./

The following Extracts are selected from a few of the many favourable
Notices which hare appeared concerning this Work : —

" A spirited translation of a celebrated work, very elegantly gotten up." — Spectator.

" Mr. Houghton Kingdon has here commenced a very acceptable service to the
British botanist, by undertaking a translation of De Candolle's celebrated work on
Vegetable Organography. . . We have to state that it is executed with a perfect
knowledge of the author and his language ; and promises to be an indispensable ad-
dition to the Botanical Library." — Literary Gazelle, Feb. 23, 1839.

" This edition of one of its author's most valuable elementary works cannot fail to
prove acceptable and highly useful to the English student of botany. The part before
ns is vigorously translated ; it is got up in a very handsome style, and contains a satis-
factory specimen of the plates which are to illustrate the minute parts of Vegetable
Anatomy, &c. .". . . . We are glad to learn that it is to be followed by a translation of
the same author's work on Vegetable Physiology." — Monthly Review, February 1839.

" A translation of this work has long been a desideratum to the English botanist.
One is now offered by Mr. Bot/ghton Kingdon, which, for the convenience of pur-
chasers, is to appear in Monthly 2s. 6d. Parts. The work is well printed, and is
illustrated by Plates of the organs, and numerous anatomical parts of Vegetables." —
Tait's Edinburgh Magazine, February 1839.

" We are glad to see the commencement, in numbers, by Mr. Kingdon, of a
translation of the elementary works of the celebrated De Candolle. This will, indeed,
be a great desideratum to the English reader. It is true that we have had partial
translations and comments on the speculations and views of this eminent philosopher,
but no opportunity of following him through the consecutive workings of his master
mind. The style of the translation is clear and concise." — The Botanist, February
1839.

" We are very glad that there is at length a prospect of our possessing a good
English version of De Candolle's Organography and Physiology, for the latter will be
commenced at the conclusion of the former : every candid botanist will unhesitatingly
admit that this celebrated author claims a place among the most philosophic natu-
ralists of the present day. The translation is good, and the illustrations are neatly
executed." — The Naturalist, March 1839.

" The translation of De Candolle's work on the Organography of Plants, by Mr.
Boughton Kingdon, is a valuable addition to our botanical literature. . . . We are
glad to observe that it will continue to be published in Monthly Parts, so that it will
be within every body's reach." — Gardener's Gazette, March 16, 1839.

'• We most strongly recommend the work to all our readers who have a taste for
plants, and more especially to all young gardeners." — Gardener's Magazine, April
1839.

VEGETABLE

ORGANOGRAPHY;

AX ANALYTICAL DESCRIPTION

THE ORGANS OF PLANTS.

M. AUU. F. DE CANDOLLE,

Knight Commander of the Legion of Honour ; Member of the Sovereign Council of Geneva ; Honorarj
Professor of Natural History . Director of the Botanic Garden, n-.i i President of thi

Arts of that Cits ; Member of the imperial Academy of Sciences of st. Petersburgfa

ol the Royal Academies of Sciences of Paris, Stockholm, Turin, Munich,

Harriett)* Naples, Brussels; of the Royal Societies »i' London,

Edinburgh, and Copenhagen ; and of the Linna?an

and Horticultural Societies of London,

\c. &c. \c,

fRANSLATED BY BOUGHTON KINGDON.

SECOND EDITION.
VOL. II.

LONDON :
PUBLISHED BY HOULSTOX & STONEMAN,

6.'), PATERNOSTER ROW;
AND HOl'LSTOX & HUGHES, 154, STRAND.

184:1.

Q

ClL

\l
V. 2

LONDON :
R. CLAY, PRINTER, BREAD STREET HILL,

CONTENTS TO VOL. II.

PAGE

BOOK III.

On the Reproductive Organs, gr the Parts essential

to Reproduction.

Introduction • j

Chap. I.

Of the Inflorescence 4

Sect. I. Of the Inflorescence in general 5

II. Of Axillary, or Indefinite Inflorescence 7

III. Of Terminal Inflorescence 18

IV. Of Mixed Inflorescence 22

§ 1. Of Thyrses ;/,.

§2. Of Corymbs 26

Sect. V. Of Anomalous Inflorescence 27

§ 1. Inflorescences opposite the Leaves ib.

(j 2. Radical Inflorescence 29

§ 3. Lateral, or Extra-axillary Inflorescence 30

§ 4. Petiolary Inflorescence ih.

§ 5. Epiphyllous Inflorescence 32

Sect. VI. Of Pedicels and Peduncles 33

VII. Of Bracts 40

Chap. II.

Of the Structure of the Flowers of Phanerogamous

Plants 47

Sect. I. General Observations ib.

II. Of the Calyx, or Sepals 18

VI CONTENTS.

PAGE

Sect. III. Of the Corolla, or Petals 52

IV. Of the Stamens 56

V. Of the Pistil, or Carpels 67

VI. Of the Torus and its Adhesions 74

VII. Of Abortion of Parts of the Flower, or of their Dege-
nerations 80

VIII. Of Monochlamydeous, or incomplete Flowers ... 86
IX. Of the Relative Position of the Parts of one Floral

Verticil compared with that of another . .... 91

X. Of the Multiplication of the Floral Organs .... 93

§ 1. The Multiplication of the Rows of Verticils .... ib.

§ 2. The Multiplication of Parts of a Verticil 96

§ .'!. General Examination of Double Flowers 98

Sect. XI. Of the Irregularity of Parts of a Floral Verticil, or of

Irregular Flowers 100

XII. Of the Primitive Disposition of the Parts of a Floral

Verticil, or of the ^Estivation 105

XIII. Of Flowers united together Ill

XIV. Of the absolute Number of the Parts of each Floral

Verticil 112

XV. Of Nectaries 115

XVI. Comparison of Foliaceous and Petaloid Parts . . . 118
XVII. Of the particular Analogy between the Male and

Female Organs of Flowers 123

XVIII. General Conclusions and Considerations upon the

Structure of Flowers 125

Chap. III.

Of the Structure of the Fruit of Phanerogamous Plants . 133

Sect. I. Of the Fruit in general ib.

II. Of the Carpels considered in the state of separation

from one another 135

III. Of the Carpels of the same Flower united together . 148

IV. Of Carpels, considered with regard to their relation to

the other Parts of the Flower which are persistent,

or united around them 162

V. Of the Organs situated outside the Flower, and which

sometimes seem to form Part of the Fruit .... 170
VI. Of the Aggregation of Fruits which proceed from dif-
ferent Flowers 172

VII. Of the Umbilical Cord and its Expansions .... 177

< (INTENTS. VII

PAGE

Chap. IV.

Of the Structure of the Seed of Phanerogamous Plants . 183

Sect. I. Of the Seed in general ib.

II. Of the Spermoderm, or Skin of the Seed .... 188

III. Of the Albumen 192

IV. Of the Embryo 197

Chap. V.

Or the Organs of Reproduction without Fecundation in

Phanerogamous Plants 214

Chap. VI.

Of the Organs of Reproduction in Cryptogamous Plants . 219

Sect. I. General Considerations ib.

II. The Equisetaceae 223

III. Of the Marsileacae or Rhizospermae 226

IV. Of Ferns 227

V. Of the Lycopodiaceae 233

\ I. Of Mosses 236

VII. Of the Hepaticae 244

VIII. Of Lichens 247

IX. Of Fungi 249

Of Algae 251

§ 1. Of the Characeae 252

* 2. Of the Thalassiophyia- 254

. § 3. Of the Confervae 257

HOOK IV.

Of the Accessory Organs, or, of the Modifications of the
Fundamental Organs which render them capable of
serving as the means of protection for the other
Organs, or of fulfilling other Accessory Uses . . . 260

Chap. I.
Of Thorns 261

Chap. II.
Oi Tendrils 267

Vlll CONTENTS.

PAGE

Chap. III.
Of Fasciculated Expansions 273

Chap. IV.

Of Deposits of Nourishment, or the Fleshy, Succulent,
etc. Degenerations which modify the Texture of
Organs 275

Chap. V.
Of Scales 281

Chap. VI.
Of Buds 283

BOOK V.

Conclusion, or General Considerations 295

Chap. I.
Of the Individual Plant 296

Chap. II.
Of Vegetable Symmetry 302

Chap. III.
General Summary of the Structure of Plants .... 309

General Index to the Natural Orders and Genera men-
tioned in the Work 319

Description of the Plates 329

VEGETABLE OKGANOGKAPHY.

BOOK III.

OF THE REPRODUCTIVE ORGANS, OR THE PARTS
ESSENTIAL TO REPRODUCTION.

INTRODUCTION.

As soon as an organized being, and particularly a
plant, or a part of one, commences its visible existence,
it only presents to us a development of organs; whence
it has been concluded sometimes as a reality, at others
as a figurative expression, that all these beings proceed
from a germ. This name of Germ has been given to a
body imperceptible to our senses, which is supposed to
exist in organized bodies, and to be or to contain in
miniature the body, or the part of one, which proceeds
from it. The germs may be considered either as being
formed by the organ, or by the being upon which they
are developed, or by that which is transmitted to it at
the period of fecundation ; and in this case, the force
which causes this creation of germs, is termed the Plastic
Force : or, it is supposed that the origin of these germs
dates from the origin itself of organized beings, that they
were all inserted into one another ; so that all the germs

VOL. II. B

2 VEGETABLE ORGANOGRAPHY.

of a given species, which are and will be developed, were all
contained in one another, and all in the first which existed.

These two contradictory theories are so vast, that
they seem to contain all the opinions of which the
subject is susceptible, and, consequently, one of the
two must be true. However, if we reflect, one is almost
as unintelligible as the other; for — 1st, on the one
hand, nothing among known facts can make us compre-
hend the creation of a germ, since we never see anything
more in the Inorganic Kingdom than transformation of
compositions ; and, in the Organic, than developments ;
and, 2d — Nothing can make us conceive, either in
imagination, or in reason, an indefinite insertion of pre-
existing beings.

If we separate these questions, which are more meta-
physical than physical, and limit ourselves to general
facts, we shall see that an insertion, (if this idea be not
extended too far,) is proved by evident examples, as in
Volvox. We shall perceive, 2d, that the germs, or rudi-
mentary beings, are often visible a long time before their
ordinary appearance ; as, for example, when we find in the
centre of the trunks of palms the floral scapes which would
be developed externally after several years. 3d. We
shall be obliged to acknowledge that all beings are deve-
loped, as if their nutritive matter, deposited in an
invisible and pre-existing receptacle, had its place, thus
to speak, fixed before hand.

Thus, whether the expression of Germ be taken either
as a reality, or as an image, it will serve equally well to
describe the origin of organized beings. In the two
hypotheses, these germs arise from certain organs ; in
the theory of Plastic Forces they are formed by them, in
that of the pre-existence of germs they are simply
nourished and developed by their action. Whichever it
be, they are presented in plants in two different states ;
either they are dispo0ed in such a manner as to be

INTRODUCTION. O

developed as a natural effect of the laws of nutrition, as
takes place in the development of branches, tubercules,
offsets, layers, and suckers, for all these bodies may be
considered as resulting from the development of germs
more or less latent ; or their development requires a
preliminary operation, which has been named fecunda-
tion, which tends to give the germ a proper life, and
this is performed by means of a complicated apparatus
of organs, which collectively constitute the flower. Re-
production without fecundation, not presenting any
organic apparatus which is peculiar to it, will not occupy
us in detail before we treat of the functions of plants ;
whilst reproduction by fecundation, or sexual reproduc-
tion, being caused by numerous and various organs, will
henceforth engage all our attention. It has a double
claim in this respect, for it is important that all the
floral organs should be well understood, not only because
they perform one of the principal functions of vegeta-
tion, but also because it is upon their constant forms
within certain limits, varied ad infinitum in the different
species, and remarkable for their symmetry, it is, I say,
upon these forms, that all the classification of plants rests.
With regard to the general appearance of the organs
of fructification, plants are distinguished into Phanero-
gamous and Cryptogamous : the former are those
which have their flowers visible to the naked eye, more
or less symmetrical, and with distinct sexual organs ; the
latter are those which have the flowers (if indeed they
have any) visible only to the microscope, scarcely, if at
all symmetrical, and the sexual organs not distinct. The
former comprehend all Exogens, and the greatest part
of Endogens ; the latter all Cellulares, and some Endo-
gens. "We proceed now to study the organs of Re-
production, following this fundamental division of
Phanerogamous and Cryptogamous plants.

b2

VEGETABLE ORGANOGRAPHY.

CHAPTER I.

OF THE INFLORESCENCE, OR THE DISPOSITION OF THE
FLOWERS OF PHANEROGAMOUS PLANTS.

I designate, with botanists, under the name of the
Inflorescence, the collective distribution of the
flowers of a plant ; or, as Rceper calls it, that part of
the stems or branches which bears no other branches
but the floral axes. This term must be distinguished
from that of the Flowering, which means only the
expanding of the flowers ; the study of the inflorescence
forms an essential part of Organography, that of flower-
ing is essentially physiological.

The organs of the inflorescence are the supports of
the flowers comprised under the name of Peduncles,
or Pedicels, and the accessory envelopes of the flowers,
or the Bracts. We shall commence by first examin-
ing the general disposition of the flowers, and afterwards
study separately their supports and envelopes. Through-
out the whole of this chapter, we shall be principally
guided by the excellent Memoire of Turpin ; by the
ingenious ideas which Mr. Robert Brown has occa-
sionally advanced upon this subject in different places
in his works, particularly in his remarks upon the Com-
posite ; and by a very remarkable Memoire, which
M. Rceper has been so kind as to communicate to me ;
and lastly, by several observations of my own.

OF INFLORESCENCE. 5

Section I.
Of the Inflorescence in general.

A flower, considered in an organographical point of
view, is an assemblage of several verticils (usually four)
of foliaceous origin, disposed above or within one
another, and so close that their internodes are not
distinct.

These verticillate organs being, then, lateral, it would
seem that the stem or branch which bears the flower
would be prolonged beyond it; and this prolongation
does, in fact, take place sometimes accidentally : Turpin
has figured some examples, and I have myself observed
it in the Pear and Rose. Of the latter I have given a
figure here (PI. 17); but it is not usually so, and it
almost always happens in the natural course of things
that the flower truly terminates the branch, which is so
exhausted by the abundant nourishment which the
different floral organs attract, that it has not the vegeta-
tive force necessary for its prolongation ; this only takes
place, in the cases above mentioned, when the flower,
being sterile, attracts the juices but little, and the
branch at the same time is well nourished. It may,
then, be declared as a general law, that the flower is
terminal with respect to the branch which bears it.

This branch has received the name of Pedicel (Pedi-
cellus). It is sometimes long and distinct, at others
very short and hardly visible ; in this case it is usual
to say that the flower is Sessile, which only signifies in
Organography that its pedicel is very short.

Since, then, every flower is terminal upon the pedicel,
the whole study of Inflorescences ought to rest upon

6 VEGETABLE ORGANOGRAPHY.

the different positions of the pedicels, with regard to
the organs which bear them. They may arise either
immediately upon the stem or principal branch, or
spring from parts of the stem or branches which are
more or less different from ordinary ones ; in this case
these floral stems or branches bear the collective name
of Peduncles (pedunculi).

The pedicels may arise upon the stem or branches
after two systems — either laterally to the axil, or at the
extremity of the branch which bears them. We are
about to follow the numerous and varied consequences
of these two modes of inflorescence ; but, before entering
into any detail, it is necessary to say that the name of
Floral Leaves is given to those, from the axil of
which a pedicel proceeds, provided that they do not
differ from ordinary ones; and that they are called
Bracts, when they differ in size, colour, form, or tex-
ture. Bracts differ especially from ordinary leaves, in
never having true buds in their axils ; and in this
respect they nearly approach the verticillate organs,
which compose the flower.

We shall study the different inflorescences in the fol-
lowing sections: — 1st. Axillary Inflorescences. 2d.
Those which are terminal. 3d. Those which partake of
both these modes. 4th. Those which form, or seem to
form, an exception to the preceding classes.

OF INFLORESCENCE.

Section II.

Of Axillary, or Indefinite Inflorescences, or those with
a Centripetal Evolution.

The branches bear the flowers laterally, and in a
scarcely definite number. They may be terminated by
a flower bud, which we shall examine in the following
section : or by a bud, in this case, which forms the
subject of this section, sometimes the branch does not
flower; sometimes it bears flowers in the axils of the
leaves, and the branch itself may elongate by the
development of the terminal bud. Let us follow the
details of this axillary position of the flowers, and let
us first take the most simple cases.

If we examine the vegetation cither of the Periwinkle
(Vinca Major), or Veronica hederacea, &c, we find
that their stems or principal branches give rise to a
flower from most of their axils, and that the stem or
branch is prolonged by the apex ; and as the leaves of
the axil, from which the pedicels arise, are not per-
ceptibly different from ordinary ones, and as the length
of their internodes is well marked, we are content, in
describing the inflorescence of these plants, and all which
resemble them in this respect, to say, that their pedicels
are axillary and solitary. When the development of the
leaves, and all the organs of those plants, proceeds from
the base of the stem towards the apex, we remark that
their lower flowers are developed first, and the expan-
sion of them continues from the base upwards. Now,
from this which is observed so clearly in plants with
axillary pedicels, we shall proceed to discover it with
more or less decided variations in all those which have
not a terminal inflorescence.

8 VEGETABLE ORGANOGRAPHY.

It usually happens, especially in erect stems, that the
upper leaves, even when they do not bear flowers, are
smaller, and have their internodes shorter than the
lower ones, which results from their slower develop-
ment, and their receiving less nourishment. This
double effect is greatly augmented if these same upper
leaves bear a flower in their axil, probably because this
flower attracts a part of the nourishment, which would
otherwise have been employed either in making the
leaf larger, or elongating the internode ; in this case
the leaf is called the floral one, or bract, and the top of
the stem or branch, thus organized, receives the name
of the Terminal Raceme, or Spike. It seems, in fact,
to terminate the stem, but it is only formed of axillary
flowers, and the stem only ceases to elongate by the
exhaustion which it suffers in developing the flowers
and nourishing the seeds ; it then terminates in a point
by the simultaneous abortion of the flowers and bracts.
It is well known that by much nourishment these
branches can be made to elongate beyond their ordinary
dimensions ; sometimes they are naturally prolonged
in an unusual manner. Thus in the Pine-apple and
Eucomis, the axis of the stem is prolonged at the apex,
and it ceases to bear flowers; then the leaves which
were small and membranous where they had axillary
flowers, become large and truly foliaceous here where they
have none : it is this which forms the crown which sur-
mounts the spike of the Pine-apple and the raceme of
Eucomis. An analogous phenomenon is met with in
Galiistemon, and some other Myrtaceae of New Holland;
the axis of the spike is prolonged at the apex, and
forms' above the inflorescence, a true branch with leaves :
tins phenomenon also accidentally happens in some
cones, the axis of which is prolonged into a leafy
branch (PI. 16, fig. 4). It is to the inflorescence similar

OF INFLORESCENCE. 9

to what takes place in the flower when its axis is pro-
longed, as we have seen above.

We may connect with these facts that which happens
in Hoya carnosa, although it relates to the prolongation
of the peduncle, and not to that of the stem. The
peduncle, or floral branch, arises from the axil of the
leaves ; the first year it bears a kind of umbel composed
of pedicels, which are developed in the axils of very
small bracts. These pedicels disarticulate and fall off
after flowering, but the peduncle remains several years:
at each period of flowering it is prolonged a little at its
extremity, and it bears the traces of all the successive
flowerings, as if they were the remains of a raceme of
the same year. The fact is remarkable, as presenting
the only example that I know of a peduncle which
persists and flowers several years in succession.

There is so little difference between flowers said to be
arranged in racemes, or spikes, and those said to have
axillary pedicels, that it is not rare to find stems or
branches in which both states are united ; thus in seve-
ral species of Digitalis, and a multitude of other plants,
the lower flowers are solitary in the axils of large and
distant leaves, whilst the upper ones are in the axils of
bracts, which are small and near together. Descriptive
botanists have been accustomed to designate this inter-
mediate state by the terms of a raceme, or spike, in-
terrupted AT THE BASE, or WITH LEAVES AT THE
base. In a number of cases we see the lower flowers
solitary in the axils of leaves, which gradually diminish,
approach each other, and the flowers then form a true
raceme. All the difference between this case and that
of ordinary racemes is, that sometimes the transform-
ation of the floral leaves into bracts takes place sud-
denly from the first, which bears a flower in its axil ;
sometimes only gradually as they approach the top.

10 VEGETABLE ORGANOGRAPHY.

If now, instead of studying the formation of the
raceme in a single stem, we examine what happens in
the branches of a branching one, we shall find evidently
that each of them may present the same phenomenon,
and as the branches spring from the axils of leaves,
axillary racemes will thus be formed. These kinds of
racemes then are only floral branches : sometimes they
bear also at their base a certain number of leaves, which,
having no flowers in their axils, retain their natural
forms, and then they are considered as so many distinct
racemes, or we are contented to say, that the plant
bears several ; sometimes towards the base they have
their leaves provided with flowers and changed into
bracts, so that the axillary raceme is without leaves,
properly so called, then the whole is considered as a
single inflorescence, and the name of Compound Ra-
ceme is given to it. Thus all racemes which spring
from the axils of leaves only differ from terminal ones
in their being placed at the top of a branch instead of a
stem. Their flowers arise from the axils of bracts, or
floral leaves, and the whole branch from the axil of a
leaf.

All that we have said in taking the raceme for an
example is applicable, with very slight differences, to the
different kinds of indefinite inflorescence which we are
now about to pass rapidly in review, viz. — the Spike,
the Raceme, the Umbel, and the Capitulum, with the
varieties of each.

1st. The name of Spike (spica) is given to those
indefinite inflorescences where the flowers arise from the
axils of leaves, either sessile, or borne upon a scarcely
visible pedicel, as, for example, in the Plantain. The
limit between the spike and raceme is very uncertain,
seeing that it only rests upon an appearance ; in fact,
the pedicel always exists, and its length alone is

OF INFLORESCENCE. 11

variable : thus it is not rare to find inflorescences which
are racemes at the base, and spikes above, or which are
spikes in their infancy, and become racemes when they
are older. When several floral branches have flowers
in a spike, and are so close together as to seem to form
a single inflorescence, the name of Branching Spike
is given to it ; as, for example, in Statice s])icata, a
variety of Plantarjo lanceolate., Sec.

We describe, under the particular name of Catkin
( amentum), certain spikes which are remakable for this —
that after the flowering of the male flowers, or tire fruc-
tification of the female, has been performed, the axis of
the spike dries up, and disarticulates at the base ; such
are the male inflorescences of the Hazel, Oak, &c, and
those of the two sexes of the Willow. The difference
between the catkin and spike is less distinct in reality
than it appears to be ; and it frequently happens, for
example, that in the same species of Willow the male
flowers are in a catkin, or caducous spike, and the
females in a permanent one. There are some spikes,
the flowers of which are slightly pedicellate, and
approach the raceme. In Firs we find branching cat-
kins, formed of a central branch and several lateral
ones.

The name of Cone (conns strobilus) is given to the
female spikes of the Coniferae, which are furnished with
very large bracts. The female spikes of the Hop are
kinds of cones with membranous bracts.

The flowers of almost all the Gramineas are alternate,
and closely arranged along an axis, at the base of
which are found one, or, more frequently, two peculiar
bracts, which are called glumes. We give this inflo-
rescence the name of Spikelet (spicula, locusta); and
as these spikelets are found in almost all the Gramineae,
it is usual to say that their flowers are spiked when the

12 VEGETABLE ORGANOGRAPHY.

spikelets are arranged in spikes, as in the Wheat, and
that they are paniculate when they are in a panicle, as
in the Millet and Agrostis. *

The Spadix is also a kind of spike to which a par-
ticular name is given ; it is applied to the spike of
Monocotyledons, when they are enveloped in their
young state by a large sheathing bract, which com-
pletely surrounds them, and is called the Spathe. The
spadix is simple, in Arum, for example, and sometimes
it is covered with flowers throughout its whole length, as
in Calla; at others its summit is naked, as in Caladium :
it is branched in the Palms.

