes trifoliata, the buckbean or bogbean, whose leaves are
employed as a simple bitter. This perennial herb occurs in bogs of Europe, Asia
and North America. Note the long petioled, trifoliate leaves whose bases sheathe
the rhizome and the long peduncle bearing a raceme of tubular flowers which are
pinkish-purple or white. (Reproduced from U. S. Dept. Agric. Misc. Pub. 77.)

Fic. 445.—Apocynum cannabinum or Canadian hemp, a perennial herb whose
rhizome and roots contain cardiac stimulant glucosides. Note the opposite, ovate-
lanceolate leaves, the central terminal cyme of greenish white flowers and the fruit
of paired follicles (on right). (Reproduced from U.S. Dept. Agric. Misc. Publ. G73)

APOCYNACE OR Doc Bane Famity.—Herbs, rarely shrubs,
not infrequently clambering or climbing in habit (Adlamanda).
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,

Se ae ee ee ean

'
;
:
:
3

588 PHARMACEUTICAL BOTANY

simple, entire, deciduous or evergreen. Inflorescence cymose.
Flowers regular, pentamerous, rarely tetramerous; sepals five,
gamosepalous, green, rarely subpetaloid to petaloid; petals five,
slightly to deeply gamopetalous, in shape varying from open
tubular, stellate, to elongate tubular to elongate funnel-shaped,

Fic. 446.—Strophanthus hispidus—Branch and seed with comose awn. (Sayre)

in color varying from greenish-yellow to white or from yellow-
red to crimson to crimson-purple to nearly purple-blue; stamens
five, epipetalous; pistil usually bicarpellate; ovary two-celled
with central placentation; style more or less elongate with ter-
minal brush of hairs, knobbed or multifid; stigma a circular band
or circular spur beneath terminal style swelling. Fruit two folli-

cles (Apocynum, etc.), a berry, drupe, or capsule. Seeds flattened
frequently hairy, albuminous. :

589

THE DICOTYLEDONS

OrrFiciAL DruG Part UsEpD BoTANICAL ORIGIN HABITAT
Strophanthus Seed (deprived of awn) oes eee { Attica
Strophanthus hispidus

Strophanthin —_Glucoside Strophanthus Kombé Africa

Apocynum Rhizome and roots Apocynum United States
cannabinum and Canada

UnorrFiciAL DruGc

Aspidosperma Bark Aspidosperma Argentine
Quebracho blanco Republic

ASCLEPIADACE OR MILKWEED Fami_y.—Herbs or shrubs
containing a milky juice, many
species yielding rubber. Leaves
entire, more or less fleshy, some-.
times verticillate. Inflorescence
usually a dichasial or scorpioid
cyme. Flowers regular, pen-
tamerous; sepals woolly, small,
synsepalous; petals five, rarely
four, synpetalous, elongated into
awls; the corolla varying in shape
from stellate to campanulate and
in color from pale green to
yellow, to greenish-brown, choc-
olate, or from white to yellow, to -

scarlet, to crimson, to purple, to
blue; stamens five, epipetalous,
fused in relation, forming a cylin-
drical swollen mass around the
central pistil; filaments flattened
and furnished with a crown
having various appendages;
anthers two-celled, each cell con-
taining a pollen mass (pollinium),

the

Fic. 447.—Asclepias tuberosa,
Pleurisy-root or Orange Milkweed.
The medicinal root, to left, is orange
brown and devoid of latex. Note the
alternate leaves and terminal cymes of
unbels, which bear orange-colored
flowers. To right, stem bearing follicles
two of which are discharging their seeds.
(Reproduced from U. S. Dept. Agric.
Miscellaenous Publication 77.)

adhering to the glandular prominences of the stigma; pistil
bicarpellate, superior. Fruit typically two dry follicles (Asclepias),
rarely becoming succulent or bladdery. Seeds numerous, com-
pressed, imbricate, with a comose appendage.

Druc Part Usep BoTANICAL ORIGIN Hasitat
Asclepias Root Asclepias tuberosa United States
Condurango, N.F. Bark Marsdenia Peru and Ecuador

Condurango

590 PHARMACEUTICAL BOTANY

OrDER TUBIFLORZ OR POLEMONIALES

CoNVOLVULACE2 OR Morninc Giory Famity.—Frequently
herbaceous, more rarely sub-woody or woody, perennial climb-
ing 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 cir-

Fic. 448.—Convolvulus scammonia—Branch. (Sayre.)

cumnutating in action. Vascular bundles frequently _bicol-
lateral. Leaves alternate, simple, exstipulate, varying from
cordate to cordate-sagittal, to broad-reniform to reniform,
palmately lobed to palmatifid to palmately-compound (Jpomea
shows all these transitions). Stem and leaves frequently contain
a dull, viscous, watery to milky-white, resinous juice. Inflo-
rescence a scorpioid cyme, becoming reduced in some forms to

THE DICOTYLEDONS 591

a solitary flower. Flowers pentamerous; 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 filaments 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, Convolvulus, etc.), dehiscing
septifragally, rarely a berry. Seeds scantily albuminous to
exalbuminous.

OrriciAL DruG PART UsED BOTANICAL ORIGIN HABITAT
Jalapa Tuberous root Exogonium Jalapa Mexico
Ipomoea Root Ipomoea orizabensis Mexico
UnorriciAL Druc
Tampico Jalap Root Tpomeéa simulans Mexico
Wild Jalap Root Ipomoea pandurata United States
Turpeth Root Root : Operculina Turpethum East Indies
Scammoniz Radix Root Convolvulus Asia Minor,

Scammonia Greece, Syria

HyDROPHYLLACEAE OR WATER LEAF Famity.—Annual, her-
baceous, 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. Inflorescence rarely expanded, usually scorpioid
cymes. Flowers small to large, funnel-form in Yerba Santa
(Eriodictyon californicum); 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
bicarpellate. Fruit a two-celled capsule. dehiscing usually
septicidally.

OrrFiciAL DruG PArT UsEeD BoTANICAL ORIGIN HapsiTAT
Eriodictyon Leaves » Eriodictyon California and New
: californicum Mexico

BoRAGINACE& OR BoraGE Famity.—Herbaceous (Boraginee
sub-family) or shrubby (Heliotropee sub-family) plants, forming a
primary root and a single or often branched shoots. Leaves
often divisible into expanded, sometimes large basal and alter-

592 PHARMACEUTICAL BOTANY

nate, scattered cauline leaves. Each of these simple, exstipulate,
often hairy, rarely glabrous. Inflorescence a raceme of dichasial
or scorpioid cymes, at times condensed into a dichasium of
scorpioids or a simple scorpioid cyme. Flowers pentamerous,
regular, passing to slight or marked irregularity as in Blue-weed
(Echium); sepals five, green, slightly or deeply gamosepalous,
often hairy; petals five, the corolla varying in shape from rotate

——

* +" -s
j a fa
fy yy”
* p
5 ; r y A
wt ae Ay See 7 ‘3
- i ¥ i .
4 xi
a h

Fic. 449.—Yerba Santa (Eriodictyon
californicum), an evergreen shrub found
along the coastal ranges from central
California to Oregon. The leaves are
used in expectorant preparations. (Re-

Fic.
Comfrey, a hairy perennial herb with a
mucilaginous root and purplish, blue,
or pinkish-white tubular flowers borne

in pendulous cymes. (Reproduced from

produced from U. S. Dept. Agric. Miscel-
laneous Publ. 77.)

U.S. Dept. Agric. Misc. Publ. 77.)

with shallow tube, as in Forget-me-not (M fyosotis) and Borage, to
tubular, as in Comfrey (Symphytum), to funnel-shaped in most
species; in color, all transitions, 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
(Heliotropee) or exalbuminous (Boraginea).

THE DICOTYLEDONS

593

UnorriciAL Druc Part Usep + BoranicaA ORIGIN HABITAT

Symphytum Root Symphytum officinale Europe and

(Comfrey) United States

Cynoglossum Herb and root Cynoglossum United States

(Hound’s tongue) officinale

Alkanet Root Alkanna tinctoria So. Europe and
Asia

Lungwort Leaf Pulmonaria officinalis Europe

VERBENACE& OR VERVAIN FamiLy.—Herbs, as the Vervains
(Verbena), shrubs, as the Glory-bower Cana, rarely trees,

as Premma arborea, one of the
sources of the drug Tonga, and

Tectona or Teakwood, whose |
stems and branches are usually

quadrangular and rarely scented.

Leaves generally opposite, ex- :

stipulate, simple or compound.
Inflorescence a terminal panicle
of spikes (Verbena hastata), acyme
(Beauty-berry or Callicarpa) or
head (Lippia lanceolata). Flowers
white, pink or blue (Verbena
hastata), irregular, more or less
2-lipped; calyx gamosepalous,
tubular; corolla gamopetalous,
hypogynous with a 4-5 fid limb;
stamens generally 4, didynamous
and inserted on the corolla tube
or throat; pistil of 2-4 carpels, a
terminal style and undivided
stigma. Fruit a drupe or 2 to
into as many nutlets.

Part Usep
Overground portion
Bark

UnorrIiciAL Druc
Verbena
Tonga

ferrreer

:

Fic. 451, —The Blue ed Ver-
Note the lanceolate leaves
with serrate margins and the flowers
which are usually blue, and arranged in

bena hastata.

panicles of narrow spikes. (Reproduced
from U. S. Dept. Agric. Misc. Publ. 77.)

4 celled berry, usually splitting
Seeds exalbuminous.

BoTANICAL ORIGIN HABITAT
Verbena hastata United States
Premna arborea Fiji Islands

(in part)

Lapiat# (LAMIACE2) OR Mint Famity. —Herbs producing

creeping runners that spread out and root at the nodes.

Stems

594 PHARMACEUTICAL BOTANY

quadrangular, rarely cylindrical in outline. Leaves opposite,
decussate, mainly petiolate; leaf margin nearly always serrate,
dentate or crenate. Stems and leaves further characterized
by the presence of glandular hairs containing aromatic volatile
rca. guerregem Ol. These hairs, for the greater
© 48. part, consist of a short one-celled
_ stalk and a head (gland) of six or
« cightcells. Inflorescence a raceme
.. or spike of verticillasters (double
_ dichasial cymes) or, as in the
_ Ground Ivy, a reduced verticillas-
_ ter. Flowers typically pentamer-
| ous, rarely tetramerous; sepals five,

_| synsepalous, ribbed and forming a
_ tubular, regular or irregular bilabi-
» ate (Scullcap, etc.) calyx whose

upper lip is bifid and lower trifid;
corolla of five to four gamopetalous
ict: mB Fe petals, hypogynous, frequently two-
_ Fic. 452.—Peppermint (Mentha lipped, the upper lip bifid, the
sot hee wot fee ee lower trifid; stamens four, didyna-
of eastern and central North Mous, rarely one pair alone fertile
America. Note the opposite, petio- and the other pair reduced, in some

late, ovate-oblong leaves with serrate : “
cases alm Be
margins and the flowers which are ost or quite to disappear

purple and borne on blunt spikes of ing point, as in the Sage (Salvia)
verticillasters terminating the square and the Horsemint (M. onarda);
stems. (Reproduced from U.S. Dept. pistil bicarpellate, embryologically
aes Ae: Peat 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 gynobasic, elongate, slender, with
two stigmatic surfaces. Fruit four nutlets enclosing as many
exalbuminous seeds.

i

OrrictAL Druc PART Usep BoraAnicAL Oricin Hasrrar
Mentha Piperita Leaves and tops Mentha piperita Europe
(Peppermint)

Oleum. Menthz Volatile oil from Mentha piperita
Piperita fresh aerial parts of
flowering plant

Europe

OrFIcIAL DruG
Menthol

Mentha Viridis
(Spearmint)
Oleum Menthz
Viridis
Scutellaria
(Scullcap)
Thymus (Thyme)

Oleum Thymi

Oleum Rosmarini

Oleum Lavandulz

ee ee ee eer os

Cataria

(Catnep)

Salvia (Sage)
UnorriciAL Druc
Marrubium -
(Horehound)
Hedeoma
(Pennyroyal)

THE DICOTYLEDONS

Part Usep
Alcohol separated
from vol. oil
Leaves and tops

Volatile oil from
fresh aerial parts,

of flowering plants
Overground portion

Leaves and flower-
ing tops

Volatile oil from
flowering plant
Volatile oil from

fresh flowering tops

Volatile oil from

fresh flowering tops

BoTANICAL ORIGIN
Mentha arvensis
var. piperascens
Mentha spicata

Mentha spicata .

Scutellaria lateri-
flora

Thymus vulgaris

Rosmarinus
officinalis
Lavandula Spica

Fic. 453.—Catnip (Nepeta Cataria). Note the cordate leaves with crenate
margins and the spikes of whorled flowers which are white and spotted with purple.
(Reproduced from U.S. Dept. Agric. Misc. Publ. 77.)

Leaves and
flowering tops
Leaves

Leaves and
flowering tops
Leaves and
flowering tops

Nepeta Cataria
Salvia officinalis

Marrubium
vulgare

Hedeoma
pulegioides

595
HABITAT
Europe
Europe
Europe

United States
and Canada

Southern Europe

Mediterranean
Basin

Southern Europe

Europe and Asia
Southern Europe
Europe and Asia

United States
and Canada

PPLE hdl Sete Mag HN A Sg oc

et ee ee eS any Le a oe

596

PHARMACEUTICAL BOTANY

UnorriciAL Druc PART Usep BoTANICAL ORIGIN HABITAT
Herba Majoranz Leaves and Origanum Mediterranean
(Sweet Marjoram) flowering tops Majorana regions
Collinsonia Rhizome and roots Collinsonia
(Stone Root) canadense United States
Serpyllum Leaves and Thymus Europe and Asia
(Wild Thyme) flowering tops Serpyllum
Melissa (Balm) Leaves and Melissa officinalis Southern Europe,
flowering tops Asia Minor
Monarda Leaves and Monarda punctata United States
(Horsemint) flowering tops
Origanum Leaves and Origanum vulgare Europe, Asia and
flowering tops North Africa
Hyssopus Leaves and Hyssopus Southern Europe
flowering tops officinalis
Summer Savory Leaves Satureja hortensis Southern Europe
Mountain Mint Leaves Pycnanthemum United States
montanum
Sweet Basil Leaves Ocimum Basilicum Asia and Africa
Motherwort Leaves and Leonurus Cardiaca Europe
flowering tops
Lavender Flowers Lavandula Spica _S. Europe
Rosmarinus Leaves Rosmarinus - j
ers Mediterranean
officinalis Basi
Greek Tea Herb =

Sideritis spp.

SOLANACE2 oR NiGutsHapE Famiry.—Herbs (potato,
tomato, horsenettle), shrubs (Lycium spp.), vines (Solanum dulca-
mara or bittersweet), rarely trees as in some tropical Daturas.

‘Stems and leaves possess bicollateral bundles. Many of the

plants as belladonna, stramonium, hyoscyamus and scopola
contain mydriatic alkaloids. Leaves alternate, exstipulate,
entire or more or Jess lobed, rarely compound; often glandular-
hairy. Flowers in cymes or solitary; regular or rarely irregular
(Petunia, Tobacco spp.), pentamerous, perfect, synphyllous;
sepals green (rarely petaloid), rotate to tubular, usually per-
sistent and accrescent; petals rotate (Solanum), to tubular
(Atropa), to funnel-shaped (Tobacco), and so (1) open to all
comers, or (2) to bees or wasps, or (3) to butterflies, moths;
color, greenish-yellow, or greenish-white, to white, to pink,
crimson, purple, rarely blue; stamens five, epipetalous, hypog-
ynous, along with style usually forming nectar glands. Fila-
ments short to long, anthers dehiscing longitudinally or by
apical pores; pistil bicarpellate, syncarpous, with or without

THE DICOTYLEDONS S37

nectar girdle; superior ovary, two-celled with central placenta-
tion, ovules numerous, style more or less elongate with bilobed
or bifid stigma. Fruit, a capsule dehiscing longitudinally

Fic. 454.—The Henbane (Hyoscyamus niger Linné.), a member of the Nightshade -
family, whose leaves, with or without the tops, constitute the official drug Hyoscy-
amus and are a source of the valuable medicinal alkaloids, hyoscyamine and
scopolamine.

(Tobacco, Stramonium) or transversely (Henbane); or a berry
(potato, egg-plant, tomato, red pepper). Seeds albuminous.

OrriciaL Druc Part-UseD BoTANICAL ORIGIN Hasirat
Belladonnz Folium Leaves and tops Atropa Belladonna Central and
Southern Europe,
Belladonnz Radix Root Atropa Belladonna Asia Minor and
Atropina Alkaloid Atropa Belladonna Persia
Stramonium Leaves Datura Stramonium Asia and Tropical

America

598

OrFIcIAL DrucG
Hyoscyamus
Capsicum
UnorriciAL DruG
Dulcamara
Manaca
Duboisia
Tabacum
Scopola

Paprika
Pimiento
Miré

Solanum

PHARMACEUTICAL BOTANY

Part Usep
Leaves and tops
Fruit

Twigs and stems
Root

Leaves

Leaves

Rhizome

Fruit
Fruit
Roots, stems

and leaves
Ripe fruit

BoTANICAL ORIGIN
Hyoscyamus niger
Capsicum frutescens

Solanum Dulcamara
Brunfelsia Hopeana
Duboisia myoporoides
Nicotiana tabacum
Scopola carniolica

Capsicum annuum
var. longum
Variety of Capsicum
annuum

Brunfelsia
hydrangzformis
Solanum carolinense

HABITAT
Europe, Asia
Tropical America

Europe and Asia
Tropical America
Australia.
Tropical America
Alps and Car-
pathian Mts.
Tropical Amer-
ica; cultivated
Spain

South America

United States

Fic, 455.—The Deadly Nightshade, Atropa Belladonna,—Branch.

(Sayre.)

Pe EN ee SE ee EN ee meee

THE DICOTYLEDONS 599

SCROPHULARIACE2 OR Ficwort Famity.—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, Buchnera, Rhinanthus, etc., 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

Fic. 456.—Bittersweet (Solanum Dulcamara). Note transition from simple to lobed
to compound leaves and flowers with rotate corollas. (After Gager.)

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
regular, 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 (Verbascum Blattaria, etc.) to irregular tubular to
elongate, irregular bilabiate to funnel-shaped. In color, corolla
varies from greenish to greenish-yellow or white (Scrophularia)

600

PHARMACEUTICAL BOTANY

to pure white or from red to purple to blue (Veronica). Stamens
five, fertile, equal in length in a few Verbascum (mullein) species
or unequal in other Verbascum species to stamens four with a long
sterile staminode (Pentstemon, beard-tongue) to four didyna-

peor

be

fic. 457.—Culver’s Root or Tall
Speedwell (Veronica virginica). Note the minal
rhizome and roots and portions of the
stem bearing whorls of lanceolate,
serrate leaves and terminal spikes of
usually white, often pink or blue flowers.
(Reproduced from U.S. Dept. Agric. Misc.

Publ. 77.)

OrriciaL Druc
Digitalis
Digitalis Pulverata

Leptandra
(Culver’s Root)
UnorriciAL Druc

Verbasci Flores

Euphrasia
(Eye-bright)
Verbasci Folia
Balmony

Part Usep

Leaf
Standardized
powd. leaf
Rhizome and
roots

Corollas with
stamens

Plant

Leaves
Leaves

8

mous stamens with a short petal-
oid staminode (Scrophularia,
figwort) to four didynamous
stamens with a minute often
nectariferous staminode (Linaria,
toadflax), to frequently four
didynamous stamens only, the
two lateral or two anterior
stamens stronger and_ longer
(Antirrhinum, snapdragon), to two

_ perfect stamens and two minute

staminodes (Calceolaria), to two
stamens alone developed (Veron-
ica, speedwell). Pistil bicarpel-
late, ovary two-celled with
central placentation; style ter-
with bilobed stigma;
ovules numerous, small. Fruit
a two-celled and usually many-
seeded capsule. Seeds richly
albuminous, anatropous or
amphitropous.

BorTANIcAL OrIGIN HABITAT

Digitalis purpurea Europe

United States
and Canada

Veronica virginica

Verbascum
phlomoides
Verbascum
thapsiforme

Europe and Asia

Euphrasia officinalis Europe

Verbascum Thapsus Europe and Asia
Chelone glabra North America

THE DICOTYLEDONS 601

PEDALIACEAZ OR SESAME Famity.—Tropical herbs often
thickly covered with viscous hairs. Leaves soft, usually alter-

gt

Fic. 458.—The foxglove, Digitalis purpurea var. gloxinaflora, Second year of growth.

nate, more rarely opposite, exstipulate. Flowers irregular,
pentamerous. Fruit a capsule (Sesamum, etc.), drupe, or rarely
a one-seeded, indehiscent nut. Seeds exalbuminous usually.

602 PHARMACEUTICAL BOTANY

_ OrriciAL Druc PART Usep BoTANICAL ORIGIN HABITAT
Oleum Sesami Fixed oil | Sesamum indicum __ Asia and Africa
(Benne Oil) (cultivated varieties)

ACANTHACE2 OR AcANTHUS FamiLy.—Usually herbaceous
(Ruellia), rarely sub-woody or woody plants, occasionally bushy

Fic. 459.—Plantago Psyllium, the source of an excellent variety of Psyllium seed.

in habit, containing 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, exstipu-
late. Inflorescence a raceme of condensed cymes, becoming
a simple raceme or spike, rarely condensed into a solitary

THE DICOTYLEDONS 603

terminal inflorescence. Flowers hermaphrodite, usually irregu-
lar; calyx five-cleft; corolla hypogynous, gamopetalous, 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-
pylotropal ovules; style terminal, filiform. Fruit a capsule
containing numerous curved seeds. The family is of pharma-
ceutic interest mainly because of Ruellia ciliosa, a pubescent
perennial herb growing in the Eastern United States, whose
rhizome and roots have frequently been admixed with or sub-
stituted for Spigelia.

PLANTAGINACE& OR PLANTAIN Fami_y.—Annual or perennial
herbs of caulescent (Plantago Psyllium, P. arenaria, etc.) or more
usually acaulescent habit (Plantago lanceolata), rarely sub-shrubs.
Leaves all radical in most species, or in a rosette, alternate or
opposite to whorled, simple, nerved, flat, entire or toothed.
Flowers usually hermaphroditic, regular, tetramerous, and
arranged on spikes; calyx of 4 sepals; corolla gamopetalous,
hypogynous, tubular with a 4-lobed limb; stamens 4, epipetalous;
carpels 2 or 1, the ovary 1 to 4 celled, ovules 1 to 8 in each cell,
peltate on the middle of the septum in many ovuled cells or at
the bottom in 1-ovuled cells; style filiform with 2 lines of stig-
matic papilla. Fruit a pyxis in the Plantains, rarely a bony
nucule. Seeds albuminous with a straight (Plantago) to curved
(Bouguerta) embryo. The seed coat of many species abounds in
mucilage, accounting for their laxative properties.

OrriciAL Druc Part Usep _ BorAnicAL ORIGIN HABITAT
sgt a Arts Mediterranean countries
Plantaginis Semen Seed Plantago arenaria
Plantago ovata Asia, S. Europe, N. Africa

OrDER RUBIALES

RUuBIACEZ oR MADDER Famity.—Herbs, as Cleavers (Galium)
and Partridge Berry (Mitchella), etc., shrubs as the Ipecac
(Cephaélis) and the Buttonbush (Cephalanthus), or trees (Cinchona
species) with fibrous roots, sometimes, as in Cephaélis Ipecacuanha,
annularly enlarged. Roots, stems and to a less extent leaves

Pee Poets ee aD a Rg en eee ree tren oe

604 PHARMACEUTICAL BOTANY

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;

Fic. 460.—Cephaélis Ipecacuanha—Plant and dried root. (Sayre.)

sepals five (Cinchona, etc.) but four in Galium, small, green, sub-
tended with other flowers by one or two or more enlarged peta-
loid. 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

THE DICOTYLEDONS

605

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.
OrriciAL Druc PART Usep - BoTANICAL ORIGIN
Feebly basic — arabica
ee principle Coffea liberica
Cinchona Bark Cinchona succirubra
Quinina Alkaloid or its hybrids

Quinidinz Sulfas Sulfate of Alkaloid \Cinchona Ledgeri-

Cinchoninz Sulfas Sulfate of Alkaloid{ ana, C. Calisaya

Cinchonidinz Sulfas Sulfate of Alkaloid jand hybrids of
these with other
Cinchona species

Dried aqueous
Gambir ae ee {om Gambir

from leaves and

twigs

Cephaélis

Ipecacuanha Rhizome and ‘ Ipecacuanha

root Cephaélis acuminata
Emetine Hydrochloride of
Hydrochloridum alkaloid emetine

obtained from

Ipecac.
Mitchella Entire plant Mitchella repens

Hasirat
Eastern Africa
Liberia

South America

Malay
Archipelago

Brazil

United States of
Columbia

North America

Ethylhydrocupreinz Hydrochloridum, a synthetic compound made from cuprea

bark or quinine.
UnoFFICIAL DRuG

Coffea Tosta Roasted seeds ‘aes ae { Attica
Yohimbe Bark Corynanthe Johimbi Africa
Remiji dunculata :

Guprey Hare saad eontis atic { Brazil
Loxa Bark Bark Cinchona officinalis | Ecuador and Peru
Ethylhydrocupreine Alkaloidal com-

pound prepared

from cupreine

found in bark of

Remijia pedunculata
Cleavers Herb Galium aperine Europe and Asia
Madder Root Rubia tinctorum Mediterranean region

Sweet Woodruff Entire plant Asperula odorata Europe, W. Asia

606 PHARMACEUTICAL BOTANY

CAPRIFOLIACE2 OR Honey SuckLe Famity.—Shrubs (Elder,
Viburnum, Snowberry, etc.) or rarely herbs (Twin Flower).
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 Honeysuckles

ee re

Fic, 461.—Two medicinal Viburnums. A, Viburnum Opulus var. americanum,

the ich of cramp bark. B, Viburnum prunifolium, the black haw, whose root bark
is official.

(Lonicera) and in a few Weigelas and allies; calyx pentamerous,
superior; corolla superior, gamopetalous, limb pentafid, small
in Viburnum and Sambucus to elongate-tubular or irregular-
infundibuliform in Honeysuckles; stamens five, inserted on tube
of corolla and alternating with corolla segments; 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 (Honeysuckle, Elder, etc.) from an
inferior ovary, several celled, occasionally becoming one-celled
with several to rarely one seed and drupaceous (Viburnum),
or fruit a capsule (Diervilla, Weigelia). Seeds albuminous.

THE DICOTYLEDONS | 607

OrrFIciAL DruG PART UsEp BoTANICAL ORIGIN HABITAT

Sambucus Flower (air- Sambucus canadensis) United States
dried) 1 Sambucus nigra Europe

Viburnum Viburnum Eastern and

Prunifolium Bark of root prunifolium st United
States
Viburnum Opulus Bark Viburnum Opulus United States
var. americanum and Canada

ORDER CUCURBITALES

CucuRBITACE2 OR GourD Fami_y.—Herbaceous, succulent,
very often annual (Colocynth, Pumpkin, etc.), more rarely perennial
(Bryonia, etc.), rarely shrubby
plants, the perennial and shrubby
forms perennating by swollen
roots, some of which are heavy
and tuberous. Stemsvery usually
grooved and ridged, often pro-
vided with roughened and barbed
hairs. ‘Tendrils are frequently
produced in the axils of leaves
from tendril axillary buds (Pump-
kin, Colocynth, Watermelon,
Cucumber, Bryony, etc.). Leaves
varying from entire, simple, usu-
ally deltoid to triangular through
stages of trilobate, pentalobate,
deeply palmatifid to palmatipar-
tite to seldom approaching com-
pound (Colocynth). Venation in
nearly all cases palmate. Leaves thin, herbaceous, much expand-
ed, often hairy. Vascular bundles of petioles, branches and
stems bicollateral. Inflorescence either of loose cymes or more
frequently racemes or spikes or entire axillary inflorescence may
become solitary axillary. Flowers pentamerous, very rarely
tetramerous, moncecious (Bryonia alba) or dicecious (Bryonta dioica) ;
sepals 5, gamosepalous, adnate to ovary; corolla of 5, rarely 4
gamopetalous petals varying in size and shape from small to large
campanulate (Pumpkin) or broadly cup-shaped (Cucumber),

Fic. 462.—Colocynth—Portion of
vine and whole fruit. (Sayre.)

Meg ey ee Mee Peay cee

608 PHARMACEUTICAL BOTANY

and in color from greenish-yellow to greenish-white to pure yellow —

to yellowish-white to white; stamens typically 5, epigynous, with
anthers either joined by pairs or synantherous; carpels usually 3,
rarely 4 or 5, fused to form a compound pistil with a short thick
style and lobed stigma; ovary inferior, one- to three-celled. Fruit
a pepo (a berry from an inferior ovary with thick skin). Seeds
flat and exalbuminous.

OrrFiciAL Druc PArt UsEepD BoTANICAL ORIGIN HABITAT
Bryonia Root ea alba E
et iad urope
Bryonia dioica
Colocynthis » Pulp of unripe —Ciitrullus Colocyn- Africa and Asia
but full grown __ this
fruit
UnorriciAL Druc
Pepo Seed Cucurbita Pepo Probably Tropi-
(cultivated varieties) cal America
Elaterinum Principle from Ecballium Elaterium Mediterranean
elaterium region
Watermelon Seed Seeds Citrullus vulgaris Southern Asia
Momordica (Balsam Fruit Momordica East Indies
apple) Balsamina

ORDER VALERIANALES

This order includes the Valerianacee or Valerian Family and
the Dipsacee or Teasel Family. Both are represented by herbs
with flowers having an inferior ovary containing a single ovule.

VALERIANACEZ OR VALERIAN Famity.—Herbaceous often
low succulent plants with creeping rhizomes, frequently strongly
scented and possessing stimulating properties. Leaves fre-
quently dimorphic; radical fascicled; cauline opposite; petiole
dilated, exstipulate. Inflorescence a raceme of dichasial 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, gamopetalous, 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. Fruit an akene from inferior

Se ee ee IT enn ee ea

Eee a en eee wae

,

THE DICOTYLEDONS 609
ovary crowned by a persistent, expanded, pappose calyx rudi-
ment. Seeds anatropous, exalbuminous.

OrriciAL DruG Part Usep BotANICAL ORIGIN HABITAT
Valeriana Rhizome and roots Valeriana officinalis Europe and Asia

Fic. 463.—Valeriana officinalis—Plant and rhizome. (Sapre.)

ORDER CAMPANALES (CAMPANULALES)

This order comprises the Bluebell Family (Campanulacee),
the Lobelia Family (Lobeliacee) and the Daisy Family (Com-
posite). Its outstanding characters are the epigynous corolla,

ee ee ne a ae eee oe

610 PHARMACEUTICAL BOTANY

usually 5 stamens, an inferior ovary and a three- to two-cleft
stigma.

CAMPANULACE2 OR BLUEBELL FAmity.—Herbs of annual
or more commonly 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, exstipulate. Inflorescence primitively a race-
mose 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, elon-
gate; stigmas as many as carpels. Fruit a capsule. Seeds
albuminous. The plants contain inulin.

LoBeLiaAcEz or LoBeuia F AMILY.—Herbs, with inulin and
latex contents, corresponding with Campanulacee in their vegeta-
tive parts, but differing from that group by having irregular
flowers (pale blue in Lobelia inflata), anthers always united into
a tube (synantherous) and pistil always bicarpellate with two-
celled ovary and bilobed or bilabiate stigma.

OrrIctaL Druc Parts Usep _— BoTANICAL ORIGIN HABITAT
Lobelia Leaves and tops Lobelia inflata United States and Canada

Composir# (AsTERACER®) or Darsy Famity.—Herbs_ of
annual or perennial habit, rarely shrubs or trees, and with
watery or milky juice. Inulin is present in cell sap of paren-
chyma. Leaves alternate, rarely opposite, simple to compound,
exstipulate. Inflorescence a capitulum or a raceme of capitula,
each capitulum surrounded by an involucre or protective whorl
of bracts, and composed of numerous small flowers called florets
that may be: (a) wholly regular, tubular and hermaphrodite
(Thistle, etc.); or (6) central florets as in (a), but marginals

THE DICOTYLEDONS 611

strap-shaped or ligulate and usually pistillate (Daisy, Dahlia,
etc.); or (¢) florets all ligulate and hermaphrodite (Dandelion,
Chicory, etc.); or (d) florets in part or in whole bilabiate (Muti-
sia, etc.). Flowers small (florets), closely crowded, pentamerous,
shaped as above, with ovary inferior and other floral parts supe-
rior. Sepals rudimentary, tooth-like (Sunflower), or reduced

Fic. 464.—Capitulum of a composite. Jerusalem artichoke (Helianthus tubero-
sus). 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.)

toa pappose or hairy rudiment above the ovary that is function-
less during flowering, but that expands in fruit as a hairy fruit
disseminator (Dandelion, Thistle, etc.); or sepals wholly
absorbed (Daisy). Petals synpetalous, tubular, ligulate OF
rarely bilabiate, greenish-yellow to white, or through pink-
crimson and purple to blue (Chicory). Stamens five, epi-
petalous, filaments distinct, anthers united into an upright
anther-box (so synantherous) into which pollen is shed before or

; |
Fic. 465.—Arctium minus or Com-
: mon Burdock, a source of the drug

Lappa or Burdock Root. (Reproduced
from U.S. Dept. Agric. Miscel. Pub. 77.)

Fic. 467.—Brauneria angustifolia,
or Narrow-leaved Purple Cone-flower.
(Reproduced from U. S. Dept. Asgric.
Miscel. Publ. 77.)

612 PHARMACEUTICAL BOTANY

Fic. 466.—Grindelia squarrosa or
Gum Plant, a native of the Plains and
Prairies from Saskatchewan to Mexico.
(Reproduced from U. S. Dept. Agric.
Miscel. Publ. 77.)

Fic. 468.—Tussilago Farfara or
Coltsfoot. (Reproduced from U. S.
Dept. Agric. Miscel. Publ. 77.)

THE DICOTYLEDONS 613

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

Fic. 469.—Matricaria Chamomilla, the German Chamomile. Branch and dis-
sected flowers. (Sapre.)

stigmatic hairs for pollen reception. Fruit an indehiscent
akene, often (Dandelion, Thistle) crowned by the pappose, calyx
rudiment. Seed single, exalbuminous.

The Composite is subdivided into two sub-groups or series,
namely the Tubuliflore and the Liguliflore. The TuBuLIFLOR&
include all those Composites whose flower heads contain either

614 PHARMACEUTICAL BOTANY

tubular florets only, as the Eupatoriums and Burdocks or
marginal ligulate florets and central tubular florets, as in the
Sunflower, Chamomiles, etc. The LicutirLor# include those
Composites whose flower heads possess ligulate florets only, as

i

y

y Oa pn

i My

eZ
\4

«3

\ \\ Low
AK
Wes

=

My

Fic. 470.—Chicory (Cichorium Intybus). 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.)

the Dandelions, Hawkweeds, Chicory, Lettuces, etc. They
contain a bitter or acrid milk juice.

OrriciAL DruG Part Usep BoTANICAL ORIGIN HABITAT
Arnica Flower head Arnica montana Europe
Matricaria Flower head Matricaria Europe and

Chamomilla western Asia
Calendula Ligulate florets Calendula officinalis Mediterranean
basin
Eupatorium Leaves and Eupatorium North America

flowering tops perfoliatum

‘ml

Grindelia

Taraxacum
Echinacea

Lappa

Santoninum

OrrFic1AL Druc

THE DICOTYLEDONS

PART Usep
Leaves and
flowering tops

Rhizome and roots

Rhizome and
roots

Root of first

year plant

Inner anhydride
of santoninic acid

UnorrFiciAL Druc

Anthemis
Arnice Radix

Pyrethri Flores

Pyrethrum

Inula

Senecio
Farfara
Absinthium -
(Wormwood)
Lactucarium
Santonica

Carthamus
(Safflower)
Achillea
(Yarrow)
Tanacetum
(Tansy)

Gnaphalium
(Everlasting)

Cichorium
(Chicory)
Oleum
Erigerontis
Flores Cyani
Solidago
Silphium
Vanilla Leaf
Swamproot

Blessed Thistle

Flower head

Rhizome and roots

Flower head
(unexpanded
or partly
expanded)

Root

a

Rhizome and roots

Entire plant
Leaves

Leaves and
flowering tops
Dried milk juice
Unexpanded
flower head
Tubular florets

Leaves and
flowering tops
Leaves and
flowering tops

Leaves and
flowering tops

Rhizome and
roots
Volatile oil

Florets
Herb
Root
Leaves”
Root
Herb

BotTANICAL ORIGIN

Grindelia camporum
Grindelia cuneifolia
Grindelia squarrosa
Leontodon Taraxacum
Brauneria pallida
Brauneria
angustifolia
Arctium Lappa and
Arctium minus
Artemisia Cina

Anthemis nobilis
Arnica montana
Chrysanthemum
cinerarizfolium
Chrysanthemum
roseum
Chrysanthemum
Marschallii
Anacyclus
Pyrethrum

Inula Helenium
Senecio aureus
Tussilago Farfara
Artemisia Absinthium
Lactuca virosa

Artemisia Cina

Carthamus
tinctorius
Achillea millefolium

Tanacetum vulgare
Gnaphalium
polycephalum
Anaphalis margaritacea
Cichorium Intybus

Erigeron canadensis

Centaurea Cyanus

Solidago odora
Silphium laciniatum

Trilisa odoratissima

Eupatorium purpureum
Cnicus benedictus

615

HABITAT

Western North
America

Europe

Central United
States

Europe and Asia

Turkestan

Europe
Europe
Dalmatia
Herzegovina

Western Asia

-

Northern Africa
and southern
Europe

Europe and Asia
United States
Europe

Europe, Asia and
Africa

Europe
Turkestan

India

Europe and Asia
Europe

North America

Asia
Europe

North America

Europe

E. United States
United States

S. United States
United States
Europe

CHAPTER XXIV
ECOLOGY

Eco.oey is that department of biology which deals with the
relations of plants and animals of various habitats to their
environmental conditions.

Ecology may be divided into two branches, autecology and
synecology. AUTECOLOGY deals with the structure and behavior
of the individual organism and its parts as related to environ-
ment, whereas synecology deals with plant communities as
related to soil, light, climate and other environmental factors.

Every living thing is a creature of circumstance, dominated
and controlled by heredity and environment. Its fundamental
structural and functional characteristics are determined by the
genes in its protoplasm but its characteristics become modified
by external factors of environment. In order to exist and keep
healthy it must adapt itself to the various factors of its sur-
roundings. The environmental factors having to do with the
existence and health of plants include soils and soil constituents,
air, moisture, light, range in temperature, gravity, and sur-

ounding animals and plants of other kinds. Various aspects of

most of them have been considered in preceding chapters.

PLANT Communitirs AND ASSOCIATIONS

An entire group of plants of similar habits occurring in a
common _ habitat constitutes what is termed a plant community.
A plant association is | a plant community having a definite com-
position 7.e., made up of a definite aggregation of plants deveiop-
ing under similar conditions. Thus, a desert where the cacti
and euphorbias form the dominant plants with a definite
assernblage of other plants growing with them would represent
a cactus-euphorbia desert association. Plant communities may be
classified either from the point of view of their order of develop-

ment, as based upon the principle of succession, or upon their
616

ECOLOGY 617 |

water relation. ‘The latter method appears to be simpler, and
will now be considered.

According, therefore, to the relation plant communities have
assumed in regard to water, they may be grouped as follows:
Hydrophytes or water plants.
Helophytes or marsh plants.
Halophytes or salt plants.
Xerophytes or desert plants.
Mesophytes or intermediate plants.
.. Tropophytes or alternate plants.

Fivbroravide —The effect of an aquatic environment on
the structure of water plants is most striking. The root systems
_are reduced both in length and number of branches. The
root hairs of those immersed in the water are absent. The
supportive action of the water is such that the fibrovascular
elements of the stems, which usually function both for support
and for the conduction of crude sap, are greatly reduced in size
and strength. The leaves, stems and roots possess large air-
spaces. ‘The mesophyll of the leaves is spongy and the chloro-
plasts motile. Stomata are entirely absent from leaves that are
submerged and only present on the upper surface of floating
ones, where they are nearly always open. Some of these plants
have broad floating leaves and dissected submerged ones, often
with thread-like divisions. ‘The submerged parts are devoid of
special protective walls, e.g., those containing cutin or suberin.
The cell sap has a low osmotic pressure. The submerged leaves
often absorb more water than the roots. The free floating
microscopic plants (blue-green algze, bacteria, diatoms, desmids,
etc.) form the plankton of our ponds, rivers and lakes. The free-
swiming higher plants (the plewston) comprise certain liverworts
like Riccia and Ricciocarpus, water-ferns and such seed plants as the
water-lettuce and water-hyacinth. The aquatic plants, includ-
ing the algae, mosses, and flowering plants which live attached to
rocks comprise the /ithophilous benthos. Another class of aquatic
plants (benthos) include those with true roots, which attach the
plant to the substratum, and at most possess floating leaves.
This type includes the water-lilies, the water-chestnut, the splatter
docks, the floating-heart and the pondweeds.

An wOn es

618 _ PHARMACEUTICAL BOTANY |

HeE.opuytes.—To this group belong plants typical to marshes.
A Marsh is an area with wet soil, wholly or partially covered with
water and with annual or perennial herbs (never shrubs and
trees) which are adjusted structurally to a mucky soil, lacking
the usual supplies of oxygen. These plants likewise show an
adjustment to a partial or periodical submergence. Like
hydrophytes, marsh plants are for the most part perennial.
They produce adventitious roots and possess horizontal rhizomes,
or runners, and frequently have air chambers in roots, stems and
leaves, so that they are adapted to meet the scarcity of air in wet
soils. They also show a striking development of erect chloro-
phyll-bearing organs in the shape of leaves, in the flags, and
stems, in the rushes.

The taller seed-like plants of the marsh-land, such as seed-
grass (Phragmites), the bur-reed (Sparganium), the cat-tails ( Typha),
the blue-flags (Jris), the sweet flag (Acorus calamus) and the
papyrus (Papyrus) form associations known as fresh-water marshes,
reed-marshes or fens. The channels or pools of water in among
these amphibious plants are filled with true aquatic plants.

_Hatopnytes.—The plants of this group live in a soil which is
rich in soluble salt, usually common salt (NaCl), and, on account
of the fact that the osmotic force of the root is nearly inadequate
to overcome that of the concentrated solution of the soil, the soil
to such plants is physiologically dry. _A halophyte, in fact, is one
form of xerophyte. The most striking feature among halophytes
is that they are nearly all succulent plants. The leaves of such
plants, for example, are thick, fleshy and more or less translucent.
They are Tich in concentrated cell sap by which they are able to
counteract the osmotic pull of the concentrated saline solution
of the soils in which they live. Anatomically, they are poor in
chlorophyll; the intercellular-air-spaces are small and_ the
palisade tissue is more abundant. Coatings of wax are found
and a hairy covering, although infrequent, sometimes occurs.
Coriaceous and glossy leaves, especially in tropical halophytes,
are noteworthy, while in many salt-loving plants the stomata
are sunken. Halophytes are found in our coastal salt marshes
and on saline tidal flats in temperate and tropical countries and
on the alkali flats of the interior of continents. Notable exam-

ECOLOGY 619

ples of these plants are the Salt Marsh Samphire, Salicornia
ambigua, the Mangroves (Rhizophora), the Goosefoots (Cheno-
podium), and the Bald Cypress (Taxodium).

XEROPHYTES.—The plants of this group, like the halophytes,
are adjusted to live in a soil which is physiologically dry. The
soil may owe this condition to its physical nature, such as porosity
(sand), or to the presence of humic acids, or by chemical action,
which inhibits the absorption of water. They are adapted to
meet the conditions of strongest transpiration and most precarious
water supply. ‘To meet such conditions of physiological drought,
the plants show various structural adaptations. In deserts,
where the atmospheric precipitations are less than a certain
limit, the plants acquire a xerophytic structure, such as succu-
lency, water storage tissue, associated frequently with mucilage,
lignified tissues, thick cuticle to the leaves, depressed stomata
(frequently in pits), reduced transpiration surfaces and thorns.
Mechanical tissues like wood and bast fibers attain their highest
development in these plants. Cacti, conifers, lichens, and the
century plants (Agave) are types of xerophytes, while many bog
plants like the cranberry and Labrador tea, with noc leaves,
are xerophytic.

MEsopnyTes.—These are plants that grow in soil of an inter-_
mediate character which is neither specially acid, cold or saline,
nor excessively wet or dry, but is sufficiently well supplied with
water and rich in the elements required for plant growth.
Plants which grow under such conditions do not have structures
by which transpiration is closely controlled. They have large
leaves frequently toothed and incised, with numerous stomata
__on lower or both surfaces, a thin, moderately cutinized epider-
mis, and small intercellular-air-spaces. ‘The leaves and stems
are usually of a fresh green color. ‘Typical of the mesophytes
are the grasses, the crop plants of field and garden, and most of
the annual and biennial herbs of temperate regions.

TROPOPHYTES.—This, term was first introduced by Schimper
in 1898 for land plants which have deciduous leaves and whose
conditions of life are, according to the season of the year, alter-
nately those of mesophytes and xerophytes. The mesophytic
condition is found in summer, when the trees, shrubs and peren-

620 PHARMACEUTICAL BOTANY

nial herbs, included in this group, are in full leafage, and when,
owing to the regular supply of rain during the growing season,
the soil is plentifully supplied with water to meet the demands of
these plants during the period of active transpiration. During
the winter they are xerophytes. The cold of winter freezes the

shade plant or umbrophyte, growing in sparsely shaded woodland.

water in the soil so that the transpiration is reduced to a minimum,
and this is associated with the fall of the leaves of the trees and
shrubs and the death of the overground parts of the perennial
herbs which spring up each year from their underground parts.
The vegetation of cold temperate regions is mainly tropophytic.

The deciduous trees and shrubs, also known as the broad-
leaved plants and the summer-green plants, form the principal

a ee

ECOLOGY 621

tropophytes. ‘The deciduous forests, which include the oaks,
the beeches, the ashes, the maples, the walnuts, the chestnuts,
cover a part of eastern and western China, central Europe
(England, France, Belgium, Germany) and eastern Australia,
and are coincident with the countries occupied by the most
civilized races of man, such as the Americans, Europeans,
Chinese and Japanese. ‘The cold temperate climatic conditions
which have determined the distribution of the forest trees have
been influential also in the development of the energetic races of
mankind.

PLANT SuccEssion.—In the development of vegetation, the
same region becomes successively occupied by different com-
munities of plants. This process is called plant succession.
There are two kinds of plant succession, (1) the Aydrarch which
begins in water and (2) the xerarch which may begin on bare
rock, rocky talus slopes, or other location where the soil is
extremely dry. As the scope of this book forbids little more than
mention of this topic, the student interested in it is referred to
works on plant ecology.

RELATION OF PLANTs TO ANIMALS

The great and fundamental réle of green plants in the world’s
economy is that of constructing highly complex organic com-
pounds for food out of simple, inorganic, raw materials. This
food, part and parcel of their very bodies, is eaten by animals
whose chief and contrasting réle is that of breaking down the
plant food into its simplest elements.

