THE

PRINCIFLERS

OF

PHARMACOGNOSY

AN INTRODUCTION TO
THE STUDY OF THE CRUDE SUBSTANCES
OF THE VEGETABLE KINGDOM

FRIEDRICH A. JKIGER, Pu.D., M.D.,

AP Neds ‘ontiing
PROFESSOR IN THE UNIVERSITY OF STRASSBURG
AND

ALEXANDER TSCHIRCH, Pu.D.,

LS CTO ease,
LECTURER ON BOTANY AND PHARMACOGNOSY IN THE UNIVERSITY OF BERLIN

WITH ONE HUNDRED AND EIGHTY-SIX ILLUSTRATIONS IN THE TEXT

TRANSLATED FROM THE
Seconp AND CoMPLETELY REVISED GERMAN EDITION

BY
FREDERICK B. POWER, Pu.D.,

PROFESSOR OF MATERIA MEDICA AND PHARMACY IN THE UNIVERSITY OF WISCONSIN

| NEW YORK = og.
WILLIAM WOOD & COMPANY >
1887 2

Mo. ‘Bot Garden

1913

- THE AUTHORS’ PREFACE.

Ty the present work, the “ Principles of Pharmaceutical
Materia Medica” (Grundlagen der Pharmaceutischen Waa-
renkunde), by F. A. Fliickiger, published in 1873, appear
in a revised form, and materially extended. In this second
edition, the above-named author has undertaken chiefly the
preparation of the first part, and the newly associated author
(A. Tschirch), the second part, comprising morphology and
anatomy, so that really a new book has been produced. Al-
though the original plan has been essentially retained, the
arrangement of the material within the more extended space
has, nevertheless, necessitated numerous changes.

It seemed to us appropriate in this revision to also take into
consideration those crude substances of the vegetable kingdom
possessing an organic structure which receive technical appli-
cation. On the other hand, zoology has now been but inci-
dentally considered, for the reason that pharmacognosy has to
treat of only a very limited number of animal substances.

With regard to the classification of tissues, we have thought
it proper to follow Haberlandt’s “ Physiological Plant
Anatomy” (Physiologische Pflanzenanatomie), Leipzig, 1884,
_ which, besides the anatomical consideration of tissues, also
elucidates their physiological functions. In our capacity as

teachers, we have acquired the experience that an anatomical a

ae description becomes of much more interest to the pupil by a oe

oh reference to a relations, ee. the — :

af Oe .

iv AUTHORS’ PREFACE.

of physiological plant anatomy in its details is still in need of
completion and general recognition, the outlines of the same
appear to us, nevertheless, to be sufficiently well defined to
be able to serve, in the manner intimated, as a foundation.

In one point, however, we have departed from Haberlandt’s
classification. The cell, its contents, and its membrane have
been treated by us in a more complete manner, and placed be-
fore the tissues. This deviation recommends itself for practi-
cal and didactic reasons ; for the beginner must be instructed
regarding the cell and its contents before learning anything of
the tissues. The substances contained in the cell also possess
too great an interest for the pharmacognosist to be inserted
in the text simply in a secondary manner.

A second change, as already intimated, is that we have like-
wise made a place for technico-microscopical investigation, of
which every one will approve who takes into consideration the
extent to which the apothecary of our day is called upon as an
expert. In order to aid in furthering this service of phar-
macy, which is of general interest, we have, for example,
treated more thoroughly of starch and the textile fibres,

In the morphological portion, which has experienced a
complete transformation, we have endeavored to give a
brief sketch of the most important phenomena, whereby
the technical expressions in present use have received explana-
tion, especially those which occur in the deservedly widely
distributed Syllabus of Eichler. Here, as in the anatomical
portion, we have drawn the narrowest boundaries, since the
present work does not pretend to be a complete text-book or
manual. Nevertheless, some few sections which are of im-
portance to pharmacy have received relatively somewhat
greater development and more precise adaptation, as for ex-
ample, that relating to the receptacles for secretions. A chap-
_ ter on the galls has also been newly inserted.

_ In the selection of the woodeuts, we have chiefly considered
the drugs and crude substances of technical application. The

AUTHORS’ PREFACE. Vv

same applies to the selection of examples from anatomy and
morphology. The one hundred and four illustrations of the
first edition have, for the most part, been again introduced, al-
though increased by thirty-seven illustrations sketched by ‘the
_ newly associated author, as also by a number borrowed from
other works, the names of which, as a rule, have been men-
tioned. With some of the Thistrations which were placed at
our disposal by the publishing house of Springer, it has not
each time been stated whether these were derived from the
works of Hager, Hartig, or Méller. Very often we were also
obliged to content ourselves by referring to still other illustra-
tions, for example, to the plates of the handsome anatomical
atlas by Berg.

Tt has likewise been our endeavor, through abundant cita-
tions of the literature, to be of service to those members of the
profession who may desire more complete information. Al
though not every one will be in a position to refer to the
sources of information cited by us, we nevertheless wished, on
the one hand, to show that we have endeavored to utilize the
best and most recent acquisitions upon the wide field of the
auxiliary sciences, and, on the other, we hope thereby to afford
an impulse in many directions. The latter purpose was par-
ticularly kept in view with regard to the section devoted to the
history of drugs. Tue Avrnors.

STRASSBURG AND BERLIN, May, 1885.

TRANSLATOR’S PREFACE.

Tue generally acknowledged importance of the study of
pharmacognosy as a branch of useful knowledge, and the con-
stantly increasing recognition of its extended practical, as well
as scientific applications, will doubtless afford to all who are
conversant with the subject a sufficient vindication for the pro-
duction of an English version of a work which bears, as its
highest commendation, the names and impress of its distin-
guished authors. _

The work, here presented will doubtless at once indicate the
aim and the scope of the science of pharmacognosy, and
clearly demonstrate its intimate connection on every hand with
chemical, botanical, and microscopical science, as also the im-
pulse afforded for the further investigation of points of his-
toric interest and a more extended knowledge of the geograph-
ical or climatic and commercial relations of vegetable products
in their varied applications, either as medicinal remedies, as
food, or in the arts.

However important the consideration of the physiological
action and therapeutic uses of drugs may be from the stand-
point of medical science, it is evident that this alone should
not suffice for the professional pharmacist or for pharmaceuti-
cal students, who should be instructed in the science of phar-
macognosy in its broadest sense. It is thus to be hoped that
the principles outlined in the work in question may serve to
broaden the exposition of the — materia medica in ape .

Vili TRANSLATOR’S PREFACE.

directions intimated, and to secure for the same a still wider
recognition and better appreciation of its usefulness.

In the preparation of this edition, the translator has been
kindly favored by Professor Fliickiger with a few additional
notes, which have been suggested by the advances in literature
or in the related sciences during the short period which has
elapsed since the publication of the German edition, and a list
of the illustrations occurring in the work has also been ap-
pended.

The translator may finally be permitted to state that he has
endeavored to preserve in this edition not only the form, but
also the attractiveness of the original work, and while attempt-
ing to follow the original as closely as was consistent or possi-
ble, has spared no pains to maintain accuracy of diction and
the correct rendition of many newly adopted technical terms.

That the work may accomplish the mission designed by its
authors, receive to some degree the appreciation which it
merits, and be made available to a larger circle of readers, was
the highest purpose and the chief desire attending the labor
which the translator has been permitted to bestow.

UNIVERSITY OF Wisconsin, MADISON, January, 1887.

SYNOPSIS OF CONTENTS.

The mission of Pharmacognosy,
Treatment of the subject matter,
I. The mother-plant,
II. Geographical distribution of ths enniie,
III. Cultivation, i
IV. Collection. Preparation,

V. Commercial relations, ; :
VI. Description of drugs, : ‘ 5
VII. Organological importance of the pame, < :
VIII. Inner structure, .- ‘ .
IX. Chemical constituents, : : ee
X. Substitutions and adulterations, ‘ 4
XI. History, : E .
XII. Pharmacogucetioal ayilsine. :
Aids to Study, - ; . ‘ :
I. Collections, . i é
II. Literature, : :
Morphology, . .

1. Roots, rhizomes, runners and clone; sabes. bulbs, .
2, Stems, woods, barks, . : : :
3. Herbs, ‘ : .

4, Leaves, . ‘ ‘ ¥ : . ;

5. Flowers, calyx, corolla, androeceum, gyneeceum, oe

6. Inflorescence, . : : ‘

7. Fruits, 4 : ‘ . ‘ :

8. Seeds, . ‘ ‘ ‘ ‘
Plantanatomy, . : ‘

I. The cell, :
A. Contents of ia cell, pais
1. Protoplasm, cell-formation, celldvision, :
2 pads aa gees nteme eee

x SYNOPSIS OF CONTENTS.

PAGE
8. Chlorophyll bodies, . ; , : 100
4, Crystalloid coloring matters, . . . 104
S. Bat, ‘ ‘ : : ; 105
C: Slatch, ~ . “ 3 ; ‘ - 108
7. Inulin, “ : : 124
8. Crystals, cadotonn oxalate, , F . 129
9. Tannin, . ; . 135
10. Cell-sap (dry-weight), oui, hesperidin,
aloin, ; ‘ . 139
11. Inorganic substances, reduction to aals; ‘ 144
B. The cell-wall, . 148

1. Morphological ANE of the cell wid;
growth in thickness, pores, stratification, 148

'. 2, Optical behavior of the cell-membrane, . 158
3. Chemical behavior of the same, . . 4 159
(a) Cellulose, ; ‘ . 159
(b) Lignified membrane, acelin, 3 160
(ec) Cork, cutin, wax, F ‘ . 161
(a) Intercellular substance, . F 162
(e) Inorganic deposits, . F .. 163
(f)Gum, mucilage, . ‘ ; 163
(g) Pectin, lichenin, ‘ . . 170
II. Forms of cells, . ‘. e ‘ : 171
III. Cellular tissue, : ‘ ; ‘ é . 174
IV. Systems of tissue, . ‘ ; ‘ ; 5 175
A. Epidermal tissue, ; . 176
1. Epidermis, hypoderma,. cuthela, appendages
of the epidermis, hairs, ‘ ae
2. Periderm, : ‘ Pre tcy’§

(a) Phellogen, chelicdebn, Fi 187
(b) Cork, bork, ‘ F 5 . 188

B. The mechanical tissue, : : ‘ ae
Bast, collenchyma, libriform, stone-cells, nu-

cleus-sheath, ‘ ; ‘ . 194

C. The absorbing system, : : i) BOS

Root-hairs, haustoria, i : - 208

D. The assimilating system, . . : . 209

Leaf structure, . ‘ ; = . 210

E. The conducting system, . ‘ ae

1. Components of the vasculet. bundle, . . 216
(a) Vascular portion: vessels and tra-
cheids, wood-parenchyma, . 217

SYNOPSIS OF CONTENTS. xi

PAGE
(6) Cambium, annual rings, medullary
rays, : % 3 . 220
(c) Sieve portion: sijeve-tubes, cambi-
form, bark, . ‘ 231
2. Arrangement of the vascular and sieve por-
tions in the bundle, 5 ‘ . 234
F, The storing system, . : : 235
Reserve receptacles, wahenesleartoae = . 235
G. The aérating system, ; ee
Intercellular spaces, stomata, lenticela, . 235
H. System of secretion receptacles, . : 241

1. Cells containing oil, resin, mucilage, crystals
and laticiferous juice, glandular hairs. 243

2. Lysigenic passages: laticiferous ducts, lysi-
genic oil and balsam receptacles, . 247

3. Schizogenic passages containing oil, balsam,
resin, gum-resin, laticiferous juice

and mucilage, . . < -..
Pathological Formations, . ; i : : : . 264
Micro-chemical Reagents, : ; : : : . 268

Index, : ; ‘ - ‘ ; ; és . 281

LIST OF ILLUSTRATIONS.

FIG,
1. A bulb, median longitudinal section, 5
2. Leaves of Hucalyptus globulus Labillardiére, . é
3. Gratiola officinalis, rhizome with rootlets,
4, 5, 6, 7. Valvate, imbricate, convolute, and plicate einivation;
8. Hypogynous, perigynous and epigynous flower, .
9. Crocus sativus, stigma, . ‘
10. Schematic figure, elucidating the pees of fertilization,
11. Diagram of a flower of the Graminex, :
12. Diagram of a typical dicotyledonous flower,
18. Diagram of a typical monocotyledonous flower,
14. Diagram of a cruciferous flower, . :
15, Diagram of a papilionaceous flower,
16. Schemes of inflorescence,
17. Conium maculatum, section of fruit,
18. Pisum sativum, legumes,
19. Brassica oleracea, silique,
20. Colchicum autumnale, capsule,
21. Hyoscyamus niger, capsule, ‘
22. Forms of dehiscence of the capsule, .
28, 24, 25. Atropous, anatropous and campy lotropods ovules,
26. Myristica fragrans, longitudinal section of fruit,
27. Toxodium distichum, section of a medullary cell, . é
28. The process of cell division, seams oid ago
29. Elliptical starch granules, :
30. Semen Ricini, cells from the sims,
31, Gluten-cells and aleurone, :
32. Spectrum of chlorophyll and xanthophyli,
33. Chlorophyll granules, .
84. Crystalloid coloring matters,
_ 35, Starch granules from the potato,
_ 36. Starch granules from the rhizome of ginger,

.

. .

. .

. . . .

LIST OF ILLUSTRATIONS. xili

FIG. PAGE
87. Starch granules from Tuber Colchici, . : : . 110
38. Compound starch granules, 2 = dit
39. Compound starch granules from Radix Sareaparille, ‘ a8 bs |
40. Compound starch granule of the oat, 112
41. Bone-shaped and club-shaped starch grunales of Ruphordia; « 112
42, Starch granules of Sago, . : ‘ ‘ - 118
43. Potato starch, : . ‘ : s ; «148
44, Bean starch, , é ; é ‘ : y 114
45. Maranta starch, : ¥ ; ‘ é 2 . 114
46. Curcuma starch, . s ; : , é ; 115
47. Wheat starch, — : 3 ; : ; : - 116
48, 49. Maize starch, . : : : ‘ ; ; 116
50. Rice starch, . ; ‘ é ‘ : ‘ Sars Ys
51. Oat starch, ‘ ‘ : é 117
52. Starch granules in pobstiied light, . : ‘ ~ 119
58. Corroded starch granules, : : : : ‘“ 121
54. Spheero-crystals ofinulin, . é ; - 125, 126
55. Rhaphides from Radix Sarenparill, Fe 127
56. Bulbus Scille, transverse and longitudinal acticin: : . 128
57, 58. Crystals of calcium oxalate, monoclinic, . ‘ : 130
59. Twin crystals of calcium oxalate, . ‘ A ; ~ 18k
60. Quadratic crystals of calcium oxalate, ‘ ‘ < 132
61. Transverse section from an ordinary oak-gall,_ . : . 132
62. Rosettes of calcium oxalate, . 133
63. Aloe socotrina, transverse section Sheccnisthe the itergindd portion

of a leaf, - * ‘ 143
64. Schematic reprecsieésion of the development of the wall of a

wood-cell, . ‘ . 149
65. Polyhedral vaenchyine feos Rhizoma Grainints, ; Re |
66. Spiral and annular vessels from Bulbus Scille, . 5 . 150
67. Cells with net-shaped thickenings, . 151
68. Rhizoma Filicis, showing vessels with scalariform ilekeninas: 152
69. Porous cells, . : . 152
70. Thickened cells with ‘pore-canals; bast-fibre and stone-cells, 153
71, Spirally arranged pits, . . é . . 154
72. Areolated dots of the tracheids of fir-wood, : = . 154
78, 74. Bast-fibres from Cinchona barks, . 3 155, 156
"5, 76. Various stone-cells, : cys
77. Sections of bast-fibres and atonb-oella obsse-ved in pola

light, . . ‘ gor 158

78. The formation wi gum in herrea eed, " " cee a

79, Transverse section through tragacanth, . + + 168 oa,

xiv LIST OF ILLUSTRATIONS.

FIG. PAGE
80. Spheroidal cells, Isodiametric parenchyma, . ive Meg od
$1. Parenchymatous tissue from the pith of the elder, . ‘ 171
82. Hyphe from Fungus Luricis, . ‘ ‘ . 173:
83. Vanilla, longitudinal section through the outermost layer, 197
84, Epidermal cells of a root (epiblema), ‘ ‘ Rae i 3
85. Mace, transverse section, 4 a - ‘ . 178
86. Colocynth, rind of the fruit, ; : : . 17%
87. Semen Paradisi, transverse section, . : : ‘ 179
88, 89. Semen Hyoscyami, transverse section, : ames Wt Pa bot
90. Semen Stramonii, tangential section through the epidermis, 181
91. Chinese galls, transverse section, . ; ; - ~ 13k
92. Hairs of cotton, . ‘ 182:
93. Stinging hair of the nettle. and hairs of Nux conan, A . 183:
94. Fructus Anisi, transverse section, ; : 184.
95, Flores Verbasci, club-shaped and stellate Sindee ‘ ‘ . 185.
96, Oil-glands of the Labiate, : ‘ 186.
97. Canella alba, cells from the phelioderss of the back. P +, oem

98. Cinchona Calisaya, transverse section through the bork, . 188.
99. Juniperus communis L., transverse section through an older

internode, : : 5 j ; ee
100. Cortex Sassafras radicis, bork, . ‘ : A . 189
101. Cork of the cork-oak, . : : : +k
102. Radix Calumbe, cortical lage, . ? ; . 190:
103. Rhizoma Curcume, transverse section, ‘ : st 1 Jee
104. Cortex Cascarille, cork “—_* and primary bark, . . 191
105. Cortex Guaiaci, : F ‘ . 192
106. Radix Pyrethriromani, . . 192°

107. Jamaica Quassia-wood, transverse panies through the bark, 193.
108. Potato, transverse section through the outermost layer, . 193
109. Vanilla, transverse section through the fruit; collenchyma, . 195

110, Typical bundle of bast-cells, —. . ; ; 5 oe
111. Bast-cells of Corchorus olitorius (jute), . , ‘ . 196-
112, Alfa-grass or Esparto, transverse section through an involute
leaf, ‘ ‘ 4 " ‘ i . 197
113. Linum usitatissimum, transverse stein ofstem, . co
114. Illustrations of technical fibres, . . 198
115. Fructus Cocculi, sclerenchyma from inner ¢ legee of seed veal, 199
116. Tea-leaf, transverse section, : . : : . 200°

117. Cinchona barks, short bast-cells, 5 ‘ xx: Ot
118. Cortex Coto, transverse section through the si sieve portion, . 201
119. Rhizoma Veratri, transverse section of rootlet, t 202

420. Acteea spicata, transverse section of rootlets, eas ce a

LIST OF ILLUSTRATIONS. xV

FIa. PAGE.
121, Aconitum Napellus, transverse and longitudinal section of root-
lets, , . 204
122. Helleborus viridis, trunavaris cians of ranaiaie, ‘i ‘ 205
123. Sarsaparilla, radial longitudinal section, ‘ : . 206
124. Vera-Cruz Sarsaparilla, transverse section. ‘ ‘ " 207
125. Rhizoma Galange, transverse section, . F ‘ - 207
126. Radix Sarsaparille, longitudinal section, : F ‘ 209
127, 128. Eucalyptus globulus, transverse sections of leaf, . 210, 211
129. Mentha piperita, transverse section of leaf, . ‘ . 212
130. Leaf of Digitalis purpurea, ‘ ‘ . < . 214
131, 132. Rhizoma Filicis maris, : ‘ 214
133. Maize stem, transverse section through a collateral aaa, . 216
134, Spiral and annular vessels from Bulbus Scille, A ei aEe
135. Vessels from Rhizoma Filicis, ; ; : . . 218
186. Longitudinal section of two tracheids, < é 219
137, Lignum Quassic jamaicense, transverse section, ‘ . 219
138. Schematic representation of the activity of a cambium cell, 221
189, Libriform from Quassia wood, : é , ; . 222
140. Cells of the medullary rays, ‘ ‘ < 224
141. Medullary rays from Lignum Siniperi, : ‘ .- ae
142. Rhubarb, tangential longitudinal section, 7 ‘ 225

143, Wall-like cells of medullary rays from Lignum Juniper, - 226
144. Stipes Dulcamare, one-rowed medullary rays, Fs 226
145. Cinchona lancifolia, transverse section through the phloén,: 227
146. Rheum Rhaponticum and true rhubarb, transverse sections, 298, 229
147. Sieve-tubes from Fructus Papaveris, ‘ ; : - 230
148. Sieve-tubes of Acer,
149. Ceratonia Siliqua L., frauaveres iickions of bark, : . 232
150. Bast-fibres from Cinchonns alba Payta, . 4 : 233°
151. Rhizoma Calami, moniliform tissue, ‘. : » 236
152. Fructus Aurantiorum, transverse section from the — 236
153. Caryophyllus, transverse section, ‘ 237
154. Mentha piperita, surface view of epidermis of ee . 238
155. Mentha piperita, vertical section of a stoma, .
156, 157, 158. Stomata from dicotyledons and monocotyledons,
159, 160. Arillus of Myristica spec., cross section, . « .
161. Cinnamomum Camphora, cross section through a sub-epider-
mal oil-cell of leaf, . ‘
162. Aspidium Filia mas, fundamental tinstie of rhizome, =
163, 164, 165. Radix Taraxaci, latex-tubes, = 4 . 246,

.

166. Tuber Jalape, cells containing laticiferous j juice, .
167. Latex-tubes of the fig, i

. . . .

Xvi LIST OF ILLUSTRATIONS.

NO. PAGE
168. Formation of an oil-gland of Dictamnus Frawinella, : 250
169-175. Formation of intercellular spaces, : ‘ 252, 253
176. Balsam-passage (oil-space) of Radix Inule, : . 253
177. Rhizoma Arnice, central portion of an oil-space, . ; 253
178. Radia Sumbul, longitudinal section from the bark, : . 254
179. A coniferous leaf, transverse section through an oil-canal, . 255
180. Pinus silvestris, transverse section from the wood, ~ . 256
181. Pistacia Lentiscus, transverse section of the bark, . ‘ 256

182. Arnica montana, portion of a transverse section of the rhizome, 257
183. Fructus Foeniculi, longitudinal section of an oil-passage, . 259
184. Fructus Conii, transverse section, . - i Q . 260
185. Fructus Conii, longitudinal section through coniine layer, . 261
186. Cinnamon bark, elements isolated by maceration, é . 270

THE PRINCIPLES

OF

PHARMACOGNOSY.

THE MISSION OF PHARMACOGNOSY.

THE substances which are employed medicinally for their
remedial action are either the products of human skill or they
belong directly to the two organic natural kingdoms. Among
the medicinal substances formed through chemical operations
we meet, indeed, with such as are produced only by certain
definitely conducted chemical or physical processes, as, for
example, the acids, iodine, bromine, chloral, phenol, glycerin,
alcohol, and the alkaloids, be it that chemical industry starts

from materials belonging to the inorganic kingdom, or that its
first point of departure, as in the two last-named examples, falls
within the circle of organic nature. In some rare cases only
(mineral waters, for instance) is it sufficient to merely choose a
- suitable form for that which nature offers ready for use; more

frequently chemical skill is directed toward the isolation of
active principles from plants (exceptionally also from animals
or animal substances, or from the mineral kingdom), and to the
separation of these principles from associated substances, or,
in other words, to their purification. In all these cases the

. _ object i is to ploce at the Sisppent ot ot medical science bodies Meoae oe

2 THE MISSION OF PHARMACOGNOSY.

are sharply characterized in a chemical sense, or, briefly stated,
chemical units; for only such substances as are at all times ac-
cessible and complete in their identity can afford a sure founda-
tion for scientific medicine and pharmacy. In this direction
lies the aim of the future.

Medicinal agents of this kind are outside of the sphere of
pharmacognosy. By general concurrence in pharmaceutical
circles, there are assigned to it those substances which are di-
rectly furnished by nature, or at least such as have not actually
been submitted to chemical processes. Since the few crude medi-
cinal substances from the mineral kingdom which were em-
ployed in former times have long ago lost their significance, the
scientific knowledge which pharmacognosy has to offer is con-
fined to organic nature, or virtually only to the vegetable king-
dom; for even among the animals, and the parts and products
of animals, only castor, musk, and cantharides represent at the
present time important elements as medicinal agents. In the
cantharides cantharidin alone is of importance, which now
stands at the disposal of medical science in a pure form. Phar-
maceutical interest in the beetle itself is therefore diminished in
a similar manner as it is, at least from this standpoint, in the

“animals which afford cod-liver oil, honey, isinglass, and milk- .
sugar.

The wonderful development of the natural sciences and of
medicine, which exerts so great an influence, especially
since the second decade of the present century, has liberated
pharmacognosy of an enormous burden. In a pharmaceutical
work which was first published at Ulm in the year 1641, under
the title “‘ Pharmacopoeia medico-chymica seu ‘Thesaurus phar-
macologicus,” by the city physician Johann Christian Schréder,
of Frankfurt-on-the-Main, and which in its time was much
valued, the author also enumerated the **simplicia” then em-
ployed. Among these there were about thirty minerals, and
more than one hundred and fifty medicinal substances derived —
from the animal kingdom or representing entire animals, to-
gether with a very large number of roots, herbs, leaves, etc.
Such a superfluity of medicinal agents, with all of which it was —

_ Wissenschaft, etc.,” by A. Tschirch, Pharm. came) 1831, ate 9

THE MISSION OF PHARMACOGNOSY. 3

impossible to become accurately acquainted, characterizes the
medicine and pharmacy of the European middle ages' and the
condition of popular medicine still existing among the Asiatic
nations.

The primary object of modern pharmacognosy is to scrutinize
all the evidence offered by botany, zoology, and pharmacy re-
garding the medicinal agents under consideration, to arrange
this material in scientific form, and by an appropriate and com-
prehensive representation to subject it to a closer examination.
It is only by this means that pharmacognosy assumes the form
of a branch of knowledge which is of equal importance to both
pharmacy and medicine. Upon a thorough acquaintance with
medicinal substances, and their proper treatment, the practical
success of pharmacy is largely dependent, so that a deeper study
of the science of pharmacognosy may reasonably be expected of
the pharmacist. He will therefore confer honor upon himself
and his profession if he advances somewhat farther even than is
unavoidably required by his immediate interests. It is, how-
ever, scarcely possible to sharply define the boundary between
the daily round of duties and scientific studies, and, indeed, this
is not only impracticable, but even undesirable.

Pharmacognosy should therefore comprehend, to a certain
degree, all that pertains to a monographic knowledge of medi-
cinal substances.* The consideration of these substances from
the above-outlined standpoint of natural science is to be supple-
mented by purely historical and geographical references, and
such as are connected with the history of civilization, together
with commercial relations. All this should be made to assume
a rich, animated, and symmetrical form which, in many cases,
may become quite attractive. Among this copious material, how-
ever, those characters must be prominently pointed out which
can lead to a rapid, approximate valuation, whenever it is

* Compare ‘‘ Die Frankfurter Liste” in Archiv der Pharm., 201 (1872), _
pp. 453-511, and ‘‘ Das Nérdlinger Inventar,” ibid., 211 (1877), p. 97.

* The aims of modern pharmacognosy have recently been thoroughly : oe 2

explained in the essay: ‘‘ Ueber die Bedeutung der Pharmacognosie als

4 THE MISSION OF PHARMACOGNOSY.

practicable to do this, without resorting to an actual chemical
analysis. In an accurate knowledge of the nature of medicinal
substances lies, indeed, the best protection against substitution
and adulteration.

The most important property of medicinal substances, how-
ever, is their medicinal action, yet this must remain excluded
from pharmacognostical consideration, for it has become the
subject of an independent scientific discipline, viz., pharma-
cology. From apparent practical reasons it will, indeed, occa-
sionally be found advisable, especially in the case of particularly
interesting substances, to at least allude to their therapeutic
action. That these two domains present many points of contact,
and that pharmacology receives especial support from scientific
pharmacognosy, in fact, that it presupposes a knowledge of the
latter, is clearly evident. In England, France, and other coun-
tries, the two expressions, Pharmacology, which treats of the
therapeutic action of medicinal substances, and Pharmacognosy,
which comprehends a scientific knowledge of the substances
themselves, are occasionally employed in a sense different from
that which has just been indicated. It must be admitted that
the tenor of the two words expresses no sharp distinction.*

With regard to the chemical side of pharmacognosy, it is
necessary to restrict it within certain limits. It is doubtless
quite as appropriate that the properly isolated constituents of
drugs should be enumerated and characterized, as that it should
be stated, or at least intimated, where material deficiencies occur
in this direction, which it is desirable to supply. An exhaustive
treatment of the chemical constituents, however, falls within
the domain of chemistry, or pharmaceutical chemistry.

' Schmiedeberg, in his ‘‘ Grundriss der Arzneimittellehre,” Leipzig,
1883, says: ‘‘ The science which treats of medicinal substances (Arznei-
mittellehre) has only to do with such agents as are useful in the curing
of disease. It is, therefore, desirable that all substances which do not
serve as food, and which, through their chemical properties, produce
changes in the living animal organism, should, in order to investigate
these effects, be brought within the borders of a single science, which

may be called Pharmacology, or, since it is chiefly supported by experi-
_ ment, Experimental Pharmacology -

-

THE MISSION OF PHARMACOGNOSY. 5

Accordingly, the aim and position of pharmacognosy appear
definitely outlined when, beside the questions which it has
to answer, the boundaries are also drawn beyond which it
should not pass. It is thus to be understood, when on page
3, with a less exact expression, mention was made of a “ certain
degree” of completeness.

In the fostering of pharmacognosy, botanists and physicians in
former times have rendered service, aided, indeed, occasionally
by apothecaries." The most brilliant of such services on the
part of physicians was brought to a close by the publication in
London, 1839 and 1840, of ‘The Elements of Materia Medica
and Therapeutics” by Jonathan Pereira; for, in the mean time,
the separate and more thorough treatment of pharmacognosy in
the previously explained direction had been taken in hand by
pharmacists, and at the earliest period, and with by far the
greatest success, by Guibourt, the former teacher of this branch
of science at the Ecole de Pharmacie in Paris.? Similar, though
less prominent results were accomplished in Germany by
Johann Bartholomaeus Trommsdorff, of Erfurt, through his
‘Handbuch der pharmaceutischen Waarenkunde,” Gotha, 1822,
and particularly, also, by Theodor Wilhelm Christian Martius.
In his ‘‘Grundriss der Pharmakognosie des Pflanzenreiches,”
Erlangen, 1822, Martius says that pharmacognosy is to be re-
garded as ‘‘a part of general materia medica, or that science which
relates to the examination of the medicinal substances derived
from the three kingdoms of nature with a view to ascertain
their source and quality, to test them for their purity, and to

‘Examples in Flickiger’s ‘“‘ Pharmakognosie,” 2d edition, p. 1,013
(Pires); p. 992 (Morgan); p. 209 (Bansa).

* As precursors of Guibourt may be mentioned Nicholas Lémery,
author of the ‘‘ Traité universel des Drogues et Simples,” 1697, and
Etienne Francois Geoffroy, whose ‘‘ Tractatus de Materia Medica” ap-
peared in 1741. These two Parisian apothecaries were, however, more
properly recognized as physicians. The “ Histoire générale des

Drogues,” 1694, which is likewise to some extent worthy of considera~
tion, has as its author the druggist Pierre Pomet. The three above-

named publications demonstrate how much pharmacognosy was at that ae
time indebted to Paris. _ Ce ee

6 é THE MISSION OF PHARMACOGNOSY,

determine substitutions and adulterations.” In the year 1825,
Martius began to deliver lectures on pharmacognosy at the
University of Erlangen, which were the first, as it appears, that
were announced under this name. The credit of having aided
in securing general approbation for the new expression pharma-
cognosy’ is due to the writings and lectures of Wiggers (see
below).

The question, however, now arises: what is a medicinal sub-
stance? ‘To attempt to give a more precise definition is useless,
for this term has been undergoing a continuous change in the
course of time and in the light of advancing knowledge, not
only from the period of antiquity, but also from -country to
country, and, indeed, from one medical school to another, from
one pharmacopeia to another. We are compelled here to assume,
as it were, an intermediate standpoint, and to select those sub-
stances of importance which are employed within the circle of
observation available to us. That which has already been con-
signed to oblivion, or is only very rarely employed, and especially
that which is no longer used by scientific medicine, deserves less
attention than new drugs which may be presumed to have a
valuable future. From a pharmaceutical standpoint, however,
considerable interest may still be attached to many substances
even though they find but little medicinal application at the
present time. With regard to Nuwz vomica, Santonica, Radix
Belladonne, Galle, and Podophyllum, by way of example, a
scientifically educated pharmacist will desire to be satisfactorily
informed, even when these crude substances shall have become
banished from medicinal use to a much greater extent than is
at present the case. In proportion as strychnine, santonin,
atropine, etc., become of greater importance in their medicinal,
or forensic and technical relations, a corresponding attention
should be paid to their derivation. Neither pepper, nor piperine

1Tt occurs also as the principal title: ‘‘ Pharmakognostiche Tabel-

.

len” in the fifth edition of Joh. Christ. Ebermaier’s “‘ Tabellarische

Uebersichten der Kennzeichen der Aechtheit und Giite, etc., der a

_ Arzneimittel,” Leipzig, 1827. The first edition of this work pene? in

ae : - 1804, but whether under the above title is not known to us.

THE MISSION OF PHARMACOGNOSY. 7

or piperidine play an important part in modern medicine,
although the former constituted for centuries the most important
of all spices, and still maintains, as a condiment, a prominent po-
sition in the world’s commerce. Pharmacognosy would not
completely fulfil its mission if such circumstances were not taken
into consideration; and a similar statement might be made
regarding cacao, tea, coffee, coca, cola, guarana, and mate.

Important medicinal substances become more clearly intelli-
gible when they are compared with others which may in them-
selves be insignificant; and herein also lies both a demand and a
justification for pharmacognosy to occasionally extend its do-
main in an apparently unnecessary manner. ‘Thus the so-called
spurious Cinchona barks for a long time presented only subordi-
nate interest, but the consideration of their structure was
admirably adapted for the demonstration of the varying peculi-
arities of those barks which alone furnished quinine and the
allied alkaloids until the Cinchona cuprea appeared in the mar-
ket (since 1880), and proved that these bases are by no means
confined to the true Cinchona barks.

Hence there are various considerations which are liable to
extend the compass of pharmacognosy. On the other hand, there
are cases where substances which apparently belong here may be
permitted to remain unconsidered. This may occur in such
cases, for instance, where chemistry alone is capable of affording
an exhaustive characterization. In the fats, wax, volatile oils,
and the several varieties of sugar, the purely chemical properties
are of such eminent importance that pharmacognosy can only in
exceptional cases find motives to complete their characteristics in —
other directions. This task must be relegated to those depart-
ments which concern themselves with the chemico-technical or
commercial knowledge of such products, for even the latter has —
for some years past likewise experienced the most substantial
advancement.’ The latter has been materially aided by the

1 See Wiesner, ‘‘ Die Rohstoffe des Pflanzenreiches,” Leipzig, 1873, andl aes

the collective work begun in 1882 by Benedikt and eight associates, oS
_ * Allgemeine Waaren- und Rohstofflehre,” Cassel and Berlin, Fischer. _

The fifth volume —— “Die iiasalae und saconmoanenians aus dem :

38 THE MISSION OF PHARMACOGNOSY.

earlier development of pharmacognosy, while, inversely, pharma-
cognosy receives at the present time many impulses from com-
mercial relations, as an example of which may be quoted the fact
that it has to derive statistical and other information relating to —
important drugs from consular reports. An abundant supply
of such information is especially offered in the semi-annual re-
ports of the firm of Gehe & Co., of Dresden, which may regu-
larly be obtained from the booksellers.

Only such things fall within the sphere of pharmacognostical
consideration as cannot, for our purpose, be sufficiently investi-
gated by a single science. With regard to many leaves, flowers,
seeds, and fruits, it may indeed be contended that botany is
capable of affording a sufficient description ; it is, however,
readily to be observed that pharmaceutical requirements demand
the consideration of other than purely botanical properties. ‘The
changes which occur on drying, chemical qualities, commercial
relations, and historical facts are al! equally worth knowing,
and call forth the discriminating and classifying ability of phar-
macognosy.

It must, however, be admitted that the limitation and treat-
ment of objects of pharmacognostical study are rather arbitrary.
Pharmacognosy is by no means a branch of knowledge with
sharply defined boundaries, and herein, in fact, lies the nature,
_ and probably also a special charm of the science, that it enlists the
aid of several disciplines for the single purpose of acquiring a
more thorough knowledge of the crude substances employed as
medicinal agents, or of parts of plants or products which are
otherwise important from the standpoint of pharmacy.

Pflanzenreich,” by T. F. Hanausek, 1884, pp. 485. With one hundred —
“lecodents Ske

TREATMENT OF THE SUBJECT-MATTER.

In most cases, the following points prominently present them-
selves:

I. Naming the plant (or the animal) from which the sub-
stance is derived.

Here it happens not infrequently that a consideration of the
synonyms is indispensable in order to avoid misunderstanding;
for if we revert no farther back than to the time of Linné, we
sometimes meet with plants which, since then, have been
variously named by botanists. Thus, for instance, Hagenia
abyssinica Willdenow (1790), Bankesia abyssinica Bruce (1799),
and Brayera anthelmintica Kunth (1824), designate the tree
which furnishes us the koosso. Linné, in 1737, represented the
clove tree under the name of Caryophyllus aromaticus, but Thun-
berg, in 1788, named it Hugenia caryophyllata, properly at-
taching it to the genus Eugenia, which had existed since the
year 1729. Many examples of this character are to be found’ in
the families of the Umbellifere, Composite, and Labiate.
Since the end of the preceding century, the mother-plant of the
calumba root has received half a dozen, and the sabadilla plant
four names. On the other hand, the same name has occasion-
ally been bestowed upon different plants. Thus Linné’s Croton
Cascarilla is a different shrub from that so named by Bennett,
and the Croton Eluteria of the latter, which furnishes the

cascarilla bark, is not the same tree as that intended by Swartz
under the same name. A similar condition exists with refer- =
ence to Cassia lanceolata, under which name Nectoux under- _
stood the present Cussia acutifolia, Delile, Wight, and Arnott

the ©. angustifolia of Vahl, while Forskal’s C. lanceolata is -
identical with C. Sophera L. Pliny had ee made a dis- ae

10 TREATMENT OF THE SUBJECT-MATTER.

tinction between the Norway Spruce Fir,’ Picea, and the-
Silver Fir,’ Abies ; but Linné, in 1753, transferred the name
Pinus Abies to the first-mentioned, which, in 1771, was again
reversed by Duroi. The tree which furnishes kamala has re-
- ceived in the course of time the following names: Croton philip-
pense Lamarck (1786), Echinus philippensis Loureiro (1790),
Rottlera tinctoria Roxburgh (1798), Mullotus philippinensis
Miller Argoy. (1862), and Echinus philippinensis Baillon
(1865). ae

Of some few drugs the mother-plants have not yet been ascer-
tained with perfect certainty. For example, we do not know
precisely whether Rhewm officinale Baillon furnishes most or all
of the commercial Radix Rhei. Equally unsatisfactory is our
knowledge of the plants which afford asafetida, sarsaparilla,
olibanum, elemi, copaiva balsam, galbanum, ignatia seeds, and
Levant soap-root. The origin of tragacanth, of Asiatic salep,
and of condurango bark has also not yet been determined with
sufficient accuracy.

If. Geographical distribution of those plants which are of
importance in pharmacy.

However little of striking significance this subject may pre-
sent from a practical point of view, it highly merits consideration
in a scientific treatment of the topic. The representation of
a pharmaceutically important plant is only then truly satisfac-
tory when we are also informed regarding its habitat or the
extent of its cultivation. ‘T'he sources from which this knowl-
edge is to be obtained are, in the first place, the floras of indi-
vidual countries, works on plant geography,’ and narratives of
travel. Since many officinal plants are among those that have been
longest employed by man, or are distributed over large tracts of
territory, or are at least generally known, they find incidental con-

' Fichte or Rothtanne of the Germans.

* Weisstanne or Edeltanne of the Germans.

* A. De Candolle : « Géographie botanique raisonnée,” 1855, also ‘‘ Ori-
gine des Plantes cultivées,” 1883. Grisebach: ‘Die Vegetation der
Erde nach ihrer klimatis-hen Anordnung,” 1872,—second edition, two
vols,, 1884,

GEOGRAPHICAL DISTRIBUTION OF IMPORTANT PLANTS. 11

sideration in the comprehensive works just mentioned. From
a narrower, pharmaceutical standpoint, an illustrative represen-
tation of the distribution of officinal plants throughout the
world has been undertaken by inscribing them upon a plani-
- ’ sphere.’ A survey of the respective plants is thus obtained,
which, however, as may readily be conceived, has nothing to do
with plant geography. The distribution or, more properly
speaking, the selection of these plants can, at most, only in so
far be considered as having taken place in a normal manner, as
it goes hand in hand with the course of civilization. India,
Persia, and the Mediterranean region, the primeval seats of
civilization, have, therefore, the preponderating number of
officinal plants to exhibit, Australia not a single one; the entire,
enormous Arctic region perhaps only Polyporus officinalis and
Laminaria, to which, if desired, Cetraria islandica might be
added. A further objection to the graphic representation of the
distribution of officinal plants may also be found in this, that
the districts of production are mostly much more confined than
the geographic area of the plants, because so many drugs, on
account of their limited use, can play no considerable part in
the wholesale trade. Cetraria islandica, for instance, is col-
lected for the drug trade in the mountains of Central Europe
and the Alps, and not in the far North. Finally, the distribu-
tion of useful plants is also sua ora upon certain influences of
cultivation.
1 Barber: ‘‘The Pharmaceutical or Moadidubitanion! Map of the
World,” London, 1868. The same map is offered also by Fristedt,
‘*Pharmakognostik Charta 6fver jorden,” Upsala, 1870, as likewise in
the appendix to his ‘‘ Lirobok i organisk Pharmacologi,” Upsala, 1873.
The map projected by Schelenz, in Archiv der Pharm., Band 208 (1876),
_ suffers from being on altogether too small a scale, although it considers.
only the plants of the Pharmacopoea Germanica. This fault is avoided —
by Oudemans in his ‘‘ Handleiding tot de Pharmacognosie,” Amster-
dam, 1880, since he dedicates a special map to each of the five divisions CS
of the earth. If it is desired to go still further, then the distribution of = =
each individual plant must be brought upon a special map. This has, =
for instance, already been accomplished by Lloyd for Hydrastis cana-
densis in his “ Drugs and Medicines of North America,” Vol. I, Be. 3,
P. 82; Bao ae the Pharm, Leica sar New Bec P- 287. oars

12 TREATMENT OF THE SUBJECT-MATTER.

IM. Cultivation of officinal plants for medicinal purposes, or
also for industrial applications.

The Cinchonas of the East Indies, Jamaica, and other coun-
tries; the poppy in Asia Minor, Bengal, and Malwa; the tobacco
which is cultivated in ali temperate and warm countries; the fea
plant in Assam; the peppermint 1n Michigan; the cinnamon in
Southern China and on the island of Ceylon; the liquorice in
Calabria, Spain, and Moravia; the various species of Citrus
(agrumi) in the Mediterranean region, California, and the West
Indies; the coca in Bolivia and Peru; and the cacao, extending
from Mexico to Brazil, are examples of the transplantation and
extensive cultivation of such useful plants, without mentioning
at all the cotton plant, the Eucalyptus, the sugar-cane, and the
sugar-beet. Inaless extensive amount, Althea, Angelica, Levis-
ticum, fennel, and anise are cultivated in Thuringia, and cara-
way near Halle; the latter two to a greater extent in Central
Russia. Furthermore, the manna-ash in Sicily; the rose in Kis-
anlik on the Balkan Mountains and in Southern England; pep-
permint in the same locality; and Lobelia in the State of New
York. Finally the extended cultivation of odorous flowers, near
Grasse in Provence,’ which are, however, more largely employed
in the art of perfumery.

The English Government, through the prudent management
of the large botanical gardens at Kew, near London, has pro-
vided for the distribution of important useful plants in India

and the Colonies. ‘To these endeavors is due the establishment
of the Cinchonas in India, while recently the same has been ac-
complished with the calumba plant, ipecacuanha, jalap, and the
trees which afford caoutchouc, gutta-percha, copaiva balsam, and
cinnamon.” The results are still to be awaited, as are likewise
_ those of similar endeavors on the part of the State Depart-

ment of the United States. :

IV. Collection and Preparation.—The section which fol-

'Flickiger, Archiv der Pharm., 222 (1884), p. 468. :

* Compare Fliickiger, ‘‘ Ueber den chinesischen Zimmt,” in Arch. der
Pharm., 220 (1882), p. 835. Further, Brockmeier, ‘‘ Ueber den Einfluss
_ der englischen Weltherrschaft auf die Verbreitung wichtiger Cultur- _

__— gewiichse, namentlich in Indien.” Dissertation, Marburg, 1884, pp. 56.

COLLECTION AND PREPARATION. 13

lows later on (p. 139), relating to the liquid contents of the
plant cell, shows how great are the changes produced in the
plant, simply through the process of drying. It is evident that
these changes bave hitherto not been, by far, sufficiently consid-
ered.’ They may often be surmised through changes in odor,
as, for instance, in the case of coriander, the underground por-
tions of Orchis (salep), Iris (orris root), Veratrum and Aconi-
tum. Occasionally the drugs assume, upon drying, other col-
ors.

It is of special importance to determine accurately the proper
time of collection of each plant, or of its officinal parts; for
during the life of a plant it does not at all times contain the
active principles in equal amount—indeed, in many plants, cer-
tain constituents are, at some periods, entirely wanting. The
time of collection is to be so chosen that the maximum amount
of the desired substances is obtained. Quite independent, how-
ever, of the impossibility of insuring this by watching the
collectors, it must also be admitted that our scientific knowledge
of these conditions is still altogether too fragmentary. In
the case of Folia Digitalis, Folia Hyoscyami, Fructus Conti,
Tuber Colchici, Rhizoma Filicis, and some few other crude
vegetable substances, we are, however, in this respect well
informed.

Digitalis leaves are weaker in active constituents before the
period of flowering than afterward, consequently the leaves of
the first year are to be entirely rejected. In the case of Hyos-
cyamus, the leaves of the second year’s growth are likewise
preferred, at least in England. Schroff, in 1870, showed that
Fructus Conii contains the largest amount of coniine immedi-
ately before the period of ripening. To the same investigator
we are indebted for the proof that Zuber Colchici is only active
at the flowering period of the plant. Rhizoma Filicis, according
to all experience, should only be collected in late summer. The |
absolute age of the respective parts also comes often into con-
sideration. Thus Radix Belladonne of the second or third year

1 Experiments worthy of notice relating to this subject have
already been made by Schoonbrodt. See Jahresbericht der Pharm., 1869,

14 TREATMENT OF THE SUBJECT-MATTER.

is richer in atropine than that of seven years’ or still older
growth, which is probably chiefly caused by the fact that this
alkaloid is mainly contained in the bark, and in older roots the
latter is relatively less in amount than in younger; the constitu-
ents of belladonna leaves are not so variable in amount.' The
fact that many fruits and seeds before ripening contain starch,
and afterwards more sugar, oil, and other substances, will be
explained further on when we shall consider the subject of
amylum. In the juice of Aeballium Elaterium Richard, elate-
rin occurs abundantly in July, but in September this powerfully
drastic, crystallizable body is wanting therein.” Pepper, cubeds,
and cloves are richer in volatile oil before ripening; the Cinchona
barks may occasionally be poor in quinine, or even contain none
at all.

It is self-evident that the quality of the soil must also have
some influence upon the chemical development of the plant.
Valerian root grown in dry localities is richer in volatile oil
than that in moist ground, and in the case of Taraxacum, the
root, from a chemical point of view, shows great variations, ac-
cording to locality and the time of year. The red Flores Mal-
ve become blue, and gentian root, which in the interior is purely
white, becomes colored yellowish-brown, unless the water is
abstracted from it with the most extreme care. Flores Rhoea-
dos do not even then retain their color, while, on the other hand,
the remarkable brown coloration of Flores Verbasci may easily
be avoided. The Cinchona barks also assume, during the pro-
cess of drying, another color.*

Still other changes occur when the drugs are scalded, or are

‘Lefort, Journ. de Pharm. et de Chim., XV. (1872), pp. 268, 421; .

Gerrard, Pharm. Journ., XV. (1884), p. 153. Compare further, Dragen-
dorff, ‘‘Chemische Werthbestimmung stark wirkender Droguen,”
St. Petersburg, 1874.

* Kohler in Buchner’s Repertorium der Pharm., XVIII. (1869), p. 590.

* The changes which the green coloring-matter of. plants undergo
upon drying, and the ways and means to be resorted to in order to re-
duce these changes to the smallest compass, have been thoroughly eluci-
dated by Tschirch: “‘ Einige practische Ergebnisse meiner Untersuchun-

gen tiber das Chlorophyll der Pflanzen,” in Archiv der Pharm., 222
(1884). .

COMMERCIAL RELATIONS. 15

‘dried over an open fire, as with salep, jalap, curcuma, and many
Indian aconite tubers. This treatment manifests itself particu-
larly by the starch assuming a pasty form. The leaves of tea
and maté, which are exposed to a light roasting process, suffer
still more significant chemical changes.

Marshmallow root, rhubarb, calamus, ginger; and pepper
(white) are pared, in order to give them a better appearance,
which, however, in the three latter cases is partly effected at
the expense of the volatile oil. As a judicious form of treat-
ment, on the contrary, is to be designated the so-called sweat-
ing process of the cacao; the slight fermentation which is
thereby produced destroys a bitter principle.

V. Commercial Relations.—A very limited number of the
medicinal substances coming under consideration here claim a
position in the world’s commerce, and play a conspicuous part
therein. In this category, opiwm should be placed in the first
line, although really only about that portion of it which does
not find medicinal application, viz.: the East Indian. Of the
same rank are the Cinchona barks. Pepper, tea, cacao, tobacco,
sugar, and maté, regarded as luxuries, and counted among the
most important products, are scarcely to be considered as drugs,
which is more truly the case with ginger. Such drugs as are
also technically employed, are usually of much greater import-
ance in the latter respect. Catechu, colophonium, dammar,
dye-woods (logwood, Brazil-wood, ete.), galls, gum arabic, gutta-
percha, turpentine, ete., are employed, for example, in various
industries in infinitely larger amounts than in pharmacy. Even
for culinary purposes spices are much more largely employed
than in pharmacy, notwithstanding the departure of modern
taste from the former custom,to strongly season articles of food.

The most valuable information in this department, viz., sta-
tistical records, are especially to be found in the voluminous
publications (Blue-books) of the English Government, and,
recently also, to an increasing extent, in those of the United
States. Furthermore, in the German “ Handels-Archiv,”

and in official tables relating to the commerce of France and

other countries.

16 TREATMENT OF THE SUBJECT-MATTER.

VI. Description of the drug itself according to external
characteristics, the odor and taste, and, in the case of liquids
also the spevific gravity. Drugs provided with organic struc
ture present many more points of observation for their exami-
nation and description.

VII. The determination of the parts which here come under
consideration, according to their organological importance.
The general considerations relating to this subject are treated
of in a subsequent morphological section.

VIII. Microscopic structure of the drugs of organic con-
struction.

To this subject special sections of the present work are dedi-
cated. Among those drugs in which development does not
proceed from the activity of cell formation in the organism, or
is not regulated in accordance with morphological laws, there

are some (as for example: Aloe, Balsamum tolutanum, Benzo-
imum, Catechu, Ohrysarobinum, Elemi, Opium, Styrax, and
Terebinthina communis) which, nevertheless, with reference to
their crystalline constituents, call for microscopic examination,
in the course of which polarized light renders essential service,
because these crystals, on account of their double refraction,
appear much more distinct under the polarizing microscope
than when observed in ordinary light.

IX. Chemical constituents.—The enumeration and brief
characterization of these principles essentially belongs to the
functions of pharmacognosy. Under this heading are to be-
considered not only the principles peculiar to certain drugs,
but also the more commonly distributed constituents of plants.

_ Though their isolation and quantitative estimation forms the
object of chemical analysis,’ yet with the aid of micro-chemical
reagents (see the concluding chapter of this book) it is possible
to arrive in a short time at a series of valuable conclusions.
regarding individual constituents. This method of procedure:
especially affords information in many cases regarding the loca-
_ tion of important constituents in the tissue.
'G. Dragendorff : ‘‘ Die qualitative und quantitative Analyse von
_ Pflanzen und Pflanzentheilen.” Gottingen, 1882 (see below, page 49).

>

SUBSTITUTIONS AND ADULTERATIONS.—HISTORY. 17

X. Substitutions and Adulterations.—An acquaintance,
based on the preceding considerations, with drugs possessing an
organic structure at once affords the means for their examina-
tion. When the structure of the respective substances has been
rendered undiscernible in consequence of a fine degree of pul-
verization, the task is more difficult. In such cases, the aid of |
chemical analysis must especially be sought. This is rendered
still more necessary when liquid drugs of variable composition
are under consideration, as, for instance, Copaiva balsam, Pert
balsam, Storax, and Turpentine.

XI. History.—The knowledge of medicinal substances
would remain incomplete if their history did not also receive
consideration. The investigation should extend to the time
and place of first acquaintance with the mother-plant, to the
time of the first medicinal employment of each single drug, and
to its importance in the world’s commerce. Outside of the very
narrow domain of pharmacy, the relations of drugs to agricul-
ture, to domestic economy, and to various industries, should
also be permitted to be indicated, in order to illustrate the part
played by them among existing commodities.

A thorough historical representation of pharmacognosy in
this sense is still wanting; the preparatory labors that have as
yet been brought to light admit of the following preliminary
survey."

1. The earliest application of the products of organic nature
for medicinal purposes, as also for purposes of fumigation, re-
fers to those countries where a higher intellectual life first
became developed. In Egypt, numerous monuments of the
earliest times have been preserved, which prove an acquaintance
with a number of drugs at a very remote period of antiquity.
Illustrative representations on temple walls, which originated in
the seventeenth century B. C., inform us of Egyptian sea voyages
to the provinces of northeastern Africa and Arabia, which were —
in part undertaken in order to procure gum arabic, frankincense, e

1 For appropriate descriptions we are also indebted to Schir, “Die |

Altesten Heilmittel aus dem Orient.” Schaffhausen, 1877, pp. 24 and .

oe - — der Gifte.” Basel, 1883, a 48.

18 TREATMENT OF THE SUBJECT-MATTER.

and myrrh. It is, indeed, possible that through these primeval
trade relations spices and medicinal substances found their way
to Egypt’ even from southern and eastern Asia. In the
Inscriptions of temples,” and in the rolls of papyrus, frequent
reference is made to such things, the correct interpretation of
which is still in part uncertain; only a few of them have actually
been brought to light from burial-vaults.* Especially in the
oft-recurring, but not corresponding recipes for Kyphi, a medi-
cinal compound which served for manifold purposes, there occur
a large number of drugs, as, for example, mastic, cardamom,
curcuma, ladanum (resin of Cistus ladaniferus) and fenugreek."
Tt will only be after a still more extended investigation of
Egyptian antiquities that the extent of the knowledge in ques-
tion among the ancient Egyptians can be established. Through
their highly developed knowledge of agriculture they also
engaged in the cultivation of several of the plants which here
come under consideration, as, for example, coriander, fenugreek,
flax, poppy, ricinus, and sesame.*

‘ Diimichen, ‘* Die Flotte einer agyptischen K6nigin,” 1868, and

*‘ Historische Inschriften,” 1869. Mariette-Bey, ‘‘ Deir-el-Bahari,”
1877. Flickiger, ‘‘ Pharmakognosie,” second edition, pp. 6, 35, 41, 560,
935. Schumann, ‘‘Zimmtlander,” Gotha, 1884 (Supplement No. 73 to
Petermann’s Mittheilungen).
_ _* Dimichen, “ Das Salbélrecept aus dem Laboratorium des Edfu-
Tempels.” Zeitschr. fiir agypt. Sprache und Alterthumskunde, Decem-
ber, 1879.
_ * Braun (Ascherson and Magnus), “Ueber die im Museum zu Berlin
aufbewahrten Pflanzenreste aus altigyptischen Grabern.” Zeitschrift.
fiir Ethnologie, IX., Berlin, 1877, pp. 289 to 310, and Schweinfurth,
‘‘ Ueber Pflanzenreste aus altdgyptischen Grabern.” Ber. d. deutsch.
botan. Ges., IIT. (1884), p. 351.

_. * Lepsius, in the Zeitschr. fiir Agypt. Sprache und Alterthumskunde,

October, 1874, p. 106: « Kyphirecept aus dem Papyrus Ebers.”
*Compare further Thaer, “Die altdgyptische Landwirthschaft,”
Berlin, Parey, 1831, 4to, pp. 36, with six plates. Unger, ‘‘ Botanische
Streifziige auf dem Gebiete der Culturgeschichte,” Sitzungsberichte
der Wiener Akademie, 1857 to 1859, especially in volume 45, “Inhalt —
eines gyptischen Ziegels an organischen Kérpern,” and volume 54

_ (1866): “Ein Ziegel der Dashurpyramide in Aegypten nach seinem —

_ Inhalt an organischen Einschliissen.” Sehweinfurth, ‘ Blumen-

*

HISTORY. 19

2. The industrious trading nation of the Pheenicians,’ and
through them probably the Israelites, were as well acquainted as
the Egyptians with the above-mentioned drugs, to which from
the Old Testament Scriptures may still be added, aloes, cinna-
mon, coriander, saffron, ginger, olive-oil, sugar, and pepper.®
In their religious ceremonies these people evidently employed
aromatic substances in large amount. Drugs which at that time:
were exceedingly highly prized, but which for a long time past
have become completely obsolete, were Radix Costi* and the
Aloeswood of Aquilaria Agallocha, Roxburgh.‘ The high es-
teem with which these two aromatic substances were regarded
from that time until the eighteenth century is scarcely intelligi-
ble to us: and in India and China it still continues undiminished.

3. The Chinese were also, without doubt, at a very early
period, familiar with the medicinal substances which were native:
to that country, as, for instance, with camphor, with remedies:
from their animal world, and with such from the mineral king-
dom. Since cinnamon was certainly exported in the earliest
times, it may be presumed that foreign drugs were probably at.
that time also imported into China. The respective ancient lit-
erature of this country, however, has still been too little seruti-
nized to afford reliable information regarding these conditions.
schmuck agyptischer Mumien,” Gartenlaube, Leipzig, 1884, 628 and loc.
cit. (note 3). [Also, particularly, Woenig, ‘“‘ Die Pflanzen im alten
Aegypten,” Leipzig, 1886. F. B. P.]

1 Flickiger, ‘‘ Pharmakognosie,” pp. 119, 120, 560.

* The plants mentioned in the Bible have led to widely-extended dis-
cussions. It is sufficient here to name the following modern writings.
relating to the subject: Cultrera, ‘* Flora biblica ovvero spiegazione delle
piante menzionate nella S, Scrittura,” Palermo, 1861; Ursinus, ‘‘ Arbore-. |
tum bibl. c. contin. hist. plant. bibl.,” November, 1865; Duschak, “‘ Zur
Botanik des Talmud,” Pest, 1871; Hamilton, ‘‘ Botanique de la Bible,”
Nice, 1872; Smith, ‘‘ Bible Plants, their History, a Review of the Opin-
ions of Various Writers regarding their Identifications,” London, 1878;

Wilson, ‘‘ The Botany of Three Historical Records (Genesis, New Testa-
ment, Assise of Weight and Measure),” Edinburgh, 1879; leunios ore a

d@ Histoire naturelle de la Bible,” Paris, 1885, 4to, pp, 116, 112 plates. —

3 Fliickiger, ‘‘ Pharmakognosie,” pp. 444, gees sopmiae “Materia
_ Medica of Western India,” 1884, P. asia? poeta
_ 4 Flickiger, loc. cit., 195,

20 TREATMENT OF THE SUBJECT-MATTER.

So much is certain, however, that the principal work of the
Chinese, the herbal Pen ¢’sao kang mu, which is, indeed of a
much later date, is based in part upon very much older sources.’
The highly developed condition. of popular medicine of this
people,’ which adheres so tenaciously to primeval customs, refers
to a period of very remote antiquity. For information relating
to many of the pharmaceutically important plants of China,
pharmacognosy is indebted to the great Venetian traveller,
Marco Polo (toward the close of the thirteenth century), as
also in the eighteenth century to the missions of the Jesuits in
China.* |
That an early acquaintance with medicinal plants and
drugs existed in Japan has not yet been proved, but it may cer-
tainly be assumed, for instance, as regards menthol, ‘‘ Hakka.”
4, As to how far this is the case with regard to India has like-
wise not been established. Sanskrit literature possesses in
“Susruta” and “Charaka” information relating to medicinal
substances, which are probably in part of more ancient origin;
but recent investigation assigns to these writings a much less
ancient date.“ The extent of the pharmacognostical knowledge
of Indian antiquity therefore requires more exact investigation; °
the application in India of many medicinal substances and pro-
ducts of the vegetable kingdom adapted to fumigation, such as
awhite sandal-wood, camphor, cinnamon, and cardamon may cer-

1 Fliickiger, loc. cit., p. 1012.

* Hanbury, ‘Science Papers,” 1876, 211 to 272. Fliickiger, Archiv der
‘Pharmacie, 214 (1879), p. 9.

*The activity of this order deserves further mention in connec-
tion with the history of some other drugs, as, for instance, the cinchona
‘barks, ginseng root, maté, and ignatia seeds. The sassafras tree appears
to have been brought by Jesuits from Canada to France; the earliest in-
formation regarding the tape-worm remedy koosso probably likewise
originated from a member of this order. In Rome, Manila, Paris, and
South America they maintained pharmacies, which were probably al-
ways conducted by Jesuit friars,

* Compare Flickiger, ‘‘ Pharmakognosie,” p. 1020.