Besides these modifications of the structure of spikes,
to which it is thought fit to give particular names, they
also differ from one another: — 1st. In the distance of
the flowers, or the length of the internodes ; thus, the
flowers are very close in Plantago lanceolata, and very
distant in P. sparsiflora ; frequently the lower ones are
more distant than the upper (spica basi interrupta).
2d. In the relative position of the flowers : opposite, as
in Crucianella ; verticillate, as in Myriophyllum ; disti-
chous, as in Gladiolus ; or in a simple, double, or mul-
tiple spire ; characters always connected with the
disposition of the leaves. 3d. In the size and nature
of the bracts ; when these are large and foliaceous, the
spike is leafy (spica foliosa). 4. In the form of the
central axis, or rachis ; which may be cylindrical, com-
pressed, angular, or marked with depressions, in which
the flowers are as it were inserted. 5. In the general
form, which is usually cylindrical or conical, but some-
times oval or globular, and then it may be confounded
with the capitate flowers, of which we shall afterwards
speak.

2d. The Raceme (racemus) only differs from the
?pike in having longer pedicels, arising from the axils

OF INFLORESCENCE. 13

of bracts. In general those at the base of the raceme,
being older and more nourished, are the longest, and
they diminish in size as they approach the top. The
inverse takes place in a small number of cases ; thus, for
example, in Hyacinthus comosus the upper flowers are
sterile, with their pedicels coloured and very long,
forming a bunch or crown at the top of the raceme.
All the differences which we have just now seen found
in spikes when compared together, are also met with in
racemes, but without having any particular names. We
shall only say a few words on those which seem suffi-
ciently important to deserve special names.

W e have already said that those racemes are called
compound, or branched, which are formed by the union
of several partial ones into one inflorescence. When
these, or their partial branches, are very long, much
branched, and diffuse, the name of Panicle (panicula)
is given to them collectively ; for example, in Kcclreute-
ria. If the axis be very short, and the branches very
long and diffuse, as is seen in Juncus, the inflorescence
has been designated by the name of Anthela.

It sometimes happens in a simple raceme that the
lower pedicels are very long, and the upper very short,
whence it results that the flowers, although springing
from different points, nearly reach the same level. This
kind of raceme, confounded with other very different
inflorescences, has received the name of Corymb (co-
rymbus) ; the Ornithogalum, said to be umbellate, and
some species of Iberis, are examples of this.

The same thing may happen in compound racemes,
either because the lower lateral branches or racemes are
longer than the upper, or because each of them, con-
sidered separately, presents the same phenomenon, as
to the length of its pedicel ; this disposition is observed
in Viburnum, the Elder, &c. ; to this also the name of

14 VEGETABLE ORGANOGRAPHY.

Corymb is given, and when it is wished to distinguish it
from the preceding, we may, from analogy with the
racemes, of which they are modifications, designate
the first by the name of a Single Corymb, and the
second by that of Compound Corymb. But as several
very distinct inflorescences have been confounded under
the name of Corymb, I have reserved this name for a
particular class, which we will shortly examine, and I
shall designate by the name of Simple Corymbiform
Raceme, simple racemes, with the flowers placed on
the same level, and by that of Compound Corymbiform
Raceme, those which being compound present nearly
the same disposition. The reasons for this manner of
speaking will become evident when we treat of true
corymbs.

3d. The inflorescence in appearance the most distant
from the raceme, is the Umbel (ambella). This name
is given to an assemblage of one-flowered pedicels,
which all spring exactly from the summit of a branch, or
common peduncle. We distinguish the Simple Umbel,
as, for example, in the Cowslip or cultivated Cherry ;
and the Compound Umbel, which exists in almost all
the UmbelliferEe — it differs from the simple umbel in
its common peduncles being themselves disposed in
umbels ; we here distinguish consequently the General
or Universal Umbel, which is formed by the peduncles,
and the Partial Umbel, or Umbellcle, which is
formed by the pedicels. The umbel really differs less
from the raceme than it seems at first siffht. If we
compare, in fact, the different racemes together, we
shall find some, it is true, which have the axis very
long, as in the Ornithogalum of the Pyrenees ; but some
are also found with the axis much shorter, as in the
Ornithogalum said to be umbellate, which has its
flowers really in a raceme; lastly, we see racemes, the

OF INFLORESCENCE. 15

axis of which is so short that all the pedicles might
appear to spring from the summit, as in the Candytuft
(Iberis) ; and thus on comparing the corymbiform
raceme, we come to consider that the latter is a raceme,
the axis of which is wanting, or nearly so. I shall,
perhaps, be better understood by a rude metaphor.
Let us suppose a floral branch formed as a telescope,
which bears a pedicel above each of the tubes of which
it is composed ; when all the tubes are pulled out and
lengthened we have a raceme : let us push them half
way back, it will still be a raceme, but very short; let
us shut them up completely, and we shall have a terminal
umbel. When we compare together the inflorescences
of Eryngium and other Umbellifera?, &c, it is difficult
not to see this extreme analogy of umbels, with racemes
with a short axis. This analogy is also remarkable in
another respect ; at the base of each pedicel of an
umbel is found in general a bract, or small leaf, and
another is found at the base of the oeduncles of the
general umbel: thus, in this case, as in those we have above
analyzed, it is true to say, that the pedicels arise from
the axil of a leaf, and that the compound inflorescences
are formed by floral branches, which also arise from the
axils of their proper leaves.

4th. Botanists have confounded, under the name of
Capitulum, or Head, several inflorescences in reality
very different, and which present nothing in common
but very close flowers, either with the pedicels absent
or very short. Keeper has begun to give some exact-
ness to this incongruous assemblage, distinguishing the
glomerule, of which we shall speak under definite inflo-
rescences, and the capitulum, which makes part of the
indefinite ones. We may also even say that under the
name of capitulum we unite a particular state of each of
the preceding inflorescences.

16 VEGETABLE ORGANOGRAPHY.

Thus, when a spiked flower, instead of having the
axis elongated, is found to have an oval or globular
one, and as the flowers are very close around it, there
results an oval or globular spike which has often been
called capitulum ; such are the head of flowers of
the Plane-tree, Conoca?yus, &c, the female heads of
Sparganium, the globular spikes of several Plantains,
and some species of Phyteuma, &c.

When a raceme presents a very short axis, with
numerous flowers, and very short pedicels, there may
result from this union of circumstances a kind of globu-
lar head or capitulum ; this happens in Cephalanthus.
Likewise when an umbel has its pedicels very short and
its flowers very compact, it may resemble a true capitu-
lum ; this happens in several species of CEnanthe.

Capitula differ principally from one another in the
form of the axis : this is sometimes more or less elon-
gated, as in the examples mentioned as being derived
from the spike, sometimes it is short and more or less
expanded, as in those which appear to be derived from
the corymbiform raceme, or the umbel ; but all inter-
mediate degrees are found in the same families, such as
the Dipsacese for example. "When the axis is reduced
to a disc, much expanded, the name of Receptacle,
Phoranth, or Clinanth, is given to it, and then the
inflorescence receives that of Anthodium or Cala-
thide ; but although the terms to designate this kind
of inflorescence peculiar to the Compositae, and some
neighbouring groups, have been multiplied, it would be
difficult to establish a definition which would separate
those inflorescences from other capitula. The conical
receptacle in Anthemis, the oval one in Sphcsranthus, and
the oblong one in Rudbeckia, approach the elongated
ones of the capitula of Eryngium and Phyteuma, whilst
the flat ones of the Artichoke and Thistle would seem

OF INFLORESCENCE. 17

analogous to the discs which support the partial flowers
of the Umbel] iferae.

In all the inflorescences which we have enumerated the
law of expansion is simple and uniform ; everywhere the
lower and external flowers open first ; and the flowering
proceeds consequently from the base upwards in the
spike and raceme, and from without inwards in the
corymbiform racemes and the umbel ; it proceeds from
the base upwards in the spiciform or elongated capitula,
and from without inwards in the flat ones. This regular
progress of the expansion Rceper has called by the name
of Centripetal. It must, however, be remarked that,
as regards compound spikes or racemes, the central axis,
which is the prolongation of the stem or principal
branch, flowers before the lateral ones, each of which
follows in its turn its development in the same direction.
The only exception which I know to this mode of
development, is that which certain Dipsaceae present, the
flowering of which often commences in the middle of
the spike : this anomaly must be connected with some
peculiarity in the vegetation of these plants ; for, as to
their form, they cannot be separated from centripetal
inflorescences, and this mode is presented regularly in
the other species of the family.

Some of the inflorescences which have been enume-
rated in this section may be combined together. Thus,
the flowers of the Gramineae are, as we have said, dis-
posed in small distichous spikes, which have been called
spikelets ; and these, more or less pedicellate, are dis-
posed in panicles, sometimes fvery lax, at others
more or less compact ; the flowers of Carex are dis-
posed in close spikes, which are arranged in a raceme
along the central axis ; those of Papyrus are dis-
posed in spikelets/Avhich are pedunculate and arranged
in an umbel at the top of the stem ; the flowers in

VOL. II. C

18 VEGETABLE ORGANOGRAPHY.

Juncus and others are frequently found in capitula,
which are disposed in a contracted panicle. Thus may
not only these primitive dispositions be either simple or
branching, but the branch itself may present a dispo-
sition sometimes resembling, at others differing from,
that of the central axis.

A second difference which goes to prove the little
real importance of these divisions, in appearance so dis-
tinct, is, that it happens in plants with separate sexes
that the male and female flowers often present different
dispositions ; thus, the male flowers of the Indian Corn
are in branching spikes, the female in simple ones ;
the males of the Fir in catkins, the females in cones ;
the males of the Hop in panicles, the females in a kind
of cone or spike ; the flowers of Hura crepitans, although
springing from the same axil, are disposed in two man-
ners,— the females solitary, the males in a spike, &c.
Generally, in all cases of inequality of the two sexes, the
male flowers are more scattered and with longer pedi-
cels, and the female more sessile and compact.

Section III.

Of Terminal Inflorescences, or those with a Centrifugal

Evolution.

In this second class of inflorescence, the stem or prin-
cipal branch, instead of being prolonged indefinitely in
a straight line, and only bearing the flowers laterally, is
found terminated by a flower, which, instead of arising
from the axil of a bract, is found to have at the base of
its pedicel two opposite, and sometimes several verticil-

OF INFLORESCENCE. 19

late bracts. Let us examine the first case on account
of its greater simplicity.

In the axil of each of the two bracts there may arise
a branch, which, as happens in the primitive one, is
found in the same manner terminated by a flower with
two bracts, which, in their turn, may produce two
brandies, and thus indefinitely. There results from
this disposition a series of bifurcations, in the centre of
each of which is found a solitary flower : the inflores-
cence in this sense is terminal, since each flower termi-
nates its branch; it is indefinite in this respect — that
each branch may, from the axils of the two bracts, give
birth to new branches, which enjoy the same faculty ;
whence it results, that in this case, as in the preceding,
there is no limit to the elongation of the plant and the
development of flowers, except by the exhaustion pro-
duced either by the paucity of nourishment or the
greediness of the floral organs. We designate under
the collective name of Cyme (cyma) all the inflores-
cences of this kind; calling those Diciiotomous Cymes,
where the flower is provided with two bracts, and where
the branches go on bifurcating without cessation ; this
is most frequently the case in Dicotyledons ; for ex-
ample, in Erythrcea, Kalanchoe, most CaryophylleEe, &c.
In the same manner we call cymes Triciiotomous,
Tetrachotomous, Pentachotomous, &c. where each
terminal flower has under it three, four, or five bracts
which give birth to as many branches ; the Euphor-
biums present examples. Sometimes, in these different
systems, the central flower is abortive, and thus we
might at first sight confound them with umbels or com-
pound corymbiform racemes : the order of evolution,
which will engage our attention presently^, is sufficient
to remove this doubt ; here the central flowers, or those
which terminate the branch, expand first, whilst, in

c2

20 VEGETABLE ORGANOGRAPHY.

corymbiform racemes and umbels, the lateral ones are the
first to expand.

A second remarkable difference which is observed in
cymes, especially in dichotomous ones, is, that of the two
branches which ought to be developed in the axils of
the two bracts, one is sometimes abortive, and then the
terminal flower seems lateral ; this is very clearly ob-
served on comparing the Silenes said to have spiked
flowers, with those with an evidently dichotomous inflo-
rescence. In this case the flowers are generally dis-
posed upon only one side, either by a tendency of the
branches to be abortive on that side, or by a torsion of
the axis. The branches or stems in which this dispo-
sition takes place are in general, before their develop-
ment, rolled up : this is observed in Drosera, the cymes
of which have the flowers unilateral ; in the Silenes,
said to be spiked ; in the branches of the cymes of
Sedum ; in those of Echium, and other Boragineae. I
give to these cymes, the flowers of which seem unilateral,
the name of Scorpioid Cymes, which makes allusion
to their mode of development.

The different dispositions of cymes of which we have
spoken, may be combined together ; thus, several species
of Sedum present a general cyme, the central flower of
which is abortive, and which is divided into several
lateral branches, some of them dichotomous at the base,
others simple, and with unilateral flowers, on account
of the abortion of the secondary ramuscules. When a
cyme has the lateral branches very short, the flowers
are found agglomerated together ; this is seen, for ex-
ample, in the Sweet- William. Rceper gives to this
disposition the particular name of Fascicle (fasciculus),
which, on acount of its vague nature, is found applied
by different writers to other dispositions of flowers.
That of Contracted Cyme (cyma conlracta) would

OF INFLORESCENCE. L2l

possess some advantage, in ray opinion, because it would
make known the nature as well as the appearance of
this disposition.

Lastly, the same naturalist proposes to call Glome-
RULE (glomerulus) those cymes which are so contracted
that their ramification is scarcely apparent, appealing
at first sight to be true capitula ; but they differ, be-
cause the flowering commences at the centre instead of
the circumference. This disposition, less frequent than
that of true capitula, is observed in Corymbium and
some other Composite. Cardojiatum and the Euphor-
biums have the flowers in glomerules which are dis-
posed in cymes.

In all the inflorescences which I have pointed out,
the central flower of each ramification always expands
before those which terminate the branches situated
above it ; so that, where the flowers are near together
in a fascicle or glomerule, or corymbiform or umbelli-
form cyme, the flowering proceeds from the centre to
the circumference, and the evolution, for this reason,
has been called Centrifugal by Rceper.

When centrifugal inflorescences are reduced to a
single flower, it seems impossible to distinguish them
from the one-flowered pedicels of indefinite inflores-
cences ; but there are almost always means of recog-
nising them ; and, in particular, the pedicels of the
indefinite have only one bract at their base ; those of
terminal inflorescences have two opposite ones, and
sometimes a third lateral one, when the cymes them-
selves arise from its axil.

Notwithstanding the extreme difference which is
found between the two systems we have explained, there
are numerous cases where the two modes are combined
in the same plants ; we now proceed to examine this
in the following section under the name of Mixed Inflo-
rescences.

22 VEGETABLE ORGANOGRAPHY.

Section IV.

Of Mixed Inflorescences, or those which partake of the

two preceding.

Inflorescences may be mixed after two different sys-
tems; viz. either, 1st, because the central axis proceeds
in the manner of indefinite inflorescences, and the lateral
branches follow the progress of terminal ones ; or 2d,
because the central axis proceeds in the manner of ter-
minal inflorescences, and the lateral branches follow
the progress of indefinite ones.

To the first of these divisions belong the true Thyrses ;
to the second, the true Corymbs. Let us examine the
modifications of these two inflorescences, and their par-
ticular affinities with the two preceding classes.

§ 1.— Of Thyrses.

If I examine one of the Labiata?, I see that the stem
or branch is prolonged indefinitely by its extremity, and
that the pairs of leaves may be there developed succes-
sively, one after the other, without any other natural
termination than that of the vegetation ; all the inflo-
rescences proceed from the axils of the leaves, and each
is a di- or trichotomous cyme. Thus, the whole inflores-
cence of the Labiatae is a Thyrse, interrupted by the
distance of the internodes and the great development of
the leaves from the axils of which the cymes are deve-
loped. When these cymes are very lax, this disposition
is very evident ; when they are close and compact, so
as to form axillary fascicles, the union then of two

OF INFLORESCENCE. 23

opposite ones forms a kind of ring or false verticil
around the stem ; and for this reason the Labiatae have
been frequently confounded with true verticillate flowers,
which are very rare in nature. It sometimes happens
that the cymes of the Labiatae are composed of a very
small number of flowers ; they may even be reduced to
one, without the fundamental type of the inflorescence
being altered; for in this case the floral pedicule pre-
sents two opposite bracteoles, from the centre of which
rises the proper pedicel, and from the axils of which
ought to proceed the lateral ramuscules. It happens in
some Labiatae that the cymes only arise from the upper
axils ; then the leaves are reduced to the state of bracts,
and the internodes are much shortened. These pheno-
mena cause the continuous thyrse to be rather compact,
sometimes in the form of a raceme, as in Clinopodium ;
at others, in that of a spike, as in Lavandula. In these
racemiform or spiciform thyrses, it sometimes happens
that the upper branches cease to bear flowers, are more
or less coloured, increase much in size, and form at the
summit of the thyrse a bunch, which recalls to mind
that of Eucomis : this is observed in Salvia Horminum,
Lavandula slcechas, <xc.

All that I have said of the Labiatae is equally appli-
cable to the Lythrariae, in which are found sometimes,
as in Ammannia, lax lateral cymes ; sometimes, as in
Lythrum, short ones, which collectively resemble either
a terminal spike (Salicarice) or simple axillary flowers
(Hyssopifolice).

The comparison of the species of Eugenia together
presents a clear example of the apparent modifications
which may result from the system of interrupted thyrses.
We here find species which seem to have a simple one-
flowered pedicel ; but this apparent pedicel bears two
opposite bracteoles, and it ought to be considered as a

24 VEGETABLE ORGANOGRAPHY.

pedicule, from the apex of which proceeds a terminal
flower, and frequently two lateral branches from the
axils of the bracteoles : when these branches are deve-
loped, the cyme is bifid and three-flowered ; when
subsequent ramifications take place, then a true di- or
trichotomous cyme is formed. If, however, this last
state takes place towards the top of the branches, as the
leaves may be but little developed, the internodes near
together, and the principal stem but slightly disposed
to elongate, then the union of these lateral cymes forms
what is called a terminal panicle, which is only a true
thyrse, with ramified branches, or a Paniculiform
Thyrse.

The example of the thyrse of Eugenia leads us to
understand several inflorescences which resemble also
racemes or panicles ; such are the thyrses of the Lilac.
Here the floral branch only bears at its base a small
number of leaves ; the axis elongates after the system
of indefinite inflorescence ; and the lateral branches (in
the axils of leaves reduced to the state of very small
bracts) are true cymes, the union of which forms a
thyrse : this also happens in the Vine, &c.

Several Leguminosas present analogous phenomena ;
thus, there is a great number of species where we see
racemes apparently simple and resembling true ones ;
they resemble them, in fact, by the possibility of inde-
finite elongation, and the axillary position of the pedun-
cles ; but each peduncle bears two opposite bracts, from
which arise either only a terminal pedicel, or one with
lateral ramuscules ; thus, in all these plants, the raceme
is in appearance almost indifferently simple or branched,
and ought to be considered as a true Racemiform
Thyrse.

Several Monocotyledons are found, which, it is said,
have their flowers in spikes, with three bracts at the

OF INFLORESCENCE. 25

base of each flower — as Pitcairnia. Of these bracts,
the lower one represents the true leaf; the two others
are the bracteoles of a very short peduncle, and the
whole forms a Spiciform Thyrse. In neighbour-
ing species we see these peduncles elongated into
true cymes.

Lastly, we have seen, in speaking of Cymes, that their
branches seem, from the abortion of one of them, to
bear lateral flowers : when this phenomenon is combined
with that which I have mentioned, there results a sin-
gular kind of inflorescence, which may be seen in the
woody kinds of Echium ; the axis lengthens indefinitely
by its summit, and bears branches laterally, which arise
from the axils of leaves changed into bracts. These
branches are true c\mes, reduced by abortion to only
their terminal flowers which appear lateral, or scorpioid
cymes ; the whole then is a thyrse with scorpioid cymes,
or, if we wish, for the sake of abbreviation, a Scorpioid
Thyrse.

Thus, Thyrses are systems of inflorescence, of which,
1st, the central axis follows the laws of indefinite inflo-
rescence, and may present all its modifications, such as
the states of spikes, elongated corymbiform racemes,
umbels, &c. ; and of which, 2d, the lateral branches
follow the laws of terminal inflorescence, and may pre-
sent all its modifications, viz. the state of dichotomous,
trichotomous, scorpioid cymes, fascicles, and glome-
rules. The evolution of these two systems follows the
laws of each : — the development of the central axis and
its parts proceeds from the base upwards ; that of each
of the lateral branches commences at the centre and
follows a centrifugal course.

26 VEGETABLE ORGANOGRAPHY.

§ 2.— Of Corymbs.
The inverse to all that we have yet observed takes
place in the Corymb {corymbus). This term has
hitherto, in all the writings of botanists, been used in a
vague sense, and solely founded upon appearance. I
propose to limit it to a very distinct case, which deserves
a special name, viz. the case of inflorescences where the
central axis follows the laws of terminal inflorescence,
and the lateral branches, that of indefinite ; almost all
the Composite are examples of this system, and the
name of Corymbiferae has for this reason been given to
several of them. If we follow the development of Tolpis,
or of most of the Composite, we see that the central
axis terminates in a capitulum, and that the lateral
branches are developed in a centrifugal order ; those
which are nearest the central capitulum (which may
be here provisionally considered as a flower) expand
first ; but all these successive capitula, which, compared
together, follow the centrifugal evolution, are indivi-
dually subject to the laws of the centripetal, in each of
them the expansion of the flowers proceeding from the
circumference to the centre. When the corymbs are
very near together, as, for example, in Cardopatum, the
expansion appears perfectly irregular, because the
flowers of each capitulum, or the capitula of the whole
corymb, follow two different systems of evolution.
When the partial capitula are reduced to a single flower,
the whole evolution is centrifugal, in which this head,
composed of capitula, differs much from true capitula ;
this happens in Echinops. When the capitula are soli-
tary, or, in other terms, when the lateral branches are
not developed, the single capitulum flowers after the
system of indefinite inflorescence alone, and then the
head of a monocephalous compound flower does not
differ from that of other flowers in capitula.

OF INFLORESCENCE. 21

Section V.

Of Anomalous Inflorescences, or those which seem to form
an exception to the preceding laivs.

The general systems of inflorescence which we have
examined in the three preceding sections, might appear
to include all phanerogamous plants, but there are some
which form an exception to the general laws ; such are
the inflorescences said to be opposite the leaves, radical,
extra-axillary, petiolary, epiphyllous ones, or those
which are modified by unions, abortions or degene-
rations.

§ 1. — Inflorescences opposite the Leaves.