The raw materials which constitute the ordinary diet of
green plants represent salts of various metals, such as nitrates,
phosphates, sulfates, etc., the water which they absorb from the
soil, and carbon dioxide which they inhale from the air during
sunlight.

Carnivorous PLANTS.—There exist, however, about 500
species of green plants which, in addition to the common habits
of nutrition possessed by their relatives in the vegetable world,
have acquired the luxurious appetite for the flesh and blood of
animals. These exhibit a variety of devices for the allurement,
capture, imprisonment, digestion and absorption of their prey.

622 PHARMACEUTICAL BOTANY

Chief among these carnivorous plants are the sundews, fly-
traps, pitcher-plants, bladderworts and butterworts. In each
it is the leaves which have become modified for the purposes
indicated.

Dead
Organisms

“Bacterial .
; Decay...

Carbohydrates.
Plant proteins
Fats

Products

a

Fic, 472.—The carbon cycle. (After Gager.)

Intermediate
Decomposition

Tue SuNDEWS

The sundews are curious, small, bog plants that are mainly
members of the genus Drosera which comprises 84 species, occur-
ring in tropical, warm temperate and cool temperate regions of
both hemispheres. Each of these possesses a rosette of leaves
that arise from a greatly reduced root system. The shape of the
leaves varies depending upon the species, although it is fre-
quently rotund, spatulate or filiform. The whole upper surface of
the leaf-blade is covered with glandular tentacles often of a wine-
red color. The glandular heads of these are covered with a
viscid secretion that glistens in the sunlight like dewdrops, a
phenomenon that accounts for the common name assigned to the
members of this group. The tentacles in the central part of the
leaf-blade are short and erect and their stalks are green. Toward

the margin they become longer and more inclined outward and
their stalks are of a wine-red to purple color.

ECOLOGY 623

When examined under the microscope each tentacle shows a
slender pedicel terminating in a gland. A conducting bundle
containing spiral vessels and simple vascular cells is observed to
come off of a fibrovascular bundle in the leaf-blade and run

Fic. 473.—Drosera rotundifolia, the Round-leaved Sundew. (From Jenkins’ ‘Inter-
esting Neighbors.’’)

through the center of the pedicel to the gland. Upon entering
the gland it is enlarged and spread out into a number of tracheids.
These centrally placed tracheids are surrounded by a protective
sheath (endodermis) outside of which are two layers of secretory
(epidermal) cells with wine-red, granular contents, the outer
layer of which is palisade-like. These glands are commonly oval
excepting the extreme marginal ones which are considerably

624 PHARMACEUTICAL BOTANY

elongated. They secrete, absorb and are acted upon by various:
stimulants.

Darwin! has shown that when a small object is placed on the
tentacles in the center of the leaf these transmit an impulse to the
marginal tentacles. The nearer ones first respond and slowly
bend toward the center and then those farther away until finally
all become bent over the object. The time essential varies from
10 seconds to 5 or more hours depending upon the nature, con-
tents and size of the object, upon temperature and also upon the
age of the leaf. Again, if the glands are repeatedly touched or
brushed or if chemical substances are placed on these, the mar-
ginal tentacles curve inward. The bending part of each tentacle
is confined to a limited space near the base. Not only the ten-
tacles but the blade of the leaf becomes much incurved when any
strongly exciting substance is placed on the blade.

The time during which the tentacles and blade remain curved
over the object varies according to the temperature, character of
object, age, etc. Dr. Nitschke? found that during cold weather —
both the blades and tentacles re-expand within a shorter period
than when the weather is warm. Darwin* found that the ten-
tacles remain clasped for a much longer period over objects
which yield soluble nitrogenous matter than over those yielding
no such matter. After a period varying from 1 to 7 days the
tentacles and blade re-expand and are then ready to again
respond. It has been shown by the same authority that as soon
as the tentacles become inflected over an object yielding soluble
nitrogenous matter their glands pour out an increased amount
of secretion which becomes acid in nature. These glands, more-
over, continue to secrete as long as the tentacles remain closely
inflected.

In nature small insects catch sight of the glittering drops on
the tips of the reddish tentacles and, mistaking these for honey,
alight upon the leaf. They become instantly entangled by the
viscid glandular secretion. They try to stroke the viscid fluid off
of their legs but only besmear themselves more. Soon they

' Darwin: Insectivorous Plants, p. 9, 1884.
* Darwin: Insectivorous Plants, p. 13, 1884.
* Darwin: Insectivorous Plants, p- 13, 1884.

ECOLOGY 625

become covered with the sticky substance which occludes the
orifices of their breathing tubes (tracheae) and they perish in a
short time (one-quarter of an hour according to Dr. Nitschke)
from suffocation. The surrounding tentacles bend over the
insect’s body and clasp it on all sides. A digestive juice con-
taining a proteolytic enzyme of peptic character and an acid is
now poured out of the glands. This digests the flesh and blood
of the insects caught. The water-insoluble chitinous parts of the
body are left on the surface of the blade. If the insect be large,
the bending of the tentacles is augmented by the inflexion of the
whole surface of the leaf-blade which assumes a concave shape.
With the tentacles also curved over, the whole leaf simulates
a closed fist. The glands then absorb the soluble nitrogenous
material which is assimilated by the plant.

Among the insects caught by the sundews are gnats, flies, ants,
beetles, small butterflies and dragon flies. The last named are
captured by the cooperation of 2 or 3 adjacent leaves.

Tue Fry Traps

The fly traps comprise two species, viz.: the Venus Fly Trap
(Dionea muscipula) and the Submerged Dionzea (Aldrovanda
vesiculosa).

The Venus Fly Trap is restricted to damp localities of a coastal
plain strip extending for about 55 miles north and 45 miles south
of Wilmington, North Carolina, and nowhere over 15 to 20 miles
in width. an

_ Each plant shows a rosette of modified leaves which together
are rarely more than 6 or 7 inches across. From the center of
these arises a flower stalk bearing a cyme of white flowers which
open from April to June.

Each leaf consists of a winged petiole that is studded on both
surfaces with small, brown, stellate hairs. The petiole is trun-
cated in front and contracted to the midrib which suddenly
broadens out into a blade composed of 2 symmetrical halves that
can fold together along the line of the contractile and irritable
midrib region. Along the margin of each half of the blade are
long, stiff, non-irritable bristles which, on closure of the blade,
interlock with each other. On the center of the upper surface

626 PHARMACEUTICAL BOTANY

of each half of the blade are three spine-like, sensitive hairs
disposed as angles of a triangle. Each of these sensitive hairs
is composed of elongated cells whose protoplasmic contents show

Fic. 474.—Dionza muscipula, The Venus Fly Trap. (From Jenkins’ “Interesting
Neighbors,’’)

movement. At the base of each is a cylindrical mass of small
cells which permits the stiff hair to be bent over. These sensitive
hairs are highly irritable and capable of receiving and transmit-
ting a stimulus. Further, over the entire upper surface of

ECOLOGY 627

the blade are numerous, closely set, sessile, glandular hairs
which, after repeated irritation, secrete an acid digestive juice.
Each of these, as observed under a microscope in surface view,
consists of a rosette-arranged set of cells composed of 4 to 12 cell
radii filled with a crimson-claret pigment.

Each sensitive hair consists of a somewhat elongated structure
composed of 3 parts, viz., base, highly sensitive joint and insensi-
tive shaft. The base consists of large epidermal cells enclosing a
prolongation of mesophyll cells within, that can receive and
propagate a stimulus from the sensitive joint to the leaf interior.
The joint consists of elongated columnar cells that enclose
similar columnar mesophyll cells with soft, elastic walls. The
shaft cells are elongated, thick-walled and almost insensitive.

When a stimulus is applied to any sensitive hair, this affects
the joint cells and causes upsetting of turgidity and exudation of
liquids with a consequent contraction of the elastic cells. ‘The
action is propagated to the midrib region where the cells, by
contraction along the upper surface and expansion along the
lower, cause closure of the halves.

Macfarlane! has shown that when an insect alights on a leaf of
Dionea, one hair must neither be stimulated twice or 2 hairs on
the same leaf at slight intervals apart in order to cause closure.
The caught insect produces a summation of stimuli, and gradual
tightening of the lamina occurs under the repeated stimuli until
the halves become closely locked. The glandular hairs now pour
out an acid digestive secretion which digests the flesh and blood
of the insect’s body. The soluble nitrogenous substances are
then absorbed by the glands and assimilated.

Dr. J. S. Hepburn? found that the secretion of the leaves of
Dionga contained a protease which was active in the presence
of 0.2 per cent. hydrochloric acid. This enzyme, therefore,
resembles pepsin of the gastric juice of man, which also acts in a
0.2 per cent. hydrochloric acid medium.

Dr. J. M. Macfarlane, who with Dr. Canby and Charles
Darwin have separately investigated the structure and phys-
iological activities of this plant, reports that 2 touches one-fourth

1 Macfarlane: Contrib. Bot. Lab. U. of Pa., 1: 7-44, 1892.
? Hepburn: Jour. Franklin Inst., 194: 780, 1922.

a ne oe

628 PHARMACEUTICAL BOTANY

to one-third of a second apart will not produce closure of the
halves, though a second stimulus that is from 1 to 120 seconds
apart from the first will effect closure, also that it is not essential
for the stimulus to be on one hair, that a prick sets off the trap
at once due to liquid escaping from turgid cells and, further, that

all muscle stimulants, mineral acids and ammonia effect closure

of the trap.

Among the insects caught by Dionea are earwigs, millipedes,
flies, ants, wood-lice and dragonflies. The length of time
required for the digestion of their softer parts and the absorption
of the soluble products of digestion varies. During this time the
trap remains closed. The power of digestion possessed by
Dionea is more limited than that of Drosera, which has been
known to capture and digest many insects in a shorter period of
time.

Aldrovanda vesiculosa, which might well be called a ““Submerged
Dionza,” is a relative of the Venus Fly-Trap. It is a floating
aquatic plant found widely distributed in shallow ditches and
ponds over the old world from Europe to Australia but nowhere
abundant. This plant is entirely devoid of roots, but possesses a
slender stem which bears whorls of modified leaves at its nodes.
Like the Venus Fly Trap, each leaf shows differentiation into a
winged petiole ending in 5 narrow processes that connect with a
terminal, rounded, incurved blade, divided into equal halves by
a sensitive midrib, but the midrib projects beyond the summit of
the lamina as a bristle. Long, rigid, spiny bristles extend from
the petiole and are thought to prevent the approach of animals
unsuitable as prey. The margins of each half of the blade are
bent inward and their rims are studded with short conical teeth.
Projecting from the upper surface of the midrib and along a line
describing the inner third of the upper blade surface are a
number of sensitive hairs and short-stalked, disc-shaped glands,
while over the outer portion of the surface are to be noted a
number of scattered stellate hairs.

Larvz of aquatic insects and small species of Crustaceans
such as Cyclops, Daphnia and Cypris, swimming by, occasionally
brush against the sensitive hairs and the 2 halves of the blade
close together, just as in Dionga, and the animals are entrapped.

ECOLOGY 629

When they attempt to escape through the place where the
margins of the blade meet, they find the conical teeth prevent
their egress. ‘They die in the trap, and when the latter are forced
open and examined a couple of weeks later they only contain
the chitinous skeleta.

Tue PircHer PLANtTs

The pitcher plants are for the most part found in bogs.
Their main representatives belong to the genera Heliamphora,
Sarracena, Darlingtonia, Nepenthes and Cephalotus. The pitcher
itself represents the hollowed out midrib of the leaf in the first
four named genera and an inpouching of the leaf in the last ©
named genus.-

Heliamphora’ is represented by a single species (H. nutans)
which is only found on and around the base of Mount Roraima,
between British Guiana and Venezuela. Here it flourishes in
widespreading dense tufts in wet places where the grass is short.
It has a rosette of red-veined pitcher-leaves and delicate white
flowers raised high on red-tinted stems. Along the entire length
of the pitcher are 2 broad wings. The pitcher itself represent a
hollowed out midrib which is tubular in shape, becoming grad-
ually broader from base to mouth and ending in a small lid.
Over the entire outside of the leaf are nectar glands and upward
directed hairs. The nectar glands secrete a sweet fluid which
entices insects so that this surface can be termed the alluring
surface. Nectar glands also occur on the inner surface of the lid.
Next, the upper one-third to one-half of the inner surface of the
pitcher is covered with downwardly projecting hairs and nectar
glands. This surface can be called the attractive and conducting
surface. A smooth surface follows, which in turn is succeeded
by a detentive surface in the lower part of the pitcher, composed
of mostly smooth-walled cells, a few of which bear short hairs.

Sarracenia! is represented by 7 species all of which are con-
fined to Eastern North America except Sarracenia purpurea, which
extends west to Western Minnesota and West Central Canada.
_ Their pitchered leaves are of varying form and color design,

‘Macfarlane: Sarraceniacee in Engler, Das Pflanzenreich, 34 Heft, (iv, 110)
39 pp., 1908.

630 PHARMACEUTICAL BOTANY

depending upon the species. Frequently they are long funnel-
shaped, tubular or vase-shaped. The color of every species
varies with its age as well as extent to which it is exposed to

Fic. 475.—Sarracenia purpurea, a pitcher plant from a northern bog. (From Jenkins’
“Interesting Neighbors.)

bright sunshine. In some respects the form of pitchers resembles 7
Heliamphora, but there is only a single wing present and further,
the lid is considerably larger. * : | -

In all we find the pitchers showing an alluring outer surface :
covered with nectar glands, but devoid of the upwardly directed

ECOLOGY 631

hairs seen in Feliamphora, an attractive inner lid surface, marked
with numerous downwardly directed hairs and nectar glands
which lead the insect toward the next or conducting surface.
This is either nearly smooth (S. purpurea) or presenting delicate,
downwardly projecting processes and on which the insect loses its
foothold, also (excepting S. purpurea, which next shows an inter-
vening glandular zone) a lower deéntentive zone, in the bottom of

Fic. 476.—A colony of Sarracenia purpurea growing along the border of a pond in
Burlington County, New Jersey. Chamecyparis in background.

the pitcher, which contains a watery fluid and whose surface,
Save its lowest portion, is marked by the presence of many elon-
gated, downwardly directed, thick-walled hairs, which prevent
the ascent of insects that have fallen into this zone from above.
Glands are also present on the upper part of the detentive surface.

Darlingtonia' is represented by a single species, Darlingtonia
californica, which grows in mountain swamps and borders of
small streams at altitudes of 1000 to 6000 feet from Plumas
county in the Sierras of California northward to Jackson and

! Macfarlane: “‘Sarraceniacez,” in Engler, Das Pflanzenreich, Heft (iv, 110),
39 pp., 1908.

Gon: _ PHARMACEUTICAL BOTANY

Josephine counties in Oregon. Its underground rhizome gives
rise to pitchered leaves which are from 1} to nearly 3 feet in
height and spirally twisted in about a half revolution. Each

E
:
i

Fic. 477.—Sarracenia flava, a pitcher plant from a North Carolina bog. (From
: Jenkins’ “ Interesting Neighbors.”)

pitcher expands near the summit into an inflated hod which
exhibits a circular opening up to an inch in diameter on the
under side. The dome of the hood is spotted with large, thin,
translucent areas. A wing extends along the pitcher from

ECOLOGY 633

rhizome to its orifice. At the upper and outer edge of the open-
ing is a moustache-shaped appendage which possesses stiff hairs
all of which point toward the pitcher opening. ‘The color of
the pitcher is green, blotched esa ;
with red and yellow. Within
and about the opening and on
the bi-lobed appendage are
alluring glands which secrete a
nectar attractive to insects.
The nectar is also secreted* by
glands along the wing. An
insect creeping over the exterior
of the pitcher is enticed to the
nectar along the wing which it
follows up to the orifice where
the honey is sweetest. Flying
insects are attracted to the
pitcher by its mottled and
colored features. They alight
on one of the flaps of the appen-
dage and follow its curve which
narrows to the orifice. Sharp
bristles in the path pointing
toward the orifice make it the
natural direction for the insect
to travel. Upon reaching the
end of the path, it is tempted
farther by honey glands within
the opening of the pitcher
which it next visits. When eae
satisfied and ready toleave, the yg. 478.—Darlington’s Pitcher Plant
translucent areas on the hood (Darlingtonia Californica) from mountains
above, like illuminated win- im California. Fram fanns " Ealereseens
pee Neighbors.”

dows, entice it from the open-

ing by which it entered. The insect sees no means of escape,
nothing but hairs on the inner surface of the pitcher pointing
downward and which direct it to the dark detentive pit below,
where it drowns in the watery secretion.

634 PHARMACEUTICAL BOTANY

By far the most beautiful and gracefully shaped pitchers are
those belonging to the genus Nepenthes which is represented by
about 68 species that are mainly indigenous to the East Indies
with headquarters in North Borneo and with Mount Kina Balu
as a center. Many of the plants are continuously exposed to a
moist dripping atmosphere.

Fic. 479.—Nepenthes Rajah. (From St. John’s “Life in the Foresta of the Far East.’’)

Each of the modified, pitchered leaves shows a petiole (usu-
ally winged) which widens into an expanded blade, the midrib of
which is continued as an elongated tendril which expands into a
terminal winged pitcher. The pitchers show a large variety of
shapes and color designs depending upon the species. Fre-
quently they are tubular, goblet or cornucopia shaped. Each
pitcher has a hinged lid which varies from small elliptic to large
heart- or kidney-shaped. The tendril is sensitive to contact
stimuli and often winds about a limb of a tree. The pitchers,

partly filled with a viscid watery secretion, either hang suspended
in the air from the tendril or rest on the ground.

ECOLOGY 635

One of the largest pitchers found in this group is that of
Nepenthes Rajah, a species found on Mount Kina Balu along the
north coast of Borneo at an altitude of 5000 feet. It has leaves
which, exclusive of petioles, are 18 inches long, and the pitchered
portion is 6 inches in diameter and 12 inches in length with a
circumference of 19 inches. The lid is 10 inches long and 8
inches broad. ‘The pitchers rest on the ground in a circle.
The color of the pitchers is deep purple. The color of the lid
portion is lavender, shading to green at the edges. One pitcher
noted by Spenser St. John’ held 4 pint bottles.

The lower surface of the petiole, lamina and the whole
external surface of the tendril and pitcher are covered with
alluring nectar glands. ‘These alluring glands also occur along
the stem. Their secretion entices insects to the pitcher mouth or
lid. Around the mouths of the pitchers of nearly all of the
species is a corrugated rim on which are parallel ridges, which are
usually extended as teeth that are inclined downward into the
pitcher cavity. The surface of this rim is smooth and polished,
so affording a poor foothold for insects. Around this rim, in the
region of the teeth, are the openings of deeply sunken, marginal
glands which pour forth a very sweet nectar, attractive to insects.
The inner surface of the lid is usually dotted with attractive
glands. :

In many species of Nepenthes, the entire inner surface of the
pitcher cavity constitutes the detentive surface. Itislined witha
smooth epidermis that is uniformly studded with glands which
secrete a digestive juice containing a proteolytic ferment and
absorb the soluble products of digestion. In many others the
upper surface of the cavity is smooth, forming a conducting
surface, while the lower one-half or one-third is alone detentive.
Insects lose their foothold on the conducting surface and fall
into the watery secretion at the bottom of the pitcher.

Cephalotus has but one species, C. follicularis, which is native to
swamps of South West Australia. It has short creeping rhizomes
which produce annually one set of 5 foliage leaves and later a set ~
of pitcher leaves which rest on damp soil. The pitchers exter-
nally are equipped with winged ridges which provide a handy

1 St. John, S.: Life in the Forests of the Far East, 1: 334, 1863.

Be Re I Se EE eine aE eT eee i Rh ORE tig eid EE OTC eE COMM pg RSP Reh ae

636 PHARMACEUTICAL BOTANY

means of ascent to the mouth for creeping animals, while their
lids are half-closed and mottled with white areas and purple
veins and are often mistaken for flowers by high flying insects.
In minute details of structure the pitchers are surprisingly like
those of Sarracenia and Nepenthes. ‘The exterior of the stalk is
studded with alluring glands. The margin of the pitcher is
corrugated and inflexed, and all over this area and exterior of
corrugation as well as on the inner lid surface are an abundance
of honey glands (attractive surface). Just inside of the margin
is a circular shelf-like ingrowth consisting of downward-directed
cells which resemble the conductive surface of Sarracenia. The
remainder of the inner surface constitutes the detentive surface.
At the bottom of the pitcher on either side are two beautiful
crimson to crimson-claret patches. ‘These have sunk in them
huge, many-celled glands which have a fibrovascular bundle
terminating in their base. The glands secrete an acid digestive
juice which partly dissolves the animal captives. A_ viscid
secretion partly fills the pitcher cavity. ;

As to relations of the pitchers of these several types of curiou
plants to animals, it may be said that insects, spiders, etc., are
attracted to the plants partly by the brilliant colors of their
pitchers and partly by the nectar drops exuded by honey glands.
They make their way to the exterior of the lid (or appendage in
Darlingtonia) where the secretion along the outer margin is
plentiful. From here they move to the inner surface of the lid
and sip the honey. Tempted further by the sight of nectar
ahead and often by hairs pointing downward, they step inside
the pitcher on to the conducting surface. Upon reaching this
surface they waver, slip off and fall into the lower part of the
pitcher which is the water holding region. They make numerous
attempts to escape but find exit impossible on account of the
downward-projecting hairs. They drown in the liquid and.
their bodies are either decomposed or digested by a digestive
juice secreted by glands lining the lower portion of the pitcher
cavity.

The animals caught by these pitcher plants are various.
While insects constitute their chief prey, slugs, spiders and rodents
have been recorded by observers as having been captured by a

ECOLOGY 637

number of them. Burbidge’ observed several Nepenthes in
North Borneo visited by a small rodent, which, while perched on
the margins of the pitchers bends its head and neck and scoops
out the caught insects and devours them. The same writer
states that if it attempted such action with WN. bicalcarata, the

Fic. 480.—Nepenthes Edwardsiana. (From St. John’s “Life in the Forests of the Far
East.’’)

two sharp spurs with which the pitcher of this species is provided

catch it by the neck and tumble it into the pitcher.
Macfarlane? has carefully observed the relations of Mepenthes

to animals. He states in part that “running insects such as ants

and cockroaches are their principal prey. Cockroaches run up

the stem and may pause to sip nectar from the alluring stem

glands. Reaching the base of the leaf, they may pass along it,
* Burbidge: Gard. of Sun, pp. 40-344, 1880.

* Macfarlane: Nepenthacez, in Engler, Das Pflanzenreich, 36 Heft., 88 pp.,
1908,

638 PHARMACEUTICAL BOTANY

attracted by the presence of honey drops there. ‘They almost
invariably run along its under side to shelter themselves from
enemies and the hotsun. Moving on restlessly and sipping from
nectar glands as they advance, they reach the tendril. The
ventral wings and the areas between are more beset with alluring
glands than is the outer part of the pitcher and along this they
often run till they reach the orifice or lid. ‘The lid glands of the
inner surface prove a greater attraction but their secretion does
not compare with the marginal glands. Straining to reach the
orifices of the glands, the insect visitors often overreach them-
selves after a few efforts and tumble into the pitcher cavity, and in
rare cases is escape again possible.”

Mr. Low who accompanied Spenser St. John on his travels in
North Borneo found a drowned rat in one of the pitchers of
Nepenthes Rajah.'

The character of the enzymes found in the pitcher secretions
has been investigated by Hepburn.” ‘This author found a tryptic
enzyme in Nepenthes which was active in a slightly acid medium.
No proteolytic enzyme was found by him in the secretions of
Darlingtonia while the Sarracenias he investigated were found to
possess proteolytic enzymes. The protease in the pitcher liquor
of Sarracenia flava and S. minor acts best in 0.2 per cent. hydrochloric
acid while that present in S. purpurea, S. rubra, S. Sledgei and S.
Drummondii acts best in an alkaline solution containing 0.5 per
cent. or less of sodium carbonate. Dakin* found a protease in
the pitcher liquor of Cephalotus.

Tue BLADDERWORTS

The bladderworts belong to the genus Utricularia. They are
aquatic plants which are found in pools and bogs of various parts
of the world with their centers in S. America and the East
Indies. .

The plants are rootless and according to the season of the year
sink to, the bottom or rise to near the water surface. They have
finely dissected submerged leaves, some of which are specialized

Ree John, S.: Life in the Forests of the Far East, 1: 335-336, 1863.
: Hepburn, St. John and Jones, Jour. Frank. Inst., 189: 152, 1920.
Dakin: Jour. and Proc. Roy. Soc. W. Australia, 4: 37-53, 1919.

ECOLOGY 639

as bladder-like traps which allow the entrance but not the escape
of small animals. Each of the traps represents a greatly
modified, inflated leaf of vesicular form and usually about 1% to

1g inch in diameter. From the margin of its opening project

:

Fic. 481.—A Bladderwort (Ultricularia Grafiana). Note the submerged stems
bearing modified, bladder-like leaves. Small animals become entrapped in
these bladders where they are decomposed and the water-soluble portions of their
remains are absorbed as nourishment by the plant. In the lower right-hand cor-
ner is a bladder-like leaf of another species (Utricularia neglecta) shown in section.
The entrance to the bladder is protected by numerous hairs and by a trap door

(t). Drawing based upon Kerner. (Gager.)

stiff bristles. There is an upper and lower lip present. The
lower lip is greatly thickened and provided with a cushion-like
process that extends into the bladder. From the upper lip is
suspended a thin elastic valve, the free edge of which rests upon
the inner surface of the lower lip cushion and closes the opening.

640 PHARMACEUTICAL BOTANY

Small animals, including crustaceans like Cyclops and Daphna,
as well as larvae of small insects and worms, enter the bladders
either because they seek a sheltered harbor for a time from larger
prey or because they expect to find food within the bladder. In
entering they have to press upon the valve and push it back. As
soon as they stop pressing upon the valve, the elastic character of
the valve causes it to close by bringing it back on the under lip
cushion. ‘The entrapped animals struggle to escape by pressing
themselves against the valve, but their efforts are in vain, for it is
impossible for the prisoners to force the valve outwardly over the
. cushion, and they die in time either from starvation or suffoca-
tion. Their bodies decompose and the water-soluble products of
decay are absorbed by quadrifid cells lining the entire inner wall
of the bladder.

Tue BurreRrworts 4

The Butterworts comprise an interesting group of flesh-eat-
ing plants which are placed in the genus Pinguicula. About 40
species exist of which probably the best known is Pinguicula vul-
garts, a member of the Uftriculariacee. which occurs in bogs or
other damp places generally in mountainous districts of northern
North America and Europe. Each plant exhibits a root system
of from about 5 to 16 short, submerged, unbranched roots from
which arises a rosette of oblong-ovate, yellowish-green leaves, the
younger central ones being concave and more or less erect, the
older marginal ones being flat or convex with their lower surfaces
resting upon the moist ground. From the center of the leaf
rosette there arises a slender scape bearing a single flower of
violet blue hue that is spurred in its corolla portion.

Alike with other investigated species of Pinguicula, the leaves
of this plant have somewhat upturned margins which give them
the form of a broad trough, the upper surface of which is covered
with a viscid secretion. Microscopical examination of this sur-
face discloses 2 types of glands (excepting along the margin), the
first being of toadstool shape and consisting of a cylindrical stalk
bearing a disc-shaped head of 8 to 16 cells arranged radially,
the second consisting of a shorter stalk bearing an 8-celled head.

ee a ee

ECOLOGY 641

Darwin' and others have experimented with the leaves on
living plants of several Butterworts and found that drops of water,
gum or sugar, and scratching of the surface of these leaves pro-
duced no response, that insoluble solid bodies such as grains of
sand and powdered glass allowed to remain for some time caused
slight incurving of the margins and a slight increase in the quan-
tity of mucilage secreted by the glands, but that when nitrogenous
bodies such as bits of cartilage, meat, clotted blood, etc., are
placed near the margin in contact with the glands they are excited
and pour forth abundant mucilage and an acid secretion.. More-
over, prolonged contact with these
bodies caused a marked incurving of the
margin which pushed them slowly
toward the middle of the leaf where
more glands are present and so increased
secretion. This acid secretion com-
pletely dissolved these substances which
were later absorbed by the leaf.

When small insects alight from the
air upon the leaf of the butterwort, they
become fastened by the mucilage and in
struggling to extricate themselves only
become deeper enmeshed by it. Within
a short time the acid juice, secreted by
the glands as a result of the stimuli, |
digests their bodies and all excepting
their hard, indigestible outer skeleta — P16. 482-—Pinguicula aul-
: garis, a butterwort. An ex-
(exoskeleta) is absorbed. tract of this plant containing

If small creeping insects come upon a proteolytic enzyme is used
the margin of the leaf they stimulate the in certain sedative expecto-

: rant compounds. (After Le-
edge to curl over and the marginal aie ad Dae

glands to secrete. While the insect is

enmeshed within the adhesive mucilage, it is slowly pushed by
the curling up of the edges into the middle of the leaf where the
acid secretion glands are most abundant. In about 24 hours the
softer parts of its body are digested and absorbed and the leaf
expands to its normal condition.

1 Darwin: Insectiv. Plants, pp. 371-390, 1899.

PHARMACEUTICAL BO TANT

SIGNIFICANCE OF THE CARNIVOROUS Hapsit

In all of the animal-eating plants which have been investi-
gated structurally and chemically, sufficient evidence has been
recorded to warrant the conclusion that these forms have devel-
oped their peculiar habit and correlated structures with the
object of providing adequately for their nutrition.

CHAPTER XXV

GENETICS AND EVOLUTION

GENETICS is the science which deals with the coming into.

being _ of organisms. It deals with heredity, variation, sex
determination, and the origin of individual plants and animals.

Herepiry is the faithful transmission of characters from
parent to offspring. While similar to genetics it is not entirely
the same, since the heredity complex, representing what the
_ offspring receives at the time of fertilization, may be changed by
environment. Genetics deals with both the parental complex
and the environmental “complex. “Modern genetics is based
upon the law that the chromatin material of germ plasm is
inherited.

All organisms are described in the literature in terms of
characters. These are the units employed to distinguish each
individual, variety or species from another. Characters are
evident in the outlines of leaves, color of leaves, size of seeds or
plants, etc., and are the result of hereditary material acquired
from the parent, modified by environment. They are mainly
hereditary. Hereditary substance is.chromatin material in the

nucleus of the cell.

MENDEL’s LAWS OF ALTERNATIVE INHERITANCE

The methods of heredity which have been confirmed in the
main by the experiments and observations of many biologists
were discovered by Gregor Johann Mendel (1822-1884), an

Austro-Silesian monk of Briinn, Austria.t_ Mendel was the first

to determine that the genes or “unit characters,” as he ‘termed

them, can separate from one another and recombine in such a

way that the characters which they represent appear to act as

_ Separate units that can often be removed from one strain and
- introduced into another.

1 Although Mendel’s Laws were published in 1866, they were disregarded until

1900 when DeVries unearthed them and brought them to the attention of or.
643

Sey cnn eee

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4
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644 PHARMACEUTICAL BOTANY

Mendel experimented with the hereditary characters of
various kinds of garden peas because these were the best materials
he could find to test out his ideas and obtain light on the prob-
lems of heredity and variation in which he was so keenly
interested. Hestudied seven pairs of constant differentiating char-
acters in the peas he worked with, namely, tall vs. dwarf stems;
round (smooth) vs. wrinkled (angular) seeds; yellow vs. green
seeds; colored flowers and seed coats vs. white flowers and seed
coats; parchmented vs. non-
parchmented pods; green vs.
bright canary yellow, unripe
| pods; and normal round stems
| with axillary flowers vs. fasci-
ated (flattened) stems with ter-
j} minal or umbellate flowers.
The members of such pairs of
contrasting (alternative) char-
| acters were called allelomorphs
by Bateson. Each member of
two different strains of plants
which can be crossed, as for
example, a tall stemmed pea
with a dwarf stemmed pea, is
Fic. 483.—Gregor Johann Mendel at the -said to be allelomorphic to the

ee other. :

Mendel’s Laws embody the following principles:

1. Tue Princiete or Unrr CHaracters.—The inheritance of

THE PURITY OF THE GERM CeLis (GaMETES).—Every germ cell is
pure with respect to any given unit character. In the germ cells _

i YT o ” o

GENETICS AND EVOLUTION 645

of hybrids there occurs a separation of the determiners of con-
trasting characters resulting in the production of different kinds
of germ cells, each of which is pure in respect to any given char-
acter. ‘This principle was later called “the ‘Splitting | of hybrids”
by DeVries.

Probably the most outstanding principle in the Mendelian
Law of heredity is the fact that the heritable characteristics of

organisms are independent one of another and may be inherited

_independently, The determining substances of these character-

istics must, accordingly, occur in the germ cells as independent,
self perpetuating particles. Such substances are called genes, or
factors or unit characters.

STRUCTURE OF THE CHROMOsOME.—In the resting nucleus the
nuclear reticulum consists of chromatin threads. These threads
as often observed are longitudinally double. Each unit of the
double thread appears as a chain of beads and consists of a
linear series of granules called the chromomeres and these in turn of
one or more.smaller particles called chromioles. ‘The chromomeres
lie opposite each other forming pairs. The determiners within
the germ cells are believed to occupy the same arrangement, so
that chromomeres or their components may represent genes.

MALE AND FEMALE CHROMOSOMES.—Every sexually produced
organism is a double being. Every cell of a plant or animal is
double, one half having been derived from the male sexual cell
and the other half from the female sexual cell. Each of these
cells contains two sets of chromosomes, one set from the sperma-
tozoon and one set from the egg. ‘These two sets of chromosomes
contain the genes derived from both parents.

During the maturation of the germ cells the chromosomes
become unravelled. Each paternal chromosome is matched
by a homologous chromosome of maternal origin. During
maturation homologous chromosomes and also their chromo-
meres lie side by side in pairs. At the time of reduction of the
chromosomes each pair of chromosomes (maternal and paternal)
appear as two parallel strings of beads, the beads representing
chromomeres and containing the genes.

DisTRIBUTION OF CHROMOSOMES DURING MATURATION AND
FERTILIZATION.—Maturation involves the reduction in the

646 PHARMACEUTICAL BOTANY

10

Fic. 484.—Diagrams illustrating the behavior of chromosomes during the reduc-
tion division which occurs at the time of spore formation in ferns and in most other
plants which have sexual reproduction. The maternal chromosomes are shown in
black, the paternal in white. Homologous chromosomes (corresponding maternal
and paternal chromosomes) are of corresponding length. As a result of the process
here shown, the spore mother cell (in the resting condition in 1) gives rise to a tetrad
or group of 4 spores as shown in 10, The spore mother cell nucleus (1) has the
diploid chromosome number (in this case 6) which is characteristic of the cells of the
sporphyte. These six consist of 3 pairs of homologous chromosomes, one of each
pair having come from the male gamete and one from the female gamete in the last
fusion of gametes. Note that in 2 the three pairs with the chromosomes of each
pair in close contact are lying within the nuclear membrane. In 3 the paired
chromosomes have shortened and have become closely “‘knotted” whereas in 4 they
have become “unraveled” and have undergone further shortening. In 5 the mem-
bers of each pair have separated but are still close to one another. In 6 the nuclear
membrane and the nucleolus have disappeared and the six chromosomes, each
showing evidence of a later lengthwise splitting, have been separated into two groups
of three. This is the actual reduction division, Note that in 7, only one chromosome of
each homologous pair is present in each of the resulting nuclei which are therefore
different in the chromosome “stock.” Each of these nuclei now undergoes a second
or homotypic division (8) with a splitting of each of the chromosomes, so that there
are two kinds of nuclei in the tetrads (shown in 9), two having the same chromosomes
as the upper nucleus in 7 and two having the same chromosomes as the lower nucleus

shown in 7. (From ‘A Text Book of General sy ‘ail Rebicus by peril
sion of the Publishers, J. Wiley @ kg me) Botany” by Holman and Robbins by per

GENETICS AND EVOLUTION | 647

numper of chromosomes to one-half that of the somatic cells.
In the primary spermatocyte and primary odcyte of animals at
synapsis the chromosomes unite in pairs. One member of each
pair was derived from the mother, the other from the father.
When division occurs, the chromosomes do not divide, but entire
chromosomes separate and are distributed in pairs to two daugh-
ter cells. At the division resulting in spermatids and mature
egg, the pairs separate and each daughter cell receives univalent
chromosomes. A similar condition prevails during the division
of the mother cells of pollen grains and embryo sacs in plants.
Each spermatozoon and ovum is thus provided with one-half the
number of chromosomes found in the somatic cells or primordial
germ cells.

DIsTRIBUTION OF GENES DURING MATURATION AND FERTILIZA-
TION.
of homologous chromomeres. Since the homologous chromo-
somes carry the genes and the genes are also paired, the genes will
be distributed as were the chromosomes.

ALLELOMORPHS.—Genes that occupy corresponding positions

_in homologous chromosomes are called allelomorphs. ‘The mem-

bers of a pair of allelomorphs may be the same or they may differ;
if they are the same, the individual bearing them is said to be
homozygous as far as the character is concerned that is controlled
by the particular genes; if they are different, the individual is said
to be heterozygous. _
DomiNANTs AND ReceEsstvEs.—In a heterozygote the genes
are of two kinds, dominants and_recessives. A gene is said to be
dominant when the character. it represents appears in the hetero-_.

zygote; its allelomorph in such a case is said to be recessive.

In _ peas, tallness is dominant, dwarfness recessive; in the sun- |

flower, the branched habit is dominant and the unbranched habit
is recessive. The result of dominance and recéessiveness in
allelomorphic genes is that heterozygotic individuals do not
exhibit all that they inherit from their parents, since some of the
genes are recessive.

GENOTYPE AND PHENOTYPE.—The term genotype is used to
express the entire genetic make-up of an organism. Individuals
having a similar eine constitution are said to be of the same

Tae een ye Nae eee eee ee

648 PHARMACEUTICAL BOTANY

genotype. ‘The term phenotype is used to describe that part of the
inheritance that appears in the individual. Forms alike in body
characteristics irrespective of their genetic constitution are of the

Mononysrips.—A monohybrid is an individual whose
parents differ with respect to one character. ee

Fic. 485.—Diagram illustrating the results obtained from crossing a variety

of garden pea breeding true to smooth cotyledons with one breeding true to wrinkled
cotyledons. (Gager.)

RESULTS OF CROssING INDIVIDUALS WITH ONE PAIR
OF CONTRASTING CHARACTERS

Mendel first made sure the characters with which he was to
work were constant for certain varieties and then began crossing
one variety with another which differed from each other in some
of the previously mentioned characters, studying the offspring of
several successive generations with respect to the behavior ot the
contrasting characters.

In every instance he found that the plants that developed
from the cross involving a single pair of alternative characters

GENETICS AND EVOLUTION 649

showed only one of the characters occurring in the parental
plants. The obvious character he termed “dominant,” the
hidden or latent character, “recessive.”? In one of his experi-
ments, he crossed tall pea plants with dwarf pea plants by trans-
ferring the pollen from the anthers of the tall plant to the stigma
of a dwarf plant; the seeds resulting grew to be tall plants like the
tall parent. ‘These offspring plants represented the first hybrid
generation called the first filial generation and are represented by
the symbol F;. Accordingly, Mendel designated tallness the
dominant character and dwarfness the recessive character. -He
next permitted the tall Fi: hybrids to be self fertilized and their.
offspring called the second filial generation (F2) were found to be
tall and dwarf in the ratio of three tall to one dwarf. With seed
obtained from self fertilized /, plants he grew an F3 generation.
He found that all the dwarfs (recessives) of the F2 when so inbred,
gave only dwarfs which bred true indefinitely, while all the tall
plants of the F, generation, when inbred, proved to be of two
_kinds, one-third pure dominant talls which bred true indefinitely,
and two-thirds hybrids similar in number of kinds and ratio to
that obtained by growing seed of F; plants, 7.e., three dominants to
one recessive. ‘These results may be diagrammatically repre-
sented as follows:

PARENTS TxD
F, (first filial hybrid generation) rae (all tall)

F, (second filial generation) 1 TT: 27(D): DD (3 tallto 1 dwarf)
| ;
F; (third filial generation) All 7 37 1D All D

When there is only one pair of contrasting characters usually
but two types of offspring occur in the Fz generation, dominants
and recessives.

REsuLts oF Crossinc INDIVIDUALS wiTH Two Pairs
oF CONTRASTING CHARACTERS

Dihybrids are individuals whose parents differ in respect to
two pairs of contrasting characters. When Mendel set out to
prove the phenomenon of a 3 to 1 ratio in the number of domi-

sis Cat igh iti pat Bi

PES. 2c oe ape 8 ee

650 PHARMACEUTICAL BOTANY

nant characters and recessive characters appearing in the
progeny of crosses between plants examined for one set of con-
trasting characters such as tallness and dwarfness, he found that
by the same method experiments could be carried on in which
two sets of contrasting characters would be involved. His
variety of Pea plants that differed in one set of contrasting char-
acters, when crossed, produced monohybrids or factors repre-
sented by a suitable symbol such as Dd (Tallness D, Dwarfness
d). Two pairs of contrasting characters such as a variety of Pea
plant which Mendel found to produce yellow, wrinkled seeds
and another plant producing the characters for smooth, green
seeds, when cross-pollinated, developed spores which gave rise to
four kinds of sperm and four kinds of eggs and not the character-
istic two kinds of sperm and eggs as of the monohybrids. This
condition of a progeny in which there are two separate con-
trasted characters is called a Dihybrid. In the cross of a variety
of Pea plants having green color, smooth surface seeds with
another variety having yellow color, wrinkled surface seeds the ©
following progeny appears in the first filial generation: Factors for

smooth (indicated by S$) and yellow (indicated by Y) are domi-
nant; whereas factors for wrinkled (indicated by s) and green

(indicated by y) are recessive. The F; diagrammatically is:

sex eT
ar
SsYy (all smooth and yellow)

In the second filial generation or Ff, after self-fertilizing the
progeny of the first filial generation or F\, it is then noted that
there are four possible kinds of male gametes and four possible
kinds of female gametes. During mitosis, the male gametes
(indicated by o*) unite with the female gametes (indicated by @ )
and there is a union of four different male characters (indicated
by $,s,Y%y) with four different female’ characters (indicated by
S,5,Yy). The multiple of such a union of gametes is sixteen, and
indicates that there wil] arise an F, generation of zygotes with
sixteen dihybrid characters in a definite proportion. The
actual composition of the Ff, zygotes of this dihybrid may be
represented by the following checkerboard diagram:

GENETICS AND EVOLUTION 651

ef e
SsVYyXSsVy
Male gametes— SY Sy sf sy

S| SSTY- | ESV | SEX 1- SeXy
ila Sy | SSYy | SSpy | SsYy | Ssyy
—— Eh! MYX Ss Vy ssY¥Y ss Vy

SsYVy

Ssyy ssYy ssp

Fr, Generation (Showing progeny with a 9-3—3-1 ratio)

In the checkerboard diagram there are evident homozygous
(like) characters and heterozygous (unlike) characters. Any
zygote with one or both characters for dominant (SS, YY or Ss,
Yy) will have the appearance of smooth surface and yellow color.
Zygotes with one dominant character in the dihybrid but with
2 or 3 other recessives (Ssyy or SSyy) will appear with smooth
surface and green color. Likewise, zygotes homozygous for one
contrasting character, if that character is recessive, will appear
recessive for that particular character regardless of how the
second contrasting character appears. For example, if a zygote
has the factors for ssYY, the character ss is homozygous for
wrinkled surface, a recessive character, and homozygous for _
yellow color, a dominant character; the dihybrid is, then, yellow
and wrinkled. If the dihybrid appears with characters ssYy, the
character is still homozygous for wrinkled surface but heterozy-
gous for yellow color; it will then appear also yellow and
wrinkled. Although the result seems to be the same as far as
external appearance is concerned, the second contrasting char-
acter of Yy, being heterozygous, will not breed true for yellow in
succeeding generations. In the like manner, if the first pair of
contrasting characters were Ss instead of ss, again there would be
a heterozygous character which would not breed true should that
zygote be self-fertilized.

With this fact as a basis, the F: generation of the cross between
varieties of Pea plants having yellow color, wrinkled-surface
seeds and plants having green color, smooth-surface seeds bear
zygotes in the ratio of 9 plants with dominant.characters for both _

652 PHARMACEUTICAL BOTANY

yellow color and smooth surface; 3 with homozygous characters 2

for the recessive factor of green color but either homozygous or
heterozygous for the dominant factor of smooth surface and thus
all appearing with green color and smooth surface; 3 with homo-
zygous characters-for-the recessive factors of wrinkled surface
but either homozygous or heterozygous for yellow color; and 1
with both sets of characters homozygous for the recessive green
color and wrinkled surface which breed true when self-fertilized.
The four classes when grouped as to the appearance of seeds are
then as follows:

9 Yellow color, smooth surface
3 Yellow color, wrinkled surface
3 Green color, smooth surface
1 Green color, wrinkled surface

Ga.ton’s Law or ANCESTRAL INHERITANCE.—Francis Galton |
was the founder of the scientific study of inheritance. He
concluded from a study of several selected traits of certain families
that the two parents contribute between them on the average
one-half of each inherited faculty, each of them contributing one-
quarter of it. The four grandparents contribute between them
one-quarter, or each of them one-sixteenth; and so on, the sum
of the series } plus 14 plus 1 plus 144 . . . being equal to 1.

VARIATIONS

VaRIATIONs represent the differences which exist between the
individuals of a species. They are those changes in organisms
which make them different from their parents or from their
species. No two organisms of any species are exactly alike.
This is very apparent from the examination of children in the
same family or plants reared from seeds of the same fruit.
While having similar characteristics in the main, differences
may readily be made out whereby we can distinguish the indi-
viduals. Examples of kinds of variations are variations in size, _
symmetry, outline of organs as leaves, the appearance of a new _
Pigment, etc.

Other "kinds of variations are (a) modifications due to differences
in environment to which organisms are exposed; these are _

GENETICS AND EVOLUTION : 653.

called fluctuations. (2) Variations due to new combinations of ances-
tral characters which in some cases were latent in the parents
but present in the grandpabents or in more distant ancestors.
Such instances are called ‘‘reversions.”’ (3) Mutations or sudden
and frequently large differences which appear among individuals
not of hybrid origin. It is believed that many varieties of
economic plants had their origin in seed mutants or “sports.”
The mutations seen in plants reared from seed sports are due to
hereditary causes in which there probably occur a combination
of parental characters. Bud mutations or bud sports sometimes
occur on normal plants as in some varieties of fruit trees espe-
cially in certain citrus fruits. The Boston Fern, Nephrolepsis
exaltata bostoniensis is a bud sport of Nephrolepsis exaltata. This
form of mutation is due either to the alternation of a simple gene
in a certain chromosome or to abnormalities in the distribution of
the chromosomes during mitosis. Continuous variations are small
additions or diminutions of certain parental characters. Dis-
continuous variations are sudden marked variations which arise
without the appearance of transitional stages, as for example
where variations occur in the normal number of parts, as in the
four-leaved clover. De Vries called these kinds of variations
mutations.

EVOLUTION

Organic evolution is the established fact that all forms of
organisms existing today have been derived from others pre-
viously existing either by direct descent or common ancestry.
It did not originate with Charles Darwin. It was first conceived
by Anaximander, Empedocles and other ancient Greeks. ‘The
more important theories as to the method of evolution will now
be discussed.