> Only a few points of guidance in this direction have as yet been
_ Offered by Lassen’s ‘‘ Indische Alterthumskunde,” Bonn, 1847-1852.

HISTORY. ME

tainly be traced to a very remote period. It may be presumed
that musk has also been in use there for a very long time.

5. The centuries representing the flowering period of Greek
and Roman civilization considerably increased the number
of medicinal substances, not only through such as were obtained
from the region of the Mediterranean, but also through others
of Oriental origin. Among these are especially: Amygdale
dulces, Bulbus Scille, Cantharides, Carice, Castoreum, Cortex
Granati,' Euphorbium, Fructus Anisi, Fructus Cardamomi,'
Fructus Feniculi, Fungus Laricis, Galle, Herba Sabine,
Indigo, Mastiche, Opium,’ Piper longum, Radix Glycyr-
rhize, Radix Rhei (?), Rhizoma Filicis, Rhizoma Tridis,
Sandaraca, Scammonium, Semen Feni greci,’ Semen Lini,
Semen Sinapis, Succinum, Siliqua dulcis, Succus Glycyrrhize,
Terebinthina and Tragacantha.

That the number of plants employed in classical antiquity
was very considerable is shown particularly in the writings of

-Dioscorides and Pliny,’ to which the following centuries until
the close of the European middle ages continually refer, and
indeed, almost without any advancement, on their own part, of
the existing knowledge. Many plants of the Italian flora which
are employed medicinally have often been thoroughly considered
by Roman writers on agriculture, namely, by Cato, Columella,
Palladius and Varro. Columelia * (in the years 35 to 65 a.D.)
had also made observations in Spain and Syria. The principal
contents of their writings, so far as they come into consideration
here, are to be found compiled in a very complete and systemati-
cally arranged form in Magerstedt’s ‘‘ Bilder aus der rémischen

Landwirthschaft.”* Hehn also, though with much more spirit _

1 These might have perhaps also been quoted under division 2, page 19.

? To the editions of Pliny mentioned in Fiiickiger’s ‘‘ Pharmakognosie,’””
pp. 997 and 1,014, the new translation by Wittstein may be added. |

* Fliickiger, loc. cit., p. 991, etc. Compare also, Meyer, ‘‘ Geschichte —
der Botanik,” Bd. I. and II., Kénigsberg, 1854, 1855.

4 Heft IV., Sondershausen, 1861: ‘‘ Die Obstbaumzucht der Romer;” i

Heft V. (1862): ** Der Feld-, Garten- und Gemiisebau der Rémer;” oe ae

(1863): “‘Die Bienenzucht und die Bienenpflanzen der Romer.” The
author does not enter upon the consideration of the —— of sana :
plant names. S

22 TREATMENT OF THE SUBJECT-MATTER.

and taste, incidentally touches upon this subject in his publica-
tion: “‘ Kulturpflanzen and Hausthiere in ihrem Uebergang
aus Asien nach Europa.” Berlin, 4th edition, 1882. <A very
remarkable light is thrown upon the intercourse of the Occident
with the Orient through the Periplus of the Erythrean (Red)
Sea. This survey of the coast of the East Asiatic-Indian Ocean,
which was accomplished in the first century after Christ, enu-
merates a number of the commodities: which were to be met
with in those ports, and among these many of pharmaceutical
interest, such as myrrh, Sanguis Draconis, Styrax liquidus,
sandal-wood, pepper, frankincense, and saffron.

The continued importation of the Indian spices mentioned
under division 2, page 19, is proved by a list dating from the
years 176 to 180 a.p. In this the commodities are enumerated
which arrived from the Red Sea,” and which were subject to the
Roman duty at Alexandria.

It is quite natural that even at that time the art of adultera-
tion was also applied to drugs. A single sentence from Pliny
is amply sufficient for the confirmation of this fact; for, when
treating of saffron, the well-versed Roman encyclopeedist em-
ploys the expression: “ adulteratur nihil zeque.”” ®

Even with regard to the relation of prices of some few drugs
in those early times, Pliny has furnished us some information.
Thus, for example, a pound of black pepper was estimated at 4,
white pepper at ?, long pepper at 15, and indigo * (also named as

_* Enumerated in Meyer's “Geschichte der Botanik,” II. (1855), p. 85.
Compare also, Flickiger, loc. cit., p-. 1,013.

2 Tbid., p. 985.

* Flickiger, loc. cit., p. 740.

* Pliny, who in such matters was certainly quite inexperienced, must
have had an attentive observer as a source for his statements here
(Nat. Hist.,” XXXV., 27; page 470 of the edition" by Littré). After a
good description of indigo (indicum), heating of the same is recom-
mended, in order to test the vapor and the peculiar odor: ‘‘ probatur
carbone; reddit enim, quod sincer est, flammam excellentis purpure et,
dum fumat, odorem maris.”

_ When treating of verdigris, Pliny (XXXIV., 26) also resorted to a
chemical reaction, in order to recognize in it an admixture with iron;

HISTORY. 23

a medicament) at 20 denarii. A larger number of prices is
given in Diocletian’s “ Edictum de pretiis rerum venalium,” of
the year 301 A.p., which, indeed, applied more to articles of
food and other indispensable necessaries of life than to spices
and medicinal substances, and was also issued orly for the eastern
part of the Roman empire, not for Europe.’ The edict men-
tions, among other things, almonds, hemp-seed, figs, fenugreek,
linseed, olives, mustard, sesame seed, and grapes.

6. During the decline of antique civilization the fostering of
the sciences passed into the care of the Arabs, who also seized
upon the medical traditions of antiquity, and were enabled to
widely extend them, through their geographical position, from
the highlands of Asia and India to Spain and northern Africa,
as also, through their frequent connections with India, occasion-
ally to enlarge and expand the same. The most eminent repre-
sentatives of Arabic medicine in the tenth and eleventh centu-
ries, Alhervi, Avicenna, Mesue, Serapion, and others, enriched
materia medica with some Asiatic drugs, such as tamarinds,
nux vomica, cubebs, senna leaves, rhuburb, camphor, and worm-
seed (?), and, through the introduction of formulas for medicinal
preparations, exercised an exceedingly lasting influence upon
pharmacy, even in the Occident. With regard to many drugs
from distant lands, the Arabians at an early period received in-
formation through travellers, or from historical and geographical
authors,” such as Khurdadbah, Istachri, Masudi, Idrisi, and
Ibn Batuta. Ibn Alawan reported in the twelfth century upon
the flourishing condition of agriculture among his people in
Spain, from which also dates the cultivation of saffron, still
continued there at the present day. The most prolific informa-
tion, however, chiefly from earlier and often much older sources
of Arabic literature, has been brought together by Ibn Baitar
in his large encyclopedia of simple medicinal remedies and

one should employ for this purpose paper soaked in nutgall—thus prob-
ably the earliest application of test-paper !

1 Fliickiger, loc. cit., p. 997. Compare further, Burckhardt, “Die woe as

Zeit Konstantins des Grossen.” Leipzig, 1880, p. 62.
? Fliickiger, loc. cit., p. 107, Note 10. _

24 TREATMENT OF THE SUBJECT-MATTER.

foods.* The critical examination and often very difficult inter-
pretation of many statements relating to this subject contained
in Arabic literature, which is constantly progressing, admits of
the hope of some very remarkable revelations in this respect.

A pharmacentical manual of Aboul Mena, called Cohen el At-
thar (Priest and Apothecary), who lived in the thirteenth century
at Cairo, has not yet been printed.?

%. In the far West, these antique recollections were likewise
partly revived at the same time in civil and ecclesiastical
quarters. Thus the Emperor Charlemagne, through special
edicts, issued in the year 812, ordered the cultivation, north of
the Alps, of a series of long-known useful and medicinal plants,
among which may be mentioned :* Althea, Amygdalus, Anisum,
Coriandrum, Cydonia, Feniculum, Iris (Gladiolus), Levisti-
cum, Mentha, Petroselinum, Rosmarinus, Ruta, Sabina, Salvia,
and Sinapis. It is worthy of note that the following useful
plants, which are indigenous to Italy or generally cultivated
there, are wanting in the “ Capitulare,” or the principal one of
those Imperial edicts, viz.: Glycyrrhiza, Inula Helenium,
Lavandula, Punica Granatum, and Thymus vulgaris. In the
architectural plan of the convent of St. Gall, which was drawn
in the year 820, but was not executed, the place which the
medicinal plants were to occupy in the garden was designated,
perhaps in compliance with the ‘ Capitulare.” *

In the library of the University of Wurzburg, Germany, a
recipe for a mixture of powders, ‘ contra omnes Febres et contra
omnia venena et omnium Serpentium morsus et contra omnes
angustias cordis et corporis,” has been preserved which is perhaps.
a century older than the preceding. This curious manuscript

'Fliickiger, loc. cit., p. 1,003. 6
_ * According to Leclerc, ‘Histoire de la médecine arabe,” IT. (Paris,
1876), p. 215, it appears very worthy of notice.

_* The complete list is given by Pertz, ‘‘ Monumenta Germanize his-
torica,” legum tom. I. (1835), p. 186, and also in Meyer’s ‘‘Geschichte
- Botanik,” IIL, p. 401. Compare farther, Fliickiger, loc. cit., p-
1,005. ee

HISTORY. 25

is probably one of the oldest monuments of popular medicine in
Germany; init are mentioned Pimpinella and Galanga, both, to
the best of our knowledge, for the first time.
~ 8, The determining influence upon the medicine and pharmacy
of the middle ages proceeded frum the medical school of Salerno.
_ Its activity extended from the ninth century until the close of
the middle ages, although the school itself was continued, at
least in name, until November 29th, 1811, when it was sup-
pressed by an order of Napoleon. To the medical practitioners
who taught there, and in the not very remotely located Bene-
dictine monastery of Monte Cassino, is due the credit of having
transmitted the medical art of the Arabs. Thus the stock of
esteemed remedies of the Occident was increased by a number
of new medicinal substances, or at least such as were previously
but slightly accessible. To these belong Ammoniacum,' Asa-
Setida,’ Benzoinum, VCamphora, Caryophylli, Cinnamomum
zeylanicum, Cortex Aurantiorum, Cortex Limonum, Cubebee
(as a spice; first used medicinally since 1813), Folia (or prob-
ably at first only Siligue) Senn», Fructus Cocculi, Fructus
Colocynthidis, Galbanum,'’ Herba Canmabis, Lignum Sandali,
Macis, Moschus, Radix Rhei, Resina Draconis, Rhizoma Curcu-
me, Rhizoma Galange, Rhizoma Zedoarie, Semen Myristice,
Semen Strychni, Styrax liquidus, Tamarindi, and Salep. co
The writings of the most prominent Salernitans, namely those
of Constantinus Africanus, Macer Floridus, Nicolaus Prapo-
situs, Platearius, Arnaldus de Villanova, and especially the
** Regimen sanitatis Salernitanum ” must be studied in order to
obtain an insight into the science of medicine of the middle
ages. A very complete enumeration of the Simplicia and Com-
posita of that time is given by Renzi in a list published under
the name of Alphita, which is opti to belong to the thir-
teenth century.”

ae Presumably known at an earlier period.

* The history of the School of Salerno, which is not yet ssohaaoiie — :
elucidated, has been briefly but very appropriately described by Hander-

son, in the publication mpentioned in Pichignrs: sj Sen o
p. 1,018, oe

26 TREATMENT OF THE SUBJECT-MATTER.

9. While the erudition of southern Italy interested itself in
the maintenance and extension of the sciences of the Orient, the
commercial republics of Italy—Venice, Amalfi, Pisa, Florence,
and Genoa—were enabled through their fleets to obtain from the
far East and South such commodities as were required in medi-
cine as well as for the more refined enjoyment of life and the
improvements in handicraft. Venice, which was by far the
strongest of these commercial states, began already in the
eleventh century to develop the elements of its incomparable
splendor, and to become the central point for the traffic in drugs.
Up to the end of the sixteenth century there flowed into this
city, in the greatest abundance, those spices which were sought
with such extreme eagerness, but the value of which, in our time,
has become very greatly reduced. The greatest importance was
attached to pepper, the commercial history of which vividly
reflects this entire and exceedingly remarkable intercourse of
nations, as, indeed, in the middle ages, pepper represented the
symbol of the entire spice trade.*

The Levant trade of Venice, which for those times was incom-
parably large, also gave an impulse for the establishment, in that
city, of chemical industries, such as the manufacture of sal-am-
moniac, corrosive sublimate, cinnabar, soap, and glass, together
with the bleaching of wax and the refining of borax (Tinkal,
from Thibet) and of camphor.

Through the shrewd participation of the Venetians in the
crusades of the twelfth and thirteenth centuries, their commerce
and influence attained the highest degree of prosperity. Even
the continued warlike conflicts of the Occident with the Orient
must have necessarily also contributed to the knowledge and dis-
tribution of special medicinal substances in other parts of
Europe. Many historians of that time, who visited Palestine,

‘The spice dealers in various countries were known as Piperarii.
Such a ‘ Gild of Pepperers” existed already in London in the year 1345,
and the “Society of Apothecaries” still existing there, which received
its charter from James I., in the year 1617, properly dates back to
_ those pepper dealers. Compare Pharm, Journ., XV. (1884), p. 367; also

Flickiger’s “ Pharmakognosie,” p. 867,

HISTORY. 27

described, with great clearness, for instance, the sugar-cane and
sugar, while others made themselves acquainted with the agrumt
(the fruits of species of Citrus), with licorice, dates, cotton, and
cumin (Cuminum Cyminum).'

It was only subsequent to these voyages that swgar became a
regular article of commerce, chiefly in the hands of the Venetians.
The successful cultivation of saffron in England and France
during the middle ages, and the culture of roses in the province
of Champagne was started, or at least probably received a new
impulse, through the crusaders.

In Germany the great Benedictine monasteries, for instance,
those of St. Gall and Fulda, formed the central points of intellec-
tual culture, and likewise supported and distributed botanico-
medical knowledge, even though they did not actually augment
it. The first monastery of this in many respects highly meritorious
order, which was founded by St. Benedict himself, in the year
528, at Monte Cassino, northwest of Naples, and afterward be-
came so greatly celebrated, stood in the eleventh century in
close relation to the medical school of Salerno. This school was
also of influence in determining the intellectual course of St.
Hildegard, who, in the year 1148, became abbess of a convent of
the Benedictine order, founded at her own instigation near
Bingen on the Rhine. To this personage is attributed, although
not with absolute certainty, a work which is very remarkable
from a pharmacognostical point of view: ‘‘ Subtilitatum diver-
sarum naturarum creaturarum libri novem,” and is presumed to
have been written about the year 1178. By the enumeration of
a number of indigenous plants, to which some prominent
characteristic, or often also the German name,? was occasionally
added, the book of Hildegard, frequently designated as ‘‘ Phy-
sica,” reveals itself as a truly German production. Of incom-

* Compare the respective articles in the ‘‘ Pharmacographia” and in

Flickiger’s ‘‘ Pharmakognosie;” also Meyer, ‘Geschichte der Pole 2

nik,” IV., p. 110; Heyd, *‘ Levantehandel,” IT., p. 670.

‘ Couper also Pritzel and Jessen, ‘‘ Die dentachens Volksnamen a
Pflanzen. Aus allen Mundarten and Zeiten BEE ie a bic Han-
ee 1882. os z

28 TREATMENT OF THE SUBJECT-MATTER. |

parably greater importance, however, is the Dominican monk,.
known as Albertus Magnus, who from the year 1260 to 1280:
was Bishop of Regensburg (Ratisbon). In his books ‘‘ De
Vegetabilibus,” numerous medicinal plants and drugs have re-
ceived for the most part very intelligent treatment, though this.
is largely based upon other sources of information, including
Arabian.

The history of indigenous, and probably also foreign medici-
nal plants may be further pursued with the aid of the diction-
aries and glossaries of the middle ages. Compilations of this
kind’ were also specially made use of in attempting, quite in
accordance with medieval custom, to identify in the indi-
genous flora plants of classical antiquity. From a mercantile
point of view, very valuable information is afforded by the ordi-
nances and lists of the departments of customs? of that time, as.
also especially by the trade-books of the Venetians and Floren-
tines.* The extraordinary importance of the commercial inter-
course with the East during the middle ages has finally been
described in an exhaustive and captivating manner by Heyd,*
who not only represents the political and economical sides of
the subject, but also gives detailed accounts of the most im-
portant objects of that remarkable commercial era, namely:
aloes, aloes-wood, ambergris, balm of Gilead, benzoin, Brazil-
wood, camphor, cloves, frankincense, galanga, galls, ginger,
indigo, kermes (a coloring insect afterwards supplanted by cochi-
neal), lac, mace, musk, nutmegs, pearls (both as a medicament.
and ornament), pepper, precious stones (quite a series of these
were employed as medicinal agents), rhubarb, saffron, sandal-
wood, and tragacanth. All that the civil and ecclesiastical
travellers and historians of those centuries were able to report
relating to these articles has been made use of in the proper
place by Heyd, in his careful examinations.

* Flickiger, ‘‘ Pharmakognosie,” pp. 107, 330, 642, 688, 713, 892, 898.

? Tbid., pp. 781, 983.

3 Ibid., pp. 1,011, 1,012.

ne Geschichte des Levantehandels im Mittelalter,” 2 vols., Stuttgart,

HISTORY. 29

The precious goods from India were obliged, as a rule, to take
their way through the Red Sea and through the domain of the
Egyptian sultans, for which reason the Italian merchants of
the middle ages were compelled to bestow the greatest attention
upon their relations to those rulers. The ambassadors of the
latter to the doges of Venice, toa Venetian queen of Cyprus, and
to Lorenzo de’ Medici in Florence, brought rare drugs of the
Orient to Europe, as, for instance., aloes-wood,' civet, Mecca bal-
sam,’ myrobalans,* opium, and sugar, the latter of which was at
that time, or in the second half of the fifteenth century, still a
rare article. Thus in the year 1461 denzoin was brought for the
first time to Venice.*

It was only in case of necessity that the Italians permitted
their goods from the Orient to take the much longer route
through the Persian Gulf, or even through Central Asia to the
Black Sea, instead of through Egypt.

The intercourse with the Levant consisted quite especially in
the importation of numerous Asiatic products; there were but
few articies which the Italians, inhabitants of Southern France,
and Catalonians, for example, had to ship to Alexandria. The
Venetian statesman, Marino Sanudo,’ in the year 1307, mentioned
as such articles almonds, honey, hazelnuts, mastic, and saffron.

The most’ prominent Florentine houses also conducted the
Levant trade with consummate mastery. Thus, for example,
in the first half of the fourteenth century, the large trade asso-
ciation which bore the name of the leading house, Bardi in
Florence. It was in their service that Pegolotti, about the year
1340, wrote that very remarkable trade-book, ‘‘ Pratica della
mercatura,”*® which gives the most instructive informatio nre-
garding the commercial relations, coins, weights, measures, and .

1 Flickiger, ‘‘ Pharmakognosie,” p. 195, and this work, p. 19.
* Tbid., p. 180.

8 Thid., p. 244.

“Tbid., p. 118. .

5 Fliickiger, ‘‘ Pharmakognosie,” p. 1,018.

* Compare further Heyd, loc. cit., i., p. xiii,

30 TREATMENT OF THE SUBJECT-MATTER.

trade products of that time, and is far superior to similar publi-
cations of a later date.

A less comprehensive, but nevertheless remarkable book, is
that of the Venetian Pasi or Paxi, ‘‘ Taripha,” * of the year 1503.

10. Sources of information are not wanting for judging of the
supply of commodities, and also of preparations which were to
be met with in the fifteenth century in German pharmacies.
The “‘ Frankfurter Liste’ and the ‘‘Nérdlinger Register” ex-
hibit quite a series of drugs * which, at that time, were actually
kept in stock; and the inventory of a pharmacy in Dijon of the
year 1439 is also known.

The conquest of Constantinople by the Turks on the 29th of
May, 1453, as also the discovery of a sea passage to India in
. May, 1498, are events which, by its causes and results, led to
the gradual but almost complete downfall of the Levant trade,
and especially to the exhaustion of the resources of Venice. The
discovery of America completed the enormous revolution. The
age of great discoveries could at first gain no greater number of
new, important drugs from the vegetable kingdom of Asia,
which had already been the source of supply for so longa period.
The discovery of a sea passage around the Cape of Good Hope,
however, resulted in a much more abundant importation of

familiar commodities of that character. Furthermore, the

scientific world now finally received more exact intelligence relat- |
ing to the celebrated products of India. For this information, —
thanks are primarily due to a Portuguese physician, Garcia de

- Orta (Garcias ab Horto), who resided for thirty years in Goa,

India. His discourses upon Indian drugs, which appeared at

Goa in the year 1563, form a highly agreeable contrast to the

mostly confused and altogether too meagre notices of the Arabs

and of Marco Polo, the great traveller, whose reports are other-

wise so valuable.

The most substantial advancement of pharmacognostical

1 Plickiger, ‘ Pharmakognosie,’ "ps 3,08
_ * See page 3, note 1.
3 Schweizerische Wochenschrift fir Pharmacie, 1873, Nos. 6, My 8.

HISTORY. 31

knowledge at that time in Europe proceeded from Clusius,
who, after obtaining a broad preliminary education, was engaged
in- botanical pursuits, principally at Vienna, and finally, until
the year 1609, at Leyden. As early as the year 1567, this dis-
tinguished man published a Latin translation of Garcia’s ‘‘ Colo-
quios” under the title ‘* Aromatum et simplicium aliquot
medicamentorum apud Indos nascentium historia.” Clusius
freed the work of the Portuguese of its labored diction, omitted
many useless and speculative additions of the author, and, on the
other hand, added valuable notes comprising personal experi-
ences and observations, Clusius was already acquainted with
the cola nut, star-anise, gamboge, winter’s bark, sabadilla, and
vanilla.

The most instructive and abundant information relating to the
Indian flora, combined with illustrations of long-known, celebrated
medicinal plants, is presented (though, indeed, at a much later
period) in the twelve folio volumes of Rheede’s ‘‘ Hortus indicus
malabaricus,” which appeared at Amsterdam between the years
1678 and 1703, as also in the ‘“‘ Herbarium amboinense,” by
Rumphius (Rumpf), in six volumes, Amsterdam, 1741-1755.
These grand achievements of the Dutch have been followed in
modern times by the magnificent works of the English botanists.’

Less laudable, even though it is intelligible when considered
in the light of that period, appears the commercial policy of the
Dutch in the seventeenth and eighteenth centuries, through
which they were able to retain for themselves the exclusive pos-
session of Indian spices, particularly of nutmegs, cloves, pepper,
and cinnamon, occasionally even advancing the prices by the
destruction of these articles when the stock had accumulated to
too great an extent.

In the first half of the sixteenth century the knowledge of
crude medicinal substances had meanwhile been much promoted
in Germany by the talented teacher Valerius Cordus (1515-

1544), after whose premature death his most important writings _

were first published, in the year 1561, by his friend and associate

1 See Archiv der Pharm., 222 (1884), p. 249.

32 TREATMENT OF THE SUBJECT-MATTER.

Conrad Gesner. Cordus, in his annotations to Dioscorides and
in the “Historie Stirpium (s. Plantarum),” in opposition to
those of his predecessors who were involved in the study of an-
tiquity, recorded good individual observations, and described
the drugs much more carefully, particularly according to his
personal inspection; thus, for example, Vuz vomica, Cocculus
Indicus, and Lignum Guaiaci Cordus has rendered valuable
service to practical pharmacy by the compilation of a dispensa-
tory* in the years 1542 and 1543, by request of the authorities
of the city of Nuremberg. -

Under the title of « Horti Germanix,” Gesner, in the year
1560, gave interesting accounts of medicinal and useful plants,
or otherwise noteworthy species, which he or his friends, mostly
apothecaries,* cultivated in Germany. Gesner permitted this
publication to appear as an appendix to that of his friend
Cordus.

At a still earlier period, the flora of the surroundin g country
had been brought into requisition, but without much critical dis-
crimination, as was the case with the widely diffused, popular
book, ‘‘ Hortus Sanitatis.” The same degree of consideration was
accorded to the curious work on distillation by the surgeon
Hieronymus Brunschwig, of Strassburg, first published there in”
the year 1500, and which gave the crudest instruction relating

1 A still earlier monographic publication relating to Lignum Guaiaci,
which, for those times, is very worthy of notice, and written in excel-
lent Latin, likewise emanates from the pen of a prominent German, the
Knight Ulrich von Hatten: “ Virichi de Hutten Eq. de Guaiaci medi-
cina et morbo gallico liber vnus,” 4to, 26 chapters, unpaged. The first
of the numerous editions, printed in the year 1519 in Schaffer’s house at
Mayence, bears at the end of the work a woodcut likeness of the author. s
This publication, by its accuracy and instructiveness, excels most of
its contemporaries in the description of new medicinal substances.
Hutten describes the habitus of the tree, the wood, bark, and resin, and
lastly the applications. ;

* Flickiger, ‘ Pharmakognosie,” p. 994.

* Among foreign apothecaries Gesner mentions also the meritorious _
Peter Coudenberg, of Antwerp (Flickiger, loc. cit., p. 458); also —

Follietus, of Vevey, on Lake Geneva, Switzerland.

HISTORY. 33

to the preparation of distilled waters, whereby all conceivable
herbs, and things of an entirely different nature, were brought
together in a senseless manner. Here and there Brunschwig
inserted a notice worthy of consideration, relating to a plant,
for instance, one from which it may be concluded that at the
end of the fifteenth century /avender was cultivated in Germany,
and anise near Strassburg. Brunschwig’s book corresponded,
moreover, so much to the views of that time, that in the year
1597, an Augsburg physician, Adolf Occo, introduced one hun-
dred and forty different Aque@ destillate into the “‘ Pharmacopea
Augustana.”

Subsequent to the year 1531, Otto Brunfels had numerous
figures of plants prepared at Strassburg, some of which are very
good, and are still worthy of notice as the earliest good exam-
ples of the application of woodcuts to botanical purposes. His
descriptions are, indeed, of an incomparably lower order than
those contained in the herbals of his direct successors, Cordus
(see p. 32), Bock (Tragus), Fuchs and Gesner. ll these
German “fathers of botany” of the sixteenth century, though
not exclusively physicians, have also, through their writings,
furnished pharmacy with a more exact knowledge respecting the
indigenous medicinal plants, a large number of which were,
at that time, kept at the pharmacies. It was also at this period
of general reformation that Brunfels (in his ‘‘ Reformation der
Apotheken”’) brought forward a list of drugs, prepared without
much judgment, but containing those which appeared to him
worthy of commendation, under the title ‘‘ Ein gemeyne beset-
zung einer Apotheken, von Simplicibus.”

11. To the discovery and settlement of America we are in-
debted for a number of substances already in use by the
people of that country, which soon found their way to Spain,
Portugal, and other parts of Europe. The writers of those two
countries, Gonzalo Fernandez (Oviedo), Monardes, and Hernan-
dez, who were the first to devote attention to the natural pro-
ducts of the new world, also began to describe medicinal plants’
more accurately than had been done before.

Thus America gradually furnished: Balsamum Copaiva,

'3

34 TREATMENT OF THE SUBJECT-MATTER.

Bals. peruvianum, Bals. tolutanum, Cascarilla, Cinchona,
Elemi, Folia Nicotiane, Fructus Capsici, Fr. Pimente
(Amomt), Fr. Sabadille, Lignum Hematozryli, Lig. Fernam-
buci (Brazil wood), Lig. Guaiaci, Lig. Quassie, Lig. Sassa-
fras, Radix Ipecacuanhe, Rad. Sarsaparille, Rad. Senege,
Rad. Serpentarie, Resina Guaiaci, Semen Cacao, Sem. Saba-
dille, Tuber Jalape, and Vanilla.

At a later period (1636-1641), the German geographer and
astronomer Marcgraf, and the Dutch physician Piso, were jointly
commissioned by the government of the Netherlands, to pro-
mote the knowledge of Brazilian medicinal plants. Previous to
this they had already furnished descriptions of Copaiva, Elemi,
Jaborandi, Ipecacuanha, Matico and Tapioca.

12. The re-awakening of the sciences at the beginning of the —
new era also led to the gradual foundation of scientific botany,
and in the seventeenth and eighteenth centuries to the medici-
nal employment of a large number of plants of Central Europe,
as well as of some few drugs from other parts of the world,
which then arrived in Europe for the first time. Of the
long series of plants and substances which are properly to be
enumerated in this connection, there may be mentioned: Cate-
chu, Cortez Frangule, Flores Arnice, Flores Chamomilla, Folia —
Aconiti, Fol. Digitalis, Fol. Laurocerasi, Fol. Menthe piperite,
Fol. Toxicodendri, Fol. Uve ursi, Herba Chenopodii ambrosior-
dis, Herba Cochlearie, Herba Conii, Herba Hyoscyami, Herba
Lobelia, Fructus Anisi stellati, Gummi-resina Gambogia, Kino,
Lactucarium, Lichen islandicus, Lycopodium, Oleum Cajuputt,
Ol. Rose, and other distilled oils, Radix Calumbe, Rhizoma
Caricis, Rhiz. Filicis, Saccharum lactis, Secale cornutum,
Tuber Chine and Tuber Colchici.

13. In the cities of Germany, the stock, and especially the —
prices of drugs in the pharmacies, were regulated by official
ordinances, while pharmacies were also openly conducted on the
part of the magistrates, or leased by them, In no other country —
at that time were the affairs of pharmacy the subject of such
thorough official attention. The manuscripts relating hereto,
admit of a complete insight into the store-rooms and laboratories

HISTORY. 35

of the pharmacies from the beginning of the sixteenth century.
Such, for example, are the inventories of the senatorial phar-
macy at Brunswick (Braunschweig) of the years 1518 to 1658,"
the catalogues of several other pharmacies, and a large number
of price-lists which were issued in the course of the sixteenth,
seventeenth, and eighteenth centuries in all German countries
for the use of the apothecaries. It was rarely, however, that
the latter were mentioned as participants in the official control,
reference being usually made only to physicians and officials.

By the aid of these manuscripts, many of which still remain
buried in archives, the introduction and gradual distribution of
many drugs may be traced, as, for example, some of those
derived from America. These documents also afford positive
information concerning the extensive cultivation of certain me-
dicinal plants, which has long since been abandoned, as, e. Oss
the Angelica near Freiburg, in the old district of Breisgau,
Baden; Licorice, near Bamberg; Saffron, in England, Germany,
and Austria, and Cassia obovata in Tuscany.

An insight is also afforded into the obscure department of
adulterations, to which drugs, as well as articles of food and luxu-
ries, were subjected in the middle ages, no less than at any
other time (see p. 22). The magistrates of German and Italian
cities resorted to the most extreme police measures, even pun-
ishment by death,’ for such offences, and authorized physicians
to watch the apothecaries. In the ordinances accompanying the
above-mentioned price-lists, and in special manuscripts, these
regulations are quoted at great length.

The unfortunate apothecary Zanoni de’ Rossi, of Venice, in
the year 1402, when preparing the highly celebrated “ Theriac,”
was detected in the omission of Rhubarb, Amomum, Opoponaz
(“ Pharmacographia,” p. 327) and Saffron, even substituting

1 For a copy of this, one of us(F.) is indebted to the apothecary Dr.
Grote, in Brunswick. Compare also the notice of the latter in the
Archiv der Pharm., 221 (1883), p. 417.

* Flickiger, “‘ Pharmakognosie,” p. 740, Compare further the thor-
ough work by Elben, “Zur Lehre von der Waarenfalschung.” A Ti-
bingen dissertation, 1881, i

36 TREATMENT OF THE SUBJECT-MATTER.

for the latter Safflower (Carthamus tinctorius), while he also
adulterated Musk and the Syrups. The court to which such
matters pertained, ‘“‘ Avogaria di comun,” caused his prepara-
tions to be thrown over the Rialto Bridge into the canal, the
musk was burned, the offender banished from his profession,
placed in prison, and a fine of four hundred gold ducats im-
posed upon him—fortunately, in contumaciam.*

14. The proper study of drugs received an impulse from
the chief emporium of the drug market of the middle ages.* The
Signoria of Venice founded in the year 1533, in her Univer-
sity at Padua, a chair of pharmacognosy, ‘‘ Lecturam Simpli-
cium,” which she filled by the appointment of the physician,
Francesco Buonafede. The University of Bologna, as early as
the year 1534, also received such a “‘Spositore o lettore dei
Semplici” in the person of Luca Ghini, who, in the year 1544,
removed to the University of Pisa. Buonafede, “‘ primus Sim-
plicium explicator,” the first professor of pharmacognosy, dis-
tinguished himself by the foundation of the first botanical
garden, which, in accordance with his proposal, in the year 1545,
was effected by a resolution of the Venetian senate. The land
required therefor was given by the Benedictine Convent, Santa
Giustina, with a proper appreciation of the usefulness of the
purpose. The garden still remains in sight of the magnificent
cathedral of Sant’ Antonio, in Padua. The decree of the senate,
in a very admirable and appropriate manner, gives prominence
to the value of the garden for the study of medicinal plants,
and carefully provides for the management of the same. From
the possessions of the Republic on the islands of Candia and —
Cyprus were to be procured the valuable plants, as also minerals. —

_ ' Cecchetti, ‘‘La Medicina in Venezia nel 1,300,” Archivio Veneto,
xxv. (1883), 376. Ibid., xxviii. (1884), 29, it is also stated that, in the
year 1820, on the Rialto Bridge, inspectors of goods from the Levant
were appointed, ‘‘ Signori sopra le merci del Levante.”

* A description of this drug market is still wanting. As men-
tioned by Fliickiger, loc. cit., p. 634, there existed in Venice, in the year
1506, a chamber of commerce, “Cinque savii alla mercanzia,” whose
acts are still preserved in the great Venetian central archives.

HISTORY. 37

As is evident from Gesner’s “ Horti Germanixw” (see p. 32),
he stood in active intercourse with apothecaries and other friends
who were engaged in the cultivation of officinal plants. The
opportunity was thus afforded of mentioning cinnamon, cloves,
worm-seed, colocynth, and fenugreek, the mother plants of
which were cultivated in a garden at Venice, in the quarter
S. Gervasio.’ But the first public garden for scientific purposes
is that at Padua.

Buonafede also arranged in the botanical garden at Padua the
first collection of drugs, ‘‘ Spezieria,” for purposes of instruc-
tion, in which the dried, crude products of the Levant were
preserved, in order to be employedas standards for the discrimi-
nation of pure and adulterated articles. The garden and the
collection are mentioned as two superabundant sources from
which the most substantial knowledge may be obtained regard-
ing those substances which contribute to the happiness of man-
kind. Notwithstanding these efforts, Buonafede, at the age
of 76 (in the year 1549) lost his position, and, having become
blind, died in penury on the 15th of February, 1558. The pro-
fessorial chair, ‘‘lettura de’ semplici,” was taken in the year
1551 by Gabriel Falloppio.* :

The University garden at Padua was followed in the year 1547
by that of Pisa, in 1567 by Bologna, in 1577 by Leyden, and in
1593 by Montpellier. In Germany the first botanical University
garden was brought into existence in 1593 through the efforts of
the medical faculty of Heidelberg; in the years 1624 and 1625
Ludwig Jungermann ‘established the gardens of Giessen and Alt-
dorf (near Nuremberg). It was first in 1628 that Paris also re-
ceived such an one. In the year 1658 there wasa medical garden

' Flickiger, loc. cit., p. 781. As early as 1330 the subject of a garden
in Venice was agitated, which the celebrated physician Maestro
Gualtieri wished to establish ‘‘ pro herbis necessariis arti sue.” ecm:
Archivio Veneto, xxv. (1883), 375.

2 R. de Visiani, ‘‘ Origine ed anzianita dell’ orto botanico di Padova.” —
Venezia, 1839, 48 pages; and: ‘‘ Della vita e degli scritti di Francesco
Buonafede.” Padova, 1845, pp. 24.

3 Reess, ‘‘ Address of the Prorector.” Erlangen, 1884, 19.

38 TREATMENT OF THE SUBJECT-MATTER.

in Westminster (London), which is presumed to have belonged to
the “‘ Society of Apothecaries.” Prior to 1674, as it appears, the
garden of this corporation was removed to Chelsea, where it still
continues to exist.’ :

15. In the second half of the sixteenth and in the course of the ©
seventeenth and eighteenth centuries, pharmacognosy was prac-
lically applied in the unnecessarily numerous pharmacopeias of
different cities and countries of Europe, the largest number of
which probably appeared in Germany. In this country, phar-
macy had attained a high degree of development, as is shown by
the official price-lists and various other ordinances. The first
official pharmacopeeia, however, entitled ‘‘ Ricettario fioren-
tino,” was published by the city of Florence in 1498. The num-
ber of drugs brought together in these pharmacopeeias, dating
from the sixteenth to the eighteenth century, was so considera-
ble, that but little mention was made of new additions. Beside
the most important of all, the Cinchona-barks (introduced in
Spain about 1640, in England in 1655, and in Germany about
the year 1669), there are to be mentioned, by way of example,
Ipecacuanha (about 1682), Catechu (in 1640), and Senega (1735).

Efforts now began to be made in the direction of the chemical
investigation of drugs, which at the end of the seventeenth and
beginning of the eighteenth century received the attention of ~
numerous physicians and chemists; in Germany, ¢. g., by
Friedrich Hoffmann in Halle (1660 to 1742) and the distinguished
apothecary to the Court, Caspar Neumann of Berlin (1683 to
1737), in Paris by the previously mentioned Geoffroy and
we (see p. 5), and in England by Robert Boyle (1627 to

691).

Among the numerous and important discoveries of Scheele
(1742 to 1786) there are but few which relate to drugs, although
in this connection the discovery and examination of the acids
most frequently occurring in plants was of the greatest im-
portance. Scheele, in 1769, discovered tartaric acid, and in 1776
recognized the distribution of oxalic acid, which had already

' Fliickiger, ‘‘ Pharmakognosie.” 544, Note 6.

HISTORY. 39

been observed by Savary in 1773; in 1784 he first prepared citric
acid in a pure state, and in 1785 separated malic acid. Hydro-
cyanic acid was likewise discovered by Scheele in the year 1783,
although it was not allotted to this celebrated investigator to
also prepare this substance from plants.

Johann Bartholomzus Trommsdorff, who rendered such valu-
able services to pharmacy (from 1790 to 1837), and in 1795
established a Pharmaceutical School (Institute) at Erfurt, de-
voted a portion of his varied labors to the chemical investigation
of vegetable medicinal substances. Notwithstanding these ef-
forts, in his ‘‘ Handbuch der pharmaceutischen Waarenkunde ”
(Erfurt, 1799, pp. 624; third edition, 1822), no essential ad-
vancement of pharmacognosy can be recognized.

The greatest progress in this department was inaugurated in
the year 1816 by the apothecary Sertiirner, through his dis-
covery of the first alkaloid, morphine. Since this brilliant

- discovery, chemical research of the nineteenth century continues
to give expression to the most important facts; and, for the reason
previously intimated (p. 1), it more rarely occurs at the present
time that a plant is able to attain permanent and prominent
importance among valued medicinal agents. As accessions of
the last eight decades are to be mentioned: Carrageen, Cortex
adstringens brasiliensis, Cort. Granati, Cort. Monesie, Flores
Koso, Folia Coca, Galle chinenses, Guarand, Gutta Percha,
Helminthochorton, Herba Lobelie, Herba Mutico, Kamala,
Laminaria, Lupulin, Pengawar Djambi, Rad. Ratanhie, Rad.
Scammonie, Secale cornutum, Semen Calabar, Sem. Colchici
and Tuber Aconiti. The very unequal, and in part very
questionable importance of these drugs is at once apparent. As
examples of drugs which have appeared more recently, and
have directly received the condemnation of science, may be men-
tioned: Lignum Anacahuite (from Cordia Boissiert D. C., a
Mexican shrub), Cortex Condurango, the leaves of Sarrazinia
purpurea, of Peumus Boldo, of Eucalyptus globulus and of
Grindelia robusta, furthermore Cortex Coto, Cortex Quebracho
and Radix Gelsemii. On the contrary, as permanent and im-
portant acquisitions since the year 1873, may be considered:

40 TREATMENT OF THE SUBJECT-MATTER.

Jaborandi leaves, from the Brazilian Rutacea Pilocarpus pen-
natifolius, and the leaves of Erythroxylon Coca, from Peru and
Bolivia.

The zealous endeavors of the eclectic school of medicine of
North America to introduce new vegetable substances from their
flora have also met with but little success.

After the discovery of strychnine and veratrine in the year
1818, of brucine in 1819, quinine, cinchonine and caffeine in
1820, coniine in 1827 (1831), aconitine and atropine in 1833,
Liebig and Wohler, in 1837, recognized in amygdalin the first
representative of another numerous class of vegetable princi-
ples, many of which possess energetic physiological properties.
The chemical investigation of drugs, far superior to the anti-
quated method of estimation, which was mostly based upon exter-
nal characteristics, was thus correspondingly forced to the front.

16. In the mean time, not only the systematic investigation of
the vegetable kingdom, but also its anatomical and physiological
study became established more and more upon truly scientific
principles. In consequence of this advance, the condition of
pharmacognosy could not remain unchanged. Guibourt (1790-
1867), in his lectures at the Paris school, and in his writings (see
p. 5), had already, to some extent, taken a higher standpoint,
and Pereira (1804-1853), perhaps to a still greater degree, in his
large text-book (p. 5), in an introductory address in 1842, and,
furthermore, in his lectures. Schleiden, in 1844, with perfect
comprehension of the subject, demonstrated the importance of
the microscopical examination of drugs, which, in 1847, was
further elucidated in the most brilliant manner by an essay on
the structure of sarsaparilla, accompanied by microscopic
figures. When Schleiden’s “Botanische Pharmakognosie,”
which was written in this spirit, made its appearance in 1857,
the author no longer stood alone. Weddell, in 1845 and 1848,
had become acquainted with the Cinchonas in Bolivia and Peru,
* and in 1849, in his handsome work on this subject, had also
employed the microscope in an admirable manner for the
purpose of distinguishing their barks. Schleiden extended this _
examination to all the Cinchona barks which at that time oc- —

PHARMACOGNOSTICAL SYSTEMS. 41

curred in commerce. To Otto Berg (1815-1866) is due the
credit of having subjected a large proportion of the crude medi-
cinal substances employed in Germany, at least all the more
important ones, to microscopical study. Inthe figurative repre-
sentation of the anatomical structure of organized vegetable
substances he was, indeed, preceded by the no less distinguished
Dutch botanist, Oudemans, 1854-1856. But, in 1865, Berg
followed with the fifty plates, for the most part very true to
nature, of his ‘ Anatomischer Atlas zur pharmaceutischen
Waarenkunde.”

It is in this direction that pharmacognosy of the present day
is being developed, on the one hand in practical connec-
tion with botanical science, and on the other supported by the
constant progress in organic chemistry. The greater the num-
ber of active principles of the vegetable kingdom extracted by
chemical processes, or even built up artificially, the greater the
changes effected in the pharmaceutical importance of the re-
spective drugs, for which, indeed, new interest may again be
specially incited by a more searching botanical examination.

XII. Pharmacognostical Systems.—The larger number of
medicinal substances, even by the most exhaustive treatment,
will only appear of significance from some of the points of view
that have just been mentioned, while in other respects they may
present nothing at all worthy of note. It appears of less im-
portance to treat of these substances in detail than to consider
the order in which they are classified. They have been grouped
in special pharmacognostical systems, in a more or less artificial
manner, by considering either their organological importance,
or, to a greater extent, their medicinal action and most promi-
nent chemical constituents, or by utilizing at the same time all
these principles of division. In opposition hereto, their arrange-
ment in accordance with the natural families of plants, as fol-
lowed by botanists, commends itself. The employment of a
system founded upon this plan is appropriate, because a knowl-
edge of the families of plants may be presupposed, thus scarcely
leaving a doubt respecting the proper position of each drug, and
because it does not admit of the separation of the parts or pro-

42 TREATMENT OF THE SUBJECT-MATTER.

ducts which are furnished by one and the same plant. The
advantages of such a classification are greater than the disad-
vantage, which may be supposed to arise from the fact that by
this arrangement things are found in close association which are
not connected either morphologically or medicinally. The
‘* Pharmacographia,” subsequently mentioned, as also the
**Grundriss der Pharmacognosie” of one of the authors‘ (F.),
presents the drugs of the vegetable kingdom arranged in accord-
ance with the natural families of plants, while the “‘ Pharma-
cognosie des Pflanzenreiches” (by F. A. Fliickiger) bases its
classification more upon the external characters of the drugs.
An excellent example of chemical classification is presented by
Falck’s ‘‘ Uebersicht der speciellen Drogenkunde,” Berlin, 1883,
Dp. 57.

‘The ‘‘Grundriss” adheres to the well-planned system of the ‘‘ Sylla-

bus der Vorlesungen iiber specielle und medicinisch-pharmaceutische
Botanik,” by Eichler, third edition, Berlin, 1883, p. 54.

AIDS TO THE STUDY OF PHARMACOGNOSY.

The consideration of medicinal substances in their various
aspects, as has been indicated, presupposes the possession of
corresponding adjuncts or aids. These are primarily the requi-
site preliminary knowledge of botany, zoology and chemistry,
as also experience in the use of the microscope. With this
general knowledge, and the corresponding degree of skill, phar-
macognostical description is everywhere intimately connected,
as well as with the practice of pharmacy itself. The present
work has not included the chemistry of plants in its plan,
nor is it designed as a complete guide for microscopical study,
and refers, therefore, in this respect to the literary aids enu-
merated under section II., B, page 4%. In addition to the
latter, an oral and practical introduction to the methods of
microscopical examination is expressly recommended. Such
opportunities are easily obtained, and, as in other departments
of applied natural science, practical instruction proves here also
to be of the greatest utility. Those who earnestly enter this
field will soon become incited to zealous labor.

As scientific aids the following are to be considered:

I. COLLECTIONS.

A. Collections of Drugs.—The pharmacies themselves rep-
resent to a certain degree such collections, which are found more
complete, in their adaptation to scientific purposes, in many of
the higher educational institutions, and particularly in such
as are directly subservient to the interests of pharmacy.
The most instructive, and by far the most extensive collection

.

44 AIDS TO THE STUDY OF PHARMACOGNOSY.

of crude medicinal substances from the vegetable kingdom, how-
ever, is contained, in the Museum of Economic Botany,
in the botanical garden at Kew, near London. Less ex-
tensive, since it is only in process of formation, is the
botanical museum in Berlin. Collections which are devoted
to pharmacy in its broadest sense are those of the Pharmaceu-
tical Society of Great Britain, in their own building in London,
the rapidly increasing collections of the General Austrian Pharma-
ceutical Association (Allgemeiner ésterreichischer Apotheker-
Verein) in Vienna, and those of the School of Pharmacy (Ecole
de Pharmacie) in Paris.’ Aside from the museum of the firm
of Gehe & Co., in Dresden, which is worthy of considera-
tion, the want of a large pharmaceutical centre in Germany is
to be regretted. The Pharmaceutical Institutes connected
with the Universities of Germany correspond but little to the
ideal requirements which should be made of a pharmacognosti-
cal institution that is equipped in accordance with the times.
The great international expositions of modern times have pre-
sented, for the time, highly instructive and extensive collections
of crude medicinal substances from different countries, of which
reports have been published,” and in which, e. g., the medicinal
agents even of the Asiatic nations are also thoroughly illustrated.

‘ This applies also, to a greater or less extent, to the museums and cabi-
nets of some of the Schools of Pharmacy in the United States. (F.
B. P.)

* For example, the excellent official reports of the Austrian experts
who were delegated to these expositions. Compare further: Flickiger,
Schweizerische Wochenschrift fir Pharmacie, 1867, 325; also Archiv
der Pharmacie, 214 (1879), 1-48 and 97-136. Paul, Holmes, and Pass-
more, “ Universal International Exhibition,” Paris, 1878; London, 1878,
pp. 198. Schaer, “‘ Botanischer Congress und Ausstellung pharmaceu-
tisch wichtiger Pflanzenproducte zu Amsterdam,” April, 1877. Archiv
der Pharm., 212 (1878), 928. Wittmack, “Die Nutzpflanzen aller
Zonen auf der Pariser Weltausstellung, 1878; ” Berlin, 1879, pp. 112. See
also the reports of Dr. Carl Mohr in the Pharm, Rundschau, New
York, 1885, pp. 57-60, 77-83, 97-100, 126-131, 146-154, 165-170, 198-202,
227-230, entitled: “ Mittheilungen iiber die medizinisch und technisch —
wichtigen Producte des Pflanzenreichs auf der Weltausstellung von
New Orleans.”

COLLECTIONS. 45

B. Collections of plants which are either themselves offici-
nal, or which furnish the crude medicinal or technical substances,
or afford the material for preparations. Botanical gardens
present such plants in the living state with most admirable
selection and arrangement, for example, those of Kew (pages
12 and 44) and Edinburgh, the gardens of the Ecole de
Pharmacie, and of the Faculté de Médecine in Paris, and
those of Berlin, Amsterdam, Vienna, and Palermo.’ In the
garden of Mr. Thomas Hanbury at Mortola, near Mentone,
France, there are also cultivated many pharmaceutically impor-
tant plants.”

Similar and very excellent results have been attained at the
garden of the University of Breslau through the judicious and
untiring efforts of Géppert.* The large Umbellifere of Asia,
which furnish asafetida, galbanum, sumbul and ammoniacum,
are cultivated by Mr. Max Leichtlin in Baden-Baden.

Many plants of pharmaceutical importance are, as yet, to be
had only with difficulty or not at all, or are at least not readily
obtainable in the living state, so that all knowledge concerning
them must be derived from herbaria, or from descriptive and
illustrative representations. Such are presented in the greatest
degree of completeness by the previously mentioned institutes in
Kew, London and Paris; and the botanical museum (section, Her-

1 In the United States, the gardens of Mr. Henry Shaw, at St. Louis,
Mo., are deserving of special mention, while the Congressional garden
and garden of the Department of Agriculture at Washington, the
botanical garden at Cambridge, near Boston, and the Arnold Arboretum
(for woody plants) at Brookline, Mass., likewise present something of
interest. There is also under contemplation the establishment of a large
botanical garden at Montreal, under the direction of the McGill Univer-
sity and the local horticultural society (F. B. P.).

2See Flickiger, ‘‘Osterferien in Ligurien;” Buchner’s Repertorium
fir Pharmacie, xxv. (1876), 449-505; also (Fliickiger) ‘“‘ La Mortola, Der
Garten des Herrn Thomas Hanbury,” Strassburg, 1886 (privately
printed), pp. 30.

3 Compare his publications: ‘‘ Unsere officinellen Pflanzen,” Gorlitz,
1883, pp. 12; and ‘‘Catalog der botanischen Museen der Universitat
Breslau,” Gérlitz, 1884, pp. 54.

‘
46 AIDS TO THE STUDY OF PHARMACOGNOSY.

barium) in Berlin has also begun worthily to follow the example,
though not precisely in a pharmaceutical direction.

C. Microscopical Preparations. Notwithstanding the im-
portance of a knowledge of the inner structure of many medi-
cinal substances, the preparation of objects for the microscope
often requires such an expenditure of time as to interfere
with its accomplishment. This is, however, so instructive (see
p. 48) that it is only in cases of necessity, or for the purpose of
supplementing individual work, that the purchase of micro-
scopical preparations will be attended with substantial benefit.
It must, nevertheless, be admitted that these preparations are
now presented, especially in Germany, in an unusually attrac-
tive and complete form.

Il. LITERARY AIDS.

The following list contains a selection of works which are
adapted to quite extended requirements, but makes no claim to
completeness,

A. Medico-pharmaceutical Botany. Of ies iota works,
the following deserve particular mention:

Kosteletzky, ‘‘ Allgemeine medicinisch-pharmaceutische
Flora,” 3 volumes, Prague, 1831-1834 (now sold by Hoff, ©
Mannheim).

Geiger, Nees von Esenbeck and Dierbach, ‘‘ Pharmaceutische
Botanik,” 3 volumes, Heidelberg, 1839-1843,

Bischoff, ‘‘ Medicinisch-pharmaceutische Botanik,” Erlangen,
1843; second edition, 1847.

Svbleiden, ‘*Handbuch der medicinisch-pharmaceutischen
Botanik,” Leipzig, 1852.

Rosenthal, ‘‘ Synopsis plantarum diaphoricarum. Systema-
tische Uebersicht der Heil-, Nutz- und Giftpflanzen aller
Linder,” Erlangen, Enke, 1872.

These works, which are excellent in their way, may, perhaps,
occasionally still be consulted with profit; they are, however,
more than replaced by the following:

Luerssen, ‘‘ Medicinisch-pharmaceutische Botanik,” also un-

LITERARY AIDS. 4?

der the more appropriate title, ‘‘ Handbuch der systematischen
Botanik, mit besonderer Beriicksichtigung der Arzneipflanzen,”
Vol. I., Cryptogams, pp. 657, 181 woodcuts, Leipzig, 1879. Vol.
II., Phenogams, pp. 1,229, 231 woodcuts, Leipzig, 1882.

Luerssen, “Die Pflanzen der Pharmacopca Germanica,”
Leipzig, 1883, pp. 664, 341 woodcuts. Chiefly compiled from
the preceding.

Herz, “‘ Synopsis der pharmaceutischen Botanik,” Ellwangen,
1883. .

Baillon, ‘‘Traité de Botanique médicale-phanérogamique,”
Paris, 1883-1884, pp. 1,499, with numerous woodcuts,

Targioni-Tozzetti, ‘‘ Corso di botanica medico-farmaceutica,”
second edition, Firenze, 1847. Not seen by us. Furthermore:

Leunis, ‘‘ Synopsis der drei Naturreiche.” II. part, Botany.
Third edition edited by A. B. Frank, Hanover, 1883.

Schimper, ‘‘ Taschenbuch der medizinisch-pharmaceutischen
Botanik und pflanzlichen Drogenkunde,” pp. 208, Strassburg,
1886.

B. The following works give illustrations of officinal plants —
(accompanied by a descriptive text which is often unsatisfac-
tory).

Nees von Esenbeck, “‘ Plante medicinales,” Diisseldorf, 1828-
1833. 4 volumes. Folio, with 545 colored plates (12 inches
wide and 19 inches long).

Hayne, ‘‘Getreue Darstellung und Beschreibung der in der
Arzneikunde gebriiuchlichen Gewiichse, und solcher die mit
ihnen verwechselt werden kénnen,” Berlin, 1805-1846. 14
volumes, quarto, with 648 colored plates.

Berg and Schmidt, ‘‘ Darstellung und Beschreibung sammt-
licher in der Pharm. borussica aufgefiihrten officinellen Ge- —
wiichse,” Leipzig, 1854-1863. With 208 colored plates, quarto.
Of excellent artistic execution.

Kohler’s ‘‘ Medicinalpflanzen,” Gera-Untermhaus. Begun in
1883, and not yet completed. |

H. Gross, ‘Die wichtigeren Handelspflanzen in Bild und —
Wort,” Esslingen, 1880.

48 AIDS TO THE STUDY OF PHARMACOGNOSY.

Bentley and Trimen, “ Medicinal Plants,” 4 volumes, Lon-
don, 1875-1880. Small quarto, with 306 colored plates.

Artus, “ Handatlas simmtlicher medicinisch-pharmaceutischer
Gewiichse,” etc. Sixth edition, edited by G. von Hayek.
Jena, 1882. j

Cassone, F., ‘Flora medico-farmaceutica,” 6 volumes, with
600 colored plates. 8vo, Torino, 1847-1852. Not seen by us.

In America, the following illustrated works are in course of
publication, which especially represent the medicinal plants in
use in this country.

Lloyd, J. U. and ©. J., ‘Drugs and Medicines of North
America.” A quarterly devoted to the historical and scientific
discussion of the botany, pharmacy, chemistry and therapeutics
of the medicinal plants of North America, their constituents, —
products and sophistications. Cincinnati, Ohio. Vol. I., of
which the first number was issued in April, 1884, is now
(March, 1886) complete, and comprises the natural order Ranun-
culace,

Millspaugh, C. F., “‘ American Medicinal Plants.” An illus-
trated and descriptive guide to the American plants used as
homeopathic remedies: their history, preparation, chemistry,
and physiological effects (6 fasic.). Fasc. I., New York, 1884.
Royal 8vo, 30 plates with descriptive text. (Four fascicles of
this work have appeared up to the present date, July, 1886.)
F. B. P.

C. Medico-pharmaceutical Zoology.

Brandt and Ratzeburg, ‘‘ Medicinische Zoologie,” Berlin,
- 1829-1833. 3 volumes, quarto, with 60 admirably executed
copper plates.

Martiny, ‘‘ Naturgeschichte der fiir die Heilkunde wichtigen
Thiere,” Giessen, 1854. 30 plates.

Moquin-Tandon, ‘“Eléments de zoologie médicale,” Paris,
1860; second edition, 1882, with numerous woodcuts.

D. Use of the Microscope.

Nigeli and Schwendener, ‘Das Mikroskop,” Leipzig, 18675
second edition, 1877.

Dippel, “‘ Das Mikroskop und seine Anwendung,” 2 volumes,

LITERARY AIDS, 49

Braunschweig, 1867-1872; second edition, 1883. By the same
author, “‘ Grundziige der allgemeinen Mikroskopie,” Braun-
schweig, 1885.

Hager, ‘‘ Das Mikroskop und seine Anwendung.” Seventh
edition, with 316 woodcuts. Berlin, 1886.

Behrens, ‘‘ Hilfsbuch zur Ausfiihrung mikroskopischer Unter-
suchungen im botanischen Laboratorium,” Braunschweig, 1883,
pp. 398. With illustrations. American edition, entitled: ““The
Microscope in Botany,” translated by Rev. A. B. Hervey, and
R. H. Ward. Boston, 1885.

Stein, ‘‘Das Mikroskop und die mikrographische Technik,”
Halle, 1884. With illustrations.

Poulsen, ‘‘ Botanische Mikrochemie,” Cassel, 1881, pp. 83.
American edition, entitled: ‘‘ Botanical Micro-chemistry,” trans-
lated by Professor William Trelease. Boston, 1884, pp. 118.

E. Chemistry.

Husemann and Hilger, ‘‘Die Pflanzenstoffe.” 2 volumes,
second edition, Berlin, 1882-1884. This book, which is very
complete from a chemical standpoint, dlso elucidates the
physiological action of the constituents of plants.

Dragendorff, ‘‘ Die qualitative und quantitative Analyse von
Pflanzen und Pflanzentheilen,” Gottingen, 1882, pp- 285. Eng-
lish edition, entitled: ‘‘ Plant Analysis, Qualitative and Quanti-
tative,” translated by Henry G. Greenish. London,’ 1884, pp.
280.

Ebermayer, ‘‘ Physiologische Chemie der Pflanzen,” Berlin,
1882.

F. Illustrations of Drugs. Figurative representations of
drugs in an ordinary sense are of less value than collections,
perhaps with exception of the Cinchona barks, in which a com-
parison with illustrations may occasionally be desirable. Such
colored illustrations of the Cinchona barks are presented in a
very handsome form by the following:

Bergen (H. von), ‘‘ Monographie der China,” Hamburg, 1826.
Quarto, with 7 plates. ;

Weddell, “Histoire naturelle des Quinquinas,” Paris, 1849.
Folio. The plates 28, 29, 30. grey me ey
4 MISSOURI
BOTANICAL
GARDEN.

Sina

a

50 AIDS TO THE STUDY OF PHARMACOGNOSY.

Delondre et Bouchardat, “‘ Quinologie,” Paris, 1854. Quarto.
23 plates.

A large number of drugs are illustrated, and described by
valuable essays in the following:

Hanbury, ‘‘ Science Papers,” London, 1876.

With regard to some antiquated drugs reference may be made
to Gobel and Kunze, ‘ Pharmaceutische Waarenkunde,” Kise-
nach, 1827-1834. 2 volumes, quarto, pp. 240 and 300. This
work contains numerous colored plates, representing barks and
roots.

G. Figurative representation of inner structure.

Oudemans, “‘ Aanteekeningen op het systematisch- en pharma-
cognostisch-botanische gedeelte der Pharmacopwa Neerlandica,”
Rotterdam, 1854-1856, pp. 661, with 37 plates. Has long been
out of print.

Berg, ‘‘ Anatomischer Atlas zur pharm. Waarenkunde,” Ber-
lin, 1864, 50 plates. Quarto.?

Vogl, ‘‘Nahrungs- und Genussmittel aus dem Pflanzen-
reiche. Anleitung zum Erkennen der Nahrungsmittel, Genuss-
mittel und Gewiirze mit Hilfe des Mikroskops,” Vienna, 1872,
pp- 138, with 116 woodcuts.

Bell, «Die Analyse und Verfiilschung der Nahrungsmittel.”
I. and H, Berlin, 1882 and 1884. The original English work,
of which this is a translation, bears the title: ‘‘ The Analysis
and Adulteration of Foods.” .

Miller, ‘“Mikroskopie der Nahrungs- und Genussmittel,”
Berlin, 1886, 8yo, pp. 384, with 308 woodcuts.

_ Schimper, ‘Anleitung zur mikroskopischen Untersuchung
der Nahrungs- und Genussmittel,” Jena, 1886, pp. 140, with
79 illustrations.

In the various commentaries to pharmacopeeias, in the large
work of Luerssen, mentioned on page 47, as also in mono-
graphs, there occur many illustrations pertaining to this
subject. Thus, for example, in Rauter’s “Memorial Publica-

' This excellent work will be further referred to by us under the desig-
nation of Berg’s “‘ Atlas,”

LITERARY AIDS. ; aL

tions of the Vienna Academy” (‘‘ Denkschriften der Wiener
Akademie”), 31 (1872), 23, hair formations; Martinet, “ Or-
ganes de sécrétion des végétaux” in Annales des sciences natu-
relles, xiv. (1872), 91, and many other publications mentioned in
the chapter on anatomy. The structure of leaves is represented
by Lemaire in his ‘‘Détermination histologique des feuilles
médicinales,” Paris, 1882; and that of barks in MOller’s “ Ana-
tomie der Baumrinden,” Berlin, 1882. The excellent illustra-
tions accompanying Arthur Meyer’s “‘ Beitrige zur Kenntniss
pharmaceutisch wichtiger Gewiichse” in the Archiv der Phar-
macie, Band 218, 210, 220, 221 (1881 to 1883), relate to the
officinal Smilacew and Zingiberacex, to Aconitum, Veratrum,
“Gentiana and Ipecacuanha.