Inflorescences opposite the leaves appear to be always
formed by the real top of the stem. One may be con-
vinced of this by the following considerations : —

A leaf provided with its axillary bud may be consi-
dered as the point of departure, or origin of two distinct
productions: — 1st, a bud, which may be developed into
branches with leaves or flowers ; 2d, the branch, which
is the prolongation of the same stem which bears the
leaf. Two things may happen in the development of
these bodies ; the one which is the most simple, is that
the continuation of the stem is stronger and more
vigorous and forward than the axillary bud, which then
developing after the other, and weaker than it, always
retains the lateral position, and forms, consequently, an
axillary branch if it have only leaves, or an axillary
inflorescence if it have flowers ; this latter is most fre-
quently the case, and we have examined it in § 2, 3,

28 VEGETABLE ORGANOGRAPHY.

and 4. The other, which takes place in a smaller
number of plants, is that where the axillary bud grows
so powerfully and rapidly as to cause two appearances
at once, viz. that it resembles the continuation of the
stem, and that the true stem is pushed to the side oppo-
site the leaf. In this state of things, less rare than is
imagined, two different cases happen, either by the
more or less forward disposition of each of these organs,
or by their place upon the stem.

1st. Sometimes the axillary bud, thus developed into
a branch which seems terminal, receives such strength
as to flower first, attracts all the juices, and then the
real summit of the stem, thrown on one side under the
form of a branch, becomes abortive, and perishes. In
this case the raceme which is formed, although really
axillary, is said to be terminal ; it is this which takes
place in several Cruciferae.

2d. Sometimes the axillary bud, developed into a
branch and replacing the stem, has less tendency than
the latter to flower ; and then this top of the stem,
pushed on the side opposite the leaf, absorbs propor-
tionally sufficient juice to sustain it, and begins to
flower under the form of an inflorescence opposite the
leaf. All who will follow the development of the
flowers opposite the leaves in the Cruciferae, Umbelliferae,
Leguminosas, and, in general, in all plants with alter-
nate leaves, will, I think, be convinced that it is in this
manner that the phenomenon takes place. They will
even be able to give an easy account of the details of the
phenomenon ; thus one may understand by this theory,
why the stem is often bent in a zigzag manner in the
species where the inflorescences are opposite the leaves.

3d. When the fact which I have just mentioned takes
place at the bottom of the plant, neither the axillary
bud nor the stem itself being disposed to flower, it only

OF INFLORESCENCE. 29

results from the rapid growth of the bud, that the true
stem takes the appearance of a branch opposite the
leaf; and if the two productions have an equal degree
of development, the stem is said to be forked, or, when
the same phenomenon is frequently repeated, dicho-
tomous.

4th. When it takes place towards the top of the
plant, and the two productions have nearly an equal
force, or an equal disposition to flower ; then, according
to circumstances often very slight, the inflorescence
seems either terminal or opposite the leaves ; and this
explains why, in the descriptions of different authors,
we so frequently find these two expressions taken the
one for the other, especially in the families which I
have mentioned.

§ 2. — Radical Inflorescences.

Flowers are said to be radical when they seem to
spring from the root : but this term ought only to be taken
as a simple metaphor ; for the inflorescence never arises
anywhere but from the stem. Pedicels having a single
flower, or floral branches, which bear several, are said
to be radical in some cases ; viz. sometimes, when the
stem is very evident, these pedicels arise from the lower
axils only, as in Vinca herbacea ; sometimes, when it is
so short and level with the surface that it is with diffi-
culty distinguished from the root, as in Mandragora,
the leaves are very near the neck, and the peduncles
arise from their axils ; sometimes, when the stem is
entirely subterranean, the leaves are reduced to the
state of scales, either fleshy or scarious, and the pe-
duncles which spring from their axils proceed from the
earth, as if the root gave birth to them (we see this in

30 VEGETABLE ORGANOGRAPHY.

bulbous plants) ; sometimes, finally, the stem, although
long, is buried under the earth or water, and produces
true leaves, which have, as usual, axillary peduncles ;
this takes place in Nymphcca, Utricularia, Sec. Thus the
various flowers said to be radical do not differ from
ordinary ones as to their anatomical origin.

§ 3. — Lateral, or Extra-axillary Inflorescences.

It is usually said, that flowers are lateral, supra-
axillar}r, or extra-axillary, when they seem to arise from
the stem beyond the axils of the leaves. This pheno-
menon ought to be referred to two classes : sometimes,
as in Solarium, it is a true anomalous development, ana-
logous to that which renders their leaves geminate :
sometimes it is a simple case of union ; the peduncle
which arises from the axil is sometimes intimately con-
nected with the branch which gives birth to it ; then the
flower or flowers which it bears seem, according to their
direction, to arise from the branch at the point where
the junction terminates : examples of this phenomenon
are found in different families, but it nowhere presents
a more singular appearance than in a small section of
Capparis (Capparides seriales), where the flowers are
arranged three, four, or five, in succession, in a longi-
tudinal series above the flower (PI. 16, fig. 2); this is
a unilateral spike adherent to the branch.

§ 4. — Petiolary Inflorescences.

Flowers are said to be petiolary when they spring
from the petiole of the leaf: this illusion takes place in
two cases, viz. with regard to the petioles of simple
leaves, and those of compound ones.

OF INFLORESCENCE. 31

The first is also a case of union, the inverse of the
preceding. The peduncle which arises from the axil
unites sometimes with the petiole, and then the flower
or flowers which it bears seem to arise from the petiole
at the point where the union ceases ; this is very visible
in Chailletia, where the flowers are upon the same
branches, sometimes evidently axillary or petiolary,
according as the peduncle is free or adherent to the
petiole ; it is also remarked more or less constantly in
several species of Hibiscus. The union, on the con-
trary, is constant in Tapura, for example, and several
others. The position of the flowers of Thesium may
result from the union of the pedicel with the leaf or
petiole.

The second example of petiolary flowers is that of
those which spring, it is said, from the common petioles
of pinnate leaves ; as, for example, in several species of
Phyllanthus. These flowers always arise from the axils
of the organs which are called leaflets, when the axis
receives the name of petiole ; but the truth is, that what
is called a compound leaf is a branch with alternate
leaves, and consequently the flowers are axillary as
usual. "What is remarkable in this kind of branches
(which Martius has happily designated by the name of
Rami pinncpformes) is, that their base is articulated with
the stem. The Zizyphus, after some years, elucidates
the nature of these branches which resemble leaves :
when an old one is observed, we see here and there
kinds of thick knots, from which arise eight or ten small
branches in a bundle ; each of these is simple, and bears
alternate leaves, and frequently flowers, in its axil : in
the autumn, a part of these branches disarticulate and
fall off, whilst some remain and become true persistent
branches which are not capable of being disarticulated.
It is impossible for any one to have followed the vege-

32 VEGETABLE ORGANOGRAPHY.

tation of this plant without seeing the truth of what I
have just set forth ; and I have concluded that it must
he the same in those species of Phyllanthus said to have
pinnate leaves. The manner in which Martius describes
those which he has observed, proves that he has arrived
at the same results, and that consequently the pre-
tended flower-bearing leaflets of these plants are penni-
form branches. The observation of Phyllanthus Cochin-
chinensis in a living state has confirmed my opinion :
this case then enters into the general law of axillary
flowers.

§ 5. — Epiphylious Inflorescences.

Flowers are said to be Epiphyllous, or arising from
the leaves, in four cases : the first, which enters into one
of the preceding, is where the peduncle is very inti-
mately united along the petiole (if it exist), and the
middle nerve of the leaf, so that the flowers appear to
arise from the limb at the point where the union ceases :
this is what seems to take place in Polycardia, where
the union takes place as far as the top of the middle
nerve ; in these cases, if the peduncle bear any bracts at
its apex, the flowers seem to arise from a disc of the
leaf.

The second case deserves this name still less ; it is
where the floral branches are large, dilated, green, and
flattened in the form of leaves ; we see this, for example,
in Xylophylla (PI. 16, fig. 1) and Opuntia ; but it is so
true that these bodies which bear the flowers are
branches and not leaves, that when their subsequent
development is followed, they are seen to change gra-
dually into cylindrical branches, furnished with others
like what they themselves originally were.

OF INFLORESCENCE. 33

The third case is that which Moreea Northiana pre-
sents, the flowers of which are said to spring from the
edge of the leaf; but this is also an example of the
necessity of distinguished primitive and modified forms :
this Morcea has, like all the others, a peduncle furnished
with flowers, but it is surrounded as far as the origin
of the flowers by a leaf folded upon itself, and which
surrounds it so closely that the flowers seem to proceed
from it. Zostera presents in the same manner a leaf
folded lengthways, and the flowers spring from a
peduncle united to this folded leaf.

The fourth case is that of lluscus, which seems to owe
its appearance to its leaves arising from foliiform
branches of an entirely peculiar form ; when the deve-
lopment of a young plant is followed (PI. 16, fig. 3),
we see that the true leaves, as in the Asparagi, are
caducous and slightly embracing scales, whilst the very
compressed organs which spring from their axil are
true branches which are destined to bear flowers. In
some species, as in 11. Hypoglossum, this foliiform
branch bears, besides flowers, a true floral leaf, the pre-
sence of which confirms the nature of the branch.

Section VI.
Of Pedicels and Peduncles.

It is usual to designate under the special name of
Pedicel (pedicellus) the immediate support of each
flower; and to reserve those of Pediclle or Peduncle
(pediculus, pedunculus) for all the ramifications of the

VOL. 11. D

34 VEGETABLE ORGANOGRAPHY.

general axis, or rachis. It evidently results from the
ideas contained in the preceding Section, that the pedicel
ought in fact to he considered as a proper organ ; but
that the axis, and all the ramifications of compound
inflorescences, except the pedicels, can only be con-
sidered as floral branches. However, in order to con^
form to usage, and to avoid circumlocution, I shall
employ the words Pedicule and Rachis, in their ordinary
sense. I have already explained that which relates to
the general distribution of these pedicels ; it now remains
to examine their forms, articulation, and history.

Pedicels, properly so called, are always terminated by
a single flower ; unless two be joined together, as in
several Honeysuckles, and then an apparently single
one seems to bear two flowers or fruits. These sup-
ports are in general either strictly cylindrical, or a little
expanded into a reversed cone below the origin of the
flower, or slightly compressed. Their length is some-
times very considerable ; it is described either in refer-
ence to that of the calyx or flower, or in comparison
with the size of the bract or leaf, from the axil of which
it proceeds. When it is so short that no interval between
the axil and the flower can be distinguished, the latter
is said to be sessile, or to have the pedicel wanting ; but
we may say that in reality it always exists, although
sometimes hardly visible; and consequently this cha-
racter is uncertain, as are all those which relate to the
degree of development : it very frequently happens that,
in the same species, often in the same inflorescence,
either in different places or at different ages, the pedi-
cels are either so long as to be distinct, or apparently
entirely wanting.

The Pedicules present in general more varieties of
form than pedicels ; for, independently of those which
are common to both, there are several others which

OF INFLORESCENCE. Go

depend either upon the mode of inflorescence, or upon
what we confound under the name of peduncle — the
several different degrees of the ramifications. In gene-
ral they present a more cylindrical form in racemes and
spikes, and have a tendency to dilate at the top in
umbels.

In the former, the principal difference which they
present is that of being either really cylindrical, or more
or less compressed ; this compression in some cases,
goes so far as to give them a form, expanded into a
strap* as in certain species of Eugenia and Eucalyptus.

Sometimes this strap-like form appears peculiar to
the species, without our being able to determine the
cause : — sometimes it seems to result from the compres-
sion of neighbouring organs ; thus the peduncles which
proceed from several bulbs are compressed, at least at
their base, by the pressure of the coats : sometimes it
appears to be owing to the peduncle being bordered by
a foliaceous membrane ; this seems to take place in
RascuSf and more evidently in Urtica membranacea ;
in this case the flowers arise from the middle of the
membrane which represents the true pedicule. Some-
times the compression is owing to a kind of dilatation
or foliaceous expansion of the peduncle, as in Xylophylla
(PI. 16, fig. 1). When the peduncles are much com-
pressed, the pedicels usually arise from the angles, and
not from the plane surfaces — for example, in Xylophylla;
whence it results that they strictly alternate, and if
they are near together their flowers are said to be
distichous.

In umbels, or umbelliferous cymes, the peduncles have
a tendency to be dilated at the summit; and this is con-
nected with two circumstances : it is large in proportion
as the number of flowers which ought to be found placed
upon the peduncle is large ; it is also generally larger

d2

HC) VEGETABLE ORGANOGRAPHY.

in proportion as the flowers are more nearly being
sessile upon the horizontal expansion produced by it.
This expansion bears the name of the Receptacle of
the flowers (receptaculumj ; some have given it the
name of Phoranth or Clinanth.

When the flowers are not numerous, as in di- or tri-
chotomous inflorescences, or when they are provided with
distinct pedicels, as in most Umbelliferas, the receptacle
then differs so little from the other parts of the rami-
fications, that this name is but seldom applied to it ; but
in this case, as in those where it is more evident, the re-
ceptacle or common point of departure of the branches
of an umbel, is a more or less dilated portion, in which
is deposited, before the flowering, a certain quantity of
nutritive matter, which serves for the further develop-
ment of the flowers or fruits. All many-flowered
receptacles are likewise generally very thick and fleshy.
This deposition of nutriment which is stored up for the
flowers, is appropriated by men and animals for their
own use ; thus we take the receptacles of the Fig, the
Artichoke, and other syngenesious plants, precisely for
the same reason which makes us select tubercules and
fleshy cotyledons, because we find in them a deposition
of nutriment ; thus a great number of insects establish
themselves in the receptacles of capitulate or umbellate
flowers, because they not only find there a shelter, but
also abundant nutriment. When insects attack them
(and floriculturists know that this is but too frequent,)
they always direct their efforts at the point of departure
of the rays, which represents the receptacle, and which
contains the store of nourishment intended for the
flowering.

The covered receptacles of sessile flowers are of a
whitish colour, being etiolated by their being shaded
from the fight ; before flowering they are in general

|->C

OF INFLORESCENCE. 37

rather thick, and become thinner, or at least part with
a considerable portion of their store, during that period ;
therefore care is taken to select those for culinary pur-
poses before this takes place. Those of the Chicoracea*
empty themselves very early ; those of the Cinarocephalae
remain a longer time fleshy, but at maturity they only
present a tissue resembling an empty pith ; in some the
inverse takes place, as in the Fig, which becomes fleshy
on approaching maturity. It is, perhaps, worthy of
remark, that in all plants with milky juice, the recep-
tacle, at the period of flowering, is full of a juice of a
different nature ; thus the Fig, and the receptacles of
all the milky Compositae, are filled with this juice before
flowering, and cease to receive it or form it after that
period has commenced.

The receptacles are sometimes in the form of a cylin-
der or elongated cone, as in the eapitula of spiked
flowers — Dipsacus and Eryngium, for example ; some-
times in that of a short cone, or simply convex, as in a
great number of Compositae or Dipsaceae ; sometimes
flat, or even slightly concave, as in most Compositae and
in Dorstenia. Sometimes the margins of the receptacle
are elevated, and cover the flowers ; we see this in Dor-
stenia : it is more decided in the Fig, where they en-
close all the flowers in a kind of envelope, with a very
small opening at the top.

At maturity, the receptacles undergo changes of
form, which facilitate or cause the fall of the seeds.
Flat or convex ones become raised in the centre, and
thus throw off the seed; concave ones open by the
reflection of their margins, as is seen in the Fig when
left to itself, and still more in Ambora.

Peduncles which arise from a stock which is under
ground, or on a level with the surface, have received the
particular name of Scapes (scapl); they only differ

88 VEGETABLE ORGANOGRAPHY.

from ordinary brandies in their never bearing true
leaves, but only bracts or floral ones; thus the scape
which supports the head of flowers of the Daisy, or the
solitary one of the Cyclamen, or the spike of the Plan-
tain, is entirely devoid of true leaves.

Pedicels and pedicules are often furnished with arti-
culations, the study of which presents some interest,
both on account of their causing the fall of fruits, and
because they elucidate the true structure of the organs
of inflorescence.

Pedicels often appear articulated in the middle of
their length ; but it must be remarked that when this
phenomenon takes place, either near the base or apex,
or really at the middle, we observe below the articu-
lation two small bracts; which indicates that we ought to
consider as terminal or compound inflorescences, the
cases where such articulations are found, and to reserve
the name of pedicel for the part above it, and which
bears the flower. What tends to confirm this opinion is,
that it happens in a great number of plants — several
Myrtaceae, Leguminosae, &c. for example — that where
such an articulation is found, a second or third pedicel
is seen to spring from it, proving that the lower portion
is a true pedicule, and not a part of the pedicel. In
the same manner we here and there meet with articu-
lations in different parts of the system in compound
inflorescences ; but what is more remarkable is, that
sometimes the floral branch itself is articulated at its
base : this is what is observed in the spikes or racemes
of several Amentacees, to which the particular name of
Catkin (amentum) has been given — a name which,
from analogy in form, has been sometimes extended to
the spikes or racemes of some species of these families
where the articulation does not exist. We meet with
this articulation, and consequently the fall of the

OP INFLORESCENCE. 39

entire system of flowers or fruits, in the Mulberry,
Fig, &c.

The study of the vegetation of the pedicules and
pedicels forms a part of physiology rather than
of Organography: I shall limit myself to remarking
here —

1st. That we frequently see them take different and
fixed directions before, during, and after flowering : in
general they spring upright, and fall as they advance in
age; but different plants present in this respect very
curious physiological phenomena.

2d. They also sometimes, on becoming old, alter
their length in a very decided manner.

3. They also sometimes change in texture : thus, that
of Anacardium, which bears the Cashew-nut, becomes
so fleshy after flowering as to have nearly the form and
size of a pear ; that of the Fig is so pulpy as to be con-
sidered a true fruit. On the contrary, some remain
and dry up after flowering, so as to take the appearance
of true spines, as we see in Mesembryanthemum spino-
sum, Alyssum spinosum, &c.

4th. Some pedicules, when their flowers are abortive,
are transformed into elongated processes, which are
named Tendrils, and of which we shall speak in the
next Book.

5. Some peduncles, especially among those which
arise from near the neck, and which are called Scapes
(scapi), present a disposition to twist themselves in a
regular spire, in a manner analogous to twining stems,
and sometimes even more decided : this is observed in
the scapes of Cyclamen, and in those which bear the
female flowers of Vallisneria. These, as is well known,
elongate so as to elevate the flower to the surface of
the water, unrolling the turns of their spire ; and con-
tracting again after flowering, in order to bring back

iO VEGETABLE ORGANOGRAPHY.

the fruit to ripen at the bottom of the water. All
that we have said of twining stems is applicable to
these scapes.

Section VII.
Of Bracts.

Bracts, in general, are the leaves from the axil of
which the floral branches, their ramifications, or even
the pedicels themselves, proceed ; they differ from
ordinary leaves in form, size, colour, &c. or, at least,
what is more constant, in their not bearing true buds
in their axils, the flowers replacing them.

That bracts are only leaves modified by their position,
it is hardly necessary to endeavour to establish, seeing
that the slightest inspection of them suffices to show it.
This opinion is especially demonstrated by frequent
cases where the bracts change into true leaves, as hap-
pens in several Cruciferse, Plantains, &c.

In simple inflorescences, such as the raceme of a
Hyacinth, the pedicels all spring from the axils of bracts,
and there is no difficulty in distinguishing these organs ;
but in compound racemes there are as many different
orders of bracts as degrees of ramifications. The com-
mon name of bracts is given to all, except in one single
case, — that where the last ramifications of a compound
inflorescence bear pedicules terminated by a single
pedicel, or, as is commonly said, when the pedicels are
articulated in their length ; then the little bracts which
are found at this articulation are sometimes called
Bracteoles. This distinction is not strict, but it is,

OF INFLORESCENCE. 41

convenient in practice for avoiding long circumlo-
cutions.

Bracts are, as we have said, leaves modified by the
production of flowers, which, developing in their axil,
attract a great part of the sap ; whence it results that
they are in general smaller, less divided, and more
membranous than the leaves of the plant. Frequently
they participate, as well as the pedicels, in the colour of
the flower, as is seen in Hortensia, (of which that which
is commonly called the flower, is essentially formed of
coloured bracts,) in Salvia splendens, Melampyrum, &c.
The last presents the double singularity of the coloured
bracts being larger and more divided than the leaves.

The colouring of bracts takes place the more readily,
in proportion as they are nearer the flowers. When the
leaves of the plant are compound, the bracts of the first
ramifications are sometimes so likewise, but most fre-
quently they are reduced to simple scales resembling
the rudiment of a petiole.

Bracts are often triple or trifid; and in this case the
two lateral ones, or the two lateral lobes of a bract
apparently single, are rudiments of stipules : thus, in
plants where the stipules are distinct from the petiole,
we frequently find, at the base either of the floral
branches or of the pedicels, three distinct bracts, the
two lateral being the smaller. In plants where the sti-
pules are adherent to the petiole, we frequently find
bracts with three lobes : sometimes the stipules retain,
in this state of bracts, a large development, and the true
leaf is either entirely or partly abortive ; the bract is
then replaced by two lateral and opposite ones, as is
seen in Cliffortia, &c. This phenomenon is analogous
to what takes place in Lathy rus Aphaca.

There are plants where the floral leaf, in being trans-
formed into a bract, instead of taking a membranous and

42 VEGETABLE ORGANOGRAPHY.

foliaceous appearance, assumes either that of a spiny
point, (as in Barleria, Exoacantha, &c.) or of a small
tendrill, (as in some species of Bauhinia,) or of a
tubercle or gland.

As long as the bracts are, by the disposition of the
flowers, so separated from one another as not to form a
ring or particular envelope, the name of bracts is
applied to them ; but they take another appearance when
the near approach of the pedicels or peduncles compels
the branches to arise in more or less regular verticils, as
is seen in umbels, corymbs, and. capitula ; the name of
Involucrum is then given to them collectively, and to
each individually the names of Scale, Leaflet, or Bract.

In umbels, where the common pedicule is not dilated
into a receptacle, the involucra are generally composed
of as many bracts as there are rays in the umbel; and
they are distributed in a single row.

In flowers in a compact head, the number of the
leaflets of the involucrum is rarely fixed ; they form
around the flowers an envelope of one or several rows,
which surround them so closely that it seems as if all
the flowers of a head formed but one, the involucrum
appearing to be a calyx.

The bracts which compose the involucra may be
either verticillate in a single row ( uniseriales ) , or in
two (biseriales), or in several ( pluriseriales ) \ When
they are in two rows, and the outside one is perceptibly
smaller, the involucrum is said to be Caliculate, or
furnished at its base with a kind of small calyx : when
they are in several rows, and the outer ones cover over
the base of the inner ones, gradually diminishing in
size, the involucrum is said to be Imbricate. A sin-
gular mode of imbrication is accidentally presented in
some Pinks : their flower, in the natural state, is fur-
nished at the base with two pair of leaves reduced to th^

OF INFLORESCENCE. 43

state of bracts ; but sometimes instead of two pair, we
find fifteen or twenty imbricated, so as to form a kind
of elongated spike, and in this case the flower is most
frequently abortive. This monstrosity is called in gar-
dens Dianthus Caryophyllus imbricatus.

The pieces which form the involucra, especially those
in a single row, are sometimes perfectly free ; this is
most frequently the case : sometimes they are united
by their margins, so as to resemble a single leaf; such
are those of several kinds of Bupleurum, Seseli Hlppo-
marathrum, Othonna, and Nyctago. These involucra
are usually very inaccurately called monophyllous, which
term ought to be replaced by that of GaiMOPHyllous,
which expresses their true nature.