THe LAMARCKIAN THEORY OF Use AND DisusE.—Jean
Baptiste de Lamarck (1744-1829) a French naturalist, pro-
pounded his theory on the origin of species in 1809 in his “Phil-
sophie Zoologique.” He assumed that the environment changes,
that use and disuse may alter the character of the individual,
that characteristics acquired by an organism in its lifetime are
inherited by its offspring. The persistent use of a part of the

654 PHARMACEUTICAL BOTANY

body, as the muscles of the arm of a blacksmith, results in greater
development of that organ whereas the continued disuse of an
organ results in its partial atrophy or degeneration. While
these facts about the environment and the use and disuse of
organs have been recognized in the lives of many individuals,
Lamarck assumed that the changes brought about in an organism
through use and disuse of organs were passed on to the descend-
: ants. As an illustration of his theory he
used the neck of the giraffe. He assumed
the early ancestors of the giraffe were
grazing animals and that a change in the
} environment caused them to feed on the
} leaves of trees. The present long neck
was acquired, according to Lamarck, as
the cumulative result of the inheritance
of slightly greater neck-length brought
about in each generation by reaching up
SF persistently to browse upon higher
Fic. 486.—Jean Baptiste Vegetation.
de Lamarck (1744-1829) who His theory of the inheritance of
: xicalgci petit eh tpl acquired characters is largely rejected by
dividual in its lifetime are DlOlogists of the present day because of
inherited by its offspring and the lack of convincing evidence.
Feiss: anos changes. Tue Darwinian THEORY oF NAtTu-
RAL SELECTION.—Charles Darwin
(1809-1882) was an English naturalist, who made many observa-
tions on the British exploring expedition that circumnavigated
the globe on H.M.S. Beagle from 1831-1836. As an outcome of
this experience and other later observations he worked out a
theory of evolution which he published in 1859 in a book entitled
“The Origin of Species by Means of Natural Selection,” or “The
Preservation of Favored Races in the Struggle for Life.”

Darwin’s Theory is based on the following fundamental
propositions:

1. PRODIGALITY oF Nature.—Living organisms are enor-
mously fertile, yet the total number is approximately sta-
tionary. It seems well established that many more offspring are
produced by plants and animals than can actually survive. A

GENETICS AND EVOLUTION 655

very small proportion of the seeds and spores of plants actually
germinate and averagely few of the seedlings attain maturity.

2. STRUGGLE FOR EXxIsTENCE.—Owing to the production of
individuals far beyond the earth’s facilities to support them,
there is a struggle for existence. An intense competition is con-

Fic. 487.—Charles Darwin. (Gager.)

stantly taking place in nature for food, water and light among
individual plants and animals. The struggle is unequal because
of the variability in structures, activities and close relations.

3. ORGANISMS AND SPECIES DiFFER IN ResPECT TO THEIR
Fitness FOR A GIVEN ENVIRONMENT.—Obviously, a water plant is
unfitted to live in dry soil and a dry soil plant unfitted to live in
the water.

656 PHARMACEUTICAL BOTANY

4. VARIATION AND Herepity.—The individuals of a species
are not all alike and among the offspring of the same parents there
are always at least slight variations. These variations are con-
tinuous or fluctuating in character.

5. Natura SELecTion.—Through the struggle for existence
there is a natural selection which results in the survival of the
fittest. Darwin argued that in this biological competition some
organisms and species: possess advantageous variations and are
consequently preserved while
the less fortunate perish. The
favorable characters are handed
) down to the descendant, genera-
tion after generation, so long as
| they continue to be advanta-
| geous. Thus, the weak and
| poorly adapted organisms are
eliminated while the forms most
| in -harmony with their environ-
| ment survive.

6. ORIGIN OF SPpECIES.—
| According to Darwin, if certain
members of a given group adjust
themselves to the conditions of

Fig. 488.—August Weissmann. life through variation in one
(Atwood.)

direction and others in other
directions, then divergence among the forrhs through the con-
tinued action of Natural Selection generation after generation
must in the course of time be sufficient to make them rank as

separate species.
WEIssMANN’S GERM PLAsm THEORY.—August Weissmann
(1834-1914), a German biologist, developed a theory of heredity
based upon the principle of the continuity of the germ plasm.
While this principle was recognized earlier by Owen (1849),
Galton, Nussbaum and others, it remained for Weissman through
a series of brilliant essays and a book on the Germ Plasm (1892)
to bring it forcibly to the attention of scientific world.
Weissmann vigorously denied the inheritance of all forms of
acquired characters. He believed (1) that the cells of an

GENETICS AND EVOLUTION 657

organism are of two kinds, the body or somatic cells containing
somatoplasm and the germ cells containing germ plasm, (2) that
the germ cells reside in the body but are not part of the body or
soma, (3) that the germ cells arise as direct descendants of the
germ cells of the previous generation, (4) that the germ plasm
continues from the beginning of life and the soma protects it,
(5) that variations arise from combination of different characters
in the germ plasm of each parent, (6) that the determiners
originate solely in the germ plasm and migrate from the germ
cells out into the various parts of
the developing body and that |
differentiation of the organism is
produced in this way, (7) that a
new type of organism arises only
in consequence of a changed type
of germ cell.

Weissmann’s ideas about vari-
ation were just the reverse of those
held by Lamark and Darwin who
believed that variations first origi-
nate in the body and are thence
passed on to the germ cells. He pi a range
stressed the imp ortance of heredi- of Ate of ae as @ meted
ty in evolution but almost NC¥- of evolution. (After Walton and Foss.)
lected environment.

De Vries Mutation THEoRY.—Professor Hugo De Vries,
a Dutch botanist, in 1901, advanced the theory of evolution
through sudden abrupt variations. He asserted that new
species arise suddenly and become fully established from a
parent form which may continue to live side by side with the new
form. He called these sudden variations, mutations. De
Vries reached his conclusions from the striking variations he
discovered in successive generations of the Evening Primrose
(Ocnothera lamarckiana), a species he found growing wild, in 1886,
in a waste field about Hilversum in Holland. In experimenting
on the Evening Primrose in his Amsterdam gardens, he found the
mutants when self fertilized proved stable, and when crossed
behaved according to the Mendelian laws. De Vries’ principal

658 PHARMACEUTICAL BOTANY

conclusions have been verified by many subsequent investigators
and his methods extended to the discovery of mutants among
many species of plants and animals.

Curomosome Murations.—Since De Vries’ discovery, cyto-
logical studies have been applied to the study of mutations.
These have shown that many mutants possess new chromosome
numbers throughout their structure. For instance, Oe¢nothera
lamarckiana possesses 14 chromosomes while one of its mutants,
Oenothera lata, has 15. .

Po.yPpLoipy.—The normal somatic cells of the bodies of higher
plants and animals have two sets of chromosomes (diploid).
Polyploids are forms having multiples higher than two, e.g. three
or more chromosome sets. Polyploids have been found to
originate by hybridization followed by chromosome doubling.
Extreme temperatures are claimed by Sax and others to pro-
duce doubling of the chromosomes. Polyploidy has been found
to account for many mutants in wild plants as the chrysanthe-
mums, maples, “roses, apples, cherries, hawthorns, erigeron and
rumex and in cultivated plants including wheat, oats, tobacco,
sugar cane, dahlias, etc.

Tue Mopern THrory or tHe Gene.—The current theory
had its origin in the work of Thomas H. Morgan, an American
geneticist, who discovered that the fruit fly, Drosophila melanogas-
fer, was in a mutating condition. About 400 mutant characters
have been identified in this insect. Morgan showed that the
genes can be linked into four groups that correspond to four
pairs of homologous chromosomes in the germ cells. The loca-
tion of the characters or their determiners or genes on the
chromosomes has been worked out by Morgan and his associates,
and their experiments have shown that genes may cross over
from one homologous chromosome to another. They have
further shown that genes displaying linkage with each other are
located in the same pair of chromosomes, that the substance of
the chromosomes binds the genes to each other and causes one
factor of a specific character to be inherited along with another
gene on the same chromosome.

Several explanations have been advanced concerning the
causes of gene mutations or the appearance of new hereditary

GENETICS AND EVOLUTION — 659

characters. Gates suggests they may be environmental, as for
example the lowering of the temperature during periods of germ
cell maturation. Others have suggested internal causes, such
as the weakness of attraction between homologous chromosomes
and the consequent failure of synapsis.

The rate of mutations has been slightly increased in Drosophila
by Muller through raising the temperature of cultures of this
insect. In 1926 he showed that the X-ray had the power to alter
and rearrange the genes. By exposing adult Drosophila flies to
powerful X-rays he greatly accelerated the rate of the mutations,
increasing these by up to 150 per cent.

It is now thought probable by many geneticists that genes
located in the chromosomes represent the sole vehicle of inherit-
ance and that heritable variations can come only from changes
in the genes. ‘These changes represent mutations.

Appendix I

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 3 principal kinds, viz.: simple, dissecting
and compound.

THe SIMPLE MICROSCOPE

This consists simply of a convex lens or several combined into a
system and appropriately mounted. A good example of a sim-

Fic. 490.—A dissecting microscope. Description in text.

ple microscope is a reading glass. This type of simple micro-
scope is valuable in field work, in the examination of dried
herbarium material or the external characters of crude drugs,
where only a low magnification of the object is required.

Tue DissEcTING MICROSCOPE

But when flowers or other plant parts are to be dissected, it is

necessary to have both hands free. To meet this need various
661

662 PHARMACEUTICAL BOTANY

forms of stands have been devised which have been combined
with an arm and lens to constitute what are known as ‘‘Dissect-
ing Microscopes.” One of the simpler forms of these is shown

Fic. 491.—Zeiss stereoscopic dissecting microscope with built in illuminating
equipment.

in Fig. 490. It consists of a metal stand with convenient hand
rests; a glass stage plate, full size, on which the object to be dis-
sected is placed; a double faced mirror adjustable for reflecting
light from various angles, the concave face for concentrating all of
the light at the plane of the object. A jointed lens arm is carried

APPENDIX I 663

on a rack-and-pinion focusing adjustment having ample range
to suit a variety of thicknesses of objects. A black-and-white
metal stage plate serves for use as a contrast background. A
more elaborate type of dissecting microscope of the twin objec-
tive binocular type is shown in Fig. 491. This gives erect images
and a stereoscopic effect.

. THE CompounD MICROSCOPE
A. Its CONSTRUCTION:

The principal parts of a compound microscope are:

1. The base, generally horseshoe shaped, which rests on the
table.

2. The pillar, an upright bar, which is attached to the base
below, supports the rest of the instrument and incorporates the
inclination joint.

3. The stage, a horizontal shelf upon which is placed the
preparation 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. The mirror
generally has two faces; the one is plane for initial light intensity,
the other concave for concentration of light on the object.

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.

6. The body tube, a cylinder which holds the draw tube and
lenses and moves up and down perpendicularly above the open-
ing 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 and
is used for finding the focus.

7. The fine adjustment, a micrometer screw back of the tube,
which, on being turned, produces a very slow motion of the entire
framework which holds the body tube. It permits exact focus-
ing of the higher power lenses.

8. The oculars or eyepieces which slip into the upper end of the
draw tube. Each of these consist of two plano-convex lenses, the

:
:
‘
2
z

664 PHARMACEUTICAL BOTANY

lower one being the larger and known as the collective or 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. It determines the size of the field of view.

Fic. 492.—Illustrating the parts of a compound microscope.

According to the system adopted by the maker, oculars are
designated by numbers, as 1, 2, 3, 4, etc., or by figures which
represent focal lengths, and by magnification numbers, as 7X,
10, ete.

9. The objectives, which screw into the bottom of the body.
tube or nose piece. They consist of a system of two, three or
more lenses, some of which are simple, others compounded of a

APPENDIX I 665

convex crown lens and a concave flint lens. Objectives like
oculars are usually designated by fractions as 149, 1é, 24, etc., or
in millimeters, as 2 mm., 4 mm., 16 mm., which represent focal

y

EP Evepoiat
o era |
paiement 0; men]
wok
A
ss i
{| a!
A
y |
i! \
c \

(Eo

te,’
1
: H Enteric:
Pencil
mans 2a
Paral
Light %
; 3 3

Fic. 493.—Diagram illustrating optics of a compound microscope in use. Fy,
Upper focal plane of objective; F2, Lower focal plane of eyepiece; A, Optical tube
length = distance between F, and F2; Ou, object; Oz, real image in F., transposed
by the collective lens, to Os, real image in eyepiece diaphragm; Ou, virtual image
formed at the projection distance C, 250 mm. from EP, eyepoint; CD, condenser
diaphragm; L, mechanical tube length (160 mm.); 1, 2, 3, three pencils of parallel
light coming from different points of a distant illuminant, for instance, a white
cloud, which illuminate three different points of the object. (Courtesy of Bausch
and Lomb Optical Co.) :

lengths or numbers, as 97, 45, 10, etc. which indicate the mag-

nification of the objective alone. -
The smaller the number in millimeters or fraction represent-

ing the focal length of an objective, the greater is its magnifying
power. :

666

PHARMACEUTICAL BOTANY

The function of an objective is to take in the divergent light
from the object and change it into convergent rays that meet in

PM ee
cy ular

SS

Cx

LE LLL EE
SY/

7,
4,

YN:
Nt) nh.

D

Fic. 494.-——Sec-
tional view showing
component elements
of an apochromatic,
high power objective.

the upper end of the microscope tube in the
front focal plane of the ocular forming there a
magnified image of the object. ‘This image is
further magnified by the lens of the human eye.

OsjecTivEs are either dry lenses or zmmerston
lenses. If an air space be present between the
tip of the objective and the object, the lens is
called a dry one; if a liquid is present between
the tip of the objective and the object, the lens
is called an immersion lens. If this liquid be oil,
the objective is called an ozl immersion objective;
if water, a water immersion objective.

Dry objectives are of low, medium and high
powers. Dry medium and dry high power

objectives are the best types to use on objects in air.

When examining objects in water, glycerin or

balsam with the dry objective, the cover glass
should be slightly under 0.17 mm. in thickness
for best results.

The important properties of the microscope
objective are: (1) Numerical aperture (N.A.) which
measures the amount of light received from the
object and represents the index of resolving
power. (2) Resolving power is that by which two
small elements in the structure of an object and
which are only a short distance Aper are dis-
tinctly separated.

The higher the N.A. the greater the resolving
power of the objective and the finer the detail it
can reveal.

N.A. equal W times sin u
wherein

n = the lowest refractive index that appears

between the objective and the front of
the objective.

Fic. . 495.—A
new type of ob-
jective in a correc-

tion mount. Turn-
ing the correc-
tion collar varies the
distance between
the two upper lenses
and the two lower
lenses fixed in the
body of the mount.

= half the angular aperture of the objective.

APPENDIX I 667

The degree of angle of illumination directly affects resolving

power up to a maximum resulting from filling the aperture of
the objective with light.

Fic. 496.—A modern monocular research microscope with revolving nose piece,
side fine adjustment and substage.

A 4 mm., 0.85 N.A. objective will resolve lines separated by
distance ranging between 0.00062 and 0.00031, dependent upon
the aperture employed. For a 4 mm., 0.65 N.A. objective the
limiting values are 0.00081 and 0.000405.

668 PHARMACEUTICAL BOTANY

Two objectives of the same equivalent focal length (E.F.) and
the same N.A. should show the same illuminated area in the
back lens, when viewed without an eyepiece and illuminated
with the widest cone of light they can take in.

It also is evident that an objective cannot show its full effi-
ciency if it is not used with a condenser of an N.A. large enough
to fill the back of the objective with light.

DeptH oF Focus.—Depth of focus (known also as depth of
sharpness of penetration) depends on the N.A. and the magnifi-
cation and is inversely proportional to both. The higher the
N.A. and the higher the magnification, the less the depth of
focus. An increase of the depth of focus, for instance, by insert-
ing diaphragms above the back
lens of the objective must neces-
sarily decrease the effective diam-
eter of the back lens and thus
decrease the N.A., thereby lower-
ing the efficiency of the objective.

: | To comply with different re-
Fic. 497.—Aplanatic condenser (Zeiss) quirements in this respect the
N.A. 1.4. Focal length 10.5 mm. A
makers offer two 4 mm. objec-
tives, whose depths of focus, when used with the same magnifica-
tion, are in the ratio of 3:4.

It should be borne in mind that the finer the definition of an
objective the more sensitive it is to incorrect focusing and to
slight changes of the fine adjustment screw. This means less
depth of focus. Objectives that show greater depth of focus
than others of the same N.A. cannot be well corrected. _

Working distance of an objective is sometimes confused with the
equivalent focus. It varies for different powers being dispropor-
tionately less for high powers, and is measured from the front of
the objective to the upper surface of the cover glass. For
instance, the high dry objective (4 mm. ef. 0.65 N.A.) has a
working distance of 0.6 mm.

Some microscopes are fitted up with a nose-piece, capable of
carrying two, three, or four 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

APPENDIX I 669

object examined. When using the condenser, employ only the
plane mirror.

B. Irs 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 same time turning the mirror at such an angle that it

Fic. 498.—Microscope lamp, useful in illuminating opaque objects.

reflects light from the window or lamp up through the opening
in the stage to the objective. When opaque objects are to be
illuminated, a stronger illumination is required than that usually
afforded by an ordinary laboratory lamp or by. the light from a
window. For this purpose a microscope lamp, such as the Bausch
and Lomb No. 4578 (see Fig. 498) is very satisfactory. Mirrors
have two faces, a plane and a concave. Use the concave unless
employing the condenser, when the plane mirror should always
be used.

4. Regulate the quantity of light by the iris diaphragm. If
too bright it must be cut off somewhat. The higher powers

670 PHARMACEUTICAL BOTANY

require brighter light than the lower; the oil immersion full
opening.

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.

Fic, 499.—Binocular microscope. This is a compound microscope with a bin-
ocular body designed to relieve eyestrain for those doing microscopical research
work or otherwise engaged for prolonged periods with the microscope. Note the
parallel position of the eyepiece tubes which are adjustable for interpupillary
distance by means of a compound slide, with the distance indicated on a numbered
millimeter scale for future guidance. One tube has a spiral adjustment for correct-
ing any difference in vision existing between the two eyes. The instrument shown
is that designated CAE of the Bausch and Lomb Optical Co.

6. With the lower power in position quite close to the
object, move the coarse adjustment upward until either the
object or small solid particles on the slide appear distinctly,

APPENDIX I 671

which means that the lenses are in focus. (If two objectives
only accompany your instrument, the low power is the shorter
one.) 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 position. Lower the body tube carrying the
objective 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 while looking through the ocular because of the danger
of pressing it into the cover glass and the risk of ruining the
delicately mounted lenses.

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.

9. When the oz/ immersion objective is to be used, a small drop of
immersion oil (slightly evaporated cedar oil) should be placed on
the cover glass directly above the object, and the body tube
should be run down with the coarse adjustment until the. front
lens of the immersion objective enters the drop and comes almost
into contact with the cover glass. This should be done while
watching the objective. Then look through the ocular and
draw the objective up with the fine adjustment until the object
comes into focus.

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.

2. Never allow the dry objective to touch the cover glass or
the liquid in which the object is mounted.

672 PHARMACEUTICAL BOTANY

3. Never touch the objective or ocular lenses with fingers or
cloths.

4. Never change from lower to higher power objective with-
out first ascertaining that the body tube has been raised suff-
ciently to allow the high-power objective to be slipped into place
without injury to the objective or mounts.

5. Never clean the microscope lenses or stand with cloths
that have been used for removing surplus of alkali, acid or other
reagent from slides.

6. 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

Cnedeheted 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.

7. 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 ona
cover glass.

8. Never lift the slide from the stage, but, after raising the
objective, slide it off the stage without upward movement.

9. Never allow the stand (microscope without lenses) to be
wetted with such substances as alcohol, soap, etc., which dissolve
lacquer.

10. Keep the microscope covered when not in use.

Fic. 500.—Ocular
micrometer.

MICROMETRY

The unit of length used in microscopic measurement is the
micron (u) 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 (Fig. 500).° Itis a circle of glass suitable for inser-
tion 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

APPENDIX I 673

surface of the glass may be etched with squares (0.5 mm.), the
net micrometer (Fig. 501).

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 (Fig. 502) is a slide
with a scale engraved on it divided to hun-
dredths of a millimeter (0.01 mm.), in some
cases, also, to tenths of a millimeter (0.1 mm.),
every tenth line being made longer than Fic. 501.—Net mi-

intervening ones, to facilitate counting. crometer, about 173 X
Miricas actual size. The en-
ETHOD:

Ee graved scale is 10 mm.
1. Insert the ocular micrometer within the square and is divided

tube of the ocular by placing it on the dia- into small squares with
: 0.5 mm. sides.
phragm 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.

Fic. 502.—Stage micrometer used to standardize the ocular micrometer. The
mm. readings etched on this micrometer indicate the width of the larger and smaller
spaces of the stage micrometer scale seen in the center of the stage micrometer.

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.

4. Note the known value for each division of the stage
micrometer scale which may either be etched on the stage

674 . PHARMACEUTICAL BOTANY

micrometer or indicated on a label found pasted upon it. If
the value indicated is 0.01 mm. (1499 mm.), then each division
of the stage micrometer scale has a value of 10

microns; if 0.1 mm. (149 mm.), 100 microns.
5. Multiply the number of included divi-
sions of the stage micrometer scale by the
value in microns given for each division and
divide the result by the number of included
divisions of the ocular micrometer scale. The
Fic, 503.—Scale of quotient represents the value of each division

cals i eR ET of the ocular micrometer scale.

Each small space of 6. Note the optical combination (number
actual scale has a value of ocular, objective and tube length) used and
of 10 “tedcrons, each keep a record of it with the calculated microm-

large space between the
longest lines, 100 mi- Cter value. Repeat for each of the

crons. The length of combinations.

the entire scale is 1 mm. To measure an object by this method,
read off the number of divisions it occupies of the ocular microm-
eter scale, and express the result in microns by looking up the
recorded value for the optical combination used.

Appendix IT
HISTOLOGICAL TECHN IQUE

In this chapter the various methods which are commonly
employed in the preparation of plant materials for microscopic
examination and the reagents used in the examination of the
same will be discussed.

Makinc 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. Fresh material should always be
kept moistened in water on account of the danger of cell shrink-
age through loss of water. Dried membranous or subleathery
leaves should be kept in a moist chamber for at least 24 hours or
until sufficiently supple before they are sectioned. Dried
woody roots and stems can usually be best prepared for section-
ing by soaking segments of these in equal parts of alcohol,
glycerin and water for several weeks prior to 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. Fine shavings can
then be whittled off of these with a sharp knife. In some cases
alternatingly boiling and cooling the material will suffice.

The object to be sectioned is held between the thumb and
finger of the left hand. If tender and flexible, such as a fresh
leaf, it must be placed between the two flat surfaces of elder pith
before sectioning. A segment of pith about an inch long is
halved lengthwise with a sharp knife and a portion of the leaf is
held between the halves of pith while the section is cut through
pith and leaf. The pith is later separated from the leaf section.

Sections through other delicate parts of plants, as buds, slender
675

676 PHARMACEUTICAL BOTANY

roots, etc., may be made in the same way, only a groove should
be made in the pith of such size as is necessary to hold the material
firmly enough without crushing it. In certain instances, when,
because of the smallness of the object and its resistance to cutting,
good sections can not readily be made with the aid of pith, a
small sized cork stopper can be used with better results. A hole
just large enough to prevent the object from slipping is made in
the center of the smaller end and the object inserted preparatory
to sectioning. The upper surface of the razor is wetted with
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.
Small bits of representative parts of entire sections yield better
results when examined under the microscope than complete sec-
tions which have been cut too thick. 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, before they become dry. If
the sections are cut from fresh material and are to be stained,
they should be placed in 95 per cent. alcohol for at least several
minutes prior to staining. If from material preserved in forma-
lin solutions, they should be washed in water.

Surface sections of leaves may occasionally be prepared by
stripping off the leaf epidermis. In most instances, however,
the leaf epidermis adheres firmly to the subjacent tissue and it
becomes necessary to employ different technique. A good
practice is to bend the leaf over the index finger of the left hand
and hold it firmly between the index finger and the third finger
on the one end and with the thumb on the other. Another
method is the following: Place a representative portion of the
leaf not over }4 inch square in a watch crystal with 10 per cent.
to 25 per cent. chloral hydrate solution, warm gently for several
minutes, according to the texture of the leaf, transfer to a clean
slide, add a drop or two of chloral hydrate solution, cover with
cover slip and exert pressure on the cover slip with a rotary
movement, describing the figure 8. The epidermis will thereby
usually separate from the underlying tissues. It sometimes
becomes necessary to study large areas of the epidermis of

APPENDIX II 677

leathery leaves. Such leaves should be prepared by boiling in
10 to 20 per cent. aqueous solution of either sodium or potassium
hydroxide until the epidermis puckers up in the form of blisters
when the epidermis can be lifted off by means of fine forceps or
dissecting needles and floated on the slide.
SECTIONING IN PARAFFINE OR CELLOIDIN.
When it is necessary to study the micro-
scopic structure of very delicate plant parts,
superior results can generally be obtained by
imbedding the material in paraffine or
celloidin (see pp. 692 and 699), which is
-subsequently hardened, and sectioned by
means of a sliding or rotary microtome.

Kinps 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.

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 the plane
of the radius.

3. A tangential-longitudinal section is one
made parallel to a plane tangent to the
cylinder. This type of section is therefore Fg. 504.—Showing

prepared by cutting parallel to the outer the planes in which sec-
long surface tions are cut, A, trans-
i ‘ . versely; B, longitudinal

4. A surface section 1s one made by Glin’ CO tencieadk:

cutting or stripping off the outer protective nal tangentially. (After
covering of the plant organ. Stevens.)
MICROTOMES
Microtomes are instruments employed to facilitate the cut-
of organic tissues. The three most commonly

hand, sliding and rotary microtones.

ONE.—Lhis type is shown in Fig. 505. If the
hard to bear the strain, it is placed directly
end of the tube that is tightened by the

ting of sections
used types are the

Hanp MicrotT
object is sufficiently
in a clamp of the upper

678 PHARMACEUTICAL BOTANY

screw seen on the side of the tube, or it may first be inclosed in
elder pith or cork and then clamped in. The object to be sec-
tioned is raised a little at a time through the hole in the glass
plate at the top by turning the finely graduated feed near the
base of the tube. ‘The section razor is then laid flat on the glass
plate and pulled across the object with a long sliding motion.
The upper surface of the razor blade is
kept wet with 50 per cent. alcohol and
after several sections have been cut they
can be swept by the finger or camel’s
hair pencil to a dish of water. Each
division of the feed represents 10 microns,
so that the thickness of sections desired
can be regulated by moving the feed,
accordingly, just before each stroke of
the razor.

Suipinc Microrome.—This type of
microtome (see Fig. 506) is adapted for
cutting all kinds of sections. It consists
of an iron supporting frame of horizontal
and upright portions. The horizontal
base rests on the table and is hollowed
out to accommodate a drip pan that
Fic. 505.—Hand microtome. can readily be removed and cleaned.

Descuipeiodt tp text. The front of the upright portion
exhibits a frame which accommodates a sliding feed mechanism
to which is attached the object carrier, ‘The top of the upright
portion shows a flat bed which carries a solid iron block
which can be readily slid along the bed when the latter is lubri-
cated with paraffin oil. The upper surface of the block js
grooved to accommodate the thumb screw. The microtome
knife consists of a blade portion that is flat on its lower and
hollow ground on its upper face. The latter is placed in the
clamp on the top of the microtome, and its position adjusted.
Sections of woody material can be cut directly on this microtome
and placed in dilute alcohol. When paraffine sections are cut
the cutting edge of the knife should be parallel to the motion; but
when celloidin sections are desired the knife must be set at an

APPENDIX II 679

oblique angle to the frame and drawn across the block with a
long sliding motion. The knife and the top of the celloidin
block must be constantly kept wet with 80 per cent. alcohol.

The feed mechanism is covered to protect the

Fic. 507.—Rotary microtome.
a (Courtesy Spencer Lens Co.)

wearing parts from dust.
The object is placed in the object carrier and clamped in.

By means of the graduated disk at the base of the feed mecha-

nism the thickness, in terms of microns, is regulated after each

stroke of the razor.

680 PHARMACEUTICAL BOTANY

Rotary MicroroMe.—When paraffin ribbons are desired,
especially for the study of serial sections of material, the rotary
microtome surpasses by far the efficiency of the sliding type of
instrument. The Spencer Rotary Microtome No. 820 is shown
in Fig. 507.

REAGENTS AND STAINS AND METHODS OF PREPARATION
OF MATERIALS

The following reagents will be found of value in the micro-
scopical examination of plants, powdered vegetable drugs and
foods: (additional reagents and formulas for their preparation
are presented elsewhere in the text in connection pith methods
of examination).

Acetic Acip.—This reagent should contain not less than
36 per cent. nor more than 37 per cent. of pure acetic acid. It is
used for various operations such as distinguishing between cal-
cium carbonate which dissolves in it with effervescence and
calcium oxalate which is insoluble in it. A 2 to 3 per cent.
solution is occasionally employed as a mounting medium for
microscopic plants.

Acip ALCOHOL.—Mix 149 cc. hydrochloric acid with 100 cc.
of 70 per cent. alcohol. This is employed in removing the excess
of stain from overstained material. When used, it should be
thoroughly. washed out with diluted alcohol or water.

ALCOHOL.—Useful in the form of various percentage solu-
tions for dehydrating, preserving, hardening and solvent pur-
poses. Absolute alcohol (Dehydrated alcohol) should contain
not less than 99 per cent. by weight of ethyl hydroxide and must
be kept in well stoppered bottles because of its property of rapidly
absorbing moisture from the air and so becoming reduced
in strength. It should always be used before cedar oil or xylol
in making a balsam mount, but if clove oil is used for clear-
ing instead of cedar oil or xylol, 95 per cent. alcohol may be
employed.

ALCANNA TINCTURE.—Macerate 20 Gm. of alkanet root for a
week in 100 cc. of 90 per cent. alcohol, boil for several minutes
and cool. Dilute with an equal volume of water just before

SST ae On eee Te Oe

a i

APPENDIX II 681

using. ‘This reagent imparts a red color to fixed oils but may
stain other contents as well.

AmMoniéA WATER.—An aqueous solution of gaseous ammonia
containing in each 100 cc. not less than 9 Gm. and not more
than 10 Gm. of NH;. Useful in clearing highly colored vege-
table powders, such as ground roasted coffee, when mixed with
equal parts of peroxide of hydrogen. Material should be
macerated in this mixture within a tightly corked vial over
night or longer and washed with water before examination.

AMMONIA WATER, STRONGER.—This is an aqueous solution
of NH; containing not less than 27 per cent. nor more than
29 per cent. by weight of ammonia gas. It is employed in the
preparation of ammonia water and cuoxam.

ANILINE CHLORIDE.—A saturated aqueous solution acidified
with hydrochloric acid is useful in staining lignified walls which
are colored a golden yellow. :

ANILINE SULFATE.—A saturated aqueous solution is em- ©
ployed as a test for lignified walls. Sections are mounted in .
this reagent and a drop of sulfuric acid added. A yellow color
is imparted to lignified walls.

Anne BuLue.—A saturated aqueous solution is useful in
staining sieve tubes. Sections should be placed in this solution
for 24 hours and then washed to remove excess of stain. A one
per cent. solution in 90 per cent. alcohol is used in double-stain-
ing with safranin. It stains cellulose walls blue.

Bismark Brown (ANILINE Brown).—A saturated aqueous
solution is useful in double staining with gentian violet. A dilute
aqueous solution made by dissolving 0.2 Gm. of Bismark Brown
in 100 cc. of distilled water is of value in staining transparent
tissues.

CutLorAL Hyprate.—A solution prepared by dissolving 25

Gm. in 10 cc. of water is an excellent clearing agent. It dis-
solves starch, resin, protein, chlorophyll, etc., and causes more
ess expansion of shrunken cells. ;
Cutorav IopinE.—Saturate the previously prepared chloral
hydrate solution with iodine by adding crystals of resublimed
iodine to it and shaking. Useful for the detection of minute
traces of starch or small starch grains, which are colored blue.

or |

682 PHARMACEUTICAL BOTANY

CHLORZINCIODINE SOLUTION.— Dissolve 25 Gm. of anhydrous
zinc chloride and 8 Gm. of potassium iodide in 8.5 Gm. of water
and add iodine crystals to saturation. Keep in dark colored
containers. Useful in distinguishing between cellulose and
lignified walls. Cellulose walls are colored blue or violet and
lignified walls yellow when mounted in this solution. It causes
starch grains to swell and colors them blue.

Curomic Acip SoLution.—Dissolve 5 Gm. of chromic acid
in 45 cc. of diluted sulfuric acid (U.S.P.). A valuable reagent
for separating the component cells in sections of plant organs.
The sections are placed in this solution in a watch-glass and
removed one by one, as required, in the course of 10 to 15
minutes to a slide, washed with a few drops of water and sub-
jected to pressure with a glass rod.

CorRALLIN SODA SoLuTion.— Dissolve 15 Gm. of carbonate of
soda in 35 Gm. of distilled water and add sufficient corallin to
produce a pale bright pink color. This reagent stains the
callus plates of sieve tubes pink. It should be freshly prepared
as needed.

Eosin.—An aqueous solution prepared by dissolving 1 Gm.
of eosin in sufficient water to make 100 cc. is useful in the staining
of cytoplasm, aleurone grains and other cell contents. If
permanent mounts are desired, transfer to 1 per cent. acetic
acid for a minute, wash with water, then to glycerin or through
glycerin to glycerin jelly.

A saturated alcoholic solution of eosin (using absolute alcohol
as solvent) is employed when eosin-stained material is to be
mounted in balsam. Acetic acid is omitted in this case and the
sections passed from alcoholic eosin to a clearing agent as cedar
oil or xylol, thence into balsam.

ErHer.— Useful in defatting sections or powders of oil seeds.

Fast GREEN.—A one per cent. solution in 95 per cent. alcohol
is useful in staining tissue with cellulose walls green.

FEHLING’s SoLuTION. (Alkaline Cupric Tartrate T.S.).

Copper Solution.—Dissolve 34.66 Gm. of small, uneffloresced
crystals of cupric sulfate in sufficient distilled water to make
the mixture measure 500 cc. Keep in well stoppered bottles.

APPENDIX II 683

Alkaline Tartrate Solution.—Dissolve 173 Gm. of crystallized
potassium and sodium tartrate and 50 Gm. of sodium hydroxide
in sufficient distilled water to make the solution measure 500 cc.
Keep in rubber-stoppered bottles.

Mix exactly equal volumes of the two solutions when required,

This solution yields a precipitate of cuprous oxide with reducing
sugars.
FERRIC CHLORIDE SoLuTIon.—A 1 per cent. solution of ferric
chloride in distilled water is used to detect the presence of
tannin. It yields a dark green to bluish-black precipitate with
tannin.

Fucustn (Basic).—Dissolve 1 Gm. of fuchsin in 100 cc. of
95 per cent. alcohol and add an equal volume of water. This
solution will impart different tints of red to the various tissues.
It has been found especially useful in staining sections which are
to be photomicrographed.

Fucusin, Actb.—A 0.2 per cent. aqueous solution is useful
in staining plastids and the phytoglobulins of aleurone grains.
A 1 per cent. solution in 50 per cent. or 70 per cent. alcohol
is occasionally employed in conjunction with methyl green or
fast green in the double staining of tissues of plants.

GentiAN VioteT (METHYL VioLeT).—An aqueous solution,
made by dissolving sufficient of the dye in 1 per cent. acetic acid
solution to impart a deep violet color, is useful in fixing and
staining pollen grains, spores and other fresh plant material.
A 1 per cent. solution in distilled water is used in conjunction
with Safranin and Orange G. solutions in staining material used
for the study of nuclear divisions.

Griycerin, DituTe.—Equal parts of pure glycerin and dis-
tilled water. Used as a mounting medium.

GrycerInE GuM.—Dissolve 20 Gm. of gum arabic and 20
Gm. of glycerine in 15 cc. of water. Useful for fixing small
fruits or seeds on pith before sectioning.

Ha@MaTOXYLIN, DELAFIELD’s.—Used for double-staining, see
p. 701. ie ‘ :

H&MATOXYLIN, HEIDENHAIN’S.—This consists of two solutions,
a mordant and a stain. The mordant (Solution I) is prepared

684 PHARMACEUTICAL BOTANY

by dissolving 3 Gm. of Ferric Ammonium Sulfate in sufficient
distilled water to make 100 cc. The stain (Solution II) is pre-
pared by dissolving 0.5 Gm. of hematoxylin in 100 cc. of distilled
water. The mordant and the stain are used separately on the
sections or tissues to be treated.

Hyprocutioric Acip.—The concentrated C.P. acid is used
with an equal volume of Phloroglucin Solution in determining
lignification. Dilute solutions are employed for neutralizing
alkaline solutions, etc.

Iopine WAtTER.—Add as much iodine to distilled water as it
will dissolve. Stains starch pale blue to purplish-blue.

IopINE AND PorasstuM Iop1bE SoLution.—Dissolve 1 Gm. of
iodine and 3 Gm. of potassium iodide in 50 cc. of water. Keep
in glass stoppered bottles. This reagent stains cellulose, ligni-
fied and corky walls and proteins yellow and starch blue to
bluish-black.

LABARRAQUE’S SOLUTION.—Triturate 50 Gm. of fresh chlor-
inated lime with 250 cc. of water so as to form a uniform mix-
ture. Dissolve 35 Gm. of monohydrated sodium carbonate in
250 cc. of hot water, and add this solution to the chlorinated lime
mixture in a suitable vessel. Stir or shake thoroughly. (Warm
if solution gelatinizes.) ‘Transfer the mixture to a wetted muslin
strainer, returning the first portion until the liquid passes through
clear, and when no more liquid drains from it, wash the precipi-
tate with sufficient water to make the product weigh 500 Gm.
Keep in amber colored, glass stoppered bottles in cool place.
It gradually loses in strength on standing. ‘This reagent is
used for bleaching highly colored sections and powders. The
materials bleached should always be washed with water before
mounting.

Licnut Green.—This is used as a combination stain with
safranin. Dissolve 1 Gm. of light green in 100 cc. of clove oil;
25 cc. of absolute alcohol and 75 cc. of clove oil may be used in
place of pure clove oil, if the light green is found to precipitate:
out when xylol is subsequently employed as a clearing agent.

NaAPHTHOL SOLUTION.—Dissolve 5 Gm. of alpha-naphthol in
50 cc. of alcohol. Used in conjunction with concentrated sul-
furic acid for the detection of inulin. <A violet coloration is

APPENDIX II 685

produced if sections containing inulin are first treated with a
drop or two of this reagent and then, after a minute, a drop or
two of sulfuric acid be added.

- PerROLEUM EtHEer.—Useful in clearing sections or powdered
material of oil.

PHLOROGLUCIN SOLUTION.—Dissolve 1 Gm. of phloroglucin
in 50 cc. of 95 per cent. alcohol. This solution gradually darkens
and loses strength with age. It is unfit for use after three or
four months. The lignified walls of elements first treated with
several drops of this solution and subsequently with a drop or
two of concentrated hydrochloric acid are colored red.

PorasH SoLutTion.—Several percentage solutions are em-
ployed in histological technique for varying purposes. ‘The most
commonly used one is the 5 per cent. aqueous solution which is
used for clearing sections of starch, protein and tannin. Ten to
20 per cent. aqueous solutions are used for the purpose of sepa-
rating the epidermis of leathery leaves. The leaf material is
boiled in the strong potash solution until the epidermis puckers
up in the form of blisters, when it should be removed to a slide
and the epidermis lifted off by means of a fine forceps or dissect-
ing-needles. A 2 per cent. solution is used for the isolation of laticiferous
vessels from plant organs. Pieces of stems, barks or roots containing
these elements are placed in a test tube and digested on a water-
bath for a half hour or longer until the parenchymatous tissues
are sufficiently soft. The potash solution is then poured off and
the material washed with water, transferred to a slide and the
woody or other undesirable parts removed with the aid of dis-
secting needles. The softer parts are then stained with iodine
and potassium iodide solution; a cover slip is adjusted, the slide
placed on stage, and the laticiferous tissues are identified by the
presence of branching and anastomosing tubes stained yellow to
yellowish-brown with granular contents. A 1.5 per cent. solu-
tion is used for the digestion of ground cereals or wheat middlings
for the purpose of clarification preparatory to microscopical
examination of these materials.

PoTasstUM FERROCYANIDE.—A 10 per cent. solution is
ed in studying the localization of alkaloids in plant

employ
materials. Sections of fresh material are fixed, in order to pre-

686 PHARMACEUTICAL BOTANY

vent postmortem filtration of the alkaloids, by placing them in
this solution for 5 minutes, and the excess of this reagent washed
out with water. The K,Fe(CN). solution precipitates the
alkaloids in the cells which can then be colored by adding a
drop or two of FeCl; solution. Clearing of the sections may then
be effected with chloral solution.

RutTuHentuM Rep AnD LEAD ACETATE SOLuTION.—This is an
excellent reagent for the detection of mucilage, to which it
imparts a pink color. Add sufficient ruthenium red to a 10 per
cent. solution of lead acetate until it shows a wine-red color.
The lead acetate solution should be prepared with distilled
water and the ruthenium red added to quantities of it as needed,
as the reagent is unstable. In pharmacognosy, it is especially
useful in detecting powdered cacao shells in powdered cocoa and
chocolate, since the cacao shells are the only portions of the
cacao seed which contain mucilage.

SAFRANIN.—Several kinds of safranin solutions are in use,
viz., Alcoholic Safranin, Anilin Safranin and Aqueous Safranin.
Alcoholic Safranin is prepared by making a saturated solution of
safranin (alcohol soluble) in 95 per cent. alcohol and then adding
an equal volume of water. Anilin Safranin is made by mixing
equal parts of saturated alcoholic solution of safranin with aniline
water. Aqueous Safranin represents a 1 per cent. solution of
safranin (water-soluble) in water. Safranin solutions per se
stain both protoplasmic contents and cell walls red. When
properly used in contrast staining they stain all suberized walls
and lignified walls red.

ScHWEITZER’s REAGENT (Ammoniated Cupric Oxide Test
Solution).—This is a valuable reagent for dissolving cellulose.
It is prepared by dissolving 10 Gm. of copper sulfate in 100 cc.
of distilled water, adding sufficient 20 per cent. solution of sodium
hydroxide to precipitate the copper hydroxide which is collected
on a filter and washing the precipitated copper hydroxide free
from sulfate with cold distilled water. The precipitate, which
must be kept wet during the entire process is then dissolved in
the minimum quantity of ammonia water necessary for complete
solution.

APPENDIX II 687

Sutruric Acrp.—Concentrated sulfuric acid, containing
from 93 to 95 per cent. of H2SO, is useful in the detection of
suberized or corky walls. These resist its action, while lignified
and cellulose walls are completely dissolved by it. This acid as
well as dilutions of it with water are also employed fer se or with
various chemicals in a number of microchemic tests.

Additional reagents and stains are discussed under appropri-
ate headings in this and other chapters of the text.

Tue TECHNIQUE OF Maxkinc A TEMPORARY Mount

1. Place a drop or two of water (or reagent) in the center of a
clean glass slide.

2. With the aid of a forceps take the section or very small
quantity of the material to be examined and spread it on the drop
of water.

3. Place a clean cover-glass over the material. In placing
the cover-glass, do not drop it flat upon the drop of water, but
place one side of it down first and allow it to squeeze the water
along under it.

4. Keep the top of the cover-glass dry.

When filamentous algze or molds are to be examined, the
material tends to cling together and must be carefully separated,
in the drop of water, with dissection needles before the cover-
glass is placed over the material. In case a coarse ground drug is
to be mounted, the coarser particles should be first crushed in
the water on the slide and subsequently teased apart with dissec-
tion needles.

Care should always be taken to see that the water or mounting
medium used is not contaminated with foreign substances. This
can best be practiced by examining the mounting medium under

the microscope before the material to be examined is placed in it.

Tur TecHuNniqgue oF MAkinc PerMANENT Mounts

1. Tue Mountinc Meprum.—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 conditions can be met by adding an antiseptic (2 per cent.

688 PHARMACEUTICAL BOTANY

acetic acid, or formaldehyde) to the water used in mounting and
carefully sealing the cover glass with asphaltum or zinc white.
As a rule, a better way is to use a mounting medium that will not
evaporate, e.g., glycerine, glycerin gelatin or Canada balsam.
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 trans-
parent 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. Glycerin-gelatin has the advan-
tage of mixing readily with 50 per cent. glycerin in which the
object should be placed before being mounted in this medium.
It should be warmed on a water bath before using and the
cover glass applied quickly after it is placed on the specimen. It
cools rapidly and constitutes the quickest and simplest means of
effecting a durable permanent mount. Its disadvantage is due
mainly to its jelly like consistency which is frequently responsible
for damaged mounts when the cover-glass above the preparation
is too greatly strained. Canada balsam slowly becomes solid, so
that the mount is exposed to no accident short of actual breakage.
Balsam has the disadvantage 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, a
clearing oil such as cedar oil, clove oil or xylol must be used as an
intermediate agent between alcohol and balsam.

2. STAINING.—For two reasons it is generally better to stain
plant tissues before mounting. Transparent tissues may become
almost invisible in glycerine, glycerin-gelatin, 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 section with a different
stain (double staining), so that the different parts may be brought
out like areas on a colored map. The most common stains are
hzematoxylin derived from logwood, and various anilin stains—
safranin, fuchsin, eosin, crystal violet, fast green, methyl violet,
light green, iodine green, methyl-green, malachite green, etc.

A number of useful methods of staining and mounting plant
materials will now be discussed. For additional methods the

APPENDIX II 689

reader is referred to “Stain Technology,” published by the
Commission on Standardization of Biological Stains, Geneva,

Di Ys

METHODS FOR THE PREPARATION OF CANADA BALSAM Mounts
No. 1. SAFRANIN AND Fast GREEN

1. Safranin. By means of an eye-dropper put 10 to 15 drops
of a strong alcoholic solution of safranin into a watch glass of
distilled water. Allow the sections to lie in this mixture until
deep red.

_ 2. Transfer the sections to another watch glass containing tap
water and rinse off excess of stain.

3. Dehydrate and differentiate in 50 per cent. alcohol.

4. Dehydrate in 95 per cent. and absolute alcohol (1 minute
in each).

5. Fast Green. Place 5 drops of a solution of fast green (fast
green, 1 Gm., absolute alcohol 25 cc., clove oil, 75 cc.) in a watch
glass of absolute alcohol. Allow the sections to remain in this
solution until the green color stains the softer tissues properly.

6. Clear sections in xylol.

7. Mount in Canada balsam dissolved in xylol.

8. Label slide, indicating subject and stain combination.

This method has given excellent results in staining sections of
a wide variety of fresh and macerated roots, stems and leaves
which have been cut free hand or on the sliding microtome.

No. 2. SAFRANIN AND LicHuT GREEN

1. Stain object with either alcoholic safranin or anilin safranin
(1 per cent. solution in 50 per cent. alcohol) for 1 to 24 hours or by
gently warming object in the safranin solution until vapors arise.