H. Special Pharmacognostical Text-books and Manuals.

Berg, ‘* Pharmaceutische Waarenkunde.” Fifth edition, Ber-
lin, 1879, pp. 696. This first appeared in 1850 as the second
volume of the second edition of his ‘‘ Pharmaceutische Botanik ”
(the first edition of the latter was issued in 1845).

Fliickiger, ‘‘ Lehrbuch der Pharmacognosie des Pflanzen-
reiches,” Berlin, 1867, pp. %58. Second edition, 1883,- pp.
1,049.

Flickiger, ‘“‘Grundriss der Pharmacognosie,” Berlin, 1884,
pp. 260.

Flickiger and Hanbury, ie Pharmacographia: A History of
the Principal Drugs met with in Great Britain and British
India,” London, 1874. Second edition, 1879, pp. 803.

Flickiger et Hanbury, ‘‘ Histoire des Drogues d’origine végé-
tale.” Traduction de l’ouvrage anglais ‘‘ Pharmacographia,”
par J. L. de Lanessan. 2 volumes, with woodcuts. Paris, 1878.

Fristedt, ‘‘ Larobok i organisk Pharmacologi,” Upsala, 1873.

Guibourt, ‘‘ Histoire abrégée des Drogues simples,” Paris,
1820. Sixth edition by G. Planchon: “ Histoire naturelle des
Drogues simples.” 4 volumes, Paris, 1869-1870. With wood-
cuts.

Marmé, ‘‘ Lehrbuch der Pharmacognosie des. Pflanzen- ‘iad
Thierreichs,” Leipzig, 1885.

Oudemans, ‘‘ Handleiding tot de Pharmacognosio van het

52 AIDS TO THE STUDY OF PHARMACOGNOSY.

Planten- en Dierenrijk,” Haarlem, 1865. Second edition, Am-
sterdam, 1880.

Planchon, ‘‘ Traité pratique dela Détermination des Drogues
simples d’origine végétale,” 2 volumes, Paris, 1875. With illus-
trations.

Vogl, ‘‘ Commentar zur ésterreichischen Pharmacopée,” Band
I. ‘ Arzneikérper aus den drei Naturreichen,” Vienna, 1869.
Third edition, 1880, pp. 516, with 164 woodcuts.

Wigand, “Lehrbuch der Pharmacognosie,” Berlin, 1863,
with 141 woodcuts. Third edition, 1879, pp. 447, with 181
woodcuts.

Wiggers, ‘‘Grundriss der Pharmacognosie,” Gottingen, 1840,
pp. 429. Fifth edition, “Handbuch der Pharmacognosie,”
1864, pp. 800.

Wittstein, ‘ Handworterbuch der Pharmacognosie des Pflan-
zenreichs,” Breslau, 1883, pp. 994.

The numerous drugs in use in India, especially those of the
western portion of the peninsula, have been thoroughly consid-
ered by:

Dymock, “‘ The Vegetable Materia Medica of Western India,”
Bombay (London), 1883-1884, pp. 786. (Compare the reviews
in the Archiv der Pharm., 222 (1884), 249, and Pharmaceu-
tische Zeitung, Feb. 23d, 1884. Supplement to No, 16.)

All that has been accomplished in relation to the useful plants
of India in the broadest sense, including the drugs, is compiled
in the following large work in course of publication at Calcutta:

Watt, «A Dictionary of the Economic Products of India.”
The first volume of this work, comprising the letter A, was pub-
lished in 1884. Large 8vo, pp. 353.

: The drugs employed in the United States have been elaborated
y:

Maisch, ‘“‘A Manual of Organic Materia Medica,” Philadel-
phia, 1882, pp. 459, with illustrations. Second edition, 1885,
pp- 511, with 242 illustrations.

Lloyd (J. U. and C. G. )s “« Drugs and Medicines of North
America.” See page 48, —

_Of the resources of Brazil, an idea is given by:

LITERARY AIDS. 53

Peckolt, ‘* Catalog der pharmacognostischen, pharmaceut-
ischen und chemischen Sammlung aus der Brasilianischen
Flora.” Zeitschrift des Allg. ssterreichischen A potheker-
Vereines, VI. (1868), 518. Extended pharmacognostical essays
by the same author are to be found in the periodical mentioned.

Representations and descriptions of drugs may be found,
although for the most part very deficient, in the various com-
mentaries to the pharmacopeias, and in the manuals of the
practice of pharmacy.

J. An exhaustive work on the History of Pharmacognosy,
based upon an adequate study of sources of information, is still
wanting. In the circle of German literature the following are
worthy of consideration, however, in this respect:

Meyer, “Geschichte der Botanik,” 4 volumes, Kénigsberg,
1854-1857. This work has, indeed, remained uncompleted,
since it extends only to the last quarter of the sixteenth century,
but nevertheless includes the periods which, in many respects,
are the most interesting for the history of pharmacognosy.

Jessen, ‘‘ Botanik der Gegenwart und Vorzeit in culturhisto-
rischer Hinsicht,” Leipzig, 1884. This work supplements the
last named, and contains, notwithstanding its brevity (495
pages), a large amount of substantial information, although
pharmacognosy does not properly fall within its sphere.

Hehn, ‘‘ Kulturpflanzen und Hausthiere in ihrem Uebergang
aus Asien nach Griechenland und Italien, sowie in das iibrige
Europa,” fourth edition, 1882.

Sigismund, ‘‘ Die Aromata in ihrer Bedeutung fir Religion,
Sitten, Gebriiuche, Handel und Geographie des Alterthums bis
zu den ersten Jahrhunderten unserer Zeitrechnung,” Leipzig,
1884, (Reviewed in the supplement to the Pharm. Zeitung,
Bunzlau, May 31st, 1884, p. 377.)

A. de Candolle, “‘ Origine des plantes cultivées, Bibliothéque —
scientifique internationale,” Paris, Germer Bailliére & Co.,
1883.

K. A Bibliography of Pharmacognosy is also still want-
ing. The beginning of such a work is presented in Pereira’s
** Elements of Materia Medica,” II. (1857), 833-869. Some

54 AIDS TO THE STUDY OF PHARMACOGNOSY.

references may furthermore be found in the previously cited
(p. 51, under H), ‘* Pharmacognosie,” by Fliickiger (especially
pages 983-1022), and in the ‘‘ Pharmacographia.” A manu-
script bibliography of pharmacy by Piper has been preserved by
the Pharmaceutical Society of Great Britain in London since
1883.

L. That valuable aids to the study of pharmacognosy are con-
tained in the various scientific journals and in monographs
needs scarcely to be mentioned. In Germany, even previ-
ous to the year 1825, the Archiv des Apotheker-Vereins im
nordlichen Teutschland inaugurated the stately series of volumes
of the present Archiv der Pharmacie, while many other German
professional journals have since ceased to exist. In Paris, the
Journal de Pharmacie et de Chimie has likewise appeared regu-
larly since that time. In England, Jacob Bell, who rendered
such valuable service to English pharmacy (1810-1859), issued
in July, 1841, the first number of the “Transactions of
the Pharmaceutical Meetings,” which now, under the title of
The Pharmaceutical Journal and Transactions, continually pre- |
sents a rich supply of pharmacognostical information, com-
mensurate with the commercial status of England. The Amert-
can Journal of Pharmacy, in Philadelphia, stands already
in its fifty-eighth year of publication.

The “ Annual Reports,” finally, summarize all that is of value
in current literature. Thus, in Germany, the ‘‘ Jahresbericht
der Pharmacie,” which was begun in 1841 by Theodore Martius
(1795-1863), and continued by Wiggers from 1844 to 1865. In
1866, Theodore Husemann, together with Wiggers, undertook
its elaboration, and from 1867 to 1873 August Husemann was
associated with them. In 1874, Dragendorff took the place of
Theodore Husemann, and continued the “Jahresbericht” until
1878, when, in 1879, Marmé and Wulfsberg associated them-
selves with him. The ‘ Bericht” of 1880 was edited by the
latter, and the two following by Beckurts. A similar service
has been rendered in England by the ‘‘ Yearbook of Pharmacy,”
which has been issued since 1870, and much more punctually
than the German “ Bericht,” by the British Pharmaceutical

LITERARY AIDS. ou

Conference, and in America, since 1857, by the annual “ Re-
port on the Progress of Pharmacy,” contained in the ‘‘ Proceed-
ings of the American Pharmaceutical Association,” in which
trade relations also find consideration.

‘The Botanisches Centralblatt (Fischer, at Cassel), a journal
of reference, which has appeared since 1880, under the editor-
ship of Behrens and Uhlworm, and (since 1873) Just’s Bota-
nischer Jahresbericht (since 1883 under the editorship of Geyler
and Kéhne), also consider the literature of pharmacognosy,
even though only in a secondary manner.

MORPHOLOGY.’

_The few officinal alge, lichens and fungi, such as Chondrus
(Irish-moss), Cetraria (Iceland-moss), Secale cornutum (Ergot),
and the unofficinal Fungus laricis (Larch Agaric), afford ex-
amples of entire plants’ which serve for medicinal purposes;
_ the remaining drugs consist of parts of their mother-plants.
Among the underground or, at least, half underground organs
which fall within the province of pharmacy, the following are
to be distinguished:

Roots (Radices). We confine this expression to the axes
of endogenous formation, in which the capability of producing
leaves, and mostly: also chlorophyll, is wanting, but which
possess a root-cap. This latter is a delicate, slightly extended
tissue on the growing end (tip) of the root. The cells of the
root-cap increase in number by division, but remain uniform;
and farther back, within the cap, the point is reached where
the formation of the different systems of tissue begins (see Ana-
tomy, under formative tissue).

Roots are designated as main or primary roots when they
represent the direct (underground) descending continuation of
the base of the stem. Frequently the primary root becomes
divided into root-branches, which, when they are very thin, are

'A thorough treatise on morphology, based upon modern views, is
contained in Leunis’ “Synopsis,” Bd. I., 1882, Edited by B. Frank.
Compare also Th. Liebe, ‘‘ Die Elemente der Morphologie,” Berlin, 1881.

*The sclerotium of Claviceps purpurea, which constitutes the Secale
cornutum, is the passive condition of the fungus. Fungus laricis is the
fruit-bearing portion of Polyporus officinalis Fr., and is thus, strictly
considered, also only a portion of a plant, even though quantitatively
_ by far the preponderating portion.

RUNNERS AND STOLONS. 57

also called root-fibres. Examples of primary roots are afforded
by Radix Pyrethri, Rad. Scammonie, and Rad. Tarazaci ;
excellent root-branches are possessed by Rad. Ratanhie, and
root-fibres by Rad. Angelice.

Secondary roots are such as originate laterally from the pri-
mary root or from parts of the stem, and which, upon the
whole, since the main root is often suppressed, occur much more
frequently than the primary root; in those cases, therefore,
where the distinction between primary and secondary roots is
not sharply characterized, as with Rad. Angelice and Rad.
Levistici, they may be designated simply as roots.

Occasionally, only the secondary roots are collected, but for
the most part the primary and secondary root together are offi-
cinal.

Above the tip of the root, mostly confined to small tracts, and
advancing with the growth in length, are found the root-hairs,
which individually (see Anatomy) become firmly attached in
their growth to the particles of earth, and convey the dissolved
nutritive substances of the soil to the root.

The function of the root is to fix the plant in the soil (hence
the central vascular bundle cylinder; see Anatomy) and to ab-
sorb the inorganic nutritive substances and water from the soil.
- The roots, as well as many rhizomes, are provided with a
bark which, in some cases, is removed by paring before the
drug is made use of (Althea, Glycyrrhiza, Iris), and in others,
where it is rich in active constituents, it is allowed to remain
(Calamus). Barks separated from the root are also officinal,
thus, e. g., Granatum (Pomegranate), the bark of the root of
Punica Granatum Lin. The root-bark of Krameria (Rhatany)
was also formerly in use.

Beside the true roots, there is still a series of underground
formations, which belong, however, morphologically to the
organs of the stem, even though they also fulfil physiologically
the function of roots. These are the runners or stolons, rhi-
zomes and tubers.

The Runners (flagella) and Stolons (Stolones) which
emanate from the root or from the base of the stem lie on the

58 MORPHOLOGY,

surface of the ground or at a slight depth, and develop hori-
zontally, often to a considerable length. They possess under-
leaves, at least in a rudimentary state, often contain chlorophyll,
and inclose a clearly defined medulla. At some distance from
the point of origin, the runners are able to develop roots and
sprigs of foliage; if their connection with the mother plant is
subsequently severed, they develop new individuals. Radix

Glycyrrhize and Rad. Saponarie consist to a large extent of
runners.

The typical form of stolon is shown very nicely in the so-called —

Rhizoma Caricis (Carex arenaria Lin.). The sheath-like under-
leaves here surround the nodes, which are provided with root-
bundles. Rhizoma Graminis (Triticum repens Lin.) may also be
classed with the stolons.

The runners are botanically connected with the creeping or
repent and decumbent stems.

Of the underground stem formations which bear under-leaves
there are furthermore to be mentioned: the Rootstocks or
Rhizomes (Rhizomata'). These are perennial stems or
branches of vascular cryptogams and phenogams, growing half
or entirely under the surface of the ground, which are provided
with the rudiments of leaves, the remnants of leaf-sheaths or
leaf-nodes, and send out roots (secondary roots). We also re
tain the name rhizome for those cases where the roots (secondary
roots) are collected with it, or even when these preponderate, as
with Valeriana. Radix Sarsaparille alone forms an exception;
_ for although its rhizome occurs in part in commerce, it is never-
theless inadmissible for medicinal use.

The rhizome is provided at its apex with a bud, and annually
develops new, ascending, herbaceous stems (leaves).

The peculiarity of the rootstock and the bulb, so definitely
characterized by nature in opposition to the root, had already
been rendered prominent by the Greek naturalist Theophrastos *
(371 to 286 B.c.), Las

For the most part the rootstock alone is collected (Jris,

1“Pil@ua root, stem. eta eg pee
_ * Compare Jessen, ‘* Botanik der Gegenwart und Vorzeit,” 1864, 26.

t

TUBERS—BULBS. 59

Galanga). When besides a central rootstock a secondary or
lateral rootstock is present, both of them occur in commerce
(Zingiber), although rarely as two separate commercial varie-
ties (Curcuma rotunda and longa, the latter of which forms the
secondary, the former the primary rootstock). Of Zedoaria
only the primary rootstock is in use.

The roots of a rhizome (either alone or connected with the
latter) form the sarsaparillas of commerce. ’

Branched rootstocks (many-headed) are not of rare occur-
rence, and then possess many buds (Galanga’).

The rhizomes serve also as receptacles for reserve substances,
and are nearly always densely filled with reserve nutritive sub-
stances (starch, etc.).

Still more pronounced receptacles for reserve materials are:

The Tubers (7udera).—These are underground portions of
the stem, or ramifications of the roots of phenogams (Orchis),
which are so thickened that their diameter approximates their
development in length or exceeds it. This growth in thickness
stands in connection with the periodical accumulation of
constructive materials, especially starch. They present a fleshy
structure, which after drying is mealy and horn-like, not woody,
as, ¢. g., Tuber Aconiti, Tuber Jalape and Tuber Salep.
Tuber Chine deviates from the ordinary type of the tuber by its
often very considerable length and strongly developed vascular
bundles. *

It is, however, not only the stem which participates in the
formation of underground organs, occasionally it is also leaves
(under-leaves) which form these to a predominating extent, as,
for example, in the case of the bulbs.

Bulbs (Buldi).—These are fleshy, thickened under-leayes or
parts of leaves, serving for the storage of starch and other sub-
stances serving as nourishment, which, similarly to buds, are
severally inwrapped, scale-like, about a very short axis pro- —

‘Compare A. Meyer, Archiv der Pharm., 218 (1881), 280, with cs
illustrations, .

* A. Meyer, loc. cit. 425, with iitiateatiods, ae

* Compare A. Meyer, Archiv der Pharm., 218 aan ae

60 MORPHOLOGY.

vided with’roots, and stand above the surface of the soil or are
half or entirely imbedded therein. The axis of the bulb is mostly
flat, disk-like (Fig 1, 7). The only officinal bulbs of the U.S.
Pharmacopeia (1880) are Scilla and Allium.

By the accumulation of reserve substances, aid is given to the
respective plant, corresponding to the climatic conditions, ren-
dering it capable of attaining its full development in the shortest
time and during the most favorable periods of the year.

A Corm (Cormus) is intermediate between the tuber and the
bulb. Itisa bulbo-tuber or bulbodium tunicatum, or atrue tuber,
which is provided with under-leayes and enveloped by them (Col-
chicum).

The thickened, and for the most part subterranean organs of

Fig, 1.—Median longitudinal section through a tunicated bulb. 1, the disk-shaped,
expanded axis; v, the bud; t, under-leaves; 7, roots. _
the stem, above described, lead us to the consideration of the
proper organs of the stem, growing above ground.

Stem structures are formed either by the development ot
the stem portion of the plumule of the germinating plant (see
below), or as lateral shoots on other organs of the stem, in the
axils of leaves.

That portion of the stem which lies between the point at
which the cotyledons are affixed and the point of attachment of
the root is called the hypocotylous' member, caulicle or radicle.
This remains mostly ;short (in the Hypogee,” i. ¢., those

‘*Txé under, and xorvAndar (cotyledon) cavity.
* "Tmo under, and yara earth.

STEMS OR TWIGS—WOODS. 61

plants whose cotyledons in germinating do not appear above the
surface of the ground); it can, however, also become extended,
and then elevate the cotyledons far above the ground (in the
Epigee,' for example, Linum, Vicia faba).

The form of a transverse section of the stem structure is
cylindrical or angular. In the latter case the position of the
leaves stands in direct relation to the number of the angles
(Labiate, with a quadrangular stem and cross-like arrangement
of the leaves). On cylindrical stems the leaves are arranged in
spirals.

If the stems are developed in a leaf-like form, they are called
phyllodia or phyllocladia’? (Ruscus, Australian species of Acacia
Phyllocactus).

In a pharmacognostical relation, the following organs of the
stem come under consideration:

Stems or Twigs (Stipites). By these names are desig-
nated the weaker biennial and triennial, overground axes of
dicotytedons, which are covered with an epidermis or cork, and
contain chlorophyll. The only example properly considered
here is presented by Stipes Dulcamare.

Woods (Ligna). By wood is understood the mostly ‘‘lig-
nified ” tissue (see below) of overground axes (or also of dicoty-
ledonous roots), located inside of the cambium ring, which, with
exception of the medullary rays, has attained considerable solidity
through a material thickening of the cell walls. Only wood of
many years’ growth of the Gymnosperms, as also that of the
Angiosperms, serves for medicinal purposes, with or without
the bark, occasionally also either entirely or partially freed from
_ the outermost, younger layers, the so-called sapwood or albur-
num (see Anatomy).

Wood upon which the bark still remains, is frequently found in
the two varieties of Lignum Quassie and in Lignum Sassafras,
while Lignum Juniperi and Lignum Guaiaci are nearly always
freed therefrom.

The numerous dye-woods, as, ¢. 9-, Lignum Homgicey (log:

!Exi upon or over, and yaia earth. wee.
* SUAAor leaf, and xAaddos shoot orstem.

62 MORPHOLOGY. _

wood) and Lignum Fernambuci (Brazil wood) arrive in commerce
in the form of heartwood, freed from the colorless sapwood. In
the case of Lignum Guaiaci, the valuable constituent, the resin,
is also confined to the heartwood, although the dealers do not
make a practice of removing the sapwood.

In a botanical relation, the herbaceous stems (caulis) are dis-
tinguished from the woody trunks (trunci), which, with the
dicotyledons, as a rule, form branches (rami).

Barks (Cortices). The organs of the stem, as well as
roots, are invested with a bark, The monocotyledons do not
possess a true bark, since they show no secondary growth in thick-
ness. ‘The barks of dicotyledons—and such are officinal—con-
sist in their young condition of a predominating parenchymatous
tissue, derma, or primary bark, which is covered by the epider-
mis. Only barks of several years’ growth, however, are brought
into use, where the epidermis is replaced by cork. Where the
latter is in a state of active growth, and represents a coherent
layer, it is called periderm, outer bark, or exophleum." :

Upon further development of the overground axes (or roots),
which here come into consideration, the growth of the original
bark is very soon increased through the activity of the cambium.
Within the primary bark there is developed the secondary bark,
inner bark, liber (bast), or endophlwum, the boundary of which
is very frequently sharply defined from the primary bark. The
bark then presents also a middle layer, the so-called middle bark
_ or mesophleum,* which, at a later period, often becomes very
neocon constituting the remainder of the primary
- When, however, the formation of cork does not remain con
fined to the periphery, but forms also in the interior of the
bark tissue, the middle bark, by means of cork bands, ca)
become entirely separated and thrown off, This process, the
_ formation of bork or rhytidoma,’ can also extend to the inner
bark. If, through peeling, the bork-scales themselves are als?

' Ex, & out of, from, and ploror bark,
* Mé6o5 middle, _
*"Puris, puridos fold, wrinkle; Scuda I build.

BARKS—HERBS. 63

removed, such a bark then consists at last almost exclusively of
the inner bark, as, e. g., the Cinchona calisaya and Coto bark.

The inner bark is intersected by medullary rays (bark rays)
which, upon a transverse and longitudinal section, are often
plainly perceptible, without being magnified, as delicate lines.

The fracture of barks [that is, the appearance of a fractured
surface], which is frequently a very useful characteristic, on the
other hand, chiefly depends upon the fibres in the secondary
bark (bast-bundles), upon the degree to which the elements of
the same are thickened, and upon the form of combination of
their prosenchymatous cells.

It is to very long, soft, and intertwining bast-fibres that Cortex
Mezerei (Mezereum), as likewise the bark of Radix Althee, owe
their eminently fibrous character. The Cinchona barks are brit-
tle, because their strongly thickened bast-tubes remain short and
are usually isolated. The outer surface of Ceylon cinnamon
admits of the ready recognition of the long, wave-like bast-bun-
dles, which here and there intersect each other.

The cork (outer bark), is usually rejected, since it contains
no active constituents, If it be thin, it often remains preserved
on the bark, and can afford good means of distinction. Of the
cambium, itis but seldom that remnants are preserved, since its
cells possess very delicate walls.

Occasionally entire annual plants, or those of a few years’
growth (without the roots) are employed. In pharmacognosy,
these are then spoken of as herbs. “3 :

Herbs (Herba). These are the leafy shoots of phenogams
which, beside the leaves and tenderer portions of the stem (the
main axes are often removed) may possibly contain individual
flowers and fruits, or the entire inflorescence and collection of
fruits. There are no reasons for the exclusion of these struc-
tures which often accompany the leaves, and frequently it would
be virtually impossible, as, ¢. g., in the case of Herba Centaurtt,
Herba Meliloti, Herba Serpylli, ete. ee

Not unfrequently, the form of ramification of the shoots is of
diagnostic importance in the case of herbs. The different systems
of ramification may, therefore, be considered here ina few words,

64 MORPHOLOGY.

The formation of homologous axes is called ramification. An
axis, together with its branchlets, is called a system of ramifica-
tion. The systems of ramification appear in two principal
forms.

1, THE DICHOTOMOUS SYSTEM (the axis ceases to increase in
length at the apex):

(a) ForKED DicHoTomy.’ Directly beside the apex which
extends no farther in growth, there arise two new axes which
develop equally,.and can form the bases for further ramifica-
tions.

(4) SympopiumM®* (in a more restricted sense), With each fur-
cation an axis is developed, ultimately stronger than the other.
The basal portions of the successive furcations form a pseudo-axis
(sympodium) on which the more feebly developed axes are at-
tached as lateral branches. Suchasympodium can consist either
of the forked branches which are always of one side (bostryx),
or of forked branches which are alternately of the right and left
side (wickel or cincinnus).

2. THE MONOPODIAL SYSTEM * is produced by the main shoot
continuing to grow, and forming lateral branches below the
apex.

The monopodium can be either racemose, ti. e., when the
main axis, even at later periods of growth, is always most
strongly developed, or cymose, when the lateral axes overtop the
main axis. In the latter case a pseudo-axis may be formed.
Regarding this compare the subsequent remarks under inflores-
cence.

Exogenous * formations which arise laterally on the organs of
the stem in acropetalous * succession, and have a different form
from the stem which produces them, are called leaves.

The leaves arise densely crowded on the summit of the stem.
If the parts of the stem located between two leaves become ex-

* Atyoréuos cut in two (diya twofold) parts (réuva@ cut).
* Zur entire, united, and xovs, rodcs foot.

’ Movos single, and zovs foot.

‘Ex, 2 outside, and yéros formation.

5’ Aupcs point, and petere to strive after.

THE LEAVES. 65

tended, there are formed the internodes.'. The parts of the
stem bearing the leaves are thus called the nodes, These are
occasionally swollen (Polygonacee, Piperacee, and the grasses).
The place of attachment of the leaf is called the point of inser-
tion.” According to the distribution of the leaves on the stem,
there are distinguished the spirally arranged (Chenopodium),
crossed or decussate’ (Labiate), and whorled leaves (Juni-
perus).

The stalked leaves admit of the distinction of the leaf-stalk or

Fig, 2.—Leaves of Eucalyptus globulus Labill. (Heterophylly). a, sabre-shaped
leaf of an older branch; }, heart-shaped leaf of a younger branch. The oil cells are
indicated by dots. (Tschirch, in Pharm, Zeitung, 1881, No. 88.)

petiole ( pefiolus) and the leaf-blade or lamina; * on the boundary _
of both these parts there often appears a small membrane or |

ligule (ligula). The base of the leaf-stem is often ea Bo

* Inter, between, and nodus, node or joint.
_ ? Inserere, to insert.
* Decussare (decussio = the number ten), to divide in form of an x
(i. e. , crosswise),
. Lamina, : thin plate.

66 MORPHOLOGY.

a broad sheath (vaginw), and surrounds the stem in a tube-like
form. With the leaf-sheath are also to be classed the stipules
(stipule),’ which are mostly of different, but occasionally of
the same form and color as the main leaves (Asperula), and oc-
cur at the base of the latter.

The form of leaves is very multifarious, although those which
are. developed in a broad, flat form decidedly predominate.
If one and the same plant possesses differently formed leaves
at different places, it is called heterophyllous* (Eucalyptus,
Fig. 2). Since the leaves serve principally for the assimilation
of carbonic acid in the light, they present their broad, green
surfaces to the sun. This applies, however, only to the true

<<

MIDNA FAG

Fig. 3.—Rhizome of ‘Gratiola officinalis, with roots. a—b, rhizome; at 6, the ter-

minal bud; d, under or rudimentary leaf; c, a shoot issuing from the axil of a rudimen-
tary leaf, and likewise provided with rudimentary leaves,

FoLIaGe LEAVES (Folia), which are chiefly met with in the
region between the root and the flower. It is otherwise with the
_ UNDER-LEAVES or CATAPHYLLA (squame):, which are inserted
chiefly on the basal portion of the stem, and on the rhizomes,
stolons, etc., with a broad surface (Fig. 3). They are for the
most part brownish or pale in color, and to be regarded as
protecting envelopes for the tender, growing portions. The
fleshy bulb-scales are likewise rudimentary or under-leaves

’ 1 Properly halm or straw, on account of their dry, membranous con-
sistence. In German, Nebenblitter.

*”Erepos the other, and puAAor leaf.

5 Squama, scale.

THE FLOWER. 67

(Fig. 1,7). Such appear, however, also as bud-scales, on the
upper portion of the stem, enveloping and protecting the young
leaf-buds; they have here the same function.

The germinating leaves or cotyledons’ must also be regarded
as rudimentary leaves. These possess, however, as we shall sub-
sequently see, quite another function.

The third form of leaves are the HIGH LEAVES or HYPSO-
PHYLLA (bractee).* These belong to the inflorescence, are like-
wise small and mostly delicate, and serve often as protecting in-
teguments (the glumes of grasses).

The leaves, particularly foliage leaves, are often entirely (as
with the stipules* of Smilax) or in part (many Papilionacee)
converted into tendrils, although, on the other hand, all tendrils
are not metamorphosed leaves ( Vitis, Ampelopsis, Passiflora).

The most manifold forms and colors are assumed by the ele-
ments of the last leaf formation, the flower. While the leaves
of the calyx (sepals) still possess a purely leaf-like nature and
are mostly green, the leaves of the corolla ( petals) appear colored
and often of a particular form, the staminal leaves become con-
verted into structures of quite another form (stamens), and the’
pistil-leaves closed together to form peculiar receptacles (ova-
ries). Nevertheless, the flower must be considered as the sum-
mit of a branch, encompassed by manifold leaf organs.

The summit of a branch or axis bearing the flowers (receptacu-
lum, hypanthium, torus) is, as a rule, either conical or flat, but
can also be enlarged in a broad or cushion-like form and be-
come a disk * (Rutacew). If the disk lies within the circle of
stamens, it is called an intrastaminal disk.*

There occur, however, also extrastaminal disks (sculine).
The disk generally bears honey-glands or nectaries*; the latter,

' KorvAy cavity, pan.

* Bractea, a thin, small leaf of wood or metal. In German, Hoch- |
bldtter or Deckblitter.

3 In German, Nebenblitter.

4 Aiéxos disk.

5 Intra, within. ree

° Nectar, the drink of the gods, honey, a sweet substance.

68 MORPHOLOGY.

however, also appearing as appendages of the perigon (Ranuncu-
lus), or as metamorphosed parts of the perigon itself (spur), or
even as appendages of the stamens (Crucifere, Lauracee).
The form of the nectaries is also exceedingly variable. In the
case of Huphorbia Cyparissias, for example, they have the
shape of a half-moon.

The flower-stalk or peduncle bears the flower (los) which, in
consequence of the bright colors of its corolla, attracts the insects
for fertilization. This is the physiological significance of the
color and fragance of the flower. Insects are especially attracted
by the labellwm of flowers of the Orchidacex (the posterior leaf
of the inner perigon circle) and of the Zingiberacese (the meta-
morphosed lowest stamen of the outer staminal circle), as also
by the vexillum of flowers of the Papilionaces (the uppermost
petal).

When in a flower both the calyx and corolla are present,
the leaves of the calyx or the sepals (sepala') are mostly green,
while the leaves of the corolla, or the petals ( petala*), are of an-

other color. When both cannot be distinguished, or one of them
is wanting, the floral envelopes are spoken of as a perigon.§
When the individual leaves are united or coalescent, the floral en-
velopes are termed sympetalous,* gamopetalous* (synsepalous,
gamosepalous); when they are separate, choripetalous° (chori-
sepalous), or also spoken of in general as gamophyllous and
choriphyllous.

According to the arrangement of the floral leaves in the bud,
there are distinguished the following forms of wstivation :

Valvate (Malva, Vitis), Fig. 4a;

Imbricate (Geranium, Veronica, Rosa), Fig. 4b and 6;

Convolute (Gentiana, Phlox), Fig. 5;

Plicate or plaited (Campanula), Fig. 7 ;

'Separ, divided (?)

* Tlérador leaf.

* ITepz around, and yéros ‘brood, seed.

* Dur together, and zérador leaf.

’Tauos marriage, and zérador leaf.

° Xwpil@ to divide (yapis separated), and zérador leaf.

CALYX AND COROLLA. 69

and, according to the disposition of ordinary leaves in the bud,
the conduplicate, plicate or plaited, involute, revolute, circinal
or circinate, and corrugate or crumpled forms of vernation (ver-
natio). Compare also the diagrams in figures 11-15.

O

Fie. 4 a.—Valvate estivation (Vitis). Fie. 4 b.—Quincuncially imbricate
‘estivation (Rosa).

After flowering, the perigon, as a rule, falls off. In some
cases, however, it takes. part in the formation of the fruit. It
remains either herbaceous (Chenopodiacex, Polygonace), or
becomes soft like a berry (Morus), or grows out in a hair-like
form (riophorum). The calyx, by subsequent enlargement,
often becomes the calyx of the fruit (Hyoscyamus, the Borragi-
nace, Physalis), and serves them for the protection of the lat-

Fie. 5. Fie. 6. Fie. 7.
Fia. 5.—Convolute eestivation (Phlox) ; a, elevated view ; 6, outline.
Fia. 6.—Imbricate ewestivation (Veronica), a and bas in Fig. 5.

Fic. 7.—Plicate or plaited sestivation (Campanula), a and 6 as in Fig. 5.

ter. Occasionally, however, there appear in its place structural

appendages which are comprehended under the name of pappus*

(the Compositw, Valeriana). The latter are either bristly or
‘ [Iaxmo3 crown of hairs on the fruits of the Dandelion and the Let-—

70 MORPHOLOGY.

hair-like, and serve as organs of distribution, as well as of attach-
ment of the fruits.

In plants of the Malvacexw, there even appears, outside of
the true calyx, an involucre of high leaves or bracts, forming an
outer calyx (calyculus). In Fragaria, the outer calyx is formed
from the stipules of the calyx leaves or sepals.

Occasionally the perigon is reduced to small scales (Jodicule),
as in the case of many grasses (compare Fig. 11, 7), certain spe-
cies of Aconitum, and others.

The andreceum' of the flower consists of the stamens.”
They form the male or fertilizing organs, and bear upon a long
stem (filament), which is occasionally branched or expanded in
a leaf-like form at the base, the receptacles of the pollen grains,
namely, the pollen-sacsin theanthers.? These receptacles, which
are mostly two in number, constituting the so-called halves of the
anthers or thec@,* are held together by the connective,° and, as a
rule, contain, in each, two pollen-sacs, lying one above the other.
There occur also so-called monothecous anthers (Ricinus, the
Malvacew). It is rarely the case that one-half of the anther is
fertile, and the other not (Salvia). The connective, rwhich is
mostly short and but slightly developed, becomes occasionally
long and thread-like (Salvia), and then holds the anthers wide
apart. In other cases (Zilia), it forms a wide bridge. Rarely
are the anthers destitute of filaments, or sessile.

In order to discharge the pollen, the anthers open mostly by
a longitudinal line or chink (cleft, rima), more rarely by valves
(Lauraces). If they dehisce toward the interior, they are called
' introrse ;° if, on the contrary, toward the exterior, eztrorse.’. The
. pollen is mostly granular, and readily becomes dust-like ; in the
Orchidacex and Asclepiadacez, it forms glutinous masses ( pol-

'’Ayyp aman, and o7xos house.

* Stamen, warp on a loom, thread,

5” Av Snpds blooming.

4Onun a receptacte,

5 Connectere, to connect.

* Introrsus (introversus), toward the interior,
1 Extrorsus, toward the exterior.

THE GYNZCEUM. V1

Zinaria, pollinia). The pollen grains, like to the spores of
of the cryptogams, possess a variously marked outer coat (exine),
provided with bristles or prickles ; by the development of the
pollen tube, the intine projects in a sack-like form through the
exine.

If the stamens have all coalesced to form one bundle, they are
called monadelphous’ (Geraniaces, Linaceew, Oxalideex); when
coalesced to the number of two or several, polyadelphous?
(Papilionacee). The stamens may, however, have coalesced
with the perigon (Symphytum) or with the gyneceum, when
they are called gynandrous*® (Orchidacee, Aristolochie). In
the latter case, the style, stigma, and anthers form the fructi-
fying column (gynostemium).*

Occasionally the stamens develop leaf, horn, and pocket-like
appendages which, in the case of Asclepias and Vincetoxicum,
assume the form of an inner crown (corona staminea).

Sterile stamens are called staminodiums (Lauracee, Linum,
Zingtberaceex).

The gyneceum ° (pistil®) of the flower, the female sexual
organ of the plant, is formed of the fruit leaves or carpels,’
which are mostly closed to form one vessel, the ovary, contain-
ing in its interior the seeds. If but one carpel takes part in
the formation of the ovary, the latter is called monomerous;°
the side traversed by the mid rib of the carpel is then the pos-
terior side, the line of coalescence of the leaf-edges the ventral
suture. Through the formation of false dissepiments or parti-
tions, the monomerous ovary, which is’ usually unilocular, can
become multilocular. When several carpels become united, the
ovary is polymerous.’

1 Mévos one, and &deA@os brother.

* ToAvs many, and adeAgos,

3 Dury and av7p.

4Tvyyn a woman, and 6r7uw@yv stamen.,
5Turvy wife, yvvaixetov female household.
® Pistillum, pestle. .

' Kapros fruit.

8 Mévos one, and”“épos part.

* ToAvs many, and sépos part.

92 MORPHOLOGY.

If the ovary is superior, i. ¢., if it occupies the uppermost part
of the flower, and the stamens and perigon are inserted below it,
the flower is called hypogynous ' (Potentillee) (Fig. 8, a); if,
however, the andreeceum and perigon are elevated above the
ovary upon a cup-shaped, annular wall, by means of an axial
ring of the hypanthinm located below the ovary, which latter
then remains at the base of the cup, the flower is called perigy-
nous * (Rosex) (Fig. 8, 2); if the cup is now firmly closed at the
top, there is formed the epigynous flower * (Pomew) (Fig. 8, c).
In the two latter cases, the ovary is ¢nferior.

The ovary is unilocular (Fig. 21) when the carpels -are united
by their edges; multilocular when ‘the carpels are turned in-

Fra. 8—a, hypogynous; 6, perigynous; c, epigynous flower, longitudinal section
(Prantl).

ward, and are in contact by their surfaces (Fig. 20). Through

_ the formation of false dissepiments, a polymerous ovary can also
become further divided (Labiate : 2 carpels and 4 compartments .

or cells [nutlets or achenia], Linwm), or a unilocular ovary can

become multilocular (Crucifere, Papaver). The false dissepi-

ments, which, moreover, often produce but an incomplete divi- —

1"Y'x6 under, yorn a woman (pistil),
tg Tepi’ about, yurn
*"Exi upon, yur7.

THE FLOWER. V3

sion (Papaver), are for the most part extended growths of the
placenta.

If a flower contains stamens and an ovary, it, is called herm-
aphrodite’ (%, as in most of the higher plants); when these
are contained in separate flowers, they are called diclinous or
unisexual® (Urticine). If diclinous flowers, the male ( 2 ) as:
well as the female (2), are found on one and the same plant, the
term monecious is applied * (Juglandez); if the male and female
flowers occur on different plants, they are called diwcious *
(Saliciner, Cannabis, Humulus); when upon the same.plant
there are found unisexual as well as hermaphrodite flowers, the
plant is called polygamous *° (many Composite).

When several carpels occur, these are mostiy coalescent in the
ovary, as also in the style (syncarpous gyneceum, as in Malva).
They can, however, be free from each other (apocarpous gyne-
ceum, as in Rubus), and generally in all polycarpic flowers),
or only coalescent in special places (partially apocarpous, as in
Asclepias Cornutt).

The style, the upward extension of the carpels, bears the
stigma. The stigma iseither simple (Fig. 10) or branched (Fig.
9) (Crocus, Euphorbiacex), feathery, tufted (Graminez), often
capitate, or even extended in a disk-like form, as in Asclepias
and Vincetozicum.® It is, as a rule, provided with papille
(Fig. 9 in 3), and secretes a glutinous liquid. If the pollen
grains of the anthers fall upon the stigma, they develop long
tubes (pollen tubes, Fig. 10, uw), which penetrate downwards in
the conducting tissue of the style (g), as far as the ovules, and
here effect the fertilization. The fertilized ovules then develop
to form seeds.

Fertilization with other plants of the same species affords,

'Epuagpposiros bisexual.
Ags double, and Ary7 bed.
3 Mévogs one, and ofxo0s household.
4 Ais double, two, and otKos household.
5 [ToAvs many, and ya@uos marriage.
_ In the last-mentioned plants, the stigma also bears clamplike
bodies, to which the pollinia are attached. ei

4 MORPHOLOGY.

even in hermaphrodite flowers, a greater supply of seed. Fer-
tilization with the pollen of plants of other species produces,
when successful, so-called hybrids. The transmission of the
pollen takes place either through the agency of the wind (ane-
mophilous plants), or more frequently, by insects (entomophilous
or zoidiophilows * plants).

If a flower is projected on a plane, 7. ¢., if represented in such
a manner that, when seen from above, only the points of inser-

Fie, 10.

Fie. 9.—Crocus sativus. 1, the three-limbed stigma ; 2, summit of a limb of the
phe sores strongly magnified ; 3, a section of the edge of the stigma, with the papil-

Fig. 10.—Schematie figure for the elucidation of the process of fertilization. n stigma
with the papillze (w), and three pollen grains already provided with protruding pollen
tubes (n). Of the latter, one has already penetrated through the micropyle (m), and
fertilizes the embryonal vesicle (6) contained in the embryo-sac (q). p outer, s inner
integument ; ¢ inner, h outer funiculus ; ¢, nucleus ; at a later period, occasionally pass-
ing into perisperm (the endosperm originates from the embryo-sac) ; 0, pericarp.

tion of the separate parts of the flower are delineated by corre-
sponding outline figures, a diagram? of the flower is obtained.

"Av Enos wind, Sdov animal, and giAos friendly, loving.
* From éza@ through, and ypa@evr to write,

THE FLOWER. 4D

In the diagram the stamens are represented by small circles, the
staminodia or abortive stamens by crosses (Fig. 11), and the
perigon leaves by segments of circles (Figs. 12 and 13). The
abortive,’ 7, e., the defective or suppressed parts of the flower,
are designated by dotted lines.

Ordinarily, the individual parts of the flower are arranged in
circles (cycles),? the members of which alternate with each other
(Figs. 12 and 12), so, indeed, that each member of a successive
circle, with regard to economy of space, always lies between two
members of the preceding circle. Ina typical dicotyledonous
flower (Fig. 12), for example, the five corolla leaves lie between

Fie. 11.

Diagram of a flower of the Graminez. 6, palea inferior; v, palea superior ; /, lodi-
culze or perigon leaves developed as small scales (Tschirch),

the five calyx leaves ; the first circle of stamens lies between the
corolla leaves—therefore in front of the calyx leaves or sepals,
the second circle in front of the corolla leaves or petals, etc.
Such a flower is called diplostemonous,* as in most of the phe-
nogams. If, however, the first circle of stamens lies above the
corolla leaves (thus epipetalous), and the second above the calyx

1 Abortus, miscarriage.
2 KvxAos circle, ~
8 ArmAdos double, 6r7ucv stamen, thread.

76 MORPHOLOGY.

leaves (thus episepalous), the flower is called obdiplostemonous ’
(Gruinales, Crassulacew, Saxifragacez).

According to the number of circles of the parts of the flower,
the latter is spoken of as tri-, tetra-, or pentacyclous ;? and accord-
ing to the number of members of the circle, as tri-, tetra-, or
pentamerous’ circles. The typical dicotyledonous flower is
pentacyclous-pentamerous (Fig. 12), the typical monocotyledo-
nous flower (Fig. 13) pentacyclous-trimerous.

In addition to a drawing (projection of the flower upon a
plane), or by means of a diagram, the structure of the flower

i O
*

f x
i 1
' ,
A US77 7 oe
J - O f .
Fia. 12. Fie. 18,

Fie. 12.—Typical diagram of a dicotyledonous flower.
Fia. 13,—Typical diagram of a monocotyledonous flower (Liliacese) (Tschirch).

can also be expressed by formulas,‘ as e. g., the typical dicotyle-
donous flower :
K (calyx) 5
C (corolla) 5
A (andreceum) 5 +5
: G (gyneceum) (5 )
which indicates that the androeceum consists of 2 pentamerous
circles, and the gyneceum of a superior ovary [ (5) ], formed of
5 coalesced carpels (Fig. 12). Ifa circle is deficient, it is de-
1 Ob, opposite.
* Tpis, rérpa, wévre (three, four, five), xvxAos circle.
*Tpis, rétpa, wévre (three, four, five), uépos part.
*See under Androeceum in Eichler’s ‘* Syllabus.”

THE FLOWER, res

signated by the mark 0, for example, in the flower of the
Grasses (Fig. 11):
K 90
C 2
A 3+0
G (2)
or, since there is usually no differentiation of calyx and corolla
in the monocotyledons': P (perigon) 0+2, A 3+0, ete.
If the ovary is inferior, it is expressed by a stroke above, for

example, G (3).
} {
(Eo %))
a af!

|

Fia. 14. Fia. 15.
Fic. 14.—Diagram of a cruciferous flower, with diagonal corolla.
Fie. 15.—Diagram of a papilionaceous flower (descending estivation in the corolla,
ascending in the calyx). v, vexillum or standard ; aa, alee or wings ; c, carina or keel
(Tschirch).

Coalescence is indicated by parenthetical marks, as (2) = two
coalesced leaves ; deduplication or chorisis,? French dédouble-
ment,* i. e., division of an organ into two or several parts by

1 Mévos alone, single, and xorvAy.
? Xoptl@ to separate,
5 Dédoublement, duplication.

78 MORPHOLOGY.

an added exponent, which denotes the number of the parts, for
example, 2°,

If a flower can be divided into two equal halves (reflex images)
by several dissecting lines drawn through its central point, it is
called regular or actinomorphous * (®) (Figs. 12 and 13). Such
flowers, however, which can only be symmetrically divided by
one section, are called monosymmetrical or zygomorphous* (A)
(Figs. 11, 14, 15).

If the only possible symmetrical section lies in the median
plane (|), the flower is called median-zygomorphous (7), as
in the Labiate and Papilionacer, otherwise oblique-zygomor-
phous (A), as in Hyoscyamus, or transverse-zygomorphous
(—>), as in Fumaria. Vf flowers can in no manner be symmetric-

ally divided, they are called asymmetrical,’ as in the Zingiber-
acez.

The place where the bract accompanying the flower is located
is represented in the diagram either below or in front. The
first leaf of the flower is then located for the most part opposite
the bract, either above or in the rear.

Through subsequent inversion (resupination), the position of
the flower to the bract is occasionally reversed. Thus the label-
lum of an orchideous flower in its natural disposition is located
in the rear and above, and only subsequently, through reversion
of the ovary or of the peduncle, is brought forward and below.

The individual flowers often combine to form the so-called
inflorescence. According to the form of ramification, the fol-
lowing are distinguished :

___ IL Racemose Ixriorescence.—The main shoot (axis, rachis)
is not over-topped by any of the lateral shoots produced thereon,
in an acropetalous manner.
f 1. Spike (spica). The flowers sessile on the axis
Elongated { (Fig. 16 a) (Carex), to which belongs also: the
axis. catkin (amentum), when pendulous and falling
off as a whole (Juglans).

1” Aurts ray, and zopp7 form.
? Zuyov yoke, and opp form.
* A privativum, and éUusuerpos symmetrical.

THE FLOWER. 79

inclosed by a sheath (spathe) (Aroide).
3. Raceme (racemus). The individual flowers have
long pedicels (Crucifere) (Fig. 16 6).

(4. Head or Capitulum. The flowers sessile on a flat-
tened or head-shaped axis (Composite) (Fig.
16 d and e).

5. Umbel (umbella). On the axis, which is abbrevi-
ated to 0, originate numerous flowers with stalks

Shortened 4° 0F pedicels of equal length (Umbellifere). The
axis. umbel is often compound, so that small um-

bels (umbellets) stand at the end of every ray.

The umbels and umbellets are then mostly sur-

rounded by a circle of bracts, called the involu-

cre or involucel (involucrum, inyolucellum)

(Fig. 16 c).

II. Cymose INFLORESCENCE.—The main axis is over-topped
by one or several more strongly developed lateral axes.
(1. Cyme (cyma’). Below theterminal flower origi-
nate numerous, mostly equally strong lateral
Without | shoots (Zuphorbie) (Fig. 16 q).

Pseud-axis. } 2. Dichusium.* Below the terminal flower origi-

nate two equally formed lateral shoots (false

| dichotomy) Valerianella (Fig. 16 m).

(3. Bostryx.* The over-topping lateral shoots of

| successive members arise on the same side of
their main axis (Fig. 16 7).

4, Cincinnus.* The over-topping lateral shoots
arise alternately on opposite sides of their

| main axis (Asperifoliacer) (Fig. 16 # and 7).
By the combination of several types of inflorescence with each

other is formed a compound inflorescence. To this belong the -

corymb (Sambucus) and the anthela.

Elongated

2. Spadix. The flowers sessile, axis fleshy, and
axis. |

A With
Pseud-axis.

pes ©

1 Kove the young stalk of the cabbage.

* From 65 twofold, and ya@67s cleft, separation.
* Boérpvé curl, tendril.

4A curl of hair.

30 MORPHOLOGY.

Finally, there is also to be mentioned the particular form of

cyme ; hk, andi, cincinnus; k, true dichotomy ; 1, bostryx ; m, false dichotomy.

Fia. 16.—Schemes of inflorescence. a, spike ; b, raceme ; c, compound umbel ; dand e, heads; /, compound raceme ; g,

inflorescence of the Euphorbie, which is designated as a cya-

- THE FRUIT. 81

thium. In this case, the cup-shaped hypanthium bears numer-
ous male flowers (each with one stamen), and a long-stalked
female flower.

Pharmacognostically, there belong to the flowers and forms
of inflorescence, the developed, complete, officinal, separate
flowers of the phenogams, and likewise the buds of individual
flowers, for example, Caryophylli. Furthermore, undeveloped
forms of inflorescence, Flores Cine (Santonica), as well as the
expanded inflorescence, such as Flores Arnice, Flores Chamo-
mille. Flores Koso (Brayera) consist of the inflorescence from
which the petals have fallen. In the case of the compo-
site flowers, the involucral scales are still present in the drug,
perhaps with the sole exception of Flores Arnice. Finally,
Flores Rhewados, Flores Verbasci and Flores Rose represent
only the petals or corollas, and Orocws only the stigmas,

From the ovary with the ovules there is produced, after fer-
tilization has taken place, the fruit containing the seed.

Fruits, aggregate or collective fruits, or parts of fruits of the
angiosperms and gymnosperms are officinal with or without the
seeds. For the rind (pericarp) of the Aurantiex, which in the
fresh state is juicy, the customary, though incorrect designation
of cortex may be retained here, in order to avoid the introduc-
tion of a new term. :

By the term “fruit” we mean here only the ovary which, as
a result of fertilization, is in process of maturing, or has already
become fully matured. Its outer wall, and the dissepiments and
placenta, may thereby suffer the most manifold changes, by
which other parts, including those not belonging to the flower, are
also frequently affected, as, e. g., in the case of the fig, Fructus
Juniperi, the apple and strawberry, which are therefore to be
designated as pseudo-carps.

In the jig, as likewise in the strawberry and the apple, the
upper -part of the axis participates in the formation of the
(pseudo-) fruit (hence the name : hypanthodium'), which in all
three cases assumes a fleshy character. In Juniperus, however,
_ it is the three bracts of the flowers which become fleshy.

'From uo under, and aeyS$os flower. |
ee ;

82 MORPHOLOGY.

The wall of the ovary which becomes developed to form the
seed-vessel is called the pericarp (pericarpium'). On the latter,
from the exterior to the interior, there may often be distin-
guished three layers, differing in their structure or their color,
the epicarp (epicarpium ’), mesocarp (mesocarpium *), and endo-
carp (endocarpium’).

The outer coating of the fruit often shows completely the
structure of the epidermis, 7. ¢., it isprovided with a very strong
cuticle and with stomata; often, however, it is formed to a
predominating extent of stone-cells (sclerenchyma). The variety
of the tissues and their contents is still greater in the middle
layer (mesocarp), which in many fruits consists of fleshy, juicy,
or even very loose tissue. When its cells contain an abundance
of juice and finally lose their coherence, it is designated as pulp,

: Fia. 17,—Cé ium maculatum. a, endosperm ; c, commissural surface ; r, coste or,
ribs with the vascular bundles (f v); f, vallecule or grooves (Hager).

as, é. g. in the legume of the tamarind. The inner layer of the
fruit (endocarp) originates from the epidermis of the cavity of
the ovary, and often develops into a hard stone-shell, as in the
case of the almond. It is, however, not always the case that a
discrimination can be made between the three layers of the
ripened seed-vessel, and their relative development is very
variable.

Of aggregations and forms of fruits, the following are distin-
guished :

' IZept about or around, xapzds fruit.
*Exi upon.

4 Mé6os in the middle.

* Evdor within.

THE FRUIT. 83

The multiple fruit (syncarpium'), formed by the coalescence
of several monomerous ovaries (Star-anise, Rubus ideus).

The mericarp (mericarpium ’), double akene, is formed by the
separation of the compartments of a multilocular ovary ; the
separate fruits thus produced are called schizocarps* (Umbelli-
fere, Fig. 17).

With these may also be classed the jointed or articulated
fruit (Raphanus raphanistrum), the mericarps of whichare
akenes.

The axis on which the two schizocarps of the Umbellifere
are suspended is called the carpophore.* The main or primary

Fic. 18.—Pisum sativum. Legumes. a, tip; b, base; v, ventral suture ; d, dorsal
; suture.

ribs of umbelliferous fruits are called coste or juge@ ;° the sec-
ondary ribs, coste secundaria ; the grooves lying between them,
vallecule* (Fig. 184). In the latter are located, when present,
the oil-tubes or vitte.’

1 Sur together, xap7os fruit.

2? Mépos part, xapzos fruit.

8 Sylow I split, xapzos fruit.

4 Kapzos fruit, and pépecr to bear.
5 Juga, ridge.

* Diminutive of vallis, valley.

' Vitta, band.

84 MORPHOLOGY.

There are furthermore distinguished :
I. Dry Fruits: Pericarp woody, coriaceous.
(a) Indehiscent fruits, not opening by valves or regular
lines.

1. Nut (nux), having a hard pericarp (Cannabis).

2. Caryopsis’ (or Grain) and Akene,* having a Jeather-
like, membranaceous pericarp (Gramine, the ce-
reals).

With these may also be classed the Samara, or so-called
winged-fruit, an akene which, through a subse-

quent growth of the pericarp, appears winged
(Ulmus, Betula).

3. Mericarps (see above).

Fia. 19.—Silique of Brassica oleracea. 1, closed ; 2, opened, one valve removed ; v,
the other valve ; d, partjtion with the seed. -

(6) Dehiscent fruits: opening by valves or regular lines,
containing several seeds. .

1. Follicle (folliculus*), formed of one carpel, and
dehiscent by the ventral suture (J/licium ani-
satum).

2. Legume (legumen), formed of one carpel, but also
dehiscent by the dorsal suture (Fig. 18), often
with false dissepiments (Leguminose).

3. Silique (siliqua), of two carpels. The latter sepa-
rate from each other first at the base ; on the parti-

1 Kapvor nut, ovis appearance.

*tAyaivior (from @ privative, yazva@ I open), a fruit which does not
open

5 Diminutive of follis, a sack or tube.

-

THE FRUIT. 85

tion which remains attached, the seeds are loca-
ted (Crucifere) (Fig. 19).

4. Capsule (capsula), formed of several carpels, dehis-
cent longitudinally from above (Fig. 20), more
rarely from below, or ultimately opening by a lid
(Pyxidium, Fig. 21: as in Hyoscyamus, Anagal-
lis), or by holes (pore-capsule, as in Papaver
somniferum).

The dehiscence (dehiscentia) of the capsule is septicidal' (or
through the dissepiments, in fruits opening by their sutures,
Melanthiew, Fig. 22 ~), when in the case of a multilocular ovary
_ the coalesced dissepiments become separated from each other

Fie. 20. . Fie. 21,

Fie. 20.—Colchicum autumnale, capsule dehiscent (septicidally) from above (Hager).
Fic. 21.—Hyoscyamus niger, capsule dehiscent by a lid. a, closed ; b, opened.

(Colchicum, Sabadilla); loculicidal? (infruits dehiscing through
the cells of the pericarp: Liliew), when each carpel becomes
split in the middle (Lilium, Scilla, Aloe). If in the latter case
the column of the dissepiments with the seed, separated from
the wall of the capsule, remains standing in the middle, the
dehiscence is called septifragal* (Fig. 22 c). In septifragal
dehiscence, the opening may take place from below (Geranium),
or from above (Balsaminex, Epilobium).

II. Fleshy Fruits : Pericarp mostly fleshy.

1 Septum, dissepiment, and cedere, to cut or to break.
* Loculus (diminutive of locus), compartment, and ccedere,
’ Septum, and frangere, to break.

86 MORPHOLOGY.

1, The Stone Fruit or Drupe (drupa’), endocarp very

hard, not dehiscent (Amygdalus, Juglans).
2. The Berry (bacca), endocarp and mesocarp fleshy, epi-

carp often hard (the grape, currant, date).
Occasionally the fruit is inclosed or surrounded by a body
which is mostly cup-shaped, termed a cupule (cupula). In the
case of the oak, this is formed from four coalesced bractlets.
The tanning material known under the name of “ valonia,” con-
sists of the cupules of the fruits of Quercus Vallonea Kotschy.
In the seed-vessel, formed of the carpels, are contained the
ovules (ovula*). The latter consist of the fwniculus* or podo-
sperm, with which they are attached to the wall of the ovary,
or to some special part of the ovary called the placenta,’ which,

Re Ne

Fia. 22.—Forms of dehiscence of the capsule. a, septicidal ; b, loculicidal ; c, septi-
fragal.

according to its position, is termed basal, central, or parietal ;*

the integuments," forming one or two coats (Fig. 10 p s), which

in front do not close completely, but leave an opening (the
micropyle,’ Fig. 10 m); and the kernel or nucleus (Fig. 10 #),

containing the embryo-sac (Fig. 10g), in which the embryo*®

is formed (compare Figs. 23, 24, 25).

' Drupus, ripe and ready to fall.

*Diminutive of ovum, egg.

* Diminutive of funis, a rope or cord.

* Placenta, a cake, from a remote analogy with the placenta of the
higher animals,

5 Paries, wall,

° Integumentum, a covering or skin,

* Mrxpos small, Uy gate or entrance,

*“EuZpvor the unborn fetus in the womb.

THE SEED. 87

These ovules, after having been fertilized, constitute the seed.

That portion of the fruit of phanerogams which is formed
from the ovule and contains the developed embryo is called the
seed. Before the latter are utilized, they are, for the most
part, completely freed from the seed-vessel. In the case of
some seeds, the seed-shell or testa and the inner membrane are
also removed.

The seed consists of the seed-coats or integuments and the
embryo, and frequently, in addition thereto, of the albumen.’
The former usually consists of an external, firm, occasionally
very hard seed-shell or testa,” whichis invested with a thin, but
often very tough inner membrane ; this may readily be removed,
especially after softening in water,-as in the case of the
almond, coffee and Semen Ricini, so that the kernel of the seed
alone remains. Semen Quercus consists, in the commercial
form, exclusively of the kernel, the two cotyledons without the
membrane of the seed. With Semen Myristice or the nutmeg
(as also with Cacao), on the contrary, the membrane penetrates
into the kernel or nucleus, and in the case of the former, for
example, cannot be separated in a connected form.

The testa is formed from the integuments of the ovule. For
its first development the embryo requires a special supply of
nutritive substances, which may be stored in the tissue of the
embryo itself. In this case, a particular albumen is not present,
the seed being thus destitute of albumen, or exalbuminous, as,
é. g., Semen Quercus, the almond and mustard (and in general
all the Cruciferz). ;

If, however, there is developed, simultaneously with the em-
bryo, a special tissue filled with reserve material, this is called
albumen. If this tissue, with regard to its origin, belongs to the
embryo-sac, as is usually the case, it is termed endosperm *
(Umbellifere, Fig. 17 a); if, however, a portion of the nucleus
(kernel of the ovule) has been converted into albumen, it is dis-

1 Albus, white (the white of egg).
2 Testa, a vessel, or also a shell.
’”Eyvéov inside, 6repua seed.

88 ‘MORPHOLOGY.

tinguished as perisperm.! The seeds of the cardamom and
of pepper represent simultaneously both forms of albumen, peri-
sperm as well as endosperm. The chief contents of the
albumen-cells belong nearly always to the class of protein
substances, and are frequently developed, in part, in a crystal-
loid form. With these is usually associated fat, and not rarely
also amylum, sugar, and mucilage. This abundance of contents,
which, moreover, are very commonly deposited in thick-walled
cells, mostly imparts to the tissue of the albumen a firm, horn-
like character. In the German usage of the word, there is
accordingly sometimes understood (rather ambiguously) under
the expression “‘albumen,” the entire tissue of the seed which
contains the previously mentioned reserve substances, and some-
times, in a chemical sense, that class of nutritive substances
which are also termed protein bodies.

The degree of development of the albumen is very variable.
It is often much more extensive than the embryo, as in Semen
Myristice (the nutmeg), Semen Oolchici, and in Nuzx vomica;
in other cases, it appears only as an insignificant appendage, as
in Semen Lini, or perhaps also disappears at a later period, so
that it is no longer observable in the ripe seed.

The embryo contains, in a more or less advanced state of
development, the rudiments of the axis and leaf-organs, the
former shortly attenuated in one direction as the radicle or cau-

‘licle (this is always directed toward the micropyle), and in the
opposite direction often bearing the rudiments of the stem and
leaf structures, or plumula ;* the latter may very plainly be seen,
for instance, in the almond, and also in Nuzx vomica.

The leaf-organs, embryonal leaves, seed-lobes or cotyledons,°*
usually form the preponderating portion of the embryo, and
occur, especially in many dicotyledons, already developed in a
delicate, distinctly leaf-like form, as in Vux vomica and Semen
Ricini. In seeds destitute of albumen, e. g-, in the almond,
bean, pea, and acorn, on the contrary, the cotyledons are of a

* Hep around or about, and 6zépua seed.
* Diminutive of pluma, feather.
* KorvAy cavity, xorvAydwy cavity of a bone, pan.