When the involucra enclose several flowers, there can
be no doubt upon their nature ; but when they enclose
only one, it is often difficult to affirm if the envelope
be an external calyx or an involucrum : this doubt is
especially very great when the leaflets are united toge-
ther, as the sepals of a calyx. Though in the Marvel of
Peru the involucrum has been almost constantly taken
for a calyx, we are assured that it is an involucrum,
because in several plants of the same family this organ
contains several flowers, which is never the case with a
true calyx : the same illusion has also existed for a long
time in the Euphorbia, where the involucrum has been
called by the name of calyx, until it was known that
what was thought to be a single flower was an assem-
blage of several in a head. We know now in the same
manner that the spiny envelope of Chestnuts, the
cupule of the Acorn, or Hazel-nut, are involucra, and
not calyces. The question is more delicate in the
Malvaceae, which frequently bear, outside the calyx, a
row of verticillate leaflets: some call them the external
calyx, because they spring from the base of it. There

44 VEGETABLE ORGANOGRAPHY.

are some who have considered them as representing the
stipules of the calycinal leaves : some believe them to
be one-flowered involucra ; founding this opinion upon
the irregularity of their presence, number, position, and
shape, which appears to indicate that they form part
rather of the organs of inflorescence than of the flower,
properly so called. The question will be resolved in
the affirmative, if a Malvaceous plant should be found
bearing1 more than one flower within this external
covering.

The bracts which arise at the base of the partial
umbels form what is called the Partial Involucrum,
or the Involucellum : those which grow at the base of
the peduncles, or general umbels, take the name of the
General Involucrum. In flowers in heads, we fre-
quently find one or more within a primary envelope ;
this is the involucellum : for example, in Echinops the
involucellum is one-flowered, with several imbricated
leaflets ; and in Lagasca it is also one-flowered, but the
leaflets are combined. These involucella, in the ex-
amples which I have mentioned, are collected into a
compact head, which is itself surrounded by an invo-
lucrum, to which the name of General Involucrum, or the
involucrum properly so called, is given, and they are
themselves sometimes surrounded by others more ex-
ternal. There is in general but little exactness in the
manner in which these parts are designated and com-
pared together, and great errors in description fre-
quently result.

In a great number of capitate flowers, we find, besides
the scales of the involucrum, other bracts, situated be-
tween the flowers, and springing from the receptacle :
the leaflets of the involucrum are analogous to the bracts
which arise at the base of compound inflorescences ; the
scales of the receptacle represent the bracts belonging to

OF INFLORESCENCE. 45

each flower, or the bracteolcs ; and what, among other
circumstances, tends to prove the analogy of these organs
is, that these scales are always situated on the outer
side of each flower, which corresponds to the lower one
of racemes, and consequently their position is the same
as that of the bracteoles. When the gradations of form
from the leaflets of the involucrum to the scales of the
receptacles are carefully followed, no one can doubt the
identity of these organs. As they are situated between
very close flowers, it frequently happens that they are
abortive ; or reduced to a perfectly scaly state, and of
very small dimensions ; or, lastly, they are united either
with each other, or with the flower. When they are
united together by their margins, it results that each
flower seems as if it were inserted into a little cup ; this
is observed very well in Syncarpha : when they sur-
round the calyx, and are combined with it, they seem
to make part of the flower, as is seen in Scolymus angio-
spermus : but when the two phenomena take place at
once, the whole head of flowers does not make more
than a single body, apparently hollowed out into semini-
ferous holes, and its structure can only be unravelled
by very delicate analogy ; we see this in Gundelia and
Opercularia.

When the pieces of the involucrum are large and
sheathing at the base, the name of Spathe is given to
it, and the pieces of which it is composed are improperly
called Valves. This organization is only met with in
Monocotyledons ; and when it is mentioned in Dicotyle-
dons, it is as if one said that an involucrum had the
form or appearance of a spathe. In true spathes there
are sometimes one, sometimes two valves ; but in the
latter case they are never opposite, but alternate ;
and the lower one, which is the larger, embraces
by its base the upper one. This organization is met

46 VEGETABLE ORGANOGRAPHY.

with in all the compound inflorescences of Monocoty-
ledons. The little bracts situated at the base of the
pedicels which spring from the spathes, bear the name

of SPATHELL.E.

Among Spathes, we distinguish by the name of
Glumes those which are of a more scaly and dry
texture ; they are peculiar to the immense family of the
Gramineas. In this sense, the glumes which grow at
the base of the locustse of grasses are analogous to
spathes, or involucra ; those which are found around each
flower, and which are called Glumell^e, are, according
to some, analogous to involucella or spathellae, and ac*
cording to others, to the true integuments of the flower.
The opinion of the former is founded — 1st, Upon the
analogy with the Cyperacese, where the scale is evidently a
bract ; — 2d, Upon the fact that the outer glumella is
always situated a little below the inner one ; whence it
results that these valves are not verticillate, as the true
floral integuments, but alternate, as the leaves of the
Gramineas. These reasons appear to me to be strongly
in favour of this opinion.

Lestiboudois has been willing to corroborate this
theory by a third argument, viz. the quaternary num-
ber, which he admits without mentioning his reasons,
in the glumellae ; but it appears evident to me, with
Mr. Robt. Brown, that the glumes and glumellae pre-
sent the ternary number peculiar to Monocotyledons,
the outer one being a single piece, the inner one two
united.

We sometimes find in the Aroideae and Palms, very
large spathes composed of a single sheathing leaf; an
organization possible in Monocotyledons, where the
leaves are essentially alternate, but which cannot take
place in the involucra of Dicotyledons, the pieces of
which are essentially opposite or verticillate.

STRUCTURE OF FLOWERS. 47

Bracts approach more or less the sepals, or pieces of
the calyx, either when they are coloured or when they
are verticillate ; and the transition of the organs of ve°-e-
tation to those of the flower is found to be so gradual,
that the more it is studied the more we see this unity
of composition, which forms the base of philosophical
Organography. This observation will become more
clear from the examination of the structure of the
flower itself, which will be the object of the following
chapter.

CHAPTER II.

OF THE STRUCTURE OF THE FLOWERS OF PHANERO-
GAMOUS PLANTS.

Section I.
General Observations.

The flower, considered physiologically, is the appa-
ratus of organs which perform the sexual fecundation,
and of those which serve as their immediate envelopes.
Considered with regard to Organography, we shall see
that it is an assemblage of several (usually four) verticils
of leaves, variously transformed, and situated, in the
form of a bud, at the extremity of a branch called the
pedicel.

The organs which perform the fecundation, are, — the
female organs, or Pistils, which enclose the ovules ; the
male organs, or Stamens, which fecundate the others.

48 VEGETABLE ORGANOGRAPHY.

The immediate envelopes are the Corolla, which is of
an analogous nature to the genital organs ; and the
Calyx, which serves as the external covering, and is of
a foliaceous nature. To these four organs must be added,
for the sake of clearness, the Torus, which serves as
the common base of the corolla and stamens ; and the
Axis, which is the prolongation of the pedicel. These
six parts spring from the top of the pedicel, and consti-
tute all the essential organs of the flower. All that is
found outside the calyx, belongs to the bracts or invo-
lucra, which I have already mentioned ; and that which
is met with in the interior, and does not form part of
these six organs, is reduced to some nectariferous glands
which do not appear essential to the flowering.

We shall first describe each of these parts in its most
simple and least complicated state, joining only to their
description the cohesions* which they contract. We
shall examine the modifications which they present, in
their adhesions, abortions, or affinities with neighbour-
ing organs ; and we shall conclude with some observa-
tions upon the whole structure of flowers.

Section II.

Of the Calyx or Sepals.

The Calyx is the external envelope, usually folia-
ceous, which is observed in almost all complete dico-
tyledonous flowers, and which forms the sole envelope

* From analogy with medical language, I call Cohesions the unions
of organs of the same nature, and Adhesions those of different organs :
thus the sepals united together are Coherent ; united to the ovarium,
Adherent.

STRUCTURE OF FLOWERS. 41)

in most of those which are incomplete. It is formed
of pieces which are disposed in verticils in one or two
rows, and which bear the name of Sepals, (sepala.)

The sepals are evidently organs very analogous to the
nature of leaves: and it might be said with some reason,
that they are bracts which constantly exist, and form
an integrant part of the flower. The identity of nature
of the sepals, bracts and leaves, is derived from the fol-
lowing facts : — 1st. Their internal anatomy presents
vessels and tracheae as in leaves, and their tissue presents
most frequently a great analogy in the distribution of
the fibres. 2d. Their surface, as that of leaves, presents
stomata, most frequently distributed in the same manner
in the same plants. 3d. When they have glands or
hairs, these organs are similar in their nature, form, or
position, to those of leaves. 4th. They are almost
always green, as leaves, and endowed, as they are, with
the faculty, both of becoming etiolated in darkness, of
decomposing carbonic acid gas, and of exhaling oxygen
when placed in water under the sun. 5th. Lastly,
they take, under several accidental circumstances, an
extraordinary development, and then they absolutely
resemble true leaves, as is frequently seen in Roses,
for example, (PI. 17.) We may regard the sepals,
then, rs bein? of a foliaceous nature: and it may
be said that they are kinds of floral leaves, which by
their position take particular forms, and serve as the
outer covering of the flower. They are, as leaves, some-
times articulated at their base, and then they become
detached and fall off spontaneously, either at the com-
mencement of the flowering, as in the Poppy, or at the
conclusion of it, as in Ranunculus ; sometimes they are
continuous, or adherent at their base, and then they do
not drop off, and are said to be Persistent. But then
they either dry up after flowering, and are termed Mar-

vol. II. E

50 VEGETABLE ORGANOGRAPHY.

cescent, as in the Broom; or they become fleshy, as in
certain Ficoids; or they increase in size, remaining folia-
ceous, and are said to be Accrescent, as in the Alke-
kengi. Some calyces present a singular mode of fall;
viz., their upper parts remain close or united together
after flowering, and the tube is cut transversely by a
rupture, either near the base or at the origin of the
lobes : it is thus that the cap-like calyx of Eucalyptus
is formed, and in an analogous manner, that of Scutel-
laria galericulata is cut transversely near the base, when
the seeds are ripe.

When the sepals are articulated at their base, they
can never be united together, and are therefore con-
stantly distinct. When, on the contrary, they are con-
tinuous with the stem, they are presented under two
states ; sometimes free with regard to each other, some-
times united together at their margins by a kind <of
natural union, taking -place before they are visible exter-
nally. In this case, we easily recognise, in the greater
number of species, the existence of partial sepals, either
by the disposition of the nerves, or because the union is
hardly ever so complete as for one part not to remain
free near the top, forming the lobes of calyces with
united sepals.

Calyces with free sepals are said to be Polysepa-
locs; and when we wish to express at the same time
their number as well as freedom, we say Bi-, Tri-,
Tetra- &c. sepalous.

Those with united sepals are called Gamosepalous,
or improperly Monosepalous, because, on account of
their union, they seem to form only a single body. If
they are only united at their base, their free parts are
designated by the name of Partitions, and the calyx
itself is said to be Bi-, Tri-, Quadri-partite : if the
union proceeds as far as the middle, they are named

STRUCTURE OF FLOWERS. 51

Divisions, and the calyx is said to be Bi-, Tri-, Quadri-
fid : if the union proceeds nearly to the apex, the free
parts are called Teeth, and the calyx is said to be Two-,
Three-, Four- &c. toothed : if the union reaches
the apex, the calyx is said to be Entire: if the sepals
are unequally united together, there may result, among
other combinations, the following : — Two or three may
be united on one side ; and there remains on the other,
either a single sepal not united so far, or two or three
united together beyond the point which connects them
with the preceding : the name of Lips is given to these
kinds of parcels, and the calyx is said to be Two-lipped
when there are, as is most frequently the case, two se-
pals united into an upper lip, and three in a lower one :
we say of a calyx that it is One-lipped, when all the
sepals are united together and thrown on one side, be-
cause the union on the other only takes place in a very
incomplete manner. Lastly, there are some rare cases
where the sepals are so united by their apices, that they
cannot be separated, and the calyx can only open by a
rupture at its base ; it is this which takes place in Eu-
calyptus, where it is broken transversely near the base
of the limb, and is detached in the form of a hood. For
a long time the calyx was considered to be a single
organ, which was cut or divided, without any reason
being given for these pretended divisions ; its structure
was then almost unintelligible, and unfortunately it was
at this period that the terms were established by which
its modifications are designated. I do not propose to
change them, for the fear of causing too great a confu-
sion ; but it is necessary to know, once for all, that all
that has been said of the divisions of calyces must be
understood to relate to sepals differently united together.
The use of the calyx is evidently to serve for the
covering and protection of the other floral organs during

e2

52 VEGETABLE ORGANOGRAPHY.

flowering, and sometimes to the young fruit, where it is
persistent. It is probable, seeing the foliaceous nature
of this organ, that it serves also to elaborate the juices
destined either for the flower or the young fruit.

Section III.
Of the Corolla or Petals.

The Corolla is the more or less coloured inner en-
velope, which is observed in the greatest number of the
flowers of Dicotyledonous plants, but which is entirely
absent in several others. It is formed of pieces disposed
in one or several verticillate rows, and which bear the
name of Petals (petalu).

The petals differ more in nature from leaves than the
calyx does, and very much resemble, as we shall see, the
stamens : they usually have neither stomata nor tracheae ;
they are of all kinds of colour, except green ; and even
when they are greenish, they do not appear to owe it to a
cause analogous to that of leaves, for they are not capable
of being etiolated. They do not disengage oxygen when
exposed to the sun, but, on the contrary, they have a ten-
dency to diminish the free oxygen of the air by forming
carbonic acid : they frequently exhale very different
odours from those of leaves; and these odours, when they
are concentrated, act in a peculiar manner upon the nerv-
ous system of the human species. Lastly, when they bear
glands or hairs, these organs are very different from those
of foliaceous parts. All these different characters of the
petals are found in the stamens and torus, and hereafter I
shall endeavour to prove the identity of the nature of
these organs.

STRUCTURE OF FLOWERS. 53

The petals are almost always articulated with the
torus, and consequently Deciduous : frequently they
fall off very early, that is to say, before fecundation ; they
are then said to be Caducous : sometimes they are con-
tinuous or adherent, and then they are termed Persist-
ent— for example, in Campanula.

When they are completely distinct from each other
down to the base, the corolla is said to be Polypeta-
lous, or, if it is wished to express the number of petals,
Di-, Tri-, Tetra-petalous. When they are more or
less united together, it is most frequently said, but in an
improper manner, that the corolla is Monopetalous, a
term for which I substitute that of Gamopetalous,
which signifies that the petals are united. This union
is seen to be the case, by the distribution of the vessels
in the greatest number of cases ; it is perceived
also more clearly in some plants, such as JRhodora
Canadensis, Campanula medium, or Phlox amccna,
(PL 18, figs. 8, 9, 10, 11) in which we frequently
find every degree of union which can possibly be met
with, whether in the flowers of the same individual, or
in the petals of the same flower. It happens in the
Compositor, that the five petals united into a tube, pre-
sent in certain flowers a lateral fissure, so deep that the
corolla, instead of appearing tubular, becomes strap-
shaped. This phenomenon is constant in the Chicoraceas,
where some ingenious philosophers attribute it to the
presence of glands which unite the lobes together at the
apex : in the other Compositae it only takes place in the
marginal flowers of the capitula, but here also we recog-
nise the original nature of ligulate flowers, in their
sometimes happening to retain their tubular form.
Thus, I have observed in gardens an accidental monstro-
sity of Tagetes erecta, where the semi-florets were
changed into tubular ones, larger than those of the disc.

54 VEGETABLE ORGANOGRAPHY.

The degree of coherence of the petals, is expressed by
the same terms as those which refer to the calyx ; thus
we say of a corolla, that it is Partite when the petals
are united at their base ; Divided, when united as far
as the middle ; Dentated, when the union proceeds
very nearly to the apex; Entire, when the union is
complete.

The petals, more frequently than the sepals, present
natural cohesions in their upper parts, remaining free at
the base ; this is seen in the two petals united at the
summit, which form the keel of the Papilionacese, or
still more in the union of the five petals in the common
Vine, which takes place at their summits whilst their
bases are distinct. They are also sometimes united by
their base and apex, the middle parts remaining free, as
in some kinds of Phyteuma.

The petals arise, in the greatest number of cases, in a
single row, equal in number to the sepals, and situated
between each of them : when they are in two rows, the
external one is between the sepals, and the inner one,
which alternates with the first, has its petals opposite
the sepals : when the number of rows is more than two,
the third has its petals opposite those of the first, the
fourth opposite those of the second, &c. Among the
exceptions to these laws, we must mention the few cases
where the petals are opposite the sepals, as in the Ber-
beridese.

Each petal, considered with regard to its structure,
sometimes presents, as leaves, a kind of support formed
by the union into a narrow process of all the bundles
which afterwards spread out to form the limb of the
petal ; this kind of petiole bears the name of the Claw,
(unguis,) and the dilated portion that of the Lamina or
Limb. There is a less number of unguiculate petals
than of petiolatc leaves. The partial bundles of vessels

STRUCTURE OF FLOWERS. 55

which spring from the top of the claw, and spread out
into the limb, are in general less distinct, thick, and
regular, than the nerves of leaves ; the same terms of
Penninerved, Palminerved, &c. may be applied to
them, but there is seldom any necessity for describing
them. The corollas of the Composite present a dispo-
sition of the nerves which is peculiar to them ; viz. each
petal, instead of having as usual a middle nerve, is fur-
nished with a very perceptible one upon its two margins,
as far as the apex ; whence it results, that in the tube of
the corolla, where the petals are united, we see five large
nerves, each of which proceeds from one of the sinuses,
and is formed by the union of the two marginal ones.

When the petals are free and furnished with claws, as
in the Pink, these claws are usually straight and near
together, thus forming a kind of tube with five slits. If,
on the contrary, they are united, as in the Tobacco,
their union forms a true entire Tube ; and the limbs of
these petals united by their claws, may be either entirely
free, or united half-way, or as far as the apex.

The entrance of the tube bears the name of the
Throat (faux) : we may here frequently remark little
petaloid appendages, which have received the name of
Scales or Appendages when they are free, and which
are called the Crown when they are united together, or
when it is wished to designate them collectively.

The use of the corolla is evidently to serve for the
protection of the sexual organs, especially before their
complete developement. Its fugacity, and its nature, so
distant from a foliaceous one, do not permit us to assign
it any use with regard to the nutrition.

5G VEGETABLE ORGANOGRAPHY.

Section IV.
Of the Stamens.

The Stamens, (stamina,) or the male organs of
plants, arise from the torus, and are placed in one or
several rows or verticils between the petals and pistil.
Rceper has proposed to give them collectively the name
of Andrceceum, in order to have a collective term ana-
logous to those of corolla as concerns the petals, and
calyx relative to the sepals, and pistil to the carpels.

The number of stamens is very variable in the
different genera — from two to a hundred: when but one
is found, it results only from the abortion of the other
or others which form with it the regular verticil.

When they are disposed upon only one row, their
number is usually equal to that of the petals or sepals;
and they arise either opposite the sepals and between the
petals, or, what is more seldom, between the former and
opposite the latter. We sometimes find several stamens
which arise in one row before each sepal; then the total
number is the product of the number of sepals, multiplied
by that of the stamens placed before each of them.

When they are disposed in two circular rows, there is
almost always one opposite each petal, and one before
each sepal ; and the total number is double that of each
of these organs. Sometimes, when the petals are absent,
the stamens are found situated alternately opposite and
between each sepal : if there be more than two rows,
the third is situated opposite the first, the fourth oppo-
site the second, so that the whole number of a regular
flower is the product of the number of rows multiplied
by that of one of the rows, which is usually equal in
number to that of the petals or sepals.

STRUCTURE OF FLOWERS. 57

In this sense there is never an indefinite number of
stamens, to speak truly ; but the greater the number is,
the more chances are there of accidental abortions or
multiplications, and the more it is neglected to count
them exactly : it is usually said that the number is
indefinite when it exceeds twenty.

Each stamen is composed of a filament and an anther.

The Filament (filamentum) is a body which springs
from the torus ; and its form is sometimes cylindrical, or
subulate and very long; sometimes compressed; more
rarely expanded towards the apex into a kind of scale
or cap, as in Borago laxijlora. In texture it is very
analogous to petals, especially to their claws. It is
coloured as they are ; bears the same kinds of hairs and
glands; is devoid of stomata and tracheae, &c. Their
length seems only to be determined by the necessity of
supporting the anther at a sufficient height for it to be
placed in a favourable position with regard to the stigma.
Thus, in the greatest number of cases, the filament is of
the necessary length for the anther to be nearly of the
height of the stigma, or a little above it ; but frequently
in drooping flowers, the filament is shorter, and the
anther is found above the stigma. This law is often
modified by the torsion or peculiar movements of certain
flowers. The filament, as it only serves for the support
of the anther, may be, then, either very long or very
short ; in this last case, it is sometimes so short that it
is said to be completely wanting. When it is articulated
at its base with the torus, the stamen detaches itself and
falls off after fecundation ; when it is not articulated, as
in Campanula, it perishes and dries up without falling.

The Anther (anthera) is a kind of purse borne by
the filament, containing a powder which is called the
Pollen. As the pollen itself contains the fecundating
matter, and is consequently the essential part of the

58 VEGETABLE ORGANOGRAPHY.

organ, the anther which protects and nourishes it is also
a very important one. It is situated upon the top of the
filament in three different ways : — 1st, It is frequently
attached by the middle of the back to the very tapering
extremity of the filament; and then, as it is placed in a
vertical position before the flowering, and afterwards
takes an horizontal one, it is said to be Versatile. 2d,
In several cases it is attached by its base to the apex of
the filament of which it appears to be the continuation,
it is then Erect. 3d, In some families it is adherent to
the filament by its whole dorsal face, so as to have no
movement; it is then said to be Adnate or Adherent
to the filament. In this last case, it frequently happens
that the top of the filament is prolonged beyond the
anther into an appendage, either strap-shaped or in a
filiform point, as in the Oleander, or as a gland, as in
Adenanthera. Sometimes it is the connectivum which is
prolonged in the same manner, sometimes the cells
themselves ; so that the terminal appendages result from
verv different anatomical causes.

Anthers are generally composed of two membranous
sacs, applied to one another and joined by a body which
is called the Connectivum. This body is sometimes so
small, and scarcely apparent, that it is neglected in
descriptions ; sometimes so large and well developed,
that the two cells of the anther are separated from each
other, as is seen in the Sages.

We find a certain number of anthers which have but
one cell : sometimes this conformation appears natural
to the plant, and then it is only met with in those anthers
which are attached by their base to the top of the
filament ; sometimes it results from an accidental abortion
of one of the cells, which especially takes place when the
connectivum is very large and the cells very distant, as
in the Sage; sometimes it is owing to the filament

STRUCTURE OF FLOWERS. 59

being, as it were, split into two, or divided, so' that in the
place of one stamen with a bilocular anther are found
two with unilocular ones : this is observed in one of the
stamens of Impatiens noli-tangere.

I do not know that any anthers exist which have
naturally more than two cells, but this appearance
results in several cases from two causes : — 1st, Each cell
is most frequently divided into two small ones by a
longitudinal fold of the dorsal part, and this frequently
gives the anther the appearance of being four-celled :
2dly, It sometimes happens that two or more neigh-
bouring anthers are united together, so that the ap-
parently simple body which results, appears to be four-,
six-, or many-celled ; for example, in the Salix impro-
perly called Monandra, and probably in the Yew. We
shall revert to this subject on speaking of the Cohesion
of the Stamens.

The cells open at maturity in four different ways : —

1st. The most frequent case is that where the opening
takes place by a longitudinal fissure through the middle
of each cell ; and when the anthers are two-celled, we
say that they are Birimose.

2d. It happens more rarely that the opening takes
place bv transverse fissures, as in the Lavender.

3d. Several anthers open at their apex by two pores
(Biporose) situated at the top of the two cells — in
Solarium, for example ; or by one pore at the top of a
unilocular one, as in Amaranthus. This kind of dehiscence
may result from the longitudinal fissure remaining united
except at the apex.