2. Wash out excess of stain with 50 per cent. alcohol.

3. Dehydrate with 95 per cent. alcohol for two minutes.

4. Further dehydrate by placing material in absolute alcohol

for 20 to 30 seconds. ‘ : :
5. Stain with a saturated solution of light green in clove oil for

from 3 to 30 minutes.
6. Clear in xylol for 1 minute. Blot up excess from around

edge of section.

690 PHARMACEUTICAL BOTANY

7. Mount in Canada balsam.

8. Label slide.

The safranin will impart a red color to the suberized and lig-
nified walls, while the light green will stain cellulose walls green.

Should the light green precipitate upon clearing in the xylol,
add 3 cc. of absolute alcohol to each 100 cc. of xylol.

Should air-bubbles be detected in the balsam shortly after
mounting, heat a dissection needle in a flame and touch each
with its tip, when they will be found to disappear.

If too much Canada balsam has been used, some of it usually
spreads beyond the edge of the cover-glass, or on its surface. In
this event wait until the balsam hardens, when it can be scratched
off with a knife, and the surface of the glass cleaned with a rag
moistened with turpentine oil or xylol. i

Should the Canada balsam become too thick, it can be
thinned down with either xylol or benzol.

No. 3. SAFRANIN AND ANILIN BLUE

1. Stain sections with either alcoholic or anilin safranin (1 per
cent. sol. in 50 per cent. alcohol) for from 1 to 24 hours or by
heating them in the staining fluid until vapors arise.

2. Wash out excess of stain with 50 per cent. alcohol. Be
careful not to remove too much of the safranin. Washing should
stop as soon as the stain is reduced to a pink in cellulose walls.

3. Stain with Anilin Blue (1 per cent. sol. in 90 per cent.
alcohol) for from 3 to 10 minutes.

4. Wash in 95 per cent. alcohol for a few seconds.

5. Rinse with slightly acidulated 95 per cent. alcohol for a few
seconds.

6. Wash for 1 to 2 minutes in 95 per cent. alcohol containing a
trace of an alkali to remove the acid.

7. Dehydrate with absolute alcohol for two minutes.

8. Clear in cedar oil for one minute. Blot up excess of cedar
oil from around section. Clove oil or xylol may be substituted
for the cedar oil.

9. Mount in balsam.

10. Label slide.

If the sections are properly stained, the lignified and suberized
walls will be stained red and the cellulose walls a brilliant blue.

APPENDIX II 691

No. 4. SAFRANIN AND METHYL GREEN

1. Stain sections with either alcoholic safranin or anilin
safranin for from 1 to 24 hours or by warming them in the safra-
nin solution until vapors arise.

2. Remove excess of stain by moving sections about in a vessel
of 50 per cent. alcohol until the cellulose walls are pink in color.

3. Stain with 0.5 to 1 per cent. solution of methyl green for
from 30 seconds to 5 or 10 minutes. Good results with this stain
requires experience with the materials to be stained. The
average hand cut section requires 10 minutes.

4. Remove excess of stain and dehydrate in 70 per cent.
alcohol for several seconds.

5. Dehydrate with 95 per cent. alcohol for 10 seconds.

6. Further dehydrate by placing material in absolute alcohol
for one minute.

7. Clear in cedar oil for one minute. Blot up excess of oil
from around edge of section.

8. Mount in Canada balsam.

9. Label slide.
If the sections are properly stained, the lignified and suberized

walls will be stained red and the cellulose walls green.

METHOD FOR THE PREPARATION OF A GLYCERIN-GELATIN Mount

1. Stain object with an aqueous solution of eosin.
2. Transfer to 1 per cent. acetic acid for a minute.
3. Wash out acid and excess of stain by moving the section

about in a dish of water.
4. Transfer object to weak glycerin (glycerin 10 parts, water

90 parts) for 3 to 5 minutes.

5. Transfer object to 50 per cent. glycerin for 3 to 5 minutes.

6. Transfer object to concentrated glycerin for 5 minutes.

7. Remove excess of glycerin around object and mount in
glycerin-gelatin. The slide and cover slip should be warmed
_ before the glycerin-gelatin is dropped over the object and the
cover slip quickly lowered. The preparation of Glycerin-
gelatin is as follows: Macerate 14 grams of gelatin in 84 cc. of
water for 2 hours, add 76 cc. of glycerin and warm; add 2 cc. of
liquefied phenol, warm and stir for 15 minutes untilclear. Filter

692 PHARMACEUTICAL BOTANY

while hot through glass-wool or filter paper and collect the
filtrate in a wide mouthed bottle. Keep well stoppered so as to
exclude dust.

Glycerin-gelatin becomes solid when cool. For use warm the
bottle in a water bath after first removing the stopper. A glass
rod sufficiently long to reach to the bottom of the bottle can be
inserted in the cork and used for transferring the material to the
slide.

8. Ring mount with zinc white, balsam or asphaltum at the
edge of cover slip. If the cover slip is circular, this can best be
done by means of a centering turn-table. A camel’s hair brush is
dipped into the zinc white, balsam or asphaltum and held to the
margin of the cover slip while the slide, fastened with clips to the
turn table, is rotated with it.

9. Label slide.

If the objects or sections are such as not to be liable to shrink they can
be transferred from water or weak glycerin directly to glycerin-gelatin.

TECHNIQUE OF Fixinc, DEHyDRATING, HARDENING AND
IMBEDDING IN PARAFFIN

When the intention is to study the protoplasts in their natural *
form or the processes of cell division, the fresh material must be
put through the various stages of fixation, hardening and imbed-
ding before it is sectioned. The steps will now be considered in
the order in which they must be carried out.

Fixation.—This is the process of killing and coagulating the
protoplast. The essence of good fixation is in rapid killing. It
should be simultaneous with coagulation or hardening so that the
protoplast will not be modified by later treatment. Fixing
fluids are always substances unknown to protoplasm, ¢.g.
poisons. The coagulation of protoplasmic structures is due to
the fact that these are alkaline in reaction whereas the fixing fluid
is acid. Fixing fluids must be judged not only as to killing and
hardening but also as to reaction of tissues to stains afterward.
Fluids that are mixtures make the best fixing agents. Among the °
fixing agents employed are the following: Osmic acid (OsO,)
comes in sealed glass tubes containing 0.5 gm. or 1 gm. It has a
very powerful odor and is easily affected by organic materials.

APPENDIX II 693

It is used in 1 to 2 per cent. solutions and should be made up in
distilled water. It fixes cytoplasm well but the nucleus not as
good. Its disadvantage lies in its inability to penetrate rapidly.

Chromic acid (CrOs) in 0.5 to 1 per cent. aqueous solution is
very favorable for nuclear structure but like osmic acid penetrates
rather slowly. |

Picric acid CsH2(OH)(NOz)s; is one of the most penetrating
fixing fluids but has very little hardening power. It is employed
in saturated aqueous solution.

Corrosive sublimate (HgClz) in 0.2 per cent. aqueous or alcoholic
solution penetrates and hardens rapidly but does not give as sharp
optical differentiation as the others considered.

Absolute alcohol can be employed for very small objects that
are dry. If the objects are moist, shrinkage will follow.

Carnoy fluid, consisting of 6 parts absolute alcohol, 3 parts
formaldehyde and 1 part of glacial acetic acid, can also be used
for fixing small objects. It has the advantage of fixing these in
about 10 minutes. Moreover, the objects can be carried directly
to absolute alcohol, thence to hot melted paraffin and imbedded.

Chrome-acetic-formalin fluid (Craf Fluid). This comparatively
new fluid has been found to give very favorable results in fixing
materials for chromosome studies. The chromosomes are better
defined after staining with Heidenhain’s hematoxylin and give a
good contrast with the cytoplasm. It is prepared by mixing
equal volumes of the following solutions designated as Solution A
and Solution B just before using. Solution A contains chromic
acid, 1 Gm., glacial acetic acid, 7 cc. and distilled water 92 cc.
Solution B contains neutral formalin (40 per cent. formaldehyde
solution) 30 cc. and distilled water 70 cc. The material to be
fixed should be placed in this fluid for from 12 to 24 hours after
which it can be directly transferred to 75 per cent. ethyl alcohol,
changing the solution every 15 mi. for 3 or 4 changes, thence to
85 per cent. alcohol (1 hr.) to 65 cc. of 80 per cent. ethyl alcohol
and 35 cc. n-butyl alcohol (1 hr.) to 45 cc. of 90 per cent. ethyl
alcohol and 55 cc. of n-butyl alcohol (1 hr.) to 25 cc. absolute
ethyl alcohol and 75 cc. n-butyl alcohol (1 hr.) to n-butyl alcohol
for 2 or 3 hours with hourly changes. The material in butyl
alcohol can be infiltrated with paraffin by adding melted

694 : PHARMACEUTICAL BOTANY

paraffin (m.p. 54° to 55°C.) directly to the beaker or other
receptacle containing it and placing the beaker in a paraffin
oven at 56° for several hours. Then replace the butyl-alcohol-
paraffin mixture with pure. melted paraffin for 3 hours and
embed.

Formalin-alcohol, consisting of from 2 to 6 per cent. of formalin
in 50 to 70 per cent. alcohol, is a good fixing and preserving agent
for materials which are not of very delicate texture. It should
not be used for unicellular or filamentous alge and fungi. For
these 2 to 6 per cent. aqueous solutions of formalin have given
good results. Material can be left in the formalin solutions
until required for use, provided care be exercised in storing it in
air-tight containers.

For most materials the Flemming fluids have proven very satis-
factory and are the most generally employed. They are of two
classes, viz.; 1. Those that simply involve chromic acid and acetic
acid (the chrome-acetic fluids) and 2. Those that involve chromic
acid, acetic acid and osmic acid (the Chrome-Osmium-Acetic
Fluids). The formulz follow:

CHROME-ACETIC FLUuIps

Song : per cent. Chromic acid solution......... 100 ce.
Glacial meetie atid sat oe eres ice
1 per cent. Chromic acid solution......... 1Oee;
Medium f per cent: Glacial acetic acid) vi): toeei
Distilled Water ss SE Oa A oes 29 ec
1 per cent. Chromic acid solution......... 25.ce:
Weak 1 per.cent. Glacial acetic acid #2 occur. . 10 cc.
Distilled water oss, bia dae 8 65 cc.
CHROME-OSMIUM-ACETIC _FLUuIDs
1 per cent. Chromic acid solution......... 7500:
Strong , per cent. Osmic acid solution........... 20 cc.
Glacial acetic acid 6.1) ar, ee aes 7c.
1 per cent. Chromic acid solution......... 25 cc.
Weak ss cent. Osmie acid solution; 210.5 10 cc.
ea ee ;
1 per cent. Acetic acid solution. .......... 10 cc.
Disalled waters.) hn ee ee 55:cc.

APPENDIX II 695

The acetic acid in all of the Flemming fluids is of great
advantage since it penetrates very rapidly, carrying the chromic
acid or chromic and osmic acids into the tissue depths, thus
insuring complete fixation. :

The material to be fixed should be cut into small pieces not
longer than 5 mm. nor broader than 20r 3mm. The amount of
fixative to be used should not be less than 15 times the bulk of the
material to be fixed. For delicate material, the weak fixing fluid
is used, while for the slightly woody the medium strength fluid
and for the more woody, the strong fluid is indicated. The
material should be placed in the fixing fluid immediately after it
is gathered. One or two drachm homepathic phials are con-
venient for the process. The material is kept in the fixing fluid
for from 12 to 24 hours and then washed in small cheese cloth
bags which are placed in running tap water for from 6 to 12
hours or over night.

DeHYDRATING AND HARDENING OF SOFTER PLANT MATERIALS.
After washing, the material, still kept in the bags, is placed in 10
per cent. alcohol for 1 hour and is then carried through a series of
alcohols. Each of the series 10 per cent. stronger than the one
before it, remaining in each grade for 11% to 2 hours until 70
per cent. alcohol is reached. Take out of bag and place in phial
in 70 per cent. alcohol. If the material is not to be imbedded in
paraffine immediately, it can remain in 70 per cent. or 85 per
cent. alcohol (if very delicate) until needed. It is not safe to
leave very valuable material in a grade below 70 per cent. over
night. From the 70 per cent. alcohol it is carried to 85 per cent.
to 95 per cent. to absolute alcohol, remaining in each at least 6
hours with 2 or 3 changes of the last.

CLEARING AND IMBEDDING OF SOFTER PLANT MATeErIALs.—In
order to get the material from absolute alcohol into paraffine,
some medium must be used which mixes with absolute alcohol
and which also dissolves paraffine. Either oils such as cedar,
clove or bergamot or substances like xylol, chloroform or benzol
satisfy this requirement. To clear with xylol—transfer material
from absolute alcohol to a mixture of 34 absolute alcohol and 14
xylol for 12 hours, then to mixture of equal parts of absolute
alcohol and xylol for 12 hours, then to 34 xylol and 14 absolute

696 PHARMACEUTICAL BOTANY

alcohol for 12 hours to pure xylol for 12 hours. To phial con-
taining material in pure xylol add paraffine in small pieces and
put on top of paraffine bath sufficiently long until paraffine is
melted. Then add more paraffine and put phial in paraffine
bath at 56°C. over night. Pour fluid off and add pure melted
paraffine and repeat 2 or 3 times until rid of all trace of xylol. A
tray is then prepared by taking a piece of paper and folding up
its edges all around to the height of about a half inch. Half fill
this on a cool surface with melted paraffine. Heat two dissection
needles in bunsen flame and with these dispose pieces of material
in orderly fashion over the crust which has by this time formed at
the bottom of the tray. Blow upon the surface of the paraffine to
harden it more quickly and as soon as the surface crust will bear
it, plunge the tray into cold water. The material can now be
left imbedded in paraffine until required for sectioning.

If cutting is to be done in a cool room, softer grades of paraffine
with melting points between 40° and 50°C. should be used for
imbedding. If, on the other hand, cutting is to be done at sum-

mer temperatures, the harder grades melting at between 55° and
70°C. should be employed.

Use or N-BpuTyYL ALCOHOL IN [MBEDDING Woopy PLANT ORGANS

This method, originally devised by Dr. Conway Zirkle, is
designed to eliminate the hardening and shrinking of tissues
induced by the ethyl alcohol and xylol used in the common
procedure of paraffin embedding. While the common method,
heretofore described, is suitable for the softer plant materials, it is
unsuited for plant materials containing abundant lignified
elements. The higher concentrations of ethyl alcohol and the
xylol so harden the woody elements that they are almost impos-
sible to cut, but break and chip the microtome knife.

Woody plant organs or tissues may be dehydrated and cleared
with a mixture of ethyl and N-butyl alcohols, eliminating the .
use of the higher ethyl alcohols and xylol. N-butyl alcohol dis-
solves sparingly in water but in all proportions in paraffine. A
mixture however of equal parts of ethyl and butyl alcohols is
completely miscible with water. By substituting butyl alcohol
for xylol, the processes of dehydration and paraffine infiltration

eee oe en

APPENDIX II 697

are simplified while hardening and shrinking in many materials
are avoided.

DeEHYDRATION.—For small cubes of woody material, keep one
hour in each of the following solutions (over night in No. 6):

ee ee OS oe ee Se I

Water's cir r Celta Wee to ee eS 25 ARS
95% :B. Ale, ) Hoo. eo ers B40; 20: 30: 35:50: 45... 40. 25
Abs. butyl (03.3), 2 tgsgae 7 ee eee a 208 989) 90, 7 ot LOO 100

Water oe 90 8 75 65 45 25 15 15 5
95% EB. Ale. / Acai ede 5 10 20 30 50 70 80 50 40
Abe. 8, Ales i; a ee ees sly is AS re sae Oi RP Sancza
Abs. Butyl... .i35 6 re Re aS 80, 18 On

Material should remain in pure butyl alcohol until all water is
removed. ‘This usually requires several hours.

INFILTRATION.—Pour paraffin into vial (one-half or two-thirds
full) and allow to cool. Place material in butyl alcohol on top
of paraffin and put vial into oven. After material has stood in
the melted mixture for an hour or so, pour off mixture and add
pure melted paraffin. Repeat after another hour. ‘Two or more
such changes are necessary, depending on the size and nature of
the pieces of material. All the butyl alcohol should be removed
before imbedding, although traces of it do not interfere with
sectioning as do similar amounts of xylol.

TECHNIQUE OF SECTIONING AND MountInG MATERIAL
IMBEDDED IN PARAFFINE

Strip off the paper tray from the imbedded material and cut
out a block of paraffine containing the object which is to be
sectioned, taking care to include at least 2 or 3 mm. of paraffine
on all sides beyond the specimen. ‘Take a segment of pine wood
about an inch long and with a surface at one end about 3 in.
square, and coat the square area with melted paraffine. Warm
the paraffine on the piece of pine wood and quickly press the
paraffine block containing the specimen into this melted paraffine
in the desired position for cutting. Heat a dissecting needle and
apply this all around the base so that the paraffine block is
firmly sealed to the wood. Dip paraffine block in cold water to

698 PHARMACEUTICAL BOTANY

harden. Now trim the paraffine block with a sharp scalpel so
that the faces form right angles with each other. Adjust the
wood in the clamp of the microtome and the microtome blade
so that the top of the paraffine block just touches the near surface
of the microtome knife. Make certain that the knife edge and
the two opposite faces of the paraffine block are perfectly parallel.
Now trim the remaining two sides of the block close to the object.
Adjust the automatic feed of the rotary microtome by moving
dial to number on scale representing thickness in microns desired

of sections and turn wheel of microtome. It will be observed that
the carrier moves up and down and with each downward move-
ment slightly forward, causing the knife to cut sections which
adhere in ribbons.

Transfer the ribbons by means of a camel’s hair pencil or
dissecting needle to a piece of dust free paper with the side down-
ward which was next to the knife. The ribbons are now ready
to be mounted on slides.

The slides to be used should only be those which are devoid of
grease or dirt of any kind particularly on the surface upon which
the ribbons are to be mounted. A very good plan is to keep a
number of slides intended for this purpose submerged in a
saturated solution of potassium dichromate in concentrated
sulfuric acid. ‘These can be taken out as needed and thoroughly
rinsed with water.

With a clean cloth stretched over the forefinger vigorously rub
one surface of each slide until perfectly dry and free of lint. Then
place a small drop of Mayer’s albumin fixative on clean surface
and rub over the surface. (The formula for Mayers Albumin
Fixatiwe is as follows: Egg white and Glycerin, equal parts,
Salicylate of Soda, 1 Gm. to each 100 cc. Mix thoroughly and

- filter.) Now flood the surface with water and cut the ribbons
into segments of the desired length and arrange in rows on slide,
being careful to have the segments somewhat shorter than the
length of the cover slip because of tendency of paraffine to stretch
when warmed. Warm slide gently by holding high above a
bunsen flame or flame of an alcohol lamp until ribbons stretch
out in smooth fashion. Absorb superfluous water, from beneath
ribbons with blotting paper held to their edges and at same time

APPENDIX II 699

push the sections into even rows. Then leave the sections to
dry for several hours or over night.

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.

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. If impossible to extract excess of
safranin by this procedure, use acid alcohol and remove acid by
washing with 50 per cent. alcohol.

6. Place the slide in a well of gentian violet or fast-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.

10. Label slide.

IMBEDDING IN CELLOIDIN

Whenever material is unsuited for free hand sectioning and
will not give good results when imbedded in paraffine on account
of size, hardness, or brittleness, celloidin may be resorted to as an
imbedding medium.

The technique employed is similar to that of the paraffine

method so far as the preliminary fixing, hardening and dehydrat-
ing are concerned up to and including the 95 per cent. alcohol

700 PHARMACEUTICAL BOTANY

stage. From this point the various succeeding steps in the
procedure are as follows:

1. Place material in equal parts of 95 per cent. alcohol and
ether (known as ether-alcohol) for several hours.

2. Transfer to a 2 per cent. solution of celloidin in ether-
alcohol, for 2—5 days.

3. Transfer to a 6 per cent. solution of celloidin in ether-
alcohol, for 2—5 days.

4. Transfer to a 12 per cent. solution of celloidin in ether-
alcohol, for 3-10 days.

5. Prepare a pine block sufficiently large in cross section to
support the material and otherwise adapted to its being clamped
in the object carrier of the microtome. Soak one end of this
block in ether-alcohol for a while and then dip it in the 2 per cent.
celloidin solution.

6. ‘Take the material from the thick celloidin and set it in
proper position, for cutting the sections desired, on the prepared
end of the block and allow the celloidin to thicken for a few
seconds only.

7. Dip the celloidin end into the thick solution; remove and
hold upright so that the new coating may spread out over the
end of the block and solidify the union. :

8. As soon as the celloidin has hardened a little to form a sur-
face film, drop the preparation into a vessel of chloroform and
allow to remain here 1 day.

9. Transfer preparation to a vessel containing equal parts of
glycerin and 95 per cent. alcohol until required for sectioning.

SECTIONING CELLOIDIN MATERIAL

Clamp the block in the sliding microtome and set the knife
obliquely so that the sections can be cut with a long sliding stroke.
Keep the knife and top of the block wet with the alcohol-glycerin
mixture and as soon as the sections are cut, sweep them with a
camel’s hair pencil into a dish of 70 per cent. alcohol. The sec-
tions can be attached to a slide by placing the slide in a closed
chamber over ether. The ether vapor dissolves the celloidin and
causes the sections to adhere to the slide.

APPENDIX II 701

STAINING AND MountTInG CELLOIDIN SECTIONS

1. Place sections in safranin solution for 1 day. This safranin
solution should be made by dissolving as much safranin in 95 per
cent. alcohol as it will take up and then diluting with an equal
quantity of water. '

2. Rinse sections in.50 per cent. alcohol to remove excess of
stain.

3. Transfer them to Delafield’s haematoxylin for 10 minutes
(Delafield’s haematoxylin is made by dissolving 1 gm. of hama-
toxylin in 6 cc. of absolute alcohol and adding this gradually to
100 cc. of a saturated aqueous solution of ammonia alum. This
is left exposed for a week, filtered, 25 cc. each of methyl alcohol
and glycerin added, allowed to stand 6 hours, again filtered, and
ripened about 2 months before using).

4. Rinse sections thoroughly first in water, then in 35 per
cent. alcohol, then in 50 per cent. alcohol.

5. Put them quickly through acid alcohol (1 drop of HCl in
50 cc. of 70 per cent. alcohol).

6. Transfer to 70 per cent. alcohol for about 2 minutes.

7. Transfer to 85 per cent. alcohol for about 2 minutes.

8. Transfer to 95 per cent. alcohol for about 2 minutes.

9. Transfer to absolute alcohol for about 2 minutes.

10. Clear sections in a mixture of equal parts of cedar oil and

phenol for at least 2 minutes.
11. Remove excess of clearing solution and mount in balsam.

12. Label slide.
DestLiciFICATION OF Harp Woopy MarTeErIALs

It frequently happens, even after prolonged maceration or
boiling in alkaline solutions, that thin sections of hard roots,
stems, woods or fruits are difficult or impossible to procure.
This is due to the presence of deposits of silica and other mineral
substances that usually occur in woody tissues. Therefore, it is of
prime importance that these substances be removed as thoroughly
as possible. For this purpose a 10 per cent. aqueous solution of
commercial Hydrofluoric Acid (or stronger solutions up to the
pure acid for very hard materials) is most useful. Small fruits

ge ee oe ge isd ea a gen eg Se ee ee ee RT ee a ee ees

eT ae ei ere ee ee

702 PHARMACEUTICAL BOTANY

or short segments of other hard material are placed in this acid
(which should be kept in a bottle coated internally with a thick
layer of paraffine) for from 3 days to a week, depending on the
size of the objects, with one or two changes of the acid. The acid
is then. washed out thoroughly with running water for 2 to 5
hours. This treatment completely frees the tissues of all mineral
deposits without affecting the organic structure.

ScHULZE’s MACERATION PROCESS

This method is employed for the separation of cells. Radial-
longitudinal sections, that may be cut with a pen knife, are placed
in a beaker or test tube containing 50 cc. of nitric acid of specific
gravity 1.3 (about 2 volumes of nitric acid and 1 volume of water
will serve the purpose). To this add 1 gm. of chlorate of potash
crystals and heat gently until the reddish color which first
appears in the tissues has disappeared. Stop the action by pour-
ing the whole of the contents into a vessel containing water and
wash well with water. The cells can now be readily separated
with dissection needles and mounted in water for examination.
Do not mount in glycerine, for it makes the already bleached
elements too transparent.

Classified List of Reference Works

GENERAL BOTANY

Bergen, J. Y., and B. M. Davis. Principles of Botany. Ginn and Co., Boston,
1906.

Coulter, J. M., C. R. Barnes, and H. C. Cowles. A text-book of botany for colleges
and universities. 3v. American Book Co., New York, 1931.

Eyster, W.H. College Botany. R. Long and R. R. Smith, Inc., New York, 1932.

Gager, C. S. General Botany. P. Blakiston’s Son and Co., Phila., 1926.

Ganong, W. F. A text-book of botany for colleges. Parts 1 and 2. Macmillan
Co., New York, 1919-20.

Holman, R. M., and W. W. Robbins. A text-book of general botany for colleges
and universities. 3 ed. John Wiley and Son’s, New York, 1934.

Kerner von Marilaun, A. The natural history of plants. Their forms, growth,
reproduction and distribution. Transl. by F. W. Oliver. 2v. H. Holt and
Co., New York, 1895.

Mottier, D. B. A text book of botany for college students. P. Blakiston’s Son &
Co., Inc., Phila., 1932.

Robbins, W. J. and H. W. Rickett. Botany. 2ed. D. Van Nostrand Co., Inc.,
New York, 1934.

Sinnot, E. W. Botany: principles and problems. McGraw-Hill Book Co.,
New York, 1923.

Smith, G. M., J. B. Overton and others. A text-book of general botany. Revised
ed. Macmillan Co., New York, 1928.

Strasburger, E. A Textbook of Botany. Rewritten by H. Fitting, H. Seip, R.
Harder and G. Karsten. 6th English ed. Macmillan Co., Nz Ys, 1930.

PLANT MoRPHOLOGY

De Bary, H. A. Comparative morphology and biology of the fungi, mycetozoa
and bacteria. Eng. transl. by H. E. F. Garnsey, rev. by I. B. Balfour. Claren-
don Press, Oxford, 1887.

De Bary, H. A. Comparative anatomy of the vegetative organs of the phanero-
gams and ferns. Transl. and annotated by F. O. Bower and D. H. Scott.
Clarendon Press, Oxford, 1884.

Eames, A. J. Morphology of vascular plants (lower groups). McGraw-Hill
Book Co., N. Y., 1936.

Eames, A. J., and L. H. MacDaniels. An introduction to plant anatomy.
McGraw-Hill Book Co., New York, 1925.

Goebel, K. Organography of plants. Part I: General organography. Eng. ed.
by I. B. Balfour. Clarendon Press, Oxford, 1900. :

Goebel, K. Outlines of the classification and special morphology of plants.
Eng. transl. by H. E. Garnsey, rev. by I. B. Balfour. Clarendon Press,

Oxford, 1887.
703

704 PHARMACEUTICAL BOTANY

Haberlandt, G. Physiological plant anatomy. Transl by M. Drummond.
Macmillan and Co., London, 1914.

. Hitchcock, A. S. The genera of grasses of the United States. U.S. D. A. Bul.
No. 772, 1920.

Jeffrey, E.C. The anatomy of woody plants. Univ. of Chicago Press, Chicago,
1917:

Molisch, Hans. Anatomie der Pflanze. G. Fischer, Jena, 1922.

Scott, D. H. and F. T. Brooks. An introduction to structural botany. Part 1.
Flowering plants. 11led. A. and C. Black, Ltd., London, 1927.

Scott, D. H. and F. T. Brooks. An introduction to structural botany. Part 2.

- Flowerless plants. 9%ed. A. and C. Black, Ltd., London, 1924.

Solereder, H. Systematic Anatomy of the Dicotyledons. Transl. by L. A.
Boodle and F. E, Fritsch. Rev. by D. H. Scott. 2v. Clarendon Press,
Oxford, 1908.

Stevens, W. C. Plant Anatomy from the standpoint of the development of the
tissues and handbook of microtechnic. 4ed. P. Blakiston’s Son and Co.,
Phila., 1924,

Rusby, H. H. A manual of structural botany. Lea and Febiger, Phila., 1911.

PLANT HistoLocy AND Microscopy

Ballard, C. W. Elements of vegetable histology. J. Wiley and Sons, New York,
1921, ;

Chamot, E. M. Elementary Chemical Microscopy. 2ed. New York, 1921.

Gathercoal, E. N. and E. H. Wirth. Pharmacognosy. Lea and Febiger, Phila.,
1937;

Greenish, H. C. The microscopical examination of foods and drugs. 3ed.
J. and A. Churchill, London, 1923.

Hanausek, T. F. and A. L. Winton. The microscopy of technical products. John
Wiley and Sons, New York, 1907..

Mansfield, W. Histology of medicinal plants. John Wiley and Sons, New York,
1916.

Matthews, J. M. The textile fibers. New York, 1924.

Rusby, H. H., and S. E. Jeliffe. Morphology and Histology of plants. New
York, 1899.

Wallis, T. E. Analytical Microscopy, its aims and methods. E. Arnold and Co.,
London, 1923.

Whipple, G. C. Microscopy of drinking water. 3ed. John Wiley and Sons,
New York, 1914.

Winton, A. L. and J. Moeller. Microscopy of vegetable foods. 2ed. J. Wiley
and Sons, New York, 1916.

Youngken, H. W. A textbook of Pharmacognosy. 4ed. P. Blakiston’s Son

and Co., Phila., 1936,

MicroscopicAL TECHNIQUE

Chamberlain, C. J. Methods in plant histology. 4thed. University of Chicago
Press, Chicago, 1924.
Conn, H. J. Biological Stains. 3ed. Geneva, N. Y., 1936.

Bailey, I. W. Méicrotechnique for woody structures. Botanical Gazette 49:
57-58, 1910.

ee Ee Og eee

CLASSIFIED LIST OF REFERENCE WORKS 705

Molisch, H. Mikrochemie der Pflanze. 2te Aufl. F. Fischer, Jena, 1921.

Schneider, H. and A. Zimmermann. Die botanische Mikrotechnik, ein Hand-
buch der mikroskopischen Arbeitsverfahren. 2te Aufl. G. Fischer, Jena,
1922.

Strasburger, E. Das botanische Prakticum. 7te ed. Rev. by Max Koernicke.
G. Fischer, Jena, 1923.

Zimmermann, A. Botanical microtechnique. Transl. by J. E. Humphrey.
H. Holt and Co., New York, 1893.

PLANT TAXONOMY

Abrams, L. An illustrated flora of the Pacific States in 3 volumes. Vol. I,
Ferns to Birthwirts. Stanford University Press, 1923.

American Joint Committee on Horticultural Nomenclature. Standardized plant
names. Salem, 1923. :

Beal, W. J. Michigan Flora. Mich. Agr. Col., 1905.

Britton, N. L. and Addison Brown. An illustrated flora of the northern United
States, Canada and the British possessions. 2nd ed. C. Scribner’s Sons,
New York, 1913.

Coulter, J. M., and A. Nelson. New manual ‘of botany of the central Rocky
Mountain region. (Vascular plants.) American Book Co., New York,
1909.

DeDalla Torre, C. G., and H. Harms. Genera siphonogamarum ad systema
Englerianum conscripta. W. Englemann, Leipsig, 1900-1907.

Engler, A. and K. Prantl. Die Naturlichen Pflanzenfamilien nebst ihren Gattun-
gen u. wichtigeren Arten, insbesond. der Naturpflanzen. 23v. W. Engelmann,

Leipsig, 1900-1925; 2ed., 1930-
Engler, A., and E. Gilg. Syllabus der Pflanzenfamilien. 4te and 10te Aufl.

_ Gebruder Borntraeger, Berlin, 1924,
Engler, A. Das Pflanzenreich regni vegetabilis conspectus. (Issued serially).
Heft 1-90, 1900-1927. W. Engelmann, Leipsig, 1900— is
Gerth Van Wijk, H. L. A dictionary of plant names. 2v. M. Nijhoff, 1916.
Gray, Asa. Gray’s new manual of botany. Handbook of the flowering plants
and ferns of central and northeastern United States and adjacent Canada.
7th ed. Revised by B. L. Robinson and M. L. Fernald. American Book

Co., New York, 1908.

Gray, Asa. Synoptical flora of No
& Co., 1886.

Hitchcock, A. S. Methods of d
Sons, New York, 1925.

Hutchinson, J. Families of cpaping hasan

. Macmillan Co., London, ; .
eae ed Av, 1893-1895 and 8 supplements, 1911-1936. Clarendon

rth America. 2ed. Ivison, Blakeman, Taylor
escriptive systematic botany. John Wiley and

I. Dicotyledons. II. Monocoty-

Press, Oxford.
Jackson, B. D. Guide to the literature of botany. Longmans Green and Co.,
London, 1881. of the fowerlog plants of California. Associated

Jepson, W. L. A manual ak
? San Francisco : a
Le Ryssctetesones' ae Decale. General system of botany, descriptive and

analytical. Transl. and ed. by J. D. Hooker. London, 1872.

706 PHARMACEUTICAL BOTANY

Lyons, A. B. Plant names, scientific and popular. Nelson Baker, Detroit, 1900.

Pfeiffer, L. Nomenclator botanicus. 2v. T. Fischer, Kassel, 1873-1874.

Pool, R. J. Flowers and flowering plants. led. McGraw-Hill Book Co., N. Y.,
1929.

Pritzel, G. A. Thesaurus literaturae botanicae . .. 2ed. F. O. Brockhaus,
Leipsig, 1872.

Rydberg, P. Flora of the Rocky Mountains and dinates plains. 2nd ed.
New York, 1922.

Small, J. K. Flora of the Southeastern United States. New York, 1903.

Swingle, D. B. A textbook of systematic botany. led. McGraw Hill Book Co.
New York, 1928.

Wettstein, R. Handbuch der Systematischen Botanik. 3te Aufl. ‘F. Deuticke,
Leipsig und Wien, 1924.

Wiegand, K. M. and A. J. Eames. The flora of the Cayuga Lake Basin, New
York. Cornell Univ. Ag. Exp. Sta. Memoir 92, 1925.

CyToLocy AND GENETICS

Babcock, E. B. and R. E. Clausen. Genetics in relation to agriculture. 2nd ed.
McGraw-Hill Co., New York, 1927.

Castle, W. E. Genetics and eugenics. 4ed. Harvard Univ. Press, 1931.

Meyer, A. Morphologische und Physiologische Analyse d. Zelle d. Pflanzen u.
Tiere. G. Fischer, Jena, 1920.

Newman, H. H. Evolution, Genetics and Eugenics. 3ed. Univ. of Chicago
Press, 1932.

Sharp, L. W. An introduction to cytology. 2nd ed. Rev. McGraw-Hill Book
Co., New York, 1926.

BACTERIOLOGY AND MICROBIOLOGY

Bergey, D. H. Manual of Determinative Bacteriology, 4th ed. Williams and
Wilkins Co., Baltimore, 1934.

Conn, H. W. and H. J. Conn. Bacteriology. 2nd ed. Williams and Wilkins
Co., Baltimore, 1924.

Jordan, E. O. A text-book of general bacteriology. 11th ed. W. B. Saunder’s
Co., Phila., 1935.

Marshall, C. E. Microbiology. A text-book of microorganisms general and
applied. 2ed. Rev. P. Blakistons Son and Co., Phila., 1917.

Park, W. H., A. W. Williams and C. Krumwiede. Pathogenic microorganisms.
7th ed. Rev. Lea and Febiger, Phila., 1920.

Rice, T. B. Text Book of Bacteriology. W. B. Saunders Co., Phila., 1935.

Schneider, A. The microbiology and microanalysis of foods. P. Blakiston’s
Son and Co., Phila., 1920.

Zinsser, H. A pent Hiei of bacteriology. 5th ed. D. ae and Co., New
York, 1925,

Druc PLANT CULTIVATION

Corbett, L. C. The propagation of plants. Farmers’ Bulletin No. 157, U. S.
D. A. Government Printing Office, Washington, 1917.
Corbett, L.C. Pruning. Farmer’s Bul. No. 181, U.S. D. A., 1915.

CLASSIFIED LIST OF REFERENCE WORKS 707

Dillman, A.C. The production of seed flax. U.S. D. A. Farmer’s Bul, No. 1328,
1924.

Henkel, Alice. Weeds used in medicine. U. S. D. A. Farmer’s Bul. No. 188,
1917,

Henkel, Alice. American root drugs. U.S. D. A. Bur. Pl. Ind. Bul. 107, 1907.

Hood, S. C. Commercial production of thymol from horsemint (Monarda
punctata). Bul. No. 372, U.S. D. A.

Kraemer, H. Cultivation of medicinal plants, pp. 727-748, in Applied and
Economic Botany, Phila., 1914.

Kilmer, F. B. The cultivertoll of medicinal plants. Am. Jour. Pharm., 87:
343-359 and 421-435, 1915. (This article includes an extensive citation of
the literature on the subject.)

Rabak, F. The production of volatile oil and perfumery plants in the United
States. U.S. D. A. Bur. Pl. Ind. Bul. No. 195, 1910.

Rixford, G. P. Smyrna Fig culture. U.S. D. A. Bul. No. 732, 1918.

Russell, G. A. Drying crude drugs. U.S. D. A. Farmer’s Bul. No. 1231, 1921.

Stansel, R. H. and R. H. Wyche. Fig culture in the Gulf Coast region of Texas.

Tex. Agr. Exp. Sta. Bul. no. 466, 1932.
Stockberger, W. W. Drug plants under culivation: Farmer’s Bul. No. 663,

U.S. D. A, 1915.
Stockberger, W. W. Ginseng culture. U.S. D. A. Farmer’s Bul. No. 1184, 1928.
Stockberger, W. W. Growing and curing hops. U. S. D. A. Farmer’s Bul.
No. 304, 1928.

Shrader, J. H. The castor oil industry. Bul. No. 867, U. =. 2. A. 1920.
Sievers, A. D. Some effects of selection on the production of alkaloids in bella-

donna. Bul. No. 306, U. 8. D. A., 1915. °
Sievers, A. F. American medicinal plants of commercial i jcilpia tice: iS: DA:

misc, publ. 77, 1930.
Sievers, Ae Peppermint and Spearmint as farm crops. U. S. D. A. Farmer’s

Bul. No. 1555, 1929.
Teetgen, A. B. Profitable herb growing and collecting. Country Life, London,

1919, ee : os.
Tschirch, A. Handbuch der et ci 2ed. Chr. H. Tauchnitz, Leipsig,
1930, ;
Van Fleet, W. Goldenseal ae cultivation. U.S. D. A. Farmer’s Bul. No. 613,

Wided ak N. Almond varieties in the U. S. U. S. D. A. Dept. Bul. No. 1282,
1924,
! Pant Ecotocy AND GEOGRAPHY

Campbell D. H. An outline of plant geography. Macmillan and Co., New

xy Nee ee . methods in ecology. University Publishing Co.,
ements,

Lincoln, , Neale ~~ on of Natural Conditions of the Ecological Society
Committee on te a ide to the Americas. Edited by V. E. Shelford

_ Naturalists’ gui
a ae veilliaans and Wilkins Co., Baltimore, 1926.

Engler, A., and O. Drude. Die Vegetation der Erde. 15v.
gier,
Leipsig, 1896-1923.

W. Engelmann,

708 PHARMACEUTICAL BOTANY

Fernald, M. L. Persistence of plants in unglaciated areas of Boreal America.
Memoirs of the American Academy of Arts and Sciences. 15: No. 3, 2a9=
342, 1925.

Grisebach, A. Die Vegetation der Erde nach ihrer Klimatischen anordnung.
2v. 2te. Aufl. W. Engelmann, Leipsig, 1884.

Hilgard, E. W. Soils, their formation, properties, composition, and relation to
climate and plant growth. Macmillan and Co., New York, 1906.

Knuth, P. Handbook of flower pollination. Transl. by J. R. A. Davis. 3v.
Clarendon Press, Oxford, 1906-1909.

Scharff, R. F. Distribution and origin of life in America. Macmillan Co., New
York, 1912.

Schimper, A. F. W. Pflanzen-Geographie auf Physiologischer Grundlage. G.
Fischer, Jena, 1898.

Warming, E. and M. Vahl. Ecology of plants. An introduction to the study of
plant communities. Transl. by P. Groom and I. B. Balfour. Clarendon
Press, Oxford, 1909.

Weaver, J. E. and F. E, Clements. Plant Ecology. McGraw-Hill Book Co.,
Inc., New York, 1929.

PLANT PuysIoLoGy AND PLANT CHEMISTRY

Detmer, W. Practical plant physiology. Transl. by S.A. Moor. Swan, Sonnen-
schein and Co., London and Macmillan Co., New York, 1909.

Duggar, B. M. Plant physiology with special reference to plant production.
Macmillan Co., New York, 1911.

Green, J. R. An introduction to vegetable physiology. 3rd ed. J. and A.
Churchill, London. P. Blakiston’s Son and Co., Phila., TOLL,

Haberlandt, G. F. J. Physiologie und Okologie. B. J. Teubner, Leipsig, 1917.

Hass, P., and T. G. Hill. An introduction to the chemistry of plant products.
3rd ed. 2v. Longmans, Green and Co., 1921 and 1922.

Henry, T. A. The plant alkaloids. 2nded. J. and A. Churchill, London, 1924.

Jost, L. Lectures on plant physiology. Transl. by R. J. H. Gibson and supple-
ment. Clarendon Press, Oxford, 1907 and 1913.

Maximov, N. A. Textbook of Plant Physiology. McGraw-Hill Book Co., Inc.,
New York, 1930.

Oppenheimer, C. Die Fermente und ihre Wirkung. 5te Aufl. Issued serially.
Georg Thieme, Leipsig, 1924- :

Palladin, V. I., and B. L. Livingston. Palladin’s Plant Physiology. 3rd Amer.
ed. P. Blakiston’s Son and Co., Phila., 1926.

Pfeffer, W. The physiology of plants. A treatise upon the metabolism and
sources of energy in plants. Transl. by A. J. Ewart. 3v. Clarendon Press,
Oxford, 1900 and 1906.

Tummann, O. and L. Rosenthaler. Pflanzenmikrochemie. Gebr. Borntrzger,
Berlin, 1931.

HortTICcULTURE AND AGRONOMY
Bailey, L. H. Standard Cyclopedia of Horticulture. 6v. Macmillan Co.,

New York, 1900-1917.
Bailey, L. H. A manual of cultivated plants. Macmillan Co., New York, 1924.

CLASSIFIED LIST OF REFERENCE WORKS 709

Harshberger, J. W. Textbook of pastoral and agricultural botany. P. Blakiston’s
Son and Co., Phila., 1920.

Meehan, T. The native flowers and ferns of the United States. 2v. Robson,
Phila., 1880.

Rehder, A. Manual of the cultivated trees and shrubs hardy in North America.
Macmillan Co., New York, 1927.

Robbins, W. W. Botany of crop plants. 2ed. P. Blakiston’s Son and Co.,
Phila., 1924.

MycoLocy

Atkinson, G. F. Mushrooms, edible, poisonous, etc. Andrus and Church,
Ithaca, 1900.

Buller, A. H. R. Researches on Fungi. 3v. Vol. 1, 1909, Vol. 2, 1922, Vol. 3,
1924. Longmans Green and Co., London.

Duggar, B. M. Fungous diseases of plants. Ginn and Co., Boston, 1909.

Harshberger, J. W. A text-book of mycology and plant pathology. P. Blakis-
ton’s Son and Co., Phila., 1917.

Henrici, A. T. Molds, yeasts and actinomycetes. J. Wiley Sons, Inc., N. Y.,
1930.

Heald, F. D. Manual of plant diseases. 2ed. McGraw-Hill Co., N.Y;, 1932.

Mcllvaine, C. One thousand American fungi. Bowen-Merrill Co., Indianapolis,
1900.

MEDICINAL AND Potsonous PLANTS

Bentley, R. and H. Trimen. Medicinal plants. 4v. jJ. and A. Churchill,
London, 1880.

Bernhard-Smith, A. Poisonous plants of all countries. Bailliere, Tindall and
Cox, London, 1923.

Long, H. C. Plants poisonous to live stock. Univ. Press, Cambridge, 1917.

Millspaugh, C. F. American medicinal plants. 2v. Boericke, New York, 1887.

Nees von Esenbeck, T. F. L. Plante medicinales. 3v. Arnz, Dusseldorf,
1828. ;

Pammell, L. H. A manual of poisonous plants. The Torch Press, Cedar Rapids,
Ia. Part I, 1910, Part I, 1911.

Wood, H. C., C. H. LaWall, H. W. Youngken, I. Griffith, A. Osol and L. Gershen-
feld. The United States Dispensatory. 22nd ed. J. B. Lippincott Co.,
Phila., 1937.

Woodville, W. Medical botany. 5v. 3ed. Bohn, London, 1832.

Youngken, H. W. A text-book of pharmacognosy. 4th ed. P. Blakiston’s Son

and Co., Phila., 1936.
History OF BOTANY
Harvey-Gibson, R. J. Outlines of the history of botany. A. and C. Black,

London, 1919.

Green, J. R. History of botany, 18
Green, E. L. Landmarks of botanical history. Smithsoni

ington, 1909.
von Sachs, J. History of botany, 1530-1860. Rev. by I. B. Balfour. Clarendon

Press, Oxford, 1906.

60-1900. Clarendon Press, Oxford, 1909.
an Institution, Wash-

Glossary

Axor’TION.—The imperfect or non-development of an organ.

ACAULES’CENT.—Without an obvious aerial stem.

Acuene’ (akene).—A small, dry, one-celled, indehiscent fruit in which the seed
coat and pericarp (fruit wall) are not firmly attached.

AcHLAMY’DEOus.—Destitute of calyx and corolla.

Acic’ULAR.—Applied to crystals of calcium oxalate, etc., that are needle-shaped.

Acips (Plant Acids).—A widely diverse group of cell constituents having the prop-
erty of uniting with radicals to form salts or esters.

Acrop’ETAL.—Development from outside (below) toward the inside (above).

Actinomor’pHic.—Applied to flowers like those of the Mustard, etc. which are
capable of division into equal halves in more than one direction. Cf.
zygomorphic.

Acu’mInATE.—Tapering gradually to a long point.

Acutr’.—Sharp-pointed, the point being less than a right angle.

Ap’NATE.—Applied to the growing together of unlike parts.

Apventi’T10us.—Applied to roots and buds that are out of their ordinary position.

Arstiva't10on.—Arrangement of the parts of the flower in the bud.

A.su’men.—Nutritive material stored in the embryo, endosperm, or perisperm.

ALBuUR’NUM.—Sapwood.

ALEU’RONE.—Applied to small protein bodies in which are usually embedded one
or more crystalloids, one or more globoids and at times a small rosette of cal-

cium oxalate.

ALKALoIs.—Nitrogenous organic bases.

ALLELO’MORPHS.—Alternative forms of a gene which are located at the same point
on each one of a pair of chromosomes.

AL'TERNATE.—Applied to leaves, buds, etc., that are arranged singly (one after

another) at the nodes.
Am’ent.—A scaly, spike-like inflorescence. Another name for catkin.

Amor’pHous.—Without definite shape.
Ampuit’Ropous (ovules and seeds).—Half-inverted and straight, with the hilum

about the middle, and micropyle terminal.

AmpLex’/ICAUL.—Clasping the stem.

ANAL'ocy.—Resemblance in function.