THE SEED. 89

thick, fleshy character. With monocotyledonous plants, the
embryo in the seed is usually less clearly developed ; in Semen
Colchici, Semen Sabadille, and in the cardamom, the cotyledon
is not yet really distinctly leaf-like, and quite as little in the
pepper and cubeb. The tissue of the embryo throughout is built
up of more delicate cells than that of the albumen, and this
difference is also readily apparent without being magnified.

The cotyledons and the radicle are often bent in a character-
istic manner, as is evident upon a longitudinal section through
Semen Stramonii, while the fruits of the Umbellifere present
an example of a tolerably straight embryo. A very remarkable
folding is shown by theembryos of Semen Faenugreci and Semen
Sinapis, as in general with all Orthoploce, Spirolobee and
Diplecolobeew. Outside of our sphere of consideration, there
occur remarkably complicated foldings in the cotyledons of the
cotton-seed.

The seed is connected with the placenta by means of the
funiculus ; the place where the latter enters the testa usually
remains characterized by its color, a depression, or an elevated
line, and is distinguished as the hilum (Fig. 10 h, Fig. 24 h).
Less frequently the terminal point of the funiculus is also per-
ceptible in the base of the seed ; if this is the case, it bears the
name of inner hilum or chalaza’ (Fig. 24 ch). This is readily
recognizable, among other instances, in Semen Ricini.

The seed is straight, atropous or orthotropous,” when the apex
of the ovule, the orifice (micropyle), lies opposite the hilum,
whereby the funiculus remains short. It is thus, e. g., with the
Piperacese, where the seed forms the termination of the flower
axis. More frequently, however, the ovule together with its
coats, 7. e., the entire seed, is reversed, whereby its apex, the
micropyle, is moved close beside the hilum. This form, with
the funiculus running along the back, which is the most usual
with the angiosperms, is designated as a reversed, anatropous*
seed (Fig. 24). The ovule is here coalescent with the funiculus

1 X@Aate hail, but also a sty on the eye-lid.

* From a@ privative (and dpSds straight), and rpéz@ turn, direct.

* Ava against, rpéro I turn. oe

90 MORPHOLOGY.

whereby a suture, rhaphe,' is produced (Fig. 24 7), which is more
or less apparent, e. g., in Semen Tiglii and in the cardamom.

The reniform or kidney-shaped seeds, on the contrary, are
mostly produced from so-called campylotropous * or curved
ovules. With these, the nucleus as well as the integuments are
curved (Fig. 25). They therefore also possess, as a rule, a curved
embryo.

With relation to their attachment, the ovules are sometimes
pendulous, sometimes erect or ascending, and sometimes hori-
zontal. If in an anatropous (pendulous) ovule the funiculus lies
toward the interior or the middle-line of the fruit, such an ovule
is termed epitropous* (Umbellifer, Euphorbiacex), If the funi-
culus is directed toward the outer wall, it is called apotropous
(Vitis, Rhamnus, Cornus).

Fie, 28. Fia. 24. Fig, 25.
@
() b, 7 \ fs
Atropous ovule. Anatropous ovule, Campylotropous ovule.

m, micropyle.; ch, chalaza ; h, hilum ; r, raphe.

Appendages of the Seed.—Many seeds are provided at the
hilum with an indurated appendage (as in the pea), which, in
the case of Semen Ricini, Semen Chelidonii (obdurator, carun-
cle or caruncula ‘*) and Semen Colchici, still remains perceptible,
even after drying, while in Semen Tiglii, on the contrary, it
readily falls off.

A peculiar, compact, fleshy outgrowth is developed on the
nutmeg, and is designated as the seed-covering or arillus (Fig.

1*Pagy suture,

? KaumviAos bent, and TPETO,
*’Exi upon, and rpézo.
+Caro, flesh.

THE SEED. 91

26 ar). This arillus, which is known in commerce under the
name of mace, represents the only structure of this character
which is properly treated of here. A relatively still more devel-
oped arillus, but consisting only of a thin membrane, incloses
the seed of the cardamom. The small, red, cup-shaped body
of Taxus fruits is also to be regarded as an arillus.

The arillus always originates at the base of the seed, and is to
be considered as an outgrowth of the funiculus.

The pappus, a form of appendage of fruits (page 69), is
formed by a subsequent outgrowth of the calyx.

The germination of the seed proceeds in this way, that with a

Fie. 26.—Fruit of Myristica fragrans, longitudinal section. ar, arillus; s, seed
(Hager).
simultaneous evacuation of the endosperm which may be pres-
ent, the plumule and radicle break through the testa of the
seed, the former developing to form the stem, and the latter to
form the root. Thereby the cotyledons, which are sometimes
fleshy and thick (as in the Bean), sometimes thin and leaf-like
(Ricinus), are either elevated above the ground and become
green (Epigea), or remain in the ground until their evacuation
and rejection (Hypogea, compare page 69). Occasionally, the
cotyledon still incloses for some time the young leaf-bud
(Maize).

PLANT ANATOMY.’

In order to obtain a satisfactory knowledge of vegetable
drugs, an accurate anatomical study of them is in most cases
indispensable. This part of pharmacognosy is therefore based
upon an acquaintance with the principles of plant anatomy.
The following lines may serve for a preliminary acquaintance |
with this very extended department, more complete information
being contained in the text-books of anatomy.? The beginner
should, nevertheless, continually bear in mind that anatomical
_ study, unless accompanied by work with the microscope,* must
always remain poor and deficient inits results. It would, there-

1From ava and réure@ cut.

*De Bary, ‘‘ Vergleichende Anatomie der Vegetationsorgane,” Leip-
zig, 1877. The most comprehensive and fundamental work, which,
with regard to the amplitude of its contents, can be compared with no
other, An excellent English translation of this work bears the title:
““Comparative Anatomy of the Vegetative Organs of the Phanerogams
and Ferns;” by A. De Bary. Translated by F. O. Bower and D. H.
Scott, 1884 (F. B. P.).—Sachs, ‘“ Lehrbuch der Botanik,” iv., Leipzig,
1874 (at present only to be had through antiquarian book-sellers). An
English edition of this work bears the title : ‘‘ Text-book of Botany,” by

Julius Sachs. Translated by A. W. Bennett, assisted by W. T. T. Dyer,
Oxford, 1875. Second edition, 1882. F. B. P.—Haberlandt, ‘‘ Physio-
logische Pflanzenanatomie,” Leipzig, 1884.—Weiss, ‘‘ Anatomie der
Pflanzen,” Vienna, 1878.—Leunis, ‘‘ Synopsis,” newly edited by Frank,
One volume, Hannover, 1882, For our purpose, there may also be men-
tioned: Hanausek, ‘‘ Anatomische, physikalische und chemische Ver-
haltnisse des Pflanzenreiches, mit besonderer Riicksicht auf Waren-
kunde und Technologie,” Hélder, Vienna, 1882,

*In microscopic work, the following are very useful: E. Strasburger,
“Das botanische Practicum,” Jena, 1884; and, by the same? author,
“* Das kleine botanische Practicum,” Jena, 1884.

THE CELL. 93

fore, really seem most proper to provide this chapter with an
introduction to the construction and use of the microscope.
There have recently appeared, however, so many publications
relating precisely to this subject,’ that we may omit further ref-
erence to it in this place.

The experience acquired at the preparation table is in the
end the most important of all ; and as chemical analysis cannot
be learned without a laboratory, so also the methods of micro-
scopical investigation cannot be learned without a microscope
‘and dissecting needles.

I. The Cell.

~The elementary organs from which the body of the plant is
constructed are the cells. Although it is not necessary for the
. formation of the idea of a cell that the same should be inclosed
by a membrane (naked swarm-pores), nevertheless, by far most
cells are provided with such.

Most plants (all the more highly organized ones) consist of
numerous cells. Among the lower plants there are, however,
many which are formed of but single cells, some of which assume
the most manifold forms, branch abundantly (the mould fungus
Mucor Mucedo), and, indeed, without being in any manner
divided by lateral walls, imitate a stem, leaf, and root ( Caulerpa).
Of such one-celled plants, there are none which come under
consideration in pharmacognosy in a restricted sense, although
the yeast fungus (Saccharomyces cerevisiw), on account of its
fermentative action, and the various pathogenic fungi (dacteria),
which claim an increased degree of interest in consequence of
the recently observed relations between them and the most dan- —
gerous diseases, as well as the Diatomezx, whose siliceous coats

1 Behrens, ‘‘ Hilfsbuch zur Ausfiihrung mikroskopischer Untersuch-
ungen,” Braunschweig, 1883. The American edition of this work has
been noticed on page 49, F. B. P.—Dippel, ‘‘ Das Mikroskop,” II., Braun-
schweig, 1883-84.—For the theoretical part, the following is very valua- —
ble: Nageli and Schwendener, ‘‘Das Mikroskop,” Leipzig, 1877.—
Further: Hager, ‘‘Das Mikroskop und seine Anwendung,” Berlin, —
1879.—J. Vogel, ‘‘Das Mikroskop,” Leipzig, 1885. .

94 PLANT ANATOMY.

form the so-called infusorial earth, are of the greatest import-
ance in practical life.

THE CELL-WALL AND CONTENTS OF THE CELL.

I. Contents of the Cell.

The cell consists of the cell-wall, and the contents of the cell.
The most essential constituent of the cell while exercising the
functions of life, is the protoplasm (plasma’). This repre-
sents a turbid, semi-liquid mass, which completely fills the inner
portion (lumen) of cells located at the growing point, or which
are otherwise in a state of active development. At a later
period, there appear in the protoplasm cavities or vacuoles.*
The latter, filled with colorless cell-sap, constantly continue to
enlarge as the cell becomes older, coalesce with each other, and
finally, while the protoplasm gradually contracts toward the
wall of the cell (protoplasm-sac, primordial utricle, cell-sac),
form a large, central cavity, filled with cell-sap (Fig. 27 p). If
the cell has ceased to grow, the protoplasm will also have disap-
peared, with the exception of a delicate film attached to the
membrane. The protoplasm takes the most active part in all
formative processes in the cell, and is the most important sub-
stance in the cell; the formation of the cell-wall proceeds from
it, and to it the other constituents of the cell, for the most part,
owe their origin.

The cells increase in number only by division. A cell (mother-
cell) becomes divided by a septum (which is mostly median)
into two daughter-cells (Fig. 28). And it is considered asa law
that with the division of the cell (of the protoplasm) a division of
the nucleus (see below and Fig. 28) is also always associated. In
the case of reproductive cells, a rounding of their form is also
generally associated with their division.

The protoplasm, which always possesses a semi-liquid, and
(with the exception of the outermost and innermost layer, hya-
loplasm) a granular character (microsomes), is a body of compli-

' p@roy the first, and 2la6ua organization or form.
* Vacuum, empty.

THE CELL. 95

cated composition, very rich in nitrogen,’ which contains several
substances belonging to the albumen group (protein substances),
together with water and inorganic salts (phosphates and sul-
phates of the light metals). Itis not devoid of structure, but
possesses a fine organization.*

Substances capable of abstracting water (such as sugar and
glycerin) contract the protoplasm, 7. ¢., in consequence of the

Fic. 27.—Transverse section through a medullary cell of Toxodium distichum. a,
nucleus ; b, nucleolus ; c, protoplasm-sae contracted toward the wall (separated from
the latter by reagents) ; i, primordial utricle (hyaloplasm) ; p, cell-sap; -m, correspond-
ing tips of adjacent cells ; d, the cell-wall ; e-s, the cell-walls of adjacent cells ; g, inter-
cellular space (Hartig), :

elimination of water from the contents of the cell, the proto-
plasm-sac is drawn from the cell-wall. Protoplasm is colored

1 Compare Reinke, “Studien tiber das Protoplasma,” Berlin, 1881.

2 Very many investigations have recently been published, relating to — : :

the structure of protoplasm (especially by Strasburger, Schmitz, Tangl,

96 PLANT ANATOMY.

yellowish-brown * by iodine,* rose-red by Millon’s reagent, violet
by Trommer’s reagent, and red by sugar and sulphuric acid.
Dead protoplasm abundantly absorbs coloring substances (espe-
cially fine with eosin). Imbedded in the protoplasm of young
cells (Fig. 27), or suspended by threads of protoplasm (Fig. 28,
b, c), there is found the nucleus (Fig. 27 a, and Fig. 28), which
mostly occurs singly. The nucleus possesses one or two nucleoli
(Fig. 27 5), and consists likewise, for the most part, of a proto-
plasm-like substance, in which, however, the nuclein is con-
tained ina granular form. By treatment with coloring sub-
stances (hematoxylin, aniline-green, alum-carmine), the nuclei
are rendered more clearly visible.

Since the protoplasm has contracted in old cells to the mini-

Sear

Fic. 28.—The process of cell division schematically represented in its individual phases
(Hartig). :

mum and the-nucleus has entirely disappeared, both of these play
but a subordinate part in pharmacognosy, which occupies itself
mostly with organs consisting of completely developed tissues,
notwithstanding the importance of the protoplasm in the econ-
omy of the plant itself, and in the estimation of the value of
herbs as fodder.

‘ All protein substances are colored yellow by iodine, thus protoplasm,
gluten, protein crystalloids (for the latter, iodine in glycerin is em-
ployed), the fundamental portion of the chlorophyll granules, etc.

* For these micro-chemical reactions, the compilation by Poulsen may
be highly recommended (‘Botanische Microchemie,” Cassel, 1881.
American edition by Trelease, see page 49). Compare also Tschirch,

_ ‘Microchemische Reactionsmethoden im Dienste der technischen Micro-
scopie,” Archiv der Pharm.,, 1882.

ALEURONE. 97

There is another body very closely related to protoplasm, and
like this consisting of protein substances, which is likewise of
the greatest importance, pharmacognostically. This is aleu-
rone,’ or protein granules, which are found in numerous seeds of
the Umbellifere, and Euphorbiacex, in Vitis vinifera, Silybum
Marianum, Myristica, Amygdalus, Cardamomum, and the Brazil

os Ss

Fig. 29.—Elliptical, plainly stratified starch granules, st, with a broad, central hilum,
from the cotyledon of a seed of Pisum sativum, after the addition of water. a, protein
substances (aleurone); i, intercellular spaces (Sachs).

nut (Bertholletia excelsa).? In many cases the granular contents
of the cell, when more strongly magnified, may be resolved into

Adevpoy the fine fiour of grain, gluten (German: Kleber) of Hartig,
in distinction from amylon. Aleurone was discovered by Hartig
{Botanische Zeitung, 1855, p. 881, and 1856, p. 257). For the most
thorough examination of it we are indebted to Pfeffer, see Pringsheim’s
Jahrbiicher fiir wissenschaftliche Botanik, viii. (1872), 429.

*The following investigators have contributed to the knowledge of the
crystallized, vegetable albuminous bodies: Ritthausen (many publica-
tions in the Journ. fiir prakt. Chemie of the last few years), Maschke,
Nageli, Sachsse, Weyl, Schmiedeberg, Barbieri, Schimper, Drechsel, De
Luynes, Griibler (Journ. fiir prakt. Chemie, 1881); in the latter, the lit-
erature is collated. Compare also Husemann and Hilger, ‘‘ Die Pilan-
zenstoffe.”

98 PLANT ANATOMY.

numerous separate granules of a roundish or polyhedral form * (as
in the cotyledons of the pea’ and bean, Fig. 29 a), which fill the
intervening spaces between the starch granules (in the pea), or the
entire cell (the glutinous layer in the seeds of cereals). To
these small granules the name of alewrone may likewise be
applied. In a more restricted sense, the name of aleurone is
applied to those large granules which, imbedded in a homo-
geneous mass of albumen, replace starch granules to a certain
extent, and which consist of a fundamental mass of an albu-
men-like substance, inclosing crystalline (calcium oxalate) or
seemingly crystalline, roundish bodies (glodoids). The albumen-
like, fundamental mass is either amorphous or crystalline (erys-
talloids); in the latter case, it is surrounded, together with the
inclosed substances, by an enveloping, amorphous mass.

Fic. 30.—Cells from the albumen of Semen Ricini (Sachs). A, a single cell in concen-
trated glycerin ; the contents show but indefinitely formed masses. B, the game section
with a little water added, whereby crystalloids, fine granules of protein substances and
drops of oil are rendered visible. C, the same section warmed with more diluted glyce-
rin, whereby the drops of oil become expelled, and the crystalloids attacked and gradu-
ally dissolved.

"The _globoids (phosphates of calcium and magnesium) are
never wanting. Crystalloids * oceur handsomely developed in

‘Mounting mediums containing water must, however, be nvoided ii in
the preparation, since the granules thereby become destroyed, as has
occurred, for instance, in Fig. 29. In the examination of aleurone gran-.
ules with inclosed substances, concentrated glycerin or fatty oil is.
always to be employed.

* Compare Tangl, *‘ Das Protoplasma der Erbse.” Sitzungsbericht der
Wiener Akademie, 1887.

*From xpv6raddos crystal, and 7805 similarity. They owe their

ie name to C. Nageli, _ er wage sptanie eo Akad.,” 1862, p. 121.

ALEURONE. 99

the aleurone granules of the seeds of Elwis guineensis, dithusa
Cynapium, and all Euphorbiaces (Ricinus, Croton); they are
wanting in the aleurone granules of umbelliferous seeds. Crys-
talloids occur, together with crystals, in Asthusa Cynapium.
Occasionally an aleurone granule in each cell is distinguished
from the others, either by its size alone, or also by inclosing
crystals of a different formation or larger size (Fig. 31 A at ¢).
Such a granule is termed solitary (German, solitdér, Hartig).
The erystalloids are doubly refractive,’ their angles, however,
are inconstant; they are insoluble in water. The aleurone
granules free from crystalloids, on the contrary, dissolve for

Fie, 31.—A, Two gluten-cells from the seed of the raisin. In the cell at the left much
granular protoplasm anda nucleus cis present, The cell at the right, after complete
ripening, with a large, solitary granule (c), and numerous, small aleurone granules. B,
Aleurone from the seed of Ricinus communis with ecrystalloids. C, Aleurone from
Euphorbie, Myristica (c), Croton (b), Phyllantus (bb). D, Aleurone from the seed of
Bertholletia excel: , f dissolution of a crystalloid into several crystals. Z, Aleurone from
the seed of Liipinis (ec) and Contum (d) (Htartig). :

the most part in pure water (Pwonia, Lupinus), and all of thine oe

in feebly alkaline water. The fundamental mass, consisting of
protein substances, is insoluble in alcohol, ether, benzol, chloro- foe

form, and paraffin; it is colored yellow by iodine. ae
‘They therefore appear more clearly in polarized light. Ds ape ee

ee oe oe emp aay

100 PLANT ANATOMY.

The glodoids dissolve in inorganic acids, also in acetic and
tartaric acids, but not in a dilute solution of potassa.

There occur, moreover, in the vegetable kingdom, crystal-
loids which are not inclosed in aleurone granules (the potato,
Fig. 108 a).

The aleurone granules, as is already indicated by their exclu-
sive occurrence in seeds, belong to the reserve substances which
have the function of presenting to the germinating plant, in its
first development, and before it is capable of assimilating inde-
pendently, sufficient material for building up its organs. They
are, therefore, of the greatest importance in the economy of the
plant. How abundantly protein substances are contained in
some seeds is shown by the following figures, showing their per-
centage: Nux vomica, 11; Cacao, 13 ; Black Mustard, 18 ;
Almond, 24; Linseed, 25 3 Lgnatia seed and White Mustard, 27.

These numbers are calculated from nitrogen estimations, with
the presupposition that albuminous bodies contain 15 per cent
of nitrogen.

With the albuminous bodies are directly connected the ehlo-
rophyll bodies, the fundamental mass of which likewise consists
of an albumen-like substance.' This fundamental mass (stroma),
which, moreover, is also very soft, is of a sponge-like structure
(Fig. 33 a), and contains in the meshes of the frame-work a
small amount of the mixture of coloring substances, to which
Pelletier and Cayentou? in the year 1817 gave the name “ chlo-
rophyll.”* The crude chlorophyll consists of two coloring mat-
_ ters, chlorophyll in a more restricted sense, or pure chlorophyll,*
and xanthophyll.* The former is bluish-green, the latter yel-

E ' Sachs, ‘‘ Flora,” 1862 and 1863.—Mohl, ‘‘ Verm. Schriften.”
* Journ. de Pharm., 1817, p. 486.
* XAwpos green, and guAdor leaf.
“Compare Tschirch, ‘* Untersuchungen fiber das Chlorophyll,” Berlin,

1884, In this publication, the entire literature relating to chlorophyll
‘to the year 1883 is ¢

ritically sifted aud reviewed. At the close of the
work, a catalogue of the literature is given, comprising nearly 600 in-
vestigations,

° Zav Sos yellow, and pvAAor leaf.

CHLOROPHYLL. 101

ow. The emerald-green color of leaves is thus a mixed color,’
and the spectruin of the leaf a mixed spectrum. While chloro-
phyll only presents bands in the less refractive half of the spec-
trum (red-green), and shows a continual absorption of the vio-
let : in the case of xanthophyll there occur no bands at all
(Fig. 32 s) between red and green, but only in the blue. Pure
chlorophyll can be prepared, as one of us (‘T.) has shown, by the
reduction of chlorophyllan, a crystallizable body.
oBG 86D Et £ . f
m5 ik 6s io et | 5D 45 j
i) Ir

Q 1 2
i i Wit 4
i HF ie 2
u 4 3
> vy
t 2
| | Bie j | ] |

Fig. 32,—1. Spectrum of 2 leaves.
2. Spectrum of 5 leaves.

3. Spectrum of a dilute t alcoholic solution of pure chlorophyll.
4. Spectrum of a concentrated

5. Spectrum of an alcoholic solution of xanthophyll.
In the leaf spectrum, band 2 of the xanthophyll is mostly covered by the projecting ter>
minal absorption of the pure chlorophyll; at least it is always rendered unclear
(Tschirch).

The chlorophyll granules of the higher plants always appear
as roundish, disk-like bodies (Figs. 33, 109, 129, 161), which,

'By means of benzol, as has been shown by G. Kraus (‘‘ Zur Kenntniss
der Chlorophyll-Farbstoffe,”’ Stuttgart, 1872), an alcoholic tincture of
chlorophyll from leaves may be split into two layers, a yellow lower
layer containing xanthophyll, and a green upper layer which contains
the chlorophyll. The separation is, however, not quantitatively exact.
(Tschirch, ‘‘ Untersuchungen iiber das Chlorophyll,” Berlin. Parey,
1884.)

102 PLANT ANATOMY.

when they lie close beside each other, become flattened polyhe-
drically, without, however, coming in contact, for the reason
that they are provided with a thin membrane of protoplasm,
They are the organs in which the most important process of
plant life is effectuated, viz., the assimilation of carbonic acid
under the influence of light, with the formation of organic or
carbon compounds. Only. organs containing chlorophyll are
capable of effecting this change. Indeed, we also find in the
chlorophyll granules an abundant accumulation of assimilation
products, and especially starch (Fig. 33, 0, c,d). Ifa leaf of
the peppermint, after the coloring matter has been extracted by

Fie. 33.—a, Chlorophyll granule, the sponge-like structure indicated by punctations ;
5, c, d, inclosures of starch in the chlorophyll granule ; e, a cell with chlorophyll gran-
ules located along the wall (Tschirch).

alcohol, is placed in iodine-water (see Micro-chemical Reagents),
it assumes at one a bluish-black color ; every chlorophyll gran-
ule contains some starch granules of a black color (see the
iodine-starch reaction). oe P
The chlorophyll granules always lie imbedded in the proto-
plasm-sac, on the inner wall of the cell, and shrink by the
_ contraction of the sac, by the addition of reagents, or by the
_ death of the cell. Since the fundamental mass of the chlorophyll
granules is very soft, many of these flow together by this process
____ to form larger masses. Thus in drugs (green leaves and stems)

CHLOROPHYLL. 103

the chlorophyll granules are seldom found unchanged, mostly
forming within the cells shapeless masses, in which the granular
structure can be recognized only with difficulty.

A similar condition also exists with regard to the maintenance
of the coloring matter, the chlorophyll.’ If, namely, leaves are
quickly dried, the plant. acids act but slightly upon the chloro-
phyll, only a little brownish-yellow chlorophyllan- (an oxi-
dation product of chlorophyll) is formed, and the leaves remain
handsomely green. If, however, they are dried without care
and slowly, brownish-yellow leaves are obtained, in consequence
of the abundant formation of chlorophyllan.* Some leaves, how-
ever, become brown even with the most careful drying (Vico-
tiana, Juglans).

Since the formation of chlorophyll is dependent upon light,
_ it is found only in those parts of plants which grow above ground
and are exposed to the light.* Leaves developed in the dark are
yellow (<‘ etiolement” etiolation ; the coloring matter is called
etiolin’). All leaves and green shrubs contain chlorophyll,
although it is sometimes conceaied by red coloring matters dis-
solved in the cell-sap (Dracena leaves). We meet with it also
in seed-vessels (Juglans), and in barks, especially in the thinner
ones (Rhamnus, Salix, etc.). Since, however, it occurs only in
cells which still possess the functions of life, it is wanting in
such barks as consist entirely of permanent tissue, or in which
the peripheral layer is wanting (Cinchona, Cinnamon).

The chlorophyll coloring matter, being a harmless green color,
is of practical importance.

'The name enssisiy aber must remain confined to the coloring sub-
stance.

* These circumstances have been thoroughly considered by Tschirch:
**Einige practische Ergebnisse meiner Untersuchungen tiber das Chloro-
phyll,” in Arch, der Pharm., 1884.

3 Nevertheless, the half-underground leaf-bases of Rhizoma /filicis are
also green. Exceptions are presented also by many green embryos en-
closed by untransparent fruit-casings, and by the small embryos of the —
Coniferze, developed in the dark.

‘From the French word étioler, to eticlate or become blanched,
which is derived from the latin stipula, halm.

104 PLANT ANATOMY.

The chlorophyll of leaves (crude chlorophyll) is insoluble in
water, but soluble in alcohol, ether, carbon bisulphide, acetone,
benzol, volatile and fatty oils (Olewm Hyoscyami of the Phar-
macopoa Germanica is colored by chlorophyll), chloroform, and
dilute solutions of caustic potassa (in the latter, with chemical
change), forming emerald-green solutions, which are dichroic
(green-red), and also show a magnificent fluorescence.

A conyenient method for distinguishing chlorophyll from
other green coloring matters, is as follows: The alcoholic solu-
tion of the coloring matter is shaken with concentrated hydro-
chloric acid and ether, the acid solution then becomes blue, the
ethereal yellow. Noother green coloring matter shows precisely
the same deportment. Pure chlorophyll dissolves with a blue
color in hydrochloric acid, and is soluble in the same solvents
as crude chlorophyll (see above). Pure chlorophyll appears to
stand chemically in close relation to the lecithines, or to be itself
a lecithine.

The colored erystalloids of many flowers and fruits (Capsicum
annuum, Rosa, Crocus, Carthamus, Tropeolum, Chrysanthe-
mum) should also be considered here. The development of
these proceeds mostly in such a manner that the chlorophyll
bodies—in the beginning the flowers and carpels of the fruit.
are mostly green—by a disturbance of their form and loss of
their original color, pass into the crystalloid coloring matters.
The yellow coloring matters (anthoxanthin ') especially occur
often in the form of handsomely developed crystals (Fig. 34),
as in the Carrot, and they probably always possess, besides the
coloring matter, a plasmatic basis.” Occasionally these coloring
matters also appear in the form of granules.

The red and violet coloring matters (anthocyan *) are, as a

‘From &y Sos flower, and Zav 34s yellow.

* They are capable of swelling. These crystalloids have recently been
repeatedly examined, thus by Hildebrand, Pringsheim’s “ Jahrbiicher,””
1861.—Nageli, ‘‘Sitzungsberichte d. Minch. Akad.,” 1862.—Weiss,

** Sitzungsber. d. Wiener Akad.,” 1866.—Schimper, Botan. Zeit., 1883.—
A. Meyer, Ibid., 1883.

*From a&v os flower, and xvayeos blue.

FAT. 105

rule, dissolved in the cell-sap (red potatoes, red foliage leaves
and petals).

In the fundamental protoplasmic mass, there is very frequently
found liquid or solid fat, for instance, in the embryo of the Gra-
mine, in the endosperm of Ricinus, and in the cotyledons of the
Crucifere. The fat appears to be combined with the proto-
plasm in the finest state of division. Microscopical sections of
seeds which are very rich in fatty oil (Ricinus, Tiglium, Amyg-
dalus, Corylus) show, when observed under water, a number of
small oil-drops, which are not visible when alcohol or glycerin,
instead of water, is used as the mounting medium. It is only
after gradually diluting the alcohol or glycerin under the cover-

IR: (30? LG
sin ve.

(LAYS

Fig. Gon! coloring matters (anthoxanthin bodies) from flowers and fruits
Tschirch).

glass with water, that the oil-drops are brought to view. From
this it may be concluded that the fatty oil is contained in the
dry seed in combination with another substance, which pre-
vents the oil from flowing together in drops. This evidently
very loose compound (perhaps containing albumen) is destroyed
by water, and the oil then unites in the form of drops.

This result may, however, be so interpreted that the fatty oil
occurs in the cells very intimately mixed with the protoplasm, —
that the albuminous body mixed therewith becomes dissolved
by the water, and that thus the oil is caused to form larger
drops. However this may be, the fatty oil is evidently very
effectually protected by this highly remarkable manner of

106 PLANT. ANATOMY.

its storage ; it is, indeed, sufficiently well known that the oil
becomes quickly rancid when the seeds are comminuted or even
moistened. —

Many cells of other tissues contain, moreover, free fat in a
liquid or solid form. In the former case, the drops of oil admit
of especially easy recognition on account of their remarkable
refraction of light, e. g., in Secale cornutum, and in Senega
root. ‘I'he fats deposited in a solid form are crystalline, which
may be seen with special clearness, among other examples, in
Cacao, Cocculus Indicus, and in the Nutmeg.’ The fat con-
tained in the kernels and shells of the Cocculus fruits consists
almost entirely of free stearic acid.”

In Stillingia sebifera (Nat. Ord. Euphorbiacexe), there is
found upon the surface of the black seeds a coating of fat, and
in Peckia (Cybianthus) butyrosa (Nat. Ord. Myrsineacesx), each
_of the four nuts has a pericarp several millimetres in thickness,
the inner portion of which forms a yellow, leafy substance.

Fats are found, not only in the seeds, but occasionally also in
the fleshy portion (sarcocarp) of fruits {a great deal in that of
the Oil-palm, Eleis guineensis, the Japanese wax-trees Rhus
succedanea and Rhus vernicifera, the olive Olea europea), im
pollen, spore-cells (Lycopodium, Pollen Pini), in some roots
(Cyperus esculentus), and in the passive state of fungi (Secale
cornutum).

As has already been mentioned, fats and oils are found in
small amounts in almost all tissues which exercise the functions
of life; they occur regularly in seeds. This is readily seen
when a section is treated with concentrated sulphuric acid ;
the protoplasm and membrane become immediately destroyed,
and the small drops of oil, which are otherwise scarcely visible,
flow together to form larger drops, which are not attacked by
the sulphuric acid.

This is, in general, the best method for the detection of small
amounts of fatty oil in microscopical preparations. In this
__ ‘Compare also Miller, ‘‘ Ueber Muscatniisse.” Pharm. Centralhalle,

1880, No. 51-53. :
_ *Schmidt and Rémer, Archiv der Pharm., 221 (1883), 34.

FAT. 107

way, one may readily succeed in rendering visible, for instance,
the fatty oil of Lycopodium, of which the latter contains near ly
50 per cent, but which does not admit of recognition by simple
microscopical observation ; it is only necessary to crush the
grains, and then to add the sulphuric acid or concentrated solu-
tion of calcium chloride.

In seeds, the fats play the part of reserve substances, and in
living tissues, particularly those containing chlorophyll, they
are an assimilation product, which evidently finds at once fur-
ther application in building up the tissues.

The fats are soluble in boiling alcohol, in ether, carbon bisul-
phide, benzol, paraffin and volatile oils, and are colored brown-
ish-black by osmic acid.

With those seeds which contain fatty oil most abundantly,
this may exceed half the weight of the kernels (after the removal
of the seed-shells). Thus in Amygdalus, Cacao, Papaver,
Ricinus, Sesamum, and Croton Tiglium ; in the latter, the oil
amounts to nearly 60 per cent. For the most part, however,
the amount of fat of other seeds which concern us here is
small; linseed and black mustard afford about 33 per cent of
oil.

‘The fat of the olive, the yellow palm-oil, as also the so-called
Japan-waz, are contained in the fleshy portion (sarcocarp) of |
the fruits of the respective plants from which they are derived.
‘The remaining solid and liquid fats of the vegetable kingdom,
which are brought in large amounts into the markets of the
world, are furnished by seeds.

The fats are esters (compound ethers) of propenyl or glycerin.
The acids combined with this radical belong mostly to the
series of the ordinary fatty acids, although a portion of very
many fatty oils and even of. solid fats consists of olein, 7. e., of
the propenyl ester of oleic-or elaic acid, which belongs to the
acrylic acid series. Nowhere has a propenyl ester been proved
to exist singly in nature ; every fat is a mixture of several such
esters. When a fat is dacounposed (saponified) by means of a

caustic alkali, the base is therefore —— found to becombined — 2
with more than one acid. 7 as so ag

108 PLANT ANATOMY.

For us the most remarkable and important constituent of the
cell-contents is the starch (amylum ').

The latter occurs abundantly, in the form of characteristic
granules, in seeds and other receptacles of reserve substances
(rhizomes, tubers). The seeds which are provided with starch
(reserve-starch) are, however, very much less numerous than
those which contain none. It appears also in the conducting
tissues (transitory starch), and in the interior of the chlorophyll
granules (assimilation-starch, autochthonous starch), but then
mostly in very small granules. For our purpose, the starch
granules of the reserve-receptacles are especially important.

Between the starch and other constituents of the cells there
exist manifold, but as yet only slightly explained relations.
Thus in the case of Radix Belladonne, Budde’ has found cer-
tain relations to exist between the amount of starch contained
in the root and the amount of alkaloid. ‘The amount of atro-
pine is most considerable in roots which are very rich in stareh,
and least in those which are free from starch (compare also
page 13).

Starch is organized and appears in the form of more or less
distinctly stratified granules? (Figs, 35, 36, 37, 39, 42, 43, 44,
45, 46).

Some drugs are exposed, in their fresh condition, to a higher
temperature in order to dry them more quickly. If these parts
of plants are juicy, the amylum thereby suffers that change
which is known as the formation of paste. The granules swell
to a high degree and flow together, to form structureless masses
or balls of paste. Thus in the case of Curcuma,* Jalap, Salep,
some varieties of Sarsapariila, and the East Indian Aconite
tubers. Sago is nothing more than swollen and dried starch.

The layers (which are especially handsome in the granules
from the potato and leguminous seeds) are arranged around a

'From a (a privative), and “vAy mill—flour prepared without a mill.

* Archiv der Pharm., 220 (1882), 414.

*Nageli, ‘Die Starkekérner, Pflanzenphysiologische Untersuchun-
gen,” 1858. The most comprehensive work relating to starch,

* Berg, ‘‘Anatomischer Atlas,” Taf. xix., Fig. 48,

STARCH. 109

common central point of the granules, which, in consequence of
unequal growth, are mostly not uniformly round; in granules
of very eccentric construction, however, the layers are in the
form of immeasurably thin shells on the side having the slight-
est growth. The layers (Fig. 35) originate through an abrupt
variation in the amount of water of the separate zones. An
outermost layer containing but very little water is followed by
one with an abundance of water, then again by one poor in
water, e¢ cetera. The centre of the granule, the nucleus, is very
rich in water,

Fic. 35.—Starch granules with very distinct layers and hilum, from the potato, very
highly magnified.

If the starch granule advances no farther in growth, a cavity
(hilum) generally remains in place of the nucleus. This space is
often confined within a very small compass, and therefore appears
asa small, dark point (nucleus-point) i in the starch of the potato
and of the rhizomes of some Zingiberacee (Figs. 36 and 46). In
the starch granules of Zuber Colchici, Maranta (Fig. 45),
Maize (Figs. 48 and 49), Radix Calumbe and others, the some-
what larger hilum often assumes the form of a star or a cross
(Fig. 37), and in many seeds from the family of Leguminosex, as
in Semen Calabar and the Bean, the hilum is proportion-

110 PLANT ANATOMY.

ately very wide, and extended in the direction of the axis of the
frequently elliptical granules (Fig. 44).

The nucleus, as a rule, is located eccentrically, although it is
central in the large granules of the cereals (Fig. 47) and in the
small, round granules of very many plants." Occasionally sev-
eral nuclei are found in one granule.

The layers disappear in consequence of the abstraction of water,
when the granules are observed under liquids free from water,
such as benzol, paraffin, volatile oils, fatty oils, glycerin, or when
they are warmed. Glycerin loses this property to a degree pro-
portionate to the amount of water it contains. On the other
hand, the distinction of the layers is also obliterated through
their intumescence (by the addition of water), even by water
at 60 to 70° C. ora still higher temperature, but even in the

ae
0074

Fie. 36.—Starch granules from the rhizome of ginger (Hager).
Fic. 37.—Starch granules with a stay-shaped hilum, from Tuber Colchici.

cold by means of saturated solutions of. many bodies which are
very readily soluble in water, such as caustic potassa or soda,
potassium iodide, calcium chloride, sodium nitrate or acetate,
and chloral hydrate. These substances increase the capacity of
absorption of water by the starch to an enormous degree, far
beyond the distinction of the separate layers just explained, so
that these swell up to a uniform mucilage.

If starch granules are pressed under the cover-glass, fissures
and clefts are formed, which, proceeding from the cleft of the
nucleus or from the periphery, form cracks having a course
mostly at right angles with the layers.

According to the very thoroughly founded and developed

_ 1 The eccentricity in the case of Cyperus esculentus amounts to i,
in Canna lanuginosa to #5.

STARCH, EEE

views of Carl Niigeli,’ starch grows in such a manner that the
formative material inserts itself between the layers of the
granule, and is by no means added externally through “ apposi-
tion.” The objections raised in opposition to Nigeli’s view of
“intussusception,” * especially by Schimper? and by Arthur
Meyer,’ are deduced from the supposition that starch possesses a.
crystal-like character. Its crystalloids, as in other carbo-hydrates
(see text to Fig. 54 a and 4), are united in the form of spheres,
sphero-crystals, but are highly characterized by the capability of
swelling. Their stratification is the result of alternate solution
and renewed deposition of solid substance. That the granules
are less dense toward the interior is shown by the penetration
of the solvent.°

Occasionally two nuclei surrounded by a separate series of

Fig. 38. Fie. 39.

Fie, 38.—Amylum, compound granules with a common integument (Dippel).
Fie. 39.—Compound starch granules from Radix Sarsaparille.

layers (Fig. 38) are formed in a single starch granule ; if these
nuclei continue to separate from each other, a high tension is
produced in the layers common to both, which leads to the dis-
solution of the double granule into two separate ones (fractured
granules). If, instead of two, a still larger number of granules
appears, compound granules are produced, which may consist of

1 Die Stirkekérner,” Zurich, 1858. Large octavo, 624 pages and 10
plates (mentioned also on page 108, foot-note 3).—‘ Sitzungsberichte d. _
Minch. Akad.,” 1863 and 1881; Bot. Zeitung, 1881, 633; also Nageli and —
Schwendener, ‘‘ Das Microscop,” 1877, p. 423.

* Intus, in or within ; suscipere, to take up.

* Botanische Zeitung, 1881, 185.

4 Tbid., 1881, p. 841, and 1884, p. 508. Pons tee ae

*Compare further the respective references thereto in Just’s Bot _
Jahresbericht for 1881, I., 398-400, pk te

112 PLANT ANATOMY.

very numerous individual granules (Avena, Fig. 40, Spinacia,
Sarsaparilla, Fig. 39).

The false compound granules are formed by several separate
granules becoming firmly agglutinated with each other through
mutual pressure (frequently occurring in the starch in chloro-
phyll, Fig. 33 d).

The shape of starch granules is very varied.' The fun-
damental form is the sphere. All of the smaller, isolated starch
granules possess this form, such as the small granules of
the wheat (Fig. 47), of the potato (Fig. 43), and the so-called
transitory starch. When the granules fill the cell and are
densely crowded, they are always flattened through mutual
pressure, and are then mostly polyhedric (dodecahedron and

i Ihe

Fie. 40. Fia. 41.

Fie. 40.—A compound starch granule of the Oat, resolved into its separate granules,

Fic. 41.—Starch in bone-shaped and club-shaped granules from the milky juice of
Euphorbia antiquorum ; more difficult to obtain from the commercial ‘ Euphorbium ™
(gum-resin of Euphorbia resinifera).

allied forms) (Maize, Fig. 49, Rice, Fig. 50). In the chloro-
phyll granule the amylum is mostly spindle-shaped (Fig. 33 c,
ad). Club-, staff-, or bone-shaped structures are found in the
milky juice of many Huphorbie (Fig. 41), club-shaped in the
rhizome of Galanga,? and branched in the root-stock of Nelum-

‘Compare the illustrations and descriptions of forms of starch by
Vogl, ‘‘ Die gegenwartig am haiufigsten vorkommenden Verunreinigun-
gen, etc., des Mehles,” Vienna, 1880.—R. von Wagner, ‘‘ Die Starkefabri-
kation.” Braunschweig, 1876.—KGnig, ‘* Die menschlichen Nahrungs-
und Genussmittel,” Berlin, 1883.—F. yon H6hnel, “ Die Starke und die
MahlIproducte, etc.,” Cassel and Berlin, 1882.

* Berg, “‘ Anatomischer Atlas,” Plate xix., 46.

STARCH. 1s

dium speciosum, Willd. Sago-starch’ is provided with swollen
protuberances (Fig. 42). Independent of exceptions of this
character, the spherical and ovate, or often flattened forms? pre-
dominate.

Although not susceptible of strictly mathematical definition,
the size and shape of the starch granules are nevertheless char-
acteristic for individual plants. A knowledge of these peculiar-
ities is therefore indispensable in the examination of flour and
varieties of starch. Besides the smaller structures, which usu-
ally occur, each granule possesses a predominating typical form.
It is only when the latter has been confirmed, in its definite
shape and size,* by a large number of granules, that one can
assume that a certain variety of starch is present.

In the examination of starches, this is the only means for

Fic. 42.

Fic. 42.—Starch granules of Sago (Hager).

Fie. 43.—Potato starch (Koenig). a, the nucleus. Compare also Fig, 108.
their identification, but it is otherwise with varieties of flour.
The latter consist of ground fruits and seeds, and thus con-
tain, besides the starch granules, remnants of cells of the inner
tissue as well as of the integuments, and often also remnants of
hairs (Zriticwm). In the case of flour, these remnants may
therefore be very well used as a guide in their examination.*

* Compare also Wiesner, ‘‘ Die Rohstoffe des Pflanzenreiches,” Leipzig,
1873,

* For example in Zingiber, Berg’s Atlas, Plate xx., 49.

* Measurements of the size of the granules (by the aid of the ocular
micrometer) must always be undertaken. The linear diameter is deter-
mined. The unit ‘of measure is the micro-millimeter ( or mic.) =
reop Mm. = 0.000001 m. a

* Compare ied nies, ‘* Anleitung zur Erkennung organischer und

114 PLANT ANATOMY.

The most important forms of starch are the following :*

1. Porato Starcu (Solanum tuberosum, Figs. 43 and 108).
Type: large, eccentric, very distinctly stratified, quite irregu-
lar granules, which are rounded by three or four corners; they
are often rhombic and wedge-shaped, but never flattened. The
nucleus is at the smaller end.

Fie. 44.—Bean starch (Tschirch).

Secondary form: small, roundish, and medium sized, half or
completely compound granules.

Fia. 45.—Maranta starch (Tschirch),

2. Bean Srarcu (Physostigma, Vicia and species of Phase-
olus, Fig. 44). Type: bean-shaped to kidney-shaped, dis-
tinctly stratified granules, which are always simple. The
eo Beimengungen im Roggen- und Weizenmehl,” Leipzig,

.

tie: good key for use in determining the varieties of starch has been
given by Vogl: “ Nahrungs- und Genussmittel aus dem Pflanzenreich,”

Vienna, 1872; compare also Konig, “ Nahrungsmittel,” II., p. 405,
_ Wagner and others,

.

STARCH. 115

nucleus is not visible, since the granules are nearly always tra-
versed by a broad, radiately branched, longitudinal cleft.

Secondary form : small, roundish granules.’

3. Maranta Starcu, Arrowroot starch (Maranta arundina-—
cea, Fig. 45). Almost entirely typical forms: more or less flat-
tened, nearly quadrangular, rhombohedral, triangular, club- or
pear-shaped granules, with a distinct nucleus located at the
broad end, and often traversed by an occasionally three-rayed,
lateral cleft. Stratification distinct.

4. East Inptan ARrowroort (Curcuma leucorrhiza and C.
angustifolia, Fig. 46). Type: distinctly stratified, flat, tabular
or oval and lengthened, triangular granules, drawn out to a point.
on one side, which contains the nucleus, without a cleft.

Secondary form: small, triangular granules.

Fie. 46.—Starch of Curcuma leucorrhiza (Koenig).

5. Wueat Srarcu (Triticum vulgare),’ Fig. 47. Type:

(az) LARGE GRANULES, flatly lenticular, almost exactly circu-
lar, without a cleft or a distinct nucleus. They are four times.
larger than the following :

(6) SMALL GRANULES, roundish or polyhedral, often con-
nected in pairs.

1 Specific characters for the discrimination of bean- and pea-starch
have been given by Tschirch, ‘‘ Staérkemehlanalysen,” in Archiv der
Pharm., 222 (1884), p. 921.

®The rye and barley have similar granules. For further details.
regarding them, compare the previously mentioned monographs om
starch,

116 PLANT ANATOMY.

Secondary form: slightly roundish granules, having a form
intermediate between the two preceding.

6. Maize Starcu (Zea Mays, Figs. 48 and 49) presents only
typical forms.

(a) Horn endosperm (Fig. 49): sharply angular granules,

Fie. 47.—Wheat starch; v, face view ; s, marginal view ; t, parts of adouble granule ;
r, & granule of rye starch with a three-rayed cleft (Tschirch).

without distinct stratification, and mostly provided with a

Fie, 48, Fie. 49.

Fie. 48,—Maize starch granules from the farinaceous endosperm (Tschirch),
Fie, 49.—Maize starch from the horn endosperm (Tschirch),

cleft ; they are in close contact with each other, and completely

fill the cell. Inthe flour also, several granules are often still
coherent.

STARCH. Et}

(4) Farinaceous endosperm (Fig. 48): granules more round-
ish, occasionally without a cleft. They do not completely fill
the cell, and therefore are not sharply flattened polyhedrically
against each other.

7. Rice Starcu (Oryza sativa, Fig. 50), only typical forms ;
very sharply angular, almost crystal-like, fractured granules,
occasionally several still connected, without a nucleus cleft.

Fie. 50.—Rice starch (Tschirch).

8. Oat StarcH (Avena sativa, Fig. 51). Type: large, oval
aggregations, as much as 73°; mm. (50 yu) in size, composed of
from two to three hundred granules and their components
(Fig. 51 6). The latter are polyhedric and sharply angular,
without a distinct nucleus. ‘

Fig. 51.—Oat starch. a, Secondary form—filling granules ; b, component granules
of the aggregation (Tschirch). :

Secondary form: small, roundish, spindle-shaped,’ similar to
the fractured granules, the so-called ‘‘ filling starch.”
Like the cell-membrane, the starch granules, in consequence

1 Moeller (‘Die Mikroskopie der Cerealien,” in- the Pharm. Central-
halle, 1884, No, 44-48) declares these to be the characteristic forms.

118 PLANT ANATOMY.

of their stratified structure, are also doubly refractive. In pola-
rized light each granule displays a black cross, the arms of
‘ which intersect at the hilum (Fig. 52).

When the structure of the granule is destroyed, either through
tumefaction or by torrefaction, it immediately loses its optical
properties, though agents which produce swelling, but which
have neither an acid nor an alkaline reaction, produce no
change, at least ina chemical sense, in the substance of the
starch. The optical properties therefore appear to be dependent
upon the manner of construction of the granule. Nevertheless,
Nigeli entertains the view that the micelle’ of starch (a term
which he applies to the complex of atoms, surrounded by a film
of water, which through intussusception become separated from
each other), like those of the denser cell-membranes, are origi-
_ nally crystalline, and show the deportment of optically uniaxial |
crystals, but only so long as they have not become disintegrated,
which Nigeli assumes to take place upon swelling.

Starch, as the most important reserve nutritive substance, is
contained in an extraordinarily large number of reserve recepta-
cles, thus in seeds (endosperm of the cereals, the cotyledons of
many Leguminose), Semen Cacao, Sem. Myristice, Sem. Para-
disi, Sem. Piperis (v. Piper album), Sem. Quercus, while Sem.
Cydonia, Sem. Lini, Sem. Sinapis albe and probably others
contain starch, at least before ripening. Furthermore, in rhi-
zomes (Maranta, the Zingiberacexw, Aspidium Filiz-mas, Asa-
rum, Calamus), in roots (Althea, Sarsaparilla, Krameria, Ipe-
cacuanha, Rhubarb, Belladonna), and in tubers (Potato, Salep,
Jalap, Colchicum).

A remarkable exception is presented by the roots and rhi-
zomes of the Composite, which contain no starch, or only tran-
sitorily, and then but extremely small amounts of it. It is fur-
thermore wanting in Radix Gentiane, Rubie, Saponaria,
Senege, and in the rhizome of Triticum repens, at least in the
Stages of development which here come under consideration.
In all these organs its place is supplied by other substances.

Unjustifiable diminutive of mica, a small crumb.

RCH.

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rhizomes).

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120 PLANT ANATOMY.

starch granules, since the above-named transformation products
possess the inclination, under suitable conditions, to again
become deposited as solid starch (transitory starch). These
small starch-granules, which are in process of migration, are
thus found in the so-called starch-sheath, and in the sieve-tubes
and medullary rays. They are also contained in some fruits
before ripening (Olives, Fructus Conti, Fructus Juniperi,
likewise the Fig).

How the formation of starch is effected in the chlorophyll
granules (assimilating-starch, p. 108) is, meanwhile, still an
enigma. Only so much is certain, that for its production light
is required,’ while potassium also appears to be indispensable
for it.*

The manner of circulation of starch may be elucidated by a
single example. The species of Orchis which afford salep pos-
sess, after the close of the period of vegetation, a tuber which is
filled with starch and mucilage. The tuber is quiescent during
the winter, and in spring develops a stem bearing the leaves and
flowers. During the entire first period of development, the
tuber provides the young shoot with nutritive material ; the
starch migrates from the tuber upward into the shaft. In the
course of further development, the leaves unfold, and now under-
take on their own part the new formation of starch. But even

now the plant provides for a future year. Beside the old, and
now entirely exhausted tuber, a new rudimentary one is formed,
into which the starch formed in the leaves migrates downward,
in order to furnish the constructive material for the young plant
during the next year.

If the starch is dissolved in a reserve-receptacle, the gran-
ules do not disappear at once, but solution gradually takes
place, whereby a peculiar corrosion often occurs. Such corroded

‘Compare Sachs, ‘“ Experimentalphysiologie der Pflanzen,” 1865;
furthermore, Botan. Zeit., 1862 and 1864, ‘‘ Flora,” 1863, and researches
of the Botanical Institute of Wirzburg, 1884.—Godlewski, Krakauer
Akadem., 1875.—Béhm, Botan, Zeit., 1876, and others.

? Proved by Nobbe in 1871.

STARCH. 121

starch-granules are particularly well observable in the first
stages of germination (Fig. 53).

We are acquainted with starch in the plant only in the solid
form, although compelled to assume that it is formed, or ina
manner crystallizes out, from a liquid.

Starch forms a glistening powder, the specific gravity of which
varies according to its origin, but does not deviate much from
1.5. In the air-dry condition, it incloses from 13 to 17 per cent
of water, after the removal of which its density increases, accord-
ing to its derivation, to from 1.56 to 1.63. While air-dry starch
floats upon chloroform, it sinks therein after having been
deprived of its water by heating to 100° C. Dried starch quickly
absorbs again from the air the eliminated water.

The small amount of incombustible substances which it con-

sae
os Gy 4

Fie. 53.—Corroded starch-granules from the endosperm of a young maize plant 8
centimeters (about 3 inches) high, in process of solution. a, a granule still intact
(Tschirch).

tains, about 0.5 per cent, can 1 probably be explained only by
supposing them to be deposited mechanically.

The composition of anhydrous starch corresponds to the
formula C,,H,,0,,, although Musculus (1861 and 1870) and W.
Nigeli have shown that the formula O,,H,,0;, or O,,H,,0,, cor-
responds still better to the facts.*

Leuchs (in 1831) found that starch granules are attacked by
saliva. ©. Niigeli, who pursued the subject further, came to
the conclusion that the granule is built up from cellulose and a
peculiar starch substance, granulose. According to his view, the

1 For information relating to the elementary composition of starch, —
we are indebted to W. Nageli, Sachsse, Pfeiffer, Tollens and Salomon.
See also Husemann-Hilger, ‘‘ Die Pflanzenstoffe.”

122 PLANT ANATOMY.

saliva acts upon the latter substance, dissolving it, and leaving
a frame-work or skeleton of cellulose behind.

On the other hand it is to be remembered, as one of us has
shown,’ that the “ granulose” has lost all the characters of
starch. Furthermore, the acceptance of cellulose in the residue
is based upon its solubility in ammoniacal solution of oxide of
copper, the loss of its property of swelling in hot water, and the
non-appearance of any coloration by treatment with iodine. But,
on the one hand, amylum is itself soluble to a slight degree in
ammoniacal oxide of copper, and, on the other hand, ‘‘ granu-
lose” is just as little colored by iodine as the here accepted
cellulose, while the swelling property of starch may also be
destroyed by boiling with glycerin and water. There are thus
not sufficient reasons presented for concurring in Nigeli’s pro-
position.

Starch containing water, but not that which has been deprived
of the latter, possesses a highly remarkable attractive power for
iodine. It is capable of so combining with it, that the granule,
the mucilage, or the solution of starch thereby assume colora-
tions which correspond to those peculiar to iodine itself in its
different conditions of aggregation and in its solutions. The
blue, violet, or reddish color which amylum presents when it is
brought in contact with iodine was first observed by Colin and
Gaultier de Clanbry* in March, 1814; the other shadings in
violet, red, reddish-yellow, yellow, and brown were studied in
1863, and later by C. Niigeli with great thoroughness. These
shadings of color are limited by the varying reciprocal relations
in the amounts of iodine and starch, as also by the presence of
hydriodic acid and other substances.

Slight amounts of very small starch granules as in chlorophyll
(page 102) may be made more distinctly visible by causing them
to swell by means of caustic alkali, subsequently washing the
fe cea “ Starke und Cellulose,” in Archiv der Pharm, 196 (1871),

** Annales de Chimie ” 90, p. 93. —Stromeyer, at Géttingen, in Decem-

ber, 1814, pointed out the delicacy of this reaction in Gilbert’s‘* Annalen
der Physik,” 49, p. 147,

STARCH. 123

‘section with acetic acid and then with water, and finally
adding iodine solution. '

Though the force with which starch appropriates iodine is
‘quite considerable, it can, nevertheless, not be proved that the
product is a chemical compound. Even dialysis, as also gen-
tle warming, and even simple exposure to the air, is capable of
eliminating the iodine from the compound.

Only cellulose, under certain conditions, shares this behavior
of starch to iodine. Beside lichenin (see Index), there is to be
mentioned here also the amyloid of Schleiden, a form of cellu-
lose which is capable of swelling, is colored blue by iodine, and
occurs in the cotyledons of many of the Leguminose, for
instance, in those of Zamarindus. Certain membranes of the
hyphe of lichens also assume with iodine solution a blue color.

The amount of amylum, even in such plants and parts of
plants as are abundantly provided therewith, must necessarily
be subject to great fluctuations when the before-mentioned
function of starch as a reserve substance is taken into considera-
tion.

Potatoes and the rhizomes of the Maranta (Arrowroot)
afford, for instance, from 9 to 26 per cent of amylum with ref-
erence to air-dry substance, and Sarsaparilla is likewise a good
example of the fact that the percentage of starch is very variable.
The statements relating to the quantity of starch present in
‘drugs can therefore be of value only under definite conditions.

The size of starch granules is very variable, although, for the
Same kind, remaining within narrow boundaries. The largest
are found mostly in the underground receptacles of reserve sub-
stances (Solanum tuberosum as much as 90 yu,’ Canna lanugi-
nosa as much as 170 ), the smallest in the seeds of some
Species of Acacia (about 1 y).

An approximate representation of the relative dimensions is
given by the following figures: |

14 or mic. = micromillimeter = r/ya mm. = 0.000001 m. (compare
Page 113),

124 PLANT ANATOMY.

According to According to

AVERAGE MEASUREMENTS. Wiesner, Héhnel, Wagner, Tschirch *

Length in yz, Length in jz,
Potato, 60 56.0
Wheat (large granules), 26.9-28 33.0
Wheat (small granules), 6.8 6.0
East Indian Arrowroot, 50-60 —_—
Maranta, 27-54 32-46
La, ' 32-79 99-39
Bean, :
Maize, 15-20 13-19
Rice (divided granules), 5 5-6
Oat (divided granules), 4.4 7-8°
Rad. Calumba, As much as 90 yu :
Rhiz. Zedoaria, she 70 pet ate hapee Se
Tuber Jalape, re Ss .60 j BORE

Another constituent, related in its es to starch, is”
inulin,’ which Valentine Rose, in 1804, first observed as a de-
posit from the decoction of the root of Inula Helenium; Thom-
son® designated it as inulin. It occurs chiefly in the roots of
plants of more than one year’s growth belonging to the family of
the Composite, and has been detected elsewhere in but few cases.°
Prantl’ has obtained, for example, quite a considerable amount
of inulin from the roots of the flowering Campanula rapuncu-

‘The average from 100 measurements. Regarding the relations of
shape as well as size, see the very detailed statements of Kdnig,
‘** Nahrungs- und Genussmittel,” IT., 403 et seq.

* I found (as did Wiesner, in opposition to Kénig), the oat granules to
be always larger than those of rice (T.). See A. Tschirch, ‘‘Starkemehl-
analysen” in Archiv der Pharm., 223 (1885), pp. 521-532.

: me oe eeeieanens des Piiaikenreiches; ” first edition (1867), pp. 237,

» U1.

‘Compare, regarding inulin, Sachs, Bot. Zeit., 1864. Holzner, ‘‘ Flora,”
1864, 1866, 1867, and the publications cited below.

“System of Chemistry,” IV. (London, 1817, fifth edition), 75; also
in earlier editions, previous to the year 1811.

°That the Australian Lerp-Manna, in opposition to former assump- .

- tions, contains no inulin, is now definitely established. Wittstein’s

Vierteljahrsschrift fiir prakt. Pharm., XVII. ee 161, and XVIII. 1.
*** Das Inulin,” Munich, 1870, 43.

+>

INULIN. 125

loides L., and Kraus’ found it also in the families of the Cam-
panulaces, Lobeliaceew, Goodeniacee, and Stylidiacew, which,
from a systematic point of view, are each and all connected
with the Composite. Inulin has, moreover, been proved by
Kraus to occur in the roots of Jonidium Ipecacuanha, of the
family of Violacew.* In the family of Composite, inulin pos-
sesses the function of amylum ;* it is distinguished, however, in
general from the latter by the following main points.‘

Fia. 54 a.—Globular aggregations of crystals (sphzero-crystals) from Radix Inule, by
keeping fresh pieces ‘of the root for a long time in glycerin. B, cells filled with
Inulin; 4, separate, strongly magnified aggregations (Sachs). Fic. 52, VI., represents
such an aggregation in polarized light (Dippel). '

1 Bot. Zeitung, 1875, 171.

? Fliickiger, ‘‘ Pharmakognosie,” 1883, 396.

* Starch has been found in but a few roots of the Composite. Vogl,
“Kommentar zur ésterreich. Pharmakopée,” 1869, p. 847 and Dippel,
‘Das Mikroskop,” II. (1869), 27. Nevertheless, according to Kraus,
the chlorophyll granules, the stomata cells, as also the starch-sheaths
and sieve-tubes of plants which form inulin contain starch throughout.

“Compare further: Dragendorff, ‘‘ Material zu einer Monographie

126 PLANT ANATOMY.

In living roots or leaves, inulin does not separate out in a
solid form ; it is only when water is abstracted from the solu-
tion, in which it is there contained, that it forms either glass--
like,.amorphous masses, or fine, soft, needle-like crystals of the.
rhombic system.’ The latter may combine to form larger,
radiated, spherical aggregates or sphwro-crystals (F ig. 54 a and
6), which are best obtained when entire Dahlia tubers are placed
in absolute alcohol or concentrated glycerin. After some days,

Fie, 54 b.—Spheero-crystals of inulin from Dahlia tubers.

in consequence of the slow abstraction of water, the inulin
crystallizes in aggregations, which cannot be obtained by simple
drying. The leaves of the Composite must be prepared for
dehydration by previously boiling them with caustic potassa.

Crystallized inulin, when observed in polarized light, is seen
to be doubly refractive (Fig. 52, p. 119, No. VI.), though less
strikingly so than amylum ; the crossed arms do not appear
des Inulins,” Petersburg, 1870. Kiliani, Liebig’s Annalen, 205 (1880)
145-190.

1 Bot. Zeitung, 1876, 368,

INULIN. | 127

very distinctly on the sphero-crystals, and the amorphous
masses are neither doubly refractive nor stratified.