4th. The most singular case is that where the cells
open by valves which detach themselves from the base
upwards, as is seen in the Barberry and Laurel, and
in the two families to which these plants belong — the
Berberidac and Laurineae.

60 VEGETABLE ORGANOGRAPHY.

The position of the anthers with regard to the pistil
ought also to be observed. The most frequent and natural
ease is that where the anther, whatever be its mode of
attachment, has its back towards the outer, and its cells
on the inner side of the flower : it is this that is meant
whenever it is not expressed to the contrary in descrip-
tions; when it is necessary to express it, we say that
it is Introrse (introrsa, antica). In some plants it is
placed in the contrary way, so that its back faces the
pistil, and the cells open towards the calyx ; this kind of
anther is called Extrorse {extrorsa, postica) : we observe
it very clearly in certain adnate anthers, such as those of
the Magnoliaceas, or in those with flat filaments, as in
Iris ; but when the filaments are slender, or when the
anthers are erect or versatile, the observation of their
true direction presents some difficulties, because it often
happens that the filament is so twisted as to place the
anther in a situation different from its natural one ;
thus we may often mistake their, true position in the
Ranunculaceae.

The form of each of the cells is sometimes round,
and then we say of the anther that it is Didymous :
more frequently oval; sometimes long or linear; this
last case especially takes place in adnate anthers, as
those of the Magnoliaceas : when these cells are erect, they
present two convex bands, separated by a rectilinear
furrow: sometimes they are at the same time adnate,
linear, and more or less curved, and then they present
inequalities and curves of a remarkable appearance ; we
see this in the Gourd, in Durio, Eriodendron, &c. ; these
anthers are called Anfractuose. Those with round
cells frequently open by a transverse fissure, those of a
linear form by a longitudinal one, and oval anthers
present all kinds of dehiscence.

The colour of the anther is often yellow; sometimes

STRUCTURE OF FLOWERS. 61

orange, brick-coloured, violet, purple, or white; never
green or a true blue. But we must take care not to con-
found the colour of the anther with that of the pollen ;
and it must be remarked that the colour of the former
differs before, during, and after fecundation.

The Pollen is a mass of granules contained in the
anther, and coming out of it at its dehiscence. According
to Guillemin, who has recently studied this important
organ with care, these grains, considered before their
exit, appear disposed in a regular series, following the
direction of the walls, and floating in a viscid fluid : at
whatever age they have been examined, they have
hitherto been found completely free ; and, if it be really
so, we may suppose that their nutrition takes place by
the simple absorption of the surrounding fluid by their
walls ; but it may be believed, with some likelihood,
that in their first stage they adhere to the walls of the
anther by a filament, which escapes our sight on account
of its fugacity or shortness, but which is sufficient to
keep them in continuity with the rest of the tissue ;
Turpin goes so far as to designate the part projecting-
into the interior of each cell as being that which bears
the pollen, and proposes to call it by the name of
Tropho-pollen. The adherence of the grains of pollen
with the anther, in their earliest state, seems to me
probable — 1st, on account of the general analogy of all
the organs of plants ; — 2dly, on account of the special
analogy of the grains of pollen with the ovules; an
analogy which is so great, that, as we shall afterwards
see, one half of the anther is sometimes found bearing
pollen, and the other half ovules.

The grains of pollen at their maturity are sometimes
completely free, which is most frequently the case ;
sometimes kept in a kind of network formed of fine
filaments, as in the Onagrarise : and sometimes united

62 VEGETABLE ORGANOGRAPHY.

in compact masses, as in the Orchideas and Asclepi-
adeas.

The surface of the grains presents an important
difference : sometimes it is perfectly smooth and not
viscid, as in Justicia quadrifida and a multitude of other
plants : sometimes it is invested with a viscid layer
differently coloured, and which appears to be produced
by true secreting organs ; we see this in most of the
Cinarocephalae, Heliantheae, &c. : the surface of some
bears mamilliform eminences, as in Cobeea : in the
Cucurbitaceas it is furnished with pointed projections,
which Amici considered as kinds of lids terminating in
a point.

The general form of the grains of pollen is most
frequently globular, oval, or elliptical in Dicotyledons,
and lengthened elliptical in Monocotyledons. It is
frequently remarked, that the oval or elliptical ones
are marked on one side with a longitudinal furrow,
resembling that in a grain of wheat. They are fre-
quently found polyhedral, as in the Chicoraceas ; some
are nearly triangular, as those of the Proteaceaa and
Onagrariae ; those of Colutea are oval and truncated at
the extremities. Among elliptical or elongated pollens
there are sometimes found curved ones : Grew has even
mentioned ramified ones, but this observation does not
appear to have been verified. In general, plants of the
same family have their pollen nearly alike ; but analogous
forms are found in very different families.

Each grain of pollen contains a fluid which appears
to be slightly -viscid, and has been named the Fovilla ;
it is the true fecundating liquor of plants, and is full of
little grains, endowed with a peculiar motion, and which
are called Granules.

The pollen grains open at maturity either by a
regular dehiscence, or, most usually, by an irregular

STRUCTURE OF FLOWERS. G3

rupture : this opening is particularly caused by the
application of water, which is sometimes absorbed by
them ; and the diluted fovilla appears to come out in an
insensible manner. It is generally thought that the
grains, falling on the stigma, which is almost always
viscid and humid at the period of fecundation, then
open and deposit the fovilla; this appears to be absorbed
by the stigma, and, reaching the ovules, excites the
young embryo and conveys to it the vital movement.
Others think that the granules penetrate into the ovules
and there form the embryo. But in a subject so delicate,
it is difficult to form a decided opinion : I shall only
observe, that the constant presence of animalcule in the
fecundating liquor of animals, and of granules in that of
plants, is not sufficient to destroy the opinion of the
pre-existence of germs in the ovules ; and we might
believe, with some likelihood, that the animalcule and
granules are the exciting agents of a germ pre-existing
in the ovule. This question, more physiological than
anatomical, would be out of place here ; and I hasten
now to resume the description of the male organs of
plants.

The stamens may be coherent or united together in
three different manners ; viz. by the filaments, by the
anthers, or by both organs at the same time.

When the cohesion takes place by the filaments, it is
usual to say that the filaments are Monodelpiious
when all are united together; Diadelpiious, when they
form two parcels; Polyadelphous, when there are
several ; but these terms fall far short of giving a com-
plete idea of the variety of the real cases.

The cohesion of the filaments together is so much the
more easy, as they are naturally of a more spreading
form, or nearer one another in their primitive position :
it may take place in every degree, as in the petals ;

64 VEGETABLE ORGANOGRAPHY.

thus, there are some stamens whose filaments only
cohere at the base — for example, several Pinks ; others
where they do so as far as the middle, as the two
stamens of Salix incana ; others, where the union
proceeds very nearly to the top, as in most Meliaceae,
some Malvaceae, &c. We also find some free at their
bases at the time of flowering, and united at the top, as
in Lobelia. In this respect we find as many differences
among stamens cohering together, as in petals united
into a gamopetalous corolla.

The cohesion of the filaments may take place in a
uniform manner among all the stamens, whether they
be in a single row, as in most Amaranthaceas and
Meliaceae, or in several united by their bases, as in
Malvaceae.

Sometimes the stamens are distributed regularly in
parcels united at the base, and equal in number to the
petals or carpels ; thus, in Hypericum and Melaleuca
there are five parcels alternate with the petals, and each
composed of a nearly definite number of stamens. Some-
times, the stamens being almost in a single row, and
double the number of the petals, as in the Papilionaceae,
their filaments are all united, either all together, in a
cylindrical sheath, as in Cytisus — or nine together, in a
split sheath, on the upper side, and the tenth remaining
free, as in Colutea — or five on one side and five on the
other, forming two semi-sheaths, as in JEschinomene —
or in two lateral parcels, and a single one free, as in
several kinds of Dalbergia. Of other cohesions, the
most singular are observed in some other families : thus,
in several Cruciferae, such as CEthionema and Sterigma,
of the six stamens, the two lateral are always free, and
the four others united by the filaments, either partially
or wholly. It must be remarked, that in the genera
which present this disposition, when these filaments

STRUCTURE OF FLOWERS. G5

remain free they have a tooth projecting from the side
where they have a tendency to unite with the neigh-
bouring one. In the Gourd, of the five filaments, one
is most frequently free, at least at the base, and the
others intimately united in pairs. In the Fumaria and
the other genera of the same family, we find two parcels,
each bearing three anthers, of which the middle one has
two cells, and the two lateral one cell ; whence it might
be presumed that the real number of filaments is four,
united two and two.

When the stamens are united by their anthers, they
bear the name of Synanthefous, or Syngenesious.
This phenomenon, although less varied than the pre-
ceding, presents also some differences ; in general it
takes place in all the anthers of the flower at the same
time, and their cohesion then forms a kind of ring
through which the style passes. In this case, the
anthers thus united are introse, opening by longitudinal
fissures; and the stigma, elongating in the interior of
the ring, receives the pollen of the anthers. This is
observed in the immense family of the Compositae and
Lobeliaceae. Sometimes the two cells of the anther,
being separated by the bifurcation of the filament, or an
elongation of the connectivum, each unites with the cell
of the neighbouring stamen, and thus form bilocular
parcels which might be taken for true anthers : this
singular conformation is presented in Stapelia, and per-
haps in the bilocular anther of the Fumariaceae.

Finally, the third combination is that of the stamens
cohering both by their filaments and anthers. As yet
but a very small number of examples is known. Bar-
nadesia, a genus of the Compositae, has the filaments
united into a complete tube, and the anthers form a ring
with the cells opening internally : the Symphonieae (a
tribe or family between the Guttiferae and Meliaceae)

VOL. II. F

66 VEGETABLE ORGANOGRAPHY.

have the filaments united into a tube, and the anthers
open by longitudinal fissures on the outside of it, a
phenomenon slightly resembling the structure of the
Cucurbitaceae. Salix monandra presents two stamens
united by the filaments and anthers. Morina persica
has four fertile stamens, joined two and two, in the same
way. Lastly, the Yew appears to have eight to ten
stamens united by the filaments and anthers, forming a
bundle which expands at the top, and bears the anthers
on its inner side ; this form of the filament some
authors have named Androphore.

The stamens, as I have said above, always arise from
the torus very near from where the petals spring ; they
easily contract adhesions with these organs ; but these
adhesions are very frequent when the petals themselves
adhere together, and very rare when they are free.
Thus, in all gamopetalous corollas, the filaments are
united to them, except in the Campanulaceae, and in
polypetalous ones the stamens do not adhere to the
petals except in the Malvaceae, Caryophylleae, &c. and
then but slightly.

As the function of the stamens is connected with their
being the male organs, it will be explained in detail in
the Physiology ; it is sufficient here to state that this
opinion, almost unanimously admitted, is founded, — 1st,
upon the examination of their structure ; 2d, upon the
fact that when flowers are deprived of them either
naturally, as in dioecious plants, or artificially, as in
those mutilated by insects, &c, they are constantly
sterile ; 3d, upon the mules or hybrids which are
produced when the pollen of one plant is applied to the
stigma of another.

STRUCTURE OF FLOWERS. 07

Section V.
Of the Pistil or Carpels.

The pistil, taken collectively, is evidently the female
organ of the flower, since we see it after flowering
changed into fruit and containing the seeds. It has for
a long time been considered as a single organ ; but its
structure, and especially that of the fruit which succeeds
it, only becomes intelligible when it is considered in the
same manner as all the other organs of the flower, that is
to say, as composed of elementary organs, sometimes free,
at others cohering together, and which I call Carpels.

The Carpels spring from the centre of the flower, and
are arranged in different manners, of which the following
are the principal, viz. : —

1st. They are verticillate around a real axis, which is
the prolongation of the pedicel, and are adherent to it
by their inner angle ; this takes place in the Malvaceae,
where we see five or more carpels around a column which
arises from the pedicel. This column is expanded (in
Stegia, for example) into a kind of terminal disc, and
the carpels adhere to it by their inner angles. We
observe an analogous organization in the Euphorbiaceae;
it is also met with in certain foreign genera, such as
Gyrostemon, which seem intermediate between these two
families, otherwise so different.

2d. We also find the carpels verticillate at the top of
a central column, but pendulous from it, and con-
sequently only adherent by the summit of their inner
angle. We see this in the Geraniaceoe : the carpels do
not adhere to the column by their edges, but hang from
its apex by a long pedicel.

3d. The carpels may also be verticillate at the apex of
the axis, but erect and adherent by the base of their inner

f 2

08 VEGETABLE ORGANOGRAPHY.

angle ; this axis is sometimes so short that it is considered
absent, as in all the Crassulaceae, Aconitum, Aquilegia,
Illicium, &c. The place of the column is then vacant
in the centre of the verticil formed by the carpels.
Sometimes the axis is slightly prolonged, and the verticil
of carpels is, as it were, elevated, as takes place in
several Rutaceae.

4th. The carpels may be disposed in a spike around
the central column, as is seen very clearly in the Tulip-
Tree, Magnolia, several of the Ranunculaceae, Myosurus,
Sec. These carpels sometimes present at their base little
scales, which may be considered as true carpellary
bracts : I have observed them in some Ranunculaceae.
The nature of these organs deserves the attention of
Naturalists.

5th, If the column be very short or round instead of
being elongated, the carpels, instead of being in a spike
as in the preceding case, may be agglomerated into a
head more or less compact around this column, as is
seen in the Bramble, the Strawberry (where the column
is fleshy), the Anona, most kinds of Ranunculus,
Alisma, &c.

6th. The carpels may be dispersed upon the walls of
the torus adherent to the calyx, as is seen in Rosa, the
only example perhaps of this conformation in the whole
vegetable kingdom.

All the preceding dispositions suppose a plurality of
carpels, and this is in fact, in my opinion, the natural
and normal state of flowers ; but by abortions and cohe-
sions all may be reduced to a single one, either in reality
or in appearance.

Each carpel may be considered as a little leaf folded
upon itself, and containing the germs which are developed
after fecundation. These germs bear the name of
Ovules, and the carpels which contain them the Ovary.

STRUCTURE OF FLOWERS. 69

The carpels are usually sessile, sometimes furnished at
the base with a small support independently of the
central column, and which represents the petiole of the
leaf; this little support receives the name of Theca-
phore. It is visible in several species of Sterculia, in a
great number of Leguminosae, Capparideae, (especially
in Cleome longipes, where it is nearly a foot long,) &c.

The ovules are almost always attached to the margin
of the little leaf which, by folding, forms the ovary, or,
which is to say the same thing, on both sides of the inner
angle of the carpel; the portion, usually slightly thick-
ened, where they adhere, bears the name of Placenta ;
the apex of which and of the carpel is prolonged into a
filament, sometimes very long, at others very short,
called the Style, and this bears a glandulous organ,
glutinous at the period of fecundation, which receives
the pollen, makes it burst, and then imbibes the fovilla :
it is a kind of pistillary spongiole, and is called the
Stigma. Let us rapidly review these different points,
which we shall be obliged to revert to more in detail on
speaking of the fruit.

The analogy of the carpel to the leaves is deduced
from the following facts: — 1st. It is frequently of the
same texture and colour, and has the same faculty of
decomposing carbonic acid gas when exposed to the
light. 2d. It frequently bears stomata, and when it has
hairs or glands, these organs are often analogous to those
of leaves. 3d. It very frequently presents nerves very
analogous in their distribution to those of leaves. 4th.
The ovules are situated in most carpels in the same
places which correspond to the germs, which in some
leaves, as in Bryophyllum, are developed without fecun-
dation. 5th. We not unfrequently see the carpels, by
degeneration, develop into true leaves, as I have ob-
served it in Lathyrus latifolius. We also very easily

70 VEGETABLE ORGANOGRAPHY.

see this analogy in certain monstrosities of Cherries,
which, instead of having but one carpel, bear several,
sometimes in the state of ordinary ones, at others in the
state of leaves folded upon themselves.

The ovary being formed by the folding of a leaf, pre-
sents in different cases forms connected with its origin.
Thus when it is free from all pressure or cohesion with
the neighbouring ones, it is either compressed and flat-
tened, when the two halves of the leaves are even and
applied to one another, as in the Pea ; or swollen out, but
with a dorsal nerve, when the leaf has a middle nerve
and its sides are curved upon one another, as in the
French-Bean and Bladder-Senna ; they are nearly in the
form of a horn when the leaf has no middle nerve, as in
Colchicum. It sometimes happens that the margins of
the leaf are folded back upon themselves in the interior,
and form two-celled carpels, as in Astragalus. When
the carpels are verticillate and close to one another, they
then take, in consequence of their pressure, a triangular
form, the two lateral parts being fiat and inclined in-
wards, and the dorsal surface being flat or convex, or
even angular ; this is seen in the Crassulaceae. This is
still more evident when they are united together by their
lateral faces.

The style arises from the carpel originally near the
apex, but sometimes from the middle or base of the inner
margin, as is seen in Alchemilla ; the point whence it
proceeds is always that where the placenta terminates ;
its length is determined by the proportion which ought
to exist between the position of the stigma and that of
the anthers ; when it is absent the stigma is sessile upon
the summit of the ovary. The form of the style is
usually slender, cylindrical, and simple. But as the
ovules are generally disposed upon two rows, or placentae,
each of them has its stylary prolongation, and that of

STRUCTURE OF FLOWERS. 71

each carpel may be considered as formed of two partial
styles, sometimes perfectly free, at others more or less
united together ; and then we say that it is bifid, and
that it has two stigmata. The Euphorbiacese show very
well these different states of the carpellary styles, both
simple and forked.

When the styles are detached or raised above the
carpels, they are most frequently free ; at times united
with the central column, as in Geranium.

The stigma is, as we have said, a kind of spongiole
supported by the pistil. It is usually situated at the
extremity of the carpellary styles, and we say that there
is but one when they are united the whole way, and that
there are two when they are only united part of their
length. This manner of expression has often caused the
branches of the style to be confounded with the stigmata,
which in reality are only the glandular part, whatever
place it occupies. Thus, for example, in several Legu-
minosae this portion is naturally near the extremity ; in
the Iris the branches of the style are flat, petaloid, and
two-lipped, the upper very long and often bifid, the
lower very short; it is in the transverse fissure which
results from the position of these two lips that the glan-
dular part or true stigma is found.

The stigma, whatever be its position and form, is be-
set with viscid papilla?. The pollen, when it falls upon
it, experiences the action of this humidity — it bursts ;
the fovilla is absorbed by the spongioles, and on applying
coloured fluids to them, as Bulliard did, we see that they
follow the vessels in the interior of the style, penetrate
into the placenta, and thus reach the ovules. It is by
this means that vegetable fecundation is performed.
The collection of vessels which go from the stigmata to
the seeds bears the name of the Pistillary Cord ; we
shall revert to it on speaking of the fruit.

72 VEGETABLE ORGANOGRAPHY.

The style bears, moreover, in some plants, hairs which
have been called Collecting Hairs (pili collectores);
they are found in the Compositae, and they serve to ex-
cite the anthers, cause their dehiscence, and attract the
pollen to the stigmata. The Campanulaceae also present
collective hairs, which in position and structure appear
very similar to those of the Compositae ; but it is not
impossible that their function is slightly different. In
fact, the part of the pistil of the Campanulas which we
call from analogy by the name of stigma, appears com-
pletely inaccessible to the pollen at the period of flower-
ing, and Cassini supposes that the hairs perform perhaps
the function of stigmata ; this subject deserves to be
studied with care.

The carpels have a greater tendency to unite together
than the more external organs, which is connected,
doubtless, with their nearer approach to one another,
caused as well by their position as by the pressure of the
outer organs. We ought then to study carefully the
new appearances which result from these cohesions.
This union may take place by the ovaries alone ; by the
ovaries and styles ; by the ovaries, styles, and stigmata ;
by the styles and stigmata, the ovaries remaining free ;
and, lastly, by the stigmata alone.

"When two or more carpels are joined together by the
ovaries, there results an ovary composed of several partial
ones, which produce as many cells as there are carpels ;
this union takes place generally only in verticillate car-
pels, and there results a general ovary with cells verti-
cillate around a real or imaginary axis. These cells are
triangular, with the inner angle acute and the external
face convex. We shall see, in speaking of the fruit, the
internal conformations which result from these unions ;
I shall only remark, for the present, that every ovary
with opposite or verticillate cells, is formed by the union

STRUCTURE OF FLOWERS. 73

of the ovaries of several carpels. It is usual in this case
to say, very improperly, that the plant is monogynous
and polystylous, or with one ovary and several styles ;
whilst, perhaps, it would be better to say that it is
Gamogastrous, or has its ovaries united. The union
of the ovaries may take place by the base only, as in
Nigella orientalis ; or halfway, as in Nigella arvensis; or
as far as the apex, which is the case most frequently ;
the partial ovaries united half way form those which are
said to be split or branched.

When, besides the ovaries, the partial styles are also
united together, there results from their cohesion a style
apparently single, but formed in reality of as many par-
tial ones as there are carpels. It is then said that the
flower is Monostylous, which would be more correctly
expressed by the word Gamostylous. In this case the
stigmata or the branches which bear them are distinct ;
they are always equal to, or double the number of that
of the cells of the ovary ; they are equal in number when
the styles which arise from each placenta are united
together as far as the apex ; double the number when
they remain distinct towards the top. Thus the Euphor-
biacese have indifferently three or six stigmata, when
there are three primitive carpels.

Lastly, when the partial stigmata are all united to-
gether, there results an apparently simple one, some-
times round, at others more or less divided into angles
or protuberances, the number of which is equal to, or
double that of the carpels, which are then entirely united.

The cohesion may take place in an inverse manner.
Thus, for example, in several Asclepiadea? the ovaries
remain free and distinct, and the partial styles are united
into a single body, as in Asclepias ; sometimes the styles
are so short, that the union only takes place in the stig-
mata, as is seen in Stapelia. This kind of organization

74 VEGETABLE ORGANOGRAPHY.

can be so little explained in ordinary terms, that no par-
ticular name has been given to it, and the flowers where
this phenomenon exists are sometimes placed among
those with one, and at others among those with two
pistils. Several of the phenomena which the parts of the
pistil present, will only be intelligible when we examine
the structure of fruits.

Section VI-

Of the Torus and the Adhesions which it causes among
the parts of Flowers.

The torus, or proper receptacle of flowers, appears to
be an expansion of the top of the pedicel, from which
arise the petals and stamens, and which may be consi-
dered as the base of all the male or corollary parts of
flowers. This base of the petals and stamens being
formed by the abortions or partial developments of these
organs, does not really deserve the name of organ ; but
we are obliged to describe it by such a name in order to
avoid much circumlocution. Turpin, who also admits
that it is formed by the bases of abortive stamens, has
described it under the name of Phycosteme, which it
would be very convenient to admit, if that of Torus
had not been proposed by Mr. Salisbury some years be-
fore.

It is generally (perhaps always) devoid of stomata ex-
ternally, and tracheae internally ; it is coloured with
various tints, white, red, yellow or blue, but scarcely
ever green ; it does not decompose carbonic acid gas, and
does not become green when exposed to the light ; it
sometimes bears glands and hairs, but these are of a very

STRUCTURE OF FLOWERS. 75

different nature from those which are found upon foli-
aceous organs ; it destroys the oxygen of the surround-
ing air, and transforms it into carbonic acid, furnishing
the carbon at the expense of its substance.