Anastomo’sinc.—Applied to veins that are connected with others by cross veins,
so forming a network, as with the marginal veins of Eucalyptus.

Anat’ropous.—Inverted ovules or seeds with micropyle adjacent to hilum.

Apre’crum.— The male system of organs in a flower.

Anproa’ynous.—Applied to inflorescences composed of both staminate and

pistillate flowers.

i’Lous.—Wind pollinated.
ce as arewas sein ovules and seeds borne within a box-like covering, the

se 71

712 PHARMACEUTICAL BOTANY

ANn’NUAL.—Producing flowers, fruit and seed within a year from the time the seed
germinated and then dying completely.

AN’NULAR.—Ring-like.

ANTE’RIoR.—The front region.

An’THER.—That portion of a stamen which bears the pollen. ‘

AN’THERIDIUM.— Male sexual organ of Thallophytes, Bryophytes and Pteridophytes.

AN’THEROzOID.—A male sexual cell formed within an antheridium.

An’THOPHORE.—A lengthened internode of the receptacle between calyx and corolla.

ApeEtT’ALOus.—Without petals, as in the oaks, etc.

Apocar’pous.—Carpels separate and distinct.

Apoc’Amy.—The direct development of a sporophyte from a gametophyte without
the functioning of sexual organs or cells.

ApopeEt’ALous.—Petals separate and distinct.

APposEP’ALOUs.—Sepals separate and distinct.

Apotue’crum.—A disk like or cup-shaped fruiting organ of a lichen and some
cup-fungi.

ARCHEGO’NIuUM.—A multicellular female sexual organ found in the bryophytes,
pteridophytes and gymnosperms.

Ar’tL.—An accessory seed covering outside of the testa and arising at or about
the hilum, as in Evonymus.

Ar’ILLopE.—A fake accessory seed covering outside of the testa, as in Nutiinby;
and arising from the dilatation of the micropyle.

Aris’TATE.—Having a stiff bristle-like termination.

ASCEND’ING.—Growing obliquely upward.

As’cus.—A sac-like spore case of an Ascomycete fungus.

Art’AvismM.—Reversion to ancestral type. The reappearance of a character after a
lapse of one or more generations.

Auric’/ULATE.—Ear-like.

Awn.—A bristle-like structure that branches along its axis. .

Ax’1.—Upper angle formed where the leaf joins the stem.

Ax’ILLARY.—In the axil.

BAc’cATe.—Berry-like.

BAL’sAM.—A mixture of resin with cinnamic or benzoic acid or both and a volatile
oil.

BArs.—A short bristle usually bent back.

BARK.—All that portion of an exogenous plant axis outside of the cambium line.

BAst.—Applied to the phloem region but mainly to the fibrous portion thereof.

Beav’ep.—Applied to a thickened, jointed cell wall showing in sectional view some-
what circular areas in the wall like a row of beads, due to a close proximity of
the pores in the wall.

BeArp’Ep.—Furnished with long hairs.

Ber’ry.—A fleshy fruit whose mesocarp and endocarp are fleshy and frequently
succulent throughout, and with seeds imbedded therein, as tomato, capsi-
cum, belladonna, etc.

Br.—A prefix of the Latin language indicating two, twice or doubly.

Br’coLLAT’ERAL Bun’pLE.—A collateral bundle with the xylem strand enclosed
between two phloem strands, cambium being present or absent on either outer
or inner face of the xylem or on both faces,

GLOSSARY 743

Bren’NtAL.—Applied to plants that live for more than one year but not longer than
two years.

BiLA’BIATE.—T wo lipped.

BLApE.—Expanded part of a leaf.

BLoom.—The whitish and waxy secretion of epidermal cells, as in the stems of
Sugar Cane or the leaves of Cabbage.

Borke.—The dead bark tissues which arise when secondary phellogen forms cork in
deep regions of the bark.

BrAct.—A modified leaf, frequently scale-like, appearing on inflorescence axes.

Brac’TEOLE (bracteolar leaf).—A modified leaf found on pedicels.

Bup.—A rudimentary stem.

BuLs.—A very short scaly underground stem.

Buv’si_s.—Small underground bulbs, as in garlic,

Buv’BLeEtTs.—Small above ground bulbs, as in the tree onions.

Bun’pLe.—See Vascular Bundle. Also applied to a more or less cylindrical group of
any parallel, elongated cells or other objects.

Bun’pLE SHEATH.—An endodermis or a border parenchyma enclosing a single
bundle.

Capvu’cous.—Falling with the opening of the flower, as the calyx of Papaver.

Ca’/Lyx.—The outermost whorl of floral leaves,

Cam’sium.—A secondary meristem lying between the phloem and xylem in certain
collateral bundles (fascicular or intrafascicular cambium) or between the
phloem and xylem portions of medullary rays (interfascicular cambium).

CAmpAn’uULATE.—Bell shaped.

Campy.o’rropous.—Applied to ovules or seeds that are curved so as to bring the
apex and base near together.

CANEs’cENT.—White or gray from a coating of fine hairs.

Caour’cuouc.—A tenacious, elastic hydrocarbon found in the latex of many plants.

Capruur’1um.—A network of filaments among spores, as in the sporangium of
slime molds, the fruit body of puff balls, etc.

Cap’rrATEe.—Shaped like a head.

Cap’RiFICATION.—The process of pollinating figs artificially.

Cap’suLe.—A dry, dehiscent fruit of a compound pistil consisting of two or more
carpels.

Carpo’Hy’DRATE.—A substance consisting of carbon and of hydrogen and oxygen in

the proportion of two parts of hydrogen to one part of oxygen.

Car’PEL.—A transformed leaf bearing one or more ovules; a simple pistil; a part of a

nd pistil.

Gust unicellular, female sexual organ of certain red alge and fungi
consisting of a cell enlarged at its base, its terminal part ending in a long
filament called a trichogyne. ‘

Car’poruore.—A slender stalk, the prolongation of the receptacle, to which the

(mericarps) of the Umbellifera are attached.

i ior akenes
ee warty excrescence at or about the hilum of certain seeds, as that of

il seed, etc.
ee dry, indehiscent, one-seeded fruit of the grasses or cere Pare

the fruit wall (pericarp) and seed coat firmly adhere.
Car’xin.—A scaly spike of flowers.

714 PHARMACEUTICAL BOTANY

Cau’paTe.— Tailed.

CauLgs’cENT.—With an obvious aerial stem.

Cau’LINE.—Pertaining to the stem.

Ce...—The structural unit of plants and animals consisting of a protoplast which in
most plants is surrounded by a cell wall secreted by the protoplast.

Crentrir’UGAL.—Applied to a flower cluster in which the terminal or central flower
blossoms first.

CentTRIP’ETAL.—Applied to a flower cluster in which the lower or outer flowers
bloom first.

Cuarr.—The glumes and palets of grains; the scaly hairs on the stipes of ferns;
the bracts subtending each floret in some heads of Composite.

CuaAa’zA.—That portion of the ovule marked by the junction of the integuments
with the nucellus. 5

Cnasmo’GAMous.—Pertaining to flowers that regularly open.

Cuiamy’DosPoRE.—Thick walled spore formed within the hyphz of smuts.

CuLo/ROPHYLL.—The mixed pigments forming the green color of all green plants
and found in the chloroplastids.

Cutoropias’tIp.—A protoplasmic body in the cells of green parts of plants con-
taining chlorophyll.

Curo/MATIN.—That portion of the nucleus which is readily colored by a basic
dye. The substance that carries the hereditary characters from parent to

offspring.

CHROMO/MERE.—One of the chromatin granules, arranged linearly like the beads on
a string, comprising the chromosome.

Cxuromoptas't1p.—A protoplasmic body in the cells of certain parts of plants con-
taining a pigment other than chlorophyll.

Curo’/MosoME.—One of the bodies into which the chromatin of the nucleus is
resolved during indirect nuclear division. The chromosomes carry the genes,
linearly arranged, which control the.development of Medelian characters.

Ci14.—Vibratory, hair-like, protoplasmic outgrowths of zodspores, bacteria,
gametes, etc.

CrrcuMnuTA’TION.—The repeated bending in different directions of the growing

tips of stems of climbing plants.

Crr’cinATE.—Rolled inward from apex toward base, as the young leaves of ferns.

Circumscis’stLE.—Applied to the splitting open of capsules transversely into lid and
pot portions.

CLAp’ope.—The flattened branch which somewhat resembles a leaf.

CiAw.—The narrowed base of some petals, as those of the Pink Family.

Ce1sroc’AMous.—Applied to flowers that never open but are self fertilized, as in
some Polygalas and Violets.

Cocn’LeA.—A spirally coiled legume.

Cor’nocyTe.—A multinucleate cell.

Coue’ston.—The union of parts of the same whorl.

Co’Hort.—A group of natural orders.

Co.rorui’zA.—A root sheath through which the radicle bursts in germination.

CoLLAt’ERAL.—Applied to fibrovascular bundles in which the phloem and xylem
masses are arranged side by side.

CoLLen’cHYMA.—Tissue composed of elongated prismatic cells thickened at
their angles with a colloidal substance.

GLOSSARY 715

CotuMeL’LA,—The end cell wall of an aerial hypha that bulges into the sporan-
gium; also applied to the axis of a capsule.

Co.’umn.—The united stamens and carpels in Orchids.

Co.’umMN CELis.—Hour-glass shaped cells forming a layer beneath the palisade-
epidermal layer of leguminous seeds.

Co’ma.—A tuft of hairs, as found on the seeds of Milkweeds.

Com’missuRE.—The contiguous surfaces of two carpels as in the flowers and fruits
of the Parsley Family. ;

CompAn’Ion CELLs.—Parenchyma cells associated with sieve tubes and having a
narrow cavity, abundant protoplasm, large nuclei and the wall adjoining the
sieve tube marked by transversely elongated pits.

ConcEn’TrIic.—Applied to several circles or whorls one within the other. Con-
centric vascular bundles are those in which the xylem mass surrounds the

_ phloem mass or vice versa.

Concep’TACLE.—A sac bearing the fruiting organs in certain Alga and Fungi.

Con’puct’Inc Bun’pLE.—(See Vascular Bundle.)

Con’pucT’ING PAREN’cHYMA.—Somewhat elongated parenchyma cells serving
primarily for the conduction of fluids.

Conpu’PLICATE.—Folded together lengthwise, as for example the bud leaves of
the oak or peach.

Con 'tA.—Asexual spores cut off from the ends of hyphz or sterigmata by Penicil-
lium, Aspergillus, Peronospora, Claviceps, etc.

Conw’1opHore.—A hypha bearing conidia.

Conyuca’r10n.—One of the sexual methods of reproduction where two like sexual

cells unite to form a zygospore.
Con’NATE.—Applied to parts that have grown together, as the bases of two opposite

leaves.

ConnEctT’IvE.—The continuation of the filament of the stamen that connects the
two lobes of the anther. .

Connt’vENT.—Brought close together; converging.

Con’voLutrE.—Rolled lengthwise from one edge, as the leaves in the buds of the
Wild Cherry and Plum.

Cor’DATE.—Heart-shaped.

CorrA’/cEous.—Leathery in texture.

Cork (Su’BEROus TissuE).—A protective tissue derived from the phellogen and with
cell walls waterproofed with suberin.

Cork Camp’1um.—(See Phellogen.)

Corm.—A solid, swollen, fleshy underground stem.

Coro’LA.—The inner whorl of floral envelopes composed of petals.

Coro’na.—A crown like appendage in the throat of the corolla, as in the flowers
of Narcissus and Silene. :

Cor’TEx.—That region in dicotyl and gymnosperm roots of primary growth and
in roots and stems of monocotyledons between the outer protective tissue or
epidermis and the pericycle; in dicotyl and gymmosperm roots of secondary
growth or in barks between phellogen and phloem or pericycle. The Primary
Cortex is the region lying between the pericycle and the epidermis or the
secondary tissues produced by the phellogen. The SeconpAry Cortex is
all that tissue laid down by the phellogen on its inner face.

Cor’TICAL PAREN’CHYMA.—The parenchyma of the cortex.

716 PHARMACEUTICAL BOTANY

Cor’yms.—A flat topped or convex centripetal inflorescence with the lowermost
pedicels the longest.
Cos’tA.—A rib.
Coty.e’pon.—A seed-leaf of the embryo.
Crem’ocArp.—The peculiar fruit of Umbellifere, consisting of two inferior akenes
(mericarps) separated from each other by a carpophore.
_Cre/NATE.—Applied to leaf margins having rounded teeth.
CrEN’ULATE.—The margin with fine rounded teeth.
Crip’/RIFORM.—Sieve-like.
Cru’/crrorM.—Applied to the corolla or the calyx of flowers, the parts of which are
arranged in the form of a cross.
Crusta’ceous.—Applied to the thallus of a lichen that closely adheres to the
substratum.
Cryp’rocAM.—A plant belonging to one of the divisions of the vegetable kingdom
below the Spermatophytes.
Crys’tAts.—The solid geometrical forms assumed by many chemically homo-
geneous bodies.
Crys’TAL SAND.—See Microcrystals.
Crys’TALLOID.—A protein body found in the aleurone grains of seed or under-
ground parts.
Cu_m.—A jointed stem of a grass or sedge.
Cu’NEATE.—Wedge-shaped.
Cu’puLte.—Applied to the concave involucre enclosing the glans of an acorn but
also to other cup-shaped parts of plants.
Cu’ticLeE.—A thin covering of a waxy substance called cutin on the outer wall of
epidermal cells.
Cu’t1n.—A fatty cell-wall constituent serving to waterproof the wall.
Cutini’zEp.—Applied to a cell-wall infiltrated or covered with cutin.
Cus’pIDATE.—Tipped with a sharp rigid point.
Cyme.—A more or less flat topped determinate inflorescence.
Cy’mose.—Cyme-like.
Cys’ro.irHs.—Cell bodies consisting of deposits of calcium carbonate or silica or
both on an extention of the cell wall into the cell. ;
Cy’TrAsE.—An enzyme which digests cellulose.
Cyto.’ocy.—The study of cells and their contents.
Cy’ropLAsM.—The cell protoplasm outside of the nucleus.

Decan’pRous.—Having ten stamens.

Dectp’vous.—Applied to leaves which fall in autumn, to plants bearing such leaves
and to the calyx and corolla which fall shortly after blossoming, before the
development of the fruit.

Dec’LinATE.—Curved or bent downward.

Decompounp’.—Several times compounded, as the leaf-blades of Cimicifuga.

Decum’sent.—Erect at base, then lying on the ground, with the end rising.

Decus’sAte.—Applied to opposite leaves when the pairs stand at “ee angles to
each other along the stem.

Dents’cence.—Splitting open.

De iqueEs’cenT.—Applied to a tree whose trunk or main stem is lost in branches.

Dev’trom.—Having the shape of the Greek letter A.

GLOSSARY 747

Den'TrATE.—Having broad, acute, marginal teeth pointing outward.

Dentic’ULATE.—Finely dentate.

DermAt’oceN.—The generative tissue that gives rise to epidermis.

DertTER’MINATE.—Applied to inflorescences on which flowering begins with the ter-
minal] bud, thus ending the elongation of the stem bearing the flowers.

D1AvEL’pHous.—Applied to stamens whose filaments are united at their edges into
two sets.

DrAcGeor’Rropic.—Applied to a plant organ that assumes a horizontal position.

DrAn’pRous.—Possessing two stamens.

Dr’AstAse.—A ferment found in germinating seeds and fungal hyphae which changes
starch into maltose.

DicuLAMyp’£0us.—Pertaining to flowers that possess both calyx and corolla.

Dicuoc’AMy.—The maturation of one set of sexual organs before the other.

Dicuot’omous.—F orked.

Dic’L1Nous.—Pertaining to the stamens and carpels being found in separate flowers.

Dicor’yLe’pon.—A plant whose embryo possesses two seed leaves or cotyledons.

Dic’1raTe.—Referring to a compound leaf whose leaflets come off at the end of the
petiole.

Dimor’puisM.—Having two forms of flowers, one with long styles and short stamens,
the other with short styles and long stamens; the occurrence of two distinct
forms.

Dic:/crous.—Applied to species having two kinds of individuals, male and female.

DissEct’ED.—Cut deeply into numerous divisions.

DissEPi’MENT.—A partition separating cavities in a compound ovary or fruit.

Dis’r1cHous.—Pertaining to the arrangement of leaves in two rows.

Drvi’pEp.—Segmented to the mid-rib or base..

DorstvEN’rRAL.—Having distinct upper and lower surfaces.

Dor’sum.—The back of an organ. The lower surface of a foliage or floral leaf.

Down’y.—Covered densely with soft hairs.
Drupe.—A one-celled, one-seeded fruit whose endocarp is stony.

Drupe’Let.—A small drupe. ;

Ducr.—A tubular element found in the xylem region of a vascular bundle. See
trachea.

Dura’MEN.—Heartwood.

E- or Ex-, a prefix meaning devoid of, outside of, or away from.
Eccen’tric.—Deviating from the center. Applied to the hila of starch grains
which are outside of the center, also to woody plants which develop more —

rapidly on one side than on the other. ;
Ecuin’uLATE.—Beset with small prickles or spines.
Ecu’InATE.—Beset with prickles or spines.
Ec’roptAsm (Outer Plasma Membrane).
limiting layer of the cytoplasm.
Eoc-AppARA’TUs.—The ovum and two
embryo-sac.
ELABORATED SAP.
cells of leaves which
istributed to needing it.
sitios eds which stores oil or fat.

—The specialized, non-granular, outer
synergids at the micropylar end of the

The nutrient solution of grape sugar, etc. yanriefactared tn the
passes downward through the phloem region to be

718 PHARMACEUTICAL BOTANY

EL’ATER.—An elastic, spiral filament attached to the spores of some Liverworts
and Horsetails and aiding in their dispersal when mature.

EMAR’GINATE.—Notched at the apex.

Em’sryo.—A rudimentary plant found within the seed.

Emsryo.’/ocy.—The study of the embryo and its development.

Em’sryo-sAc.—A large cell within the nucellus of the ovule in which the embryo
is formed after fertilization.

En’pocArp.—The inner layer of the pericarp. The inner epidermis of the fruit
wall.

_ Enpo’permis.—A layer of cells forming the innermost boundary of the cortex and
surrounding the central cylinder. Also applied to the bundle sheath surround-
ing concentric bundles in the stems and leaves of pteridophytes and some
monocotyledons.

En’pocen.—A Monocotyledon.

Enpoc’enous.—Applied to the axes of Monocotyl plants that do not increase
materially in diameter.

En’popuyTE.—A plant which grows within the tissues of another.

En’pospeRM.—A storage tissue of the seed, consisting of the matured embryo sac
around the embryo, and formed from the endosperm nucleus.

En’posporE.—The inner wall of a spore.

EnpoTHE’ciumM.—A zone of one or more layers within the exothecium of an
anther.

En’sIFORM.—Sword-shaped.

EntTomMopH ILous.—Insect pollinated.

ENn’TOPHYTE.—See Endophyte.

En’zymes.—Nitrogenous, colloidal bodies closely related to the proteins and con-
cerned with the chemical changes occurring in all living cells.

EpHEM’ERAL.— Lasting for a brief period (a day or so).

EpicA’Lyx.—A whorl of bracts resembling the calyx but below it.

Epr’cArp.—The outer layer of the pericarp or the outer epidermis of fruits.

Epicot’yL.—The portion of the embryo-axis above the cotyledon or cotyledons.

Epwer’mis.—The primary outer covering layer of cells of plants, sometimes later
replaced by cork. :

Epic’ ynous.—Applied to floral leaves that appear to be inserted upon the ovary.

EpipetT’ALous.— Upon the corolla.

Ep’IpHyTe.—An air plant. A plant growing on another plant but not necessarily
nourished by it.

EpiruHe’L1um.—A delicate layer of cells lining an internal cavity.

Eg’urraAnt.—Applied to leaves, as in Iris, when they all spring from a rhizome and
are successively folded on each other toward their bases.

EryTH’ROPHYLL.—The red coloring matter of leaves.

EstivaA’t1ion (Aestivation).—The arrangements of the floral organs in the flower
bud.

Erz#’r10.—An aggregate fruit like the Raspberry or Blackberry, the product of a
single flower, consisting of an aggregation of drupelets on a receptacle.

E’r10LAT1IoN.—The bleaching of green parts of plants when kept in the dark for
some time.

Evorvu’tion.—The presumable theory that all forms of living things existing today
have been derived from others previously existing, either by direct descent
or by common ancestry.

GLOSSARY 719

ExALBu’Minous.—Applied to a seed in which the nourishment is stored in the
embryo during the growth of seed from the ovule stage.

Excen’tric.—See Eccentric.

Excres’cENCE.—A morbid outgrowth.

Excre’T10n.—Getting rid of nitrogenous waste.

Excur’RENT.—Applied to trees, the main stems of which do not disappear in
branches but grow erect to the summit, ending in a terminal bud. The
opposite of Deliquescent.

Exro’LiATe.—To shed layers of bark. To cast off layers of tissue.

Ex’1nE.—The outer wall of a pollen grain.

Ex’ocArp.—The outer layer of the pericarp.

Exoc’rnous.—Applied to the axes of Gymnosperms and Dicotyledons which
increase materially in diameter.

Ex’ocens.—Plants with exogenous axes.

Exospor’tum.—The outer wall of a spore.

ExsERT’ED.—Applied to stamens that protrude from the throat of the corolla.

Ex’t1neE.—The outer wall of a pollen grain.

Exrrorse’.—Applied to anthers which face outward, away from the gynoecium.

Face.—The free surface of an organ.

FAL’cAte.—Scythe or sickle-shaped.

Fam’1ry.—A sub-division of an order.

Farina’ceous.—Starchy or mealy.

Fas’cicte.—A bundle or cluster.

Fascic’uLAR.—Belonging to a bundle.

Fascic’uLAR CAMBIuM.—See Cambium.

Fascic’uLATE.—Clustered.

Fats.—Glyceryl esters of fatty acids that tend to solidify and crystallize at ordinary
temperatures.

Fec’uLA.—The nutritive part of a cereal.

Fer’t1e.—Producing fruit or reproductive organs. Applied to flowers which
contain functionally active stamens and carpels.

FEeRTILIZA’TION.—That method of reproduction characterized by the union of two
dissimilar gametes.

Fi’ser.—A cell that is much elongated, thick-walled, usually lignified, tapering
more or less to a point at either end and primarily functioning mechanically.

Fy’srous.—Fiber-like. Referring to root systems composed of many slender
rootlets.

Frsrovas’CULAR Bun’pLE.—A stringy group of fibers, vessels or tracheids and cells
coursing through the various organs of the higher plants and serving for sup-
port and conduction of sap.

Fi.’AMENT.—The stalk of a stamen; a thread-like structure.

Fr_AMEn’Tous.—I hread-like. .

Fr’1rorm.—Lhread-like.

Frv’sRIATED.—Fringed.

Fis‘ston.—A form of division in which the cell separates into two equal or nearly
equal parts.

Fis’TULAR.—Hollow and cylindrical. oe

Fixep Ors.—Glyceryl esters of fatty acids that tend to remain liquid at ordinary
temperatures. —

720 PHARMACEUTICAL BOTANY

FLAGEL’LuM.—A _ whip-like protoplasmic outgrowth of certain organisms, as
Flagellates or of zodspores, serving as an organ of locomotion.

FowtA/cEous.—Leaf-like.

Fou’LicLte.—A one-chambered, dry fruit that dehisces along one suture only.

Fovit’/LA.—The contents of a pollen grain.

Frac’rureE.—The manner in which a root or other plant part breaks when sub-
jected to sufficient pressure.

Fronp.—The leaf of a fern. The leaf-like, branching thallus of marine algz.

Fruir.—A matured pistil, or ovarian portion thereof together with any closely
adhering part.

Fru’tT1cose.—Shrubby.

Fuco’xAn’THIN.—The brown pigment of the brown alga (synonymous with
phycophzin).

FucA‘cious.—Falling off early.

FuNDAMEN’TAL Tis’suE.—Ground-tissue. The tissue of plants through which
the fibrovascular bundles course. 5

Funic’uLus.—The stalk of an ovule.

Fur’cate.—Forked.

Fu’strorm.—Enlarged in the middle and tapering toward either end.

Gau'suLus.—A berry-like cone, as in Juniperus, formed by the coalescence of fleshy
scales and carpels.

GaA’/LEATE.—Helmet shaped.

GAMETAN’GIuM.—The cell or organ lodging the Eat

Gam’ETE.—A sexual cell.

GAm’ETOPHYTE.—The sexual generation.

GAMoPET’ALOus.—Applied to a flower whose corolla is composed of petals which
are more or less united at their edges.

GAmosEP’ALOus.—Having the sepals more or less united at their margins.

Gem’MA.—An asexual bud-like structure found in the cupules of Liverworts.

GremMA’TI0ON.—The process of budding as seen in the yeasts.

GeEne.—The unit of inheritance which is found in the chromosomes of the germ cells.

Gen’ERA.—Plural of genus.

GeEnIc’ULATE.—Kneed.

Geot’ropisM.—Response to the stimulus of gravity.

GeERMINA’TION.—The sprouting of a spore or seed.

Gero CELL.—A reproductive cell as distinguished from a somatic or Peay: cell.

Giiis.—The spore-bearing plates of a peeels

Gia’srous.—Smooth.

GiLanp.—A cell or group of cells whose principal function is secretion.

GLAND’ULAR.—Pertaining to or bearing glands.

Gians.—A nut.

G.avu’cous.—Covered with a bloom.

GLo’sows.—Small spherical se ac of calcitum-magnesium-phosphate found in
aleurone grains.

Gtos’uLAR.—Spherical.

GLom’ERULE.—A head-like cyme. A determinate head.

Guvu’coswwE (Gly’coside).—A cell constituent which under the influence of an
enzyme or of heat, dilute acids or alkalies splits up into glucose and some
other substances.

GLOSSARY 721

G.iume.—A floral bract of the grasses and sedges.

Guvu’TEN.—The proteid matter of cereals.

Gonip’tum.—Applied to the algal cell in lichens as well as to many forms of asexual
reproductive bodies in flowerless plants.

Gon’opHOoRE.—An upgrowth of the receptacle between the corolla and stamens,
as in Passiflora.

Guarp CE.is.—The pair of special epidermal cells which surround the pore of the
stoma.

Gyn’czci'uM.—The female sexual system of a flower.

Gyn’oPpHORE.—An upgrowth of the receptacle between gyncecium and androecium,
as in Geum. 3

Gynoste’m1um.—The united stamens and style. The column of orchids.

Has’1rat.—The original home of a plant.

Ha’/prRomME.—The conducting tissue of the xylem, comprising trachez, tracheids and
wood parenchyma.

Has’tATE.—Shaped like the head of a halberd, the basal lobes diverging.

HeaAp.—An indeterminate form of inflorescence, as seen in the Daisy family, in
which the flowers are in a dense cluster on the receptacle.

HEART’ woop.—The inner portion of the wood, often containing resinous and color-
ing matters and which no longer functions in the vegetative processes of the
plant.

He iorT’ropisM.—Response to the stimulus of sunlight.

Hersa’rium.—A classified collection of dried plant specimens.

Herep'1ry.—The faithful transmission of characters from parent or parents to
offspring. :
HeERMAPH’RODITE.—Applied to flowers which contain both sets of essential organs,

not necessarily functionally active.

Hesperiw’1um.—A large, thick-skinned, succulent fruit like the orange, lemon or
grape-fruit.

Her’erocyst.—A cell of different size and content from other cells in the filaments
of Nostoc and some other blue-green algz.

HeETEROPHYL’Lous.—Having more than one kind of foliage-leaves on the same
plant.

HeETEROs’PoROus.—Producing asexual spores of more than one kind, as in Selagi-
nella and the rusts.

HeterorypP’ic.—Applied to the first cell division in meiosis which involves the
reduction of the chromosomes from the diploid to the haploid member.

HeEtTERozy’GOTE.—An organism to which its two parents have contributed unlike
genes with respect to any given allelomorphic pair governing contrasted charac-
ters, and which in turn produces two kinds of germ cells with respect to the
characters.

Hex.—A prefix of Greek origin meaning six.

Hexac’ynous.—Having six carpels or styles.

Hexam’Eerous.—Having the parts of the flower in 6’s.

HeExAN’pRovus.—Having six stamens.

Hiserna’TIoN.—Passing the winter in a dormant state of existence.

Hi'Lum.—The scar of a seed, after the stalk of the ovule has fallen off. Also applied
to the point of origin or growth of a starch grain.

722 PHARMACEUTICAL BOTANY

Hip.—The fruit of a Rose, consisting of a number of akenes surrounded by a ripened
concave receptacle.

Hirsute’.—Covered with numerous long coarse hairs.

His'prp.—Beset with erect stiff hairs, as Borage.

Hisro.’ocy.—The study of tissues with the aid of the microscope.

Ho.prast.—The base of the stipe or of the plant body of a thallophyte which is
differentiated for attaching the plant to its substratum.

Homo ’ocous.—Having the same structural nature,

Homos’Pporous.—Producing asexual spores of only one kind.

Homozy’cote.—An organism whose parents contributed to it a similar member of
any given pair of genes and whose germ cells are all alike with respect to the
genes for that character.

Hormocon’1um.—A segment of a filament of a blue-green alga which may detach
itself from the rest of the filament becoming a new organism.

Hy’srip.—A cross between two varieties or species, rarely between two genera
of the same family.

Hypatn’oprs (Water Pores).—Organs for the exudation of water from plants under
conditions of low transpiration and abundant soil moisture.

Hypropu'iLous.—Applied to flowers that are pollinated through the agency of
water currents.

Hy’propHyTe.—A water-plant.

Hyprot’ropisM.—The response of a plant organ to the stimulus of moisture.

Hycroscop’1c.—The property possessed by certain cells or substances of absorbing
moisture with avidity.

Hyme’nrum.—A spore-bearing membrane of a fungus.

Hy’pua.—A filament of the mycelium of a fungus. _

Hypo.—A prefix of Greek origin meaning under.

Hy’pocotyt.—That part of an embryo plantlet below the cotyledon or cotyledons.

HypocraTER'IrorM.—Applied to a calyx or corolla when the tube is long and
slender and abruptly expands into a flat limb.

Hypoper’mis.—That portion of a plant organ directly beneath the epidermis.

Hypoce’ous.—Beneath the surface of the soil.

Hypotue’crum.—That portion of a thallus of a lichen directly beneath or around
the apothecium.

Hypoc’ynous.—Applied to the insertion of various floral parts on the receptacle
and beneath the pistil.

Ip’iosLast.—A cell which differs materially in form, size, character of cell wall,
or contents from its neighbors in a tissue, frequently applied to colorless stone
cells.

Imsrsr’T10n.—The taking in of water by organic bodies in such a manner as to
cause them to swell up.

Im’sRIcATE.—Overlapping like shingles.

ImMERsED’.—Growing entirely under water. -

IMPARIPIN’NATE.—Applied to a pinnately-compound leaf terminating with a single
leaflet.

InDEF’INITE.—Applied to stamens and other organs of the flower, when too numer-
ous to be conveniently counted.

InDEHIs’cENT.—Not splitting open in a definite manner when ripe.

InpIG’ENOus.—Native.

GLOSSARY — : 793

Inpu’stuM.—An outgrowth of the lower epidermis of many ferns that covers the
cluster of sporangia.

INEQUILAT’ERAL.—Having unequal sides.

InFrer’10R.—Applied to an ovary when the other floral parts are inserted above it.

INFLORESs’CENCE.—The arrangement of the flowers on the stem of a plant.

INFUNDIB’ULIFORM.—Funnel shaped.

InNATE’.—Applied to anthers that are attached by their base to the summit of the
filament.

INTEG’UMENT.—A covering.

INTERCEL’LULAR.—Between the cells.

INTERFASCIC’ULAR Cam’pruM.—Applied to a cambium layer which extends from
one fibrovascular bundle to another in the stems of Dicotyledons and
Gymnosperms.

INTER’NAL GLANDs.—More or less globular or elliptical shaped secretion reservoirs.

INTER’NAL PHLoem (Intraxylary Phloem).—A strand of phloem on the inner face
of the xylem, as found in bicollateral bundles.

In’TERNODE.—That portion of the stem between two nodes.

INTERRUP’TEDLY-PIN’NATE.—Applied to a pinnate leaf that has either smaller or
larger leaflets between those of usual size.

In’TINE.—The inner coat of the pollen grain.

In’tRA.—A prefix meaning within.

INTRAPET’IOLAR.—Applied to stipules that are between the petiole and the stem;
also to buds that are beneath or inside of the base of the petiole.

IntRorsE’.—Applied to anthers that face toward the gynoecium.

InTUssusceP’TION.—The formation of additional particles of protoplasm between
those already present.

In’uLin.—A carbohydrate isomeric with starch found in the cell sap of the Compos-
ite, Lobeliacez and some other families, from which it crystallizes in
lumps (in air-dried material) or in sphaerocrystals (in alcohol-preserved

material).
In’voLucreE.—A whorl (or whorls) of bracts subtending a flower or flower cluster.

Invot’uceL.—A secondary involucre.

In’voLute.—Applied to the arrangement of leaves within a bud when they are
rolled inward from both sides. :

IrrirAsit.1ry.—That property of living matter whereby it responds to a stimulus.

Iso’prAMET’RIC.—Applied to cell having nearly the same length, breadth and thick-

ness.
Isoc’amy.—The union of sexual cells of similar form.
Isom’EROUs.—Having the same number of parts in each whorl.
IsostEM’oNous.—Having the stamens and petals each in one whorl and of the same

number.
IsrH’mus.—Applied to the constricted portion between the two half cells i in certain

desmids.

KARYOKINE’sIs.—Indirect nuclear division.

KaTAs’ouisM.—Destructive metabolism.

ac sAgiellid to a longitudinal ridge or elevation of cortical tissue of Senega
root which extends from the crown downward. Also applied to the two

inferior petals of a papilionaceous corolla which are more or less united into

a body resembling the keel of a boat.

724 PHARMACEUTICAL BOTANY

Kner.—A form of knot which projects upward into the air from the roots of certain
trees that grow in wet soil, notably the bald cypress.

LaseL’LuM.—The large, lip-like, lower petal in the flower of an orchid.

La’BIATE.—Two lipped.

La’srum.—The lower lip of a labiate flower.

Lacin‘1ATE.—Applied to the margins of leaves which are deeply cut into irregular
narrow lobes.

LameEt’L#.—Little plates or layers. Applied to the layers of carbohydrate material
in a starch grain which surround the growing point; also to the gills of
a toad-stool as well as to the more or less regular markings found in cell walls
formed by delicate layers of material of different water content or of different
composition. :

Lam’tnA.—The blade or expanded part of any leaf.

La’nATE.—Covered with long, curled, wool-like hairs.

LAN’cEOLATE.—Lance-shaped.

La’tex.—The milk juice of a plant.

Laticir’ERous.—Applied to the latex carrying tissue of a plant.

LAtiFo’LIATE.—Possessing broad leaves.

Lrear.—An expansion of the stem or branch in whose axil one or more branches
arise.

Lear’Let.—A division of a compound leaf.

Lear-TRAce.—A vascular bundle while on its way from the stem bundle to the leaf.

Lec’ume.—A dry, simple, capsular fruit formed of a single carpel and dehiscent by
both ventral and dorsal sutures.

Len’TIcELs.—Fissures in the cork or epidermis of stems formed by the swelling up
and rupture of subjacent cells which were cut off by the phellogen in places
where cork was not laid down.

LenTIc’ULAR.—Having the shape of a double convex lens.

Lrep’trome.—The conducting tissue of the phloem, comprising sieve tubes, phloem
parenchyma and sometimes companion cells.

Lev’cop.ast.—A colorless plastid found in the cells of plants not exposed to light.

Li’ANE.—A woody climber or twiner of tropical forests.

Li’ser.—The inner bark or phloem region of Gymnosperms and Dicotyledons.

Li’srirorM-Cetis.—Those cells of the xylem that are thick-walled and resemble
bast fibers.

Lic’NEOus.—Woody.

Lic’niF1ep.—Covered with deposits of lignin. :

Lic’n1n.—A mixture of substances that adheres to the cellulose walls of certain cells

’ and which is characterized by taking on a reddish coloration with phloroglucin
and hydrochloric acid reagent.

Lic’uLATE.—Strap-shaped.

Lic’utr.—A membranous appendage at the summit of the leaf-sheath in many
grasses and cereals; a strap shaped corolla of a Composite.

LicuLirLo’Rous.—Applied to Composite flower heads, as those of Dandelion and
Chicory, which contain ligulate florets only, ;

Lims.—The spreading portion of a gamosepalous calyx or a gamopetalous corolla.

Line.—One-twelfth of an inch.

Lin’EAR.—Many times longer than broad and with nearly parallel margins. .

GLOSSARY | 725

Losr.—A division of a leaf or other flattened organ which is larger than a tooth but
which is not a leaflet.

Loc’uLAr.—Having a cavity or cavities.

Locutici’pAL.—Applied to the deshiscence of a capsule when it splits open along
the dorsal suture.

Loc’u.us.—A cell or cavity of an anther, ovary, or fruit.

Lo’ment.—A modified, jointed or multilocular legume that breaks open wane:
versely into segments when mature.

Lvu’ciw.— Clear.

Lu’men.—The cell cavity. The open space bounded by the walls of a ua,

Lu’ntrorM.—Half-moon or crescent shaped.

Lu’R1w.—Dingy-brown.

Lutes’cenT.—Yellowish.

Ly’rate.—Applied to a pinnatifid leaf, as that of the Turnip, in which the terminal
lobe is the largest and the rest decreasing in size toward the base.

Lysic’gNous.—Applied to the formation of a type of intercellular-air-space which
originates through the breaking down of cell walls common to a group of cells;
also applied to secretion reservoirs which are formed by the complete breaking
down of contiguous secretory cells which leave their products in the cavity
formed.

Macro.—A prefix of Greek origin meaning large.

MaAcrosporan’GiumM.—A _ spore ‘case containing one or more macrospores. (The
nucellus in Spermatophytes. )

Mac’rospores.—The larger of the two different kinds of spores produced by some
of the higher Pteridophytes and the Spermatophytes. (The embryo-sac in
Spermatophytes.)

Macrospo’ropHy_i..—The leaf bearing the macrosporangium. (The carpel in
Spermatophytes.)

Mac’uLATE.—Spotted.

Ma/MILLATE.—Bearing teat-like protuberances.

Marces’cENT.—Withering but not falling, dropping off.

Marine’.—Applied to plants which grow in the sea or ocean.

Meput’LA.—Pith.

Mep’uLLARY.—Pertaining to the pith.

Mep’utLAry Rays.—Vertical, radial plates of parenchyma connecting the cortex
with the pith or a portion of the xylem with a portion of the phloem,

Mep’uLLARY SHEATH.—The inner ring of xylem abutting on the pith.

Mecaso’rus.—The ovule.

MeGAsPoRAN’GIUM.—See macrosporangium,

Mec’ASPORE.—See macrospore.
MEGASPO’ROPHYLL.—The carpel of seed plants. A leaf bearing one or more

megasporangia.
Maero’sts.—The reduction divisions during the maturation of the gametes, in which

the members of each chromosome pair separate, and the chromosome number

in the resulting daughter cells is reduced to half the somatic number. erie
Mem’sRANE Crys’TALS.—Crystals enclosed within a membrane developed within

the cell. ;
Mem’pRANOUs.—Thin, soft and flexile.

726 PHARMACEUTICAL BOTANY

MEeEr’1cArP.—One of the two inferior akenes which are found with the carpophore
making up the cremocarp in Umbellifere.

MER’IsTEM.—Formative or undifferentiated tissue consisting of cells which in the
living plant are in an active state of division.

MERISTEMAT’Ic.— Consisting of generative cells or meristem.

Mes’ocArp.—The middle region of the fruit wall or pericarp, lying between the
epicarp and endocarp.

Mes’opHyL_.—The region of leaf parenchyma within the epidermis or, in flat leaves,
between the two epidermises.

Mes’tome.—The conducting portion of a fibrovascular bundle.

Metap’o.isM.—The sum total of all the chemical changes which take place in a
living plant.

METAGEN’EsIS.—Alternation of generations. The production of sexual indi-
viduals by asexual means and asexual or neutral individuals by sexual means.

MEeETAmMoR’PHOsIs.—A change in the form or function of an organ or organism.

METAPHLO’EM.—Phloem formed by the cambium. Secondary phloem.

METAXy’LEM.—The secondary xylem, formed by the cambium on its inner face.

Micro.—A prefix of Greek origin meaning small.

Mr’crosE.—A minute vegetable or animal organism,

Mr’crocrys’TAts (Sphenoidal Crystals)—Very minute, arrowhead- or deltoid-
shaped crystals frequently packed in great numbers in a single cell.

Mr’cropyLe.—The opening between the coats of an ovule through which the pollen
tube enters. The orifice or foramen in the seed coat through which the hypo-
cotyl passes during germination.

Microso’mes.—Applied by Strasburger to minute particles in the protoplasm which
have a high degree of refringency.

Microso’rus.—A lobe of the anther.

MicrosPporANn’ciIuM.—A spore case containing microspores. An anther sac.

Mi’crosporE.—A small spore found in a microsporangium. The pollen grain of
a seed plant.

Microspo’RoPHYLL.—A leaf bearing microsporangia. The stamen of seed plants.

Mi'pie Lamev’tA.—A layer of calcium pectate or of some other base in com-

bination with pectic acid, appearing within the vacuole between two plasma
membranes during the last stage of cell division and appearing in suitably
mounted matured tissues as a dividing line between the cellulose walls of
adjacent protoplasts.

Miw’r1s.—The large main central vein of a pinnately-veined leaf which is con-
tinuous with the leaf stalk.

Mrro’sis.—Indirect nuclear division.

MonapDEL’PHOUs.—Applied to stamens which are united by their filaments into one
set, as in the Malvacee.

Monan’prous.—Possessing only one stamen.

Monan’tHous.—Having only a single flower on the peduncle.

Moni’trorM.—Resembling a chain of beads.

Mono.—A prefix of Greek origin, meaning one or single.

Monocar’PELLARY.—Of one carpel.

Monocu.Amyp’£ous.—Possessing but one perianth whorl.

Monocuin’ic Prisms (Monoclinic Crystals) .—Crystals possessing three axes of
unequal length, two of which are obliquely inclined to each other and the
third forming right angles to these two.

GLOSSARY 727

Monoc’.inous.—Having both andreecium and gyncecium.

Monocory.ep’onous.—Having only one cotyledon or seed leaf.

Mone@’cious.—Having separate staminate and pistillate flowers on the same plant.

Monony’sriw.—The offspring of two parents differing in only one mendelian
characteristic.

Mownotoc’uLAR.—One chambered. :

Monom’erovus.—Applied to flowers having one part running through each whorl.

Monopo’pium.—A plant axis which elongates at the apex and sends off lateral
branches in acropetal sequence.

Mownos’t1cHous.—Arranged in one vertical row.

Mvu’cronaTE.—Terminating abruptly in a small soft point.

Mutt1.—A prefix of Latin origin meaning many.

MUuLTIcEL’LULAR.—Consisting of many cells.

Mutticip’1rAL.—Many-headed; applied to a rhizome or root from which numerous
stems arise.

MuttirA’R10us.—Composed of many diverse parts.

MuttiLoc’uLAr.—Many celled or chambered.

Mut’tieLe Fruir.—A fruit composed of many small fruits, each the product of
a separate flower, as in the Fig or Hop.

MoutA’tion.—A sudden variation that is later passed on through inheritance and
that results from changes in a gene or genes.

Myce’L1uM.—The vegetative body of a fungus consisting of intertangled hyphe.

Mycov’ocy.—That branch of Botany that treats of the Fungi.

Mycorrui’zA.—An association between the roots of certain plants and the myce-
lium of certain fungi which forms an investment about their tips.

Na’pirorM.—Turnip-shaped. Somewhat globular, becoming abruptly slender
and then terminating in a conical tap root.

NAT’URALIZED.—Applied to plants that have been introduced from another
country.

NAvic’ULAR.—Boat-shaped.

Nec’TAR.—A sweet secretion by the flower.

Nec’TtARyY.—The part of the flower which secretes nectar.

NEIGH’BORING CrELLs (Subsidiary Cells)—The epidermal cells which abut upon
the guard cells of the stoma.

NervA’TION.—The arrangement of veins in a leaf.

NetTep.—See Reticulate.

Nevu’TRAL.—Said of flowers which possess neither stamens nor carpels. Also
applied to the asexual generation of plants.

Nrv’gous.—Snow-white.

Nope.—The place on the stem which normally shows outgrowths of a leaf, whorl
of leaves or leaf modifications.

Noposr’.—Having swollen joints or knobs.

Nop‘uLe.—A small rounded body as a root tubercle.

Nor’MAL.— Usual.

Non.—Not.

Nucet’Lus.—The body of an ovule.

Nucir’ERous.—Nut-bearing. ek

Nu’cieus.—A dense, definitely circumscribed region of protoplasm within the cell
containing linin, chromatin and nuclear sap.

728 : PHARMACEUTICAL BOTANY

Nucte’oLus.—A small body of dense protoplasm within the nucleus.

Nur.—A dry, indehiscent, 1-celled, 1-seeded fruit with a stony or leathery pericarp.

Nut’Let.—A small nut. The characteristic fruit of the Ladzate.

Nutri’'t1ion.—That branch of Physiology which includes the absorption, distribu-
tion and assimilation of food stuffs.

Osn.—A prefix of Latin origin signifying inversion.

Oxcon’IcAL.—Inversely cone-shaped.

Oscor’DATE.—Inversely heat-shaped.

OBLAN’CEOLATE.—Lance-shaped with the broadest part toward the summit.

Os.ate’.—Flattened at the ends or poles.

Op’LIGATE.—Necessary, indispensable.

Os.igue’.—Taking a position between erect and horizontal, as in the case of many
stems. More developed on one side than on the other, as in certain leaf
blades.

On’Lonc.—Longer than broad with nearly parallel sides.

Oxo’/vATE.—Ovate with the attachment at the narrower end.

Opsruse’.—Having a blunt or rounded end.

O’cureA (o’crea).—A sheathing stipule.

Ors.—See Fixed Oils; Volatile Oils.

OLrores’1ns.—Secretions composed of resins dissolved in volatile oil.

Ontoc’eny.—The history of the development of an individual.

O’ésporeE.—The fertilized egg.

Open COLLATERAL BunpDLE.—A collateral bundle in which a cambium zone lies
between the xylem and phloem strands.

Oper’cuLuM.—The transversely dehiscent lid or cover of a moss capsule.

Orsic’uLAR.—Circular.

Or’pER.—A division of a class containing one or more families.

Ortnor’rRopous.—Applied to ovules or seeds which are erect, with the micropyle at
the apex and the hilum coinciding with the chalaza.

O’vary.—The lower part of a pistil or carpel containing the ovules.

O’vate.—Shaped like a lengthwise section of a hen’s egg and having the attach-
ment at the broader end.

O’vu.Le.—A transformed bud destined to become a seed after fertilization.

O’vum.—The female sexual cell.

Pau’ATE.—A convex projection on the base of the lower lip of a personate corolla.

Pa’LeA (Pal'et)—An inner bract of a Grass inflorescence which with the lemma
incloses the flower.

PALEA’cEOus.—Chaffy.