With this deficiency of organic structure is connected also
a lesser capacity of combining with water. In opposition to
amylum, the composition of which corresponds to the formula
(C,H,,0,),+3H,0 (= 14.2 percent of water), air-dry inulin con-
tains only from 5 to 10 per cent of water. On the other hand, it
dissolves readily in hot water and separates therefrom unchanged
upon cooling, provided the solution had not been exposed for a
long time to a higher temperature. In the latter case, the inu-
lin very readily passes into uncrystallizable, levogyrate sugar.

Fig. 55.—Groups of fine needle-shaped crystals (rhaphides) from Radix Sarsaparille.

The solution of inulin itself likewise deviates the plane of
polarization of a ray of light to the left; solutions of starch,
which are obtained by the aid of chloral hydrate or by certain
salts (page 110), rotate tothe right, as does also the crystallizable
grape sugar obtained from starch. The aqueous solution of
inulin is never paste-like ; it is an actual solution in the ordi-
nary sense, while the paste of starch is produced only through a
swelling of the granules.

Tnulin is not colored by iodine. Indeed, we possess no heat
for it, and are only capable of recognizing it by confirming sey-
eral of its physical properties.

The amount of inulin contained in the Composite is very

128 PLANT ANATOMY,

SS

Fic. 56.—A, Transverse section; B, Longitudinal section from Bulbus Scillce, with
numerous:prisms of calcium oxalate, which are often approximately 1 mm, in length.

CALCIUM OXALATE. 129

variable, in many cases very slight, as for instance in Rhizoma
Arnice. From dried Radix Inule, on the other hand, Dragen-
dorff obtained 44 per cent of inulin; from the root of Taraza-
cum, gathered at Dorpat, Russia, in October, and dried at
100° C., 24.3 per cent, while the same root in March afforded
only 1.7 per cent of inulin.

The great periodical fluctuations, and the want of a reagent,
may explain why it has not yet been possible to detect inulin
in many roots of perennial Composite.

Although inulin never occurs in crystals in the living and
dried plant, there are other crystalline substances which occur
not infrequently in the tissue of the cell. Calcium oxalate
especially is widely distributed.

In yery many plants calcium is deposited in the cells in the
form of distinctly crystallized oxalate. ‘his salt mostly corre-
sponds to the formula CaC,0,+ H,O, and belongs to the mono-
clinic (clino-rhombic) system of crystals. Occasionally, how-
ever, forms of the quadratic or tetragonal system are also to be
seen ; this variety of oxalate contains 3H,0.

When calcium oxalate is prepared artificially, and the salt
Separates rapidly, the first-mentioned compound is obtained
either as an indistinct crystalline precipitate, or in well recog-
nizable monoclinic forms; the quadratic oxalate, on the con-
trary, crystallizes during the slow evaporation of a hydrochloric
acid solution, or also upon the admixture of very little calcium
chloride with an extremely dilute solution of oxalic acid.‘ Fre-
quently, under slightly changed conditions, a mixture of both
compounds is produced.

The two forms of calcium oxalate are of exceedingly frequent
occurrence in the vegetable kingdom. The needle-shaped crys-
tals, rhaphides* (Fig. 55), appear to belong to the monoclinic

' With regard to the more precise conditions, compare Souchay and
Lenssen, Annalen der Chemie u. Pharm., 100 (1856), 311-325.

* From fais, the needle. A. de Candolle, in 1826, introduced the
term raphides, in order to avoid the use of the word crystals, as he sup-
posed (erroneously) those deposits of oxalate of calcium not to consist
of crystals,

130 PLANT ANATOMY.

system; these occur separately or in groups, particularly in the
root formations of monocotyledons, very notably in Bulbus
Seille (Fig. 56), and in Radix Sarsaparille (Fig. 123'),but also
in stems and leaves, as, for instance, in the Aloé (Fig. 63, cr).
The undeveloped, crystalline, powdery oxalate, which is met
with, for instance, in the Cinchona barks, in Stipes Dulcamare,
and in Radix Belladonne, should probably also be considered
here. Such deposits become better recognizable when the sec-
tions, freed as much as possible from air, are observed in pola-
rized light. More distinctly and variously developed are the
Fie. 57, Fia. 58.

Fie. 57,—Fundamental form of the monoclinic crystals of calcium oxalate, with only
one molecule of water. This form, hendyohedron, resembles in appearance a rhom-

Dohedron of the hexagonal system, and is therefore often designated as ‘‘ rhombohe-
-dron-like oxalate.”

Fic. 58.—a, Hendyohedron ; b and c, crystals of the monoclinic system in Cortex
Frangul, derived by truncation from the fundamental form (from Dippel).
crystals which in form approach that of the hendyohedron
(Fig. 57), which may be regarded as the fundamental form of
the monoclinic salt. Very handsome and very regularly devel-
oped crystals of this kind occur in Radiz Calumba, Folia
Hyoscyami and Cortex Frangule (Fig. 58), and particularly
also, in a considerable variety of forms, in the non-official
bark of Liqguidambar orientalis Miller, which yields the Styrax
Uiquidus. In Cortez Aurantiorum the crystals are likewise
quite large and are inclined to be sharpened ina striking manner.

- " Compare also Schleiden, Archiv der Pharm., 1847, Plate I., Fig. 5.

CALGIUM OXALATE. 131

Forms of peculiar appearance, produced by hemitropy, and
recognizable by their inwardly inclined angles (Fig. 59), occur
in the bark of Guaiacum officinale and Quillaia Saponaria.*

Much less widely extended, at least within the sphere of drugs,
are well-devoloped forms of the quadratic system (Fig. 60), such
as are found forinstance in oak-galls (Fig. 61). Oxalate crys-
tals of this system also occur in many leaf-stalks, and are par-
ticularly handsome in the Cactex, in species of Begonia and in
Paulownia imperialis Siebold, furthermore in Urceolaria seru-
posa Ach. and other lichens.

ve

Fie, 59.—Twin crystals of calcium oxalate from Cortex Guaiaci or Cortex Quillaiw
Saponarie; a, lying on the lateral surface; c, somewhat turned; b, more strongly mag-
nified and turned to the extent of 90° (Dippel).

In Rhiz. Rhei, Rad. Saponaria, Rad. Althee, in Cortex
Granati Radicis, in figs, cloves, and in a very large number of
other parts of plants belonging to our department (Fol. Eu-
calypti, Figs. 127, 128), the oxalate crystals are most densely
crowded together in the form of clusters (Fig. 62); each of which
generally occupies a single cell. In these cases only the points of

' Further details are given by Holzner, ‘‘ Krystalle in den Pflanzen-

zellen,” Flora, 1867, 499. Sachs, ‘ Lehrbuch der Botanik” (IV.), 66.
Compare also Figs. 102 and 159,

132 PLANT ANATOMY.

the individual crystals just project, but the true erystallographi-
cal shape of the latter has not yet been determined with certi-
tude. The fact that in Cortex Cascarille, in Cortex Frangule,
in the outer surface of Fungus Laricis, in the above-mentioned
Styrax bark, and in other cases they are accompanied by dis-
tinctly recognizable monoclinic crystals of oxalate, argues possi-

Fia. 60.—Fundamental forms of calcium oxalate‘crystallizing in the quadratic system
with three molecules of water of crystallization.

bly for the assumption that these aggregates or rosettes also.
belong to this system, although in the above-mentioned leaf-

Fic. 61.—Transverse section from an ordinary (Aleppo) oak-gall; d, sclerenchyma-
tous layer in the centre ; ¢, tissue outside of and in proximity to this layer, filled with
quadratic crystals of oxalate ; e, tissue in the interior of the chamber formed by the
sclerenchyma, which contains starch and resin. *

stalks all transitional forms may also be observed, from the
quadratic octahedron to imperfectly developed rosette-shaped
aggregates of crystals. Hence it is probable that the oxalate

crystallizing in rosettes sometimes belongs to the quadratic and
sometimes to the monoclinic system.

CALCIUM OXALATE, 133

The proof that the plant formations in question are really
calcium oxalate is readily afforded. The crystals are not solu-
ble in acetic or oxalic acids, but soluble, and without efferves-
ence, in hydrochloric acid; this solution gives, upon the addition
of potassium acetate, an abundant precipitate of indistinctly
crystallized calcium oxalate. ;

After short contact with concentrated sulphuric acid, the
oxalate crystals are converted into long, lance-shaped crystals of
gypsum,

The oxalate crystals are presumably formed in the plant by
the gradual confluence of dilute solutions of oxalates with calcium
salts. In many cases this occurs with the co-operation of or-
ganized structures. The rosettes often inclose an uncrystellized
nucleus, and the needle-like tufts of oxalate are fixed, for in-

Fig, 62.—Rosettes of calcium oxalate from Rhubarb (see Figs. 142, 146) and Radix
Saponaric. ;

stance in Sarsaparilla and many other cases, in a mucilaginous
(plasmatic) integument; if the oxalate be dissolved in hydrochloric
acid (spec. gray. 1.1), the integument remains behind, and may
easily be made recognizable by staining, for example, by means
of carmine or aniline-red. This covering of protoplasm may
be detected also with greater distinctnessin Bulbus Scille. Ifa
fine section of this is moistened with alcohol, a contraction of
the mucilaginous contents of the cell ensues, in the middle of
which darker granules will appear, which are seen to be crystal-
line in polarized light. Water dissolves the mucilage and leaves
the crystals behind, which, without doubt, are to be regarded as
the first rudiments of the oxalate prisms often so handsomely
developed in the Sgwill. The latter are surrounded by a sac,
and frequently become enlarged to such a degree as to extend

134 ; PLANT ANATOMY.

through several cells, after their transverse walls are destroyed”
These crystals often attain nearly 1 mm, in length, so that they |
become visible even to the unaided eye. The latter character
applies also to the imperfectly developed, rhombohedron-like
erystals in the wood-parenchyma of Lignum Sandali rubrum,
the axes which are scarcely less than $ mm.

According to Emmerling* it would appear probable that
erystals of calcium oxalate are also formed in the plant through
the action of free oxalic acid upon calcium nitrate.

In the cases here referred to, calcium oxalate always occurs as.
one of the contents of the cell; it has been shown, however, by
Count Solms-Laubach * that these crystals may also be deposited
in the cell-wall itself, especially in the outer wall of epidermis
cells.

In regard to the amount of oxalate, the microscopical estimate
may lead toinaccurate statements. Bulbus Scille is apparently
rich therein, and nevertheless,a direct estimation of the oxalic
acid afforded but 3 per cent of oxalate; in a good rhubarb * one
of us found 7.3 per cent. The greatest abundance of oxalate in
the domain of pharmacognosy is presented perhaps by guaiac
bark, nearly 20.7 per cent. Some lichens are likewise charac-
terized by a large amount of oxalate; thus Lecanora esculenta
Eversmann, contains 22.8 per cent.

The oxalate crystals are deposits which remain withdrawn
from the sphere of vital action (secretions); in the cells which
contain them, as a rule, no further developments take place.

Other crystalline compounds of inorganic bases are of exceed-
ingly rare occurrence in plant tissues. Calcium phosphate,

? “ Berichte der Deutschen Chemisch. Gesellsch.,” 1872, p. 782.

* Botanische Zeitung, 29 (1871), 458. Plate VI.— Also in Sachs, ‘‘ Lehr-
buch der Botanik,” 1874, 68.
_ 8 One of the numerous important observations of the eminent apothe-
cary Scheele, who also discovered oxalic acid, relates to the crystals of

rhubarb, which, in 1782, he recognized as calcium oxalate. Anton van
Leeuwenhoek (1716) had, indeed, previously seen the oxalate of the

sarsaparilia root and of orris ont —Flickiger, ‘‘ Pharmakognosie,”
2d edition, 226, 315, 373.

CALCIUM OXALATE. 135

CaHPO,+2H,0, is found abundantly in a crystalline form’
in the Indian Yeak-wood (Tectona grandis L., Nat. Ord. Ver-
benacee). Calcium carbonate, which is contained in some
plasmodiums, in the cell-membranes of many marine alge, and.
in cystoliths (Ficus, Cannabis, Humulus*), does not show dis-
tinctly crystalline forms,’ or is manifestly amorphous. Crystals
of gypsum appear not to be present in plants; since they are
soluble in 400 parts of water, the conditions are probably want-
ing for their formation and maintenance.

Crystals of organic compounds, which are met with in the
tissues of drugs, are, however, no rarity. Thus, asparagin,
cubebin, hesperidin, picrotoxin, theobromine, and piperine,
which, however, may be presumed to first crystallize during»
the process of drying the respéctive drugs. Furthermore, crys-
tallized fats, probably for the most part palmitin and stearin,
which are found in many seeds, as, for instance, in the nutmeg,
in Cocculus Indicus, ete. Finally, vanillin in the parenchyma
and upon the outer surface of the Vanilla (Fig. 83). The erys-
tals which become visible in Cinchona barks, after warming their
sections in caustic alkali, first appear as a result of this treat-
ment. By very long preservation in glycerin of sections of tis-
sues rich in tannin, crystals of gallic acid also occasionally
appear, which were not originally present. After a very long
preservation of the respective sections, one may also observe the
gradual crystallization of amygdalin, filicic acid, and strychnine..

Small granules are frequently found deposited in cells, which
acquire with ferric chloride in aqueous, or often better in alco-
holie solution, a blue or greenish coloration, so that we may con-
sider them as tannin, or as tannin-like formations. On the other
hand, they also often become colored blue by iodine, as if they

' Kopp-Will’s Jahresbericht der Chemie, 1860, p. 531, and 1879, p. 937;
“Berichte der Deutschen Chemisch. Gesellschaft,” 1877, p. 2,234.—Com-
pare further Just’s Bot. Jahresbericht, 1881, I., 402, Reference No. 75.

* Fliickiger, ‘‘Pharmakognosie,” 1883, 710; Sachs, “Lehrbuch der
Bot.,” 1874, 70; Kny, ‘‘ Botan. Wandtafeln.”

*Thus also the aggregates in Castoreum; compare Fliickiger,

Grundriss der Pharmakognosie,” 1883, 237; further Just’s Bot. Jahres-
rericht, 1881, I., 402, 403. :

136 PLANT ANATOMY.

inclosed starch or had originated therefrom, as indeed both
appear in the same tissues simultaneously, or still more often
alternately. Nevertheless, tannin does not exist to any consid-
erable amount in seeds. The amount of tannin contained in
certain organs, such as barks and fruits, is subject to considera-
ble periodical fluctuations.* :
Tannin which is deposited in the purest form dissolves when
subjected to examination under water. In order to bring it to
view, the sections must therefore be observed under benzol,
volatile or fatty oils, or other liquids which do not dissolve the
tannic matter; even glycerin suffices, since it dissolves but little
tannin when concentrated. Thus in galls, shapeless masses are
found which almost completely fill the cells.?. The tannic mat-
ter also very frequently penetrates the cell-membranes, so that
the walls of entire tissues become colored after being moistened
with a solution of iron, thus, for instance, the parenchyma of
the Cinchona barks, the fibro-vascular bundles and the surround-
ings of the oil-cells in cloves, ete. Thick, hard cell-walls, which
do not become thoroughly moistened by an aqueous solution of
iron, often assume, nevertheless, the blue or green coloration
upon the simultaneous addition of alcohol. These reactions,
however, are perhaps more often produced by derivatives, decom-
position products of the tannins, or bodies otherwise related to
them, such as ellagic acid or gallic acid, the presence of which
in nature can, moreover, not yet be accepted with complete cer-
tainty. Morin and morin-tannic acid, which react in the same
manner with iron salts, have also not yet been met with in those
parts of plants to which we here devote attention. Further-
more, pyrocatechin, quercitrin, and rutin must not be omitted
here, which likewise color solutions of ferric salts green. The
_ first-mentioned substance can, indeed, be cited here only asa
very subordinate constituent of kino, and quercitrin is contained

‘Compare Wiegand, “ Satze iiber die physiologische Bedeutung des
Gerbestoffes und der Pflanzenfarben.” Botanische Zeitung, 1862, 121;

and Kutscher, ‘* Ueber die Verwendung der Gerbsdure im Stoffwechsel
der Pflanze.” Flora, 1883, _

* Berg’s ‘‘ Atlas,” Plate 49, Fig. 136.

TANNINS. 137

in Flores Rose gallice, but the latter substance, as well as
pyrocatechin, is undoubtedly widely distributed in the vegeta-
ble kingdom, and by more exact investigation will probably be
. found in many other drugs.

Of very frequent occurrence also, and probably quite general
in barks in a definite phase of life, is phloroglucin,’ C,H,-
(OH),, belonging to the class of phenols.

Resembling the tannic acid of galls, or tannin,’ there are
some other tannic matters, not of the same composition, which
produce in solutions of ferric salts a blue-black precipitate, thus,
the tannin of Folia Uve ursi, of oak-bark, of the bark of pome-
granate-root, etc. Many others, however, as the tannic acid of
the Cinchona barks, of willow and elm barks, that of Radix
Ratanhie Peruviane, of Rhizoma Filicis, Rhiz. Tormentille,
of Coffee, and also Catechu, produce with solutions of ferric
chloride or ferric salts a green precipitate, while the tannic
acid of rhubarb gives a blackish-green. In two varieties
of Ratanhia (Krameria), that from Para and that from
Savanilla, the tannic acid forming a green coloration with
iron salts is accompanied by a predominating amount of acid
producing a blue coloration. For the correct discrimination of ~
these colorations, thin sections of the respective drugs must be
moistened with a little solution of ferric chloride of the dilu-
tion stated under ‘Microscopical Reagents,” and the slides
upon which this reaction is carried out, laid upon a sheet of
white paper. The experiment is also performed at the same
time with the application of a solution of ferrous sulphate,
which permits the colorations to appear gradually, in proportion
to their oxidation, but often with a greater degree of purity.
A highly remarkable occurrence of a substance which affords a
magnificent blue color with ferric chloride as well as with fer-
rous salts is presented by the large cells of the fleshy portion of
the fruit of Siliqgua dulcis.

Between the tannic matters or tannic acids of the two classes

~1 Compare the statements of Tschirch, in Pringsheim’s Jahrb. f. wiss.
Bot., 1885, and Poulsen’s ‘‘ Botanical Micro-chemistry.” se
? From the French word tanner, to tan, of unknown origin.

138 PLANT ANATOMY.

above indicated, sharp chemical distinctions exist, which are ren-
dered evident, especially, upon dry distillation. When subjected
to this treatment, the tannic matters which produce a blue color
with ferric salts afford pyrogallol (pyrogallic acid), while those
producing a green color with ferric salts, on the contrary, afford
pyrocatechin. If the tannic matters are melted with caustic:
potassa, those giving a blue color with iron salts afford pyro-
gallol, as in the former case; the other tannic matters, on the
other hand, produce protocatechuic acid.

The knowledge of the different members of the chemical
family of tannic matters, in their details, is still very fragment-
ary. A method is also still wanting which meets all demands
for the quantitative estimation of tannic acids in all-the numer-
ous Cases where they cannot be extracted with a tolerable degree
of purity by ether-alcohol, as, for example, from nutgalls. Be-
sides, if it be considered that the amount of tannic matter is
subject to the fluctuations of vegetation, it cannot be greatly
wondered at that the analytical statements relating to it deviate
widely from each other. Many such estimations have been
made from a technical standpoint, as in the case of oak-bark,
so that the literature on this subject is quite extensive.’ The
oak-bark appears to be capable of containing a maximum of
twenty per cent of tannic matter, or more than any other part.
of a plant which concerns us here,’ unless we take into consider-
ation the galls (see the chapter at the end of this work: Patho-
logical Formations). The latter, namely, are to be regarded
simply asa morbid accumulation of tannic acid. The gallo-

' It may suffice to mention here the following: ‘‘ Bericht iiber die Ver-
handlungen der Commission zur Festellung einer einheitlichen Methode
der Gerbstoffbestimmung, gefiihrt am 10. November 1883 zu Berlin.
Redaction und Einleitung iiber die bisherigen Verfahren der quant. Be-
stimmung des Gerbstoffs von C. Councler. Nebst Untersuchung tiber
die Léwenthal’sche Methode von J. V. Schroeder,” large 8vo (IV., pp.
79), Cassel, Fischer, 1885.

* The bark of the Australian Eucalyptus corymbosa is stated to con-
tain twenty-seven per cent of tannic acid (Jahresbericht der Chemie,
1868, 807), the Myrobalans forty-five per cent: Dividivi, the pods of
Cesalpinia coriaria Willd., fifty-five per cent,

TANNINS, 139

‘tannic acid, which is present in these malformations to the ex-
tent of as much as seventy per cent, is remarkable as a specially
distinct member of the family of tannic matters; at least the ex-
ceptional cases in which it is supposed to have been elsewhere
recognized (in the Myrobalans and in sumach) may still be re- .
garded with doubt.

There may be distinguished physiological and pathological
tannic matter. The former is produced normally in the vital
process of the plant (thus the tannins of barks,’ such as that of
the oak, quebracho, and the willow). The pathological, on the
contrary, is first produced in consequence of an external influence
(the puncture of-an insect, etc.), that is, in the course of a mor-
bid process (galls). Both forms are also chemically and physi-
cally different. Skins are only tanned by the physiological tan-
nin (as in the formation of leather).

The contents of the cells which have so far been treated of, if
we except inulin, may be regarded as the organized contents.
Besides these, however, there appear a number of unorganized
bodies in the cells of plants, which are dissolved in the cell-sap,.
or deposited in the membranes, and which are not amenable to
direct microscopical observation.

In the cell-sap there are dissolved, for example, a portion of
the inorganic salts, dextrin, sugar, plant acids—the cell-sap al-
ways has an acid reaction—and tannic matters, many glucosides*
and bitter principles (Aloé, Fig. 63), coloring matters, amides,
etc.; in the membranes are deposited many alkaloids (quinine ?).

The chief solvent of most of these substances, water, evapo-
rates to a large extent upon drying the drugs.* How considerable
its amount may often be is shown in a striking manner by many

' Compare in this connection, F. von Hoehnel, ‘* Die Gerberinden, ein”
monographischer Beitrag zur technischen Rohstofflekre.” Berlin, Op-
penheim, 1880,

_ * PAvuvs sweat, and 7605 likeness.

* To this fact, as it appears, is referred the expression drug, German
droge; the u, which is still frequently inserted in the latter word
(drogue), is derived from the Romanic languages, which have appro-
priated the word. Fliickiger, ‘‘ Geschichte des Wortes Droge,” in Archiv
der Pharm., 219 (1881), 81.

140 PLANT ANATOMY,

roots. ‘The younger roots of Belladonna lose as much as eighty-
five per cent of water; Radix Tarazaci, seventy-seven per cent,
and juicy fruits still more. All parts of plants, however, retain
water which we are wont to designate as hygroscopic water, but
which by no means exists in the cells in a liquid form. The
amount of this varies very considerably according to the nature
of the tissues, and presumably also according to their con-
tents.

The squwiil, which is rich in sugar and mucilage, retains four-
teen per cent of hygroscopic water, Radix Gentiane sixteen to
eighteen per cent, and saffron about twelve per cent. If these
substances are completely deprived of water in a drying-closet,
or in the cold over sulphuric acid, and are again exposed to the
ordinary conditions of preservation, they quickly absorb again
about the same amount of water. Drugs which do not have a cel-
jular structure likewise contain definite amounts of water; per-
fectly air-dry starch, gum arabic, and tragacanth, for example,
thirteen to seventeen per cent. Seeds, on the contrary, and es-
pecially those provided with a hard testa, are capable of retaining
but a few per cent of water.

After the evaporation of the water, dissolved substances are
deposited in a solid form, as has already been mentioned when
speaking of inulin. Only a limited number of substances in-
soluble in water are capable of preserving in the dry tissue so
liquid a form as to flow in drops. Such are the volatile oils, the
boiling point of which lies from 70 to 150 degrees or more
above that of water, in consequence of which they evaporate only
to a slight extent with the water at ordinary or only slightly
elevated temperatures, and are still further retarded in this re-
~ spect when they contain resins in solution.

It is remarkable that the milky juice of Jalap also still pos-
sesses, in the dried drug, a liquid form, and indeed the resin pre-
pared therefrom is capable of retaining water very obstinately.

Besides the loss of water, and probably also of a portion of the
volatile oil, many plants experience, upon drying, chemical
changes, regarding which we are indebted to Schoonbroodt ' for

* Wiggers-Husemann’s Jahresbericht, 1869, 9.

AIR—SUGAR. 141

some valuable information, and which deserve to be further
studied. Drying changes the properties of many drugs. With
some, peculiar substances first appear during the process of dry-
ing, while others lose certain principles or acquire a different
odor (compare also page 15).

The amount of residue which remains upon drying vegetable
objects at from 100 to 110° C., until of constant weight, is
termed the dry weight. Drugs dried at ordinary temperatures
(about 15° C.) are called air-dried.

While most parenchymatous cells during life contain, besides
protoplasm and cell sap, only little or no air (bast-cells, vessels
and intercellular spaces contain it abundantly), the cells of dry
drugs are generally more or less filled with air, since upon dry-
ing this takes the place of water. It is evident from the nature
of the case that a complete replenishment of the cells with air
is not perceptible by direct observation. In cells which are
still succulent and vitally active, on the contrary, and in such
which are impregnated with liquids for the purpose of examina-
tion, as is necessary in making microscopical preparations, the
air bubbles escape as dark rings from the liquid, in consequence
. of the total reflection of the rays of light. With these the be-
ginner in microscopical observation soon becomes sufficiently
acquainted, so as not to mistake them for something else.
Tissues filled with air (cork, wood) float upon water, notwith-
standing the fact that the specifie gravity of cellulose and of
cork is greater than that of water. Tissues free from air (for
instance, the heart-wood of Guatac), or such from which the air
is removed, sink in water, as is also the case with thin lamine
of cork, or with Lycopodium, as soon as the air has been ex-
pelled therefrom by boiling.

Of the dissolyed substances contained in the cells the follow-
ing may yet be considered:

Sugar is a very widely distributed constituent of drugs.
Cane-sugar, and the other varieties of sugar, are so abundantly
soluble in water, and probably also in most cell-juices, that
even after drying they appear but rarely in a crystallized form
or otherwise as a solid constituent of the cells. The more spar-

142 PLANT ANATOMY.

ingly soluble milk-sugar, which does not, however, require more
than seven parts of water for solution at the ordinary tempera-
ture, has as yet been found but once (1871) in the vegetable
kingdom, in the fruit of the tropical Achras Sapota L.

Grape-sugar (Dextrose),’ deviating to the right, is of most
frequent occurrence in the vegetable kingdom; it occurs, for
instance, in grapes, figs, pears, cherries, in liquorice-root and in
tamarinds.

Fruit-sugar (Mucilage-sugar, Levulose *), deviating to the |
left, is contained in honey, and often mixed with grape-sugar.

Cane-sugar (Beet-root sugar, Saccharose), deviating to the
right, is contained in the sugar-cane, sorghum, the sugar-beets,
in carrots, and in the sap of the sugar-maple. By inversion ° it
passes into a mixture of one molecule of grape-sugar (dextrose)
and one molecule of fruit-sugar (levulose), the so-called invert-
sugar‘; the latter is found in fruits and in honey.

Mycose® (Fungus-sugar) is found in fungi, for instance, in
ergot.

Melitose* is found in the Manna obtained from the leaves of
species of Hucalyptus (Australian Manna).

Grape sugar is detected in the cells micro-chemically by
Trommer’s reaction.’ The sections are placed successively in a
concentrated solution of sulphate of copper in water (they
should be well washed, but not too long), and in dilute caustic
potassa, and boiled in the latter, If sugar be present, there is
formed in the cells a red, granular precipitate of cuprous oxide,
Cu,O. (The execution of this reaction requires experience. )

As aresult of incisions, there are formed in the Manna-ash

1 Dexter, right.

2 Leevus, left.

5 By boiling with dilute acids.

* Invertere, to invert, for the reason that invert-sugar deviates to the
left, that is, in an opposite direction to cane-sugar,

5 Moxos fungus.

6 MéAz honey.

‘Sachs, “ Microchem, Reactionsmethoden.” Wiener Academie.
Sitzungsberichte, 1859. :

HESPERIDIN. 143

crystalline exudations (Manna), which contain as much as
eighty per cent of a sweet principle, mannite C,H,(OH)..

The glucoside hesperidin’ is contained in unripe fruits of
the Aurantiex, dissolved in the cell-sap, especially in the various
Species of Citrus (very abundantly in Fructus Aurantii imma-

Fie. 63.—Transverse section through the marginal portion of a leaf of Aloé socotrina.
-ep, epidermis (c, cuticle); sp, stoma; a, respiratory cavity; p and g, assimilating tissue; cr,
crystal cells (with raphides); a, cells containing aloes (the large ones contain chromo-
en); gfb, vascular bundle; m, medulla containing mucilage.

turi). By immersing the fruits in alcohol, there are produced in
the cells sphero-crystals, similar to those of inulin (Fig. 54),
1 Pfeffer, Botan, Zeit., 1874, p. 481. Tiemann and Will, “Ber. d.

Deutsch. Chem. Ges.,” 1881, 946. Virgil named the Seville oranges the
apples of Hesperides, the daughters of night in Grecian mythology.

144 PLANT ANATOMY.

which are soluble in slightly alkaline water aud in alcohol.
Crystals of this character have also been detected by Adolph
Meyer * in the leaves of Coniwm maculatum.

The peculiar bitter substances of Aloé leaves, aloes, are con-
tained in special cells, which are located directly in front of the
vascular bundles (Fig. 63 @), and are confined toward the exterior
by a nucleus-sheath in a single row and with bitter contents. The
before-mentioned cells are short, and occasionally their contents.
are crystalline. The entire remaining tissue of Aloé leaves con-
tains an abundance of mucilage, but no bitter substances.

The kino, from species of Pterocarpus, also occurs as a con-
stituent of longitudinally extended cells.’

When a part of a plant is incinerated, there remains in the cru-
cible a white residue—the ash. Since by careful ignition, in
very many cases, the general outlines of the consumed portion
of the plant remain preserved (leaves of the Graminex, hemp
leaves, the shells of diatoms), it follows that the inorganic con-
stituents of the membrane which resist the action of heat (at least
in part) are so finely deposited that the molecule of the mem-
brane can be removed therefrom by incineration, while the direct:
connection of fhe inorganic particles is not destroyed thereby.
Not only in the membrane, however, do we meet with deposits
of mineral constituents, but the contents of the cells are also
abundantly provided therewith. It has already been shown that.
protoplasm contains an abundance of salts, that crystals of
inorganic bases occur in the cell-sap, and also that the globoids
(p. 98) consist of inorganic double salts. The cell-sap, more-
over, also contains not inconsiderable amounts of such mineral
constituents of plants as are soluble in water, and these are in
fact the most important, namely, nitrates, phosphates, and
salts of potassium and calcium.

Accordingly, the ash of plants contains all those substances
which are known to, be the necessary nutritive materials, name-
ly: potassium, magnesium, calcium, iron,’ phosphoric, sul-

' Compare also Fliickiger, ‘‘ Pharmakognosie,” 1883, p, 663.

* See the subsequent chapter on Receptacles for Secretions.

* A deficiency of iron is shown in leaves by their becoming yellow
(chlorosis).

ASH. * 145

phuric and nitric acids, and chlorine. There also occur in it
silicium, sodium, manganese, aluminium, iodine, bromine,
fluorine, lithium, and other elements.’

Plants rich in silicium (grasses, diatomer)—they contain it
always in the membrane—leave upon incineration a so-called
skeleton of silica.*, The Halophytes (salt-plants) especially con-
tain sodium. Manganese is less extended, but is nevertheless
found regularly, even though in small amount, for instance, in
drugs from the family of Zingiberacex.* It suffices to reduce
to ash a single seed of the cardamom, or a still smaller fragment
of the fruit-capsule, by heating on the looped end of a plati-
num wire in the oxidation flame of an ordinary alcohol lamp, and,
if necessary, fusing with a little sodium carbonate and a trace
of saltpeter, in order to obtain a bead which is colored green by
the manganate of the alkali, and which when moistened with
acetic acid affords the red permanganate. The same deport-
ment is shown by the root-stocks of this family. The ash of
ordinary cork (from Quercus suber) and that of other species of
cork is also green from the same cause.

Aluminium is of rare occurrence, but is found in not incon-
siderable amounts in the leaves and stems of species of Lyco-
podium,

Lodine and bromine occur in the vegetable (and animal)

?To interpret the composition of the ash is far more difficult. Weare
not yet capable of explaining the great differences found therein accord-
ing to some general law. A very extensive compilation of figures re-.
lating to this subject may be found in Wolff’s ‘‘ Aschenanalysen von
landwirthschaftlich wichtigen Produkten, Fabrikabfillen und wild
wachsenden Pflanzen,” Berlin, 1871. .

* Such skeletons may be prepared by warming small pieces of coarse,
firm leaves with concentrated sulphuric or nitricacid and potassium
chlorate, expelling the acid, and heating the residue upon platinum-foil
(preferably in a current of oxygen) or upon a very thin cover-glass until
it becomes white. Tissues which have not previously been treated in
this manner often fuse together in consequence of the amount of alkali
contained therein.

*Flickiger, Pharm, Journ., III. (London, 1872), 208. Ibid., 1886, p.
621; also Amer. Journ. Pharm, 1886, p. 147.

10

146 PLANT ANATOMY.

inhabitants of the sea; fluorine in the testa of the seed of varie-
ties of grain, and lithium in tobacco.

As silicium has in recent times been introduced into organic
compounds in the place of carbon, the supposition is not entirely
unjustifiable that the silicium contained in the cell-wall may be
present in the form of an organic compound.*

As is already naturally evident from mechanical principles,
an increased thickening and solidity of the cell-walls corresponds
by no means to a greater amount of incombustible substances.’
The delicate tissue, containing air, of peeled colocynth, dried at
100° C., afforded 11 per cent of ash, the seed only 2.7 per cent.
Quassia wood from Surinam yields 3.6 per cent, the bark 17.8
per cent of ash; guaiac wood, which is so exceptionally dense,
and which consists almost exclusively of strong wood-cells, gives,
nevertheless, scarcely 1 per cent of ash. Leaves very frequently
contain more than 10 per cent of inorganic constituents, for
instance, Folia Stramonii as much as 17, and tobucco leaves
occasionally 27 per cent of the substance dried at 100° C.

The developing or merismatic, and the assimilating tissues
(cambium, mesophyll of leaves) and organs (leaves* and barks)
are richer in ash than the completely developed and non-assimi-
lating (wood). The mineral substances migrate from the fin-
ished tissues to the places where development is going on. It
is also seen from this that they must play an important part in
the formative processes of plants.‘

To obtain the ash in a condition suitable for weighing is often
somewhat difficult, from the fact that many parts of plants,
and especially secreted substances, as gum, resin and sugar,
undergo complete combustion only very gradually. The incin-
eration of such substances, especially of those rich in nitrogen,

'“ Berichte d. Deutsch. Chem. Ges.,” 1872, 568.

*That the strength of flexure, for instance, of the grasses, is entirely
independent of the silica contained therein, is shown by rematmieanens
in solutions free from silicium,

* About the time of the falling of the foliage the leaves become con-
mabey poorer in mineral substances.

‘This is also evident from the experiments with solutions of nieve
_ substances,

ASH. 147

may be very much accelerated when the objects to be examined
are heated on a channeled piece of platinum-foil in a combustion
tube in oxygen gas. The same purpose may be attained in a
more simple manner, though also more slowly, when the sub-
stance which has been carbonized in a platinum capsule is moist-
ened with water, again carefully allowed to dry without decant-
ing the water, and again heated. The water conveys the soluble
salts to the unoccupied places of the capsule, and the subsequent
admission of air facilitates combustion. If this procedure is
repeated several times, a residue free from carbon will in most
cases be obtained. Too high a temperature has a retarding
effect when salts, such as phosphates of the alkali metals, are
present, which fuse together and envelop the carbon; many
substances are incinerated more completely by a very moderate
degree of heat than at a higher temperature. Very hard shells
of seeds offer obstinate resistance to the above procedure of
moistening, which may be overcome by triturating the carbonized
substance in the capsule, or in the crucible itself, with the aid
of a very smooth agate pestle, being careful to avoid loss, and
afterwards treating with water. By the strong ignition, which
ordinarily is necessary towards the end, carbonic acid is ex-
pelled, which must be replaced before the ultimate weighing of
the ash, in order to obtain figures which will admit of compari-
son. ‘This purpose is accomplished by moistening the ash with
a little concentrated solution of ammonium carbonate and
again drying. It scarcely requires to be mentioned that for
reduction to ash the substance employed should previously be
dried at 100° C.

The addition of ammonium nitrate or ammonium sulphate
also facilitates combustion, especially with substances rich in
albumen. :

The estimation of the residue left upon combustion is of the
greatest practical value, especially for the examination of vege-
table powders. For, since every portion of a plant, and thus also
every drug,. furnishes an amount of ash which fluctuates within
definite and often quite narrow limits,’ the weight of the same

‘Thus, by way of example, lycopodium affords 4, pure kamala about

148 PLANT ANATOMY,

may therefore afford information whether an adulteration with
other vegetable or even inorganic powders has taken place.

The estimation of ash must, of course, always be preceded bya
microscopical analysis of the substance itself.

II. The Cell-wall.

The integument of the cell is called the cell-membrane or cell-
wall.

The cell-wall of young cells is a thin membrane which con-
sists of cellulose, and only at a later period becomes variously
changed, either chemically through the deposition of other sub-
stances, or morphologically through the insertion of molecules
of the same kind. Even the membrane of young cells, however,
is not perfectly pure cellulose, since it owes its first formation to
the protoplasm, remains for a long time in contact with the
nitrogenous substances of the latter, and is penetrated by them.
Only cellulose which has been purified by means of chemical
solyents corresponds to the formula C,,H,,0.,.

In living cells, the wall is in most intimate contact with the
protoplasm-sac. To this contact is to be referred the growth
of the cell.

The growth of the cell takes place in a twofold manner, on
the one hand by a change of form, and on the other by a trans-
formation of the chemical nature of the cellulose, which, in the
course of deyelopment of the cells, is capable of assuming a series
of new chemical and physical properties.

The change in form of the cell concerns either chiefly its out-
line, and in this case may be considered surface-growth, or the
development of the cell is specially expressed by a thickening of
the wall, so that the growth in thickness determines the appear-
ance of the cell. Although both directions of growth are not
sharply to be separated, and are essentially based upon the

same processes, they nevertheless deviate widely from each other
in their results.

2, lupulin about 8, starch less than 1, cacao about 4, mustard seed and
flaxseed from 4 to 4.5, and pepper about 5 per cent of ash.

THE CELI-WALL. 149

If through a perfectly uniform deposition of new particles of
cellulose the mass of the cell-wall becomes equally enriched all
around, but not actually thickened, it is compelled to assume a
spherical form, and the cells become isodiametric,’ as in many
young tissues. Their mathematical regularity, however, is.
altered, as soon as the reception of constructive material takes.
place more energetically in certain places. The outline of the

Fic. 64.—Schematie representation of the development of the wall of a wood-cell. a,
youngest state; f, the finished condition. It is only in the first three stages that the
nucleus of the cell is preserved (Hartig). A film of intercellular substance closes the
pits of the cells.

cells is also very essentially controlled by the fact that they
mutually oppose their free expansion. In such cases the form
of a sphere becomes flattened to that of a dodecahedron, which
is the most uniform of those forms of cells of so frequent
occurrence which we designate as spherically-polyhedral, since

Iéos equal, and é:a@uyrep diameter.

150 PLANT ANATOMY.

from their variety and slight regularity they preclude a more
precise definition (Figs. 29, 30, 31, 56, 63, 65, 76).

When the deposition of new cell material does not take place
chiefly in a direction tangential to the cell-wall, but in such a
manner that the latter grows in thickness, this growth can take
place more largely either toward the exterior or toward the
interior. In the first case, prominences of various kinds are
formed (spores and pollen cells, the outer wall of epidermis
ells), in the latter, the cavity of the cell becomes contracted,

Oy

if

Fie. 65, Fie. 66,

Fie. 65.—Polyhedral parenchyma from Rhizoma Graminis.
Fie. 66.—Uncoiling spirals and an annular vessel from Bulbus Scilla.

often almost entirely filled up (some bast cells and stone-cells).
A perfectly uniform thickening, however, never takes place, but
the cell-membrane retains in some places its slight thickness.
The appearance of the cells which are subjected to a considera-
ble extent to the growth in thickness is chiefly determined by
the relative extent of the thickened places and those which have
remained thin. If the thickened places are in about the same

THE CELL-WALL. 151

proportion as those which have remained thin, the membrane
presents, upon a transverse section, a necklace shaped (monili-
form) appearance. Such cells are, for instance, highly charac-
teristic of the coffee-bean.'| Where the thickened places do not
attain great extension, and appear especially upon the inner
Surface, they often assume the form of rings or spiral bands.
Thus originate the spirals in many fibro-vascular bundles, as,
for instance, in the sguiil (Fig. 66), as also the net-like and
scalariform thickenings (Fig. 68) of the vessels and parenchyma
cells (Figs. 67, 83, and 182). :

Fra. 67.—Cells with net-shaped thickenings (Dippel). Compare also Fig. 182.

When thickening of the cell-wall extends over the largest part
of the inner surface, and exempts but a few expanded dot-shaped
places, pores are produced (Fig. 69). With a considerable thick-
ening of the cell-wall such places appear as dots, or by a still
greater increase in the thickness of the wall as pore-canals
(as in the stone cells, Fig. 70). Frequently a spiral-shaped
arrangement of the dots may be observed (Fig. 71), and the
course of the pore-canals also often approaches that of a
spiral line. Many true bast-cells have cleft-shaped dots

1 Berg’s ‘‘ Atlas,” Plate 49, Fig. 131.

152 PLANT ANATOMY.

arranged in a spiral inclining to the left (compare the subsequent
references under Mechanical System of Tissue). A special

Fie. 68.—Longitudinal section bhrougt. vessels with scalariform thickenings (/v).—
Rhizoma Filicis (Berg.) ig

form of thickening is represented by the bordered pits or
areolated dots (Fig. 72). When the cell-wall becomes thickened

Fic. 69.—Porous cells.

toward the interior around a place which remains thin, a canal
will remain open, which in form must approach that of a very

AREOLATED DOTS. 153

obtuse cone, in so far as the walls of the canal are not superposed
perpendicularly to the wall of the cell. If in this manner the
canal becomes narrower toward the interior, it finally corre-
sponds in form to a somewhat dilated funnel. The upper edge
corresponds to the place in the wall which has remained un-
thickened, and within this circle or border the aperture of the
funnel toward the cell-cavity appears as a pit.

Similar areolated pits are wont to appear simultaneously at
such places where two cells come in contact by the surface of
their walls; the intervening wall, which, moreover, does not
always lie in the median line of the pit, but is often, as in

Fie, 70.—Thickened cells with pore-canals. A, Bast-fibre of a Cinchona Bark, B,
Stone-cells from a nut-shell. (8 from Dippel.)

Fig. 64, d, e, f, pressed against one of the apertures (wood-cells
of the Conifers), disappears by age, so that the space occupied
by the pit establishes a direct connection between the two cells
(Fig. 72, A, C). These hollow spaces, which sometimes resem-
ble two funnels, one inyerted over the other, and which are |
sometimes arched in a more lens-shaped manner (Fig. 72), are
easily recognizable where they occur more isolated. If, however,
they are formed in larger number closely beside each other, and if,
through increasing thickening, they become gradually contracted
in a cleft-like form, more complicated relations are produced,

154 PLANT ANATOMY.

which are clearly disclosed only in thin and carefully prepared
sections (Semen,Colchici).

In those cases also where thin places of the cell-membrane
remain preserved only in extremely slight amount and extent,
the growth in thickness does not take place through the simple
deposition of new encircling scales or layers of cellulose. The

ro)
7
™~ v

Fie, 71.
Fia. 71.—Spirally arranged pits,

Fie, 72.—Areolated dots of the tracheids of fir-wood. 4, transverse section through
the tracheids or wood-cells, the pits shaded ; Band C, schematic longitudinal sections;
the spherical lines denoting the circumference of the pit and the border; D, two adja-
cent pits cut in the direction of length, with the partition-wall still retained (Sachs).

stratification, which is often highly remarkable, depends upon
variations in the amount of contained water and in the condi-
tions of tension of the individual layers ; those containing less —
water and which are denser, stand out distinctly in consequence

STRATIFICATION. 155

of their greater capability of refracting the light. Chemical
distinctions (various degrees of lignification) also play a part in
this. The function of the contained water may be proved by
complete desiccation or by more complete swelling, both of
which equalize the differences, and often either break up the
stratification or obliterate it toa large extent. The bast fibres
of the Cinchona barks possess almost entirely thickened walls,
with distinct stratification (Fig. 73). When they are softened
by means of energetic reagents, such as caustic soda, concen

= SSS
SSA
SQ
=
=S

SES

Fie. 73.—Bast-fibres from Cinchona-barks.

trated sulphuric acid, or ammoniacal oxide of copper, and the
tension of the particles of cellulose becomes equalized, it is dis-
tinetly seen that the thickening is not due to a simple concen-
tric succession of layers, but to far more complicated processes,"
In the Cinchona fibres, particularly, there is brought to view in

1A more precise elucidation of these remarkable conditions is given
by Nageli, «‘ Bau der vegetabilischen Zellmembran.” Sitzungsberichte
der Miinchener Akademie, June, 1864, page 145. Also Sachs, ‘‘ Lehr-
buch der Botanik,” 1873, p. 30 et seg. Wiggersand Husemann, Jahres-
bericht, 1866, 89.

156 PLANT ANATOMY.

the manner indicated a screw-shaped disposition of the thick-
ening (Fig. 74). Hofmeister’ found by maceration of these
fibres in nitric acid and potassium chlorate, and subsequent.
pressing, a more scale-shaped arrangement of the layers.

Cells, which have walls of considerable thickness when com-
pared with the diameter of the lumen (cell-cavity), that is such
in which the latter is contracted to a very small cleft, are desig-
nated as dast cells when they are extended in length (Figs. 110
and 111), or as stone-cells (Figs. 75 and 76°) when they are but

Fic. 74.—4A, Bast fibres from Cinchona barks, boiled with hydrochloric acid,; P, the
same softened in ammoniacal oxide of copper after treatment with hydrochloric‘acid,
(P from Dippel) ; 7, original size of the cell ; s, the swollen layers,

short. The latter, particularly, show a very distinct stratifica-
tion of the membrane. ;

1 « Verhandl. d. Sachs. Gesellsch, d. Wissensch.,” X., 1858, p. 32.

* They may also be called sclereids (derived from 6%Anp6s hard) in op-
position to the proper mechanical cells, the stereids (from érepe6s mas-
sive). Compare Tschirch, ‘“Beitrige zur Kenntnis des mechanischen
Gewebesystems,” Pringsheim’s Jahrb., 1885, and ‘Ber. d. deutsch.
botan. Ges., ITI. (1885), No. 2,

STONE-CELLS. 157

The thickening layers build themselves up, over the places
which remain thin, in such a manner that the small canals run-
ning toward the centre or the axis of the cell often present a
sort of star-shaped arrangement (Fig. 76).

Fie. 75,—Various stone-cells.

The stone-cells or sclereids (see also subsequent references
under the section: Mechanical System of Tissue) are widely
distributed in many barks, the testa of seeds, seed-vessels, etc.
A series of remarkable and manifold forms of them is readily

A. B

Fie. 76.—Stone-cells, whose cavities, t, though radiately arranged pore-canals, p, are
brought into connection with the outer surface, or even with adjacent cells, 7 1, 2, 3,
thickening layers (Dippel). ;
afforded, for example, by the star-anise. The fruit-stalk con-
tains branched stone-cells,‘ the wall of the capsule such as are
nearly cubical.

' Vogl, ‘‘ Nahrungs- und Genussmittel.” Vienna, 1872, 111.

158 PLANT ANATOMY,

The thickening of the cell-wall may also, under certain con-
ditions, confine itself to the corners, thus forming the so-called
collenchyma,' which is met with in the barks and seeds of very
many plants.

The relative thickening of individual cells and forms of cells
is, moreover, very manifold. While the parenchyma remains
mostly thin-walled until the close of life, the wood- and bast-
cells become provided with strong walls at a very early period.

Bast-tubes and stone-cells when observed in thin sections
under glycerin in polarized light, are seen to be doubly refrac-
tive (Fig. 77). A transverse section through cinchona fibres

—

A B

Fic. 77.—Thin sections through bast-fibres and stone-cells, showing double refrac-
tion in polarized light (Dippel), p, s, s’, layers of different density.
shows four dark arms of a cross upon a brightly shining ground
(Fig. 77, I.).

In the preceding pages, those morphological changes of the cell-
wall have been considered which take place in the process of
vegetation. It still remains for us to subject the chemico-phy-
sical changes to closer consideration.

’ Derived from x0AAaq, glue, since it was formerly, but incorrectly,
believed that the collenchyma cells could become mucilaginous.

* With regard to the chemistry of the cell-membrane, compare partic-

ularly the more recent researches of Cross and Bevan, ‘‘ The Chemistry
of bast-fibres,” in the Chem. News, 1882; Webster, ‘* On the analysis of

CELLULOSE. 159

The cell-wall is subject to chemical and physical changes,
either through the deposition of woody matter, lignin’ or cork-
fat (suberin),? as also through a retrograde metamorphosis of the
cellulose into gum and mucilage.

All young cell-membranes, and most of the walls of cells, with
which we shall become acquainted under the designation of
parenchyma, as also many appendages of seeds which are de-
veloped as hairs (Cotton, Asclepias, Eriodendron, Salix), consist
of purecellulose. ‘The phloém, Jeptom (sieve-tubes and cambi-
form tissue), always remains unlignified. Membranes consisting
of cellulose show, even by superficial microscopical observation,
an entirely different capacity for the refraction of light from
lignified and suberified membranes; they appear clearer, more
strongly refractive, and jelly-like (collenchyma, the cell-mem-
branes of Macis). Cellulose membranes are digestible.

As already intimated on page 123, there are some exceptional
cell-walls which, in contact with iodine-water, are colored in a
similar manner to amylum. By the treatment of pure cellulose
with mineral acids, this faculty may quite generally be imparted
to it. The respective sections or objects (for instance, cotton)
are moistened for an instant with sulphuric acid of the specific
gravity 1.84, washed without delay with much water, and then
powdered iodine strewn upon the moist preparation, or it is
impregnated with iodine-water (see Micro-chemical Reagents).
The reaction succeeds with still greater certainty with phos-
phoric acid, which is first concentrated as much as possible on a
water-bath. When hydriodic acid has been formed in an iodine
solution which has been long preserved (see Micro-chemical
Reagents), such a solution can effect the blue coloration of cellu-
lose without the co-operation of other acids, The reaction admits
of demonstration, without further preparation, with moistened
parchment-paper, which is sprinkled over with finely-powdered

certain vegetable fibres,” Ibid., 1882; Schuppe, ‘ Beitrige zur Chemie
des Holzgewebes.” Inaugural Dissertation, Dorpat, 1982, in which the
older literature is also to be found.

1 Lignum, wood,

2 Suber, cork.

160 PLANT ANATOMY.

iodine. A solution of chloride of zinc with iodine colors cellu-
lose membranes violet.

The cellulose of fungi, and suberified and lignified membranes
are not colored by the above-described treatment; in the case of
the latter two, however, this may be brought about if they are
previously boiled with nitric acid (specific gravity 1.185) with
the occasional addition of a few crystals of potassium chlorate
(‘‘ Schultze’s maceration”). The cellulose of fungi, however,
even after this treatment, is not colored by iodine.

Concentrated sulphuric acid alone (specific gravity 1.84)
dissolves cellulose with complete chemical change. This is not
the case when cellulose is dissolved in ammoniacal oxide of cop-
per (see Micro-chemical Reagents). In contradistinction to the
lignified membrane, pure cellulose possesses an exceedingly
slight inclination to take up aniline colors (page 161).

Pure cellulose may be prepared by the successive treatment of
tissues consisting of this substance (cotton, the pith of the elder
and of Aralia papyrifera) by means of caustic potassa, acids,
water, alcohol, and ether, or by the precipitation of its solution
in ammoniacal oxide of copper by means of water.

One of the most widely distributed modifications of cellulose
is formed through the deposition of lignin (xylogen’). A
membrane thus altered is termed lignified.”

Lignification appears at very early stages in the so-called wood-
cells. The wood-cells which, in dicotyledons, are separated
toward the interior by the growth in thickness, already possess
lignified membranes long before they become thick. The bast-
cells and many stone-cells (sclereids) are also often lignified. A

1 EvAor wood, and yervaa produce.

*Compare in this connection Stackmann, ‘‘ Studien iiber die Zusam-
mensetzung des Holzes,” Inaugural Dissertation, Dorpat, 1878, and the
previously mentioned dissertation of Schuppe.—M. Niggl, ‘‘ Ueber die
Verholzung der Pflanzenmenbranen ” (an historical survey). Jahres-
bericht der Pollichia, Kaiserslautern, 1881.—Ebermaier, ‘‘ Physiologische
Chemie der Pflanzen,” 1882. In this work (p. 175) will also be found
statements relating to the amount of lignin contained in some woods.

Thus (according to Schulze) oak-wood contains 54.12 per cent, and fir-
wood 41.99 per cent of lignin.

CORK, 161

regular lignification of the membrane is also found in the walls
of vessels. Lignified membranes refract the light to a less
extent than those consisting of pure cellulose, and mostly
appear light-yellow under the microscope; they are hard and
elastic, and but little capable of swelling.

Lignified membranes are characterized micro-chemically by
the fact that with a solution of iodine in chloride of zine they
become yellow (not violet). In ammoniacal oxide of copper and
Schultze’s macerating liquid (page 160) they do not dissolve.
With aniline sulphate and dilute sulphuric acid they become
straw-yellow, and with phloroglucin and hydrochloric acid
cherry-red; the aniline colors are greedily taken up by them.
By boiling with Schultze’s mixture or with alkalies, the lignin is
removed, and the membranes thus treated then show the cellu-
lose reaction. Morphological alteration, however, by no means
goes hand in hand with a change of physical and chemical
character. While, for example, the wood-cells, even in quite a
young condition when their walls are but very little thickened,
are strongly lignified, and the very thin-walled cork-cells are
always suberified, the strongly thickened collenchyma, and many
bast-cells which are thickened so as to cause the lumen or cavity
to disappear, remain unlignified.

The third modification of cellulose is cork.

This is formed by the deposition between the cellulose mole-
cules of swberin, which latter consists for the most part of the
glycerin (propenylic) esters of stearic acid and of phellonic’
acid, C,,H,,0,.2. Suberin appears to be identical with cwtin.*

The @pidermis-cells of the more delicate organs of all land
plants are covered by a delicate film termed the cuticle. The
older organs, especially those of the stem, on the contrary,
develop on their outer surface a layer consisting of tabular cork-

1 @édXAor cork.
* Definitely established by Kiigler at least for the cork of Quercus
Suber (‘* Ber. d. deutsch. botan. Ges.,” I., p. xxx. , and Inaugural Disser-
tation, Strassburg, 1884),
’ Beside suberin, there is also found in cork a wax-like body, cerin.
The cuticle appears to contain more of the latter than the cork.
11

162 PLANT ANATOMY.

cells. The cuticle and cork are both produced through suberi-
fication (the respective deposition of suberin or cutin in the
wall of cellulose).

The cuticle may occasionally become very thick (the leaves of
Eucalyptus, Agave, Aloé) and in the same manner many layers
often develop cork (the Oak"). Since both the cuticle and the
cork are but slightly penetrable by aqueous vapor, they serve as
a protection for the organs of the plant against too strong eva-
poration.

Suberized membranes are mostly brown. They are just as
little digestible as lignified membranes, but resist putrefaction
very energetically, as does also the cuticle.

Micro-chemically, suberized membranes are characterized by
the fact that they dissolve neither in concentrated sulphuric acid
nor in ammoniacal oxide of copper. The cork and cuticle there-
fore remain behind when tissues are treated with sulphuric
acid.

It is, however, to be observed in this connection that the mem-
branes of the tissues of drugs, in consequence of their strong in-
filtration with the constituents of cells, which takes place dur-
ing the process of drying, often resist very obstinately the action
of reagents, even when no suberification, etc., has taken place.
It is only after repeated boiling with alcohol, water, and ether,
that such membranes are made accessible to reagents.

Suberin cannot be removed from the membranes by the ordi-
nary solvent of fats, but only through the action of alcoholic
potassa. It is therefore very firmly (perhaps chemically ?) com-
bined with the cellulose.

Closely related to the lignified and suberized membranes is the
so-called intercellular substance (middle lamella), or that
substance which cements the cells to each other (x in Fig. 64).

The intercellular substance ? is insoluble in concentrated sul-
phuric acid and in ammoniacal oxide of copper; on the other

‘Compare also the section : Epidermal Tissue.

* Compare herewith, among others, R. F. Solla, “« Beitrage zur naheren
Kenntniss der chemischen und physikal. Beschaffenheit der Intercellu-
larsubstanz,” in Oesterr. botan. Zeitschr., 1879.

WAX—GUM—MUCILAGES. 163

hand, it is soluble in nitric acid with the addition of potassium
chlorate (though not without decomposition) and in a hot solu-
tion of caustic potassa. Aniline colors are strongly absorbed by
it.

The cell-membrane of the living plant (and to an increased de-
gree that of drugs) contains, however, beside the deposited sub-
stances just mentioned, not only the organic constituents of the
cell which enter it by infiltration, but also abundant amounts of
inorganic compounds,’ as has already been explained (page 144
et seq.). These are mostly deposited molecularly (as silicium),*
more rarely in form of crystals (as in the spicula-cells of Welwit-
schia mirabilis and the epidermis of Dracena leaves).

As has previously been mentioned, the cuticle contains not:
only suberin, but also wax-like bodies. * Occasionally wax
issues from the membrane, and then forms coatings consisting”
of granules, small staffs, or crusts. If these are distributed im
slight amount over the epidermis, the plant organs assume the
appearance of being covered with hoar-frost (pruinosus), thus
the leaves of Lucalyptus, Ricinus and Cabbage, Plums, and many
other allied fruits, and Juniper berries. In some plants, how-
ever (especially several palms, Anacardiaceew, Myricacee), the
secretion is so considerable that the wax may be collected in
large amounts, as, for instance, the Carnauba-wax, from the
young leaves of the East Brazilian palm, Copernicia cerifera
Martius.

The different varieties of wax are esters (compound ethers) of
the fatty acids; upon saponification they do not, however, afford
glycerin, but other (monatomic, not triatomic) alcohols.

The varieties of gum and the mucilages, being closely related
to cellulose, must be considered in connection with the latter.*

? The amount of mineral constituents of the membranes is very vari-
able. The best quality of cotton, dried at 100° C., affords but 1.12 per
cent of ash.

* In the siliceous coatings of diatoms, and in the grasses.
* Compare De Bary, Botan. Zeit., 1871, Nos. 9, 10, 11, 34; and ‘* Ana-
tomie, ”p. 87 et seq.
aoe also Valenta, ‘‘ Die Klebe- und Verdickungsmittel.”—Cassel,
1884,

164 PLANT ANATOMY.

The relations of these bodies to each other, as also to cellulose,
have, however, not yet been made clear.

According to Giraud’ these substances may be grouped in the
following manner:

1. Ordinary varieties of gum: arabin, bassorin, cerasin;
2. Pectose: gum tragacanth (adragantin);
3. Plant mucilages in a more restricted sense:

(a) insoluble in alkalies and dilute acids (the cellulose of
quince-mucilage);

(6) insoluble in alkalies, and forming with acids glucose
and a variety of dextrin: flaxseed, mucilage of Irish
moss;

(c) soluble in hot, concentrated alkalies, and converted by
acids into dextrin and glucose.

Mucilaginous substances, in the broadest sense, are also dis-
tinguished by their behavior to nitric acid; some afford with
it mucic acid, C,H,(OH),(COOH),, while others do not.
Furthermore, the aqueous solutions of many varieties of gum are
precipitated by the normal acetate of lead, while others are only
precipitated by the basic acetate.’

Gum arabic and cherry-tree gum (cerasin), as well as traga-
canth, are formed by a retrograde metamorphosis of the cell-
membrane,’ that is, by a pathological process; the former two by
a conversion of the membranes of the peripheral layers of the
“‘horn-bast prosenchyma ” * into gum, the latter through a meta-

‘Compt. rend., 80, 477; compare also Husemann and Hilger, ‘“ Die
Pfianzenstoffe,” I. (1882), 131.

* Compare Kirchner and Tollens, Liebig’s Annalen, 175 (1874), 205.

* In reference to this and the following statements compare: Mohl,
Bot. Zeit., 1857, 33.—Frank, Jour. f. Pract, Chem., 95, 479; idem, Prings-
heim’s Jahrb. fir wissenschaftl. Botanik., V., 25.—Wigand, ‘‘ Ueber die
Desorganisation der Pflanzenzelle,” in Pringsheim’s Jahrb., III., 115.—
Prillieux, ‘‘ La formation dela gomme.” Ann. des sc. nat., 6 Ser., Bot.
B;..1 76.

* With regard to horn-bast, compare Wigand, Flora 1877, p. 369, and

“‘ Lehrbuch der Pharmacognosie,” 1879, pp. 9 and 38; further Flickiger,
“ Pharmakognosie,” 349, The word “horn-bast” should be expunged

pre the modern terminology.

GUM—MUCILAGES. 165

morphosis of the membranes in the medulla and medullary rays
(Fig. 79).

In the Amygdalex, however, gum also occurs abundantly in
the vessels and other elements of the wood, and often even en-

ee

}

Fic. 78.—The formation of gum in cherry-wood; g, aggregates of gum formed through
metamorphsis of the cell-membranes; r, vessels more or less filled with gum; m, medul-
ary rays; jf, annual ring, spring wood; jh, annual ring, autumn wood (Tschirch).

tire groups of cells of the woody structure suffer a conversion
into gum (Fig. 78 g), as in the so-called gum discase.