This organ performs especially an important function
in the structure of flowers on account of its productions
and its connexions. Its productions are : — 1st. The
stamens and petals which we have described above in
their ordinary state ; — 2d. Nectariferous glands, to which
we shall presently revert ; — 3d. Different expansions,
which present a great resemblance to the petals or
stamens, and which have often been confounded with
one or the other of them. Thus, for example, we
remark in the Columbine little lanceolate scales, flat
and pointed, situated between the stamens and the
pistil, and which might be called either abortive stamens
or inner petals ; these organs arise from the torus, and
remain sometimes around the base of the fruit. Organs
analogous to these, but of a more petaloid appearance,
larger and more numerous, arise between the stamens
and carpels of Eupomatia Laurina, and are also produc-
tions of the torus. We find in Vceonia Moutan these
same organs united together, and forming a kind of
petaloid involucrum around the ovaries, and in the variety
of this plant which Andrews has named papaveracea
they cover the carpels without adhering to them. Mr.
Brown has remarked that these appendages sometimes
bear anthers, and we are thus authorized in considering
them as abortive stamens. If I have mentioned them here
as productions of the torus, which is true, it is that
their structure will presently serve to enable us to under-
stand the developments of this organ.

The torus, in a very great number of plants, is but
little enlarged, and is strictly reduced to the narrow cir-
cular space which is found between the calyx and pistil.

76 VEGETABLE ORGANOGRAPHY.

It is, then, from this zone, situated below the ovary, that
the petals and stamens proceed ; we call them Hypo-
gynous, and the plants which have this organization
Thalamiflor^e. In this case all the principal organs
of the flower, — the calyx, the ovary, and the productions
of the torus, are necessarily distinct and not adhering
together. But it frequently happens that the torus
extends either inwardly upon the pistil or its support,
or outwardly upon the calyx, or upon both at once, and
it contracts an intimate adhesion either with one or both
of these organs. Let us follow the details and the conse-
quences of these adhesions of the torus.

In a great number of the Leguminosse the torus is
prolonged around a very slender pedicel, which supports
the ovary, and forms a kind of small sheath, sometimes very
short, as in Peraltea, sometimes as long as the pedicel,
and reaching the base of the ovary in Neurocarpi/m
ellipticum and Martiusia. In several Capparideas the
torus is prolonged, and intimately surrounds the support
of the fruit, — for example, in Gynandropsis, — and the
stamens arise above this sheath. In the Aurantiaceae,
the torus, which is thick and glandular, is prolonged and
intimately applied to the verticillate and membranous
carpels of these plants, and increasing with the fruit
forms the glandular, yellow, and valveless envelope
which incloses the carpels. The same thing occurs in
the Poppy, except that the prolongation of the torus is
thin, strongly adherent, and does not reach entirely to
the top of the carpels, so that at their maturity they open
at the apex. It is the same in the fruit of Nuphar ; and
we see that these examples only differ from that of
Pceonia Moutan, above mentioned, in that the prolonga-
tion of the torus does not adhere to the carpels in this
Pasony, whilst it does so in the Poppy and Nuphar.
The torus of Nymphcea presents another peculiarity —

STRUCTURE OF FLOWERS. 77

the stamens adhere by their base to that portion of the
torus which adheres to the ovary, so that they have the
appearance of springing from its sides.

In all these examples, which might be easily multi-
plied, there are evident proofs of this prolongation and
adhesion of the torus with the carpels or their support.
It is only in plants with a free ovary and numerous sta-
mens that we can expect to meet with any evidence of it.

The second case, which is more frequently met with
than the preceding, is where the torus is adherent with
the base of the calyx, as it is from this portion of the
torus that the stamens and petals arise. These organs
seem to spring from the calyx, and the plants in which
this organization takes place are, for this reason, called
CalyctfloRjE ; as in this case the base of the stamens
is a little above that of the ovary, the name of Perigy-
nous has therefore been given to them. We may see
this in the Salicariac-, most Leguminosas, Rosaceae, &c.
The portion of the torus united to the calyx presents
the appearance of a membrane, either petaloid, indu-
rated, or glandular, and perceptibly differing from that
portion of the calyx which is not lined with it.

The immediate consequence of this adhesion of the
torus with the calyx is, that the sepals necessarily
adhere together at their base, into a gamosepalous
calyx. Sometimes this adhesion is extended very far, as
in the Salicaria?, and then the petals and stamens arise
from near the top of the tube ; sometimes it is very
slight, and then they proceed from the base ; in the last
case, which is remarked in the Leguminosas and Tere-
binthaceae, it is sometimes difficult to perceive otherwise
than by analogy whether the stamens be hypogynous
or perigynous. There are cases where the portion of the
torus, united to the calyx, is thickened at the top, and
forms a kind of disc, from which the petals and stamens

78 VEGETABLE ORGANOGRAPHY.

take their origin ; this is seen in several Rhamneae and
Celastrineae.

It may be remarked in general, that when the torus
is not adherent to the calyx, or, in other terms, in hypo-
gynous flowers, the petals of plants of the same family
are either constantly free, as in the class Thalamiflorae,
or constantly united, as in the Corolliflorae ; whilst on
the contrary most families of the Calyciflorae present
almost indifferently the petals free or cohering, as we
see in the Rhamneas, Leguminosae, Cucurbitaceae, Cras-
sulaceas, Portulaceae, Caprifoliaceae, &c.

We have seen what takes place when the torus ad-
heres to the ovary or to the calyx alone ; let us examine
what happens when it is adherent to both organs at once.

It may be prolonged and united to the two organs,
without their being joined together ; this is observed,
although in a very imperfect manner — 1st, In some Le-
guminosae, where the torus adheres to the calyx by the
side where it bears the stamens, and is prolonged on the
other into a little sheath which surrounds the base of
the ovary. 2d, In the Capparidea?, where it is prolonged
along the base of the ovary, and where it often happens
that it adheres also to the base of the calyx, although by
a scarcely apparent prolongation. But this organization
is especially visible in the Passifloreee. Here the torus is
much developed; it expands and is united on one side to
the base of the calyx, which it lines with a petaloid
lamina, and then gives origin to one or more rows of
coloured filaments, free in Passiflora, and more or less
cohering in Murucuja. Besides this expansion, it is
prolonged upon the base of the ovary, which it narrowly
surrounds, and it is from this portion that the stamens
spring. Thus the Passifloreae are calyciflorous, inas-
much as their torus adheres to the calyx ; but they
differ from all the other Calyciflorag, and approach the

STRUCTURE OF FLOWERS. 79

Capparidese, in their stamens arising from that portion
of the torus which does not adhere to the calyx.

Except the small number of examples which I have
mentioned, it generally happens that when the torus ad-
heres to the calyx and the ovary, it tends to unite them
together throughout the whole extent, where they are
found contiguous; we say then that the ovary is adher-
ent to the calyx, or the calyx to the ovary, or simply
that these organs are adherent. This union of organs
so distant from each other only takes place by their
uniting with the intermediate organ. The torus, reduced
to an indistinct lamina throughout all the united part,
is developed above, at the point where the limb of the
calyx becomes free : sometimes it forms an adherent
lamina to this limb, which is then prolonged a little into
a tube, as is seen in several Rubiaceee, as Gardenia;
sometimes it is expanded into a kind of disc, which
partly covers the ovaries, and gives origin to the stamens,
which are then (hardly correctly) said to be Epigynous,
such are the Umbelliferas and RhamneaD ; most fre-
quently it is not perceptibly prolonged either along the
tube of the calyx, or upon the ovary, and then the
petals and stamens arise from the circular line which is
found at the point of separation of the ovary and calyx.
This position has caused the name of Inferior ovary to
be given generically to all those which are adherent, be-
cause they seem in fact below the petals ; or to the co-
rolla that of Superior, because it seems above the
ovary; but the numerous cases where the torus is
prolonged upon the calyx, without the latter adhering
to the ovary, and where, consequently, the corolla ought
to be called inferior, although it is evidently above the
ovary, have caused these expressions, founded upon ap-
pearances, to be abandoned, and that of Adherent
ovary or calyx to be employed, which expresses the fact
without any ambiguity.

80 VEGETABLE ORGANOGRAPHY.

Section VII.

Of Abortions of Parts of the Flower, or of their
Degenerations.

All the parts of flowers may either disappear more or
less completely, or present themselves in unusual forms ;
and it is of great importance, in order to appreciate the
true symmetry of plants, to recognise them under their
different appearances ; it is this that we are now about
to do. very rapidly, examining first the cases where all
the similar parts, that is to say, those which compose
the same organ, undergo the same change.

The calyx is more rarely absent than any other organ,
probably because its external position causes it but sel-
dom to suffer in its development by the pressure of
neighbouring organs. In plants where it is free, I only
know Nemopanthes in which it appears to be entirely
absent, or in which it is reduced to a simple, scarcely
apparent rim. In those where it is adherent, the tube
is united with the torus and ovary, so as to be hardly
visible ; and the limb or part not united is sometimes
absent. Thus, for example, in the Umbelliferae, when
this limb exists, it appears under the form of five little
teeth, as in (Enanthe ; but in a great number of cases it
is completely abortive, and is replaced by a small circu-
lar rim, analogous to that of Nemopanthes.

When the flowers are collected into compact heads,
and inclosed in an involucrum, the calyx becoming as it
were an internal organ, and being submitted to the pres-
sure of the neighbouring flowers or bracts, presents
abortions more frequently. This is but seldom the case
in flowers where it is not adherent ; but Diplolama, one

STRUCTURE OF FLOWERS. 81

of the Rutaceae, presents an example of it; here the five
sepals are reduced to scales, because the flowers are in a
compact head. We find more numerous and decided
examples in families where the ovary is adherent, and
the flowers capitate, such as the Dipsacess and Compo-
sitae. In these plants, the tube of the calyx is reduced
to a thin membrane adhering to the ovary, and the limb
appears under different forms. Sometimes it has five
foliaceous teeth, like those of ordinary calyces, as in
Catananche. These teeth are sometimes changed into
membranous scales, free, as in Centaurea Cupina, or
united together, as in Hymenopappus and Favonium ; or
into almost spiny points, as in a species of Cnicus ; or
into bundles of simple hairs, as in Sonchus; united
together, and thus appearing branched, as in Stcehelina ;
or feathery, as in Scorzonera.

It is so certain that the pappus of the Compositae is
the true limb of the calyx, that it sometimes puts on
every appearance of it; thus M. Dufresne has shown
me a plant of Podospermum laciniatum, (PI. 18, fig. 1, 2,)
the pappus of which was replaced by five linear, slightly
foliaceous lobes.

I shall revert to these different forms of the pappus
when speaking of the fruit, and I shall only remark here,
that every organ called Pappus is only the limb of the
adherent calyx of plants with capitate flowers, in which
this organ is half abortive, or deformed by the pressure
of neighbouring flowers ; sometimes even it is entirely
abortive, and we then say that the pappus is absent : it
is replaced by a small circular rim, as in most Umbel-
life rae.

The Valerianeae, although having their flowers dis-
tinct and not capitate, present also a true pappus ; this
results from the limb of the calyx being rolled inwards
during flowering, and submitted consequently to a pres-

VOL. II. <;

82 VEGETABLE ORGANOGRAPHY.

sure and etiolation as powerful as that which in the
Dipsaceae results from the vicinity of other flowers. The
genera where the limb is not rolled inwards have it
developed into foliaceous teeth, as in ordinary calyces.

The abortion or complete absence of the sexual
organs of plants, or of one of them, is a phenomenon
which constantly happens in all plants said to be Uni-
sexual, and accidentally in several others. Thus, to
commence with the latter case, which is the clearer,
Lychnis dioica, although belonging to a family usually
hermaphrodite, presents certain individuals where the
female organs are well-developed, and then the stamens
are reduced to simple rudiments ; and others where the
stamens are well developed, and where the pistil is
abortive, so that in its place is only seen a little pro-
tuberance with the rudiments of five stigmata. The
same phenomenon takes place in Spircea Aruncus and
Sedum Rhodiola, Sec. All plants which present this
phenomenon accidentally are said to be Dioecious by
abortion ; thus, in several Compositae, part of the
flowers of each head are devoid of ovary, style, and
stigma by abortion, and others have no perfect stamens,
so that they are Moncecious by abortion. Thus, in
Diospyrus, Gleditsia, &c, a part of the flowers are
devoid of pistils, another of stamens; and we find,
besides, others where the two organs exist together,
constituting that state of flowers which botanists have
called Polygamous by abortion.

These three systems of flowers, unisexual by abortion,
are frequently met with in almost all the families where
we also find hermaphrodite ones ; such are the Caryo-
phyllea?, Compositae, Valerianese, Ebenaceae, Thyme-
laeae, Leguminosae, &c. ; and in all these cases it is
evident that the two sexes exist typically, and that one
of them is not developed.

STRUCTURE OF FLOWERS. 83

When the female organs are not entirely abortive, we
find in their place, sometimes, a portion of the ovary
deformed for want of fecundation, and at other times, a
glandular body. When this is the case with the male
organs, we find in their place a portion of the filament,
or a glandular body which indicates their disappearance.

But we find whole families, or nearly so, in which the
flowers are unisexual, and where no rudiments of the
abortive organs are perceived ; whence several naturalists
have concluded that there are flowers where one of the
sexes is essentially wanting. There is no reason that
this cannot be the case, and that flowers cannot be met
with which are formed of only two or three verticils,
one or two of which serve as the protecting organs, and
the innermost only is changed into the sexual ones.
However, I am inclined to believe that if this pheno-
menon take place in phanerogamous flowers, it is very
rare ; for there is hardly any family said to be unisexual
in which we do not find flowers constantly hermaphrodite:
such are the Elm among the Amentacese ; Melothria in
Cucurbitacea? ; Agdestis in Menispermea?, &c. We
even find individuals accidentally hermaphrodite in
certain species in families which pass for unisexual ones :
such are several Poplars and Willows among the
Amentacese ; the Hemp in Urticese, &c. As to families,
such as the Coniferae and Euphoi-biacea?, where we find
no example of hermaphrodite flowers, we may consider
them as presenting an abortion more constant than the
preceding, or as being essentially formed of a smaller
number of verticils.

There are other cases where the sexual organs cease
to perform their functions, and take an extraordinary
development. Thus the styles of the Anemone some-
times become large and petaloid by culture ; the branches
of the style of the Tris, although furnished with a

o2

84- VEGETABLE ORGANOGRAPHY.

stigma ill the form of a transverse fold, are usually in a
petaloid state ; a great number of double flowers also
have their styles developed into petaloid laminas ; thus
proving the peculiar analogy of the styles, stamens, and
petals.

The degenerations of the male organs are still more
frequent. When the anthers are abortive, the filaments
are changed into laminae perfectly like the petals of the
plant : this is what we see in common double flowers.
When the anthers themselves remain, although becoming
sterile, it happens sometimes that they are developed in
the form of horns : this happens in several Ranunculaceae.
The Columbine presents this very remarkably. By
culture we obtain two double monstrosities : the one
with all the petals flat, owing to the development of the
filament and the absolute abortion of the anther, as
Aquilegia vulgaris stellata ; the other with all the petals
horn-shaped, resulting from the non-development of the
filaments, and the extraordinary increase of the anther,
as A. vulgaris corniculala.

The degenerations of the petals are the more difficult
to recognise, as they themselves are constantly in an
intermediate state between the primitive one of a leaf
and that of a stamen. Every form is met with in these
organs. The principal modification is owing to the pre-
sence of certain glands, which cause the origin of spurs :
it happens in certain gamopetalous flowers that the un-
equal union of the petals is very manifest, and produces
different appearances.

The abortion of the petals is more difficult to reduce
to general laws than the preceding phenomena. Let us
first take the most simple cases. That there are plants,
the petals of which are accidentally abortive, it is
difficult to doubt : thus Sagina apetala sometimes
presents very small ones, sometimes it is entirely devoid

J

STRUCTURE OF FLOWERS. 85

of them. Thus a great number of apetalous plants are
so analogous in their whole symmetry to those furnished
with petals, that it is impossible not to think that their
absence is only owing to their non-development. Let
us observe here, that they are thus accidentally wanting
only in polypetalous flowers, and that there is no
authentic example known of an abortive corolla in
gamopetalous plants, if it be not so, perhaps, in some
cases where the stamens are wanting at the same time,
as in Gymnostyles and Fraxinus : when the petals are
abortive, there sometimes remains in their place either
a petaloid rudiment or a glandular body. We also say
that they are absent when they are accidentally changed
into stamens, as in the singular variety of Capsella
Bursa pastoris, of which M. De Jacquin has kindly
communicated to me a specimen and a drawing, which
I here give (PL 18, fig. 3, 4). In this monstrosity,
which can be perpetuated by seeds, we find the flowers
with ten stamens, instead of six of these organs and
four petals : I have found an analogous fact in a mon-
strosity of the common Bean, where the two wings of
the corolla were changed into stamens. We shall treat
more at length upon this kind of transformation, and
revert to the case where the petals are constantly wanting.
This will form the object of the following Section.

86 VEGETABLE ORGANOGRAPHY.

Section VIII.

Of Monochlamydeons, or incomplete Flowers, or those
which have but one envelope.

When a flower presents a single envelope, is it a
corolla, calyx, a union of both, or an organ different
from either ? All these opinions have been maintained,
and deserve to be examined.

Tournefort, who made the character of the calyx to
consist in its being persistent, and that of the corolla in
its being caducous, found himself compelled, from this
false definition, to give different names to organs
evidently similar in analogous plants: thus he called
corolla in the Tulip the organ which he named calyx
in the Narcissus. Linnaeus did not attach any im-
portance to this definition, perhaps on account of that
which he had adopted ; he admitted, in fact, that the
calyx was the prolongation of the bark, and the corolla
of the liber. This distinction is hardly capable of being
maintained either in Monocotyledons, where there is
neither liber nor bark, or in Dicotyledons, when the
liber is nothing but the younger cortical layers. Thus
in practice Linnaeus usually called calyx that which was
green, and corolla that which was coloured; thus the
single envelope in Monochlamydeous Dicotyledons was,
according to him, calyx in Chenopodium, corolla in
Daphne ; and in Monocotyledons, calyx in Juncus,
corolla in the Liliacea? ; often he says " calyx nisi
corolla mavis," Sec. Lamarck, in his first works, defined
the corolla as the organ nearest the stamens, and con-
sequently named all single envelopes as such ; but he
afterwards abandoned this opinion. These different
modes of expression might perhaps suffice when it was

STRUCTURE OF FLOWERS. 87

a question of purely artificial order ; but it is important,
as regards both the natural order of classification and the
physiology and comparative anatomy of plants, to fix
our ideas upon this subject, and to be able to compare
together organs truly analogous.

Those who have paid any attention to this subject
have thought that the floral envelope, when it is single,
is not a corolla, because it is often green and foliaceous,
and frequently adherent to the ovary, which true
corollas never are, and because the corolla appears more
disposed to be abortive than the calyx ; I only truly
know Nemopanthes of which it might be said with any
reason that it has a corolla and no calyx, but this is
simply because the latter is reduced to a circular rim.

Jussieu, uniting in the definition of calyx the con-
ditions of Tournefort and Linnaeus, has established that
the floral envelope, when single, is always a calyx.
This opinion cannot be called in question when it occurs
in Dicotyledonous plants which belong to families
usually furnished with calyx and corolla, but which are
devoid of one of these organs ; in this case it is
evidently the petals which are absent, as, for example,
in Clematis, the apetalous Capparideas, Caryophyllese,
Rutaceas, Rosaceaa, and Ficoids. The analogy with
neighbouring genera evidently shows it; and if one
wished to maintain that some of these organs could not
be calyces because they are coloured, I would call to
mind that the calyx, and even the bracts, of Hortensia
and Salvia splendens are as much coloured as the most
brilliant corollas ; I would add, that these single en-
velopes perform the part of true calyces, both in
bearing the stamens in calyeiflorous plants, and not
doing so in thalamiflorous ones, and in their being fre-
quently adherent to the ovary, Sec.

The question is more difficult when it takes place in

88 VEGETABLE ORGANOGRAPHY.

Dicotyledonous families which have constantly or usually
a single envelope to the flower. Jussieu, deciding the
question, has given them the name of ApetaLjE, and to
their envelope that of calyx ; on account of the un-
certainty which exists, I have decided to call these
plants Monochlamyde^e, and their envelope a Peri-
gone, neutral terms, which express a fact without
declaring an opinion.

The reasons for which this envelope may be compared
to a calyx, are: — 1st, its extreme analogy with the
calyces of plants which are accidentally devoid of petals ;
2d, its frequent adhesion with the ovary ; 3d, the
greenish and foliaceous appearance of several ; 4th, the
analogy of structure of several monochlamydeous fa-
milies with those usually furnished with petals, such as
Amaranthaceae with Caryophylleae, Juglandeae with
Terebinthaceas, Euphorbiaceae with Rhamneaa, Elasag-
nese with Combretaceae, &c. ; 5th, the existence in
several, especially in Thymelaeas, of small petaloid scales,
which may be true petals.

On the other hand, I consider that the external sur-
face of these single envelopes has all the characters of
a calyx : it is usually green ; it constantly presents
stomata, even when it is coloured, as in the Marvel of
Peru ; it frequently bears hairs or glands analogous to
those of the leaves, as in Elaeagnus ; but the inner
surface almost always presents the characters peculiar to
the sexual organs ; it is coloured, has no stomata, and
bears neither hairs nor glands analogous to those of the
leaves. We might conclude from these facts that it is
a calyx lined internally by the torus, or a petaloid ex-
pansion of it. This hypothesis would be confirmed by
this consideration, that with the exception of the Ama-
ranthaceae, which must, perhaps, be placed among
Thalamiflorae by the side of the Caryophyllea?, all the

STRUCTURE OF FLOWERS. 89

other Monochlamydeous families have perigynous sta-
mens, and the torus consequently adherent to the
calyx. Finally, whether we say that it is a calyx, or
that it is a double one with a petaloid lamina, all the
consequences are found the same, and therefore the
difference is of but little importance.

We shall now examine the envelope of Monocotyle-
donous flowers, and here we shall find some new diffi-
culties. Desvaux, considering that it is always formed
of two rows of pieces placed alternately, has proposed
to consider the outer one as a calyx, and the inner one
as a corolla. This mode of viewing it seems particularly
authorized : — 1st, by the structure of the Commelineas,
Alismacese, and several Amomeae, where the outer row
has a perfectly calycine appearance, and the inner one a
perfectly petaloid one ; 2d, because the aestivation of
the two rows is often very different, as, for example, in
Tradescantia, where that of the outer row is valvate, and
that of the inner irregularly twisted. This manner of
expression would often be advantageous for the clearness
of descriptions ; but as to the reality, it appears to me
hardly admissible: in fact, in the majority of cases these
two rows are perfectly similar, and especially in all
Liliaceous plants with an adherent ovary, the two rows
are equally united to that organ, whilst true corollas
never are. It must then be admitted, that the two rows
form part of a single envelope, which Linnaeus calls
corolla, Jussieu calyx, and I perigone.

The reasons which I have above mentioned, and
especially the adhesion with the ovary, prove that it is
not a true corolla. The idea of considering it as a calyx
presents the same difficulties which I have shown in
monochlamydeous Dicotyledons ; and, moreover, in these
two circumstances: — 1st, that the stamens are most
frequently hypogynous ; 2d, that when the flowers

90 VEGETABLE ORGANOGRAPHY.

become double, which is frequently the case, the stamens
are transformed into petals so similar to the pieces of the
perigone that it is difficult not to believe that they are of
a very analogous nature.

If we add to these reasons, that this envelope is often
green on the outside and coloured internally, that it
alwavs has stomata on the outer surface and none on
the innner, we shall perhaps be inclined to conclude,
that this perigone is formed of a calyx lined, thus to
speak, with a petaloid expansion of the torus. I have
only given this opinion as a simple hypothesis ; but I
think that it is more prudent, in the actual state of
the science, not to adopt terms which decide the ques-
tion too clearly, and that it is well to retain for these
doubtful cases of a single envelope a particular name.
I have adopted, after Ehrhart, that of Perigone, which
signifies, around the sexual organs ; and following the
analogy of the terms sepals and petals, I propose to give
to the pieces of which it is formed the name of Tepals.