PaL'isApE CrLis.—Radially-elongated cells with more or Jess indurated walls
found in certain seed coats.

PAu'IsADE PAREN’CHYMA (Palisade Tissue).—Elongated mesophyll cells lying
perpendicular to the surface of the leaf.

PAL’Lip.— Pale.

PAL’MATE.— Divided or lobed in radiate fashion.

PALMAT’Ir1D.—Palmately-cleft.

PAnpbvu’RIFoRM.—Fiddle-shaped.

PAN‘ICLE.—A compound raceme.

GLOSSARY 729

PapPILLionA’cEous.—Having butterfly-shaped flowers, as in the sub-family Papilio-
nacee of the Leguminosz.

PapiIL’LAE.—Small, short, nipple-shaped protuberances of the epidermal cells.

PAp’ILLosE.—Bearing small, nipple-shaped protuberances.

Pap’pus.—The calyx of a Composite flower.

PApyRA’cEOus.—Papery.

ParaApH’ysis.—A sterile filament found among reproductive organs in certain
plants.
ParAsitT’1c.—Growing upon or within and deriving sustenance from another living

organism.

PAREN’CHYMA (Fundamental Tissue).—The soft, filling-in tissue of plants consisting
of more or less isodiametric cells and usually with cellulose walls enclosing
living contents.

PARENCHY’MATOUS.—Referring to, consisting of or resembling parenchyma.

Pari/ETAL.—Situated on or pertaining to the wall of an ovary or pericarp.

Part’ep.—Incised nearly to the mid-rib or base.

PARTHENOGEN’ESIS.—The production of an embryo from an unfertilized egg.

Patruo.’ocy.—The study of diseases.

Prec’rose.—Carbohydrate cell wall material related to the hemi-celluloses and to
mucilage.

Pep’ATe.—Palmately parted or divided with two lateral lobes or divisions from each
of which more or less linear divisions arise.

Pep‘iceL.—A branch of an inflorescence axis supporting a single flower.

Pepun’cLe.—The main stalk of an inflorescence. The flower stalk.

Pe.iu’crp.—Transparent, clear.

PEL’TATE.—Shield-shaped and attached by its lower surface to the support.

Pen’puLous.—Hanging nearly vertically downward, as in the case of some ovules
that hang from the sides of a locule.

PenTAM’ERovus.—Applied to flowers having the number five or a multiple thereof
running throughout each whorl.

PENTAN’DROUs.—Having five stamens.

Pr’po.—A fruit of a Cucurbit; a gourd.

PEREN’NIAL.—Living more than two years.

Per’rect.—Applied to flowers that contain both stamens and carpels that are
functionally active.

PreRro’LIATE.—Applied to leaves which are united around the stem at their base.

Per’IANTH.—The floral envelopes, calyx and corolla or calyx alone when corolla
is absent.

PEr’IBLEM.—A region of meristem lying between the dermatogen and plerome in
the growing end of a root or stem. The meristem which gives rise to cortex.

PericAm’s1uM.—A zone of meristematic tissue lying just within the endodermis.

PrR’IcARP.—The wall of a ripened ovary or fruit surrounding the seed or seeds.

PericLA’pruM.—A sheathing petiole.
Per’icycLe.—A region lying between the endodermis and the conducting bundle

or bundles.

PER’IDERM.—All of the tissues produced by the phellogen, both externally and
internally, and including the phellogen itself.

PeRip’rum.—The outer covering of certain fungus fructifications as puff-balls.

PER’1GONE.—See perianth.

730 PHARMACEUTICAL BOTANY

PEeriG’yNous.—Applied to stamens and petals when they are adherent to the calyx
throat, and so borne around the gynoecium.

PER’IsPERM.—The nourishing tissue of some seeds outside of the embryo sac and
representing the nucellus of the ovule, which during maturation has become
laden with nutriment.

Per’IstroME.—The teeth around the mouth of the capsule in mosses.

PeriTHE’c1uM.—The receptacle containing asci in certain Ascomycetes.

Persist/ENT.—Applied to parts of the flower which remain until the fruit ripens or
to leaves which remain on the plant over winter.

PerR’sONATE.—Applied to a bilabiate corolla which has its throat closed by a convex
projection on the base of the lower lip.

Pet’AL.—One of the floral leaves of the corolla.

Pet’ALomp.—-Of some other color than green. Having the color of a petal.

Pet’1oLe.—A leaf stalk.

Per‘1oLuLe.—tThe stalk of a leaflet.

PuEL’LODERM.—Secondary cortex containing chloroplasts and formed by the cork
cambium or phellogen on its inner face.

Pue.’LocenN.—The meristem which gives rise to cork and frequently secondary
cortex; cork cambium,

Puio’em.—That part of a vascular bundle which contains sieve tissue and fre-
quently bast (phloem) fibers.

Putoroctu’cin.—A white crystalline substance having the formula of CsH»Os,
obtained by the decomposition of phloretin and from certain gummy extracts
and used with hydrochloric acid as a test for lignin.

Puycocy’Anin.—The blue pigment found in the Cyanophycez (Blue Green Algz).

PuycOERYTH’RIN.—The red pigment occurring in the Rhodophycee (Red Aigz)
and Cyanophyceze (Blue-green Algz).

PuycopH#'In.—The brown pigment found in the Phezophycez (Brown Algz).

PuycoxAn’THIN.—A yellowish pigment occurring in some Algz.

PuyLLoc.a’pe.—A flattened branch which resembles a leaf, as in Ruscus.

Puy.’Lope.—A dilated petiole which functions as a leaf blade.

Puy.’/LoTAxy.—The arrangement of leaves on stems.

PHYLLOXAN’THIN.—See xanthophyll.

Puy.Loc’Eeny.—The history of the race.

Puys’10LoGy.—The science which treats of the functions of living organisms.

Puyto’GLosu.ins (Crystalloids).—Crystal-like bodies of protein composition found
in aleurone grains.

Puy’ton.—A term given by Gaudichaud to an internode with a node at its upper
extremity which bears one or more leaves, in the axils of which buds may
appear. é

Pic’ments.—Coloring substances found in cells.

Pi’‘Leus.—The cap of a toadstool.

Puuur’eRous.—Bearing hairs.

Priose’.—Covered with long, straight and scattered hairs.

Pin’NATE.—Applied to compound leaves when the leaflets are ne ie the
mid-rib.

PINNAT’IFID.—Pinnately-cleft.

PINNATIPAR’TITE.—Pinnately-parted. \

PrnnaT’isect.—Pinnately-divided. ;

GLOSSARY 731

Prin’NULE.—A secondary pinna.

P1’strorM.—Pea-shaped,

Pis't1..—The central female organ of a flower consisting of one or more united
carpels.

Pis’r1LLATE.—Applied to flowers that possess one or more carpels but no fertile
stamens.

Prru (Medulla).—The core of soft parenchymatous tissue at the center of stems or
roots.

PLAcEN’TA.—The nourishing tissue which connects the ovules with the wall of
the ovary.

PLACENTA’TION.—The arrangement of the placenta within the ovary or the pericarp.

PLAsMA Mem’BRANE (Outer Plasma Membrane).—See Ectoplasm.

PLAsMo’D1uM.—A multinucleated, naked mass of protoplasm having amoeboid
movement. The vegetative body of a Slime Mold.

PLAsMOL’ysis.—A contraction of the protoplasm of a cell due to the extraction of
contained water under the arises of reagents of greater density than the
protoplasmic sap.

PLAs’tTrps.—Dense, porous protoplasmic bodies of various shapes scattered about in
the cytoplasm. :

PLe’rRomME.—A meristem found in the apical regions of plant axes which gives rise
to fibrovascular tissue.

Pui'cAte.—Folded like a fan.

Piumose’,—Feathery.

Piu’mMuLe.—The rudimentary bud between the cotyledons.

Piurioc’uLAR.—Having more than one chamber or cell.

Po’LAR Bopy.—A portion of a gamete budded off before fertilization.

Pot’LeEn.—The fertilizing dust composed of cells produced in the anthers of flowers.

Potten Gratin (Microspore).—A cell developed within the anther-sac and which
carries the male fertilizing element.

PotuinaA’t10on.—The transfer of pollen from anther to stigma and subsequent
germination thereon.

Poutin’tum.—A coherent mass of pollen grains in Orchids and Milkweeds, arranged
as to be carried by insects.

Po.y.—A prefix of Greek origin meaning many.

Potyape.’pHous.—Applied to stamens which are united by their filaments into
many sets.

PoLtyAn’pRous.—Having many stamens.

PoLyAn’THOUs.——Many flowered.

PoLyarcu’.—Said of a radial vascular bundle having many — and phloem
rays.

PoLycAR’PELLARY.—Composed of 3 or more carpels.

Potycar’pic.—Fruiting successively.

PoLycrPHAL’1c.—Bearing many heads.

Po.ycoryLep’on.—A plant such as a Conifer which possesses more than 2 cotyle-
dons or seed leaves.

PoLyEM’BRYONY.—Producing more than one embryo within a seed.

Po.ya’Amous.—Applied to species in which staminate, pistillate and hermaphrodite

flowers are borne on the same plant.
Po.yc’oNnAL.—Having several or many angles.

132 PHARMACEUTICAL BOTANY

PoLyMor’PHOous.—Having several to many different forms.
PoLypet’ALous.—Having distinct, disjoined petals.
PoLypH’yLLous.—Many-leaved.

Potysep’ALous.—Having distinct, disjoined sepals.

Po.ys’tAcHous.—Having many spikes.

PoLysTEM’oNOUs.— Possessing many more stamens than petals.

Pome.—A fleshy indehiscent fruit, two or more carpelled, with fibrous, cartilag-
inous or stony endocarp, the chief bulk of which consists of an adherent torus.

Pore CANALs.—Pores through much thickened, usually lignified walls.

Pores (Pits).—Small thin spots or small holes in the cell wall.

PREFLORA’TION.—See Aestivation.

PREFOLIA’TION.—See Vernation.

Prick’Le.—A sharp, rigid outgrowth from the epidermis and cortex.

Primor’pIAL.—First formed.

Primor’p1AL U’TrRicLe.—The outer plasma membrane. The outer layer of proto-
plasm adjacent to the cell wall. .

Procam’s1uM.—The first formed fibrovascular tissue of any organ before differ-
entiation has taken place into xylem and phloem,

Procum’sENT.—Lying flat on the ground.

Proem’sryo.—The primary stage in the development of Chara consisting of a
single filament and a long rhizoidal cell. The suspensor in flowering plants.

Promyce’L1uM.—A short hyphal growth from resting spores of smuts or rusts upon
which basidiospores are borne.

PRosEN’cCHYMA.—Tissue composed of elongated, taper-pointed cells.

Protan’pRous.—A condition of hermaphrodite flowers in which the stamens
mature before the carpels. ,

Protoc’ynous.—Applied to hermaphrodite flowers in which the carpels are mature
before the stamens.

ProTHAt’tus (Prothal’lium).—A thalloid body bearing antheridia and archegonia,
produced by the germination of a spore of a Pteridophyte into a protonema
which later undergoes differentiation.

Protone’MA.—A simple or branched green filament formed by the germination of
a spore of a moss or fern.

PRoToPHLO’EM.—The first-formed phloem elements in a fibrovascular bundle.

Pro’ropLAsM.—Living matter.

Pro’ropLast.—A term applied by Hanstein to the smallest body of protoplasm
capable of individual action, either with or without a cell-wall, and either
associated with other like units in a tissue or independent.

Protoxy’LeM.—The first formed elements of xylem in a fibrovascular bundle.

Prox’tmMAL.—Applied to the basal extremity. The attached end of an organ as
opposed to the free or distal end.

Pseupo.—A prefix of Greek origin indicating spurious or false.

Pseupo-BuLB.—The fleshy bulb-like internode of an epiphytic Orchid.

Pseu’pocarp.—A fruit which represents the product of the ripening of a single
ovary as well as one or more accessory parts.

. PsEUDOPAREN’CHYMA (Plectenchyma).—A tissue found in the ace: bodies of
many fungi and consisting of compact hyphae somewhat resembling paren-
chyma in cross section.

PuBER’ULENT.—Covered with a fine, soft, hairy coating.

GLOSSARY 733

Puses’CcENT.—Covered with soft, short hairs.

Putvi’nus.—An enlargement at the base of the petiole or petiolule of some leaves
or leaflets, as in numerous Leguminosae.

Pune’TATE.—Dotted with small spots or minute pits.

Pus’ruLAR.—Applied to surfaces having blister- or pimple-like elevations.

PutA’MEN.—The stony endocarp of a drupe.

Pyre’Norps.—Small, rounded, colorless, refractile granules embedded in the
chromatophores of numerous Algz and thought to be starch forming centers.

Pyx’1s.—A capsule which dehisces transversely into pot and lid portions.

Quap- or QuApriI.—A prefix of Latin origin signifying four.

QuADRAN’GULAR.—Four-angled,

QuaADRIFO’LIATE.—Applied to palmate leaves which have four leaflets arising from
the summit of the petiole.

QuINQUEFOL’IATE.—Applied to any compound leaf that has five leaflets.

RaAceme’.—An indeterminate inflorescence having pedicelled flowers arranged
along a lengthened axis.

RAc’emMose.—Arranged in racemes.

* Ra’cuis.—The extended portion of a peduncle.

Rap’1cAL.—Arising from the root or base of the stem.

Rap‘icLe.—The rudimentary root of an embryo plantlet.

RaA’mMAL.—Pertaining to a branch.

Ra’mus.—A branch.

Ramose’.—Branching.

Ranx.—A row of leaves or other organs arranged vertically on a stem.

Ra’pue (RHA’PHE).—The adherent portion of the ovule stalk in inverted and half
inverted ovules and seeds.

Rapu’iwes.—Bundles of needle-shaped crystals.

Recrp’rAcLE.—The shortened stem upon which the whorls of floral leaves are
inserted.

RecepTac’ULAR.—Pertaining to the receptacle.

Rec’LINATE.—Bent downward,

Rec in‘Inc.—See Reclinate.

ReEcURVED’.—Curved outward or backward to a moderate extent.

REFLEXED’,—Turned outward or backward more abruptly than Recurved.

Rec’mMA.—A capsular fruit of 2 or more carpels that first splits into separate parts
and then each of these dehisces.

REJUVENES’CENCE.—Applied to a mode of reproduction in which the protoplasm
of the cell becomes rounded out, escapes by rupture of the cell wall, forms
cilia and moves about, in time developing into a new plant.

REN’1rorM.—Kidney-shaped.

Repanp’.—Having a slightly undulating margin.

Re’pent.—Creeping. ‘

Re’pLuM.—A spurious membranous septum seen in Cruciferous fruits that per-
sists after the valves have fallen away.

Revic/uLATE.—Applied to markings or veins which are in the form of a network.

Retusr’.—Having a broad, shallow sinus at the apex.
Rev’o.ure.—Said of leaves in the bud when their margins are rolled backward.

734 PHARMACEUTICAL BOTANY

RuA’pHE.—See Raphe.

Ru1’zoms.—Absorption organs of certain plants that are analogous with roots of
higher plants but which consist of outgrowths of epidermal cells.

Ruizome’.—A creeping underground stem.

Ru’zomorpHs.—Root-like structures composed of united hyphz and seen in
certain fungi.

Ris.—A prominent vein or ridge.

Rin’Gent.—Applied to the corolla of a bilabiate type whose throat is pent and
lips separated.

RipA’R10us.—Growing along the banks of rivers or other water-courses,

Roserte’.—A cluster of leaves or other organs.

Ros’TRATE.—Beaked.

RosTEL’LuM.—A process found on the column of orchid flowers which supports the
disks of the pollinia.

Ro’rATE.—Wheel-shaped.

Rotunp’.—Rounded in outline.

Ru’Fous.—Brownish-red.

Rucosr’.—Wrinkled.

Ru’MINATE.—Applied to the albumen of certain seeds when the perisperm is found
coursing through the endosperm in irregular fashion.

Run’cinATE.—Applied to a pinnately-cleft leaf whose lobes are directed backward
as in the Dandelion.

Run’NER.—A stem or branch which roots at intervals as it trails along the ground.

Sac’cATe.—Pouch-like.

Sac’IrTATE.—Arrow-shaped.

SAMA’RA.—A winged fruit.

SAP’/ROPHYTE.—An organism that lives upon decaying or dead organic matter.

Sar’cocarp.—The fleshy portion of a drupe or other fruit.

Sca’BRous.—Said of leaves, etc., that are rough or harsh to the touch.

ScALAR’IFORM.—Applied to trachez or tracheids whose walls show transversely
arranged bars, resembling the rongs of a ladder.

ScaAn’DENT.—Climbing.

Scape.—A naked peduncle arising from a root or underground stem.

Sca’R1ous.—Dry and membranous.

Scuiz’ocarp.—A fruit that separates when mature into 2 or more indehiscent
parts.

Scuizoc’eNnous.—Said of intercellular-air-spaces or of reservoirs that are formed
by the breaking down of the middle lamella of cells where several come to-
gether and the later separation of the cells at these places.

Scr’on.—A shoot intended for grafting.

ScLEREN’CHYMA.—A tissue composed of stone cells, fibers or both.

ScLERO’T1uM.—A hardened mass of mycelium, -

Scor’pior.—Applied to certain cymes whose flowers are situated on alternate
sides of the floral axis.

ScuTEL’LuM.—A shied-shaped portion of the conten of Graminea, which absorbs
nourishment from the endosperm during germination and bales it out to the
rest of the embryo. :

Sec’unpinE.—The outer coat of the ovule.

Seep.—A fertilized and matured ovule containing an embryo.

GLOSSARY 735

Se’pAL.—A leaf of the calyx.

Sep’TATE.—Possessing one or more partitions.

Septici’DAL.—A mode of dehiscence in which the opening occurs along the line of
junction of the carpels.

SEPTIFRA’GAL.—A method of dehiscence in which the valves of a capsular fruit
break away from the partitions or septa.

Sep’ruM.—A partition between cavities in an ovary or fruit or between cells in a

tissue,

Seric’Eous.—Silky. Having a covering of fine, soft, appressed, silky, hairs.

Ser’RATE.—Toothed with teeth projecting toward the apex.

SER/RULATE.—Finely serrate.

Ses’stLE.—Without a stalk.

Se’tA.—A bristle-like structure.

Setic’ERous.—Bristle bearing.

Sit/1cuLe.—A short silique.

Sm/1qguE.—The characteristic fruit of the Crucifere, consisting of a capsule of 2
valves which separate from the replum in dehiscence.

Sin’UATE.—Wavy margined.

Sorr BAst.—The unlignified portion of the phloem.

Somat’ic Cetts.—The body cells of an individual, in distinction from reproductive |
cells.

Sore’pium.—A scale-like structure found on many lichens and consisting of a
group of alge cells surrounded by a network of hyphae. When detached
from the parent-plant it has the power of developing vegetatively into a
mature lichen.

Soro’sts.—A multiple fruit, as represented by the Mulberry and Osage Orange,

’ consisting of a swollen up, condensed and mature spike.

So’rus.— An aggregation of sporangia.

Spa’p1x.—A fleshy spike more or less surrounded by a bract called a spathe.

SpatHE.—A large bract that encloses or subtends an inflorescence.

SPAT’ULATE.—Said of flat leaves that are narrow at the base and become gradually
broader toward the summit, which is rounded.

SpeRMA’TopHYTE.—A seed plant.

SPERMATOzO'ID.—A male sexual cell. See Antherozoid.

SpERMATOzO’oN.—Another name for Spermatozoid or Antherozoid.

SPER’MODERM.—The covering of the seed.

SpHAcr’LIA.—The conidia stage of Claviceps.

Spic’ATE.—Arranged in a spike.

Spic/ULE.—A small pointed outgrowth. A needle-shaped crystal.

Sprxe.—An indeterminate inflorescence consisting of sessile florets arranged along
a lengthened axis.

Sprke’LET.—A secondary spike.

Spine.—A sharp, rigid termination of a branch as in the Honey Locust. A thorn.

SpINESs’CENT.—Spiny in structure.

SporAp’ic.—Scattered.

SporAn’GIOPHORE.—The stalk or support of a sporangium.

SporAn’ciuM.—A spore case. :
Sporr.—An asexual or sexual reproductive cell usually with a highly resistant

cell wall.

Sporoco’NIuM.—The asexual generation in Bryophytes and Pteridophytes.

736 PHARMACEUTICAL BOTANY

Spo’RoPHYLL.—A spore bearing leaf.

Spur.—A tubular or saccate appendage of some part of the flower, usually con-
taining nectar.

Soquamose’.—Scale-like.

STA’MEN.—A male organ of the flower producing pollen.

Sram’inopE.—An abortive and sterile stamen, or any body without an anther
occupying the normal place of a stamen.

Srrc’ma (pl. stegma’ta).—A cell containing a warty silicious body adherent to the
surface of a sclerenchyma fiber, and occurring in various palms, orchids, the
Scitaminea, etc.

Str’LE.—All the structures within the usual type of endodermis.

STEL’LATE.—Star-shaped.

SteM.—The ascending axis of a plant bearing leaves or leaf modifications.

Sre’REOME.—The supporting elements of a fibrovascular bundle.

Srer’1Le.—1. Unproductive, as a stamen without anther, flower without pistil,
or pericarp without seeds. 2. Devoid of living organisms.

STERILIZA’TION.—The process of riding an object of all living organisms.

Stic’/mA.—That part of a pistil or carpel which receives the pollen.

Stipe.—The stem of a moss; the stalk of a fern frond; the stalk of a toadstool or
other fungus.

Srip’ULATE.— Possessing stipules.

Srip’ute.—A modified leaf, usually blade-like and situated at the base of the leaf-
stalk. :

Sro’Lon.—A slender running branch above or below the surface of the soil, either
capable of taking root or bearing a bulb at its end.

STOLONIF’EROUs.—Bearing stolons.

Sro’/MA (Stomate).—A minute opening for the passage of gases and water vapor m
the epidermis, surrounded by two guard cells, which usually have the power to
enlarge and reduce the opening. Also employed for the opening in the epi-
dermis with the guard and accessory cells.

Srom’ATA.—Plural for stoma.

SromAt’AL ApPARA’TUS.—The stoma with its surrounding guard cells and neigh-
boring cells.

Sromat’AL CHAmM’BER.—The intercellular-air-space directly beneath the stoma.

SrRAND.—Applied to any tissue or group of tissues forming a narrow, more or less
cylindrical mass extending longitudinally through the plant or plant organ.

Strri’Ate.—Marked with fine longitudinal lines or grooves.

Srricose’.—Covered with sharp and rigid appressed hairs.

Srros’1Le.—A scaly multiple fruit consisting of a scale-bearing axis, each scale of
which encloses one or more seeds. A cone.

Stone Cexis (Sclerenchyma Cells, Sclereids).—Isodiametric or moderately elon-
gated parenchyma cells modified by thickenings, lignification of the cell
wall and loss of protoplasmic contents.

Stry_e.—That portion of a pistil connecting the ovary with the stigma.

StyLopo’ptum.—The fleshy disk directly above the ovarian portion of an Um-
belliferous fruit, formed by the expansion of the bases of the two styles.

Sus.—A prefix of Latin origin meaning under, below, subordinate, nearly or
partially.

Su’ser.—Cork tissue,

GLOSSARY 137:

SupeRiN.—A fatty cell wall constituent serving to water-proof the wall.
Susert’zED (Corky).—Applied to a cell wall infiltrated with suberin.
SuBTERRA’NEAN.—Beneath the surface of the soil.
Su’BULATE.—Narrow and tapering to an acute end.

Suc’cULENT.—Soft and juicy or fleshy.

Suc’KER.—A shoot from the root or lower part of the stem or underground stem.

SurrRru’tTICosE.—Applied to stems or plants that are woody at their base and
herbaceous above.

SuL’cATE.—Having longitudinal grooves.

Supe’RIoR.—Said of an ovary that is not adherent to and above the calyx; also
applied to a calyx which is situated on the upgrown receptacle above the
ovary or to a tubular calyx whose limb appears to spring from the top of the
ovary.

SuspEN’sor.—A row of cells, representing the first development of the fertilized
egg of a seed plant, upon the end of which an embryo is formed.

Su’rurE.—The line of union of two carpels. The line of dehiscence.

Swarm Spore.—A spore which possesses one or more cilia for movement.

Sycon’1uM.—The characteristic multiple fruit of the Fig, which consists of a fleshy,
invaginated receptacle bearing numerous akenes.

Syaro’sts.—The living together of two individuals having a communion of life
interests.

SYMMET’RICAL.—Said of flowers when the parts of each whorl are of the same num-
ber or multiples of the same number.

SympET’ALous.—See Gamopetalous.

Sym’puysis.—A union of parts.

SynApsis.—The union in pairs of homologous chromosomes, respectively of
maternal and paternal origin, to form bivalents.

SyncAr’pous.—Said of fruits and gyncecia when they are formed of two or more
united carpels. —

Syner‘cips.—Two nuclei in

nucleus.
SyNGENE’sIous.—Said of stamens when their anthers are united.

Syn’onym.—Another name for the same thing.
SynpHyLL’ous.—Applied to sepals or petals which are united by their margins.

SynsEp’ALous.—See Gamosepalous.

the upper region of the embryo sac above the egg

TAp’ULAR.—Flattened from above downward.
TANGENT’IAL.—Applied to cell walls more or less parallel to a tangential plane of

the plant organ. The walls which are at right angles with the radial walls.
Tapr’rum.—A layer of cells lining the cavity of an anther sac.
TAp-Root.—TLhe main root coursing directly downward.
Taxon’omy.—The science of classification.
Trc’meNn.—The inner seed coat.

TeLEeu’TOsPORE.—A spore produced by the Rusts toward the close of the season

which forms a promycelium the next year.
Ten’prit.—A modified stem, stipule, leaf, or leaflet which has taken on the form

of a slender appendage that is capable of coiling spirally around some object.
Trerato.’ocy.—The study of monstrosities.
Trrete’.—More or less cylindrical.

738 PHARMACEUTICAL BOTANY

TrER’MINAL.—Pertaining to the end or apex.

Ter’NATE.—In threes.

TeRrRES’TRIAL.—Growing on land.

Trs’sELLATED.—Marked like a checkerboard.

Tes’rA.—The outer seed coat.

Tretra.—A prefix of Greek origin signifying four.

Trt’rAp.—A group of four immature pollen grains cut off by pollen mother cells.

- 'TeTRACAR’PELLARY.—Having four carpels.

TeTRADYN’AMouS.—Having six stamens, four of which are longer than the other
two.

TreTRAG’ONAL.—Four-angled.

TETRAM’EROUS.—-Said of flowers that have the number four or multiple thereof
running through their various whorls.

TetTRAN’pROus.—Having four stamens.

TrTRAPET’ALOUs.—Having four petals.

TEeTRASEP’ALOUS.—Having four sepals.

Tr’ TRARCH.—Said of a radial vascular bundle having 4 xylem and 4 phloem arms
alternating with one another.

Trr’rAsporEs.—Applied to the asexually produced spores of the Mi loridee group of
Red Alga on account of being formed in groups of four in the mother cell.

Trerras’ticHous.—Said of leaves when they are arranged in four vertical rows
upon a stem.

TuHaA.’amus.—Another name for receptacle.

TuHAL’Lus.—A plant. body showing no differentiation into root, stem, or leaf.

Tuermor’RropisM.—Response of living matter to the stimulus of heat or cold.

THorN.—See Spine.

THroAT.—The opening into the tube of a gamosepalous asta or gamopetalous
corolla.

Tuyr’sus.—A compact panicle of flowers like the Lilac or Sumac.

Tis’suz.—An aggregation of cells of similar source, structure and function in inti-
mate union.

To’MENTOsE.—Covered with dense, matted, wooly hairs.

Tor’ruous.—Bent or twisted irregularly.

To’rus.—Another name for receptacle.

Tra’cHEeA.—An elongated cylindrical or prismatic tube wath open ends, found in
the xylem of the fibrovascular system and serving for the conduction of crude

Sap.

Tra/cuer.—An elongated, taper ended, tubular cell with closed ends and pitted
walls found in the xylem and serving for water conduction and support.

‘TRANSPIRA'TION.—The giving off of watery vapor by the plant.

Tri.—Three.

TRIADEL’PHOUs.—Having the filaments in 3 sets.

TriAn’pRous.— Possessing three stamens.

Tri’ARcH.—Applied to a radial fibrovascular bundle having three xylem and three
phloem arms alternating with one another.

TrICAR’PELLARY.— Possessing three carpels.

Tricu’ostAst.—An internal hair, like those projecting into the intercellular-air-
spaces of the stems of certain Water Lilies.

Tricu’ocyne.—A slender appendage to the carpogonium.

GLOSSARY 739

TricH’omMe.—A plant hair. ©

TricHor’omous.—Three-branched or forked.

Triro’LIATE.—Said of a compound leaf having three leaflets.

TRimor’PHOus.—Possessing three kinds of hermaphrodite flowers in the same
species, differing in the relative length of their stamens and carpels.

Tris’tr1cHous.—Three ranked.

TRITER’/NATE.—Applied to a compound leaf whose petiole divides into three sec-
ondary petioles, each of which again divides into three tertiary petioles, each
division bearing 3 leaflets,

Trun’cATE.—Ending abruptly as if cut off or flattened at the summit.

Tu’BER.—A short excessively thickened end of an underground stem.

Tu’BERCLE.—A small wart-like outgrowth upon the rootlets, roots or subterranean
stems of various plants.

Tu’BEROUs.—Bearing or resembling tubers.

Tu’NnIcATED.—Covered with successively overlapping coats as the bulb of an Onion.

_Tur’BINATE.—Top-shaped.

TurRGEs’cENT.—Swelling.

Ty1o’sts.—A protrusion of the wall of a cell through the pit in the wall of an
adjacent vessel and appearing in the cavity of the latter.

Typre.—An individual possessing the essential characteristics of the group to which
it belongs. :

Um’seL.—The typical inflorescence of the family Umbellifere. A more or less flat-
topped indeterminate inflorescence in which the pedicels spread like the stays of
an umbrella.

Un’ cirrormM.—Hook-shaped.

Un’‘prersHrus.—A low shrub-like plant whose base is woody and upper portion
herbaceous.

Un’puLaTE. —Having a wavy margin.

Un1.—A prefix of Latin origin meaning one.

UntLat’ERAL.—One-sided.

Uniioc’uLAr.—One-celled.

Unisr/RIATE.—Arranged in a single row, as the cells of some plant hairs.

Ur’/ceoLATE.—Urn-shaped.

Ure’pospore.—A one-celled spore produced during the life history of a Rust.

U’tricte.—An akene with a bladdery pericarp or calyx as Chenopodium fruit.

Vac’voLe.—A cavity within the protoplasm of a cell usually containing cell sap.

VALv’ATE.—Applied to the leaves of a flower in the bud stage when their margins

meet but do not overlap.
Va.ve.—One of the halves of a diatom. One of the parts of a pericarp that splits

open when ripe. ae
VartA’tTION.—The occurrence of differences among the individuals of a species or
variety.
Vari'Ety.—A group of individu
from the rest of the species.
Vas’cuLAR Bun’pLe.—A group of tubular cells frequently combined with fibers
or other kinds of cells coursing through the plant and serving primarily
uction and secondarily for mechanical support, storage, etc.

als within a species that differ in some minor respect

for cond

740 PHARMACEUTICAL BOTANY

Vas'cuLum.—A collecting case used by botanists.

VeGc’ETABLE.—A plant.

Vern.—A strand of fibrovascular tissue in a leaf.

VELA’MEN.—An absorptive tissue composed of several layers of dead cells cover~-
ing the roots of some tropical epiphytic orchids and aroids.

VENA’TION.—The arrangement of veins in a leaf.

VEN’TER.—The enlarged basal portion of an archegonium.

VEN’TRAL CANAL’ CeLL.—A cell beneath the entrance of the neck portion of an
archegonium.

VeERMIC’ULAR.—Worm-shaped.

VERNA’TION.—The manner in which leaves are disposed in the bud.

VeER’RUCOSE.—Wart-like.

VERTICILLAS'TER.—A pair of dense cymes in the axils of opposite leaves.

VERTIC'ILLATE.—Whorled.

Ves’sEL.—See trachea.

Viiiosr’,—Covered with soft, thin, rather straight hairs.

Viriwes’cent.—Greenish.

Vis’ciw.—Sticky.

Vrrra.—An oil tube in the fruit of an Umbelliferous plant.

Votatite Or..—An odorous, liquid hydrocarbon not leaving a greasy stain on
filter paper.

Vor'vA.—The covering extending from the margin of the cap to the base of the stipe

in some toadstools, which is ruptured during growth and adheres around the
bulbous base.

WATER Pores.—See Hydathodes.
_Waxes.—Esters of the fatty acids, other than glyceryl esters, usually found in plants
as a layer over the cuticle of various stems, leaves and fruits.
Woop.—All that portion of a woody exogenous plant axis lying inside of the
cambium line; also applied to the xylem region of conducting bundles.
Woop CircLe.—In exogenous stems and roots of secondary structure, all the region
within the cambium line except the pith and internal RL if present, and
consisting of the xylem and the xylem portion of medullary ra
Woop Fisers.—Fibers found in the xylem.
Woop Ray (Xylem Ray).—All that portion of a medullary ray between two wood
wedges.

Woop Wepces (Xylem Wedges).—That portion of the oes region between two
wood medullary rays.

Xyv’Ltem.—That portion of a vascular bundle which contains trachez or tracheids
or both and often wood parenchyma and fibers.

Zoéc.oer’A.—A gelatinous mass of bacteria.

Zo’6sProrE.—A ciliated spore having the power of movement.

ZyGomor’pHic.—Applied to irregular flowers like those of the Broom and Pea
which can only be divided into similar halves in one direction. cf.
Actinomorphic.

Zyc’osporE.—A spore resulting from the union of two like gametes.

Zy'cote.—A cell resulting from the union of two sexual cells or their nuclei.

Zy’mocEN.—A microorganism capable of producing fermentation.

Index

Abies, 465 Actinomyces Myricarum, 179, 373, 500
Abies balsamea, 466 Adder’s Tongue, 434
Abies sibirica, 466 Adhesion, 268
Abrus precatorius, 540 Adiantum, 434, 435
Absinthium, 615 Adiantum Capillus-Veneris, 436
Absorption of nutrient salts, 155 Adiantum pedatum, 436
selective, 155 Adnation, 265
Acacia, 301, 536, 539 Adonis N, F., 519
Acacia Catechu, 540 : Adonis vernalis, 519
Acacia Senegal, 95, 539 cidiospores, 401
Acajou Gum, 552 AEcidium, 401
Acanthacea, 602 AEcium, 401
Acanthus Family, 91, 602 Aigle Marmelos, 546
Accretion, defined, 27 Aerobe, 34
Acer, 554 : Aerogens, 364
Acer spicatum, 554 AEthallia, 374
Aceracea, 554 Agar, 361
. Acetic acid, 680 Agaricacea, 405
Achillea, 615 : Agaricales, 403
Achillea millefolium, 615 Agaricus, 404
Achras Sapota, 582 Agaricus campestris, 405
Acid alcohol, 680 Agathis australis, 466
Acidum Gallicum, 507 Agathis loranthifolia, 466
~ Acidum Tannicum, 507 Agave, 619
Aconite, 162, 518 Agave americana, 75
Aconiti Folia, 519 Agavose, 75
Aconitine, micro-chemic test for, 86 Aggregation-body, 243
Aconitum, 518 Agricultural Botany, defined, 3
calyx of, 266 Agrimonia, 535
fruit of, 301 Agronomy, defined, 3
galea of, 268 Agropyron repens, 485

histology of, 175 Fig. of cross section of stem, 219

Aconitum Napellus, 86, 518, 619 Ailanthus Family, 546
Aconitum Napellus var. roseum, Fig. of Arzoacea, 514

cross sect. of stem, 130 Akene, 303
Aconitum U.S.P., 519 Albumen, defined, 314

Acorus Galamus, 487, 618 endospermic, 314
Fig. of, 488° mode of formation, 314
Fig. of cross section of root, 144 perispermic, 314
Actea, 518 Albumens, 93
Alburnum, 213

Actinomyces, 372

Actinomyces bovis, 373 Alcaligenes melitensis, 373

741

Black, 552
Alder Buckthorn, 103, 556
Aldrovanda, 530

vesiculosa, 628

Alectoria, 417
Aletris, 490
Aletris farinosa, 490
Aleurites cordata, 551
Aleurone grains, 93
Alga-like Fungi, 376
Algz, 333

blue green, 334, 413

brown, 351

green, 336, 413

red, 357

siphon, 346
Alkaloidal reagents, 84, 85
Alkaloids, 84-87, 397
Alkanet, 593
Alkanna tinctoria, 593
Allamanda, 587
Allelomorphs, 644, 647
Allium, 491
Allium sativum, 491

Fig. of bulb of, 191

_ Almond, 535
Alnus, 504
Aloe, 490
Aloe, 84, 220, 241

Serox, 490

Perryi, 490

vera, 490
Aloin, 84, 490
Alpinia officinarum, 492

Alternation of generations, 71, 423

Althzea, 561 ge
Flores, 562
Folia, 562
mucilage in, 95

Althea officinalis, 88
rosea, 562 —

Alum root, 531

Alyssum, 277

Amanita, 407
deadly, 408

* INDEX

Amanita, toxin, 409
Amanita muscaria, 407, 408, 409
_ phalloides, 408, 409

Amanitahemolysin, 409
Amaranthacea, 514

pollen of, 283
Amber, 466
Ambrette seed, 562
Ambrosia, 283
Amelanchier, 535
Ament, 259
Amino-acids, 93
Ammonia water, 681
Ammoniacum, 577
Ameba, 24

Figs. of, 25, 27
Ameeboid movement, 26
Amomum melegueta, 492
Amygdala Amara, 536
Amygdala Dulcis, 536
Amygdalacea, 535
Amygdalin, 103
Amygdalus, 535

communis, 635, 636

Amylodextrin, 82
Amyloplastids, 42
Amylum, 485

Amylum Marantz, 493
Anabaena, 335, 336
Anabolism, defined, 31
Anacardiacea, 551

Anacardium, 552

Anaphalis margaritacea, 615
Andrea, 426

Andrzeales, 426
Androeecium, 262, 274
Andromeda, 580
Andropogon squarrosus, 485
Anemone, 260, 304
Anethol, 576

Angelica Archangelica, 577
Angelica Fructus, 577
Angelica Radix, 577

INDEX

Angiosperm, development of female
gametophyte, 292, 474
development of male gametophyte,
292, 474
life history of an, 471
Angiospermz, 443
Angiosperms, 15, 443, 471
classes of, 479
distinctions from gymnosperms,
478
fertilization in, 292
resemblances between and gymno-
sperms, 478
Aniline, blue, 681
brown, 681
chloride, 681
sulphate, 681
Anise, 258, 576
Anisum, 576
Annatto, 565
Annual plants, 162
Annual rings, 205, 206
thickening, 205
Annulus, 67, 407, 433
false, 409
true, 407
Anonacea, 515
Anthemis, 615
Anthemis nobilis, 615
Anther, 274, 277
adnate, 276, 282
attachment of, 282
confluent, 276
dehiscence, 279, 475
development of, 279
extrorse, 282
Figs. of cross sect. of, 278, 280
gross structure of, 277
histology of, 277
innate, 276
introrse, 282
lobes, 277
syngenesious, 276
versatile, 276, 282
Antheridia, 346, 348, 355, 426
Antheridium, 358
Antherozoids, 339, 441
Anthocarp, 298
Anthoceros, 419

743

Anthocerotales, 419
Anthocyanins, 100
Anthophore, 266
Anthotaxy, 256
Antigonon leptopus, 512
Antipodals, 292
Apetala, 499
Apii Fructus, 577
Apiol, 577
Apium graveolens, 577
Apocynacea, 587
Apocynum, 589
Apocynum cannabinum, 587, 589
Apothecia, 388
Apothecium, 388, 416
Appendages, epidermal, 124
Apple, 305, 534
Fig. of flower, 271
Fig. of fruit, 306
Rust, 402
Appressorium, 378
Apricot, 534
Aquifoliacea, 552
Aquilegia, 269
Arabis, 528
Aracea, 487
Arales, 487
Aralia, 575
Aralia hispida, 575
nudicaulis, 575, 576
racemosa, 575
spinosa, 575, 576
Araliacea, 575 -
Arbor vite, 465
Arbutus unedo, 579
Archegonia, 422, 426
of pine, 451, 452, 453
Archegonium, 69, 418
Archichlamydez, 499
Arctium Lappa, 615
minus, 612, 615
Arctostaphylos, 580
manzanita, 582
Uva Ursi, 580, 581
Fig. of, 581
Areca Catechu, 487
Areca, 487
Arecoline Hydrobromidum, 487
Aril, 313

744

Arillode, 314

Arisaema triphyllum, 487

Aristolochia, 275, 276, 511
reticulata, 512
Serpentaria, 512
sipho, 199, 200

Aristolochiacea, 510

Aronia, 535
Arrowroot Family, 314, 492
Artemisia Absinthium, 615 ~

Asagrea officinalis, 490, 491
Asarum, N. F., 512
Asarum, 149, 511

latex cells in, 133

Black, 121, 122
Mountain, 74, 535
Asparagine, 88

differences from Pencillium, 391
fumigatus, 392—
glaucus, 391

*

INDEX

Aspergillus herbariorum, 391
niger, 393, 394
oryza, 102, 391, 392
Asperula odorata, 605
Aspidistra, 291
Aspidium, 53, 435
American, 434
Aspidosperma, 589
Aspidosperma Quebracho-blanco, 589
Asplenium, 435
Assimilation, defined, 225
parenchyma, 116
Asteracea, 610
Astragalus, 95, 539
Astragalus gummifer, 95, 639
Atriplex, 514
Atropa, 596
Atropa Belladonna, 88, 597, 698
Atropina, 597
Aurantiea, 545
Aurantii Dulcis Cortex, 546
Amari Cortex, 546

Avena, N. F., 485
Avena sativa, 485
Awn, 274, 313
Azalea, 579, 580

Azolla, 437

Bacca, 305

Bacillariex, 348

Bacilli, anthrax, 368
types of, 370

Bacillus, 365, 372

Bacteria, 362, 363
aérobic, 34
amphitrichous, 366
anaérobic, 34
chemical composition of, 369
classification of, accd. to phenom-

ena, 364

INDEX

Bacteria, disease-producing, 373
forms of, 355
higher, 372
iron, 373, 8374
lophotrichous, 366
lower, 370
monotrichous, 366
morphology due to cleavage, 367
motility of, 365
nitrifying, 182, 367
nutrition of, 366
parasitic, 366
pathogenic, 374
peritrichous, 366
products of, 367
rapidity of multiplication, 369
reproduction of, 367
saprophytic, 366
sporulation, 368
sulfur, 372, 374
thermophilic, 364
Bacteriacea, 371
Bacterial cultures, 364
Bacteriology, defined, 3
Bacterium, 372
Balata, 582
Bald Cypress, 619
Balm, 596
Balmony, 600
Balsam Apple, 608
Balsams, 97, 98
Balsamum Peruvianum, 539
Balsamum Tolutanum, 539
Bamboo, 484
Banana, 305
Banana Family, 492
Baneberry, 518
Banyan, 161
Baptisia, 540
Baptisia tinctoria, 540
Barbaloin, 84
Barberry, 519
rust on, 401
Barberry Family, 515, 519
Bark, defined, 206
Figs. of sections of stem-bark of
Cascara Sagrada, 208, 209,
210
histology of a typical, 208.

745

Bark, scale, 130
shell, 130
Barley, 75, 259
Fig. of leaf and stem, 243
Fig. of longitudinal section through
root-tip of, 156
fruit, 304
starch of, 78, 81
Barosma, 330, 545 —
betulina, 545, 546
crenulata, 546
serratifolia, 546
Basidiomycetes, 375, 397
Basidiospore, 397
Basidium, defined, 397
Basswood, 559
Bast, fibers, 122
hard, 202
Bast, soft, 202
Batrachospermum, 357
Bauhin, G., 324
Bay Leaves, 574
Bayberry Family, 176, 600
Bayberry wax, 96, 502
Bean Family, 176
Bean, Fig. of, 313
starch of, 82
Bearberry, 580
Bear’s Foot Root, 519
Beauty-berry, 593 -
Beech, 245, 304
Beech Family, 506
Beets, 514
Beggiatoa, 374
Beggiatoacea, 374
Begonia, adventitious buds of, 185
roots of, 160
Fig. of cross sect, of leaf stalk, 117
Begoniaceea, 221 5
Belz Fructus, 546
Belladonna, fruit of, 304
Belladonnz Folium, 597
Belladonnz Radix, 597
Bentham, G., 326
Benthos, 617
Benzoin, 97
Benzoin Family, 582
Benzoinum, 583
Berberidacea, 279, 515, 619

746 INDEX

Berberis, response of stamens to stimu- Bogbean, 587

lation, 36 Boldo, 248, 522

Berberis N. F., 519 Boldo Family, 522
Berry, defined, 304 Boldus, 522
Beta, 514 Boletus, 404

maritima, 514 edulis, 75, 103

vulgaris, 514 Selleus, Fig. of, 404
Betula, 504 Borage, 592
Betula alba, 505 Borage Family, 91, 591
Betula lenta, 504, 505 Boraginacea, 591
Betula populifolia, 204 Boraginea, 591, 592
Betulacea, 504 Borke, 131
Biennial plants, 163 Borrelia, 373
Bifacial leaf blade, 250, 251 Boswellia carterii, 548
Binary division, 45 Botanical nomenclature, 329
Binary fission, 28 Botany, agricultural, 3
Binomial system, 325 cryptogamic, 3
Biology, defined, 1 defined, 1
Birch Bark, 505 departments of inquiry, 2
Birch Family, 504 derivation of, 1
Birthwort Family, 510 economic, 3
Bismark Brown, 681 : geological, 3
Bistorta, 513 importance to man, 4
Bitter Almond, 103 phanerogamic, 3
Bittersweet, 188, 189, 596, 599 pharmaceutical, 3

Climbing, 552 Botrychium, 434
Bixa Orellana, 565 Botrydium, 338
Bixacea, 565 Bougainvillea, 514
Black Alder, 552 Bract, 256
Black Caraway, 519 Bract leaves, 228
Black Fungi, 395 Bracteolar leaf, 227, 256
Black Mold, 377 Bracteoles, 227, 256
Black Mustard, 529 Bramer’s reagent, 92
Blackberry, 186, 300, 535 Brake fern, 72, 433, 435
Bladder Wrack, 355 Branch Traces, 145
Bladderworts, 638, 639 diagrams of, 146
Blakeslee, 357 Branching, deliquescent, 191
Blessed Thistle, 615 - dichotomous, 355
Bloodroot, 186, 189, 524, 526 excurrent, 191
Blue Cohosh, 522 of roots, 176
Blue Flag, 492 _ Brand spores, 398
Blue Green Alga, 334 Brandy, 556

mucilage of, 95 Brassica, embryo of, 295
Blue Vervain, 593 Brassica juncea, 530
Bluebell Family, 103, 609, 610 Brassica napus, 530 © hae
Blueberry, 580, 581. Brassica nigra, 103, 629, 530 —
Blue-weed, 592 : Brauneria angustifolia, 612, 615
Bock, H., 324 pallida, 615

Bog Mosses, 425 Brayera, 259, 636

Brazil nut, 569

Brazil wood, 539, 540

Bread mold, 377

Brewer’s yeast, 382

Bridal wreath, 535

Bromelin, 103

Brooksilk, 343

Brown Algz, 95, 361

Brown, Robert, 17

Brownian movement, 365

Brucinz Sulfas, 585

Brucine, 85

Brunfelsia Hopeana, 598
hydrangaformis, 598

Bryales, 426

' Bryology, 3

Bryonia, 608

Bryonia alba, 607, 608
dioica, 607, 608

Bryony, 607
tendril of, 189

Bryophyllum, 185

Bryophyta, 13, 418

Bryophytes, 13, 418

Buckbean, 587

Buchnera, 599

Buchu, 95, 546
subepidermal cells of, 108

Buckthorn Family, 554

Buckwheat, 512
family, 314, 512
starch of, 81

Bud, defined, 184

Bud sports, 653

Budding, 45

Buds, 184, 185.