166 PLANT ANATOMY.

Beijerinck' attributes the origin of gum arabic, the “ gum-
mosis ” of species of Acacia of Africa, to the fungus Pleospora
gummipara Oudemans; another fungus, Corynewm Beijerinckit
Oudem., causes the gummosis of the Amygdalex. Frank does
not concur in this view, and Wiesner (Botan. Zeit., 1885, p. 577,
also Ber. d. Deutsch Chem. Ges., 1885. Referate p. 639), recently
attempted to show that the transformation of cellulose (and
starch) into gum or mucilage is due to a peculiar ferment, a
“diastatic enzyme.” At all events, the true gummosis, which is

certainly a pathological process, must be separated from the
gum formation which serves as a protection to tissues (see sub-
sequent references), and which is only intelligible from a physio-
logical point of view. They are also distinguished from each
other by the fact that the ‘‘ pathological gum ”—and only this
is of interest to us here—is formed through a metamorphosis of
the membrane, while the ‘‘ physiological gum” represents an
exudation of the membranes into the cell-cavities.

The transformation of cellulose into gum and mucilage can
also take place without so great an alteration of the cells and
tissues as in the case with gum arabic and tragacanth. In such
a case only one layer of the membrane becomes metamorphosed,
as is shown, for instance, by the conversion into mucilage of
the filamentous Alge. The gum mucilage of the glandular
hair (colleters) or many foliage buds, which is often mixed with
volatile oil and resin, is formed through the conversion into
mucilage of a membranous layer (collagen layer *) lying under-
neath the cuticle of the glandular hair. The mucilage of quince
seed and of flaxseed is probably also formed primarily through
a conversion into mucilage of only the secondary membrane of
the epidermis cells* of the respective seeds (Frank). The
seeds of many of the Papilionacesx, for instance, those of T’rigo-

'* Onderzoekingen over de Besmettelijkheid der Gomziekte bij
planten.” Amsterdam, Joh. Miller, 1884, 4to, 46 pages, 2 plates.

* KcAla@ glue, and yevvae to produce, Hanstein, ‘‘ Ueber die Organe
der Harz- und Schleimabsonderung in den Laubknospen.” Bot. Zeit.,
1868, No. 43,

: * Berg’s ‘‘ Atlas,” Plate XLVI., Figs. 122, 128 6.

GUM—MUCILAGES. 167

nella fenum grecum ( fenugreek seed), present an illustration of
the formation of mucilage occurring in the inner tissue, not in
the epidermis.’

In leaves, mucilage appears to be formed but rarely in larger
amounts. A very remarkable example of this character * is
presented by the Buchu leaves from Barosma crenulata Hooker
and other species.

But mucilages are also formed in the plant, apparently with-
out this direct participation of the membrane. They then fill
either all the cells of the tissue (Irish moss), often in combination
with starch (Sphwraococcus lichenoides), or are confined to indi-
vidual cells, which are often distinguished by their shape and
size (cinnamon, elm bark, salep *), or, finally, are given off by
intercullular receptacles of secretions (Cycadew). Even the gum
which exudes upon wounded places, for the purpose of closing
the vessels—and which, therefore, serves a physiological pur-
pose—is not formed through a conversion of the membrane into
gum, but is secreted by the same in the form of drops * (see
above).

The designation, bassorin, has injudiciously ° been trans-
ferred to a part of the mucilages. Solutions of plant mucilage
are not only precipitated by basic acetate (subacetate) of lead,
but also by the neutral acetate (sugar of lead). Plant mucilage
from its various sources presents, however, in its behavior to
water, all gradations, from complete solubility to mere swelling,
accompanied by but extremely slight solution. For the pur-
pose of microscopical examination of tissues containing mucilage,

those liquids are therefore useful which act to a less extent upon

1 Fliickiger, ‘‘ Pharmakognosie,” 1883, 934.

? Flickiger, Schweizerische Wochenschrift fiir Pharmacie, 1873, p.
435; Fliickiger and Hanbury, * Pharmacographia,” 1879, p. 109; Radlko-
fer, “‘Sapindaceen-Gattung Serjania.” Munich, 1875, 100.

8 Berg’s ** Atlas,” Plate XXIII, Fig. 57.

4 Frank, ‘‘ Berichte d. deutsch. botan. Ges.,” IT. (1884), 322.

5 Injudicious in so far as under the name of Bassora Gum different
and not accurately known varieties of mucilage, similar to tragacanth,
have been grouped together; the expression bassorin is, therefore, not
capable of precise definition; and should be abandoned. ©

168 PLANT ANATOMY,

the latter, such as concentrated glycerin, alcohol, and fatty or
volatile oils. The mucilage then appears contracted to a mass
which no longer completely fills the cell, as, for instance, in
Bulbus Scille. Occasionally the masses of mucilage show a
stratification, which is rendered more prominent upon the addi-
tion of alcohol, as may be seen in Radix Althee. In such cases
it is to be assumed that a gradual, even though but partial, con-
version of the cell-wall into mucilage has taken place, especially
when the mucilage, as in salep, is colored blue by iodine and
sulphuric acid, or is even soluble in ammoniacal oxide of copper,
like pure cellulose. The latter is the case with the terminal

Fig. 79.—Transverse section through tragacanth, in which may still be seen the rem-
nants of the cell-membrane which has been converted into gum, and also isolated starch
granules.

member of the cellulose series, lichenin (see pages 123 and
170), which is related to the varieties of mucilage. That the
cell-walls are capable of passing entirely into mucilage has al-
ready been noted (page 164). In the formation of tragacanth,
not only the cell-membranes participate, but also the starch

granules which were previously deposited in the tissue, and
_ which to a slight extent are still retained as such in the fraga-
canth (Fig. 79). Other constituents of the cells also occasionally
take part in the formation of gum and mucilage.

GUM—MUCILAGES. 169

Considerable differences, which have, however, been demon-
strated as yet only by a few examples,’ are also presented by the
various kinds of mucilage from an optical point of view, some of
them rotating the plane of polarized light to the left, in the
same manner as ordinary gum, while other mucilages rotate it
to the right.

With regard to their chemical character, gums and mucilages
are but little known, and are with difficulty freed from inor-
ganic constituents and nitrogenous substances.” Gum arabic
appears to be composed of the calcium, potassium, and magne-
sium salts of arabic acid. Ifthe formula Ca(C,,H,,0,,),+3H,O is
assigned to gum arabic, it must contain 13.3 per cent of water
and 1.9 per cent of calcium; these numbers nearly correspond to
the actual proportions.

It is also scarcely possible to characterize gums micro-
chemically. They mostly swell in water (not the gum produced
by wounds) and are not rendered blue by iodine, or by iodine with
sulphuric acid. The plant mucilages are colored yellow or blue
by iodine, and blue or violet-brownish by iodine with sulphuric
acid. Both are insoluble in ammoniacal oxide of copper. The
amyloid of Schleiden, which should also be considered here, is
colored blue by iodine, yellow by iodine water, and is soluble in
boiling water. In some cases, for instance, in Cydonia and
salep, the mucilage retains the capability of being colored from
reddish to blue by iodine, after treatment with sulphuric acid,
and in this respect stands one step nearer to cellulose. It does
not follow from this, however, that mucilages always originate
from cellulose. In Semen Cydonia, Sem. Lint, Sem. Sinapis albe,
and also in the seeds of Plantago Psyllium, before they ripen, and

1 Wiggers-Husemann’s Jahresbericht, 1869, 154, top,

2 Tragacanth affords three per cent of ash. In mucilage of Irish
moss, even after repeated purification, there are still contained sixteen
per cent of inorganic substances and 0.88 per cent of nitrogen (= six
per cent of albumen): Wiggers-Husemann’s Jahresbericht der Pharm., ©
1868, 88. With reference to many other varieties of mucilage compare
Frank, Pringsheim’s Jahrb. fiir wissenschaftliche Botanik., V. (1866),
161.

170 PLANT ANATOMY.

before mucilage makes its appearance in the respective cells,
there are found starch granuies, which afterward disappear—a
circumstance which very probably stands in definite relation to
the formation of mucilage.

We shall also meet with a transformation and solution of
cellulose later on, when we come to the consideration the origin
of cell fusions. In the formation of vessels, sieve-tubes and
lacticiferous ducts, namely, a resorption (solution) of the trans-
verse walls consisting of cellular substance takes place. Besides
this process of solution, there also occurs, during the formation
of the lysigenic balsam ducts (see also under “ Receptacles for
Secretions ”), a transformation of the cellulose into secretions.

Thus, for example, the membrane may become converted into
resin ’ (see Index references to the latter). Such a transforma-
tion appears also to be the case in the formation of the resin of
Polyporus officinalis Fries.

With the varieties of gum and mucilages are also connected
the pectic substances, the knowledge of which is still very
incomplete.

In close connection with the bodies which have here been
treated of stands lichenin® (lichen-starch, amylo-cellulose), a
carbo-hydrate deposited in Cetraria islandica, in Usnea, Parme-
lia and Cladonia. AccordingtoBerg, the lichenin of the first-
named lichen, the /celand moss, is a mixture of two substances,
one of which is colored blue by iodine and is dissolved by
chloride of zinc and ammoniacal oxide of copper.?

} The formation of resin can take place: 1. As a true secretion through
proper organs of secretion. 2. By the liquefaction of the outer walls of
certain cells, 3. By a metamorphosis of the entire cell-wall and con-
tents of the cell (lysigenic and pathologic receptacles for resin). 4. By
a transformation of certain constituent bodies, increasing the resin
formed according to 2and3. (Compare Hanausek, ‘‘ Jahresbericht der

_Handelsschule in Krems,” 1880). See also subsequent references under
«« Receptacles for Secretions.” :

? From lichen.

*Jahresbericht der Pharmacie, 1873, 21. Compare Flickiger,
*‘ Pharmakognosie,” second edition, 273, and ‘‘ Ueber Stérke und Cellu-

lose,” in Archiv der Pharm., 196 (1871), 27.

FORMS OF CELLS. Lit

II. Forms of Cells.

Notwithstanding the unlimited variety of forms of developed
plant cells, they, nevertheless, show very similar outlines in a
young condition. In every case where cells can develop unob-
structed, they assume a spherical shape (Fig. 80), which is the
fundamental form of all cells (Saccharomyces, spore cells, pollen,
heads of glands, the cells of soft tissue, for instance, of the
medulla).

The subsequent distinctions in form are produced either
through unequal surface growth, or growth in thickness or
length of the cell, or through the pressure of contiguous cells.

Fia. 80. Fie. 81.

Fic. 80.—Spheroidal cells. Isodiametric parenchyma.
Fig. 81.—Parenchymatous tissue from the pith of the Hider.

If the surface growth does not proceed uniformly, there are
formed a great variety of elliptical, tabular or hemispherical,
sinuate, star-shaped (Fig. 152) or plaited cells.

If the growth in thickness is unequal, all the forms are devel-
oped which have previously been mentioned when considering
the growth in thickness of the membrane (page 149): the
pitted, scalariform, annular and spirally thickened cells (vessels,
Figs. 66 and 68), the stone-cells and bast-cells (sclereids,”
Figs. 70, 110, 115, 116, 117). If the growth in thickness is
confined to the corners, collenchyma is formed (Fig. 109 4).

If the growth in length is unequal, that is, chiefly confined
to two opposite sides, there are produced elongated forms of

1 See page 156, foot note.

172 PLANT ANATOMY.

cells, bast and wood-cells (Fig. 136), sieve-tubes (Figs. 147 and
148), furthermore sickle-shaped, and S- or U-shaped cells.

Mutual pressure also causes manifold differences of form..
Thus from roundish cells (Fig. 80), polyhedral (Fig. 81), and
more or less rectilineal forms are produced. It is only in very
delicate and soft a. ea part of fruits, medulla, leaf-
cells), and in those pl ere the membrane reaches the
outer air (outer wall of the epidermis and bordering membranes
of the intercellular spaces), that the spherical outlines remain
preserved; in firm and hard tissues (wood, Fig. 180, bast
groups, Figs. 111, 112, 113) all cells are seen to possess, on a
transverse section, more or less flattened forms with a rectilineal ©
border.

There are ordinarily distinguished, according to the scheme
first proposed by Link:

1. Parenchyma,’ thin-walled, mostly roundish-polyhedral
and isodiametric cells: cells of the fundamental tissue, of the:
medulla, of the fleshy part of fruits, merenchyma‘® of the leaves,
and when extended in a palisade-like manner, palisade-cells *
(Figs. 80, 81, 85, 108, 127, 128, 129).

2. Prosenchyma,* consisting of thick-walled, more or less.
elongated, spindle-shaped cells, with the ends wedged into each
other: wood-cells, bast-cells (Figs. 139, 136, 110, 111).

Striking and convenient as the discrimination between pros-
enchyma and parenchyma appears, yet it does not admit of
sharp application.

Fungi and lichens are composed of thread-shaped cells,
hyphe,* which continue to grow at the ends and mostly divide
and branch by transverse walls (Fig. 82). They are not only

‘“Grundlehren der Anatomie und Physiologie der Pflanzen.” Gdt-
tingen, 1807.

? Ilapad beside, thereon, and eyyvua that which is poured in, the
cells considered as standing upon each other.

* An expression introduced by Meyen (‘‘ Phytonomie,” Berlin, 1830).

4 Palus, i, masc., a stake,

* IIpos towards, between, and eyyvuc (eee above), the cells consid--
ered as inserted between each other.

5 ‘Y'o7 the tissue.

FORMS OF CELLS. 173

densely interwoven, but also cling together with great tenacity,
unless they inclose hollow spaces. The tissue (pseudo-paren-
chyma)’ of sclerotiums, for instance of Secale cornutum, consists
of remarkably short hyphae, so that upon thin sections it has the
appearance of parenchyma. It is only upon a longitudinal
section, softened by a dilute solution of chromic acid (see Micro-
chemical Reagents), that the threadlike nature of these hyphe is
likewise clearly brought to view. Notwithstanding their slight
length, they are very firmly connected with each other. *
Through subsequent resorption of the transverse walls of a
more or less elongated row of cells (cell-fusions), long tubes may

y “aly
(} OF \V/ Bi </
RNA AYA
) NY Ay V
V ‘J y
\}

Wa i \ oh
Ai} 4
N Me ne

f Ay

Vii vA any

Fie, 82.—Hyphe from Fungus Laricis. a, Hyphe; b, longitudinal section through Sg

the hollow spaces (pores), (Berg.)

be produced which possess the most varied physiological func-
tions, sometimes as vessels (in the woody structure), sometimes
as sieve-tubes (in the phloém), and sometimes as lacticiferous
ducts (in the fundamental tissue); all being thus designed ir
conducting purposes.

Tf the cells lie densely upon each other, they are sfirialy cemented
together by the intercellular substance* (middle lamella) (Fig.
64 x); if, on the contrary, they are not in contact with each

1 Wevsos illusion,
* Inter, between, and cellula, cell,

174 PLANT ANATOMY.

- other on all sides, there appear between them (especially at the
corners) intercellular spaces, which are mostly filled with air
(Figs. 127, 129, 151, 152, 155).

III. Cellular Tissue.

With the exception of the one-celled plants and plant organs
(Saccharomyces, trichomes, Lycopodium) and some fungi and
alge represented only by simple cellular threads, all plants con-
sist of cellular tissue,’ that is, of cells (aggregations of cells) in
every form of arrangement. All the cells of such tissue are
never completely uniform, but the individual parts become dis-
tinguished at an early period in a more or less pronounced de-
gree. While the cell of the alge performs conjointly all the
functions which are required of the plant, in the higher plants
a division of the work takes place in such a manner that some
forms of tissue undertake one task and others another.

Through this division of work there are then produced in the
body of the plant anatomico-physiological systems of tissue.
Such a system of tissue is, therefore, a union of cells, complete
within itself, and connected by their entire physiological deport-
ment.

This differentiation of the body of the plant, however, is not
noticeable until the later stages of development. At the places
of development, the growing points (apex of the stem, tip of the
_ root), such a difference in the tissues is not yet perceptible. The

tissue here consists rather of uniform, more or less isodiametric,
thin-walled cells containing protoplasm, and in a state of most
active division. Such a tissue is termed developing tissue, or
meristem." As such it stands in opposition toall the remaining
tissues, which are also collectively comprehended under the name
of permanent tissue. The cambium (Fig. 138) is also such a
developing tissue.
_ 1 The inner juicy tissue of maturing fruits (tamarinas, juniper berries,

oranges, stone-fruits) is resolvable into individual cells, but these are al-

ways held together by their surroundings. .
2 Mepifoo I divide.

SYSTEMS OF TISSUE. 175

While, namely, the cells of the developing tissue have not yet
assumed a definite, permanent form, but become altered by di-
vision and mutual dislocation, the cells of the permanent tissue
are conclusively defined with regard to their form, or are sub-
sequently but little changed. By far the most drugs consist of
permanent tissue.

In the angiosperms there are to be distinguished at the grow-
ing point three meristem zones: the dermatogen’ trom which is
formed the epidermis; the periblem,* from which is produced
the bark; and the plerom,* from which are formed the vascular
bundles and the medulla. In angiospermous roots there is, in
addition, the calyptrogen,* deca represents the developing tis-
sue of the root-cap.

IV. Systems of Tissue.

If, in grouping the forms of tissue as systems of tissue, the
latter are viewed not alone froma purely anatomico-topographi-
cal standpoint, but if, at the same time the question arises, im
which manner the various tissues are of equal value physio-
logically, they may be divided in the following manner:

1. The epidermal system. Function: the protection of the
organs from without. )

2. The mechanical system. Function: to give stability to the
plant.

3. The assimilating system. Function: assimilation of the
carbon.

4. The conducting system. Function: conduction, especially
the conduction of water and nutritive salts from the soil, and to
conduct away the products of assimilation.

5. The storing system. Function: the storage of reserve nu-
tritive substances and of water.

6. The aérating system. Function: the aération of the or-
gans.

1 Aépua skin, and yervaéoI produce.
2 TTep¢ around, PAyua covering.

8 TAnp@ ya that which fills.

4 Kalvarpa a cap.

176 PLANT ANATOMY.

v7. The system of receptacles for secretions. ~ Function: to re-
ceive the products of secretion of the plant.’

1. The Epidermal System.

While such plants and parts of plants which consist of a single
cell, or of but a single layer, do not possess an epidermis,’ a
more or less distinct development of epidermal layers appears
already in the thallophytes and cormophytes, which have the
thickness of but a few layers of cells. The cells in these layers
become for the most part smaller and more thick-walled toward
the exterior, and often colored, even though the formation of a
true epidermis is not yet effected (Secale cornutum, Fucus vesicu-
losus, Cetraria, Usnea, Spherococcus, the small stems of
mosses). :

It is only in the higher plants that, even in the youngest
stages, a true epidermis is formed, and this is the first of any
system of tissue which is sharply defined, morphologically, from
all the others.

It consists in most cases of a row of tabular or plate-like cells,
laterally united without intervening spaces, which in the organs
of dicotyledons are, as a rule, of quadratic form, in the elon-
gated leaves and stems of monocotyledons, however, are mostly
extended in the direction of the axis of the organ (distinct upon
surface sections). On many roots the epidermis is also sharply
defined from the other tissue by a different color and strongly
sinuous outer walls. Such an epidermis, which we meet with,

for example, on the rootlets of Helleborus niger and Veratrum

' Sachs classifies the tissues as follows: epidermal tissue, fascicular
tissue, and fundamental tissue. In the above division we adhere to the
classification of Haberlandt (‘* Physiologische Pflanzenanatomie,” Leip-
zig, 1884), which is based upon Schwendener's principles. Nevertheless
the expression ‘‘ fundamental tissue ” (filling tissue) may often be permit-
ted in the following pages on account of its brevity, notwithstanding
the fact that the fundamental tissue comprises the most varied forms
of tissue.

2’ End upon, and éépue skin,

EPIDERMIS. 177

album, was formerly termed epiblema (Figs. 84, 119, 120, 121,
122). :

Occasionally, however, the epidermis consists of several layers,
as, for example, in Macis (Fig. 85), and in many leaves (/icws),
This multiple epidermis, which is also termed hypoderma,* con-
sists, in the case of delicate organs, mostly of uniform, thin-

Fie. 84,

Fig. 83,—Longitudinal section through the outermost layer of Vanilla; a, epidermal
cells, containing crystals of vanillin; b, cells with spiral fibres.

Fic. 84.—Epidermal cells of a root (epiblema) on a transverse section; the dark cells
represent the epiblema,.

walled or slightly thickened cells (leaves of the Piperacee,
Chavica and Peperomia, of the Begoniacex, and of species of

1*Yzo0 under, and 6égua skin. We use the word only for the true
multiple epidermis, not for the layers (collenchyma, bast-fibres) which
impart strength to the single-rowed epidermis,
12

178 PLANT ANATOMY.

Ficus); the outermost row, however, is also here, for the most
part, somewhat differently formed.
In most fruits and seeds only the outermost row of cells be-

SOOO Sago:
EEC COO OSB OPS:

COCCL OCS, ogee oo =
—— DO OCU OODO GRE SECU DOC
Fia. 85.—Transverse section through Macis; c, epidermis; 0, oil-cells; v, fibro-vascular
bundles.

longs to the epidermis, and in this case, to speak offa multiple
epidermis, is incorrect.

Fie. 86.—Rind of the fruit of Colocynth (in the commercial fruit usually removed by
paring); a, epidermis; b, parenchyma; ¢, sclerenchymatous layer, which mostly forms
the outer surface of the pared fruit,

In the epidermis consisting ofa single layer, the cell-walls are
mostly firmer than in the tissues lying beneath it, and more

EPIDERMIS. 179

strongly thickened on the outer side than on the inner’ (Figs.
83, 85, 63, 109, 129, 155). Occasionally the outer wall is even
of quite remarkable thickness (Caryophylli, Macis, Fig. 160).
An example where this is not the case is presented by the epi-

Fic. 87.—Transverse section through Semen Paradisi (Grains of Paradise); f, epider-
mis; gh, testa of the seed.
dermis of Hyoscyamus seeds. Here the outer wall of the epidermal
cells of the testa is formed exclusively of the delicate cuticle.
The outer wall of the epidermis cells is always covered by the

Fic. 88.—Semen Hyoscyami, .A, Transverse section; a, cuticle; b, epidermis; c, testa ;
d, albumen cells; 0, fatty oil. B. Tangential section through the cells of the epidermis,

cuticle (see page 161 and Figs. 155, 160, 63, 128, 161), usually a
delicate * film, insoluble in sulphuric acid, impenetrable by water

‘The delicate epidermal cells of the seeds of Cydonia me Linum,
which produce mucilage, form an exception (Berg’s ‘‘ Atlas,” xlvi., 122,
128).

*Compare Tschirch, ‘‘ Ueber einige Beziehungen des anatomischen
Baues der Assimilationsorgane zu Klima und Standort,” Linnea, ix.
(1881), 139,

180 PLANT ANATOMY.

and aqueous vapor, and which for the previously described reasons
is primarily adapted to fulfil the function of the epidermis
namely, protection against too strong evaporation.

The cuticle, which is often directly visible upon transverse
sections (Figs. 160,161, 155), or is easily rendered visible by
dilute chromic acid, sulphuric acid, iodine, or potassa, covers all
the organs of the plant which are exposed to the air, and, in the
case of favorable objects, can often be removed asa coherent
film.

If the outer wall is thin, as for instance in the leaves of the
indigenous foliage trees and all the officinal leaves, then the
cuticle is in direct connection with the cellulose layer (Fig. 155);
if, however, the outer wall is very strongly thickened, the inter-
vening layers of the outer membrane of the epidermal cells are
for the most part cuticularized (cuticular layers, Fig. 161
cs, 63), that is, they have become more similar to the cuticle

AI ISe

Fie, 89.—Transverse section through Fig. 88 A, more highly magnified.

itself by the deposition of cutin. These cuticular layers often
project in a somewhat cone-like form towards the interior (Figs.
161, 63).

The outer wall is frequently somewhat arched in an outward
direction (Fig. 155). These outward arches may become defi-
nitely shaped processes, whereby the exterior surface acquires
a pitted appearance. The same appearance may, however, also
be produced by the prominent development of thick lateral walls
(testa of the seed of Hyoscyamus, Fig. 88, and the leaves of
Gentiana cruciata). In the latter case, the outer wall is always
very thin, and indented (Figs, 88 and 89). Occasionally the sur-
face delineation is also produced by the projection of sharply
circumscribed groups of epidermal cells.

The outer wall (like the thick lateral walls, Figs. 88, 89)
shows, as a rule, a distinct stratification.

EPIDERMIS. 181

While the outer walls of the epidermal cells, as a rule, are
thick, the lateral walls are mostly thin. The Figs. 63, 109, 129
and 155, therefore, reproduce the type of epidermal cells.

Upon surface sections, the lateral walls appear in many cases
sinuous, as, for instance, in all leaves of the Graminee (Figs. 88
B, 90, 154, 15%, 158), so that the individual epidermal cells
which are provided with many protuberances fit into each other
in a tooth-like manner (many corolla leaves, and the epidermis
of Semen Stramonii, Fig. 90).

As a rule, the contents of the epidermal cells are colorless,

Fre. 90. Fie. 91.

Fie. 90.—Tangential section through the epidermis of Semen Stramonii.
Fie. 91.—Transverse section through Chinese galls; a, epidermis, the cells of which
frequently grow out in the form of simple hairs; b, lacticiferous cells,

without chlorophyll; occasionally, however, coloring matters
appear in them, dissolved in the cell-sap. Colored epidermal
cells produce, for example, the red color of many stems (buck-
wheat, Ricinus) and other organs (the apple). In the red
potatoes, the coloring matter is contained in cells lying beneath
the cork.

It has already been mentioned (page 180) that the epidermal
cells often project outward. If these protuberances become —

182 PLANT ANATOMY.

larger, hair formations‘ or trichomes* are produced. These
are found in the most simple, unicellular form, in Stipites Dul-
camare, on Herba Lobelia and the Chinese galls (Fig. 91), and
upon cotton-seed (Figs. 92, 114); the root-hairs (Rad. Sarsapa-
rille, Fig. 126) also belong here.

The hairs are occasionally very long (flower buds of Althea
rosea).* Long, sharp, silicified hairs are termed prickles (sting-
ing hairs of the nettle, Fig. 93 a). Firmer, short, non-secreting
trichomes are called bristles. Of the latter kind are, among
others, the hairs of Nua vomica (Fig. 93 b), of anise (Fig. 94).

The hair formations do not, however, always remain simply
hair-shaped. Many of them assume other forms (that of a
star, shield, or head, Fig. 95), throw out branches and become

Fic. 92.—Hairs of cotton.

multicellular.* Flatly- expanded multicellular hairs (chaffy

hairs), such as are of frequent occurrence in ferns (for instance,

in Aspidium Filix mas) form the so-called Pengawar Djambi.
If the terminal cell of a multicellular trichome becomes ex-

‘Compare Weiss, ‘‘ Die Pflanzenhaare,” in Nos. iv. and v. of the ‘“‘ Bo-
tanische Untersuchungen,” of Karsten, 1867. Rauter, loc. cit., 31. Mar-
tinet, Annal. d, sciences natur., xiv, (1872), 91-282. Paschkis, ‘‘ Pharma-
cognostische Beitrége.” Zeitschr. d. allg. oesterreich. Apothekervereins,
1880, Nos. xxvii. and xxviii. Hanstein, Bot. Zeit., 1868, 725. De Bary,
“* Anatomie,” p. 61, where the literature is given to the year 1877.

* Opt?, rpryos hair.

*Sachs, “‘ Lehrbuch der Botanik ” (iv.), 101.

*The unicellular climbing hairs of the hop, reposing upon a multicel-
lular cushion, are not of this kind,

HAIRS. 183

panded in a head-like form, the formation of daughter cells
frequently occurs, with the simultaneous secretion (for instance,

Fie. 93 b,

Fic. 93 a,—Stinging hair of the nettle, with a small head. The protoplasm of the hair
circulating in currents (the direction of the currents indicated by arrows).
Fic. 93 6,—Hairs of the epidermis of Nwa vomica (Berg).

in the Labiate) of a balsam or volatile oil, which is often
accompanied by the formation of mucilage’ (Figs. 96, 129,
1The morphology and the nature of the development of secreting

184 PLANT ANATOMY.

154). Such hairs are termed glandular hairs or colleters.:. To
this class belong also the glands of Dictamnus.’

In the species of Cistus of the Mediterranean flora, these hair
formations secreting resin* are so numerous and so product-
ive that, for example, the product of Cistus ladaniferus has

j\

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os OV 7)
xD — 7}, iP
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Fic. 94,—4, Transverse section through Fructus Anisi, e, epidermis, clothed with
hairs; ec, commissural surface; 0, oil spaces; t, coating of the fruit; t’ (lower ¢), seed
_ Coat; v, fibro-vascular bundles (ribs, costee); a, albumen of the seed, the parenchyma of
which is indicated by but a few cells, B, Hairs, more highly magnified.

been collected in the islands of Candia and Cyprus from ancient
times and employed for fumigating purposes. This ladanum

trichomes has been described by Hanstein (Bot. Zeit., 1866, 747); com-
pare also De Bary, “ Anatomie.”
' KodAnros glued together, :
*Meyen, “ Secretionsorgane der Pflanzen,” Berlin, 1887; Plate i.,
Figs. 28 and 29. De Bary, « Anatomie,” p. 73.
_* De Bary, loc. cit., p. 99, Fig. 36. :

HAIRS. 185

resin is probably the only example of such a drug originating
from trichomes.’

In kamala and lupulin, which, according to the nature of
their development, should also be classed with the trichomes,’
the formation of resin predominates; and in kamaia oil is en-
tirely wanting.

Glands which secrete wax are found on the leaves of Glodu-

Fic, 95,— Flores Verbasci. A, Band-like, soft, club-shaped hairs of the threefshorter
stamens, covered with exceedingly fine, spirally-arranged, projecting points. PB. Stel-
late hairs from the base of the folds of the corolla,

laria Alypum UL. and other species,’ those secreting nectar in
Melampyrum.*

™Compare Thiselton Dyer, Pharm. Jour., xv. (1884), 301.

* With regard to the development of the lupulin glands, compare
Holzner, ‘“‘ Entwickelung der Trichome der Hopfendolden.” Bayer.
Bierbrauer, 1877, No. 19. Rauter, ‘‘ Denkschr. d. Wiener Akad.,” 1870,
p. 31, Luerssen, ‘‘ Medizin.-Pharmaceutische Botanik,” ii., p. 527. —
Harz, ‘‘Samenkunde,” ii., 896.

* Heckel et Schlagdenhauffen, Comptes rendus, 95 (1882), 91.

*Rathay, ‘‘ Ueber nectarabsondernde Trichome einiger?7Melampyrum-
Arten.” Wiener Akademie, 1880,

186 PLANT ANATOMY,

The form and size of the hairs, as also the relative thickness
of the wall compared with the lumen or cavity, afford in some
cases good points of discrimination for the recognition of adul-
terations in foods and alimentary substances. Thus Wittmack :
distinguishes wheat and rye flour by the fragmentary hairs of
the so-called coma, which always occur therein in small amount;
and upon the form of the hairs Bell? bases the distinction of
tea, elder, willow and black-thorn leaves (Prunus spinosa Lin.).
. The physiological function of the hairs of organs of assimila-
tion (leaves), which when old mostly contain air, is to diminish
the extent of transpiration. The hairs of seeds, which are often

Fic. 96.—Oil glands of the Labiate, e. g.,0f Rosmarinus. A, Longitudinal section of
valarge gland. a, stem-cell; b, eight delicate-walled daughter cells which produce the
volatile oil, by the escape of which the cuticle of the parent cell, d, becomes expanded:
Jf, epidermis of the leaf upon which the gland is formed; g, palisade-cells; e, a small
gland. B, Transverse section of Fig. 4. Compare also De Bary, “‘ Anatomie,” Fig. 39.

feather-like, are means of distribution; the firm hairs of climb-
ing plants are organs which serve to fasten them.

Some prickles, for example in Rubus, are, like the hairs,
also of epidermal origin, and are thus trichomes. Jn the forma-
tion of most of the true prickles or outgrowths, however, the
tissues beneath the epidermis, and even the vascular bundles,
also participate (Rosa, Smilax #),

'* Anleitung zur Erkennung organischer und anorganischer Beimen-
gungen im Roggen- und Weizenmehl.” Leipzig, 1884.

* Bell, “ Die Analyse der Nahrungsmittel,” i, (Berlin, 1882), 36.

"De Bary, “ Anatomie,” p. 61, where the literature is given.

PERIDERM. 187

Finally, the so-called inner hairs may also be mentioned,
which occasionally penetrate into the air-cavities (star-shaped
hairs of the Nymphe, glandular hairs of Aspidium Filix mas,
Fig. 162).

The epidermis no longer suffices for older plant organs of
several years’ growth, since it is a much too delicate tissue (for
instance, for the stems and branches), and, as permanent tissue,
is not capable of keeping pace with the growth in thickness,
In these organs, therefore, there is formed beneath the epi-
dermis, and mostly independent thereof, another tissue, the
periderm.' The latter consists of a permanent tissue, the
cork, and a formative tissue, the phellogen,’ or cork-cam-
bium.

The phellogen, by a tangential division of its cells, forms the

Fic. 97.—Cells from the phelloderm of the bark of Canella alba (J. Moeller).

cork-cells; but it is also capable of contributing to the increase
of the bark parenchyma by the formation of parenchymatous
elements. Sanio* terms the aggregate of cells which are thus
produced, and occasionally thickened on one side (Figs. 97, 149)
phelloderm * or cork layer of the bark.

1 epi around, and dépuc skin.

? PedAos cork, and yevvaw I produce.

3«*Bau und Entwickelung des Korkes,” Pringsheim’s Jahrb., ii.,
(1860), 47. H. v. Mohl, ‘‘ Entwickelung des Korkes und der Borke der
baumartigen Dicotylen” (1836). ‘‘ Verm. Schrift.,” Tibingen, 1845,
225. F. von Hohnel, ‘‘ Ueber Kork und verkorkte Gewebe tiberhaupt.”
Sitzungsberichte der Wiener Akademie, 76 (1877). Hanstein, “ Bau und —
Entwickelung der Baumrinde.” Berlin, 1853. Hofmeister, ‘ Handbuch
der phys. Botanik,” i., 252. De Bary, ‘‘ Anatomie.”

* PedAos cork, and dépua skin.

188 : PLANT ANATOMY.

The location of the phellogen is.not exclusively confined to
the region directly under or within the epidermis, but may also
develop itself in the form of bands and stripes in the funda-
_ mental tissue of many barks, or even in the phloém layer. Out-

side of such layers of internal cork (Fig. 98), the most various
tissues of the bark may therefore be represented, according to
the depth at which they lie (phloém elements, Fig. 100, bast-
cells, Fig. 98, stone-cells, Fig. 101, and, indeed, even resin-

At il |
mea zealllili

Fic. 98,—Transverse section through the bork of Cinchona Calisaya. 8, outermost
cork layer; r, cork-bands in the inner tissue; J, bast-cells (Berg.).

canals and oil-spaces, Figs. 99, 100); they become pushed out
of the course of circulation of the sap by these lamelle of inter-
nal cork, and are either thrown off as scales (in a very handsome
manner in the Platanus and Eucalyptus), or they still remain
for a long time united with the stem, and appear severed and
_ torn only in consequence of the growth in thickness (207 k,
thytidoma') ee
| Puris, Avrisos fold, wrinkle, and Saud I build.

BORK. 189

The bork of our foliage trees, for example of the oak, con-
tains, in addition to cork-cells, all the elements of the outer
bark (parenchyma, stone-cells).

Fic. 99.—Transverse section through an older internode of Juniperus communis L.;
p, outer layer of the primary bark, with a resin canal; &, interior cork (J. Moeller).

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Fic. 100.—Bork of Cortex Sassafras radicis; aa, decayed surface; ss, cork-bands ;
bb, phloém; 0, oil cells; r, medullary rays.

190 PLANT ANATOMY.

Whether barks experience the formation of bork, or are pro-
vided with a simple covering of cork, appears to depend upon

A

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Hf

Aheet arte

Fie. 101.—4, Cork of the cork-oak; aa, cork-cells; bb, stone-cells; B, more highly

magnified cork-cells,
Ee ane a f
se ae " : oy
— - —
~ : ae ee. —
= =" oe 4 x
3 Lag. 2
o 4
~ OR, & % a ¢
Lee 7! POT, 20 0, 0G 7 aly
f Gs" fo, Ooo *,
\
vis
R

aN

Fic. 102.—Cortical layer of Radix Calumbe; s, cork; m, fundamental tissue, with dis-

persed stone-cells which inclose crystals of oxalate; R, 7, More highly magnified
crystals; H, twin crystal.

the peculiarities of the species. In the Cinchonas, for exam-
ple, bork is sometimes met with, and sometimes not. Roots are

BORK—CORK. 191

also capable of forming bork, which may be very plainly observed,
for instance, in Radix Sassafras.

If the periderm layers which are formed in the interior of the
bark occupy only a part of the circumference, scale-bork is pro-
duced (Robinia, Platanus, Cinchona, Pinus silvestris, Quer-
cus); if, however, the secondary periderm layers form parallel,

Fia. 103.—Transverse section of Rhizoma Curcume; s, cork.

closed rings, which embrace the entire circumference, hollow-
cylindrical sections of bark are converted into bork, and ringed-
bork is produced ( Vitis, Clematis).

Fic. 104.—Cortex Cascarille. Cork layer and primary bark, with crystals of calcium
oxalate and coloring matter.

Since the cork-cells are produced by a tangential division of
mostly tangentially extended cells, they are flatly tabular and
parallelopipedal. They are in unbroken connection with each
other and contain air, never solid matter. As the divisions in
the phellogen take place very regularly, the transverse walls of
the cork-cells frequently traverse the entire cork tissue In one
and the same line (Figs. 102, 103, 108). 5

192 PLANT ANATOMY.

The ordinary cork, of Quercus Suber, corresponds in its form
to the above type.' Deviations from this fundamental form

Fie. 105.—Cortex Guaiaci. a, thickened cork-cells; a’, phellogen layer; b, primary
bark; c, sclerenchymatous layer.

depend upon a more undulating—though on a transverse sec-
tion, generally radial—course of the transverse walls, or upon a

o/

rae

Fie. 106.—Radix Pyrethri romani. a, thickened cork-cells; b, oil-spaces; *, xylem
rays (wood-bundles),

thickening of one or all sides (as seen in Figs. 102, 103, 108)
of the ordinarily thin walls (Figs. 97, 104, 105).

* Yet in cork (bottle-cork), numerous stone-cells occur (Fig. 101).

CORK. 193

The outermost layer is often in process of decay, as for in-
stance in the potato (Fig. 108). While this perishes, new cork

8,_cork; s’, cork-cambium or phellogen; m, layer of the primary bark; m’, containing
crystals.

Fi. 108.—Transverse section through the outermost layer of a potato. s, starch
granules; er, protein crystalloid; pl, protoplasm (Tschirch).

is continually formed from the interior (Cortex Quassie jamai-
censis, Fig. 107).
13

194 PLANT ANATOMY.

Even on leaves a local formation of cork occasionally occurs’
(Eucalyptus, Fig. 128, k).

The physiological function of cork is to protect the tissues
lying beneath it from too great evaporation and from mechani-
cal injury. The former function is presented in a very striking
manner by the potato, in which the cork layers, from five to ten
in number, cause the succulence of the entire inner tissue to be
retained for a long time undiminished.

On wounds, the so-called wound cork is frequently formed
(Fig. 109, a). This produces, in the same manner as the
thylle* and the gum which incloses the vessels, a separation of
the inner, uninjured tissues from the wounds.*

2. The Mechanical System.

Leaving out of consideration the lower plants, we find that
all the higher plants which concern us are provided with
peculiarly formed and characteristically arranged cells, the ex-
clusive function of which, as Schwendener‘ has shown, is to
impart to the plant the necessary solidity. While in young and
still growing organs the collenchyma® (Fig. 109 4, 129 coll)
represents the mechanical system, in older and matured organs
this function is assumed by the bast-cells, or stereids° (Figs.

110, 111).

The bast-cells, or the specifically mechanical elements of the
matured plant, form very elongated cells, which are pointed at:

‘Bachmann, ‘‘ Korkwucherungen auf den Blattern,” Pringsheim’s
Jahrb., xii., 1880,

* Sachs, ‘Lehrbuch der Botanik,” iv. (1874), pp. 27, 782. Weiss,
** Anatomie,” 21, :

* With regard to the lenticels, see the chapter on the Aérating System:

*** Das mechanische Prinzip im anatomischen Baue der Monocotylen.
Leipzig, 1874, z

*Ambronn, “ Ueber die Entwickelungsgeschichte und die mechani-
schen Eigenschaften des Collenchyms.” Pringsheim’s Jahrb., xii. E.
Giltay, ‘‘ Het Collenchym,” Inaugural dissertation, Leyden, 1882.

* The tissue of bast-cells may be termed stereom. (The pleurenchyma
of Meyen.) ‘

BAST, 195

both ends, provided with oblique pores (Fig. 110), often thick-

Fic. 109 a,—Transverse section through a fruit of the Vanilla, which, before separa-
tion from the plant, had become injured at the point b-b by the puncture of an insect,
and the wound closed by wound-cork, a-a; g, vascular bundles (Tschirch), Compare
Pharm. Zeit., 1884, No, 22.

Fic. 109 b.—Collenchyma. ep, epidermis; con, collenchyma cells; ch, chlorophyll
granules (Tschirch),

196 PLANT ANATOMY.

ened to the extent of the disappearance of the lumen or vavity,
and contain air. They possess a supporting capacity which is
almost equal to that of wrought-iron, and a tenfold greater
ductility than the latter, and are therefore in themselves ad-
mirably adapted for the mechanical purposes of the plant.

“Fie. 110. Fie. 111.
Fie. 110.—Typical bundle of bast-cells, at a@ in transverse section, at bin longitudinal
section; c, section of a bast-cell, showing the striping of the membrane and oblique
pits (Tschirch),

Fie. 111.—Bast-cells of Corchorus olitorius (Jute), with a lumen or central cavity of
varying width, at the top in transverse, and below in longitudinal section (Tschirch).

Moreover, they are always united to- form structures which are
in nowise inferior to the best constructions of our engineers.’

‘ With regard to the firmness of bast-cells, compare Schwendener’s

BAST. 197

In the monocotyledons the bundles of bast-cells surround the
vascular bundles, either in a sickle-like manner (as in the stem
of the maize, Fig. 133), or lie embedded in the remaining
tissue, sometimes in the inner portion and sometimes in the
outer, as isolated bands, rings or bundles, according to their
function, or they surround the outer side as a connected coat?

Fie. 112.--Transverse section through an involute leaf of the Alfa grass or Esparto
(Macrochloa tenacissima). a, bast-cell coating of the outer (under) side; d@, assimilating
tissue; c, prisms of the upper side; b, hairs (Tschirch). Compare Pharm. Zeit., 1882,
No. 68, :

principal work; furthermore, Th. v. Weinzierl, ‘* Beitrége zur Lehre

von der Festigkeit und Elasticitaét vegetabilischer Gewebe.” Sitzungs-
berichte der k. Akademie der Wissensch., Vienna, 1877, Vol. 76. F.

Lucas, “ Beitriige zur Kenntniss der absoluten Festigkeit von Pflanzen-

geweben.” Sitzungsberichte der Wiener Akademie, 1882, Vol. 85, and

1883, Vol. 87.

‘In some of these cases (especially in the prairie grasses), they serve
in the mechanism which causes the involution of leaves (Tschirch,
Pringsh. Jahrb., xii.). In the dehiscence of fruits, and the phenomena
of torsion of many awns, mechanical cells are also concerned [Stein-
brinck, Inaugural dissertation, 1873, and ‘‘ Berichte der Deutsch. Bot.

198 PLANT ANATOMY,

(Alfa grass, Fig. 112). In the dicotyledons they are located in
the bark (Fig. 113).

oS 3
RETO OOS QE
"se eereer

Fia. 114.—TIllustrations of the more important fibres which are used technically. L,
flax fibres; H, hemp fibres; J, jute; B, cotton; 8, silk; A, alpaca wool; E, Electoral
wool; W, sheep's wool.

Ges.,” i. (1883), p. 27, and others, Zimmermann, Pringsheim’s Jahr-
biicher, xii. (1881), No. 4.]

BAST—LIBRIFORM CELLS. 199

The great mechanical service which the bast-cells are capable
of was also recognized at an early period. The application of
the bast-fibres of hemp and of flax (Fig. 113) for fabrics is a
very ancient one.

The textile fibres which are practically employed may be
grouped in the following manner (compare Fig. 114):

1, Animal fibres: (a) hairs, Wool (W); (b) threads, Silk (S).

2. Vegetable fibres:

(a) hairs, Cotton (B).
(b) bast-cells, Flax (L); Hemp (H); Jute (J); Esparto;
Manilla-hemp.*

In addition to the bast-fibres, the libriform cells of the

wood also assume, especially in the older stems of dicotyledons,

Fig. 115,——A, Sclerenchyma from the inner layer of the seed vessel of Fructus Cocculi
(Cocculus Indicus). B, Some branched cells of the same, more highly magnified.

a mechanical function. The bast-cells which occur in the bark
of dicotyledons (hemp, flax, Fig. 113, the linden) are only of
service in imparting strength of flexure to the stem as long as
the wood itself has not yet acquired sufficient strength.

In accordance with the various mechanical demands made of
the plant, the structures in which the mechanical elements are

‘Compare especially Wiesner, ‘‘ Die Rohstoffe des Pflanzenreiches.”
Leipzig, 18738. Reissek, ‘Die Fasergewebe des Leines, Hanfes, der
Nessel und Baumwolle.” Denkschr. d. Wiener Akad., 1852. Berthold,
“* Ueber die mikroskopischen Merkmale der wichtigsten Pflanzenfasern.”
Zeitschr. f. Waarenkunde, 1883, No. 8; 4. Dorkoupil, ‘* Materialien zu
einem Lehrbuch der chemischen Technologie fiir Gewerbeschulen.”
Jahresber, d. Gewerbeschule Bistritz, 1882.

200 PLANT ANATOMY,

united are also very manifold. We may thus distinguish struc-
tures intended to protect the organs of the plant from the in-
fluence of bending, pressure, traction, or laceration at the
edges or margins (for instance, of leaves).

The stone-cells (sclereids,’ compare also pages 156 and 171),
as one of us (T.) has shown, also possess various mechanical
functions. In seeds which do not possess a thick-walled en-
dosperm, there is found, for example, a hard endocarp, consist-
ing of stone-cells (Figs. 115, 87). This often consists of very

Fic. 116.—Transverse section through a tea-leaf, with the characteristic, branched
sclerenchyma cells. The palisade-parenchyma, which is richer in chlorophyll, is of
a darker color. At the right of the figure are sclereids, isolated by maceration
(Tschirch), _

variously adjusted rows of cells, and can therefore also endure
strong pressure and great expansion or straining (as by germina-

tion) without becoming ruptured. The function of the
sclereids, which are so characteristic of tea-leaves (Fig. 116 *), is

_ ‘The tissue of stone-cells, in accordance with Mettenius, may be
termed sclerenchyma (see pages 156 and 172), Compare Tschirch,
_ “ Beitrége zur Kenntniss des mechan. Gewebesystems.” Pringsheim’s—
dele 16

_ *In young tea-leaves, however, these astrosclereids are wanting.

STONE-CELLS. 201

doubtful. Occasionally the stone-cells contain crystals (Fig.
102) or other substances.

Fic. 118,—Transverse section through the sieve portion of Cortex Coto. st, staff-
shaped bast-cells; m, partially sclerotized medullary rays; s, bundles of collapsed _sieve-
tubes (Moeller). ;

202 PLANT ANATOMY.

The function of the isolated bast-cells and stone-cells, for
example in the Cinchona barks (Figs. 117, 145), in the Pome-
granate-root (Berg’s ‘‘ Atlas,” plate XL.), Aconite root (Fig. 121),
Simaruba, Coto bark (Fig. 118), Cort. Guaiaci (Fig. 105),
Stipites Duleamare (Fig. 144), and Oak-bark is also still in
doubt. Such cells, however, by the form of their transverse sec-
tions and their arrangement, present very useful points of dis-
crimination in the characteristics of many drugs (Cinchona').

Fia. 119.—Transverse section of a rootlet of Rhizoma Veratri. cc, epiblema (p. 177);
m, fundamental tissue, only partly represented inthe drawing; 6, needle-shaped crystals;
kk, nucleus-sheath,. :

The nucleus sheath, vascular-bundle sheath or protective
sheath (endodermis *), which occurs in the root formations of

‘Compare also in this connection, Koch, ** Beitrage zur Anatomie
der Gattung Cinchona,” 8vo, pp. 35, 2 plates. Freiburg dissertation,
1884, especially page 20.

* Evéov within, and 5épyaskin. Compare Schendener, ‘‘ Die Schutz-
imme und ihre Verstirkungen,” Abhandl. der Berliner Akademie,

NUCLEUS-SHEATH. 203

vascular cryptogams, of monocotyledons and some dicotyledons,'

F1a. 120.—Transverse sections through rootlets of Actea spicata. a, epiblema; b, fund-
amental tissue; c, phloém; e, inner bark; k, nucleus-sheath; z, cambium; 7, medullary

rays; v, xylem.
1 Compare C. van Wisselingh, ‘‘ De Kernscheede bij de wortels der
Phanerogamen,” Amsterdam, J. Miller, 1884. (From: ‘‘ Verslagen

204 PLANT ANATOMY.

ben “tt
z
ve
Zz
-P
P
we i
:
4
a
h
Fig, 121 B.

eee noah Fig. 121 C.

ai seos Fic. 121.—Rootlets of Aconitum Napellus; B, Transverse section; C, Longitudinal
_ Section in a radial direction; ¢, epiblema; m, fundamental tissue; h, stone-cells; k, endo-
dermis; p, vascular bundles; z, xylem; s, phloém.
en Mededeelingen der k, Academie van Wetenschappen,” Afdeeling

_ Natuurkunde, 8de Reeks, Deel i., pages 141 to 178, with 1 plate). .

NUCLEUS-SHEATH, 205

and likewise possesses mechanical functions, is also to be con-
sidered here. In these organs, namely, or at least in those cases
which more nearly concern us, all the bundles, or a predami-
nating number of them, are inclosed by a single row of cells
(Sarsaparilla), or, by a layer, the endodermis, which is seen to

cones veniees
ESE AT RS

Fie. 122.—Rootlet from Helleborus viridis; a, epiblema; b, fundamental tissue; c,
central bundle; k, endodermis.

be narrow on a transverse section and consists of but a few cells
(Galanga), All the vascular bundles are located within the en-
dodermis, for instance, in Radiz Sarsaparille, Rhizoma Caricis,
in the rootlets of Actea spicata (Fig. 120), Aconitum (Fig. 121),
Helleborus (Fig. 122), Serpentaria, Valeriana, and Veratrum
(Fig. 119). On the other hand, in Rhizoma Calami, Rhiz.

PLANT ANATOMY,

206

lated bundles.
The endodermis, for example in Sarsaparilla, is composed

of prismatic cells, greatly elongated in the direction of the axis.

aing iso.

Graminis, Rhiz. Iridis, Rhiz. Curcume, Galange, Zedoarie
and Zingiberis the fundamental tissue outside of the sheath also

cont

REECE Ce ee

d,

.
.

through the endodermis (k) of sarsaparilla
o, calcium oxalate.

>

(Fig. 123), forming a tube or sheath, located centrally

fundamental tissue, and contai

Fic. 123.—Radial longitudinal section
Middle bark; b, bundles; m,

the
In
e

in
¥

ning within it the bundles.
azom

as also in Rh

.

in

a Gram

:
3
Ps
a
:
4
:
cee
®
ee

vr

eties of Sa:

bundles are crowded

some ¥:

NUCLEUS-SHEATH. 207

circle, in other cases they are dispersed, as is the case in Rhizoma
Veratri and Rhizoma Caricis, or to a still greater extent in Tuber
Aconiti, or but a single central bundle is present, as in the root-
lets of Veratrum.

The cells of the sheath are not always elongated, but often

Fig. 124,—Transverse section through the endodermis of Vera-Cruz Sarsaparilla.

nearly cubical or only slightly extended. They are also often
thin-walled, contain starch, and are then called starch-layer or

starch-sheath.

Fic. 125,—Transverse section through the endodermis (k) of Rhizoma Galange; a;
fibro-vaseular bundles; r, resin-cells; m, fundamental tissue.

The walls of the endodermis which are directed toward the
axis are usually thickened; in the lateral walls this is also
sometimes the case, so that the lumen or cavity, for instance in
the Vera Cruz Sarsaparilla (Fig. 124), becomes very much con-

208 PLANT ANATOMY.

tracted. The transverse sections of these cells of the nucleus-
sheath therefore appear differently, according to the thickness of
the thickened layers, and thereby afford serviceable characteris-
tics for the recognition of the several varieties of a drug.’

While the nucleus-sheaths in most of the examples that have
been cited are built up of a single row of uniform cells, the
root-stocks of the Zingiberacew deviate considerably in this
respect. Indeed, the endodermis of the rhizomes of Curcuma,
Galanga, Zedoaria, and Zingiber is composed of several rows of
cells * (Fig. 125).

3. The Absorbing System.

The absorption of inorganic salts from the soil is effected by
the aid of the roots and especially by means of the root-hairs.
The latter, which are true trichomes, by forming manifold pro-
tuberances, become most intimately attached by their growth
to the particles of the soil. |

Root-hairs are found on but few officinal roots (for instance,
Sarsaparilla, Fig. 126). In most cases they are broken off in
the process of unearthing them, or they may have been already
absent at the-time of collection, since the formation of root-
hairs only takes place in definite and young parts of the root.

For the absorption of organic nourishment, the phanerogamous
parasites penetrate the host-plant by means of the so-called
haustoria (as in the case of Cuscuta). To these haustoria cor-
respond the surfaces of the cells lying close to the endosperm,
which consist mostly of palisade-shaped cells with protuberances
resembling root-hairs, and which are especially met with on the
scutellum of the Gramineew. They serve for imbibing the re-
serve substances.

In order to convey nourishment to the embryo during germina-

‘Compare Schleiden, « Beitrige zur Kenntniss der Sarsaparilla.” |
Archiv der Pharm., 1847, Arthur Meyer, I bid., 218 (1881), p. 280 et seq.
Berg’s ** Atlas,” Plate iv, Flickiger, “ Pharmakognosie,” p. 295.

-? Compare Arthur Meyer, Archiv der Pharm., 218 (1881), p. 419.

THE ASSIMILATING SYSTEM. 209

tion, a longitudinal cleft is also occasionally found in ‘the en-
dosperm (Strychnos Nua vomica, Coffea *).

‘Fic. 126,—Longitudinal section throngh Radix Sarsaparille; e, epiblema; p. hairs;
cc’, cells of the bark, which are thickened on one side; d, parenchyma; 0, calcium
oxalate,

4. The Assimilating System.

The assimilating tissue serves primarily for the formation of
organic substance from carbonic acid and water under the in-
fluence of light, to which procedure the name of assimilation

has been given. This tissue is filled with chlorophyll granules
‘(compare page 100), and its cells possess forms which tend, as

far as possible, to the transmission of light on all sides, and the

rapid removal of the products of assimilation.’ a

| Jager, “ Endosperm der Coffea.” Bot. Zeit., 1881, p. 336. ae

‘Compare in this connection Haberlandt, in Pringsheim’s Jahrb.,

xiii. (1881), ae pon

44

210 PLANT ANATOMY.

The surface of the leaf which is chiefly exposed to the’ light
becomes the assimilating side. With bifacial * leaves, that is, such
as are flatly expanded, and the upperand under surface of which
is differently developed (as in the heart-shaped Fol. Eucalypti,
Figs. 2 b, 127, 129, Lactuca Sativa), this is the upper surface;
with centric leaves, that is, such as are placed vertically, and both
sides of which are equally constructed (Fol. Eucalypti, sabre-
shaped, Figs. 2 a, 128, Lactuca Scariola), it is both sides.

The cells of the assimilating side, which is always of a darker
green color, are replete with numerous chlorophyll granules,
located along the walls, and are extended ina palisade-like man-

Ss /0e ‘S

Fic. 127.—Transverse section through a heart-shaped (bifacial) leaf of Eucalyptus
globulus; oe, oil-space; s, stomata; w, undersurface; 0, upper surface. After Tschirch,
Pharm, Zeit., 1881, No. 88. ,

ner more or less perpendicular to the vertical axis of the leaf
(Figs. 96 g, 127, 128, 129 pal, palisade * parenchyma).
The special development of palisade parenchyma remains un-
_affected only in typical shade-plants.* On the other hand, in
- all centrically constructed leaves, both sides are provided with

' Bis, twofold, and facies, side.

* From the French word palissade, and this from the Latin masculine
palus (not pallus !), therefore not pallisade, as it is often incorrectly
written. (This applies more especially to the German orthography.
F. B, P.)

* Globularia Alypum L. and other species present a notable example
of an homogeneous leaf tissue without a palisade layer.

THE ASSIMILATING SYSTEM. 211

palisade cells (Fig. 128). The assimilating surface is often in-
creased by a falling off of the membrane (as in the needle-shaped
leaves of the Conifere). In order to be able to take up the
products which are formed and to conduct them rapidly, the
palisade cells are now and then located upon funnel-shaped col-
lecting cells, which are in connection with the proper cellu-
lar threads (vascular bundles); the latter running into the
nerves of the leaves as a much branched radiating system, with
extremely fine terminations (Fig. 130).

‘K > s a

aE ae
—

Fig. 128.—Transverse section through a sabre-shaped (centric) leaf of Eucalyptus
globulus; oe, oil-spaces, with drops of oil; gfb, vascular bundle; s, stomata; k, corky
growths; c, cuticle (Tschirch). Compare also Fig. 2 a and b.

The under side of the leaf, which is always ofa lighter green
color, contains much less chlorophyll than the upper side, and
is traversed by wide air-canals (spongy parenchyma, leaf-
merenchyma, Fig. 129 sch).

The entire interior of the leaf, with the exception of the
vascular bundles, which is inclosed by the ~ epidermal sides,
is termed the mesophyll.’

1 Mé6os in the middle, pvador leaf.

PLANT ANATOMY.

212

“(qoargosy,) ewur04s ‘ds $(w) joyyueur Jo syeys419 pus [Io eVBIOA JO sdoap YA ‘puyys jo ‘pe {(47) TO
yo sdoap yQIA pepracad puv sernueas [AYdosopyp WLM pory ‘teuL1oy oyy Ayperoodse ynq ‘yao ‘wuAyoucared ASuods ‘yos Sonssy
apestyed ‘7nd ‘srmaepide ‘da {eurdyoueToo ‘yj09 {stjeo-4seq ‘q {(ue_Ax) uomasod aepnosva ‘6 {(utgoyyd) uoyaod sacs ‘gs tumiq =
-ureo ‘9 feypunq zepnosva ‘f6 {(N) carou o[pprud 919 4B ‘Hy24adid DYyyUaMT JO JRO, B YSNOAY) UOTOS OSLOASUVAT— GEL “DIL

”

have utilized

una—#S

YO

~@\®

ler Blatter und Krdauter.

_ | * Anatomische Charakteristik offic
Abhandl. der naturforschenden Ges., Halle, xv. (1882),

]

ine.

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ES ee a

Adolf Meyer,’ as also Lemaire (see page 51),

THE CONDUCTING SYSTEM. . 213

the anatomy of leaves, especially the epidermis and the tri-
chomes, for the purposes of diagnosis.

5. The Conducting System.

When a leaf of the plantain (Plantago) is torn off or a maize
stem is broken, there project from the fractured surface numerous
fine threads. If the fibrous, fractured place is evenly cut with a
sharp knife, it may be seen, even with the unaided eye, that
there is a large number of compact, isolated dots imbedded in
a more delicate tissue. If the maize stem is exposed to decay,
only a bundle of very long, fibrous threads finally remains, sur-
roundered by a delicate membrane, the cuticle. These threads,
as is shown by an anatomical comparison, correspond to the
dots upon the transverse section. The threads are termed
Jibro-vascular’ bundles, vascular bundles, or conducting bundles.
As is already evident from their considerable length, they
serve primarly for the conduction of substances, chiefly in the
longitudinal direction of the organ. ;

The same extended threads we meet with in the maize leaf.
If the latter (or any elongated leaf of a monocotyledonous.
plant which may be chosen) is held toward the light, a large
number of nearly parallel, lighter colored threads (nerves) may
be seen in the green tissue.

_ The nerves do not appear so regular in a dicotyledonous leaf.
Here they are variously branched, anastomose with each other,
and form a delicate network of fine lines. This is rendered
prominent, in an especially handsome manner, when leaves like
those of Digitalis (Fig. 130), Datura or Matico are rendered
transparent by long maceration in alcohol (of about the specific
gravity 0.900), or when freed by decay from the parenchymatous
fundamental tissue * (Ettingshausen’s leaf-skeleton).

That which applies to the leaf and stem is also applicable to
the roots. Upon a transverse section of Rhizoma Filicis, for
example, may be observed a double circle of such threads

? Fibra, fibre, fibre-shaped cell, and vas, vessel. =
*Fundamental tissue in the sense explained on page 176.

PLANT ANATOMY.

214

t the frame-
have been

issues

t

ise represen
ining

e root-stock when the rema

or bundles (Fig. 131, f), which likew

work of th

Fie. 131,

Fie. 132.

?

Cc, A

um Filix mas.

of

threads enter the leaf-bases (which are here
rhizome, g threads or vascular bundles.
bundle,

(Sachs), .4, Front end of the rhizome, showing in the

through the underground stem of Aspid:

Berg).

ris

18 purpurea (Planchon),
places where the

ion

‘posed piece of the
more highly magnified portion of a

Filicis ma:

Digital
Fie. 132.—Rhizoma
rhombi
cut off). B, A decom

erse sect
spaces the

LC

—Leaf of

Fie. 130.
Fie. 131.—T

Jf, threads or vascular bundles (

.