Some authors, adopting my idea, have given the single
envelope the name of Perianth ; but I think it better
to retain that of perigone : — 1st, because that of peri-
anth was understood by Linnaeus to designate the true
calyx ; 2d, because this term, which signifies around
the flower, would be more applicable to an involucrum
than to an organ which forms part of the flower ; 3d,
because the word perigone has been proposed in this
sense before that of perianth, and we ought always to
avoid useless changes of nomenclature. The term beina:
once admitted, we must apply to the perigone all that
has been said of calyces and corollas, inasmuch as
they are formed of pieces, sometimes free, at other
times cohering ; all that has been said of calyces, as
regards their adhesion with the ovary ; and all that has
been said of petals, inasmuch as they are analogous to

I

STRUCTURE OF FLOWERS. 91

the development of the filaments. Admitting this
manner of viewing it, we shall understand how the
perigone is sometimes adehi'ent to the ovary, or com-
posed of parts opposite the stamens — characters peculiar
to the calyx ; whilst in other plants, it is free, odorous,
has its lobes alternate with the stamens, and becomes
double and multiplied by the abundance of the sap —
characters peculiar to the corolla.

The perigone is sometimes reduced, by abortion, to a
simple rudiment : this is observed among Dicotyledons,
in the Euphorbiaceas, and especially in those with the
flowers in compact heads. It is this which happens
among Monocotyledons in the Graminese, where the
perigone appears represented by the lodicules ; their
number is ternary in Bambusa and Glyceria ; sometimes
the third is smaller, and its absence in several cases may
result either from an abortion more or less complete, or
from their intimate union.

Section IX.

Of the relative Position of the parts of one floral
Verticil compared with that of another.

The position of the parts which compose the floral
verticils is susceptible of every modification which
results from each of them, being either between or
opposite the pieces of the outer verticil. The first case,
that is to say, that where each piece is between the two
outer ones, is so much more frequent than all the others,
that we may believe that it is the natural state, inasmuch
as it is conformable to the disposition of the successive

92 VEGETABLE ORGANOGRAPHY.

verticils of leaves. Thus, the petals of flowers which
are regular, and the parts of which are equal in number,
arise usually between the sepals, the stamens between
the petals, and the carpels between the stamens. But
there are some exceptions to this rule : thus, we find
the petals opposite the sepals in the Barberry ; the
stamens opposite the petals in the Primulaceae, Myr-
sineee, &c. As to the real position of the carpels,
it has been much less studied than that of the other
organs, and without doubt it would give some interesting1
characters in certain families ; but the frequency of
their abortion renders their observation delicate. Some
examples recently observed, make me think that in
perfectly regular plants, where the number of parts is
equal in all the verticils, the carpels are always alternate
with the sepals, whatever may be the position of the
verticil nearest to them : thus, the carpels of the Cras-
sulaceas are alternate with the sepals, both in the genera
Crassula, Roche a, &c. which have the stamens alternate
with the petals, and in Sedum, Cotyledon, Sempervivum,
&c. which have the stamens double the number of the
petals; the one set alternate, the other opposite them.

The different dispositions of the parts of the flower
may be modified by the number of rows of each verticil,
or by the abortion of parts, or because, in several cases,
there is developed a cluster of organs resembling that
where there is usually found but one : thus, for ex-
ample, in several Homalineas we find a tuft of stamens
situated in the angle of two contiguous sepals. The
same thing takes place in the Myrtaceae, in a very
singular manner : thus, the bundle of stamens formed
by the union of several, are opposite the petals in
Melaleuca, and alternate with them in Astartea. Several
double flowers present a fasciculated development, which
deserves to be noticed : thus, it is not rare to find

STRUCTURE OF FLOWERS. 93

bundles of petals arising from the place whence there
ought to proceed but a single petal or stamen ; this is
what is very well seen, for example, in certain double
Primroses ; but this peculiar case of multiplication leads
us to examine this subject in a general manner.

Section X.
Of the Multiplication of the Floral Organs.

The organs which compose the flower may be in-
creased, as to their number, in two ways : —

1st. The usual number of verticils may be increased
by new verticils, like one of them, which are developed
in a regular but supernumerary manner.

2d. The number of pieces of one verticil may be
increased by the unusual development of organs re-
sembling those of which it is composed.

These two phenomena have been indifferently called
by the names of Doubling or Multiplication; but I prefer
the latter term, which appears to me less hypothetical
than the other.

§ 1. — The Multiplication of the Hows of Verticils.

The multiplication of the rows of one verticil is a
fact which is accidentally observed in several plants, and
which may affect all the organs. Thus, —

1st. As concerns the Bracts, there is cultivated in
gardens a variety of the Pink, which some call Dianthus

94 VEGETABLE ORGANOGRAPHY.

Caryophyllus imbricatus, and in which the number of
bracts, situated at the base of the calyx, instead of
being four, i. e. two pair, we find fifteen or twenty pair,
crossing each other at right angles, and imbricated ; on
account of this great multiplication, the flower is fre-
quently not developed. It appears to result from the
premature transformation of the upper leaves into bracts.

2d. As regards the Perigone, we find in gardens a
variety of the white Lily, the tepals of which, instead of
being in two rows, and six in number, are disposed in
an indefinite number of imbricated verticils. In this
case the stamens and carpels are absent, or transformed
into tepals ; but we cannot say that the phenomenon is
simply owing to this transformation, for the number of
verticils is much greater than the whole usual number
of the floral organs ; it is, then, a multiplication of the
normal number of verticils. In another monstrosity of
the Lily, we find the parts of the perigone multiplied,
and the stamens also existing. All Monocotyledons
with double flowers present, here and there, analogous
facts. The inner tube, or, as it is called, the crown of
the Narcissus, may be classed here.

3d. The multiplication of the rows of the Calyx, pro-
perly so called, is more delicate to confirm, because of
the difficulty of exactly distinguishing the supernu-
merary rows of sepals from simple bracts. Some calyces
of the Berberideaa and Ericaceae appear to afford ex-
amples of this sort.

4th. The Corolla frequently has multiplied rows :
one of the most curious examples is that which Datura
fastuosa (PI. 18, fig. 5) presents, where we frequently
find two or three corollas inserted, as it were, into one
another, and having their lobes alternate. The same
phenomenon has been observed in several Campanulas,
some Labiatas, &c, and it seems possible in all gamo-

STRUCTURE OF FLOWERS. 95

petalous flowers. When this multiplication is limited to
one or two inner rows, it happens then either that the
inner corolla bears the stamens as usual, or that they are
absent. In this last case we may say that the corolla
results from the simple transformation of the stamens
into petals ; but in the first it is evident that there has
been a multiplication of the usual rows. The same
phenomenon is also met with in poly petalous flowers, as
the Pink, &c.

5th. The Stamens very frequently present this mul-
tiplication of rows, especially in the genera where the
number of rows is naturally considerable ; thus on com-
paring together several flowers of the same species of
Poppy, we find that the number of their verticils is very
variable.

6th. Finally, the Carpels, which are less numerous
and more central, rarely present this accidental multipli-
cation ; however, from time to time we find double rows
among the Ranunculaceae and Rosacea^ with verticillate
carpels. I have met with a very remarkable example of
this accident in Gentiana purpurea : I give a figure of it
in PI. 18, figs. 6, 7. Here we see two rows of ovuliferous
carpels ; the outer of four, the inner of two.

But if all the floral organs can accidentally present
the multiplication of the rows of which they are usually
composed, is it not likely that this phenomenon may be
constant in certain plants, perhaps in certain families ?
And are not the genera, such as Nymphcea, Mesem-
bryanthemum, &c , where the parts of the flower are
presented in a very great and indefinite number of rows,
evident examples of this opinion ?

96 VEGETABLE ORGANOGRAPHY.

§ 2. — The Multiplication of Parts of a Verticil.

The second kind of multiplication of the floral organs
is that where the usual number of the parts of a verticil
or row is increased. This may take place in different
ways : —

1st. The absolute number of all the verticils of a
flower may be augmented by the addition of one or two
pieces to each : thus, it is not rare to find flowers of the
Colchicum with seven or eight lobes, and as many
stamens ; flowers of the Rue and Philadelphus with
sometimes four, at others five parts, &c. In these cases
we must first examine if it be not the higher number
which is the usual state, and then the diminution of
number is classed among the cases of abortion ; but in
the contrary case, the multiplication appears to result
from the natural union of two flowers, as I have else-
where explained.

2d. In the place of an organ apparently single, but in
reality composed of several intimately united, we may
accidentally find them becoming free. Dunal made
known a curious example of this in Laurus nobilis. We
know that the stamens of this tree have on each side of
the lower part of their filaments a glandular bifid body,
borne on a short filament intimately united to that of
the stamen ; it appears that this body is an abortive
stamen, and consequently that of the Laurel is a bundle
of three united, the two lateral ones being abortive : it
happens, in fact, that the three stamens are sometimes
developed, and the whole number is found tripled, and
none of them bears the glandular body on its filament.
Several peculiar facts of the history of polyadelphous
flowers appear to coincide more or less clearly with this
example, which may be considered a complication of
union and abortion.

STRUCTURE OF FLOWERS. 97

3d. In the place where, in the usual course of vege-
tation, there arises a single organ, we sometimes see a
cluster of analogous ones developed. Thus, each of the
stamens in certain monstrosities of the Primrose, instead
of being changed into a single petal, is transformed into
a cluster united at the base. It seems that this fact is
analogous to what is constantly found in certain flowers
in which we see a cluster of organs united where, from
analogy, we ought to find but one : such are the bundles
of stamens alternate with the petals in Melaleuca and
several species of Hypericum.

4th. A fact analogous to the preceding appears to take
place in certain cases, with this difference, that the mul-
tiplied organs which, from symmetry, appear to replace
a single one, are completely free at their base ; thus, in
Lagerstromia, we count five large stamens alternate
with the petals, and four or five small ones which, situ-
ated opposite each petal, seem to represent by their
union a single stamen. This fact, combined with the
abortion of the large stamens, seems to render intelligible
the structure of several By ttneriacese. This class appears
to be connected with the preceding by the example of
the Cruciferae, where the two pair of large stamens, some-
times free, sometimes more or less united, seem, from
symmetry, to replace a single stamen.

5th. Lastly, it sometimes happens that the two parts
of an organ are so separated at their base as to appear
to form two distinct ones. Thus, Impatiens Noli-tangere
has four petals and five stamens — three alternate with
the petals, and two springing side by side at the point
where the fourth ought to arise in the regular state.
For, the three solitary stamens between the petals have
two-celled anthers, and those in a pair have them uni-
locular, and seem consequently to be a stamen divided
into two. The term of doubling applies very well to

VOL. II. H

1)8 VEGETABLE ORGANOGRAPHY.

this case ; that of multiplication represents better the
preceding, where all the supernumerary organs result
from all the parts of a single organ. It is true that they
are generally of smaller size, but it is likely that this is
referable to the general law of vegetation, — that where a
large number of organs is produced within a given space,
they find there less nourishment, and take a less degree
of development.

§ 3. — General Examination of Double Flowers.

It is customary to designate under the name of
Double Flowers (Jtores pleni) all those where all the
different floral organs, or one or two of them, take the
appearance of petals, and those where the number of
petals is, or appears to be, increased by any cause.

Double flowers ought, in my opinion, to be classed in
three divisions : —

1st. Petalodeous Flowers (fiores petalodei), that
is to say, those which become double by the simple de-
velopment into petals of all or any of the floral organs :
such are those where this development affects the bracts
(Hortensia), the calyx (Primula calycanthema), the
stamens (Rosa, See), or the carpels (var. of Anemone
nemorosa, &c.) We may also distinguish two cases
among petalodeous flowers which proceed from the de-
velopment of the stamens — viz. that where it takes place
by the dilating of the filament and the entire abortion of
the anther; and that where the filament remains in its
natural state, and the cells of the anther are developed
into petals. In the first case, which is by far the more
frequent, the supernumerary petals are always flat; in
the second they are horn-shaped. The Ranunculaeea?
present this double mode of transformation in a singular

STRUCTURE OF FLOWERS. 99

manner ; the Clematideas double after the first mode,
the Ranunculeae after the second, and the Helleboreae
present both. There are species even which become
double in both ways : thus, Aquilegia vulgaris, when its
filaments are changed into flat petals, forms the variety
called stellata ; and when its anthers are changed into
horn-shaped ones, it forms the variety caniculata when
the horn is erect, and inversa when it is reversed by the
torsion of the filament.

2d. Multiplied Flowers (flores multiplicati) are
those where the number of petals is augmented by the
increase of the number of rows of floral verticils, or of
parts of these rows, and their transformation into petals.
In the preceding class the number of parts was not
augmented, and there was only transformation ; here
there is increase in number, and transformation. All
the examples mentioned in the two first divisions of this
section belong to this class.

3d. Permuted Flowers (flores permutati) are those
where the abortion of one of the sexual organs causes a
remarkable change in the form or dimension of one of
the floral envelopes ; thus, the abortion of both or one
of them in the Compositae frequently causes a change of
form in their corolla ; it sometimes, remaining tubular,
becomes larger than ordinarily, as is seen in some
varieties of Asters, African Marigolds, &c. ; sometimes
it is transformed into a flat strap, which is the most
usual case in the Compositae called double in gardens.
Similar phenomena are met with in the Guelder Rose
(Viburnum opulus), the sterile flowers of which have
the corolla larger than the fertile ones : in the natural
state the lateral flowers alone present this phenomenon ;
but in the cultivated variety all are enlarged, owing to
the abortion of the sexual organs.

Thus the name of double flowers is applied, in ordi-

ii 2

100 VEGETABLE ORGANOGRAPHY.

nary language, to very different phenomena. Organ-
ography instructs us to class them, to compare them
with natural phenomena, and to refer them to known
analogies ; but it will be for Physiology to determine,
if possible, the causes of these different metamorphoses,
which are less unworthy than has been thought of the
observations of the Botanist, since they are intimately
connected with the study of the organic symmetry of
plants.

Section XI.

Of the Inequality of Parts in a Floral Verticil; or of
Irregular Flowers.

The different verticils which compose a flower, may
be, with regard to one another, of very unequal size :
some may even be entirely wanting, without the flower
ceasing to be regular ; for each of its portions, taken
from the centre to the circumference, resembles the
others ; but the name of Irregular is given to flowers
in which one or more parts of a verticil are different
from the others in size, form, direction, situation, or de-
gree of cohesion.

In order to form a just idea of the symmetry of
flowers, we must always endeavour to refer irregular
ones to regular types, from which they seem to be degene-
rations. Each family appears to have a regular type,
which is its natural state, and from which they differ,
either accidentally or constantly, from different causes.
When these causes result from foreign influences on the

STRUCTURE OF FLOWERS. 101

plant — as, for example, mutilations owing to culture, the
unequal action of light, the pressure of neighbouring
bodies, &c. — then the irregularities are purely accidental ;
when, on the contrary, it results either from the mode
of development of the neighbouring organs, or, what is
more frequently the case, from the disposition of the
flowers, either with regard to each other or to the stem,
then the irregularity is constant, and the flower only
presents its natural state in very rare cases, and which
may, in their turn, be called accidental.

The disposition of the flowers is, of these causes in-
herent in plants, the one of which we are the best
able to appreciate the effects. Thus, for example, when
they are near together, either in compact spikes or ra-
cemes along the axis, or in heads or umbels, the inner
or upper part of the flower, i. e. that which is nearest
the axis, is restrained in its development by the pressure
of the flowers against the axis or against each other ;
whilst the other side is more at liberty; whence it
results, that there is sometimes a complete or incomplete
abortion of some of the parts bordering upon the axis,
and the development of the opposite side ; sometimes a
more lengthened and complete union of the parts in the
neighbourhood of the centre, whilst those on the oppo-
site side are more free ; sometimes a union of the two
facts which I have mentioned.

The general result of the pressure is counterbalanced,
sometimes even disguised, by another cause : viz. in a
flower when one of the parts of a verticil is entirely or
partly abortive, the corresponding part of the neigh-
bouring verticil takes a greater development than usual,
because it gains advantage either from the place or
nourishment which the other would have made use of;
whence it results, that it is extremely rare that the irre-
gularity of one of the floral organs does not produce an

1053

VEGETABLE ORGANOGRAPHY.

irregularity in the others. Let us follow the application
of these principles to the different organs of the flower,
and to the different kinds of irregular flowers.

The sepals, in consequence of their foliaceous nature
and external position, are more liable than all the other
organs to the action of external causes ; thus we find
irregular calyces even in flowers otherwise regular ; thus
one of the free portions of the sepals of Musscenda and
Pinckneya is spread out into a limb much larger than
the other. The same phenomenon takes place, although
in a less decided and constant manner, in Roses.

The petals present inequalities of size, which result
from the unequal development of the neighbouring
sepals, or from the different modes of their metamor-
phosis.

The pieces of the calyx, corolla, or perigone, are often
united together in unequal degrees ; when the inner or
upper pieces are united at a different point from where
the lower ones cohere : it results that the flower has two
lips, an upper and lower one ; and it is so true that the
flowers with a labiate calyx or corolla owe this irregu-
larity to their position with regard to the axis, that we
never find the lips lateral, but always superior and
inferior, as we see in the labiate calyces of the Pa-
pilionaceae, Labiatas, Scrophularineae, &c, in the co-
rollas of the two last families, or the perigones of the
Orchideas, &c.

The stamens are organs very subject to irregularity,
even in plants where the rest of the structure is regular :
it must, however, be remarked, that they may be un-
equal with regard to each other without being irregular;
thus, in several flowers which have the number of
stamens double that of the petals, they are alternately
long and short, early or late in coming to perfection ;
and in this case, it is those which alternate with the

STRUCTURE OF FLOWERS. 103

petals which are the longest, most forward and constant.
When they are in several rows, the rows, when com-
pared together, are sometimes very different in size ;
but as long as all those of the same row resemble each
other, the flower is regular. The inequality of the
stamens may result either from unequal degrees of co-
hesion with the corolla, calyx, or perigone ; from un-
equal degrees of cohesion with each other ; from the
inequality of the length of the filaments ; from the un-
usual development of these filaments ; from the total
abortion of the filaments or anthers, or from their being
deformed.

The place of the stamens being always determined
with regard to the petals, we may, on attentively study-
ing them, easily perceive the total abortion of some of
them ; thus, when a flower has the corolla with five
petals, free or united, if we observe that the stamens are
alternate with or opposite the petals, we shall imme-
diately perceive if there be a vacant space ; as takes place,
for example, in the Labiatas and Scrophularinea? : in this
case, the place is either completely vacant, or marked by
a small glandular point on a little filament ; and it is so
certain that these are the rudiments of undeveloped
stamens, that it is not rare to see certain flowers with
them developed into true stamens. When this pheno-
menon takes place, the rest of the flower also becomes
regular : this accident, or rather this return to symmetry,
is well known in Linaria vulgaris ; but it is not limited
to this plant, as was at first thought : it is found in
several species of the genera Linaria, Antirrhinum,
Digitalis, Sesamum, Galeopsis, Viola, Orchis; and we
are authorized in considering it as a phenomenon common
to all irregular flowers.

All these same accidents, especially that of total
abortion or nearly so, are common in the carpels. When

104 VEGETABLE ORGANOGRAPHY.

among these there is but a small number abortive, and
there remain at least two complete ones, the pistil still
presents the appearance of regularity if considered with
regard to itself; but it appears irregular when compared
with the number of the other parts of the flower. Thus,
the Cistineffi, which have five sepals, five petals, and five
carpels, are regular ; those which, as Helianthemum,
with the same floral numbers have but three carpels,
have the fruit intrinsically regular, but relatively irre-
gular. When the number of carpels is reduced by
abortion to one, this one always presents traces of irre-
gularity ; thus, when it has several seeds, they adhere
laterally on the side nearest the axis of the flower, as is
jilainly seen in the Leguminosas. It is remarked, that
when these plants have accidentally several carpels, i. e.
when the abortion is less complete, the second, when it
exists, is situated precisely opposite the first, having the
seminiferous suture also directed towards the centre, so
that a regular fruit is the result ; I have observed this in
Gleditsia. The drupaceous Rosacea? (Amygdalaceae)
also have a single carpel by abortion ; and we find
Cherries and Plums which have accidentally either two
carpels united, or several free. Auguste de Saint-Hi-
laire found in Brazil a Mimosa with five carpels ; and
when we compare the structure of the Leguminosa?, thus
considered, with the Spiraeaceae among the Rosacea?, we
shall perceive that these two families hardly differ more
than in the abortion of the carpels being frequent in the
Leguminosas, and rare in the Rosacea?.

In carpels which appear to have but one seed, the
irregularity is visible in two respects : — 1st, It is almost
certain that the seeds are only solitary by the more or
less early abortion of one of the ovules ; — 2d, From the
manner in which the seed is situated, there must be some
irregularity in the carpel. Is it attached laterally, as in

STRUCTURE OF FLOWERS. 105

the Leguminosae ? the irregularity is evident, for the
placenta is lateral. Is it attached to the base of the
carpel, as in the Compositae ? the pistillary cord follows
one of the sides of the pericarp, and causes an irregu-
larity. Is it attached to the apex of the carpel, as in the
Dipsacese ? the vessel that supplies it with nourishment
passes along one of the sides of the carpel, and makes it
necessarily irregular. Thus, every monospermous car-
pel, every solitary carpel, is naturally a deviation from
the symmetrical order, and, consequently, an irregu-
larity most probably caused by abortion.

Section XII.

Of the Primitive Disposition of the Parts of a Floral
Verticil; or of the Aestivation.

The complete and regular flower is, as we have seen,
composed of at least four concentric verticils, each formed
of several pieces, the relative disposition of which we are
about to consider. The rapid development of these dif-
ferent organs at the period of flowering is such, that we
cannot judge of this primitive disposition unless we study
it in the buds. Linnseus, who compared it with the
vernation of leaves, has given it the name of Estiva-
tion. Richard has proposed to substitute that of
Prefloration ; which perhaps would be preferable, if
it were worth while to change a term which is not erro-
neous.

This disposition of the parts is especially important to
be observed in that which relates to the envelopes of the

106 VEGETABLE ORGANOGRAPHY.

flower, viz. the sepals, petals, and tepals, where the
pieces, free or united at the base, form the calyx, co-
rolla, and perigone. Let us first examine the dispo-
sition of flowers strictly regular.

We must first observe whether the parts of an organ
are in one row, or if they be in two or more ; when they
are strictly verticillate in a single row, there happen four
cases : —

1st. These parts may be disposed in a perfect circle,
each of them being fiat or slightly convex ; then they
all touch by their margins without overlapping each
other, or without being folded inwards : this is called
Valvate ^Estivation, because it is analogous to the
disposition of the valve of Pericarps, (PI. 19, figs. 2s, 3s,
4s, 15p.) The sepals of the Lime-tree and of most
Clematideae, the petals of the Vine and of the Arali-
aceae, the outer tepals of Tradescantia, and the leaflets
of the involucra of Othonna Cheirifolia, present examples.
The pieces of integuments with valvate aestivation are
usually remarkable on account of their margins being
thick, indurated, sometimes slightly glutinous or velvety
in their infancy ; circumstances which contribute towards
retaining them in this position.

2d. These same parts may be disposed in a perfect
circle, but each with its margin folded inwards ; they
appear valvate externally, but when we open the bud,
we see the fold of each piece. This constitutes In-
duplicate Estivation (PI. 19, fig. 6) ; it has
much affinity with the preceding, as it is sometimes
found in plants very near, as regards their form, to
those with valvate aestivation, such as Clematis Viti-
cella: the portion folded inwards is usually thin and
membranous.