Buffalo Berry, 569

Bulb, defined, 192
scaly, 192
tunicated, 192

Bulbel, 189

Bulbil, 189

Bulbs, Figs. of, 191

Bunchberry, 577

Bundles, 140, 713
bicollateral, 141, 203
closed collateral, 140
collateral, 140
concentric, 141

INDEX 747

Bundles, fibrovascular, 140
open collateral, 140
phlocentric, 141
vascular, 140
xylocentric, 141

Burdock, 612

Burning Bush, 552

Bur-reed, 618

Burseracea, 547

Butomus, anthers of, 277

Buttercup Family, 518

Butterworts, 640

Buttonbush, 603

Buttonwoods, buds of, 185

Cacao Preeparatum, 559
Cactacea, 567
Cactus Family, 567
Cactus Grandiflorus, 568
Casalpinia brasiliense, 538, 540
echinata, 538, 540
Sappan, 540
Casalpiniacea, 538, 539
Ceesalpinus, 324
Caffeina, 563, 605
Caffeine, 85, 552
micro-chemic test for, 85
Calamus, 147, 487
inflorescence of, 259
Calamus Draco (see Damonorops Draco)
Calceolaria, 275, 600
Calcium, importance of, 156
Calcium oxalate, 89
Fig. of various forms of crystals, 88
how formed, 89
tests for, 89
types of crystals, 89, 90
Calcium pectate, 20
Calendula, 614
Calendula officinalis, 614
California-Poppy, 524
California Privet, 170
Calla, 231, 257
Calla Lily, 487
Callicarpa, 593
Callitris quadrivalvis, 466
Calluna, 579
Caltrop Family, 544
Calumba, 521

748 : INDEX

Calyptra of moss, 429 Canna Family, 492
of root, 158 starch, 82
Calyptrogen, 113, 156 style of, 289
Calyx, 262, 266 - Cannabis, 509
aposepalous, 266 Cannabis sativa, 91, 609, 610
bilabiate, 266 Cape Aloe, 490
Capillitium, 375
Capitulum, 259
Fig. of, 611
Caprifoliacee, 606
Capsella, 529
Capsella bursa-postoris, 530
Capsicum, 304, 598
Capsicum annuum, 598
Srutescens, 598
Capsule, 301
histology of a, 307
Caraway, 258
Carbon cycle, 622
Carboxylase, 104
Carcerulus, 302
Cardamomi Semen, 492
petaloid, 266 Cardamom, 300, 313
polysepalous, 266 Cardiospermum, 554
regular, 267 Carica Papaya, 566
rotate, 267 Caricacea, 566
synsepalous, 266 i
tubular, 267
urceolate, 267
Cambium, 114
cork, 114
interfascicular, 169, 172, 196, 201
intrafascicular, 169, 172, 196, 201
origin of, 113
ring, 169, 196, 197

Caryophyllus, 574
Caryophyllus aromaticus, 573, 574
Caryopsis, 304
Cascara Amarga, 539
Sagrada, 90, 208, 554, 556
Cascarilla, 551
Casparian spots, 128
Casparian strips, 128
Cassava, starch of, 81
Cassia angustifolia, 251
structure of leaflets, 251
Cassia Bark, 524
Buds, 524
Fagot, 524 :
Cassia Fistula, 539, 540
Cassia Senna, 251, 539
Fig. of, 538
Cassine, 552
Castanea, 507
Castanea dentata, 88, 506, 507
Castilloa elastiea, 101
Castor-oil plant, 550
Castor-oil seed, 312
Cataria, 595
Catechu, 539, 540
Cathartocarpus fistula, 301, 640
Catkin, 259
Catleya, 494
Catnip, 261, 595
Cat-tails, 618
Cauda, 274
Caulicle, 311
Caulophyllum, 519
Caulophyllum thalictrovdes, 519, 522
Caustics, 33
Ceanothus, 556
Ceanothus americanus, 556
Cedar, 465
Cedrus, 465
Celandine, 524
latex of, 101
Celastracea, 552
Celastrus, 552
Cell, as a fundamental unit, 17
body, 452, 460
formation and reproduction,
inclusions, 40
living, 17
membranes, 18, 19, 48

44

749

Cell, mother, 292

non-protoplasmic contents, 74
plate, 49

rhizoid, 341

sap, 18, 40

stalk, 452, 460

tube, 292

typical colorless, 18

typical green, 20

Cell walls, 23, 107

behavior of to micro-chemic re-
agents, 109

cutinized, 109, 123

growth in area and thickness, 108

primary, 23

secondary, 23, 107

silicified, 109

suberized, 109

tertiary, 24, 107

various kinds of and behavior of
each to micro-chemic re-
agents, 109

Celloidin sections, 700, 701
Cells, 10

asexual, 44

column, 321
companion, 134, 135
complementary, 204
egg, 69, 339, 356
epidermal-daughter, 253
germ, 50

guard, 123, 253

neck canal, 69, 422
neighboring, 251
palisade, 321

palisade parenchyma, 117
parenchyma, 116
secretion, 147
somatic, 50

sexual, 44

sperm, 339, 356
sperm mother, 422
sterile, 422
stoma-mother, 253
stone, 119

subsidiary, 251
typical green, 20
ventral canal, 69, 422

wood parenchyma, 117

750 INDEX

Cellulose, 107, 109 Chicle, 582
enzyme hydrolyzing, 102 Chicory, 611, 615
mucilaginous modification of, 107 Fig. of, 614
reserve, 109 Chimaphila, 581

Centaurea Cyanus, 615 Chimaphila, 579

Centaurium, 587 maculata, 579

Centrospermz, 513
Century Plant, American, 75

Ceriman, 487

Cetraria tslandica, 414, 417
Chalaza, defined, 287
Chamalirium luteum, 490
Chamomiles, 614

Chara, 346

Chara fragilis, life history, 347

Figs. of, 347, 348

Characea, 346

Charales, 346
'Charophyta, 333
Chaulmoogra oil, 565
Chelidonium, 302, 524, 528

Fig. of flower, 270, 534

Fig. of fruit of, 298

Sour, 534
Cherry, Wild Black, 534
Cherry laurel, emulsin in leaves of, 103
Chestnut, 506

umbellata, 579, 581
Chinese Galls, 552
Chionanthus, 583
Chionanthus virginica, 583
Chirata, 587

Chlor-zinc-iodine solution, 682
Choke Berry, 535
Choke Cherry, enzyme in bark of, 103
Chondodendron tomentosum, 521
Chondriosomes, 43
Chondrus, 359
Chrondus crispus, 369, 360
Choripetala, 499
Christ’s Thorn, 535
Chromatin, 18 @
Chromatophore, 42, 337
Chrome-acetic fluids, 694
Chrome-acetic-formalin fluid, 693
Chrome-osmium-accetic fluids, 694
Chromic acid, fixation, 693

solution of, 682

Chromosomes, 47, 48
behavior of ane reduction divi-
sion, 646
distribution dating maturation and
fertilization, 645

INDEX

Chromosomes, number of, 50
structure of, 645
Chrysanthemum cinerariefolium, 615
Marschallii, 615
Chrysanthemum, papillae on epidermis of,
124
roseum, 615
Chrysarobinum, 540
Cibotium, 435
Cichorium, 615
Cichorium Intybus, 615
Cilia, 366
Ciliary movement, 39
Cimicituga, 258, 519
Cimicifuga racemosa, 619, 520
Cinchona, 330, 604, 605
Calisaya, 605
Ledgeriana, 605
succirubra, 605
Cinchona U.S.P., 605
Cinchonidinz Sulfas, 605
Cinchoninz Sulfas, 605
Cinnamomum, Burmanii, 524
Camphora, 523
Cassia, 524
Loureirii, 524
Saigonicum, 524
Zeylanicum, 524
Fig. of, 525
Cinnamomum U.S.P., 524
Cinnamon Fern, 433
Cinquefoil, 534, 535
Cistacea, 567
Citric acid, 391
Citromyces, 391
Citrullus colocynthis, 608
vulgaris, 608
Citrus, 545
Aurantium, 644, 545, 546
var. Bergamia, 546
var. sinensis, 546
medica var. Limon, 546
Cladonia, 414
rangiferina, 417
Class, 327
naming of, 332
Classification, principles of, 323
Clavaria flava, Fig. of, 403
Clavariales, 403

751

Claviceps purpurea, Fig. of, 396
life history of, 395
Claytonia virginica, 514
Clearing, 695
Cleavers, 603, 605
Clematis, 518
Clove, 149, 282, 574
Clove Bark, 524
Clover, 538
Club Fungi, 375
Club Mosses, 431, 439
Cnicus benedictus, 615
Coalescence, 265
Coca, 241, 544
Coca Family, 543
Cocaina, 544
Cocaine, 85
micro-chemic test for, 85
Coccaceea, 371
Coccoloba uvifera, 512
Cocculus, 521
Coccus, 365
Cochlearia Armoracia, 530
Cocillana, 548
Cocoa, 85
Cocoanut, 486
Fig. of fruit, cut vertically, 306
Palm, 485, 486
Cocoanut oil, 487
Cocos nucifera, 487
Codeine, 87, 527.
Coenocyte, 325
Coenocytic latex cells, 133
Coffea arabica, 605
liberica, 605
Coffea Tosta, 605
Coffee, 85, 605
Cohesion, 265
Coix Lacryma-Jobi, 485
Cola Family, 557
Cola nitida, 559
Colchici Cormus, 490
Colchici Semen, 490

- Colchicine, micro-chemic test for, 87

Colchicum, 192, 300
seed, 313
Colchicum autumnale, 87, 490
Coleoptile, 318
Coleorhiza, 318

Figs. of, 447, 448, 449
Confervales, 341

Conidia, 390
Conidiophore, 390
Conidiospores, 390
Conifera, 84, 253, 465
Coniferales, 455, 464

root, Fig. of tease nbction, 128
seed, 315
seedling, 316, 319
smut, 397
Fig. of, 398

INDEX

Corolla, 262
apopetalous, 270
bilabiate, 271
campanulate, 272
caryophyllaceous, 273
choripetalous, 270
crateriform, 272
cruciform, 270
defined, 269
forms of, 273
gamopetalous, 270
hypocrateriform, 272
infundibuliform, 272
labiate, 271
ligulate, 271
papilionaceous, 270
personate, 272
polypetalous, 270
ringent, 272
rotate, 272
synpetalous, 270
tubular, 272
urceolate, 272

Corona, 274

Cortex, 206, 215
primary, 167, 195
secondary, 169, 202

Corticium, 403

’ Corydalis, 528

Corylus, 504

Corymb, 258

Corynanthe Fohimbi, 605

Corynebacterium, 312
diphtheria, 373

Cosmarium, Fig. of, 338

Coto, 524 :

Cotoneaster, 535

Cotton, 126, 561

Cottonwood, Fig. of twig of, 185

Cotyledons, 227, $11, 312
function of, 311

Couch grass, structure of stem of, 219

Cowhage, 540 |

Craf fluid, 693. |

Cranberries, 581, 619

Cranesbill, 542

Crassulacea, 221

Crategus, 535

Cremocarp, 303

Crenothrix, 373
Fig. of, 371
Cribiform tissue, 133
Crocus, 191, 192, 491, 492
Crocus sativus, 492
Cronartium ribicola, 401
Croton Eluterta, 551
Croton Tiglium, 551
Crowfoot Family, 243, 615
Crown, 274
Crucifere, 103, 268, 528, 530
Crude sap, 117
Cryptogamia, 325
Cryptogams, defined, 13
vascular, 431
Crystal fibers, 90
Crystal sand, 91
Crystalloids, 94
Crystals, columnar, 89
membrane, 90
micro-, 90, 91
raphides, 89
rosette aggregate, 89
solitary, 89
Cube Root, 540
Cubeba, 499
Cucumber, 607
Cucurbita Pepo, 608
Cucurbitacea, 607
Cucurbitales, 607
Cudbear, 416
Culm, 188
Culver’s Root, 600
Cup Fungi, 387
Cuphea, 570
Cupule, 297, 304, 507
Cupules, of liverwort, 420
Cupulifera, 506
Cuprea Bark, 605
Curare, 585
Curcuma, 492

~ Curcuma longa, 492

Zedoaria, 492
Currant, 189, 304
Cusparia Bark, 546
Custard Apple Family, 515
Cutin, 24, 107, 128
Cyanogenetic glucosides, 84
Cyanophycez, 334

755

Cytase, 102
Cytisus laburnum, Fig. of cross sect.
of branch, 207
Scoparius, 540
. Cytology, defined, 2
Cytoplasm, 18, 40
Cytoplasmic caps, 47
Cytoplasmic rotation, 39
Czapeck’s medium, 376

Dacromyces deliquescens, 402
Dacromycetales, 402
Demonorops Draco, 487
Daffodil, 192, 257
Dahlia, 162, 611

Daphne Mezerewn, 669, 6T0
Darlingtonia, 631

Califormea, 631, 633
Darwin, C., 624, 627, 654, 655
Darwin’s Theory, 654
Date palm, 486
Datura Stramonium, 597

fruit of, 297
De Candolle, A. P., 325
De Candolle, Alphonse, 326
De Jussieu, A. L., 325
De Lamarck, J. B., 653, 654
De Tournefort, J. P., 325
Deadly agaric, 408
Deadly Nightshade, 598
Decay, defined, 364
Decodon, 570

ude oa

Deutzia, eechomenon leaves <f, 125.
Devil’s Aprons, 354
Dewberry, 300

INDEX 755

Dextrin, white, 82
yellow, 82
Dextrose, 74, 485
Diageotropic, 36
Dianthera, 603
Diastase, 102, 485
Diatoma vulgare, 350
Diatomaceous Earth, 350
Diatoms, 91, 348
Fossil, 352
Dicentra canadensis, 528
Cucullaria, 528

Dicotyl Plant, morphology of type, 497 |

Dicotyl Root, histology of, 167
of transitional and _ secondary
growth, 169
Dicotyl Roots, abnormal structure of,
175
Dicotyl Seed, 319
histology of, 320, 321
Dicotyl Stems, exceptional types of, 203
structure of, 194-204
Dicotyledonez, 479, 497
Dicotyledons, 16, 479, 497
development of embryo in, 295
drug yielding, 499
leaves of, 232, 255
petiole of, 243
roots of, 167
stems of, 194, 199
Dictyostele, 145, 199
Dictyotales, 353
Dicypellium caryophyllatum, 524
Diervilla, 606
Differentiation, 16
Diffusion, 153
Digestion, defined, 26, 67
Digitalis, 163, 256, 275, 599, 600
Digitalis Pulverata, 600
Digitalis purpurea, 84, 600
trichomes of, 124, 125
var. gloxineflora, Fig. of, 601
Digitoxin, micro-chemic test for, 84
Dihybrids, 649
Dioecious, 264, 418
Dionea, 35, 530
Submerged, 625
Dionea muscipula, 35, 36, 625
Dioscorea villosa, 491

Dioscoreaceze, 491
Dioscorides, 2, 330
Diosma, 545
Diplococcus, 367, 368
Dipsacea, 608
Disaccharoses, 74, 75
Dissepiments, 285
Division, by cleavage furrows, 378
simple, defined, 28 —
Division of labor, 16
Docks, 283, 512
Dodder, 161, 188, 228
Figs. of, 162
Dog Bane Family, 587
Dog’s Tooth Violet, life history of, 471
Dogwood, 256, 261
Blood Twig, 577
Fig. of flowering branch of, 261
Flowering, 577
Dogwood Family, 577
Dorema Ammoniacum, 577
Douglas Fir, 214, 465
Dracena, stem of, 220
Dracena Draco, 489
Dragon’s Blood, 487, 489
Draparnaldia, 338
Drimys Winteri, 516
Drosera, 530
Drosera, 36, 622
anglica, 530
longifolia, 530
rotundifolia, 630, 623
Droseracee, 530
Drupe, 305
Fig. of, 306
Dryopteris, 435
Dryopteris Filix-mas, 53, 435
alternation of generations in, 71
comparative physiology of root,
stem and leaf, 63
diagram of life cycle, 72
Fig. of, 54
gross structure of frond, 55
sori and sporangia, 66
stem, 53
histology of growing apex, 58
mature root, 58
pinnule, 61, 62
‘root apex, 60

756

Dryopteris Filix-mas, histology of grow-
ing apex, sori and sporangia,
66
stem, 55
stipe, 60
Figs. of trans.-sect., 61
history of gametophyte genera-
tion, 67
sporophyte generation, 53
life history of, 53
origin of new sporophyte or diploid
plant from fertilized egg, 71
Dryopteris marginalis, 482, 435°
Duboisia, 598
Duboisia myoporoides, 598
Dujardin, 17
Dulcamara, 598
Duramen, 213

Earth Stars, 410
Ebenales, 581
Eberthella typhi, 373
Ecballium Elaterium, 608
Echinacea, 615
Echinocactus, 567
Echinocereus, 567
Echium, 592
Ecology, 3, 616
Economic Botany, defined, 3
Ectocarpus siliculosus, 352
Ectoplasm, 25, 43, 374
Egg Plant, 304, 597
Eichler, A. W., 326
Elaagnacea, 569
Elaioplastids, 42, 133
Elais guineensis, 487
Elastica, 551
Elaterinum, 608
Elaters, 422, 437
Elder, 606
Elettaria Cardamomum, 492
Elm, 303
Slippery, 506
Elm Family, 507
Elodea, 20
Embryo, 8, 295, 311, 452
Figs. showing development of,
294, 295

parts of, 311

INDEX

Embryology, defined, 2
Embryo-sac, 287, 449, 474

maturation of, 292, 474
Emetinz Hydrochloridum, 605
Emulsin, 103
Endocarp, 298
Endodermis, 127
Endoperidium, 410
Endoplasm, 25, 43, 374
Endosmosis, 154
Endosperm, 293, 314
Endospore, 282, 292
Endospores, 369, 378, 381
Endosporium, 68, 475
Endothecium, 278, 280
Endotoxins, 370
Engler and Prantl, 326
English ivy, 575

roots of, 161
Entada scandens, 301
Enzyme, defined, 26, 101
Enzymes, 101

classification of, 102

proteolytic, 94, 103
Eosin, 682
Ephedra, 467, 468, 469, 470
Ephedra distachya, 471
Ephedra equisetina, 471
Ephedra Gerardiana, 471
Ephedra sinica, 468, 470, 471
Ephedrine, 471
Epicalyx, 269, 535
Epicarp, 298
Epicotyl, 311
Epidermal outgrowths, 124

hairs, 124

papilla, 124 -

scales, 124
Epidermis, 122
Epigaa repens, 581
Epilobium, 569
Epiphytes, 161
Epipremnum pinnatum, 487
Epithem, 124
Equisetineae, 91, 431, 437
Equisetum, 437

arvense, 438

life history of, 43
Ergonovine, 397

ee ee es

INDEX

Ergot, 395, 397
Ergot fungus, 395
Erica, 580
Ericacea, 279, 579
Ericales, 579
Erigeron canadensis, 615
Eriodictyon, 591
Eriodictyon californicum, 691, 592
trichomes of, 125
Eriogonum, 512
Erythrea Centaurium, 587
Erythronium Americanum, life history of,
471
Erythroxylacea, 543
Erythroxylon, 543
papilla on epidermis, 132
Erythroxylon Coca, 544
Truxillense, 544
Eschscholtzia, 524
Essential elements, 155
Etzrio, 306
Ether, use of, 682
Ethylhydrocupreine, 605
Etiolin, 100
Eubatus, 536
Eucalypti Gummi, 574
Eucalyptol, 574
Eucalyptus, 574
Eucalyptus, 186, 573
globulus, 572
lamina, 250
leaves, internal glands of, 148
photomicrograph of trans. sect. of
leaf-blade, 252
rostrata, 95
Eucarya spicata, 510
Eudorina, 338
Eugenol, 574
Eumycetes, 375
Euonymus, 553
Eupatorium, 614
Eupatorium perfoliatum, 232, 614
Eupatorium purpureum, 615
Euphorbia, 133, 188, 550, 551
pilulifera, 551
resinifera, 551
Euphorbiacea, 550
Euphorbium, 101, 551
Euphrasia, 599, 600

Euphrasia officinalis, 600
Eurotium, 391
Evening Primrose, 282, 569, 657
Everlasting, 615
Evolution, 653
Evonymus, 313, 562
Evonymus Americanus, 553
atropurpureus, 553
Europaus, 552
Eye-bright, 600
Excretion, defined, 27
Exine, 281, 292
Exoascus, 381
Exobasidiales, 403
Exogonium Jalapa, 591
Exoperidium, 410
Exosmosis, 154
Exospore, 281
Exosporium, 68, 439, 475
Exothecium, 278
Exotoxins, 370

Fabaceae, 536
Fagacea, 506
Fagales, 504
Fagopyrum, 512
Fagot Cassia, 524
Fagus, 330
Fagus grandifolia, 506
Families, naming of, 332
Family, defined, 327
naming of, 332
Farfara, 615
Fast Green, 682
Fats, 95
Fatsia horrida, 575°
Fehling’s solution, 682
Fennel, 258, 576
Fenugreek, 540
Ferment, 101
Fermentation, 364
Fern, antheridium, 68, 71
apical cell,_58
apical cone, 58
archegonium, 69, 69
Beech, 435
Boston, 653
Brake, 433, 435
Christmas, 435

INDEX

Fern, Cinnamon, 433 Filament, 274
foot of, 71, 433 : gross structure of, 277
histology of, 277
Filbert, 297
Filicales, 434
Filicinea, 431
Fir, 465
Fission, 45, 367
of Amorba, 28
Fistulina, 404
Fixation, 692
of atmospheric nitrogen, 178
Fixed oils, 95

Flower, pentandrous, 274
perfect, 262
perigynous, 264
pistillate, 263
polyandrous, 274
protandrous, 291
protogynous, 291
regular, 262
staminate, 263
symmetrical, 263
tetrandrous, 274
triandrous, 274
typical, 262
zygomorphic, 263

Fly agaric, 407

Fly-traps, 530, 625

Foeniculum, 576
histology of mericarp of, 309

Feeniculum vulgare, 576

Folia Laurocerasi, 536

Foliage leaves, 228

Follicle, (See pod.), 301

Fomes, 404

Foot, 71, 428

Foramen, 287

Forestry, defined, 3

Forget-me-not, 592

Formalin-alcohol, 694

Forsythia, 583

Fothergilla, 532

Four-o’clock Family, 514

Fovilla, 475

Foxglove, 599, 601

Fragaria, 269, 535

Fragaria chiloensis, Fig. of flower, 268

Frangula, 556

Frangulin, 103

Fraxinus, 583

Fraxinus Americana, 583
Ornus, 583

Free cell formation, 46

Frond, 55, 431

Fronds, 53, 431, 432
fertile, 434
sterile, 434

Fructose, 74

Fructus Cannabis, 510

Fruit, 8, 297
definition, 297

INDEX

Fruit, dehiscence, 298
indehiscent, 298
structure, 297
sutures, 298
valves, 298

Fruits, achenial, 300
aggregate, 300, 305

Fig. of, 299
baccate, 300, 304
capsular, 300, 301
classification of, 300
compound, 300, 301
dehiscent, 298, 300
drupaceous, 300, 301
indehiscent, 298, 300
multiple, 300, 306
schizocarpic, 300
simple, 300

Fucales, 352, 364

Fuchs, L., 324

Fuchsia, 569

Fuchsin, acid, 683
basic, 683

Fucoxanthin, 100, 351

Fucus, drug, 357

Fucus, 355

Fucus vesiculosus, 355

Fumariacea, 528

Fumitory Family, 528

Funaria, 426
Fig. of, 429

Fundamental tissue, 116

Fungi, 12, 362
alga-like, 376
basidia, 397
black, 395
carrion, 411
classes of, 362
coral, 403
cup, 387
fairy club, 403
fission, 363
gill, 405
nest, 410
poisonous, 407
pore, 404
sac, 381
saddle, 394
stink horn, 411

759

760

Fungi, tooth, 404
true, 362, 375
weeping, 402

Fungi Imperfecti, 412

Funiculus, 287

Funtumia africana, 101
elastica, 101

Gaillonella, 352
Galanga, 492
Galbanum, 577
Galbulus, defined, 307
Galea, 268
Galega, 540
Galega officinalis, 540
Galen, 2
Galipea officinalis, 546
Galium, 603, 604
Galium aperine, 605
Galla, 92, 507
Galls, Chinese, 552
Japanese, 552
Galton’s Law, 652
Galvanotropism, 36
Gambir, 605
Gamboge, 563
Gamboge family, 563
Gametangia, 353
Gametangium, 344, 379
Gamete, 46, 344, 353, 379

Gametophyte, 68, 418, 420, 458

female, 292, 294, 450, 451
male, 292, 449, 450, 458
Gamopetalz, 579
Garcinia Hanburyi, 563
Garlic, 189, 277
Fig. of bulb, 191
Gasteromycetes, 410
Gaultherase, 103
Gaultheria, 581
Gaultheria, enzyme in, 103
Gaultheria procumbens, 579, 581
Gaultherin, 103
Gaylussacia, 580, 581
Geaster, 410
Gelidiacea, 361
Gelidium, 361
Gelsemium, 585

INDEX

Gelsemium, 583
sempervirens, 684, 685
stem, 204

Gemme, 421

Gemmation, 45, 385

Genera Plantarum, 326

Generative tissues, 113

Generic names, 329

Gene, modern theory of, 658

Genes, 18, 645

Genetics, 3, 643

Genotype, 647

Gentian violet, solutions of, 683

Geological Botany, defined, 3
Geotropism, 35

dia-, 36

lateral, 36
Geraniacea, 540
Geraniales, 540
Geranium, 542
Geranium, 541

fruit of, 301

maculatum, 542

petiolar structure, 245
Geranium Family, 540
Gerardia, 161, 699
Germination, seed, 318, 453
Geum, 266, 269, 535
Gigartina mamillosa, 359, 360

Glands, external, 151
internal, 148
Glans, defined, 304
Gleba, 410
Gliadins, 93
Globoid, 94
Globulins, 93
Gleocapsa, 335
Fig. of, 334
Glomerule, 261, 577
Glory-bower, 593
Gluco-alkaloids, 87
Glucoside enzymes, 103
Glucosides, characteristics of, 83
Glucosum, 485
Glumes, 484
Glutelins, 93
Glycerine, dilute, 683
gum, 683
Glycerin-Gelatin, 691
Glycine hispida, 482
Glycosides, 83
Glycyrrhiza, 275, 539
glabra, 637, 539
glandulifera, 539
typica, 539
Gnaphalium polycephalum, 615
Gnetaceea, 469
-Gnetales, 469 ~~
Gnetum, 469, 470
Golden Bell, 583
Golden Seal, 518, 620
Gonidia, 414
Gonophore, 266
Gooseberry, 305
Goosefoot, Fig. of flower, 265
Goosefoots, 619
Goosefoot Family, 514
Gossypii Radicis Cortex, 561
Gossypium, 561
Barbadense, 561
herbaceum, 560, 561, 562
Gossypium hursutum, 560, 561, 562
Gossypium Purificatum, 561
Gourd, defined, 305
Gourd Family, 607
Grain, defined, 304
Grains of Paradise, 492
Graminales, 483

INDEX 761

Gramineae, 91, 304, 484
pollen of, 267
silica in, 91
Gram’s method, 351
Granatacea, 570
Granati Fructus Cortex, 571
Granatum, 92, 571
Grape, 188, 261, 304, 557
Grape Family, 556
Grape Fruit, 305, 545
Grape Sugar, 57
Grape Vine, 202, 556
tendrils of, 189
Graphis, 414
Grass, diagram of flower, 562
petiole of, 242
Grass Family, 259, 304, 484
Greek Tea, 596
Green Algze, 336
Felt, 346
Mold, 388
Greenbrier, histology of stem, 217
Fig. of T. S. stem, 218
Grindelia, 615
Grindelia camporum, 615
cuneifolia, 615
squarrosa, 612, 615
Gross anatomy, defined, 2
Ground Cherry, 297
Ground Ivy, 594
Ground meristem, 113
Growth, 26
of perennial dicotyl stem, 199
Gruinales, 540
Guaiac, 544
Guaiaci Lignum, 545
Guaiacum, 544
Guajacum officinale, 5644, 545
sanctum, 643, 544, 545
Guarana, 554
Guard cells, 253
Guarea Rusbyi, 548
Gulf Weed, 355
Gum, 95
Cherry, 95
Kauri, 466
Red, 95
Spruce, 466

762 INDEX

Gum resins, 97
defined, 98
Gums, 94, 95
Gutta Percha, 101, 582
Guttifera, 563
Gymnosperm, life history of a, 444
Gymnosperma, 443
Gymnosperms, 15, 443
classification of, 455
differences from angiosperms, 438
Gyneecium, 262, 283
Figs. of forms of, 286
Gynophore, 266
Fig. of, 286

Haberlandt, 123
Habitat, defined, 3
Hackberry, 507

Hadrome, 142 .

Hzmatoxylin stain, 683
Hzmatoxylon, 539, 540
Hematoxylon campechianum, 540
Hagenia abyssinica, 536
Hair cystoliths, 91
Hairs, aggregate, 125
barbed, 124
branched, 125, 126
candelabra-shaped, 124
clavate, 124
climbing, 126
epidermal, 124
functions of, 127
glandular, 126
hooked, 124, 126
multicellular, 124
multiseriate, 126
non-glandular, 124, 126
root, 127, 162, 154
stellate, 124, 125
stinging, 126
tufted, 124
unicellular, 125, 126
uniseriate, 125, 126
Halidrys siliquosa, 357
Halophytes, 164, 617, 618
Haloxylon, 514 .
Hamamelidacea, 531.
Hamamelidis Cortex, 532
Hamamelidis Folium, 532

Hamamelis virginiana, 532

Fig. of, 533
Hancermia, latex in, 101
Hard Bast, 202
Hardening, of soft materials, 695
Harebell, 272, 277
Haustoria, 161

of fungi, 362
Hawkweeds, 614
Hawthorn, 258, 535
Hay Fever, pollens responsible for, 283
Hazelnuts, 504
Head, defined, 259
Heart-wood, 213
Heath, 273
Heath Family, 579
Heather, 579
Hedeoma, 595
Hedeoma pulegioides, 595
Hedera Helix, 575
Heliamphora, 629

nutans, 629
Helianthemum, 567
Helianthemum canadense, 567
Heliconia, 253
Heliotropea, 591, 592
Heliotropism, 34
Helleborus, 519
Helleborus fetidus, 519
Helleborus niger, 519
Helonias, 490
Helophytes, 617, 618
Helvellales, 394
Hemlock, 465
Hemophilus, 373
Hemp, 507, 508

seed, 510
Henbane, 597

Hercules Club, 575
Heredity, 643
Hesperidin, 83
Hesperidium, 305 |

INDEX

Heteroauxin, 104
Heterocysts, 336
Heuchera, 531
Hevea, 550
latex of, 101
Hevea braziliensis, 551
Mibiscus, flower of, 267
Hibiscus abelmoschus, 562
Hibiscus Syriacus, 560
Hickories, 504
stems of, 202
Hilum,
of ovule, 287
of seed, 321
of starch grain, 77
Hip, defined, 304
Histological technique, 675
Histology, defined, 2
of dicotyl roots, 167
of Aconitum, 175
of annual dicotyl stem, 194
of anther, 277
of Aristolochia Sipho stem, 199
of bark, 208
of California Privet root, 170
of Cascara Sagrada, 208
of Convallaria rhizome, 215, 217
Corn stem, 216
of dicotyl roots, of primary growth,
167
of secondary growth, 169
of dicotyl stems, 194
of dicotyl tuberous root, 175
of Ephedra sinica stem, 469, 470
of fern rhizome apex, 58
pinnule, 61
root, 58
sori and sporangia, 67
stem (rhizome), 55
stipe, 60
of filament, 277
of fruits, 307
of grass stems, 219
of Greenbrier stem, 217
of herbaceous monocotyl stem, 215
of leaves, 247
of Lilac leaf blade, 247
of mericarp, 309
of monocotyl rhizome, 215

763

Histology, of monocotyl roots, 165
of monocotyl stems, herbaceous,
215
woody, 217
of perennial dicotyl stem, 199. 202
of petioles, 242, 244
of seeds, 315
of sunflower stem, 194
of typical monocotyl rhizome, 215
of Vanilla capsule, 307
Holdfast, 355
Holly, 552
Hollyhock, 401
Honey dew, 395
Honey Locust, 189
Honeysuckle, 272, 291, 305, 606
Honeysuckle Family, 606
Hooke, Robert, 17
Hooker, J. D., 326
Hop, 127, 188, 259, 291, 507
Fig. of, 508
Hordeum vulgare, 485
Horehound, 261, 595
Hormogonia}; 336
Hormones, 104
Hornbeam, 504
Horse chestnut, 187, 256
Horsemint, 594, 596
Horsenettle, 596
Horsetails, 431, 437
Horticulture, defined, 3
Hound’s Tongue, 593
Houseleek, 96, 189
Huckelberries, 581
Hudsonia, 567
Humulus, 510
glandular trichomes of, 125, 127
Humulus Lupulus, 510
Hutchinson, J., 326
Huxley, 30, 31
Hyacinth, 489
Hybrid, defined, 329
Hybrids, ‘Splitting of,’ 645
Hydathode, 124
Hydnacea, 404
Hydnocarpus anthelmintica, 565
Hydnocarpus Wightiana, 565
Hydnum, 404

764

Hydrangea, 264, 531
arborescens, 531
Hydrastis, 260, 518
Hydrastis canadensis, 518
Fig. of, 517
Fig. of colony of, 620
Hydrochloric acid, 684

Hyoscyamus U.S. P., 598
fruit of, 301
Hyoscyamus muticus, 126, 126
niger, 598
Fig. of calyx, 266
Hypericacea, 564
Hypericum, 276, 565
perforatum, 564, 565
Hypha, defined, 362
Hyphe, aerial, 377, 389
rhizoidal, 377
stoloniferous, 377
submerged, 377, 389
Hyphomycetes, 333, 362
Hypocotyl, 311
Hypocreacea, 395
Hyssopus, 596
Hyssopus officinalis, 596

Iceland Moss, 414
Ignatia, 585

Imperatoria, 477
Imperatoria Ostruthium, 477
Inclusions, 74

INDEX

~ Todine, a source of, 354

Indchiscent fruit, 298, 300
Indeterminate inflorescence, 256
Indian Corn, 315, 397
Figs. of grain and seedling, 316, 317
histology of seed, 315
Indian Cress, 267

Indigo, 540

Indigofera tinctoria, 540
Indirect nuclear division, 47
Indusium, 67, 433

Ingestion, 26
Inheritance, 643, 652
Integuments of angiospermous ovule,
287
Intercellular-air-spaces, lysigenous, 147
147

Internal phloem, 204
Internal secretion reservoirs, 148
Internode, defined, 186
Intine, 282, 292
Intussusception, 27, 31, 108
Inula, 615

Inula Helenium, 615

Inulase, 103

Inulin, 82

Invertase, 75, 102
Involucre, 256

Involucres, persistent, 297

INDEX

Iodine and potassium iodide, solution of,
684

Iodine water, 684

Ipecac, 603

Ipecacuanha, 605

Ipomoea, 591

Ipomea orizabensis, 591
pandurata, 591
simulans, 591

Tridacea, 491

Tris, 300, 491
caroliniana, 492
Family, 492
florentina, 492
germanica, 492 °
pallida, 492
petiole of, 249
style of, 289
versicolor, 492

Irish Dulse, 361
Moss, 359

Iron, as an essential element, 157

Ironwood, 504

Irritability, 31

Isoetacee, 442

Tsoetes, 442

Ivy, 188
English, 575

Jacket, 451
Jack-in-the-pulpit, 192
Jalap, 590
abnormal root structure, 175
Tampico, 591
Wild, 591
Jalapa, 591
Jamestown weed, 297
Japan ivy, 189
Japan wax, 96
Japanese Galls, 552
Jateorhiza palmata, 521
Fig. of, 523
Jequirity, 540
Jerusalem Artichoke, 192, 611
'Jew’s ear fungus, 402
Job’s Tears, 485
Juglandacea, 502
pollen of, 283
Juglandales, 502

765

Juglans, 504
Juglans cinerea, 504
nigra, 504
regia, 504
Jungermanniales, 419
Juniper, 465
Juniperus, 466
Juniperus, 465
communis, 466
Oxycedrus, 466
Sabina, 466
Virginiana, 402, 466
Jussieu, A. L., 325
Jute, bast fiber of, 120

Kalmia, 580

latifolia, corolla of, 272
Kamala, 551

trichomes of, 125
Karyokinesis, 47
Katabolism, 31
Kava, 499 |
Keel, 274, 538
Kelps, 353

Figs. of, 354, 355
Kernel, 314 :
Kieselguhr, 350
Kino, 92, 540
Knot-grasses, 512
Kochia, 514
Kola, 85, 559
Krameria, 92, 540

Krameria argentea, 540

triandra, 540
Kiihne, 36

Labarraque’s solution, 684
Labellum, 271

Labiata, 261, 693
Labrador tea, 619

Lachnea, 387

Lactase, 102 ~
Lactose, 102

Lactuca virosa, 615
Lactucarium, 101, 615
Lamarck, J. B., 653, 654
Lamarckian Theory, 653
Lamiaceea, 593

INDEX

Leaf, palmately-compound, 234, 240

Laurel Family, 515, 522
Laurus nobilis, 524”

Leaves, coriaceous, 241
decompound, 240
development of lamina of, 246
dorsiventral, 246
-hydrophytic, 248
-mesophytic, 247
Fig. of cross sect. of, 247

-umbrophytic, 247
-xerophytic, 248
foliage, 228
forms of, 233
imparipinnate, 240
interruptedly pinnate, 240
lyrate, 240
origin and development of, 228
palmate, 240
paripinnate, 240
pellucid-punctate, 241
pinnate, 240
primordial, 228
runcinate, 237
scale, 228
simple, 233
stomata of, 251
ternate, 240
trifoliate, 240
types of Angiospermous, 227
_ veins of, 230
_ Lecanora, 416
_ Lecanora esculenta, 417
_ Lechea, 509
Lecythidacee, 569
Legume, 301
nose, 536
placentation in, 288
root tubercles of, 176
_ Stipules of, 245

484

INDEX 767
Levisticum officinale, 577
Levulose, 74
Liber fibers, 122
Lichens, 13, 418, 619
Licmophora, 351
Life history, of an angiosperm, 471

of Aspidium, 53
of black mold, 377
of-Chara, 347
of Claviceps purpurea, 395
of Erythronium, 471
of Ectocarpus, 352
of Equisetum, 437
- of fern, 53
of Fucus, 355
of gymnosperm, 444
of Lycopodium, 439
_f a moss, 426
of mushrooms, 405-409
of smut, 397
of Spirogyra, 343
of wheat rust, 399
of white pine, 444
of yeast, 382
of Lamia, 456
Light Green, 684
Lignin, 108, 118
Lignocellulose, 109
Ligule, 242, 245, 484
Liguliflore, 618, 614
Ligustrum, histology of root, 170
Lilac, 261, 583
Liliaceae, 489
Liliales, 489
Lilium, 489, 490
Lily, 192
Calla, 487
Canada, 191
diagram of flower, 488
Fig. of T. S. anther, 280
ovary, 473
pollen of, 281, 283
Tiger, 189, 191
Lily Family, 489.
Lima bean, Figs. of, 320
histology of, 321

Limb, 266

Limonis Cortex, 546

Linacea, 542

768

Linaria, 599, 600
corolla of, 272
Fig. of, 273
stamens of, 275
Linden, American, 559
Linden Family, 559

Liriodendron Tulipifera, 516
carpels of, 286

Litmus, 416

Little Club Mosses, 442

Liverworts, 13, 418

l-mandelonitrile glucoside, 103

Loasa hispida, hairs of, 126

Lobelia, 163, 274, 275, 610
Lobelia inflata, 610
Lobeliacee, 609, 610
Locule, 473
Locust, corolla of, 271
modified stipules of, 246
Lodicule, 484
Loganiacea, 583
Loment, 301
Fig. of, 303
Lomentum, 301
Lonchocarpus, 540
Lonicera, 606
ese sr esiier Family, 570

Macrocystis, 351, 364
, defined, 2
Madder, 605

Madder family, 603
Magnesium as an essential element, 157
Magnolia, 301, 516
Magnolia Family, 515
Magnoliacea, 515
Mahogany Family, 548
Mahonia, 519
Male Fern, 53

life history of, 53
Mallotus philippinensis, 551
Mallow Family, 559
Mallow Flowers, 562
Maltase, 102
Maltose, 75
Maltum, 485

Malus, 584, 536

Malus sylvestris, ———_ 271

Manihot, 550

Manihot Glaziovii, 101

utilissima, 103, 551

Manna, 583

Scriptural, 417

Manna Ash, 74

Manzanita, 582

Maple, 162, 303

Red, 185

Maple Family, 554

Maranta arundinacea, 493

starch, 79, 81

Maranta, petiole of, 243

Marantacea, 314, 492

Marchantia, 418

Marchantia polymorpha, 419

life history of, 419

Marchantiales, 419

Marrubium, 595

Marrubium vulgare, 595

Fig. of cross sect. of leaf blade, 249
hairs of, 124, 125

Marsdenia Condurango, 589

Marsh, defined, 618

Marshmallow, 559

Marsilea, 437

Masterwort, 577

Mastiche, 552

Maté, 85, 552

Matico, 232, 499

Matricaria, 614

_ Matricaria Chamomilla, 613, 614
Maturation of pollen grain and forma-
tion of male gametophyte in
Angiosperms, 292

in Erythronium, 474

in White Pine, 449

of embryo sac and formation of
female gametophyte in Angio-
sperms, 292

Maw seed, 528

May Apple, calyx of, 268

Mayer’s albumin fixative, 698
Mechanical tissues, 151

Medulla, 165

Medullary rays, 139

Megaceros, 419

Megasori, 285, 448

Megasporangia, 381

INDEX

769

Megasporangium, 448, 458
Megaspore, 287, 448, 449, 460, 474
Megasporophyll, defined, 283
Megasporophylls, 448, 473
Meiosis, 50

Melaleuca Leucadendron, 574
Melia Azedarach, 548
Meliacea, 548

Melibiose, 75

Melilotus, 540

Melilotus officinalis, 540
Melissa, 596

Melissa officinalis, 596

-Melochia, 558

Melon tree, 567
Membrane crystals, 90
Membrane mucilage, 94, 95
Membranes, 19

cell, 43

plasma, 19

semipermeable, 43

vacuolar, 19, 43
Mendel, G. J., 643, 644
Mendel’s Laws, 643
Mentspermacea, 521
Menispermum, 521
Menispermum canadense, 521
Mentha arvensis, 595
Mentha piperita, 594

Fig. of glandular trichomes, 125
Mentha Piperita, U.S.P., 594
Mentha spicata, 595
Mentha Viridis U.S.P., 595
Mentha viridis, 595
Menyanthes, 587
Menyanthes trifoliata, 587
Mericarp, 303

histology of a, 309
Merismopedia, 336
Meristeles, 145
Meristem, 114

_ primordial, 114, 187

Meristems, 114

apical, 115

intercalary, 115

lateral, 116

primary, 114

secondary, 114
Merulius, 404

770

Mesocarp, 298
Mesophyll, 61, 222
Mesophytes, 164, 617, 619
Mesquite Gum, 540
Metabolism, 31
Metachlamydez, 579
Metaphase, 50
Metaxylem, 143
Methyl Salicylate, 103, 505
Methylis Salicylas, 505, 581
Metroxylon Sagu, 82
Mez., Carl, 323
Mezereum, 570
Maiucrandra, 550
Micrococcus, 365
division forms of, 368
urea, 182
Micro-crystals, 90, 91
Microgametophyte, 461
Micrometer, net, 673
ocular, 672
stage, 673, 674
Micrometry, 672
Micromorphology, defined, 2
Micron, defined, 672
Micropyle, 287, 448, 474
Microscope, binocular, 670
care of, 671
compound, 663, 664
use of, 669
defined, 661 :
diagram showing optics of, 665
dissecting, 661, 662
research, 667
simple, 661
Microscopic measurement, 672
Microsorus, 274, 473
Miucrospira, 372
“Microsporangia, 277, 381, 447, 457, 473
Microspore, 280, 447
Microsporophylls, 274, 277, 447, 457
Microtomes, 677 i
hand, 677
rotary, 679, 680 :
sliding, 678, 679
Middle lamella, 20, 21, 24, 49
Midrib, 230
Mildew, 388

INDEX

Milkweed, 295, 313

pollinia of, 283
Milkweed Family, 589
Milkwort Family, 548
Milon’s reagent, formula for, 93
Mimosa, 36, 538

pudica, 37

conduction of stimuli by leaves,
37

Spegazzini, 36, 38

thigmotropic reactions in, 36, 37
Mimosacea, 538, 539
Mimusops globosa, 582
Mint Family, 118, 593
Miré, 598
Mistletoe, 161, 479, 510
Mistletoe Family, 510
Mitchella, 605
Mitchella repens, 605
Mitella, 531
Mitochondria, 43
Mitosis, 47

heterotypic, 52

homotypic, 52
Mitrewort, 531
Mnium, 426, 427
Mock-orange, 531
Mold, black, 377

blue, 388

bread, 377

green, 388
Molds, cultivation of, 376
Mollugo, 514
Momordica Balsamina, 608
Monarda, 594, 596
Monarda punctata, 596
Monilia, 412
Monimiacea, 515, 622
Monkshood, 518
Monoclinic prisms, 88, 89
Monocotyl

leaves, 242, 255

rhizome, 215

root, 165

seed, 315

seedling, 316

stems, 215-220

secondary growth in, 220

INDEX 771

Monocotyledon, morphology of typical Movement, ciliary, 39

plant of, 481
Monocotyledonez, 479
Monocotyledons, 15, 16, 481

development of embryo in, 296
drug yielding, 483
histology of leaves of, 255
histology of roots of, 165
histology of stems of, 215
petiole in, 242
Moncecious, 264, 418, 425
Monohybrids, 648
Monosaccharoses, 74
Monotropa, 580
uniflora, 580
Monstera, 487
Moonseed Family, 515, 521
~ Moonwort, 434
Moraceae, 507
Morchella esculenta, 395
Fig. of, 394
Morels, 395
Morgan, T. H., 658
Morning Glory, 188, 300
Morning Glory Family, 590
Morphine, 86, 527
test for, 87
Morphology, defined, 2
Moss, diagram of life history of, 429
Iceland, 414
Golden, 435
Reindeer, 417
Mosses, 418, 424
bog, 425
peat, 425
rock, 426
true, 426
Motherwort, 596
Mountain Ash, 74, 535
Mountain Laurel, 272, 579, 580
Mountain Mint, 596
Mountain Rose, 512
Mounts, Canada balsam, 689
glycerin-gelatin, 691
permanent, 687
temporary, 687
Movement, 39
amceboid, 26, 39
Brownian, 365

true, 365
Mucilage, 94 ;
cell content, 94, 95
membrane, 94, 95
micro-chemic test for, 95
Mucor mucedo, 380
stolonifer, 377
Mucuna pruriens, 540
Muehlenbeckia, 512
Mulberry, 307, 507, 509
Mulberry Family, 91, 507
Mullein, 259
Musa, 231
Musacea, 492
Musci, 13, 418, 424
Mushroom, common, 405
Mustard, corolla of, 270
fruit of, 302
stamens of, 275
Mustard Family, 268, 528
Mutation Theory, 657
Mutations, 653, 657, 659
Chromosome, 658