THE CONDUCTING SYSTEM. 215

removed. If sucha root-stock be placed in a liquid prepared
from decomposed meat, after a short time the parenchyma will
be destroyed, and, after washing away the remnants of the
same, the far more resistant threads alone remain behind. These
do not run parallel in this case, but are variously intertwined
(Fig. 132).

In the root-stocks of the Zingiberacew and in Rhizoma
Caricis there are numerous isolated bundles distributed
through the fundamental tissue; in Sarsaparilla and in Rhizoma
Graminis they are brought together in the form of a ring
(vascular-bundle ring). Differently constructed from these (see
below), but similar in their entirety, are the vascular bundles
in dicotyledonous roots, as likewise in the dicotyledonous stem,
which unite to form a continuous “ring.” Dicotyledonous

roots often possess only a central, axial bundle (Jpecacuanha,
Taraxacum, Levisticum, rootlets of Arnica, Valerian, and
Helieborus ; compare also the Figures 119, 120, 122).

But we meet these bundlesalso elsewhere onevery hand. The
fruit-pulp of tamarinds is traversed by such coarse, string-like,
vascular bundles, and the shell of the almond is covered with
them. They occur in the arillus of the Myristica (Macis), as well
as in the mericarps of the Umbellifere, in the calyx of the clove,
as well as in the stigma of the Croeus—everywhere forming
long threads, which serve for conveying and for conducting
away organic and inorganic building material.

The elements of the vascular bundles are so constructed that the
impediments to movement are restricted to a minimum. The
transverse walls are greatly reduced, often lacking altogether at
wide intervals, or, when present, provided with pores’ (vessels),
or even pierced with holes (sieve-tubes), thus having the dif-
fusing surfaces greatly enlarged.

Of what elements is such a vascular bundle or conducting

‘The term pores, or pits, is applied to all those thin places of the ©
membrane which are still closed only by the middle lamella. By age
the pits occasionally become actual perforations (as in the wood of the
Conifere) ; compare page 153. |

216 PLANT ANATOMY.

bundle* composed ? In the first place, it is necessary to.exclude
therefrom any bast-cells (Fig. 133), which are, however, not at —
all regularly united therewith even in the monocotyledons.* So
long as it was not yet known that the bast-borders of the vas-
cular bundles perform exclusively mechanical functions, they
could be considered, and quite properly so, from purely anatomi-

iv

in)

)
ewe
if

>

[)

4,

vy
f

i

I

I

<a.
Mf

NYY.
ii

ITN
A
Hitt

ia

ii

I

Ws
Whi

)

Mitt

Tw

ii

Fig, 133. Fia. 134.

Fig. 133.—Transverse section through a collateral bundle of the maize stem (monocoty-
ledonous type). a, outer; i, inner; p, fundamental tissue; r, annular vessel: 8, spiral
vessel; g, pitted vessel; 1, intercellular space (containing air); 2, wood-cells; v, phloém.
The entire bundle is surrounded by a sheath of bast-cells (Sachs).

Fic. 134.—Spirals capable of uncoiling and annular vessel from Bulbus Scille (longi-
tudinal section),

cal reasons, as belonging to the latter. Since it is known, however,
that all bast-cells only serve to impart firmness to the plant,

' With regard to the structure of the vascular bundles, which can
only be briefly mentioned here, compare De Bary, ‘‘ Anatomie,” p. 328
et seq. i ;

* The bast border of the cribrose or sieve portion has been termed bast,
that of the vascular portion, libriform. Compare the table on page 217.

VASCULAR BUNDLE. 217

and have nothing to do with the conduction of substances, they
must also be separated from the conducting threads or bundles.

A typical vascular bundle (Fig. 133) consists accordingly only
of the vascular portion (Fig. 133, x r sg, xylem" in part) and
the cribrose or sieve portion (Fig. 133, v, phloém? in part). If
the terms, xylem and phloém, are so comprehended that the
bast and libriform cells are not regarded as necessary constitu-
ents of it, the ‘term sieveportion” is synonymous with that
of phloém, and “ vascular portion ” with that of xylem.*

The vaseular portion consists of vessels, tracheids, and
wood-parenchyma.

The vessels (¢rachee) are very long tubes, which often traverse
uninterruptedly the entire plant organ, and the walls of which,
In consequence of unequal growth in thickness, are variously
thickened (annular vessels, Fig. 134, 133 r, spiral vessels, Fig.
134, 133 s, trabecular and scalariform vessels, Fig. 135 fv, 133
g, 142, pitted vessels). They are formed by the disappearance
of the transverse walls in a row of cells (fusion of cells), In
coniferous wood, the vessels, as a rule, are wanting (Lignum
Juniper).

1 Evdov wood.

* Protor bark of a tree.
°For the sake of clearness these terms may be arranged in a table :

Bast-cells )
(mechanical
elements)
if = ( Phloém )} % -
Zo Sieve-tubes and 3 { (Bast) S
Es Sieve portion Latticed cells fi 3
a a¢ (Leptom) — a = | 3
i $; Ls
BS ma ( +
= 5 =| Vascular portion | Vessels and | J
32 (Hadrom), Tracheids | 2
che L Wood parenchyma a Es
~ (Wood) J 2
Libriform
(mechanical
elements)
new old

division

218 PLANT ANATOMY.

The tracheids (wood-cells) are prosenchymatous and likewise
elongated cells, but these are far from attaining the length of the
vessels. ‘Their walls are mostly pitted. In coniferous wood,
which, as a rule, consists only of tracheids, areolated or bordered
pits occur (Figs. 64, 72, 136).

The last terminations of the nerves in leaves consist only of
tracheids. The vessels and tracheids convey air and water; they
form the conducting system for water and the inorganic nutri-
ment.

The wood-parenchyma is not necessarily present, but is de-

oo
Fia.' 135.—Vessels (fv) from Rhizoma Filicis, thickened in a scalariform manner
(Berg). .

_ veloped in many cases. In contradistinction to the prosenchy-
matous, tracheal tissue, the former consists of thin-walled cells,
which are connected with the neighboring vessels and cells of the
medullary rays by means of pores, and are filled with various
substances (protoplasm, starch, oxalate and tannin). The wood-
_ parenchyma serves for conducting the carbohydrates and for the
_ Storage of starch, and either traverses the vascular portion in
____ the form of more or less isolated cells or occurs in the form of
ae aay (Lignum Campechianum, Fernambuci, Guaiaci, San-
Bee 2S ae at ; 2 aN j

WOOD-PARENCHYMA. 219

The very dense wood of Lignum Quassia (Fig. 137) contains
groups of parenchyma (p), which run transversly in the form of
bands from one medullary ray to the other, and intersect the

nM

Fie. 136. Fic. 137. ‘
Fig, 136.—Schematic representation of a longitudinal section of two tracheids from
coniferous wood, with the different forms of pitting of the membrane (a-d), viewed
( .
cWas tae iris Gaee aaune tele Quassice jamaicense; the medullary
rays, which form two or three rows of cells (r), are connected by transverse bands of
wood-parenchyma (p); between them is the libriform tissue.

wood in the form of numerous, closely arranged concentric

rings.’ By their wider and not thickened cells they present a —
‘Compare also Berg’s “ Atlas,” plate xxvii., Fig. 65 (Lignum Cam-

pechianum), plate xxvii., Fig. 64 (Lign. Guaiaci). oe

.

220 PLANT ANATOMY.

very sharp contrast to the zones which surround the large ves-
sels, so that the varieties of wood just mentioned and others
derived from tropical trees appear as if they had annual rings.
A transverse section, however, soon teaches that these bands of
parenchyma do not form connected rings, and that they also differ
in other respects essentially from the annual rings. A longitudinal
section shows still more distinctly that they consist of parenchy-
matous tissue. They are designated as apparent rings.’

Such groups of wood-parenchyma never occur in the vascular
portions of monocotyledons and in the young wood of dicotyle-
dons, but they are not rare in the older wood, the general struc-
ture of which may here be considered.

The most marked distinction between the monocotyledonous
and the dicotyledonous stem depends upon the development of
a cambium in the latter (Figs. 129, 138).

The cambium (cambium ring, thickening ring) belongs to
the formative tissues, and effects by its activity the growth in
thickness. Consequently, where a cambium is wanting, as in
the monocotyledons, a (secondary) growth in thickness is also
excluded, The stems, therefore, remain slender and thin (Palms,
Bambusa*). The isolated bundles found in monocotyledons
become initially formed as such already at the growing point,
and no merismatic layer is maintained between the vascular and
sieve portions (compare Fig. 133).

It is otherwise with the dicotyledons and the gymnosperms.
Here there appears between the interior, vascular portion, and
the exterior, sieve portion, a layer consisting of very thin-walled,+

? The fine transverse undulations or horizontal stripings which are
observed upon tangential sections of many woods (Picrasma excelsa,
Pterocarpus santalinus, Guaiacum, Ceesalpinia, Diospyros, and others)
proceed, according to Von Héhnel (Berichte d. deutsch. bot. Ges., ii., 3),
from the horizontal rows of equally large medullary rays or from a trabec-
ular arrangement of the pits of the tracheids (Tamarindus), or from both
Causes at the same time.

_ * Only some tree-like Liliaceze (Aloé, Draccena, Yucca) possess & SeC-
ondary growth in thickness.

_ *In drugs the cambium cells are, therefore, either torn, or at least

_ greatly distorted and bent. Upon a transverse section of the stems and

CAMBIUM. 2971

tabular cells, rich in protoplasm, which are in a state of active,
tangential division. By the activity of this layer vessels are
separated on the inner side and sieve elements on the outer (Fig.
138).

Since this activity is not manifested throughout the entire
year with the same degree of productiveness, but is more ener-
getic inthe spring than in autumn, the elements of the spring
wood are more numerous than those of the autumn wood.’
Since in winter the activity of the cambium ceases completely,
it follows that the large spring cells must directly follow
upon the narrow autumn cells, There is hereby produced, upon

> i

Fie. 138.—Schematic representation of the activity of a cambium-cell (c). 1, before
the beginning of its activity; 2, the cell has become radially extended, and (8) divided:
a xylem cell (x’) has been separated. In 4 the cambium-cell is again extended, while the
xylem cell has already become thickened. 5, the cambium-cell by repeated tangential
division has this time separated a phloém cell (p’). While the first-formed xylem cell
(#’) becomes further thickened, the cambium cell is again extended (6), again a tangen-
tial wall appears within it, and the second xylem cell (a’') and soon afterwards (8) the
second phloém cell (p’’) is separated. The former become strongly thickened, the
latter remain thin-walled (Tschirch),

a transverse section, a visible delineation of rings, for the most part
concentric, which are designated by the name of annual rings

roots of dicotyledons, the cambium zone frequently appears as a circular
line, distinguished by a darker color (Radix Glycyrrhiza, Rad. Calum-

be, Rhiz. Rhei, Stipites Dulcamare).
! That the cells become narrower and smaller toward autumn, does

not proceed (as Sachs, De Vries, and others have stated) from an in-
creased pressure of the bark in autumn.

222 PLANT ANATOMY.

(Fig. 18079). In the case of dicotyledonous foliage trees, the
distinction between autumn and spring wood is also further in
creased by the fact that the latter is much richer in yessels.!

(Fig. 78 jf).
The so-called wood-ring of dicotyledonous stems is produced

Fie. 139.—Libriform from Quassia wood.

by the confluence of isolated vascular bundles, in their vascular
as well as in their sieve portions, through the activity of an

- (intrafascicular) cambium, the bundles being originally formed
in a loose circle,

1 Berg’s ‘‘ Atlas,” plate xxv., 60, and plate v., 21.

MEDULLARY RAYS. 223

The elementary organs of the wood,’ as the vascular portion
of stems may be briefly termed, are as follows: the vessels and
tracheids, or tubes conveying water, the wood-parenchyma cells,
which serve for conducting the carbo-hydrates, and the libriform
cells, or the specifically mechanical elements of the wood. Be-
sides these, the wood is traversed in a radial direction by the
medullary rays.

The first three forms of cells have already been considered.

The libriform cells* (wood-cells, wood-fibres) are the bast-
cells of the wood (Figs. 137, 139), and therefore, strictly con-
sidered, belong to the mechanical system (see the latter). They
are of service, however, especially in the transitional forms, to the
other elements of the wood, occasionally also for conducting and
storing nutritive material, and are therefore sometimes provided
with contents. They are prosenchymatous, thick-walled,
provided with cleft-like, oblique pores, and are never as long.
as the true bast-cells.

The medullary rays (parenchyma rays) are aggregates of cells
consisting of one or several rows, always built up of cells which
are very much extended radially (Figs. 78, 137, 144, 149), and
which pass from the medulla through the cambium and the
sieve portion, often penetrating deeply into the bark (Figs. 100
r, 118 m, 149, 137 r, 180, 181).*

When the interior of stems is filled with fundamental tissue,
the so-called medulla, it is thus, by means of these radiating
lines of cells, placed in connection with the tissue of the outer
bark, which is located on the periphery of the fibro-vascular
bundles. These rows of cells are therefore very appropriately
called medullary rays. Their development in a vertical direc-
tion is governed by the number of rows of cells placed over each

i i tilized by
'The anatomy of the wood has been diagnostically u
Nérdlinger (* Anatomische Merkmale deutscher Wald- und Garten-
holzarten,” Stuttgart, 1881). :
? Liber, bast, forma, form. Compare Sanio, “ Vergleichende Unter-
suchungen tiber die Elementarorgane des Holzkérpers,” Bot. Zeit., 1863,
101.
® Compare also Berg’s “ Atlas,” xxxvi. to xl., Figs. 86-94 r.

‘

224 PLANT ANATOMY,

other in a compartment-like form. Very frequently this num-
ber is not considerable, so that the medullary ray, upon a trans-

.

———_ cae
Giipecennes SS
$$ ZOOS oOoOvoS oS:
Seer ——

Fie. 140.—Cells of the medullary rays, consisting of one and three rows, cut trans-
versely; tangential longitudinal section of a dicotyledonous stem.

verse section, which is made vertical to its surface of length (tan-
_ gential to the surface of the stem), represents a cleft filled with

Fie. 141.—Medullary rays from Lignum Juniperi, consisting of a single row of
cells (a).

A, Tangential longitudinal section, a medullary ray.
Cc. Transverse section through the wood of the root, a medullary ray; c, annual rings.

parenchyma (Fig. 140). In breadth the medullary rays some-
__ times present but asingle row of cells, as, for instance, in Lignin

MEDULLARY RAYS. 225

Junipert (Fig. 141), L. Guaiaci, L. Quassie surinamense,*
sometimes two or three rows, as in Lign. Quassie jamaicense
(Fig. 137), and sometimes still more, as in rhubard (Fig. 142).
It follows from this that, on a transverse section, the fibro-
vascular bundles must appear either in radial rows, or separated

wv ae
Fic. 142,—Tangential longitudinal section of Rhubarb. A, Fundamental scene a
the bark with five medullary rays, which are cut transversely. B, Section from the
xylem portion, which is traversed,by netted vessels (v) of considerable size; 0, rosettes
of oxalate crystals. :
from each other by large medullary rays. The tissue of “i
medullary rays, at least within the compass of the fibro-vascular

1 Berg’s ‘“ Atlas,” plates xxv. to xxviii.
15

226 PLANT ANATOMY.

system, consists almost entirely of cubical or horizontally (radi-
ally) extended (parellelopipedal), thin-walled cells which are
joined together in a wall-like form, without intervening spaces

(ames et se [ee | em
Oooo macy

Fig. 143.—Wall-like cells of the medullary rays from Lignwm Juniperi, ona radial
longitudinal section.

(Fig. 143). This regularity is lost at the place where the me-
dullary rays pass into the bark.

a
mm
a L)
‘
Cena nod 5 )
Loto . 4 eX: Cie bE
a fe Se ere: 3 Mg a ee
i

i.”
Fie. 144.—Single-rowed medullary rays, r, in Stipes Dulcamare, which gradually be-
come lost in the bark ; i, inner bark ; m, middle bark; a, outer bark (Berg).

- In the cambial zone, when the growth in thickness is already

MEDULLARY RAYS. 227

Sse

Ore EAS Ne

A fT |S
tet

oe

Vee
<a
Ke

EEL
a Ct 1
Arter e

ve

a)

De

S
‘ mes =
Seses; ee"

SILK
eK |
“ue

a4

Fie. 145.—Transverse section through phloém portion of the bark of Cinchone

lancifolia ; r, primary medullary rays; s, secondary medullary rays; q, stone-cells; p,.
bast cells (Berg),

228 PLA NT ANATOMY.

quite advanced, medullary rays of smaller size are often subse-
quently developed, which are designated as secondary medullary
rays. They are met with frequently, for example in the Cin-
chona barks (Fig. 145), and, according as they are located in
the xylem or the phloém, are termed secondary xylem (or
medullary) rays, or bark rays.

The great variety of special features in the structure of the

Fic. 146 a.—Transverse section through the peeled root of Rheum Rhaponticum.
¥, brownish-red medullary rays; v, vessels; 6, crystals of oxalate.

medullary rays affords very noteworthy and characteristic dis-
tinctions for many drugs.’ As against roots which have a de-
cidedly radiate structure (for instance, Rad. Rhapontici, Fig.
146 a), it may suffice here, for example, to refer to such which

"In the Conifer, as Essner has shown (Abhandl. d. naturforsch.
_ Ges., Halle, 1882), the number and height of the medullary rays, as well
_ asthe form and size of their cells, possess no diagnostical value.

MEDULLARY RAYS. 229

have no distinctly marked medullary rays, either in the wood
or in the bark (Radix Ipecacuanhe and in Rad. Tarazaci).

The medullary rays of rhubard (Fig. 146 6), which only oc-
cur regularly in the cortical portion, are very characteristic
with regard to their course and contents. Within they are
variously serpentine, and ‘therefore, since they consist of cells
with reddish-yellow contents, produce the peculiar spots on the
surface of the peeled rhizome and the marbled appearance of
the interior.*

In those fibro-vascular bundles where, by the formation of
the bundies themselves, the activity of the cambium is termi-
nated already at the growing point, no medullary rays are

ele sea

ty

Fig. 146 b.—Transverse section of Rhubarb. e, remnants of the bark, which has
been pared off: c, cambium, a, medullary rays of the marginal portion; s, stellate
spots; m, fundamental tissue.

formed; hence they are wanting in the monocotyledons and in
the vascular cryptogams.

Although, for example, the vascular bundles in a number of
monocotyledonous roots are closely connected with each other,
and are arranged in the form of compact, closed rings (Rhiz.
Graminis, Rhiz. Caricis, Rad. Sarsaparille, and rootlets of
Veratrum), the medullary rays are nevertheless wanting in these
rings.

The cells of the medullary rays have the function of con-
luctiag in a radial direction. But they are also receptacles for

1 Berg's “ Atlas,” Plate xii.

230 PLANT ANATOMY,

reserve substances. Starch is, therefore, often found in them,
especially in winter (Quercus).

Fie, 147.—Sieve-tubes from Fructus Papaveris (Dippel).

In older stems and branches, the heart-wood (duramen) and
Sap-wood (alburnum) may be distinguished. The heart-wood,

Fre. 148.—Sieve-tubes of Acer. (a) more feebly, (b) more highly magnified. a,
Sieve-plate on a transverse section; e, the same as se2n from the surface; p, the con-
tracted plasmatic contents of the sieve-tubes (Hartig).
which is mostly characterized by a darker color (logwood, red-
wood, and other varieties of dye-woods, Guaiac), consists of the

SIEVE-TUBES—CAMBIFORM CELLS. 231

older layers of wood, which are no longer vitally active, while
the sap-wood is formed of the younger wood, which often com-
prises but a few annual rings.

The second component of a vascular bundle is the sieve
portion’ (phloém in part). This consists of sieve-tubes (with
latticed cells) and cambiform cells; its elements never become
lignified, which, on the contrary, occurs regularly with the ele-_
ments of the wood.

The sieve tubes’ are elongated, thin-walled cells, which
correspond, in their entire structure, to the vessels (in the
xylem) (Figs. 147, 148). They serve for conducting non-dif-
fusible, albuminous substances, which they abundantly contain,
and are, therefore, mostly divided by oblique, transverse plates,
which appear pierced by numerous holes (sieve-plates, Fig.
148 w).

The sievye-tubes are often accompanied by small latticed cells,
which, from the manner of their development, belong to them.

The ecambiform cells are elongated, thin-walled cells, very
rich in protoplasm, without pits, which are very similar in
shape to the cambium cells, and conduct the more readily diffu-
sible substances. ‘

Other parenchymatous elements besides those already men-
tioned also participate in the conduction of substances, thus,
particularly, the starch sheaths and sugar sheaths, which
surround the vascular bundles.

In the dicotyledons and gymnosperms, the medullary rays, as
a rule, also intersect the sieve portion (bark rays or phloém
rays); often, however, they remain imperceptible in the tissue
of the bark (Radix Ipecacuanhe).* If they are distinctly

1 Alsoincorrectly termed ‘‘ soft bast.” The term “ bast” should remain
confined to the mechanical elements of bast-cells. ae

2 Wilhelm, ‘ Beitrage zur Kenntniss des Siebréhrenapparates dicotyler
Pflanzen,” Leipzig, 1880. Janczewski, ‘‘ Vergleichende Untersuchungen
iiber die Siebrdhren.” Sitzungsber. d. Krakauer Akademie, .881. De

Bary, ‘‘ Anatomie,” p. 179.
* Arthur Meyer, Archiv der Pharm., 221 (1883), p. 737.

232 PLANT ANATOMY.

visible, that portion of the bark: which is traversed by them
- Is termed the inner bark (Fig. 144 7, 149 7); the portion toward
the exterior which is connected with it, and which mostly con-

Serre]
=a eI
fe ci
‘ord n

aT
rn RPh

GEES |

7 ae
Fie. 149.—Transverse section through the bark of anolder stem of Ceratonia Siliqua
L. k, sclerotized cork-cells; s, sclereids: m, medullary rays; sb, sieve-tubes; i, inner
bark; mt, middle bark; a, outer bark (Moeller),

sists of a mixture of parenchymatous cells containing chloro-
phyll (bark parenchyma) and of mechanical cells (bast-cells,

' With regard to the structure of the bark, compare especially Moeller,
“ Die Anatomie der Baumrinden,” Berlin, Springer, 1884.

BARK. 233

stone-cells, collenchyma), is termed the middle bark (green
bark, Figs. 144 m, 149 m); and the epidermis (cork) the outer
bark (Figs. 144 a, 149 a).

Naturally, barks are found only in the dicotyledons and gym-
nosperms. The name is used to designate the entire collection
of cells located outside of the cambium. Nevertheless, in the
case of monocotyledons the portion of tissue located outside of
the nucleus-sheath is also, for the sake of convenience (but in-
correctly) termed bark.

In the middle bark, entire lines of cells are often found which
are filled with crystals of oxalate. These frequently accompany

Fic. 150,—Bast-fibres from Cinchona (China) Alba Payta, with impressions of the
neighbouring crystal cells.
the bast-cells, and are occasionally in such close contact with the
latter that depressions are produced. The crystal-cells appear
indented into the bast-cells, as in ‘* Cinchona (China) alba” *
(Fig, 150) and in the bark of Aspidosperma Quebracho.”

If the activity in the entire (cambial) thickening ring pro-
gresses in a uniform ratio, then the separated elements of the
wood and of the bark are arranged in strictly radial rows.

In the wood, these rows mostly remain preserved (coniferous

? Fitickiger, Neues Jahrbuch fiir Pharmacie, xxxvi. (1871), p. 293.
* Hansen, “‘ Die Quebracho-Rinde.” Berlin, 1880.

234 PLANT ANATOMY,

wood, logwood, red-wood) in consequence of the thickness of

the walls of the cells and the absence of dislocating tensions.
In the bark, however, especially at later periods, in consequence
of the various dislocations, lacerations, and subsequent divisions
of the cells, etc., they are seldom distinct (cinnamon). Even
the bark rays have seldom (for instance, in the willow) a pre-
cisely radial course (Cinchonas). Nevertheless, the radial rows
of the elements of the bark may often be followed nearly to the
epidermis.

According to the manner in which the distribution of the
vascular portion and the sieve portion takes place in the vascu-
lar bundle, the following designations are employed. The bun-
dle is spoken of as concentric when one of the two parts sur-
rounds the other in a sheath-like manner, that is, either the
sieve portion the vascular portion (Rhiz. Filicis, Fig. 135), or
the vascular portion the sieve portion (Rhiz. Calami, Fig. 151,
Lthiz. Iridis);' as radial, when the vascular and sieve portions
are located radially, one behind the other (the typical structure
of the vascular bundles of roots, Figs. 120, 121); as collateral,
when both parts run beside each other and are in contact later-
ally (the typical structure of vascular bundles in the stem and
leaves of phanerogams).

In the leaf, the sieve portion always lies in a direction toward
the lower surface, and the vascular portion always toward the
upper surface (Fig. 129 g, sb). :

If, through the activity of the cambium, the formation of
secondary wood takes place, the primary bundles, which had
already previously been formed at the tip without the activity of
the cambium, move more and more toward the interior; and in
all those cases where they Jie relatively widely separated from each
other, they project toward the medulla in a bow-like form and
constitute the medullary-crown or medullary sheath.

The outermost layer of the fibro-vascular body of roots, which

Yemains fora long time capable of development, or, in other
__ words, in a parenchymatous condition, and in which the root-
lets become formed, is termed the pericambium.

---——- . Compare Berg's “ Atlas,” Plate xxi., Fig. 51.

STORING AND AERATING SYSTEMS. 235

6. The Storing System.

Since the plant does not immediately consume all the sub-
stances which it takes up or forms, it must possess some
arrangement whereby these substances may be stored for future
use. The plant therefore develops receptacles for reserve
Substances, in which it deposits the building materials. Such
reserve receptacles are the seeds,’ fruits, rhizomes, tubes, bulbs
and perennial roots; indeed, even the stem (especially the me- ~
dullary rays)’ can become a receptacle for reserve substances.

The substances deposited in these organs are mostly solid;
‘readily soluble substances (sugar, dextrin) are avoided, as a
rule, by the plant. In the sugar-beet, however, the sugar is a
reserve substance. They are either carbohydrates (starch, inulin,
cellulose, for instance in the Phytelephas seed), fats (oil) or
albuminous substances (gluten, aleurone); but water also is
stored for the germinating plant (bulbs), or it is energeti-
cally retained by the fully-developed plant (epidermal water-
tissue). *

v. The Aérating System.

In the development of plants, the interchange of gases plays
an important part. The plant consumes carbonic acid (and
oxygen) from the atmosphere, and gives up to the latter oxygen
(and carbonic acid), as well as aqueous vapor. In order to render
possible this interchange of gases, cavities in the tissue are
necessary. These collect the gases, and the canals, which serve
respectively for their entrance and exit, convey them from with-
out to the interior and inversely.

For these reasons the plant is abundantly provided with
intercellular spaces (Figs. 129, 127), especially in those parts

’ The seeds have received appropriate anatomical treatment by Harz,
‘* Landwirthschaftliche Samenkunde.” Berlin, 1885.

* Malpighi, ‘‘ Anatomes plantarum idea” (1671), had already recog-
nized the medullary rays as reserve receptacles,

® All bulbs (Bulbus Scille) attract water abundantly, in consequence

of the mucilage and sugar which they contain.
* Westermaier. Pringheim’s Jahrb., xiv., 43.

236 PLANT ANATOMY.

in which the processes of development are actively effected.
While the cells of the epidermis, of the cambiform, of the cam-

{ Fig. 151.—Moniliform tissue from Rhizoma Calami, Transverse section through the
nucleus-sheath; 0, oil-cells.

bium, and of the medullary rays are closely united to each other,
without intervening spaces—the wood, the bark, and the tissue

Fic. 152.—Transverse section from the sarcocarp of Fructus Aurantiorum; the
large, branched cells having sieve-like pores at their points of connection, for example,
at cr; q, branches of cells cut transversely; 5, imperfectly developed, large crystals of
oxalate. The dotted places represent the air-cavities; v, fibro-vascular bundle.

of the leaf possess an abundance of such, mostly triangular or
quadrangular, intercellular cavities. Aquatic plants, especially

AERATING SYSTEM. 237

their immersed portions, also show very large and numerous
intercellular spaces (Rhiz. Calami, Fig. 151, Carex leaves,
rhizomes of the Nymphea, Carex arenaria and Gratiola’).
The intercellular spaces produced by the disintegration of cells,
as a result of the tensions occurring through growth, may
assume the most varied forms, and finally by far exceed the
cells themselves in area.

Excellent examples of this are afforded by the thickened
margins of Siligua dulcis, the more central tissue of Fructus

x oe
ae
a

Seam,

9p rir
ws 4

' Fie. 153.—Transverse section through Caryopiytius, inner tissue; f, loose, branched
cells, interrupted by large air-cavities; ¢, d, fibro-vascular bundles; v, central fibro-

vascular bundle.
Aurantiorum (Fig. 152), the rhizome of Calamus and a portion
of the inner tissue of the clove (Fig. 153). a

Finally, wide spaces containing air (air-cavities, lacune) may
also be }found between the tissues, in consequence of the deli-
cately-walled parenchymatous tissue, especially the medulla,

not being able in its development to keep pace with the growth
in thickness of the other tissues, so that the cells of the former

1 Berg’s ‘‘ Atlas, plate xxii., Fig. 55.

238 PLANT ANATOMY.

gradually become destroyed. In this manner the hollow stems
(Graminee, Umbelliferee, Dulcamara) and roots (Rhiz. Grami-
nis) are produced. Some of the air-cavities in many reots and
leaves (aquatic grasses) may also be so produced; at least shreds
of the cell-membrane are often found on their margins. Lastly,
there may also be mentioned the cavities which are formed in
many drugs by the ultimate laceration either of lines of cells of
the medullary rays (Radix Bardane seu Lappe, Carline) or of

Fie. 154.—Surface view of the epidermis of a leaf of Mentha piperita; ep, undulated
epidermal cells; sp, stoma; 4d, oil glands (seen from above) with a group of crystals of
menthol m (Tschirch), Compare also Fig. 129.

the bark (especially of the bark-rays, Rad. Arnice, Fig. 182 6,
R. Levistici, R. Pimpinelle).

It is, however, more especially the lower side of leaves which
is abundantly aérated (transpirating tissue, Figs. 127, 129).
Here are also chiefly found the canals of exit, the stomata,
which are mostly located at the level of. the epidermis or are
only slightly depressed below the latter (Figs. 129 sp, 155), and

STOMATA, 239

which establish a direct connection of the intercellular spaces of
the inner tissue with the atmosphere. Besides on leaves,
stomata are also found on seed-vessels, young axial organs (stems),
and even on the walls of the ovary, but never on roots.

On leaves the stomata occur for the most part only on the
lower surface (Figs. 127, 129), or the latter is at least more
abundantly provided with them than the upper surface. This
applies, however, only to bifacial leaves, the centric leaves (see
page 210) having stomata on both sides (Fig. 128). In the
monocotyledons the stomata lie mostly in straight lines, the
cleft having the same direction in all (Fig. 156); in the dicoty-
ledons they are irregularly distributed (Fig. 154).

Fig. 155.—Vertical section of a stoma from a leaf of Mentha piperita; Ep, norris
¢, Cuticle; mer, leaf merenchyma; 8, guardian cells; E, eisodial opening; 0, opisthia
opening; A, respiratory cavity; n, lateral cells (Tschirch).

Their form and development ’ is very varied. Plants occur-
ring in dry climates have variously protected (depressed) stomata
(Figs. 63, 128).

There are distinguished on the stoma (Fig. 155) the guard-
ian cells (s), the central cleft (porus), and the eisodial (Z#), and
opisthial opening? (0). Beneath the guardian cells there is

. i i im’s Jahrbiicher f. wissensch. Botanik, iv., 1862, P.
Nigh Praedaisa Ibid., v., plates 34-36. Tschirch, “ Ueber ma
Beziehungen des anatom. Baues der Assimilationsorgane zu Klima sige
Standort,” Linnea, 1881. In the latter the entire literature relating
the stomata is given up to the year 1881.

* Compare Tschirch, loc, cit., p, 140.

240 PLANT ANATOMY.

always a more or less wide respiratory eavity (Figs. 155 A, 179

Fie. 156. Fie. 157,

Fias. 156, 157, 158.—Stomata from dicotyledons and monocotyledons on a surface sec-
tion, as seen from above.

3 @); and occasionally, beside the guardian cells, peculiarly formed

RECEPTACLES FOR EXCRETIONS., 241

lateral cells (Fig. 155 n). The number of the stomata varies
exceedingly. Upon a square millimeter of surface of a leaf
from ten to eight hundred of them may be present.

The distribution of the stomata, as also the shape and size of
the epidermal cells, is employed by Bell’ as a means of dis-
tinguishing tea, elder, willow and sloe, or black-thorn leaves.

The periderm, like the epidermis, is also occasionally traversed
by open gas passages. These are the lenticels ’ or bark-pores.
They are mostly produced beneath the stomata, and, as they
consist of roundish, suberized cells, they form small, sharply
circumscribed corky protuberances, which assume on the older
organs the function of the stomata, that is, they regulate the
interchange of gases. For some barks the lenticels are charac-
teristic (Rhamnus Frangulu, Solanum Duleamara). Such
corky growths occur also upon leaves (Lucalyptus, Fig. 128 hk,
Dammara).*

8. Receptacles for Hzcretions.

During the change of matter incidental to the life of the
plant, there are formed certain substances which, so far as our
knowledge extends, find no further application, and are therefore

termed excretions.‘
They may either be produced in more or less peculiarly shaped

cells or cellular passages (oil-cells, mucilage-cells, latex-tubes),

1 « Die Analyse der Nahrungsmittel,” i. (Berlin, 1882), p. 36.

° Diminutive of the Latin Jens, a lens, on account of their form. Re-
garding to the lenticels, compare Stahl, Botan. Zeit., 1873, No. 36.
Klebahn, Ber. d. Deutsch. botan. Ges., 1833, and ‘‘ Die Rindenporen.
Ein Beitrag zur Kenntniss des Baues und der Function der Lenticellen
und der analogen Rindenbildungen,” Jena, 1884, where the literature is

also given. ie ie
’ Bachmann, ‘“‘ Ueber Korkwucherungen auf Blattern,” Pringsheim’s

Jahrb., xii., p. 190. ; :
‘Regarding the limitation of the terms excretion and secretion,
compare the interesting investigation of Szyszylowicz, “ Ueber die
Sekretbehialterfder flichtigen Oele (im Pflanzenreich.” Denkwiirdig-
keiten der Krakauer Akademie, 1880 (Bot. Centralbl., viii., 1881, p. 259).
16

oo _ ‘In this place reference may also be made to the fact that the latex-

242 PLANT ANATOMY.

or secreted by special cells or aggregations of cells (organs of
secretion) in intercellular spaces (oil-passages).

The volatile oils and the resins come primarily under consid-
eration here.

ODN SOON AODae sO ooasoD ONT ATTA OOND IDO VONONDDOONDNIAUNODS
St TERY Ase SA PSS
e a cat ey Age By WEA SEES

= Joy d\
BY,
gtb oy Vee a yen a
Be Ved ROR VEE ER

be Fae El i =
MOHAN OU OOaH UeeOdNONUTNAONNNNMTUCNTOMRCICONNN mincnae acuity 10)
Fie. 159.—Cross section through the arillus of Myristica spec. (Bombay Mace); ep,

epidermis; gfb, vascular bundle; oe, oil-cells; g, a larger oil-space formed through the
confluence of several oil-cells,

The latex-tubes were also formerly considered as receptacles
for excretions, but it has recently been rendered probable, by

Fie. 160,—

.

A portion of the same cross section, more highly magnified; cut, cuticle
(Tschirch),

the discovery of points of connection with the assimilating tissue,
that they have some share in the transfer of substances, and

that their contents are not true excretions, although of more or
. Jess importance in the process of development.’

on

OIL-CELLS, 243

Nevertheless the latex-tubes should be treated of in this place..

Oil-cells. Volatile oil, in roundish or oval cells,' is found in
cloves (as much as 20 per cent), in cudebs (as much as 13 per
cent), in mace (up to 17 per cent, Figs. 85, 159, 160),’ in the
leaves and barks of the Lauraceex * (Cinnamomum, Camphora,
Fig. 161), and in the rhizomes of the Zingiberacee (Fig. 125) *
and of Acorus Calamus (Fig. 151 a). Volatile oil, mixed with

Fic. 161.—Transverse section through a sub-epidermal oil-cell in the surrounding _—
from a leaf of Cinnamomum Camphora; ep, epidermis (e, cuticle; cs, cuticle layer; cl,
cellulose layer), p, palisade tissue; 62, oil-cell; 4, drops of oil (Tschireh).

tubes are frequently connected anatomically with other conducting
organs, especially with sieve-tubes, as, for instance, in Rad. Taraxact
(Figs. 164 and 165). : ee

' All the cells dispersed through the fundamental tissue, and differing
in shape or contents (stune-cells, oil-, resin-, mucilage-cells, ico aoe
ete.), are also designated in general by the name of idioblasts ( fee
idvos peculiar, Glacros, germ, organic structure). Heinicher (Ber. d.
Deutsch. bot. Ges., ii.) has also recently found in the Papaveracez
idioblasts containing albumen. ise :

* These examples aay also be regarded at the same time as the max!~
mum amounts of volatile oil contained in plants.

3 Berg’s “ Atlas,” Plate xxxvi., Fig. 86.

4 Tbid., Plate xix,

244 PLANT ANATOMY.

other substances (coniine), occurs in peculiarly shaped lines of
cells in the fruit of the Coniuwm.

The volatile oil fills, or almost entirely fills, the individual
cells. ‘The same applies to the mucilage of the mucilage-cells ’
(see above), such as are met with, for example, in Rad. Athee,
in the Orchis tubers, in Cortex Cinnamomi, and in Cortex
Ulmi. As arule, the cells containing oil or mucilage are some-
what larger than the cells of the surrounding tissue (Orchis,
Cort. Cinnamomi,’ Figs. 160, 161, 186 ch).

As an example of the occurrence or absence of mucilage-
cavities under special conditions of growth (and therefore by no
means regularly) we may cite Laminaria hyperborea Foslie (L.
Cloustoni Edmonston).*

In addition to volatile oil and mucilage, crystals,‘ tannin ° and
resin also occur as constituents of true secreting cells. Héhnel °
has futhermore shown that kino,’ from the bark of Pferocarpus
Marsupium Roxburgh, also occurs in extended (100-500 1)
cells or tubes, united to form long bundles, which are not su-
berized. On the other hand, the oil-tubes of the Indian species —
of Andropogon (Andropogon Schenanthus L. was examined),
from which we obtain the fragrant volatile ‘‘ grass-oils,” * are
suberized.

The previously mentioned ‘glandular hairs, the terminal

1 Berg’s ‘‘ Atlas,” Plate xxiii., Fig. 57.

? Ibid., Plate xxxvi., Fig. 86 m.

*M. Foslie, ‘‘ Ueber die Laminarien Norwegens” (Christiania Viden sk.-
Selsk. Forhandl., 1884, No. 14), p. 47 of the reprint.

* The true crystals of many plants ‘(especially of monocotyledons), as
well as the cystoliths, are to be regarded as excretions.

5 We place the tannic matters here under the excretions, although, like
the constituents of the laticiferous juices, they probably still partici-
pate in the processes involving the transmutation of matter.

* “ Ueber die Art des Auftretens einiger vegetabilischer Rohstoffe in
den Stammpfianzen.” Sitzungsberichte der Wiener Akademie, 89 (1884),
January number.

* To what extent such special substances occurring in the plant are to
be considered as excretions still remains undetermined.

__ § Fliickiger, “‘ Pharmakognosie,” p. 157.

GLANDULAR HAIRS, 245

cell of which contains a secretion, must likewise be classed here.
These glandular hairs are either trichome formations (epidermal
glands, glandular hairs, tufts, scales, compare Figs. 129 éd, 154,
96), and as such have already been treated of, or they are pro-
duced by the protuberances of cells of the inner tissue, which
border on intercellular spaces (root-stock of Aspidium Filiz

mas).

Fic, 162,—From the fundamental tissue of the underground stem of Aspidium Filix

A,—Intercellular space from the younger tissue, in the middle showing a gland,
ith’a | xudation. t
seen from above and covered with’a green & se

B.—Longitudinal section through awall-cell (m), of the intercell
which the gland (g) begins to grow as a protuberance. : :
C.—Longitudinal section through the mother-cell (m) filled with starch, from.whieh
the gland, which is borne upon a stalk, projects as a daughter-cell into the ck cae
lar space. The daughter-cell has permitted ‘its green contents to appear upon
surface (as already represented from above, in A). : i
D.—A single gland more highly magnified, and freed from its covering, with the ex
_ ception of a delicate film (cc), by boiling with alcohol (Schacht),

246 PLANT ANATOMY,

The intercellular spaces in the fundamental tissue of the
root-stock and in the leaf parenchyma of Aspidium Filiz mas
(Fig. 162), which have been described by H. Schacht,’ are very
remarkably constructed. All of the boundary cells of these
cavities do not assume a special form, but some few of them
grow in a spherical manner into the hollow space, through a pro-
tuberance of the delicate wall in one or two places. The daughter-
cell, which is thus produced, becomes immediately bounded by
a transervse wall, and elevated upon a small stalk over the
mother-cell in a head-like form (the “Zottenkopf,” or tufted
head, of Hanstein). These small glandular cells thus remind

Af ‘a
. aaa = Z
i qo]

I= Fe IEA nes => sf
Ld y
— 4
Sere
me 2

Fie. 163.—Latex-tubes (I!) from Radix Taraxaci. Tangential longitudinal section
through the inner bark,

one of the more simple forms of the above-described oil-pro-
ducing trichomes of the Labiatew. In the fern-root the glands
in their terminal head-like cell at first contain protoplasm, in
which after a short time greenish oil-drops occur; these are
finally forced out upon the surface of the gland, and envelop it
as a thin greenish layer. This section consists for the most
part of the peculiar filicie acid, which, by longer preservation
_ under glycerin, crystallizes in long needles; volatile oil is want-
ing here, or is present in but very slight amounts. Such in-
_ tercellular glands have also been met with by one of us (F.) in

-_-'1 Pringsheim’s Jahrb. f. wissenschaftl. Bot., iii. (1863), p. 352.

LATEX-TUBES. 247

the (non-officinal) root-stock of Aspidium spinulosum Swartz;
in the other ferns of our region they are wanting.

A second form of secreting cells are the latex-tubes or latici-
ferous ducts.’ In most cases (especially when the tubes are
very long) they are produced, like the vessels, by the resorption
of the transverse walls of lines of cells lying over each other
(cell-fusion). In their simplest initial formation they are dis-
tinguished, however, from the neighboring parenchyma cells,
like the mucilage-cells and the oil-cells, only by their contents
and somewhat more considerable width, as for instance in jalap

or

Fie. 164, : Fie. 165,

Fig. 164.—Longitudinal section through the outer latex-zone of Radix Taraxaci, more
highly magnified; cr, sieve-tubes; /, latex-tubes. :

Fig, 165.—Longitudinal section through one of] the inner

azxaci, in which the tubes (7) are accom panied by sieve-tubes (er).

latex-zones of Rad. Tar-

1 With regard to latex-cells and latex-tubes, compare Meyen, ae Die
Secretionsorgane der Pflanzen,” Berlin, 1837.—Hanstein, “* Die Bilon
saftgefiisse,” Berlin, 1864.—Dippel, ‘* Entstehung der Milcheaftgefasse, ”
Rotterdam, 1865.—Vogl, ‘‘ Beitrage zur Kenntniss der Milchsaftorgane,
in Pringsheim’s Jahrb., v., 31.—David, “ Ueber die Milchzellen der
Euphorbien,” Breslau, 1872.—E. Schmidt, Botan. Zeit., 1882. Scott, In-

augural Dissertation,” Wiirzburg, 1831, etc.

eee

248 PLANT ANATOMY. |

tubers (Fig. 166) or in the Chinese galls (Fig. 91). In the
Cinchona barks, the latex-tubes are distinguished by their
considerable length, often also by a far greater diameter; in
other cases, as in Fructus Papaveris and in Carica! (Fig. 167),
they are developed as branched systems of canals. In this man-
ner abundantly branched latex-tubes traverse definite layers
of Radix Tarazaci (Figs. 163 to 165). Here the system of these
tubes, without regard to the overground portions, is only de-
veloped in the bark; in Lactuca virosa it extends also to the
central parenchyma of the stem, together with all other parts
of this plant.

Fic. 166,—Cells from Tuber Jalape: containing laticiferous juice.

One may accordingly speak of Jatex-cells and latex-vessels or
tubes. The former are true cells, and, when formed already in
the germinating plant, may often become very long, and, in-
deed, absolutely branched, without being divided by transverse -
walls, as for instance in the Urticacem, Euphorbiacee and
_ Asclepiacer. The milk-tubes, on the contrary, are lysigenic*
passages, that is, canals formed by the solution of the trans-
1 The latex-tubes of the fig aré so striking that by means of them one
May easily recognize an adulteration of coffee with ‘fig coffee”
_ * Ado a dissolution, and yevvae I produce.

LATEX-TUBES. 249

verse walls. With these are to be classed the tubes of the Cicho-
riacee (Taraxacum) and those of the Papaveracee and Cam-
panulacee,

If a resorption of the transverse walls does not take place, but
the short latex-cells lie in rows over each other, lactescent lines
of cells are produced, as for instance, in the Convolvulaces
(Tuber Jalape, Fig. 166); gutta percha (mostly from Dichopsis
Gutta Bentham et Hooker) also occurs in such lines of cells
containing laticiferous juice.

The contents of the latex-tubes is a mixture of very variable
composition.’ Besides the more commonly distributed substances,

v =
Fig. 167.—Latex-tubes of the fig; tangential section throngh the more central layer.
1, tubes; 4, rosette of crystals of calcium oxalate; v, vascular bundle.

such as salts (calcium malate), starch and protein bodies, very
many, if not all, laticiferous juices contain caoutchouc. Pe-
culiar bitter principles.and alkaloids (as in opium) also occur in
them, and some laticiferous juices are therefore medicinally
valuable (opium, euphorbium, lactucarium). In the latex of
species of Huphorbia is found the indifferent, crystallizable
euphorbon. The milky juices which here come under considera-
tion are white in their fresh condition.

Caoutchoue appears in the laticiferous juices in the form of
globules, which swell in volatile oils, and are not changed by

1 Compare among others S. Dietz, ‘‘ Beitrag zur Kenntniss des Milch-
saftes der Pflanzen,” Botan. Centralb., 1883, xvi., p. 133. :

250 PLANT ANATOMY.

dilute alkalies and acids, but are dissolved by chloroform and
carbon bisulphide. :.

With the latex-tubes which are formed through solution of
the intervening cell-walls are connected the receptacles for
resin and balsam, which are produced in a lysigenic manner
(see pages 170 and 248).

These appear as roundish spaces or cavities, filled with their
contents (formerly termed ‘‘interior glands”), and are pro-
duced in such a manner that those cells which occupy the
place of the subsequent receptacle become filled at an early
period with the respective secretion ; afterwards the membrane
of these cells containing the secretion disappear (Fig. 168, com-

Fig. 168.—Formation of an oil gland of Dictamnus Fraxinella; at the left is repre-
sented, below the epidermis, a group of small cells which become filled with drops of
oil, while the cells are in process of dissolution ; at the right, most of these cells have
already become dissolved, and in their place is produced a lysigenic intercellular space
containing a secretion (Rauter).

pare also the previous remarks in connection with the cell-
membrane). ‘These receptacles are not, like the resin-canals
(which will be considered directly), bordered by a circle of se-
creting cells; the cells inclosing them are not essentially dis-
tinguished from the tissue of their surroundings. With these
may be classed the oil receptacles of the Rutacew? (Ptelea,

™Compare Rauter, ‘‘ Zur Entwickelungsgeschichte einiger Trichomge-

bilde.” Sitzungsber. d. Wiener Akad., 1872, and Von Hohnel, ‘‘ Anatomi-.

sche Untersuchungen iiber einige Secretionsorgane der Pflanzen.”
Wiener Akademie, November, 1881.

INTERCELLULAR SECRETIONS, 251

Correa, Ruta, Dictamnus),’ of the leaves and fruits of Citrus,
and of Jaborandi leaves.

Thus, for example, in the very large oil-spaces in the rind of
the fruit of species of Citrus, a solution of the cell-walls is dis-
tinctly perceptible.* This is, perhaps, still more the case in the
trunks of Copaifera,? in which the balsam passages attain
an enormous development. These trees contain the copaiva-
éalsam in canals which are as much as an inch in diameter, and
which often traverse the entire trunk, so that a single one, after
being bored, is capable of yielding balsam by the pound. In
the so-called ‘‘ gummosis,” the membranes become converted
into gum (compare page 166, Fig. 78).

In the Sterculiacez, lysigenic (protogenic, page 263) gum-pas-
_ Sages are found.‘

The lysigenic passages, like the schizogenic, which will be
described directly, may be either dermatogenic, that is, produced
by the participation of epidermal cells (Citrus, Dictamnus,
Amorpha), or they may be formed under the epidermis, deeply
in the interior (interior glands in a more restricted sense).

The secreting space of the lysigenic passages is always com-
pletely closed.

The intercellular receptacles for secretions, or the so-
called oil-passages and balsam-passages, belong neither to the
true cells, nor to the tubes produced through the fusion of cells,
nor to the passages of lysigenic origin.

The formation of these receptacles, which are found in the
Myrtaceer (Hucalyptus, Figs. 127,128, Myrtus, Eugenia, Pimen-
ta), in the Leguminose (Amorpha, Hymenea, Trachylobium),
in the Umbellifere, Composite, and Conifere, in Ovalis,
Lysimachia and Myrsine,* admits, in the families of the Um-

1 Here also in the interior of hairs, Sachs, ‘‘ Lehrbuch,” p. 93. Mar-
tinet, loc. cit, De Bary, loc. cit. Fig. 22.

?Sachs, ‘‘ Lehrbuch der Botanik,” 1874, p. 92,

* Karsten, Botan. Zeitung, 1857, p. 316.

4Trécul, Compt. rend., 1862, p. 315.—Ledig, Botan. Centralb., 1881,
vi., p. 387.

5 Von Héhnel, in the investigation cited on page 250.

252 PLANT ANATOMY.

belliferee, Composite and Conifers ' (which are very rich in such
examples) of being traced back to the production, extension, and
prolongation of intercellular spaces. Very frequently four cells.

Fie. 169. Fie. 170. Fie. 171.

Figs. 169-171.—Four boundary cells, gggg, in Fig. 176 receding from each other; in
Fig. 171 the intercellular passage so produced exerts a pressure upon the boundary
cells,

Fig. 169, gggq) recede from each other in the region where they
meet together.

The boundary cells (y), which in the beginning (Fig. 170)
still project with their conyex walls into the resin-passage,

Fia. 172. Fia. 173.

Fias. 172-173.—Beginning of the depression of the boundary cells, which in Fig. 173
are strongly compressed in a radial direction (Miller).

recede (Fig. 171), and are more and more depressed in a radial
direction (Figs. 172, 173). At the same time, in the boundary
cells, and frequently also in the farther surroundings of the

' Compare also Fig. 106.

INTERCELLULAR SECRETIONS. 253

passages, a new formation of cells takes place by the division of
the older cells (Figs. 174, 175), so that finally the fully devel-

Fie. 174. Fie. 175.

Fig. 174.—Beginning of the new formation of cells in the boundary cells (Miller),

Fie. 175,—Around the intercellular passage, which has become extended as an oil-
‘space, a layer of tabular daughter-cells (secerning cells) has been formed. Transverse
section through a branch of the root of Inula Helenium (Miller).

oped resin-passage or balsam-passage is located in a special

Fig. 176. : Fie, 177.

Fie. 176.—Longitudinal section (a) and transverse section ((b) through a balsam-pas-
sage (oil-space) of Rad. Inula. Be : :

Fie. 177.—Central portion of an oil-space from Rhizoma] Arnice, on & longitudinal
section; 0, oil-space; t, daughter-cells which have not yet become depressed.

254 PLANT ANATOMY.

form of tissue (Fig. 176). From these cells surrounding the
intercellular balsam-passage (secerning cells, epithelium) the
volatile oil, and the resin dissolved therein, penetrate through
their walls into the passage.

A canal which has been formed in the manner just described,
by the parting of cells from each other, is termed schizogenic.'
In the larch, in addition to resin receptacles of this kind,
lysigenic passages are also found (page 248).

A ramification of those passages which more closely concern us
here is not perceptible on a longitudinal section through them,

Fic. 178.—Longitudinal section from the bark of Radix Sumbul (Ferula seu Eury-
—angium Sumbul); r, medullary rays; 1, phloém; b, oil-passage (the parenchyma lying
beneath it being visible). :

so that they do not represent a vascular system, like the latex-
tubes of many plants (e. g., Figs. 163-165). The more simple
form of the resin-passages is in harmony with the manner of their
formation, although this does not exclude the passages from oc-
casionally attaining a considerable length (Fig. 176), as in the
_ sumbul root (Fig. 178), in Rhizoma Imperatorie, or, as already

_ mentioned, in Copaifera, and presumably also in Dipterocarpus.*

| Byitw I cleave, and yévos production.
___ * Fliickiger, “‘ Pharmakognosie,” p. 86.

SCHIZOGENIC PASSAGES. 255

The intercullar resin-receptacles are widely distributed in the
Coniferw; here they are found not only in the bark (Fig. 181),
but also in the wood (Fig. 180), in the scales of the cones,’ and,
indeed, the leaves mostly contain two of them (Fig. 179) on the
lower surface. It is only in a few Conifer (Zazrus) that they
are entirely wanting.

Copal is also produced in schizogenic secretion-receptacles,
in the Trachylobium as well as in the Hymenea, as Héhnel has
shown.

Fiq, 179.~Transverse section through the oil-canal of a coniferous leaf: dc, oil
canal; s, nucleus-sheath (consisting of bast-cells) surrounding the same; sp, stoma; a,

respiratory cavity; ep, epidermis (Tschirch).

_ Although a longitudinal section through the root structures
of the Composite and Umbellifere does not reveal either a
regular arrangement* ora connection of the balsam-passages —

1 Hanausek, ‘‘ Ueber die Harzginge in den Zapfenschuppen einiger Co- —
niferen.” Jahresbericht der, etc., Handelsschule in Krems, 1880,
? With regard to the distribution of the resin-passages, compare Han-
ausek, ‘ Zur Lage der Harzgange,” Irmischia, ii. (1882), Nos. 3, 4, and

HAG Utiler, Tinie ee es

256 PLANT ANATOMY.

Pex DiH!
|

Cc

Fig. 180.—Transverse section from the wood of Pinus silvestris; jg,

1 annual ring; fh,
spring wood; hh, autumn wood; hg, resin-canal; c, secreting cells; ¢, pits; m, medul-
lary rays.

_ Fig. 181.—Transverse section through the bark of Pistacia Lentiscus; st, plate of

stone-cells with inclosed crystal-cells, the medullary rays sclerotized. Resin-passages.

Oele, Gummi und Gummiharze und die Stellung der Secretbehalter im
Pflanzenkérper,” in Pringsheim’s Jahrb. fiir wissenschaftl. Botanik, v.
(1867), p. 380. H. Mayr, * Entstehung und Vertheilung der Sekretions-
_ organe der Fichte und Larche,” Botan. Centralblatt, xx. (1884), 278

.

SCHIZOGENIC PASSAGES. 257

among themselves; yet aconformity to some law in the position
of these receptacles is sometimes apparent upon a_trans-
verse ‘section. ‘lhe root-stock of Arnica, for example, shows
a large balsam-passage (Fig. 182) before each fibro-vascu-.
lar bundle. A similar arrangement may be seen to exist in

aie

Fia. 182.—Portion of a transverse section from the underground stem (rhizome) of
Arnica montana. Before each xylem-ray (wood-bundle) 1, there is a very large oil
space; 0, b, cavities formed by the laceration of the fundamental tissue; a, epidermis
_ of the root (epiblema); m, medulla, The cells of the epidermis are spirally striped; in

the surroundings of the oil-spaces are drops of oil which have escaped. Compare also
Berg's “ Atlas,” Plates 8, 9, 10. ; aii

Radix Angelice and R. Levistici, in Rhizoma Imperatorie and

others, although, in the course of development, which is not =

258 PLANT ANATOMY.

always perfectly uniform, they often become disturbed in con-
sequence of the laceration of the individual tissues.

Hand in hand with the organic transformations just described,
there occur, in the surroundings of the passages, certain chemi-
cal processes, to which the resins, volatile oils and varieties of
mucilage owe their origin and also the special form which
enables them to pass into the intercellular spaces. The resins,
namely, are either dissolved involatile oils, as ** balsams” or ‘‘ tur-
pentines,” or they are emulsified by mucilage (gum). It is only
in this form that they are capable of passing through the cell-
walls into the passages formed for their reception. Although
the manner of formation of the cells and tissues’ which
have here been considered may appear quite clear, yet the
chemical side of these processes has so far not been elucidated.
In many cases, resin and volatile oil appear to be produced from
amylum. If this may be accepted with some degree of probabil-
ity, the equally just supposition is forcibly presented that under
certain conditions cellulose, which agrees with amylum in its
composition, is also capable of undergoing the same transforma-
tion. Asa matter of fact, this is also the case with the lysigenic
canals, which have previously been considered (compare page
248).

According to Frank’s investigations,’ it would appear as if
the oil-tubes or stripes, vitte@,* which are so characteristic for
many of the umbelliferous fruits, first became forced asunder
through the volatile oil which makes its appearance in them,
while the balsam-passages of umbelliferous roots present the
development illustrated by Figs. 169 to 175. But in many of
these fruits the oil-tubes also show remnants of transverse walls
(Fig. 183) which presumably indicate a solution of original
boundary cells. The effloresced appearance of the tissues which
surround the oil-tubes in Fructus Carvi, Fructus Feniculi,

1 Especially described by Miiller, loc. cit., p. 387.—Thomas, Ibid.,
_v., p. 48.—See also Frank, ‘‘ Beitrage zur Pflanzenphysiologie,” Leipzig,
1868, pp. 120, 123.
_ ** Beitr. z. Pflanzenphysiologie,” p. 128.
sss 8 Fig. 94 0.—Berg’s “ Atlas,” Plates xli., xlii., xliii.

SCHIZOGENIC PASSAGES. 259

etc., also supports this view. It has, however, recently been
shown that the oil-tubes are produced in the same manner as
the resin-receptacles of roots,’ and are thus of schizogenic
origin.

As an exception, in Fructus Conti oil-tubes* are not found
at the period of ripening (as has previously been already men-.
tioned), but rather a connected layer of cells (Fig. 184), im
which is located the volatile oil and coniine. When observed:

Fic. 183.—Longitudinal section{through an oil-passage from Fructus Foeniculi, with
transverse walls; s, dark-brown efflorescing cork-like tissue; a, albumen of the seed.

on a longitudinal section (Fig. 185), this layer represents the
cells superposed in a compartment-like form; if their transverse

1 Lange, ‘“‘ Uber die Entwickelung der Oelbehalter bei den Umbelli-
feren,” K6nigsberg, 1884. Dissertation.—Compare also E. Bartsch,
‘‘ Beitrage zur Anatomie und Entwickelungsgeschichte der Umbelli-
ferenfriichte.” Inaugural Dissertation, Breslau, 1882.

2 In young ovaries they are present as an initial formation, but do not

become developed.

260 . PLANT ANATOMY.

walls were to disappear, the figure of an oil-passage produced
by resorption would be obtained. Such a solution of the trans-
verse walls does not occur, however, in Fructus Conii.

The spaces which have just been considered contain either
volatile oil (Myrtaceze), which in drugs is often already more or
less resinified, or a mixture of volatile oil and resin, or finally
resin itself. :

Resin, which is free from volatile oil, is also met with in
Lignum Guaiaci and in Lig. Quassie in the form of brittle,

Fic. 184.—Transverse section through Fructus Conii; a, albumen of the seed; }, ©
embryo; ce, commissural surface; e, epidermis; m, tissue of the fruit casing; ¢’, inner-
most layer of the latter; ¢, layer of cells containing volatile oil and coniine; o, vitte; v,
ribs (costz), traversed by fibro-vaScular bundles. .

shapeless masses. In these two plants the resin first makes its
appearance at a later period (at an advanced age) in ordinary
wood-cells and vessels of the heart-wood; it thus shows a deport-
ment similar to that of physiological gum (see page 166), with
which it also appears to be chemically related (see also page
264).
Dees ‘a a microscopical examination, the resin and oil, especially
when they occur mixed with each other as a balsam, escape from —

SCHIZOGENIC PASSAGES. 261

these depositories in the form of small, strongly refracting
drops, which are more frequently yellowish or brownish than:
colorless, and are clearly miscible with alcohol, or at least with
absolute alcohol, ether or benzol, and with fixed or ‘volatile
oils.

A mixture of gum, resin, and volatile oils (gum-resin), as in
asafetida, galbanum, and ammoniacum, or even gum or muci-
lage alone (Cycadeex),’ is also found in schizogenic secretion-

receptacles.
However, should any of the material occurring in cells or in

Fie. 185,—Longitudinal section through the coniine layer, t, of Fig. 184, The letters
have the same signification as in Fig. 184.

intercellular spaces be found to be indifferent towards the above
solvents, it does not necessarily follow that resin is absent, since
the older lumps of resin dissolve only with difficulty.

The resins are colored red by tincture of alkanet. The
reagent of Unverdorben and Franchimont (an aqueous solution
of acetate of copper) colors the drops of resin, after several
days’ maceration in the liquid, emerald-green.

The resins are not, however, confined to these receptacles, as

1G, Kraus, in Pringsheim’s Jahrb, f. wissensch. Botan., iv., p. 305,
Plates xxi. and xxiii.