“ Mutisia, 611

Mycelium, 362, 377, 388, 397
Mycobacteriacee, 372
Mycobacterium, 372, 373
Mycology, 3

Mycorrhiza, 181, 182, 183, 394
Myosotis, 592

» Myrica, 502

Mprica caroliniensis, 96, 179, 502
cerifera, 96, 179, 502
Gale, 179
Fig. of, 503
Macfarlanei, 179
tubercles of, 179
xalapensis, 96
Mpricacea, characters of family, 501
root tubercles of, 179
Mpristica, 517
fragrans, 518
Fig. of, 516
Myristica, U.S.P., 518
Mpristicacee, 515, 517
Myrobalans Family, 574
Myronase, 103
Myrosin, 103

772

Myrrh Family, 547

Neighboring-cells, 251

Neisseria gonorrhoea, 373
intracellularis, 373

Nepenthacea, 530

Nepenthes, 241, 245, 634, 638
bicalcarata, 637
Edwardsiana, 637

Nigella, 519 2

Nightshade, Fig. of, 535
Family, 533, re,

Nitella, 40, 346

Nitrification, 367

Nitrifying bacteria, 182, 367

_ Nitrobacter, 182

INDEX

Nitrogen cycle, 182

Nitrogen fixation, 178

Nitrosococcus, 182

Nitrosomonas, 182

Node, defined, 186

Nomenclature, 329
Binomial system of, 325

Non-protoplasmic cell contents, 74

Nostoc, 336

Fig. of, 334

Nucellus, 287, 448, 474

Nuclear membrane, 19, 43

Nuclear sap, 41

Nuclei, assisting, 292
polar, 292

Nucleins, 93

Nucleolus, 18, 41

Nucleoplasm, 41

Nucleus, 18, 41

Numerical aperture, 666

Nut, 304

Nutgall, Fig. of, 505

Nutlet, 302

Nutmeg, 314

Nutmeg Family, 515

Nutrition, defined, 31

INDEX

Ocnothera lamarckiana, 657, 658

Oenothera lata, 658

Offsets, 189

Oidium, 376

Oil, benne, 602
bitter almond, 97
cassia, 97
cedarwood, 466
chaulmoogra, 565
cocoanut, 487
enzyme, 103
linseed, 542
mustard, 97
palm, 487
rose geranium, 542
Siberian fir, 466
tung, 551
turpentine, 97

Oils, fixed, 95
nitrogenated, 97
oxygenated, 97
sulphurated, 97
terpene, 97
volatile, 96

Olea, 583

Olea europea, 583

Oleacea, 583

Oleander, latex of, 101

Oleoresins, 97
defined, 98

Oleum abietis, 466
amygdale amarz, 535
amygdala expressum, 535
anisi, 516, 577
aurantii, 546
aurantii amari, 546
aurantii florum, 546
bergamotta, 546
betula, 505
betula empyreumaticum rectifi-

catum, 505

cajuputi, 574
cardamomi, 492
cari, 577
caryophylli, 574
chaulmoogre, 565
chenopodii, 514
cinnamomi, 524
citronellz, 485

773

Oleum coriandri, 577
erigerontis, 615
eucalypti, 574
foeniculi, 577
gossypii seminis, 562 |
juniperi, 466 '
lavandula, 595 Pee.
limonis, 546
lini, 543
maydis, 485
menthz piperita, 594
menthe viridis, 594
myrcie, 574
myristice, 518
olive, 583
picis rectificatum, 465
pimentz, 574
pini pumilionis, 466
ricini, 551
rose, 535
rosmarini, 595
santali, 510
santali australiensis, 510
sassafras, 524
sesami, 602
sinapis volatile, 530
terebinthine, 465
theobromatis, 559

_ thymi, 595
tiglii, 551

Olibanum, 548

Olive, 583

Olive Family, 583

Onagracee, 569

Onion, 188, 191
epidermis of bulb scale, 18

Onoclea sensibilis, 433

Oégonia, 346, 348

Oégonium, 348

Oémycetales, 376

Oéspore, 46, 293, 341, 476

Operculina Turpethum, 591

Operculum, 429

Ophioglossum, 434

Opium, 86, 87, 101, 527

Opuntia, 667, 568

Opuntiales, 567

Orange, 545
flower, 277

Orange, fruit of, 305
peel, cross section, 148
stamens of, 277

Orcein, 416

Orchid family, 494

Orchidaceae, 494

Orchidales, 494

Orchidea, 495

Orchids, Fig. of pollinia, 282
floral organs of, 494
parenchyma of, 116
stamens of, 275

Orchis mascula, 496

Orchis militaris, Fig. of flower, 272

Orchis morio, 496

Order, defined, 327

Ourouparia Gambir, 605
Outline of plant groups, 328
Ovary, 284, 473
of Poppy, 528
Ovule, amphitropous, 287
anatropous, 287, 473

INDEX

Ovule, campylotropous, 287
forms of, 287
287
Ovules, 285, 448, 473
Ovum, 69, 450, 474

Oxalic acid, production by molds, 394

Oxidizing enzymes, 104
Oxytropism, 34

Pachystima, 552, 553
Padus, 535
Palaguium, 581, 582

Panax quinquefolium, 576
repens, 576
Pandanus, roots of, 161
Pandorina, 338
Panicle, 258, 259
Pansy, 566
Papain, 103, 567
Papaver, 524
calyx of, 267
latex of, 101
placentation in, 288
Papaver Rhoras, 528
Papaver somniferum, 101, 627, 628
Papaveracea, 524
Papaverales, 524
Papaverine Hydrochloridum, 527
Papaveris Fructus, 528
Papaw, 567
Papaw Family, 566
Papayin, 103
Papilionacea, 537, 638

Papillz, 124
Pappus, 266, 297
Paprika, 598
Papyrus, 618
Paraguay Tea, 85
Paraphyses, 356, 387, 388, 427
Parasite, 362
Parasites, facultative, 367
obligate, 367
Pareira, 162, 521
Parenchyma, 116
assimilation, 116
border, 223, 224
conducting, 117
ordinary, 116
palisade, 117, 222 |
phloem, 117, 135
reserve, 117
sieve, 135
spongy, 117, 223
wood, 117
Parietales, 562
Parmelia, 414
Parmelia perlata, Fig. of, 415
Parsley Family, 118, 148, 243, 576
Partridge Berry, 190, 605
Pasque Flower, 518
Passiflora, 266, 566
incarnata, 566, 567
Passifloracee, 566
Passion Flower, 188, 567
Passion Flower Family, 566
Pasteur’s Solution, 382
Pathogens, 364
Pathology, 3
Paullinia, 553, 554
Paullinia Cupana, 554
Paulownia, 599
Paulownia imperialis, 599
Payena, 101, 582
Pea, 82, 240
tendrils of, 189
Pea Family, 536
Peach, 301, 305, 534, 535
Peachwood, 540
Peanut, 301
Pear, 535
fruit of, 305
pericarp of, 298

INDEX 775

Pear, stone cells of, 119, 120, 301

Peat mosses, 425

Pecan, 504

Pectase, 102

Pectinase, 102

Pectose, 23

Pedaliacea, 601

Pedicel, 256

Pedicularis, 599

Peduncle, 256, 262

Pelargonium, 301, 541
capitatum, 542
odoratissimum, 542
Radula, 542

Pellaa, 435

Pelletierinz Tannas, 571

Pengawar Djambi, 435

Penicillium, 388
brevicaule, 391
camemberti, 390, 391
expansum, 391
glaucum, 388
roqueforti, 389, 390

Pentosides, 84

Pentstemon, 600
stamens of, 275

Peony, 518
Fig. of flower, 284

Pepo, 305, 608

Pepper, Black, 499
Red, 597

Pepper Family, 499

Peppermint, 594

Pepsin, 103

Perennial plants, 163

Pereskia, 567, 568

Perianth, 262, 266
forms of, 270
outgrowths of, 274

Periblem, 113

Pericambium, 56, 131

Pericarp, 297

Pericladium, 243, 576

Pericycle, 56, 63, 131

Pericyclic fibers, 122, 131

Periderm, defined, 207

Peridiolum, 411

Peridium, 401, 410

776 INDEX
Perigone, 266 Phloem, secondary, 143
appendages of, 274 Phloroglucin, solution of, 685
Perisperm, 314 Phlox, corolla of, 272
Peristome, 429 Phoradendron flavescens, 510
Perithecia, 391, 395, 416 Photogens, 364
Periwinkle, 258 Photosynthesis, 64, 66
Peroxidase, 104 defined, 224
Pertusaria, 414 Phototropism, 35
Petals, 269 Phragmidiothrix, 372
Petiole, 223, 242 Phragmites, 618
gross structure and histology of, 242 Phycocyanin, 100, 334, 357
Petiolule, 234 Phycoerythrin, 100, 335, 357
Petroleum Ether, 685 Phycophain, 100
Petroselini Radix, 577 Phycomycetes, 375, 376
Petroselinum, 577 Phycophzin, 351, 352
Petroselinum sativum, 577 Phycoxanthin, 351
Petunia, 596 Phylloclades, 188
Peumus Boldus, 522 Phyllode, 244
Peyote, 568 Phyllotaxy, 229
Peziza, 387 Physalis, 297
repanda, Fig of, 386 Fig. of calyx, 267
Pezizales, 387 Physica, 414
Pfeffer, 33 Physica stellaris, Pig. of 413
Phzophycez, 351 Physiological division of labor, 16
Phzosporales, 352 Physiology, defined, 2
Phalaris canariensis, 485 Physostigma, 539, 540
Phallales, 411 style of, 289
Phanerogamia, 325, 443 Physostigma venenosum, 640, 641
Phanerogams, 14 Physostigmine: Salicylas, 540
Pharmaceutical Botany, defined, 3 Phytogeography, defined, 3
Pharmacognosy, defined, 3 Phytopalzontology, defined, 3
Phaseolus lunatus, structure of seed of, 319 Phytoglobulins, 94
Phaseolus multiflorus, cross-sections of Phytolacca, 514
hooked hair from, 126 Phytolacca americana, 514, 515
Phegopteris, 435 Phytolacca, N.F., 514
Phellem, 207 Phytolaccacea, 514
’ Phelloderm, defined, 208 Phytopalzontology, defined, 3
Phellogen, 114, 129, 131 Phytopathology, defined, 3
Phenotype, 647, 648 Picea, 465
Philadelphus, 531 canadensis, 466
Phloem, 140 excelsa, 466
defined, 143 mariana, 466
internal, 204 rubra, 466
interxylary, 204 Pickerel weed, 116
intraxylary, 204 Picrasma excelsa, 546, 647
parenchyma, 117, 135 Picric acid, 39
primary, 143 Pigments, 98
ray, 139 Pigweed, pollen of, 283

INDEX

Pileus, 404
Piliferous layer, 152
Pilocarpinzee Hydrochloridum, 546
Nitras, 546
Pilocarpus, 546
Pilocarpus, 545
Pilocarpus Faborandi, 546
microphyllus, 546
neighboring-cells of leaf epidermis,
252
Pimenta, 574
Pimenta acris, 574
officinalis, 574
Pimiento, 598
Pimpinella, 577
Pimpinella Anisum, 576
magna, 577
Saxifraga, 577
Pinaceae, 465
Pine, 465
Austrian, 250, 447, 461, 453
Drops, 580
Family, 465
inflorescences of, 464
leaf of, 446
pollen of, 282
Scotch, 448
stem, 445
tar, 465
White, 444
wood of, 212
Pineapple, enzyme in, 103
fruit of, 307
Pinenes, 97
Pinguicula, 640
vulgaris, 640, 641
Pinites succinifer, 466
Pink Family, 273, 514
Pink Root, 583, 585
Pinna, 55, 433
Pinnularia, Figs. of, 349, 352
Pinnule, 55, 433
Pinus, 465
austriaca, 447
maritima, 466
montana, 466
nigra, leaf of, 250
palustris, 465
Strobus, 401, 444, 465

777

Pinus, sylvestris, 448

Pinus Alba, N.F., 465

Pinweeds, 509

Piper, 305, 499

Piper angustifolium, 499
Cubeba, 498, 499
longum, 499
methysticum, 499
nigrum, 498, 499
officinarum, 499

Piperacea, 499
formation of endosperm and peri-

sperm in, 314

Piperales, 499

Pipsissewa, 579

Piscidia, 540

Piscidia Erythrina, 540

Pistachio, 552

Pistacia vera, 552
Lentiscus, 552

Pistil, 285, 473
bicarpellary, 285
compound, 285
monocarpellary, 285
polycarpellary, 285
system, 283
tricarpellary, 285

Pitcher plants, 629

Pith, 165, 166

Pits, 108
bordered, 136

Pix Burgundica, 466
Canadensis, 466
Juniperi, 466
Pini, 465

Placenta, defined, 287

Placentation, 287, 288

Plankton, 617

Planococcus, 371

Planosarcina, 371

Plant, acaulescent, 186
anemophilous, 291
annual, 162
aquatic, 164
associations, 616
axis, 6
biennial, 163 —
breeding, 3
caulescent, 186

778

Plant, cell, 41
cigar, 570
communities, 616
dicecious, 264

pitcher, 629
polygamous, 264
seed, 7
tissues, 111
xerophytic, 164
Plant Succession, 621
Plantaginacea, placentation in, 288
Plantaginis Semen, 603
Plantago, 44, 301, 603
Plantago arenaria, 603
Plantago lanceolata, 603
Plantago ovata, 603
Plantago Psyllium, 602, 603
Plantain, 259, 603
Plants, aquatic, 164, 617
carnivorous, 621
classification of, 323
desert, 617
general characteristics of, 6
groups of, 12, 328
intermediate, 617
marsh, 618
naming of, 323, 329
relation of, and animals, 29
relation to animals, 621
salt, 617
water, 617
Plasma membranes, 19, 22

illustrating, successive stages of, 154
Plastids, 42
Platystemon, 524

Plum, 305, 535
Plumule, 311
Pod, 301
Podophyllum, 519
Podophyllum emodi, 519
peltatum, 519
Fig. of, 521
petiolar structure of, 245
Podophyllum Indicum, 519
Poison Hemlock, 516
Fig. of, 578
Poisons, 33
Poke, Family, 514
Root, 175, 614
Polemoniales, 590
Pollen, 278
chamber, 449, 458
grain, 280
mother cells, 278, 474
stages in germination, 293
Pollen grains, 282, 457
Fig. of, 281, 450
pine, 281, 450
Pollens, Hay-fever, 283
Pollination, 290
cross, 291
in erythronium, 475
in pine, 450
self, 290
Pollinia, 282, 283, 589
Polycotyledony, 227
Polyembryony, 479
Polygala, 103, 283
lutea, 549
pollen of, 283

INDEX

Polygala Senega, 549, 550
Polygalacea, 548
Polygonacea, 246, 314, 512
Polygonales, 512
_ Polygonum, 512
Bistorta, 513
Polypeptides, 93
Polypetala, 499
Polyploidy, 658
Polypodiacee, 435
Polypodium, 434, 435
Polyporacea, 404
Polyporus officinalis, 404
Polystichum, 435
Polytrichum commune, Fig. of, 427
life history of, 426
Pome, 305
of apple, Fig. of, 306
Pomegranate Family, 570
Ponteria, 582
Poplar, 500
Poppy, calyx of, 267, 268
Family, 524 i
fruit of, 299, 302°
leaf of, 232
Populi Gemma, 500
Populus balsamifera, 500
candicans, 500
tremuloides, 500
Pore Fungi, 404
Porella, 418, 419
Pores, 108, 110
bordered, 136
germ, 282
Portulaca, 301, 514
Postelsia, 354
Potash, solutions of, 685
Potassium ferrocyanide solution, 685
Potato, 87, 192, 272, 596
starch of, 79, 81
Potentilla, 269, 535
silvestris, 536
Prefloration, 264
Prefoliation, 230
Premna arborea, 593
Prickles, 189
Prickly Ash, 546
Prickly Pear, 567

779

Primary Meristems, 113
- of root, 157

Primordial meristem, 111

of leaf, 228
Primulacea, placentation in, 288
Principes, 485
Prinos, 552
Privet, 261, 683

California, 170
Procambium, 113
Prodromus, 326
Proembryo, 348
Promycelium, 397, 398, 401
Prophases, in nuclear division, 48, 49
Proplastids, 42
Prosopis juliflora, 540
Prostele, 145
Proteacea, 254
Protective tissues, 151
Proteinaceous enzymes, 103
Proteinoplastids, 133
Proteins, 92

formation of, 65

tests for, 93
Prothallia, female, 438

Fig. of fern, 69

male, 438
Prothallial cushion, 69
Prothallium, 68

Fig. of fern,-70
Prothallus, 68, 451, 460
Protista, 30
Protoascales, 381
Protococcales, 337
Protoderm, 113
Protonema, 68, 426
Protopectin, 107
Protopectinase, 102
Protophloem, 144
Protoplasm, 10, 25, 30

properties of, 30, 31
Protoplasmic cell contents, 40
Protoplasmic movement, 39, 40
Protoplast, 17
Protoxylem, 143, 168, 169, 196
Protozoa, 24
Prune, 305
Prunum, 536

780 INDEX
Prunus armeniaca, 534 Putrefaction, 364
Avium, 534 Pycnanthemum montanum, 59%
Cerasus, 95, 535 Pycnia, 401
Fig. of flower, 270 Pyrenoid, 337, 341
domestica, 536 Pyrenomycetales, 395
Laurocerasus, 536 Pyrethri Flores, 615
serotina, 534 Pyrethrum, 615
Prunus Virginiana, 535 Pyrogallol, 507
Psalliota campestris, 405 Pyrola, 580
Pseudobulbs, 188 Pyrus, 535
Pseudocarp, 298 Pyxidium, 301
Pseudomonas, 372 Pyxis, 301
Pseudomonas radicicola, 176
Fig. of, 178 Quassia, 547
Pseudopodia, 26, 374 Quassia amara, 546, 647
Pseudotsuga, 465 Quercus, 507
Pseudotsuga taxifolia, 466 Quercus alba, 506, 507
Psyllium seeds, 95, 602 chrysolepis, 506
_ Ptelea, 545, 546 infectoria, 507
Ptelea trifoliata, 546 Fig. of, 505
Pteridophyta, 13, 431 Michauxii, 506
Pteridophytes, 13, 431 occidentalis, 506
Pteris aquilina, 433 pedunculata, 15
cross section of rhizome, 435 platanoides, 506
Pteris, Fig. of leaf of, 434 prinus, 506
Fig. showing development of em- Suber, 506
bryo, 72 velutina, 506
Pterocarpus marsupium, 540 virginiana, 506
santalinus, 539 Quillaja, 535
Pterospora, 580 Quillaja Saponaria, 88, 634, 535
Puccinia Graminis, 399 Quillworts, 431, 442
Fig. of, 400 Quince, 535
Puccinia malvacearum, 401 calyx of, 267
Puff Balls, 409 fruit of, 305
Pulmonaria officinalis, 593 pericarp of, 298
Pulsatilla, 518 receptacle of, 297
Pulsatilla patens, 519 Quinidinz Sulfas, 605
pratensis, 519 Quinina, 605
vulgaris, 519
Pulvinus, 243 Raceme, 258
Pumpkin, 607 Rachilla, 259, 484
Punica, 571 Rachis, 233, 256
Granatum, 571 _ Radial bundles, 142
Punicacea, 571 Radicle, 311, 317, 321
Purging Cassia, 332 Radish, 161, 162
Purified Siliceous Earth, 350 Ragweeds, pollen of, 283
Purple Cone-flower, 612 Ranales, 515
Putamen, 298 Ranunculacea, 243, 515, 618

INDEX

Ranunculus, 248, 518
Raphe, 287
Raphides, 89, 90
Raspberry, 189
fruit of, 300, 306
stems of, 186
Ray, J., 325
Rays, medullary, 139
vascular, 139
Reagents, 680
Receptacle, 262, 265
Receptacles of Fucus, 356, 356, 367
of Marchantia, 421
Red Algz, 95, 357
Red Gum, 90
Red Squill, 490
Reductase, 103
Reduction division, 51
Redwood, 465
Regma, 301
Reindeer Moss, 417
Rejuvenescence, 45
Remijia pedunculata, 605
purdicana, 605
Reproduction, asexual, 44, 45, 378
defined, 38, 44
sexual, 44
Reserve cellulose, 109, 487
parenchyma, 117
starch, 76
Resin duct, Fig. of, 150
Resina, 465
Resinogenous layer, 96
Resins, 97, 98
Respiration, defined, 66, 225 ~
distinction from photosynthesis, 66
Rhamnacea, 554
Rhamnales, 554
Rhamnase, 103
Rhamnose, 103
Rhamnus cathartica, 556
Rhamnus Frangula, 103, 555, 556
Rhamnus Purshiana, 554, 556
Rheotropism, 34
Rheum, 512
officinale, 513
palmatum, 512, 513
Rhinanthus, 599
Rhizoclonium, Fig. of, 317

781

Rhizoids, 65, 347, 420, 426.
Rhizome, defined, 192
histology of monocotyl, 215
Rhizophora, 619
Rhizophoracea, 569
Rhizopus, 378
nigricans, 377
strains of, 380
Rhododendron, 580
Rhodophycez, 357
Rhodymenia palmata, 361
Rhoeadales, 524
Rhubarb, 513
Rhus, 92, 305, 552
Cotinus, 551
glabra, 551, 552
hairs of, 124, 125
japonica, 552
semialata, 552
succedeana, 96
sylvestris, 96
toxicodendron, 551, 552
typhina, 552
venenata, 551
vernicifera, 96
Rhytidome, 130
Ribbon Bush, 512
Riccia, 418, 419, 617
Ricciocarpus, 617
Rice starch, 79, 80
Ricinus communis, 256, 312, 651
Rivularia, 335, 336
Robinia, 213
Roccella tinctoria, 416
Rock Rose Family, 567
Rockweeds, 354
Root, adventitious, 160
branching, 176, 177
California Privet, 170
cap, 158
classification as to form, 161
classification as to order, 160
conical, 162
consistency, 162
defined, 152
distinction from stem, 160
duration of, 162
epiphytic, 161
fibrous, 160, 162

782

Root, functions, 6, 152
fusiform, 161
generative tissues of, 158
growing point of, 158
hairs, 152, 154
histology, 163-176
lateral, 160
main, 160
moniliform, 162
napiform, 162 —
Phaseolus, 167
primary, 142, 160, 319
regions of, 158
secondary, 160, 177, 319
Smilax, 165
soil and water relation of, 152
systems, 164, 177
tap, 160
texture, 162
tortuous, 162
tubercles, 176
Rootlet, 152
Roots, 160
abnormal structure in, 175
brace, 161
Dicotyledon, 167
fascicled, 162
fleshy, 162
Monocotyledon, 165
multiple, 162
napiform, 162
nodose, 162
nodule producing, 176
of parasitic plants, 161
of primary growth, 167
of secondary growth, 169
prop, 161, 319
secondary, 160
tuberous, 162
woody, 162
Roridula, 530
Rosa, 535
Rosa canina, 536
centifolia, 536
damascena, 535
gallica, 535
Rosacea, 245, 534
Rosales, 531

INDEX

Rose, 535

Family, 534

fruit of, 304
Rosemary, corolla of, 272
Rose of Sharon, 560
Rosette aggregates, 89, 90
Rosin, 465
Rosmarinus officinalis, 595, 596
Rotation, crop, 179

Rubus Idzus, 535
Rubus Idaeus, 535
strigosus, 536
Rue, 286, 545, 646
Rue Family, 545
Ruellia, 602, 603
Ruellia ciliosa, 603
Rumex, 612, 613
acetosella, 512
crispus, 513
obtusifolius, 513
pollen of, 283
Runner, defined, 189
Ruppia, 291
Ruscus, 188
Russian Thistle, pollen of, 283
Rust, 399
Cedar, 402
Hollyhock, 401
Wheat, 399
White Pine Blister, 401
Ruta, 545
Ruta graveolens, 646, 546
Rutacea, 545
Rutea, 545
Ruthenium Red and Lead Acetate
Solution, 686 =
Rye, 259
starch of, 78, 79

Sabadilla, 490
Sabal, 305, 487

Sabbatia, 587
Sabbatia angularis, 587
Sabina, 466
Sac Fungi, 381
Saccharomyces, 381
apiculatus, 386
cerevisia, 382
ellipsoideus, 385, 386
membranifaciens, 386
pastorianus, 387
Saccharomycetacea, 381, 382
Saccharum, 484
officinarum, 485
Saddle Fungi, 394
Safflower, 615
Saffron, 492
Safranin, solutions of, 686
and Aniline Blue, 690
and Fast Green, 689
and Light Green, 689
and Methyl Green, 691
Sage, 594, 595
Sage, corolla of, 272
Sago starch, 456
Salep, 94, 496
Salicacee, 499
Salicales, 499
Salicin, 83
micro-chemic test for, 83
Salicinum, 500
Salicornia ambigua, 619
Salix, 500
Salix alba, 500
nigra, 500
retusa, 499
viminalis, 501
Salmonella, 373
Salsola, 283
Salvia, 595
Salvia officinalis, 595
Salvinia, 437
Salviniacea, 436
Samara, 303
Sambucus, 606, 607
Sambucus canadensis, 607
nigra, 607
Samphire, 619
Sandalwood Family, 510
Sandaraca, 466

INDEX 783

Sandbox, 551
Sanguinaria, 528
latex of, 101
Sanguinaria canadensis, 626, 628
Sansevieria cylindrica, leaf of, 250
Santalacee, 510
Santalales, 510
Santalum album, 510
Santalum Rubrum, 539
Santonica, 615
Santonium, 615
Sap, cell, 18
crude, 117
elaborated, 117
nuclear, 18
vacuoles, 19
Sapindaceea, 553
Sapindales, 551
Sapindus, 553 —
Saponaria, 84
Saponins, 84
micro-chemic test for, 84
Sapotacez Family, 581
Sappan, 540
Saprogens, 364
Saprophyte, 362, 383
facultative, 367
obligate, 367
Sap-wood, 213
Sarcina, 368, 371
Sarcode, 17
Sarcocarp, 298
Sargassum, 355
Sarracenia, 289, 629, 638
Drummondii, 638
flava, Fig. of, 632, 638
purpurea, 629, 630, 631, 638
rubra, 638
Sledgei, 638
style of, 289
Sarraceniacea, 530
Sarraceniales, 530
Sarsaparilla, 490
Bristly, 575
tendrils of, 189
Virginian, 575
Wild, 575
Sassafras, 523
Medulla, 524

INDEX

Sectioning, in paraffine, 677, 697
in pith, 675

Sections, celloidin, 700, 701
making of, 675
paraffine, 697
radial-longitudinal, 677
surface, 677
tangential-longitudinal, 677
transverse, 677

Sedges, 304, 483, 484

Sedum, leaf of, 250

Seed, 311
albuminous, 314
appendages, 313
Castor-oil, 312
coats, 312
defined, 311
Dicotyl, 319
exalbuminous, 314
functions of, 8, 311
germination, 318, 322, 453
histology, 315
Indian Corn, 315

Monocotyl, 315
structure, 312
vessel, 297
vitality of, 311
Seed-grass, 618
Seedling, 319
of Indian corn, 316, 319

_ Sepals, 266
epigynous, 269
hypogynous, 269
perigynous, 269
Sequoia, 465
height of, 186
Serenoa serrulata, 487
Serpentaria, 512
Serpyllum, 596
Service Berry, 535
Sesame Family, 601
Sesamum indicum, 602
Sesuvium maritimum, 514
Shad Bush, 535
Shell bark, 130
Shepherd’s Purse, 302
Shoots, 180, 247
fertile, 437
spur, 446
sterile, 437
Shrub, 191, 582
Sideritis, 596
Sieve, 133
callus of plates, 109
plates, 133
Sieve tubes, 133, 134
stages in development of, 134
Silica, 91
bodies, 91
Siliceous Earths, 350
Silicule, 302
Siliqua, 302
Silphium, 615
Simaruba, 547
Simaruba amara, 547
officinalis, 547
Simarubacea, 546
Sinapis alba, 530
Sinapis Nigra U.S.P., 530
Sinigrin, 103
Siphon Algz, 346
Siphonales, 346, 376
Siphonostele, 145, 199
Sitotaxy, 34
Sitotropism, 34
Skunk Cabbage, 487
Slime Molds, 362, 374
Fig. of, 353
Response to water, 34

INDEX 785

Slime plugs, 134

Slippery Elm, leaf of, 221, 241

Smilax, 490
histology of root, 165, 166
medica, 490
modified stipules of, 246
officinalis, 489, 490
ornata, 490

Smuts, 397

Snapdragon, 272, 600
stamens of, 275

Snowberry, 606

Soap Bark, 84

Soapwort Family, 553

Soft bast, 202

Saja max, 540

Solanaceae, 596
development of stomata in, 254
placentation in, 288

Solanee, anther dehiscence in, 279

Solanine, 87

Solanum, 598

Solanum, 596

Solanum carolinense, 87, 598
Dulcamara, 87, 596, 598, 599
nigrum, 87
tuberosum, 87, 203

Solidago odora, 615

Solomon’s Seal, 192

Sorbinose, 74

Sorbus, 535

Soredia, 416

Sorghum, 484

Sorosis, defined, 307

Sorus, 433

Sour Gum, 575, 577

Soy Bean, 540

Spadix, 259

Sparganium, 618

Spathe, 259

Spearmint, 595

Species, defined, 326

Species Plantarum, 325

Specific epithet, 330

Specific name, 329

Spermacia, 401

Spermagonia, 401

Spermatium, 358

Spermatocyte, 68

786

Spermatophyta, 14, 443
Spermatophytes, 14, 15, 443
Spermatozoid of fern, 68
Spermoderm, 312
Sperms, 16
Sphagnacee, 425
Sphagnales, 425
Sphagnum, 425
acutifolium, 424
palustre, 425
squarrosum, 424
Spharella, 338
Spicebush, 523
Family, 522
Spiderwort, 36, 40
Spigelia, 583, 585
Spigelia marilandica, 584, 585
Spike, 259
Spikelet, diagram of typical, 484
Spikelets, 259, 484
Spikenard, 575
Spinach, 514
Spine, defined, 189
Spirea, 535
Spireme, 47
Spirilla, types of, 371
Spirillacea, 372
Spirillum, 365, 372
Spirochzta, 348
Spirogyra, 343, 344
Spirosoma, 372
Spirulina, 335
Spondiea, 552
Spongy parenchyma, 117, 223
Sporangiophores, 377
Sporangium, 67, 353, 377, 379
Spore, 68
bacterial, 368, 369
brand, 398
daughter-cells, 280
germination of, 346
mother-cells, 67, 280, 447, 457
print, 405
swarm, 375
Sporogonium, 428
Sporophore, 402, 404, 406
Sporophyll, 433
Sporophyte, 53, 418, 431
Sporulation, 368

INDEX

Spring Beauty, 514
Spruce, 211, 465
Spruce Gum, 466
Spur shoot, 446
Spurge Family, 550
Spurs, sepaline, 268
Squash, 188
Squill, 192, 490
St. John’s Wort, 277
St. John’s Wort Family, 564
Staff Tree Family, 552
Stain,
Delafield’s hamatoxylin, 701
Fast Green, 682
Gentian violet, 683
Heidenhain’s hamatoxylin, 683
Ruthenium Red and Lead Acetate,
686
Safranin, 686
Staining, 688
double, 688, 689, 690, 691
of celloidin sections, 701
of material in paraffin ribbons, 699
Stains, 680
Stamen System, 274
Figs. of, 276
Stamens, color of, 277

INDEX 787

Star Anise, 516 Stems, decumbent, 187

Star Apple Family, 581 duration of, 187

Starch, 75 elongation of, 187
assimilation, 76 endogenous, 193, 194
Barley, 78, 81 erect, 186

Bean, 80, 82

Buckwheat, 81

Canna, 82

Cassava, 80, 81

characteristics of commercial kinds,
79-82

compound grains, 77

Corn, 79, 80

corrosion, 102

fill, 77

formation of, 64

Maranta, 78, 79

Pea, 80, 82

Potato, 78, 79

reserve, 76, 77

Rice, 79, 80

Rye, 78, 79

Sago, 78, 82, 456

simple grains, 77

structure and composition of, 77

transitory, 76

Wheat, 78, 79

exceptional types of dicotyl, 203
exfoliating, 187
exogenous, 193
flexuous, 188
forms of, 187
fruticose, 190
herbaceous, 130, 189
histology of aconite, 130
of annual dicotyl, 194
of herbaceous dicotyledon, 9,
130, 203
of herbaceous monocotyl, 215
of perennial dicotyl, 199
of woody monocotyl, 217
obconical, 188
perennial, 187
phylloid, 188
procumbent, 186
quadrangular, 187
reclining, 186
repent, 187
scandent, 188

Steckbeck, 36 shrubby, 190
Stegmata, 91 size of, 186

Stele, 144 spiny, 188

Stem, defined, 186 subcylindrical, 187

direction of growth, 186
distinction from root, 160
elongation, 187
functions, 6, 186
generative tissues of, 187
modification of, 188
Sunflower, 194

Stemonitis fusca, 375.
Stems, above-ground, 188

aerial, 188
aerial-tuberous, 188
alate, 187

annual, 187
ascending, 186
cactoid, 188
climbing, 188
compressed, 187
cylindraceous, 187

subterranean, 192
subterranean tuberous, 188
suffruticose, 189
tendriliform, 188
terete, 187
tortuous, 188
triquetrous, 187
truncate, 188
twining, 188
underground, 192
Sterculiacea, 557
Sterigmata, 390, 397
Sterigmatocystis, 391
Stevens, 104
Sticta, 414
Stigma, 290, 473
Stillingia, 551
Stillingia sylvatica, 551

788 INDEX
Stimuli, extrinsic, 32 Strophanthus hispidus, 83, 688, 689
intrinsic, 32 Kombé, 83, 689
Stink horn fungi, 411 Strophiole, 313
Stipe, 55, 404 Strychnina, 585
Stipule, defined, 223 Strychnine, 85
Stipules, 223, 245 micro-chemic test for, 85
axillary, 245 Strychnos Ignatii, 85, 685
caducous scaly, 245 Nux Vomica, 85, 204, 583, 686
connate, 245, 246 Fig. of, 584
free axillary, 245 toxifera, 585
free lateral, 245 Style, 288, 473
lateral, 245 arms, 288, 289
lateral-adnate, 245 collecting hairs of, 289
lateral-connate, 245 column, 289
lateral-interpetiolar, 245 Styloids, 89, 90
modified, 246 Styracea, 582 .
sepaline, 269 Styrax, 532
Stolon, defined, 189 Styrax Benzoin, 583
Stoma, 63, 251 tonkinensis, 583 4
Stoma Mother-cell, 253 Subdivision, 327
Stomata, 123, 222 Suberin, 24, 107 ‘
structure and development of, 251 Succinum, 466
transpiration, 124 Succus Pomorum, 536
water, 124 Suckers, 185, 193 d
Stomatal apparatus, 251 Sucrose, 76
Fig. of types of, 254 Sucrosum, 485, 614
Stomatal development, 251 Sugar cane, 75
Stone cells, 119 Sugars, 74 f
Stone Root, 596 Sulfuric acid, as reagent, 687
Stoneworts, 346 Sumac Family, 551
Storax, 532 Sumbul, 576
Stramonium, 50, 163, 597 Summer Cypress, 514 :
Strawberry, 300, 534, 535 Summer Savory, 596
Fig. of flower, 268 Sundew, 530 a
Fig. of fruit, 299 Sundews, 260, 530, 622
Fig. of plant showing propagation, Sunflower, 614
86 Figs. of cross sections of stem, 194,
stem of, 186 195
tree, 579 histology of stem of, 194
Streptococcus, 386, 371 Surirella, 352 —
_ erysipelatis, 373 Suspensor, 439, 452
scarlatina, 373 Sutures, 298
Streptothrix, 372 Swamproot, 615
Strobile, 259, 307 Swarm spores, 375 . :
Strobili, 457, 464 Sweet Beall, 596 = e -
Stromata, 395 Sweet Birch, 504,505 __ a
Strophanthin, 83, 589 Sweet Cicely, 576
micro-chemic test for, 83 Sweet Flag, 618 :
Strophanthus, 313, 589 jor

Sweet Potato, 162, 590
Sweet Woodruff, 605
Sweetia panamensis, 539
Swertia Chirayita, 587
Syconium, 307

Fig. of, 290
Sympetalae, 579
Symphytum, 593
Symphytum officinale, 593
Symplocarpus fetidus, 487
Syncarp, 471
Synedra ulna, Fig. of, 352
Synergids, 292, 474
Syngenesis, 265
Systema Naturz, 325
Systematic Botany, 323

defined, 2

Tabacum, 598
Tabellaria vulgaris, Fig. of, 352
Tamarack bark, 466
Tamarind, 301
Tamarindus, 539, 540
Tamarindus indica, 540
Tanacetum, 615
Tanacetum vulgare, 615
Tannins, 92
Tansy, 615
Tapetum, 67, 278, 280
Tapioca, 551
Taraktogenos Kurzii, 565
Taraxacum, 615
Taxacea, 465, 466
Taxies, 32
Taxodium, 619 -
Taxonomy, 323
defined, 2
Taxus baccata, 467
canadensis, 467
Tea, 85, 563
Tea Family, 563
Teak-wood, 593
Teasel Family, 608
Tecoma radicans, Fig. of seed, 313
Tectona, 593
Tegmen, 312
Tela contexta, 407
Teleutospores, 401
Teliospores, 400, 401

INDEX 789

Telium, 401
Telophase, 50
Temporary mount, technique of mak-
ing a, 687
Tendril, defined, 189
Tendrils, 189
Terebinthina, 456
' Canadensis, 466
Laricis, 466
Ternstremiaceea, 563
Terpenes, 97
Terra Silicea Purificata, 350
Testa, 312
Tetrads, 67, 280, 474
Tetrasporangia, 358
Tetraspores, 358, 359
Thalamus, 262, 265
Thallophyta, 12, 333
Thallophytes, 12, 333
Thallus, defined, 12
Thamnidium, 381
Thea, 563
Thea sinensis, 563
Theacea, 563
Thelephorales, 403
Theobroma, 559
Theobroma Cacao, 559
Fig. of, 558
Theophrastus, 1, 323
Thermogens, 364
Thermotropism, 32
Thigmotropism, 36
Thiothrix, 373
Thistle, 610, 611
Thorn, defined, 189
Throat, 266
Thuja, 465, 466
occidentalis, 466
Thyme, 189, 595
Thymelaacea, 569
Thymus Serpyllum, 596
vulgaris, 595
Thyrsus, 261
Tilia, 559
americana, 559
Tiliacea, 559
Tissue, 111
assimilation parenchyma, 116
cambium, 114

790 INDEX

Tissue, chlorenchyma, 116 Tracheary tissue, 135
collenchyma, 113, 117 Tracheids, 135, 196, 139
conducting, 151 Fiber, 214
conducting parenchyma, 117 Tradescantia, 82, 253
cork, 129 Tragacanth, 95
cribriform, 133 Tragacantha, 539
embryonic, 114 Trailing arbutus, 272, 681
endodermal, 127 Trama, 407
epidermal, 122 Transpiration, 65, 226
fundamental, 116 Stomata, 124
generative, 113 Stream, 226
ground, 116 Trape natans, 569
laticiferous, 131 Trapacea, 569
mechanical, 151 Tree, defined, 190
medullary ray, 139 Trees, branching in, 191
meristematic, 113, 114, 151 Trehalase, 102
ordinary parenchyma, 113, 116 Trehalose, 75, 102
parenchyma, 116 Tremellacea, 402
pericycle, 131 Tremellales, 402
protective, 151 Treponema, 373
reserve parenchyma, 116 Treviranus, 17
sclerenchyma, 113, 118 Tribes, 329
secreting, 151 Trichogyne, 358
sieve, 133 Trichomes, 124
stony, 118 Trichophyton, 412
storage, 151 Trifolium, 540
suberous, 129, 130 Trifolium pratense, 540
tracheary, 135 repens, 536

Tissues, 8, 111 ; Trigonella Fanum-gracum, 540
adult, 113 Trilisa odoratissima, 615
classifications of, 111, 151 Trillium, 490
complex, 113 Trillium erectum, 490
primary, 111, 115 Trimorphism, 289 _
secondary, 112 Triticum, 485
simple, 113 Triticum sativum, starch of, 79

Tobacco, 596, 597 Tropaolum, 267

Tolu, 539 Tropisms, 32 vi

Toluifera Balsamum, 539 _ Tropophytes, 617, 619

: Pereira, 539 True Fungi, 375 a

Tomato, 304, 596 True Mosses, 426

Tonga, 487, 593 Truffles, 394 :

Tooth Fungi, 404 Trumpet Creeper, 291, 313 _

Tormentilla, 536 Trunk, defined, 190° a

Torulacea, 381 Trypsin, 103

Torus, 262, 265 Tschirch, 96

Touranose, 75 Tsuga, 465

Tournefort, 325 Tsuga canadensis, 466

Toxins, 33 Tuber, defined, 192

Trachez, 135 Tuber, genus, 394

Tuberales, 394

Tubercles, 176, 179

Tubiflore, 590

Tubuliflore, 613

Tulip, 260, 266

Tulip Poplar, 516

Tung Oil, 551

Turgor, 155

Turnera aphrodisiaca, 566
diffusa, 566

Turneracee, 566

Turnip, 162, 240

Turpentine, 465
Bordeaux, 466

Turpeth Root, 591

Tussilago Farfara, 612, 615

Twig, 190

Twin crystals, 90

Tyloses, 213

Typha, 282

Ulmacee, 507
Ulmus, 507
Ulmus fulva, 221, 223, 506, 507
Ulothrix zonata, 341
Fig. of, 338
Ulva lactuca, 342
Umbel, 258
compound, 258
Umbbellales, 575
Umbellifera, 243, 266, 303, 576
Umbelliflorz, 575
Unit characters, 643, 644
Uredinales, 399
Uredinium, 400
Uredo linearis, 401
Uredospore, 400
Urginea maritima, 490
Urtica urens, stinging hair of, 126
Urticales, 507
Usnea, 414, 417
Ustilaginales, 397
Ustilago Maydis, 397
Ustilago Zea, 397, 398
Utricle, defined, 304
Utricularia, 248, 638, 639
Utriculariacee, 640
Uva, defined, 304

INDEX

Uva Ursi, 581
corolla of, 273

Vaccinea, 581

Vaccinium, 581

Vaccinium corymbosum, 581

Vacuole, contractile, 27
food, 26

Vacuoles, sap, 19, 40

Valerian Family, 608

Valeriana, 609

Valeriana officinalis, 608, 609

Valerianacea, 608

Valerianales, 608

Vallisneria, 291

Valves of fruits, 298

Vanilla, 496

Vanilla planifolia, 496
Figs. of plant, 495
histology of fruit, 307
roots of, 161

Vanilla Leaf, 615

Variations, 652

Varieties, 327
naming of, 331

Variety epithet, 331

Vascular bundles, 140

Vascular rays, 199

Vaucheria, 95, 346

Vaucheria sessilis, Fig. of, 345

Vaucheria terrestris, Fig. of, 341

Vegetable Kingdom, 1

Vegetative Multiplication, 46

Veil, partial, 406
universal, 408

Veins, 222, 230

Velamen, 161

Venation, leaf, 230

Venter, 69, 70

Venus fly-trap, 36, 625
Figs. of, 35, 626

Veratrina, 491

Veratrum viride, 490

Verbasci Flores, 600

- Verbasci Folia, 600

Verbascum, 275, 599
Blattaria, 599
phlomoides, 600

91

INDEX

Water Leaf Family, 591

Water Lily, 248, 617
Family, 515

Water Molds, 376

Viola tricolor, 124, 126, 666
Violacea, 565
Violet, 189, 566
Violet Family, 565
Viscum, 479, 510
Viscum album, 510
Vitacea, 556
Vitamins, 105
Vitis, 556
Vitis Labrusca, 557
Vitte, 148
Volatile oils, 96
Volva, 409 :
- Volvocales, 337 |
Volvox, 29
Volvox globator, life history or —
Fig. of, 340
Von Mohl, H., 17
Vouacapoua Araroba, 540

Wafer Ash, 303, 545, 646
Wahoo, 553

Waldsteinia, 535
Walnut, Black, 504
Walnut Family, 502
Water Chestnut, 569, 617 |
Water Ferns, 436

Water Hyacinth, 617

INDEX

Wood-rays, 170
Wood-wedge, 170
Wormwood, 615

Xanthophyll, 99, 100
Xanthorhiza, 519
Xanthoxyli Fructus, 546
Xanthoxylum, 546
Xerophytes, 164, 617, 619
Xylem, 140

defined, 142

primary, 143

ray, 139

secondary, 143

stages in development of elements

of, 138

Yam Family, 491

Yarrow, 615

Yaupon, 25

Yeast, Brewer’s, 382
Compressed, 386
vitamins in, 105, 106, 386
Wine, 386

Yeasts, bottom, 383, 386
classification of, 382
defined, 381
top, 384, 386

Yellow Jasmine, 583

Yellow Parilla, 521

Yerba Santa, 592

Yew, 467
Fig. of tracheid of, 23

Yew Family, 465, 466

Yohimbe, 605

Yucca, 220, 259

793

Kamia, 455
life history of, 456
outline of life cycle, 462
Kamia floridana, Fig. of, 459
Kanthorhiza apiifolia, 519
Kanthoxylea, 545
Kanthoxylum, 545
Americanum, 546
XKanthoxylum Clava Herculis, 546
Zea N. F., 485
Kea Mays, 485
' Fig. of, 483
starch of, 79, 80
structure of stem of, 215, 216
Zedoaria, 492
Zingiber, 492
Aingiber officinale, 492
Fig. of, 493
Aingiberacea, 492
Zizyphus, 556
Aizpphus sativa, 556
Zéogloea, 178
Zoology, defined, 1
Zoéspore, 45, 342, 345, 353
Figen. Of, 305
Zoéspore formation, 45
Xostera, 291
Zygnema, 343
Fig. of, 338
Kygnemacea, 343
Zygomycetales, 376, 377
Zygophycez, 333
Zygophyllacez, 544
Zygospore, 46, 342, 345, 353 ,379
Zygote, 16, 46, 345, 348, 380, 461
Zymase, 102, 103
Zymogens, 364