262 PLANT ANATOMY.

is already evident from the description of the latter. In young
cells, where the resin is first formed only in small amount, it is
destitute of any special color; the same is the case where the
resin forms semi-liquid granules, especially in those plants
where it occurs emulsified as a constituent of laticiferous juices,
for example, in Tuber Jalape. ‘This extremely fine division
and liquefaction of the resins is promoted in the laticiferous
juices of the Umbellifer by the volatile oil which they contain.
Under such conditions the resin may be recognized by its
tendency to absorb coloring matters. Iodine solution, or pre-
ferably aniline colors dissolved in water, carmine, etc., when
carefully added in corresponding amount, are very useful for
this purpose. To be sure, this does not always prove that the
colored bodies are resins, but the coloration, nevertheless, affords
good points of discrimination.

The schizogenic secretion-receptacles are generally closed.
There are, however, cases where the secreting space is open, and
communicates with intercellular spaces of the parenchyma,
which likewise sometimes contain a secretion (Ozalis floribunda,
Peganum Harmala, Lysimachia Ephemerum).

With regard to their morphology and the nature of their -
development, the following varieties of secretion-receptacles
should, therefore, be maintained distinct:

I. True Cells.

(a) Isolated in the interior of the tissues, and on all sides in

connection with the other elements of the tissue.

1. Containing oil: Macis, Laurus, Cortex Angosture,
roots of the Zingiberacee, Acorus Calamus, Cinna-
mon bark.

2. Containing mucilage: Cinnamon-bark, Elm-bark,
Althea root (filling entire tissues in Chondrus).

3. Containing laticiferous juice, simple: galls, from
species of Rhus, from Eastern Asia : Euphorbiacee.

4, Containing crystals, especially in monocotyledons.

5. Containing tannin.

@: ‘United in a band of cells. Fructus Conti, Tuber Jalape,
_ Dichopsis Gutta.

CLASSIFICATION OF RECEPTACLES. 263

(c) As glandular heads of hairs.
1. Upon the epidermis, glandular hairs of the Labiate.
Kamala, Glandule Lupuli (colleters).
2. Projecting into intercellular spaces (Aspidium Filiz
mas).
II. Secretion-receptacles produced through the fusion of cells:

(a) Of rows of cells: latex-tubes (Papaveracee, Cichoriacex,
Campanulacez).

(6) Of homogeneous aggregations of cells: lysigenic oil- and
balsam-passages (Rutacezx, including the Aurantiez).

(c) Of entire, and also dissimilar portions of tissue: gum-
glands.

Ii]. True intercellular spaces :

(2) Containing oil or resin, schizogenic balsam-passages (in
the Conifere, Umbellifere,.. Myrtacee, Leguminose,
Hypericinez).

(4) Containing gum-resin (the roots of some Umbellifere).

(c) Containing mucilage, schizogenic mucilage-passages
(Cycadeze).

(d) Containing laticiferous juice (Alisma Plantago).

The secretion receptacles may be profogenic, that is, they may
be formed already in quite young tissues, or they may be
hysterogenic, that is produced at a later period in old and com-
pletely developed tissue.

In transmitted light, many leaves appear finely punctate in
consequence of the presence of resin cells, resin cavities, and
crystal cells. Bokorny has utilized these ‘ pellucid points ”
diagnostically.?. The pellucid points are caused by resin cells in
the Lauracew, Piperacee, Meliacex, Sapindacee, Canellaces,
Anonacex and others, by resin-cavities in the Myrsinee, Myrta-
cee, Rutacee, and Hypericinex. :

1 The size of the resin-canals possesses diagnostic value in distinguish-
ing Rad. Levistici and Angelice. In the former they have the same di-
ameter as the vessels; in Angelica, on the contrary, they are consider-

ably wider.
2 Die durchsichtigen Puncte der Blatter in anatomischer und syste-
matischer Beziehung.” Flora, 1882.

PATHOLOGICAL FORMATIONS.

The morphological and anatomico-physiological relations of
the organs formed in the normal vital processes of plants having
been considered so far as they relate to our purpose, some other
phenomena may still be mentioned which are produced by dis-
turbances of the normal] processes.

When a part of a plant is wounded by the human hand or by
an animal, it is capable of repairing the injury. The most
usual form of protection is the formation of cork (protective
cork) on the wounded place. Figure 109 a shows, for example,
how on a fruit of the Vanilla, which has been wounded by an
insect, cork has been produced around the wounded place,
whereby access of air is excluded from the inner tissues. Cork»
is the ordinary form of protection for delicate organs, but
naturally, is only met with in such places where cells occur
which are still delicate and readily suberized. If, on the con-
trary, the stem of a dicotyledonous woody plant is wounded
down to the wood, the plant selects another means of protec-
tion, since cells here become exposed which can no longer be-
come suberized. Accordingly, in all the cells of the wood
which border on the wounded place, gum (wound-gum) is pro-
duced as an exudation of the thick membranes, which gradually
fills the cell cavity and thus directly closes the wound. The
masses of gum and resin which occur in the dead heart-wood of

the officinal woods ( page 260) are probably such protective gum.

During this process, in those portions of the bark which are
still capable of development, the plant endeavors to close the
_ wound from both sides. There are thus produced, by very

—_— = in the _—— cambial zone, om, inflated

GALLS. 265

cushions, or excrescences, which gradually constrict the wounded
place more and more, and, indeed, may finally entirely close it.

Such enlargements of the tissue as are produced by outward

influences are termed hypertrophies. hey occur not only in the
form of excrescences, but also as variously shaped malformations,
which should not be classed with the excrescences without
further distinction. While, namely, the latter are to be regarded
as a reaction produced in consequence of a single wound, the
other anomalous formations, quite generally designated by the
name of galls or cecidie, are of service to the insect or para-
sitie plant which produces the wound. Hence there may be
distinguished according to the respective organism, fungus-
galls (mycocecidie)—for of all plants only the parasitic fungi
produce such formations—and insect-galls (zoocecidix.)
Whether, however, a plant or an insect is the cause of these
formations, the symptoms are always the same. By a vigorous,
and often greatly increased formation and division of cells, the
affected part of the plant, which always consists of cells which
are still capable of development, is very considerably enlarged,
and manifold and often very strange malformations are pro-
duced. An active conveyance of sap affords abundant nourish-
ment, which is often so great that not only are the requirements
fulfilled for the new formation of cellular tissue, but numerous
surplus products (for instance, starch) also accumulate in the
cells. Moreover, peculiar bodies (such as tannic matters) are
also frequently formed in the cells of the gall, which are either
wanting in the other tissues of the plant, or are contained in
them in very much smaller amount. Upon such an activity of
development, which is increased and qualitatively changed
through irritation, essentially depends the formation of galls.

In the fungus-galls, the hyphx of the fungus, nourished by
the richly supplied -cells, penetrate the intumescence. In the
case of the insect-galls, the interior, which is mostly hollow,
serves as an abode for the insect; it there passes through its —
entire course of development, from the egg to the perfect insect
(Cynips galls of the Oak), and even through several generations.

Since the galls seriously injure the plant only when they are

266 PATHOLOGICAL FORMATIONS.

produced in quantitatively very considerable amounts, a case of
symbiosis ' (cohabitation) is thus presented here, of beings be-
longing to two different series, though, since the one lives chiefly
at the expense of the other, this borders closely on parasitism.
These formations are very remarkable from the fact that a plant
does service for an insect and constructs for the latter its
dwelling place.

‘Since insects of different classes (Hemiptera, Diptera, Hy-
menoptera) participate in the production of insect-galls—and
only these interest us here— it is impossible to make any state-
ments of general application regarding these gall formations,
which occur upon plants of all the families of phanerogams.’
Their shape, like the nature of the irritation which produces
them, is extremely variable.

Only those galls which are rich in tannin are of technical
importance, and these are also our best sources of tannin. The
oaks, especially, furnish many valuable galls.*

The galls of Asia Minor (or galls in a general sense) are pro-
duced by the puncture of the ovipositor of the female insect of
Oynips galle tinctorie Olivier (a Hymenoptera) in the young
shoots of Quercus lusitanica Lamarck. The female insect, de-
veloped in the hypertrophic tissue from the egg deposited
therein, afterwards bores for itself a passage and escapes from
the gall, by which it was sheltered during one of its phases of
life.

The so-called Chinese and Japanese galls, on the contrary,
are produced by the female Aphis sinensis (a Hemiptera) in the
younger shoots and leaf-stalks of Rhus semialata Murray. In
these galls, which are mostly very large, the numerously intro-
duced eggs become developed as plant-lice, pass through succes-

1 Sur with, and zevr to live.
*Compare especially, Frank, ‘‘ Handbuch der Pflanzenkrankheiten.”
_ ®The galls which are employed technically and pharmaceutically
have been very excellently described by Hartwich (Arch. d. Pharm.,
21, 1883, p. 820); compare also Wiesner, ‘‘ Rohstoffe des Pflanzenreiches,”
Vienna, 1873, pp. 846, with numerous illustrations.

GALLS. 267

sive generations, and finally escape. In the commercial galls,
the small insects are still frequently found which have been
killed by scalding or have otherwise been destroyed.

MICRO-CHEMICAL REAGENTS.

In the course of the preceding representation the chemical
detection of one or another substance has already been alluded
to, and, indeed, the treatment of microscopic sections with
suitable reagents affords many valuable disclosures. As in all
other cases, definite answers are obtainable, when systematic
and accurately formulated questions are propounded. For
this purpose, chemical reagents! are of service, among which
the following may be designated as especially important:

1. Chromic Acid (free from sulphuric acid) dissolved in 100
parts of water. This is adapted in general for the purpose of
loosening composite, thickened cell-walls and constituent bodies,
whereby the finer relations of structure are very often made
evident, since chromic acid is also capable of clearing up
darkly colored cell-walls, and, on the other hand separates the
layers and finer membranes, thus bringing them more clearly to
view. By means of this acid, starch granules are completely
separated into lamine, the strata of the Cinchona bast-fibres
(page 155, Fig. 73) separated from each other and lignified mem-
branes @issolved, while suberized membranes are rendered more
clear.

When employed in a more concentrated form, or allowed to
act for a longer time, chromic acid destroys the cell-walls. Its
application, therefore, requires continual observation of the sec-
tidns which are treated therewith, in order to completely survey
the result of the phenomena.

‘Compare Poulsen’s ‘“ Botanische Microchemie,” Cassel, 1881; and
Behrens’ ‘‘ Hilfsbuch zur Ausfihrung microscopischer Untersuchun-
_ gen,” Braunschweig, 1883. The American editions of these two works
are noticed on page 49; in both of them are contained numerous refer-

| _ ences to the literature of the subject.

MICRO-CHEMICAL REAGENTS. 269

2. Hydrochloric Acid of the specific gravity 1.110 acts much
less energetically upon the cell-walls; yet, without causing these
to swell in a disturbing manner, it seizes upon many of the con-
stituent substances and thereby permits the structure of the
tissue to be more clearly recognized (a stronger acid effects
tumefaction). Calcium oxalate (page 129) is readily dissolved
by hydrochloric acid.

3. Sulphurie Acid.

The dilute acid (specific gravity 1.110) causes starch and the
membranes to swell. Cellulose is converted by it into amyloid
(page 159).

Concentrated sulphuric acid (specific gravity 1.836) dissolve
the membranes and their contents; only the cuticle, cork,
the nucleus sheaths (page 202), intercellular substance, and the
oil drops contained in the cell resist its action.

In the phloroglucin reaction (page 161), sulphuric acid can be
applied in place of hydrochloric acid. When mixed with indol !
the former is a good reagent for lignified membranes.

4, Nitrie Acid of the specific gravity 1.180, either alone or
after the addition of ammonia, colors protein substances, as also
the middle lamella, yellow. Héhnel employs it for the detec-
tion of suberin (cerin reaction).

Since nitric acid dissolves starch as well as sulphuric and
hydrochloric acids, it can likewise be employed for clearing up
tissues which are rich in starch.

Nitric acid alone, or, still better, a mixture of nitric acid and
potassium chlorate (Schultze’s maceration), is the best agent for
isolating the individual element of tissue. The boiling acid, to
which small crystals of potassium chlorate are gradually added,
dissolves the middle lamella. This method of procedure is par-
ticularly applicable to the examination of vegetable powders
(Cinnamon, Cinchona, ete.); dark membranes at the same time
become bleached thereby.

When treated in this manner, boiled scraps of drugs are re-

1 A colorless crystalline principle of the composition CsH;N, ob-
tained by the action of reducing agents on the blue coloring principle of
indigo. F. B, P.

270 MICRO-CHEMICAL REAGENTS.

solved by the pressure of the glass cover upon the slide into the
separate elements (Fig. 186), which may then be conveniently
further examined. It is, however, to be considered that the
reagent produces a swelling of the membranes, as also a solution
of the bodies contained therein. It is likewise to be observed
that Schultze’s maceration dissolves the lignin from the lignified
membranes, so that the latter then show the cellulose reaction.

Before the objects that have been thus treated are brought

Fig. 186.—Isolated elements of cinnamon bark obtained by maceration with Schultze’s
liquid. b, bast-cells; sc, stone-cells (sclereids); p, parenchyma; sch, mucilage cells;
st, starch granules of the cinnamon (Tschirch).

under the microscope, it is necessary to thoroughly wash out
the reagent.

5. Acetic Acid of the specific gravity 1.040 often clears up in
a remarkable manner such sections as have previously been
treated with alkalies. Since calcium oxalate is insoluble in
acetic acid, the latter may also be employed as a confirmative
_ test when the recognition of this salt is in question (compare —
page 133). This acid is likewise of service in the examination

MICROCHEMICAL REAGENTS. 271

of protoplasm, since it causes the nucleus of the cell to appear
sharply defined.

6. Neutral Acetate of Copper dissolved in twenty parts of
water. If tissues containing resin are allowed to remain for
some days in this solution, the metal combines with the resin to
form green clots.

”. Tannic Acid is always dissolved freshly just before use in
twenty parts of water, and employed in the search for alkaloids.

By moistening thin sections with water, to which a slight
trace of acetic acid has been added, a concentrated liquid extract
of the substance is prepared, which is tested by the gradual
addition of a few drops of the solution of tannic acid upon the
slide. Ifa turbidity is produced, it may be due to the presence
of alkaloids, but may also be caused by the so-called bitter
principles, or by albumen.

8. Solution of Soda of the specific gravity 1.160, containing
15 per cent of sodium hydroxide. Instead of this solution, an
equally strong alcoholic solution may serve for many purposes.
The corresponding solutions of potassium hydroxide have the
same action.

The caustic alkali causes the cell-walls to swell, and dissolves
many of the constituent substances, especially coloring matters,
whereby the sections are rendered very much clearer; from
lignified membranes the lignin is extracted by means of warm
solutions of caustic potassa or soda. In many cases the treat-
ment of the tissue with alkali permits of the clearer recognition
of the relations of the strata. The protein crystalloids reveal
their organic structure by swelling, whereby their plain surfaces
become for the most part rounded and the angles changed.
Many yellow coloring substances (chrysophan in Rhubarb,
frangulin in Cortex Frangule, chrysarobin) become red by
alkalies, a reaction for which lime-water is well adapted.

9. Sodium Hydroxide (solid caustic soda) or potassium
hydroxide may be conveniently kept and employed in the form
of powder, when the amount to be applied does not require to
‘be more accurately measured. |

10. Ammonia Water, of the specific gravity 0.960, is often

R72 MICRO-CHEMICAL REAGENTS.

preferable to potassa and soda, since the two latter frequently pro-
duce an altogether too energetic swelling, which detracts from
the clearness of the outlines. The jelly which is formed by the
action of the alkalies upon starch also interferes very much.
By the application of ammonia neither of these objectionable
results are produced, while its power of dissolving coloring sub-
stances is not less in extent. Starch suffers no change by the
action of ammonia.

Ammonia water, still further diluted, is adapted for softening
dried plants and many drugs which it is desired to examine
more thoroughly. Sections which are treated with nitric acid,
and afterward washed and moistened with ammonia, admit of
the sharp recognition of protein substances and of the middle
lamella by means of their yellow color (xantho-protein reaction).

11. Alkaline Solution of Tartrate of Copper. The solu-
tion of the tartrate of copper and sodium in caustic alkali, the
so-called ‘‘ Fehling’s solution,” is not convenient for micro-
chemical purposes. In place of it, the following method of
procedure may be recommended: A solution of 3 parts of
sulphate of copper (blue vitriol), free from iron, in 30 parts of
hot water is mixed with a solution of 7 parts of Rochelle salt
(potassio-sodium tartrate) in 20 parts of hot water, the resulting
precipitate collected and dried. When used, alittle of this pre-
cipitate is brought upon the object-glass, a small fragment of
caustic soda added, and thereupon a few drops of water until a
clear solution is produced, or this may also be effected by the use
of the least possibly quantity of the solution of caustic alkali (No.
8). The section is then moistened therewith. This alkaline
solution of the tartrate of copper is useful in testing for sugar,
since uncrystallizable, so-called fruit sugar (page 142) imme-
diately separates therefrom reddish-yellow, hydrated cuprous
oxide. This also occurs very soon by theaid of a gentle heat
when grape sugar is present, but not even by boiling in the case

of cane sugar (or mannite). Dextrin is also capable of reduc-

, _ ing the tartrate of copper with the aid of heat.

The varieties of gum and mucilage effect no reduction in the

es alkaline solution of tartrate of copper.

MICRO-CHEMICAL REAGENTS. 273

Sachs proceeds in the following manner in the reaction for
grape sugar. He places the (thick) longitudinal sections for
some minutes in a solution of sulphate of copper (1 part of
sulphate of copper and 4 parts of water), then washes them with
water, and brings them into a boiling solution of caustic potassa
(one part of the solution No. 8 and two parts of water). Cells
containing grape sugar then appear filled with a reddish-yellow,
granular precipitate (Cu,O). In this reaction it is necessary to
accurately proportion the time that the section remains in the
liquid, the amount of washing, thickness of the section, etc.,
for which some experience is required.

The alkaline tartrate of copper imparts to the albuminous
substances deposited in the parenchyma a violet color, in con-
sequence of the formation of compounds of copper with the pro-
tein substances, as was made known in 1872 by Ritthausen.

12. Ammoniacal Solution of Oxide of Copper is obtained
by shaking copper turnings with ammonia water of the specific ©
gravity 0.960, with the addition of very little ammonium
chloride. The ammonical oxide of copper, when prepared in an-
other manner, has a different action in some cases. This liquid
is the only solvent for cellulose. It is to be observed, however,
that its action upon the cell-walls is very different, according to
their thickness and purity, and that many, as for instance the
hyphe of fungi and cork, are not attacked by it at all, or at least
not without previous boiling with caustic alkali or with potas-
sium chlorate and hydrochloric acid. The action of the ammoni-
acal oxide of copper does not occur immediately.

The ammoniacal oxide of copper is only fit for use when it dis-
solves cotton in the course of a few hours, It is expedient to
protect it from the action of light, and not to keep it for a very
long time.

13. Glycerin of the specific gravity 1. 225 is of very general
use as a clearing agent; with a higher degree of concentration —
its power of abstracting water also comesinto consideration. In
the examination of such constituent substances as would dis-
solve quickly in water (aleurone, tannic acid), concentrated
glycerin is — useful, since its solvent power is ae gradually ©

274 MICRO-CHEMICAL REAGENTS.

exerted. Thus under glycerin the gradual swelling of mem-
branes affording mucilage and the disintegration of tissues con-
taining oil may also be conveniently observed, and by the addi-
tion of water accelerated at will.

14. Absolute Alcohol is of service, for example, in rendering
mucilage visible, which would be washed away by water or would
form a clear mixture with glycerin. The volatile oils and
resins are dissolved by alcohol.

Fats and wax are but slightly soluble in cold alcohol, but, for
the most part, can be brought into solution by boiling.

Protoplasm is killed and hardened by alcohol. Since the lat-
ter has adehydrating action, its application, either alone or with
the addition of glycerin, effects a separation of the protoplasm
from the membrane.

The tissues can be freed of air by means of alcohol, since the
latter more readily penetrates into the intercellular spaces, and
is also capable of absorbing more air than water.

By placing the respective organs in alcohol, inulin (page 124)
as also hesperidin (page 143) are obtained in sphero-crystals.
Indeed, by means of alcohol, even asparagin and sugar may be
made to crystallize in organs which are particularly rich therein.

15. Aleohol of 85 per cent by weight, having a specific
gravity of about 0.830, the ordinary Spirit of Wine, accomplishes
in most cases the some purpose as absolute alcohol.

16. Aleohol of 60 per cent, besides dissolving the resins,
also dissolves the different sugars in considerable amount.

17. Ether is applied for the removal of solid and liquid fats,
whereby resins and yolatile oils are dissolved at the same time.

18. Benzol (C,H,) serves the same purpose as ether, but,
since it boils at 80° C., it admits of gentle warming (with care!),
which is often very useful. The same may be said of chloro-
form.

19. Chloroform. Resins, fats, wax, and volatile oils dissolve _
_ inether, benzol, and chloroform. These liquids, as arule, are not

_ allowed to flow upon the preparation (under the glass egverd =
_ the slide). It is preferable to place the sora in a eines ae
soak filled with the reagent. :

/

MICRO-CHEMICAL REAGENTS. 27d

Hardened masses of resin, such as are frequently found in-
older drugs, often resist for a long time the action of the solvent,.
a fact which must be considered in order to avoid incorrect con--
clusions. An alcoholic solution of caustic soda often has a bet--
ter action than alcohol and other solvents.

20. Paraffin of a low boiling point (55 to 75° C.), the s0-
called petroleum ether or petroleum benzin. This liquid serves
for similar purposes as ether, benzol, and chloroform; it has,
however, a much less energetic solvent action upon resins.

21. Fatty Oil (Almond Oil is the best) is employed with ad-
vantage as a mounting medium, when, beside the fatty oil
occurring in the cells, it is desired to examine the other con--
stituents which would become decomposed or dissolved by glycerin:
or water (aleurone). Sections of seeds rich in oil, when placed!
in the fatty oil, appear very much clearer, since the oil is taken
up by the mounting liquid.

22. Liquid high-boiling Paraffin (Parafinwm lMquidum*
of the Pharmacopoea Germanica), serves in most cases the same
purpose as the fatty oil and is cleaner in its application.

23. Todine in the form of powder, when strewn upon moist-
ened sections, often produces purer colorations than iodine
solutions; the excess of iodine is easily washed away with water..
Powdered iodine readily cakes together. It is therefore more:
convenient to use it in form of a trituration with siliceous earth
or pumice stone, and to preserve it in this form, since the latter
substances are not objectionable in many of the reactions to be-
made with iodine.

With regard to the application of iodine and its solutions for:
the recognition of starch, cellulose, and protein substances, com-
pare the respective sections preceding.

24, Iodine Water. One part of iodine shaken with 4,000:
parts of water is used as a reagent for starch and those forms of”
cellulose which show a similar reaction.

25. Iodine Solution (Iodine with Poiana Iodide) is a.

1 A clear, oil-like liquid obtianl from petroleum, free from colored,,
fluorescent, and odorous substances. F. B. P.

276 MICRO-CHEMICAL REAGENTS.

solution of 3 parts of iodine and 8 parts of potassium iodide in
1,200 parts of water. After some time, a little hydriodic acid is
formed in this solution; the deportment of the solution is then
(for example, with starch; see page 122) not precisely the same
as when the freshly prepared solution is used.

26. Iodine Tincture. A solution of 1 part of iodine in 10
parts of alcohol of the specific gravity 0.830.

27. Iodine with Glycerin. A mixture of 1 part of iodine
solution (No. 25) with 10 parts of glycerin of the specific gravity
1.230.

28. Iodine with Chloride of Zine. In 100 parts of a solu-
tion of chloride of zinc of the specific gravity 1.800 are dissolved
6 parts of potassium iodide and as much iodine (about 1 part) as
the liquid is capable of taking up.

Pure cellulose—though not that of the fungi—assumes with
the chloride of zinc and iodinea violet color (chloride of zinc
causes the formation of amyloid). Cells containing tannin as-
sume with the chloride of zinc and iodine a reddish color.

29. Potassio-mercuric Lodide (a solution of mercuric iodide
in potassium iodide) is prepared by dissolving 1.35 paris of
mercuric chloride (corrosive sublimate) and 5 parts of potas-
sium iodide in 100 parts of water. Nearly all the alkaloids are
precipitated by this reagent from their solutions, even when
highly diluted, so that it affords indications of the presence of
such substances. The precipitated compounds are mostly
amorphous, and only a few assume a crystalline form after some
hours. |

30. Ferrous Sulphate (Green Vitriol), prepared in the
form of a fine powder, by precipitating it from its solution in
water by means of alcohol, and quickly drying it by exposure to
the air. When used, 1 part is freshly dissolved in 20 parts of
water. Many substances of the class of tannins are colored by
this salt, but usually of a different tint than by ferric chloride.
The addition of lime-water to sections which have been impreg-
nated with a solution of ferrous sulphate and poring or os
= _ rinsed with water, often produces new colorations. :
a Ferric Chloride, The officinal solution [Pharm. Germ. ]

MICRO-CHEMICAL REAGENTS. 277

of the specific gravity 1.405 is diluted with 10 times its weight
of water, and the sections allowed to macerate for some time in
the liquid. If aleohol be employed for diluting the solution,
somewhat different reactions are usually obtained, and, upon the
subsequent addition of lime-water, still further changes of color
appear. The dilute solution of ferric chloride is decomposed by
long keeping (as a result of dissociation). Only the officinal
solution should, therefore, be kept ready prepared.

The dilute solution of ferric chloride serves chiefly for the
recognition of tannic matters, which are thereby colored either
green or blue. It is expedient to discriminate between these
two classes of colorations, though this is frequently difficult,
owing to the appearance of transition colors in consequence of
several tannic matters being usually present at the same time.
This assumption of the simultaneous presence of several differ-
ent tannic matters is supported by the observation that the
coloration first produced by very small quantities of solution of
ferrous sulphate in cells containing tannin is often changed by
the further addition of ferric chloride. The behavior of pyro-
catechin, quercitrin, and rutin to iron salts may here also be |
called to mind.

Instead of ferric chloride, ferric sulphate or ferric acetate may
also be employed.

32. Mereurous Nitrate, known also by the name of “ Mil- |
lon’s reagent.” One part of mercury is dissolved, without
heat, in 1 part of fuming nitric acid, and the solution diluted
with 2 parts of water. This liquid imparts a red color to pro-
tein substances, though only when the latter are present in
considerable amount. The striping of the membranes is ren-
dered clearer by Millon’s reagent. On account of its strongly
acid reaction, care must be taken not to have it come in contact.
with the microscope.

33, Aniline Sulphate, in aqueous, or better, alcoholic solu- _

tion, colors all lignified membranes yellow, especially after the oe :

addition of sulphuric or hydrochloric acid.
34. Phloroglucin is a still more delicate reagent for lignifi-

cation. The sections are thoroughly moistened with hydro~

278 MICRO-CHEMICAL REAGENTS.

chloric acid, and a freshly prepared solution of phloroglucin in
100 parts of water dropped upon them, whereupon lignified
membranes become red. Occasionally, this coloration appears
without the addition of phloroglucin, for the reason that this
principle itself occurs in some barks.

CoLoRING AGENTS.

35. Aniline Colors. Fuchsine, methyl-violet, methyl-green,
Hanstein’s aniline-violet (equal parts of methyl-violet and
fuchsine), vesuvine, as also aniline-blue and aniline-brown, re-
ceive manifold applications, particularly in bacteriological inves-
tigations, since these organisms are capable of strongly absorb-
ing the aniline colors. But in histological investigations the
above-named colors are also employed, usually dissolved in 100
parts of water.

_ 36. Eosin in aqueous solution colors dead protoplasm
intensely red, and is therefore especially applicable, for example,
in the examination of sieve-tubes.

37. Carmine Solution. The best carmine is dissolved in
ammonia-water, the clearly decanted liquid evaporated to
‘dryness, and the residue (preferably only as required) dissolved
in 100 parts of hot water. This reagent is abundantly absorbed
by many substances, for example by albumen and resins, also
by the delicate cuticle of cells, so that, by an unequal coloration
of the walls and constituent substances, many relations may be
rendered clearer.

38. Hematoxylin (3.5 parts in 100 parts of water) in com-
bination with alum is an admirable coloring agent for cell-
nuclei.

MountTinG MEDIA.
If it is desired to keep a preparation, it must be preserved in

a medium which does not evaporate, and in which the structure —
_ of the preparation may be clearly retained. The most convenient

to use for this purpose are certain preserving liquids, especially :

_ glycerin (specific gravity 1.250) or calcium chloride (one part of

MICRO-CHEMICAL REAGENTS. 279

the salt in three parts of water). These two liquids, particularly
the first-named, leave most preparations unchanged, even after
many years. Starch, however, is dissolved by calcium chloride,
even when it is neutral.

For firmer objects, sections of more compact drugs, for
Lycopodium, and also for diatoms (provided they will bear some
warming) Canada balsam may be employed as the mounting
medium. The preparations must, however, have previously
been repeatedly washed with alcohol. The section is then placed
in Canada balsam which has been liquefied with a little warm
chloroform, and finally in the slightly warmed balsam itself.

A solution of gelatin in glycerin is also adapted for delicate, as
well as.for coarse preparations. This is obtained by gently -
warming one part of colorless gelatin with six parts of water
and seven parts of glycerin. When used, the mixture is lique-
fied by warming.

When Canada balsam or glycerin-gelatin are used, it is not
absolutely necessary to specially cement the cover-glass, since
the solidifying mounting medium holds the cover-glass firmly;
but if glycerin or calcium chloride solutions are used as mount-
ing media, it is necessary to cement the cover-glass. As varnish,
either the ordinary black asphalt varnish or the yellow “‘ pre-
pared gold-size ” are employed.

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LN LD Ex

Abortive parts of the flower, 75
Aboul Mena, 24
Absorption of
ment, 208
of inorganic salts, 208
of organic nutriment, 208
Acetic acid, 270
Acid, acetic, 270
chromic, 268
filicic, 135, 246
gallic, 185
gallo-tannic, 138
hydriodic, 275
hydrochloric, 269
nitric, 145, 269
phosphoric, 144
stearic, 106
sulphuric, 145, 269
sulphuric, concentrated, 269
sulphuric, dilute, 269
tannic, 137, 271
Acids, plant, 139
Acropetalous, 78
Actinomorphous, 78
Adragantin, 164
Adulterations, 16, 22, 35
Aérating system, 235
Aistivation, convolute, 68
imbricate, 68
plicate, 68

inorganic nutri-

Albertus Magnus, 28
Albumen, 87
Albuminous substances, 235, 273

Alburnum, 230
Alcohol, 274
absolute, 274
Aleurone, 97
granules, 97
Alhervi, 23
Alkaline solution of tartrate of
copper, 272
Alkaloid, 108
Alkaloids, 139, 249
Almond oil, 275
Aloe, 144
Alphita, 25
Aluminium, 145
Amalfi, 26
Amentum, 78
Amides, 139
Ammonia water, 271
Amyegdalin, 135
Amylo-cellulose, 170
Amyloid, 123, 168
Amylum, 108
Anatomico-physiological system of
tissue, 174
Anatomy, 93
Anatropous, 89
Androeceum, 70
Anemophilous plants, 74
Aniline blue, 278
brown, 278
colors, 278
sulphate, 277
violet, Hanstein’s, 278
Annual rings, 220, 221
Annular vessels, 217
Anthela, 79
Anthers, 70
halves of the, 70
monothecous, 70.
Anthocyan, 104

282 INDEX.

Anthoxanthin, 104
Apocarpous gyneceum, 73
Apotropous, 90
Apparent rings, 220
Appendages of the epidermis, 182
of the fruit, 91
of the seed, 90
Apposition, 111
Arabin, 164
Arabinic acid, salts of, 169
Arabs, 23
Archiv des Apotheker-Vereins, 54
Areolated pits, 153, 218
Arillus, 91
Arnaldus de Villanova, 25
Arrangement of the leaves, 65
Arrowroot, 115
East Indian, 115
Ash, constituents of the, 144
estimation of the, 147
obtainment of the, 146
of plants, 144
Asparagin, 135
Asphalt varnish, 279
Assimilating surface, 175
system, 175
Assimilation, 209
products of, 102, 107, 119
starch, 108
Astrosclereids, 200
Asymmetrical, 78
Atropine, 108
Atropous, 89
Autochthonous starch, 108
Autumn wood, 221
Avicenna, 23
Axis, 78

Bacca, 86
Balsam, 251, 258
passages, se , 203
passages, schizogenic, 254
Bark, 62, 232 7
green, 233
of monocotyledons, 62
_ parenchyma, 232
»s, 241

Bast, 62, 194, 217
bundles, 68
cells, firmness of, 196
cell bundles, 197
fibres, application of, 199
horn, 164
soft, 231
tubes, 155, 156, 158
Bean starch, 114
Beet-root sugar, 142
Bell, Jacob, 54
Benedictine convents, 27
Benzol, 274
Berg, 41
Berry, 86
Bible, 19
may as ad of Pharmacognosy,
5

Bifacial leaves, 210
Bitter principles, 139, 249
Bock, 32
Bordered pits, 153, 218
Bork, 62, 188
formation, 187
ringed, 191
scale, 191
Bostryx, 64, 79
Botanischer Jahresbericht, 55
Botanisches Centralblatt, 55
Bottle cork, 192
Boyle, 38
Bracts, 67, 70
Branches, 62
Bristles, 182
Bromine, 145
Brunfels, 33
Brunschwig, 32
Bud, 58, 91

scales, 66
Bulbodium tunicatum, 60
Bulbous tuber, 60
Bundles, bast, 63
bast-cell, 197
primary, 234
vascular, 211, 213
Buonafede, 36

Calcium, 144
carbonate, 135
oxalate, 129
phosphate, 134

INDEX.

‘Calyculus, 70
Calyptrogen, 175
Calyx, 68
leaves, 67, 68
Cambiform, 217
cells, 231
‘Cambium, 220
activity of, 221
intrafascicular, 222
ring, 220
‘Campylotropous, 90
Canals, schizogenic, 254
Cane sugar, 141
Caoutchouc, 249
‘Capitulare, 24
Capitulum, 79
‘Capsule, 85
wall of, 157
Carbohydrates, 235
Carmine solution, 278
‘Carpel, 71, 78, 76, 84
Carpophore, 83
Caruncle, 90
‘Caryopsis, 84
Catalonians, 29
Cataphylla, 66
Cathkin, 78
‘Cato, 21
‘Caubis, 62
Caustic soda, 271
Cavities, 235
mucilage, 244
‘Cecidize, 265
Cell, 93 ei
aggregates,
shoune of form of, 148, 171
contents of, 94
fusions, 173, 217
growth of, 148
membrane, 148
membrane, chemical behavior

8a
wall, 94, 148

Cells, bast, 156, 171, 194
collenchyma, 194
poco gh
c ’
pa beac 94
epidermal, 181
guardian, 239
isodiametric, 149

283

Cells, latticed, 217
lines of, 244, 249
mucilage, 244
multiplication of, 94
of leaf, 172
organized contents of, 139
resin, 263
secerning, 254
secreting, 244
spherically polyhedral, 149
stone, 156, 200
*wood, 218, 223
wood-parenchyma, 218, 223
Cellular tissue, 174
Cellulose, fungus, 160
membrane, 159
‘starch, 121
Central rhizome, 59
Centric leaves, 210
Cerasin, 164
Cerin, 161
reaction, 269
Chalaza, 89
Changes of the drug on drying,
140
Charaka, 20
Charlemagne, 24
Chemical constituents, 16
Chinese, 19 .
Chloride of iron, 275
of zine with iodine, 275
Chlorine, 145
Chloroform, 274
Chlorophyll, 100
bodies, 100
coloring matter, 100
crude, 100
formation of, 103
granules, 100
granules, fundamental mass
of, 100
pure, 100
reactions, 104
spectrum, 101
Chlorophylan, 101
Chlorosis, 144
Choripetalous, 68
Choriphyllous, 68 _
Chorisepalous, 68
Chorisis, 77
Chromic acid, 268
Cincinnus, 64, 79
Circinal vernation, 69
Clusius, 31

284

Coalescence, 77
Collagen layer, 166
Collateral vascular bundles, 234
Collecting, time for, 13
Collections, 43
of drugs, 43
of plants, 45
Collective fruits, 81
Collenchyma, 158, 171, 194
Colleters, 166, 184
Colored epidermis, 181
Coloring matters, 104, 139
matters, crystalloid, 104
Columella, 21
Coma, 186
scongeee es policy of the Dutch,

relations, 15
Compound starch granules, 111
Concentrated sulphuric acid, 269
Concentric vascular bundles, 234
Conducting system, 213
Conduplicate vernation, 69
Coniine, 259
Connective, 70
Constantinus Africanus, 25
Convent of St. Gall, plan of, 24
Convolute eestivation, 68

vernation, 69
Copper, acetate, 271

ag solution of tartrate,

ammoniacal oxide, 273
Cordus, 32
Cork, 161, 187, 233
bands, 187
cells, 187
development of, 191
fat, 159
protective, 194, 264
wound, 194
Corky growths, 194
growths cn leaves, 194
layer of barks, 187
Corm,
Cormus, 60
Corolla, 68
leaves, 67, 68
Corona staminea, 71
Corroded starch granules, 121
Corrugate vernation, 69
Corymb, 79
Coste,

iss, 88

INDEX.

Cotyledons, 67, 88
Crusaders, 27
Crystal-cells, 233
Crystalloid coloring matters, 104
Crystalloids, 98
Crystals, 135, 244
Cubebin, 135
Cultivation of officinal plants, 11
Cupula, 86
Cuticle, 161, 179
Cuticular layers, 180
Cutin, 161
Cyathium, 80
Cycles of the flower, 75
Cyme, 79
Cymose, 64
inflorescence, 79
Cystoliths, 244

Decussate, 65
Dédoublement, 77
Deduplication, 77
Dehiscence, 85
of the anthers, 70
Dehiscent fruits, 84
Derma, 62
Dermatogen, 175
Dermatogenic gum-passages, 251
Description of drugs, 15 :
Developing tissue, 174
Development of cork, 191
Dextrin, 139, 235
Dextrose, 142
Diagram of flower, 74
Dichasium, 79
Dichotomous system, 64
Dichotomy, false, 79
forked, 64
Diclinous, 73
Dicotyledonous flowers, 75
Diocletian’s edict (801 a.p.), 23
Dicecious, 73
Dioscorides, 21, 32
Diplecolobez, 89
Diplostemonous, 75
Disk, 67
extrastaminal, 67
intrastaminal, 67
ert carne of the plant-cell,

Dissepiments, false, 72, 84
Dots, areolated, 152
Double akenes, 83

Drug, 139

INDEX.

Drugs, collections of, 43
of the Orient, 29

Drupe, 86

Dry fruits, 84

Dry weight, 141

Drying drugs, 139

‘Ducts, 247

Duramen, 230

Duty at Alexandria, 22

Egypt, 17
Eisodial opening, 239
Elementary organs of the wood,
223
Elements, mechanical, 217
specifically mechanical, 194
Embryo, 86
Empbryonal leaves, 88
Endocarp, 82
Endodermis, 202
single-rowed, 208
Endophleeum, 62
Endosperm, 87
farinaceous, 117
horn-like, 116
Eosin, 278
Epiblema, 177
Epicarp, 82
Epidermal cells, 181
glands, 245
tissue, system of, 175, 176
water-tissue, 235
Epidermis, 176
colored, 181
of root, 176
of several layers, 177
strengthening layers of, 177
Epigea, 91 ;
Epigynous, 72
Epipetalous, 75
Episepalous, 76
Epithelium, 254
Epitropous, 90
Ether, 274
Etiolation, 103
Etiolin, 103
Excrescences, 265
Excretions, 241
receptacles for, 241
Exine, 71
Exogenous, 64
Exophloeum, 62
Extrorse, 70

285

Falloppio, 37
False dichotomy, 79
dissepiments, 72, 84
Farinaceous endosperm, 117
Fat, 105
Fatty oil, 105, 275
Fehling’s solution, 272
Female organs of the plant, 71
Fernandez (Oviedo), 33
Ferric chloride, 276
Ferrous sulphate, 276
Fertilization, 73
by insects, 74
Fibres, 198
animal, 199
bast, 199
plant, 199
Fibrous tissue, 199
Fibro-vascular bundle, 213
Figurative representation of inner
structure, 50
Filament, 70
Filicic acid, 135
Filling tissue, 176
Firmness of bast-cells, 196
Flagella, 57
Florence, 26
Florentine Levant trade, 26, 29
Flos, 68
Flour, 113, 186
Flower, 67, 68
cycles of, 75
formulas of, 76
receptacle of, 67
stalk,
leaves of the, 67
Flowers, 63, 67, 81
dicotyledonous, 75
tetracyclous, 76
tricyclous, 76
Fluorine, 145
Folding of the membrane, 211
Folia, 66
Follicle, 84
Forked dichotomy, 64
Form of leaves, 66

Formation of organic substances,
209

Formulas of the flower, 76
Frankfurter Liste, 30
Fructifying column, 71
Fruit, 8 -

286

Fruit sugar, 142
Fruits, 81
collective, 81
dehiscent, 84
dry, 84
forms of, 82
indehiscent, 84
stone, 86
succulent, 85
winged, 84
Fuchs, 33
Fuchsine, 278
Fulda, 27
Fundamental mass of chlorophyll
granules, 100
tissue, 176
Funiculus, 86
Fungus cellulose, 160
galls, 265
sugar, 142

Gallic acid, 135
Gallo-tannic acid, 138
fad 265
un:

ihecot | 265
Gamopetalous, 68
Gamophylious, 68
Gamosepalous, 68
Garcia’s Colloquios, 31
Garcia de Orta, 30
Gehe & Co., 8
Gelatin in glycerin, 279
Soe ae 166

Genoa,

Geoffroy, 5

Geographical distribution, 10

Germinating plant, 60

Germination, 69

Gesner, 32

Ghini, 36

Glands, epidermal, 245
intercellular, 246
internal, 246, 250
secreting wax, 185

Glandular hairs, 244
hairs, internal, 245
heads, 245, 263

Globoids, 98

Glossaries, 26.

Glucosides, 139

Glume, 67

Gluten, 235

Glutinous a 98

INDEX.

Glycerin, 273
ethers, 161
Granulose, 121
Grape sugar, 142
Grass oils, 244
Green bark, 233
Growing point, 94, ihe: 220
Growth in length, 1
in thickness, thy
Guibourt, 5, 40
Gum, 163
disease, 165
mucilage, 166
pathological, 166
physiological, 166
resin, 263
varieties of, 164
wound, 167, 264
Gummosis, 166
of the Amygdalezx, 166
Gyneeceum, 71
syncarpous, 73
Gynandrous, 71
Gynostemium, 71

Hadrom, 217
Hematoxylin, 278
Hair formations, 182
formations secreting resin, 184
Hairs, 113, 182, 199
chaffy, 182
glandular, 166, 184, 245
inner, 187
multicellular, 182
root, 208
stellate, 182
stinging, 182
Half-underground organs, 58
Halophytes, 145
Hanstein’s aniline violet, 278
Haustoria, 208
Head, 79
Heart-wood, 230

Hermaphrodite, 73
Hernandez, 33
Hesperidin, 135, 148
Heterophylious, 66
High leaves, 67

Poth involucre of, 70
Hildegard, 27
Hilum, 89

INDEX,

Hilum of starch, 109
History, 17
History of Pharmacognosy, 17, 53
Hoffmann, 38
Horn-bast, 164

bast, prosenchyma, 164

endosperm, 116
Horti Germaniz, 82
Hortus Sanitatis, 32
Hutten, 32
Hyaloplasm, 94
Hybrids, 74
Hydriodic acid, 275
Hydrochloric acid, 269
Hypanthium, 67
Hypanthodium, 81
Hypertrophies, 265
Hyphe, 1238, 172
Hypocotylous member, 60
Hypoderma, 177
Hypogzea, 91
Hypogynous, 72
Hypsophylla, 67
Hysterogenic secretion recepta-

cles, 263

Ibn Alawam, 23
Ibn Baitar, 23
Ibn Batuta, 23
Idioblasts, 243
containing albumen, 243
Idrisi, 23
Illustrations of drugs, 49
of officinal plants, 47
Imbricate zstivation, 68
India, 23
Indol, 269
Inferior ovary, 72
Inflorescence, 78
compound forms of, 79
Inner bark, 62, 282
cork, 188
- fruit layer, 82
Inorganic compounds in the cell-
membrane, 139
Integuments, 86
Intercellular glands, 246
resin receptacles, 250
secretion receptacles, 251,
spaces, 174, 235
substance, 162, 178
Internal glands, 250
glandular hairs, 187
hairs, 187

287

Internodes, 65
Intine, 71
Intrafascicular cambium, 222
Introrse, 70
Intussusception, 111, 118
tiyaee 124 P
vento of the pharmacy at
Branesecie 35 :, “4
Invert sugar, 142
Involucellum, 79
Involucre, 79
of high leaves, 70
Involute vernation, 69
Iodide of mercury and potassium,
276

4
Iodine, 145, 275
in potassium iodide, 275
solution, 275 :
tincture, 276
water, 275
with glycerin, 276
Irish moss, mucilage of, 167
Tron, 144
chloride of, 276
Isodiametric cells, 149
Israelites, 19
Istachri, 23

Jahresberichte, 54
Japan, 20

Jesuits, 20

Juge, 83

Kew botanical eo 12, 44, 45
Khurdadbah,

Kino, 144

Kyphi, 18

Labellum, 68
Lacune, 237
Leevulose, 142
Lamina of leaf, 65
Lateral axis, 64
branches, 64
walls of epidermis cells, 181
Latex cells, 248
tubes, 242, 247
Laticiferous ducts, contents of,
249
vessels, 248
Latticed cells, 217
Leaf buds, 91
cells, 172
lamina of, 65

288

Leaf, margins of, 200
merenchyma, 172, 211
nodes, 65
sheath, 66
skeleton, 213
spectrum, 101
stalk, 65

Leather, formation of, 139

Leaves, 64
bifacial, 210
centric, 210
floral, 67
foliage, 66
insertion of, 65
position of, 60

Legume, 84

Legumen, 84

Lemery, 5

Lenticels, 241

Leptom, 159, 217

Levant trade in the middle ages, 25
trade of Venice, 26

Liber, 62 %:

Libriform, 217° .
cells, 199, 217, 223

Lichen-starch, 170

Lichenin, 123, 170

igna, 61

Lignification, 160

Lignified membrane, 160

Lignin, 159, 160

Ligula, 65

Linseed mucilage, 166

Liquid paraffin, 275

Literature, 46

Lithipm, 145

Loeulicidal, 85

Lodicule, 70

Lorenzo de’ Medici in Florence, 29

Lumen, 94

Lysigenic balsam-passages, 170, 251
gum-passages, 251
oil-passages, 263
passages, 248
receptacles, 248

Macer Floridus, 25
Magnesium, 144

Maize starch, 112, 116

Male organs of the plant, 70
Manganese, 145

Mannite, 143

-Maranta aie 115

¥

INDEX.

Marco Polo, 20, 30
Marino Sanudo, 29
Martius, 5
Masudi, 23
Mechanical elements, 217
system of tissue, 194
Medicinal substance, 6
Medico-pharmaceutical botany, 46
descriptive works, 47
Medico-pharmaceutical zoology, 48
Medulla, 223
Medullary cells, 172
crown, 234
rays, 223
rays, secondary, 228
sheath, 234
Melitose, 142
Membrane, cellulose, 159
conversion into gum, 166
lignified, 159
metamorphosis of, 166
suberized, 159
Mercurous nitrate, 277
Merenchyma, 172
leaf, 172, 211
Mericarps, 83
Meristem, 174
zones, 175
Mesocarp, 82
Mesocarpium, 82
Mesophlceum, 62
Mesophyll, 211
Mestom, 217
Mesue, 23
Metamorphosis of membrane, 166
Methyl green, 278
violet, 278
Micelle, 118
Micro-chemical reagents, 268
Micropyle, 89
Microscope, 48, 93
use of, 48
Microscopical preparations, 46
structure, 16
Microsomes, 94
Middle bark, 62, 233
lamella, 162, 173
layer, 82
Milk juice, 140
sugar, 142
Millon’s reagent, 277
Mineral constituents, 144 by
constituents of plants, 144, 145
Monadelphous, 71 nes

Monardes, 33
Monoclinic oxalate, 129
Monocotyledonous flower, 76
Moneecious, 73
Monographs, 53
Monomerous ovaries, 71
Monopodial system, 64
Monopodium, 64
Monothecous anthers, 70
Monte Cassino, 25, 27
Morphology, 56
Mother-cell, 94
Mounting liquids, 278
Mucilage, 163

cavities, 244

cells, 244

glands, 263

passages, schizogenic, 263

plant, 164, 167

quince, 164, 166

sugar, 142
Mucilaginous substances, 164
Mycocecidiz, 265
Mycose, 142

Naming the mother-plant, 9
Nectaries, 67
Nerves, 211
ends of, 213, 218
Netted vessels, 217
Neumann, 38
Nicolaus Przepositus, 25
Nitric acid, 145, 269
Nodes, leaf, 65
Nordlinger Register, 30
Nuclein, 96
Nucleolus, 96
Nucleus, 86, 96
of starch, 109
sheath, 202
Nutlets, 72
Nux, 84

Oat starch, 117
Obdiplostemonous, 76
Obdurator, 90
Oil, 235, 260
almond, 275
cells, 243
fatty, 105, 275
passages, 242, 251
receptacles, 250
spaces, 251
tubes, 83, 258
19

INDEX. 289

Olein, 107
Opisthial opening, 239
Optical behavior of bast-tubes, 158
behavior of stone-cells, 158
Orthoploce, 89
Orthotropous, 89
Oudemans, 41
Outer bark, 233
wall of epidermal cells, 181
Ovaries, polymerous, 71
Ovary, 67, 71
inferior, 72
superior, 72
walls of, 72, 82
Ovule, 73, 86
Oxalate crystals, 129
crystals, aggregates of, 132

Padua (Garden), 37
Pappus, 69, 91
Parasites, 208
Parasitism, 266
Palisade cells, 172

layer, 210

parenchyma, 210
Palladius, 21
Palmitin, 135
Paraffin, 275

liquid, 275
Parenchyma, 172

bands,

cells, 151

rays, 223

spongy, 211
Parietal placenta, 86
Pasi, Paxi, 30
Pathological formations, 264

tannin, 139
Pectin substances, 170

Pegolotti, 29

Pellucid — 263
Pentacyclous flowers, 76
Pentamerous circles, 76
Pen t’sao kang mu, 20
‘Pepper, 22, 26

Pereira, 5

Periblem, 175
Pericambium, 234
Pericarp, 81, 82
Pericarpium, 82

290

Periderm, 62, 187
Perigon, 68, 69, 70
leaves, 75
Perigynous, 72
Periplus of the Erythrzean Sea, 22
Perisperm, 88
Permanent tissue, 174
Petals, 67, 6
Petiole, 65
Petiolus, 65
Pharmaceutical Journal and Trans-
actions, 54
Pharmacognostical systems, 41
text-books and manuals, 51
Pharmacognosy, 6
bibliography of, 53
Pharmacology, 4
Pharmacopceias, 38
Pharmacy in Dijon (1489), 30
Phelloderm, 187
Phellogen, 187
Phellonic acid, 161
Phloém, 159, 217
Phloroglucin, 137, 277
reaction, 161
Phoenicians, 19
Phosphoric acid, 144
Phyllocladia, 61
Phyllodia, 61
Halansien Hildegard’s, 27
Physiological gum, 166
tannin, 139
Picrotoxin, 135
Piperarii, 26

Pistil, 71
Pits, areolated, 153, 218
bordered, 153, 218
dotted, 152
Pitted vessels, 217
- Place of attachment of the leaf, 65

Placenta, 86

basal. 86
central, 86
parietal, 86

Plan of the Convent of St. Gall, 24.

Plant acids, 139
fibres, 199

mucilages, 1
ages ‘collection wt. 45

| sess. 82

INDEX.

Plerom, 1'75
Plicate zstivation, 68
vernation, 69
Pliny, 21, 22
Plumule, 88
Pollen, 70
cells, 70
grains, 71
grains, receptacles of, 70
sac, 70
tube, 73
Pollinaria, 71
Poflinia, 71
Polyadelphous, 71
Polycarpic flower, 73
Polygamous, 73
Polymerous ovaries, 71
Pomet, 5
Pore canals, 151
capsule, 85
Pores, 151, 215
Porus, 239
Potassio-mercuric iodide, 276
Potassium, 144
chlorate, 269
hydroxide, 272
sodium tartrate, 272
Potato starch, 112, 114
Pratica della mercatura, 29
Prepared gold-size, 279
Prickles, 182, 186
Primary axis, 78
bark, 62
bundles, 234
ribs, 83
roots, 56
Primordial utricle, 94
Projection of a flower on a plane,
74
Prosenchyma, 172
horn-bast, 164
Protein bodies, 88, 95
erystalloids, 98
granules, 97
substances, amount of, 100
Protogenic gum passages, 263
Protoplasm, 94
circulation of, 183
reactions, 96
Pruinosus, 163
Pseud-axis, 79
Pseudo-fruits, 81
oe deere hk

INDEX.

Pulpa, 82 ‘

Pulvis contra omnes Febres, 24
Pyrocatechin, 136

Pyxidium, 85

Quadratic oxalate, 131
Quercitrin, 136
Quince mucilage, 164, 166

Raceme, 79
Racemose, 64
inflorescence, 78
Racemus, 79 :
Radial vascular bundles, 234
Radicles, 60, 88
Radicula, 88
Rami, 62
Ramification, 64
system of, 64
Raphides, 129
Rays of the umbel, 79
Reactions of fats, 107
of grape sugar, 271
of starch, 122
Receptacle, 67
of the flower, 67
Receptacles for excretions, 241
for reserve substances, 235
for secretions, system of, 251
of the pollen grains, 70
Receptaculum, 67
Reformation der Apotheken, 83
Regimen sanitatis Salernitanum,

25
Report on the progress of phar-
macy, 55
Reserve nutritive substances, 59,
100, 118
receptacles, 235

passages, 253 fe
receptacles, intercellular, 250_
Resorption of the transverse walls,
i SO NS
Respiratory cavity, 250
Resupination, 78
Revolute vernation, 69
_ Rhachis, 78
_ _Rhaphe, 90

291

Rhizomata, 58
Rhizome, 58
central, 59
lateral, 59
Rhytidoma, 62, 188
Rice starch, 112, 117
Rima, 70
Ringed bork, 191
Root, 56, 58, 59, 118, 213
branches, 56
epidermis of, 176
fibres, 57
hairs, 57, 182, 208
primary, 56
secondary, 57
stock, 58
tip of, 56
Rumphius’
nense, 31
Runners, 57
Rutin, 136

Saccharose, 142 °
Saffron, 22, 35

Sago starch, 113
Salernitans, 25
Salerno, medical school of, 25, 27
Samara, 84

Sap-wood, 230
Sarcocarp, 107
Scalariform vessels, 217
Scale-bork, 191

Scales, 245

Scheele, 38

Herbarium amboi-

Schizocarps, 83

oben! ee balsam passages, 251,
canals, 254 Be
mucilage passages, 263 :
receptacles for secretions, 254

Schleiden, 40

Schultze’s macerating liquid, 160,

969
Sclereids, 157, 171, 200

Sclerenchyma, 200
Sclerotium, 173
Scutellum, 208
Secerning cells, 254
trichomes, 184
Secondary bark, 62
bark rays, 228
growth in thickness, 220.
medullary rays, 228
rhizome, 59

292 . INDEX.

Secondary ribs, 83 Squame, 66
xylem rays, 228 St. Gall, 24, 27
Secreting cells, 244 Stamen, 67
Secretion, receptacles of, 251 Staminodia, 71, 75
receptacles of, formed by cell Starch, 108
fusion, 263 assimilated, 108, 120
receptacles of, intercellular, autochthonous, 108
263 bean, 114
Secretions, 134, 242 cellulose, 121
organs of, 242 composition of, 121
schizogenous receptacles for, formation of, 111, 119
254 granules, compound, 111
Seed, 87 granules, corroded, 121
appendages of the, 90 granules, form of, 112
integuments of, 87 granules, size of, 123
nucleus of, 87 hilum of, 109
shell, 87 lichen, 123
testa, 87 maize, 112, 116
vessel, 86 maranta, 115
yield of, 74 nucleus of, 109
Sepals, 67, 68 oat, 117
Septicidal, 85 paths of, 119
Septifragal, 85 — potato, 112, 114
Serapion, 23 reserve, 108
Sertiirner, 39 rice, 112, 117
Sheath, 66, 79 sago, 113
nucleus, 202 sheath, 120, 206, 231
Sieve plates, 231 transitory, 120
- portion of vascular bundle, 213 wheat, 112, 115
tubes, 217, 231 Stearic acid, 106
Silica, skeleton of, 145 Stearin, 135
Siliqua, 84 Stellate hairs, 182
Silicium, 144 Stem, 61
Soda, caustic, 271 organs of, 60
Sodium, 144 structures, 60
hydroxide, 271 structures, form of transverse
Soft bast, 231 section, 61
Solitar, 99 summit of, 64, 174
Spaces, intercellular, 174, 235 Stereids, 194
Si J Stereom, 194
Spathe, 79 Stigma, 73
oe mechanical elements, papillze of, 73
Stinging hairs, 182
Secceons of chlorophyll, 101 Stipites, 61
of leaf, 101 Stipule, 66
stals, 111, 126 Stipules, 66
Stolons, 57
ike, Stoma, 239
a veeney aay Stomata, 238

Stone cells, 156, 158, 171, 200
fruits, 86 2

ores, | Ponce stn -
Spring-wood, 221 ets on,

INDEX.

Strychnine, 135
Style, 73
' Stylus, 75
Suberin, 159, 161
Suberized membrane, 162
Substance, intercellular, 162, 173
Substitutions, 16
Sugar, 139, 141, 235
cane, 141
beet-root, 142
fruit, 142
grape, 142
invert, 142
milk, 142
mucilage, 142
sheath, 231
Sulphuric acid, 269
acid, concentrated, 269
acid, dilute, 269
Superior ovary, 72
Surface growth, 148, 171
Susruta, 20
Suture, dorsal, 84
_ ventral, 71, 84
Swarm spores, 93
Symbiosis, 266
Sympetalous, 68
Sympodium, 64
Symsepalous, 68
Syncarpium, 83
Syncarpous gynzeceum, 73
System, absorbing, 208
aérating, 175, 235.
assimilating, 175, 209
conducting, 175, 213
dichotomous, 64
epidermal, 175, 176
mechanical, 175, 194
monopodial, 64
of ramification, 64
of receptacles for secretions,
175, 241
storing, 175, 235
Systems of tissue, 175
pharmacognostical, 41

Tannic acid, 137, 271
- matter, 137
matter, estimation of, 138
Tannin, 125
pathological, 139
physiological, 139
Tap roots, 56

295

Tendrils, 67
Tensions, 111, 154
Testa of seed, 87
Tetracyclous flowers, 76
Tetramerous circles, 76
Thecee, 70
Theobromine, 135
Theriac, 35
Thvlle, 194
Tissue, cavities in, 235
fibrous, 199
filling, 176

* fundamental, 176

permanent, 174
systems of, 175
transpirating, 238
Torus, 67
Trabecular vessels, 217
Trachesx, 217
Tracheids, 217, 218, 223
Trade-books, 28
Tragus, 32
Transitory starch, 108
Treatment of the subject-matter, 8
Trichome formations, 245
Trichomes, 182, 185
secerning, 184
secreting nectar, 185
Tricyclous flowers, 76
Trimerous circles, 76
Trommer’s reaction, 142
Trommsedorff, 5, 39
Trunci, 62
Tubers, 59
Tuft of hairs, 246
Typical structure of vascular bun-
dles, 217

Umbel, 79
Umbellet, 79
Under leaves, 58, 66
Underground organs, 56, 58
Unicellular plants, 93
University garden, the first botani-
peace 36 |
of 4
of Padua, 36
Vacuoles, 94
Vagina, 66
Valerius Cordus, 31
Vallecule , 93
Valvate zestivation, 68
vernation, 69

294 INDEX.

Valves, 70

Vanillin, 135

Varro, 21

Vascular bundle ring, 229
bundle sheath, 202
bundles, 211, 213

bundles, typical structure of,
217

Venice, 26

hh gee circinal, 69
conduplicate, 69
corrugate, 69
involute, 69
plicate, 69
revolute, 69
valvate, 69

Vessels, 171, 217, 223
annular, 217
netted, 217
pitted, 217
scalariform, 217
spiral, 217
sribecs iia, 217

Vesuvine, 278

Vexillum, 68

Vittee, 88, 258

Volatile oils, 140, 242

Water, amount of contained, 139
cultures, 146

Wax, 163

Weddel, 40

Wheat starch, 112, 115
Winged fruits, 84
Wood, 61, 223
autumn, 221
cells, 218, 223
elementary organs of, 223
fibres, 223
heart, 230
parenchyma, 217,218
parenchyma cells, 218, 223
ring of dicotyledonous stems,

spring, 221
Woody substance, 160
Wound cork, 194

gum, 167, 264

Xanthophyll, 100
Xanthoprotein reaction, 272
Xylem, 217

rays, secondary, 228
Xylogen, 160

Year book of pharmacy, 54

Zinc chloride with iodine, 276
Zoidiophilous plants, 74
Zoocecidia, 265

Zottenkopf, 246
Zygomorphous, 78

CORRIGENDA.
Page 170, for lacticiferous, read laticiferous.
Page 172, notes 2 and 5. Here and in a few other places the Greek ac-
cents have been broken off during the printing.
Page 211, second line from top, for falling off of the membrane, read

_ folding of the membrane.

Page 235, sixth line from top, for tubes, read tubers, .