3d. We may, from analogy, admit a Reduplicate
^Estivation, which takes place when the pieces are

STRUCTURE OF FLOWERS. 107

folded or rolled backwards, as seems to happen in the
petals of some Umbelliferae.

4th. The parts of a verticil may be disposed in an
exact circle as to their position, but each of them
slightly twisted upon its axis, so that by one of its sides
it overlaps one of the neighbouring ones; and its other
side, being a little more interior, is overlapped by the
other one which is next to it. This disposition, which
is called Twisted or Contorted Estivation, (PI. 19,
figs. 2p, 4p, 5p,) is rare in envelopes which have their
pieces perfectly free ; it is seen in the petals and sepals
of the Flax, in the petals of the Pink and of the Mal-
vaceae ; but it is much more frequent in the free
portions or lobes of gamopetalous corollas, as in the
Apocyneae and Rubiaceas.

When the parts of a regular verticil are in two or
more rows, i. e. when the same verticil is double, &c, it
may also present several cases : —

1st. If the parts, placed exactly in the same direc-
tion as regards the axis, be alternate with one another,
there results the Alternate ^Estivation, (PI. 19, fig.
14,) where the pieces of the second row exactly alter-
nate with those of the first, and those of the third with
those of the second, &c. ; this is seen in the tepals of
the Liliaceae, the petals of the Nymphaeaceae, &c. Each
of these rows may itself present one of the preceding
dispositions ; but as the pieces are more distant, we are
rarely able to recognise it with precision.

2d. Under the name of Imbricate ^Estivation,
(PI. 19, figs. 9, 11,) are generally confounded all the
cases where the envelopes, being in several rows, have
no determined order, and where the pieces overlap each
other as the tiles on a roof. We see this in the involucra
of most Compositae, in the petals of most double flowers ;
but it is probable that we confound here, in the same

108 VEGETABLE ORGANOGRAPHY.

class, dispositions really distinct. When the pieces are
in two rows, and the outer one is very short com-
pared with the inner one, we designate this disposition by
the name of Calycular ^Estivation ; but this relates
to the proportion, and not to the position of the parts.

3d. It is possible that there exists in reality an Op-
posite ^Estivation, i. e. when each of the pieces of
one row arises exactly opposite that of the first row ; but
the examples which might be referred to this class are
obscure and uncertain ; such would be, for example, the
inner petals of Epimedium and Leontice, if we may
really consider them as distinct pieces of true petals.

The cases which I have enumerated seem to me to be
the only ones which exist in flowers strictly regular ; but
there are cases of slight irregularities, which it is usual
to class among aestivations. I ought, perhaps, to have
mentioned them in the preceding section, but I hope to
do it more clearly in reserving them for this.

"When the parts of the calyx or corolla are not
situated exactly in the same manner with regard to the
axis, there is an irregularity, and then one or some of
these parts have a tendency to overlap the others during
the prefloration ; it is this that a great number of
botanists designate collectively by the name of Imbricate
^Estivation, a term which, although become customary,
has the inconvenience of being taken here in a very
different sense from what is explained above ; and it
would be advantageously replaced by that of Irregular
^Estivation, if we wished to have a collective term.
It must be remarked, in fact, that this aestivation only
exists in irregular flowers, or in those having a tendency
to become so ; for it is a deviation from the symmetrical
order. We may distinguish several cases sufficiently
constant to deserve, perhaps, a special designation.

Thus, among 5-parted flowers, we often see the pieces

STRUCTURE OF FLOWERS. 109

of the calyx, corolla, or perigone, so disposed that there
are two exterior, one or two entirely interior, and one
or two intermediate, that is to say, half covered by one
side of one of the outer ones, and overlapping by the
other margin one of the inner ones ; this is very evident
in the calyx of the Rose, and I have called it Quin-
cunxial ^Estivation (PI. 19, figs. 10, 12).

The flowers of the Papilionaceae present one of their
petals more exterior, and embracing all the others, two
intermediate ones face to face, and two inner ones also
face to face ; this constitutes Vexillary Estivation,
(PI. 19, fig. 8.)

The variety of these irregular aestivations is very
great, being connected with the irregularity of the
flowers ; and in a great number of cases it may seem as
a sign whereby to discern flowers perfectly regular, or
more or less irregular. We may see different examples
in PI. 19, figs. 7, 9, &c.

Before quitting this subject, I ought also to remark,
that the aestivation of the parts of the calyx and
corolla have not any necessary affinity, even in the more
regular families : thus, the aestivation of the Malvaceae
(PI. 19. figs. 2, 7) is valvate in the calyx, twisted in the
corolla; that of the Linaceae and Cistineae (PI. 19, fig. 5)
is twisted in both organs, but the torsion of the corolla
is in a different direction from that of the calyx. This
fact, with a multitude of others, tends to prove, that it
is contrary to the nature of things to consider the calyx
and corolla as two rows of one organ which is called
perianth ; but that they are, in reality, organs as
different as all those of which the flowers are composed.
There are perigones of which the parts are in two rows,
and each row has a particular aestivation ; such is the
flower of Tradescantia Virginica (PI. 19, fig. 3), where
the outer row is foliaceous and valvate, and the inner

110 VEGETABLE ORGANOGRAPHY.

petaloid and rumpled : this seems to confirm the opinion
of Desvaux, who considered the outer row as the calyx,
and the inner the corolla. But, besides the reasons
above mentioned against this opinion, it must be added,
that the rumpled aestivation results only from an extra-
ordinary development of the organs. Thus, the petals
of the Poppy (PL 19, fig. 1), which are rumpled in
aestivation when examined collectively, appear evidently
alternate when examined separately, especially in double
flowers where their number diminishes the rumpling.

The relative position of the stamens with regard to
each other has less apparent influence upon the structure
of the flower, seeing that the form of these organs
always causes them to have sufficient space to develop
in without overlapping ; they only present, in this
respect, differences in the number of concentric rows,
the proportion of their size, the number of each row,
and the degree of their cohesion, of which I have else-
where spoken. I have already treated sufficiently upon
the position of the carpels.

The direction of the organs forms also part of the
history of their prefloration. Most of them spring
erect, as in all the examples of aestivation which I have
mentioned ; but there are some folded or rolled
inwards in a remarkable manner; thus, the calyces of
the Valerians and Centhranthus have the limb rolled
inwards upon itself, so as only to present, at the period
of flowering, a kind of ring which is developed at the
fall of the corolla: this is Involute ^Estivation.
The stamens of the Melastomaceae have their filaments
folded upon themselves, so that the anthers are pendent
in the interior of the bud ; this is an example of Re-
plicate ^Estivation. The carpels of Spircea ulmaria,
and still more those of Helicteres, are twisted spirally
upon one another, in the manner of twisted aestivations;

STRUCTURE OF FLOWERS. Ill

but which must be considered as a Spiral /Estivation,
seeing that their margins do not overlap. The bundle of
stamens oilnga Zygia presents also an extremely distinct
and extraordinary spiral torsion. Several styles, espe-
cially in the LeguminosEe, are rolled crossways upon
themselves, or spirally upon the same plane, so as to
resemble the disposition of circinnate leaves, and to
deserve the name of Circinnate ^Estivation; for
example, the style of Sabinma.

Section XIII.
Of Flowers united together.

Among the causes which tend to conceal the true
symmetry of flowers, there is one which, although very
accidental, deserves to be mentioned ; I speak of the
union of neighbouring flowers.

Sometimes two neighbouring peduncles are united so
intimately, that they appear to make but one, terminated
by two flowers : this happens naturally in the section of
the Honey-suckles which have two-flowered peduncles ;
it happens accidentally in several trees, such as the
Cherry, Apple, (PI. 21, figs. 1, 2,) &c, and in Centaurea
(PI. 21, fig. 3).

Not only can the pedicels be united, as in the pre-
ceding case, but two or more neighbouring ones may be
so as to form but one, which then presents more or less
evident traces of this union. I have remarked this
phenomenon very clearly in certain plants of Galeopsis,
where the summit of the stem was abortive, and where
was found a terminal flower formed by the union of two

112 VEGETABLE ORGANOGRAPHY.

neighbouring ones : this flower is larger than ordinarily,
and almost regular; its calyx, corolla, and stamens
present every number, from the natural one to double
that number.

An analogous phenomenon appears to take place in
some Tomatoes {Ly coper sicum). Dunal has shown in
detail, that the singular appearance of these ovaries,
and the multiplication of their cells, so contrary to
the ordinary state of the Solaneas, results from the
flowers being formed by the union of several.

There are some plants, where the union of the
flowers only takes place by the calyces, which, in this
case, are themselves adherent with the ovaries and
bracts : tins happens in Gundelia and Opercularia, and
changes these capitula, composed of several flowers, into
a mass, where during flowering we perceive the corollas
distinct, but where we find apparently only a multi-
locular fruit resulting from the union of all the partial
ones. We shall revert to this subject on speaking of
Fruits.

Section XIV.

Of the absolute Number of the Parts of each Floral

Verticil.

We have seen that flowers are composed of pieces
disposed in several concentric verticils, and that (with
some exceptions) the pieces of each verticil alternate
with those of the preceding ; whence it results, that
if we omit irregularities from partial abortions, the

STRUCTURE OF FLOWERS. 113

absolute number of organs of the same name is usually
determined by the number of similar verticils which are
developed. Thus, when there are two rows of stamens,
their number is double that of the petals ; when three
rows, triple ; and so on. A second cause of variation in
the relative number, which I have already mentioned
elsewhere, is, that sometimes, in the place which it
would seem ought to have been occupied by only one
stamen, a bundle of them is developed ; but even in this
case, the number is a multiple of the petals or sepals.
Lastly, plants frequently present a more remarkable, and,
if I may so say, a more intimate kind of numerical
aberration. It is not rare to find upon the same plant of
the Rue, flowers with four sepals, four petals, eight
stamens, and four united carpels ; whilst others have five
sepals, five petals, ten stamens, and ten carpels. We
remark in this case, and in all analogous ones, that the
flowers of the centre of cymes, which are developed
first, are five-parted, and the following four-parted, and
Linnaeus has established as a rule in his system, founded
upon the number of parts, that it is always by the first
developed flowers that the number ought to be fixed.
The examples of this kind of aberration, which affects at
the same time all the verticils without deranging the
symmetry, are repeated so frequently, that Linnaeus used
to express them by this phrase, " Quinta seu quartapars
fructificationis interdum additur." We find facts of this
kind in the Syringas, which have their flowers sometimes
on a quaternary plan, sometimes on a quinary one ; in
Asperula, the flowers are sometimes three-parted, some-
times four-parted, &c. This phenomenon is entirely
analogous to that which we have observed when speaking
of the verticils of leaves, which are so liable to vary in
number ; we should say that a branch, whether furnished
with verticillate leaves, or with verticillate floral parts,

VOL. II. i

114 VEGETABLE ORGANOGRAPHY.

is, as it were, composed of several fragments united lon-
gitudinally, and that symmetry always exists even when
one of these fragments is wanting. A monstrosity of
Iris Chinenis seems to support this theory ; we know
that the flower of this plant is formed upon the ternary
plan, i.e. that it is composed, 1st, of two verticils of
three leaves, transformed into lobes of the perigone, and
united at their base with the ovary ; 2d, of a verticil of
three stamens ; 3d, of a verticil of three carpels, united
together, and also to the perigone. But in the example
to which I allude, the flower is only composed of two-
thirds of these organs, the perigone is in two rows of
two leaves, and it has but two stamens and two carpels ;
but the other third, thus to speak, remains behind, half
developed, and we find the rudiments of it very visible
below the flower.

Does not this, which we have clearly seen in this case,
because of the abortion not being complete, evidently
exist in the cases where the abortion is more complete
and regular ; as, for example, when the flowers arranged
in a quinary manner in the Rue, Syringa, &c, pass to the
quaternary plan ? Is it not also to the same cause that
we must refer the cases where flowers, belonging by
their analogies to a certain class, have a less number of
organs than they ought to ? Thus, for example, all the
Asparagi are upon the ternary plan ; and if Mayanthemum
appears organized upon a binary one, it is probable that
a third of its organs are constantly abortive, as we have
seen to take place accidentally in the Iris. If several
Myrtaceae, Rubiaceae, &c, present a quaternary plan,
whilst others have a quinary one, is it not that a fifth of
their organs is abortive ?

We may consider that the two great classes of plants
have their floral verticils composed of a determined
number of pieces — Monocotyledons three, Dicotyledons

STRUCTURE OF FLOWERS. 115

five. The great majority of facts corresponds to this rule,
and I have little doubt that the exceptions will come to
be arranged in proportion as we know more of the true
symmetry of plants, and the great action of abortions.
We have already seen that several of these exceptions
are explained : —

1st. By the system of abortions of which I have
spoken.

2d. By the union of several partial organs : thus,
for example, if the flower of the Graminea? appears to
present a spathe with two valves, it is most probably
because the inner valve is formed by the union of two.

3d. The exceptions, with an excess of parts, may be
explained by the union of neighbouring flowers : thus,
the flowers of Paris may, with sufficient truth, be con-
sidered analogous to those of Trillium, but united in
pairs. We remark, in fact, that Paris quadrifolia pre-
sents the parts of each verticil in every intermediate
number between three and six ; and P. polyphylla, which
presents a still greater number, may result from the
union of three or four ternary flowers.

Section XV.

Of Nectaries.

There are few terms which have been so much abused
as that of Nectary. In its strict sense it means every
excretory gland situated upon one of the floral organs,
and the juice which it secretes bears the name of Nectar.
Linnaeus made use of this term to designate every kind

i2

116 VEGETABLE ORGANOGRAPHY.

of gland, tubercle, or appendage, which, placed in the
flower, did not seem to him to be an integrant part of
one of the ordinary floral organs ; since then, Botanists,
perceiving the very great difference of objects united
under this common designation, have endeavoured to
class them separately, and have given them particular
names, often more than was necessary. I shall here
examine the nectaries in a general manner, first, with
regard to themselves, and afterwards as concerns their
connexion with the organs which bear them.

The excretory glands which are observed upon
flowers, deserve a common name, principally because,
whatever be their position, or the peculiar nature of the
juices of each plant, or the form, size, and texture of
these glands, they all secrete a more or less honey-like
juice, which presents a very similar nature in all known
plants, — a remarkable circumstance, which sufficiently
proves an analogy of structure in all the glands which
produce nectar.

The nectaries, in regular flowers, may be found placed
on all the organs, but in a symmetrical manner. Their
most usual place is upon the torus. Sometimes they form
there distinct tubercles, the number of which is con-
nected with the parts of the flower ; for example, in
Parnassia, the Crassulaceas, &c. : they are situated upon
the opposite sides of the flower in the Cruciferas : some-
times the whole surface of the torus seems transformed
into a glandular and nectariferous surface ; for example,
in Cobcea.

Sometimes they are placed symmetrically on the
ovary ; such are the three glands in the Hyacinth.
Elsewhere, the parts of the corolla, calyx, or perigone,
bear nectariferous glands, either on their inner surface,
as those at the base of the tepals of Fritillaria impe-
rialis, or on their outer surface, as in the calyx of the

STRUCTURE OF FLOWERS. 117

Malpighiaceae. The stamens also frequently bear nec-
tariferous glands, particularly on the anthers or connec-
tivum, as in Adenanthera, Prosopis, &c.

In all these examples the symmetry of the flower is
not in the least altered, because the nectaries are placed
regularly ; but it very frequently happens, that we find
in irregular flowers the nectaries so placed as to have no
connexion with the symmetry. Is it the presence of
these irregularly placed nectaries which causes the irre-
gularity of the flower, or is it the irregularity of the
flower which causes that of the nectaries ? It is pro-
bable that these two causes are each true in certain
cases, but we can only observe the concordance of the
facts without determining which is the cause of the
other. Thus, in a great number of irregular gamope-
talous corollas, such as the Labiatae and Scrophula-
rinege, we find upon the torus a nectariferous gland,
placed upon one side of the ovary, but wanting on the
other.

It frequently happens that when one sexual organ is
abortive, its place is occupied by a nectariferous gland.
Thus, in the Scrophularineas, the place of the abortive
stamen is often occupied by a gland ; in several mo-
noecious or dioecious plants the pistil is replaced in the
male flowers by a nectariferous gland.

The nectaries upon the inner surface of the corolla
are always superficial, and they often cause a cavity
there, which, seen from the exterior, forms a kind of
knob or spur. It is in this sense that Sprengel has given
these organs the name Nectarotheca ; thus, the base
of the spur of Linaria, the violet, &c, always presents
a nectary more or less developed ; and when these
flowers become regular, each of their spurs contains a
nectary.

The genus Parnassia has very remarkable nectaries.

118 VEGETABLE ORGANOGRAPHY.

i

There is raised from the torus, between each of the five
stamens, a cylindrical filament, with three, five, seven,
or nine branches, according to the species, and each
branch is terminated by a globular nectariferous gland :
is this a simple form of nectary, or an indication of a
bundle of abortive stamens ? It is impossible to affirm
any thing upon this subject.

The nectar secreted is sought after with avidity by
bees and most sucking insects, which feed upon it. In
endeavouring to reach it, it frequently happens that they
excite or shake the stamens, and cause or accelerate
fecundation ; it may also happen that these insects on
coming out of a male flower charged with pollen, bear
it either upon the female ones of the same species, which
they fecundate, or upon the flowers of analogous ones,
by which they cause cross fecundations.

Section XVI.

Comparison of Foliaceous and Petaloid Parts.

We have seen, in describing the structure of each of
the floral organs, that the one kind resemble true leaves
in their intimate structure, their green colour, the pre-
sence of stomata, and the faculty of exhaling oxygen ;
such are the bracts, sepals, and most ovaries : the others
are of a more delicate tissue, adorned with various
colours, devoid of stomata, and incapable of exhaling
oxygen ; such are the petals, stamens, styles, and some
ovaries. It is necessary now to examine to what point
these limits are distinct. Let us first commence with the

STRUCTURE OF FLOWERS. 1 19

cases where the usually foliaceous organs are found in a
petaloid state.

The sepals frequently take the colouring and texture
of the petals. In this case, when the two organs coexist,
there is no doubt upon their distinction ; thus, the calyx
of a cultivated variety of the Primrose, Primula Calyc-
anthema, is expanded; into a coloured and petaloid
limb, so that the flower seems to have two corollas.
One of the lobes of the calyx of Mussoenda and Pink-
neya is dilated into a petaloid limb, whilst the others
retain their ordinary dimensions and appearance. In
several genera of theLeguminosse, Labiatae, Verbenaceae,
&c. the calyx is more or less coloured, without our being
likely to confound it with the corolla. But, if at the
same time that the calyx is coloured, the petals happen
to be wanting, or to take an unusual form, this calyx is
frequently taken for a corolla. This occurs in Anemone
and Clematis, where the petals are absent ; in Aquilegia
and Delphinium, where they exist, but deformed and
rudimentary. In all these cases, the calyx, although
coloured, is a true calyx, and it may be recognised
either by analogy with neighbouring genera where the
two organs exist, or by the study of double flowers.

It sometimes happens that the bracts themselves,
although more distant from the petals, participate in the
same tendency, and are coloured wholly or in part, acci-
dentally or constantly ; but this phenomenon never
happens but when the calyx is coloured. Thus we find
now and then individuals of the Anemone, where the
involucrum is partly foliaceous and parti}' coloured : the
bracts of several Liliaceae, Leguminosae, &c, and the
involucra of several Umbelliferae present the same fact
more or less constantly. The bracts of Salvia splendens,
Monarda, and several other Labiatae, are ornamented
with the most beautiful colours. The involucrum of

120 VEGETABLE ORGANOGRAPHY.

Comus florida is so large, so coloured, and enjoys so
much the function of petals, that it has given this pretty
under-shrub its specific name; the bracts of Hortensia
are so coloured and so near the flower, that there are
few beginners who do not take them for true petals.

If we are surprised at seeing bracts or sepals taking a
petaloid appearance, and if we wished to deduce from it
that these organs were not originally foliaceous, we should
be quickly undeceived, both by the great number of ana-
logous organs which present the appearance of leaves,
and by the examples of leaves which take petaloid
colours. Thus, several species, such as Atriplex hor-
tensis, are indifferently wholly green, or entirely red;
others, such as different species of Amaranthus, and
especially A. tricolor, take under different circumstances,
or in different places of the same plant, very decided
yellow or red tints ; others, such as Caladium bicolor,
have the centre of the leaf constantly marked with a
large rose-coloured spot, as brilliant as the petals. There
are some, such as Tradescantia discolor, Begonia discolor,
in which one of the surfaces of the leaf is of the most
beautiful red colour ; in other cases the leaves are
marked here and there with red spots in some species of
Caladium, with white ones in Begonia argyrostigma,
and with black ones in Arum vulgare. Let us also ob-
serve, that towards the end of their life the leaves of a
great number of trees take red or yellow tints, which are
usually connected with the colour which the fleshy fruits
of the same trees take when ripe.

All these examples, which could be easily multiplied,
and where we see parts naturally green become coloured,
tend to prove that this difference is far from being as
essential as might be supposed. If chemists should
show that the resinous colouring matter, or the Chro-
mule, is not perceptibly different when green (and then

STRUCTURE OF FLOWERS. 121

it is called Chlorophylle) or otherwise coloured,* we
should easily understand that very slight physiological
modifications can cause changes of colour, and conse-
quently that petals were only simple degenerations of
foliaceous organs.

What we have said of the leaflets, calyx, and involu-
crum, may as justly be said of the carpels, for the ovaries
are sometimes in a foliaceous state, sometimes coloured,
without there being any other essential difference in
their organization ; thus, very nearly related plants, as
Ornithogalum and S cilia, have the ovary green and of a
foliaceous appearance in the one, and coloured and
petaloid in the other. There are few families where
we do not find the same disparity between analogous
genera.

But if the bracts are leaves, which no one has con-
tested ; if the sepals are leaves, as can hardly be doubted ;
if the carpels are leaves, as seems to me to have been
demonstrated above ; if all these organs, although of
foliaceous origin, be however more or less susceptible of
being coloured and becoming petaloid; how can the
petals themselves be different? Why cannot they be
leaves more constantly metamorphosed than the others ?
This supposition will presently acquire more force when
we pursue our researches in the opposite direction, i. e.
when we have examined if the organs usually petaloid,
can present themselves in a foliaceous state.

I may mention, as an example of this, the carpels
changed into leaves, which are observed in Lathyrus
latifolius, in a variety of the Cherry, &c. which I have
already mentioned. But these examples may be doubt-
ful, because the carpels, in the ordinary state, are almost

• Since writing the above, this supposition seems verified by the expe-
riments of Macaire, from which it appears to result that the coloure d
chromule only differs from the green by being more oxygenized.

122 VEGETABLE ORGANOGK APHY.

foliaceous. The examples of petals changed into leaves,
though more rare, are more demonstrative. The follow-
ing are some cases : —

1st. In gardens there is cultivated a monstrosity of
the Gilliflower (Hesperis matronalis), the flowers of
which are replaced by a multitude of foliaceous organs
which are in an intermediate state between petals and
leaves, so that we can only consider these flowers as
double ones with semifoliaceous petals. Several double
varieties of Anemone, Ranunculus, &c. present this phe-
nomenon ; and I have observed flowers of the Fraxinella
{Dictamnus albus) of which all the floral organs, increased
in number as in the preceding case, have taken the ap-
pearance of leaves.

2d. Simple flowers also present this phenomenon,
although more seldom : I have found in the salt marshes
between Dieuze and Moyenvic a monstrosity of Ranun-
culus Philonotis, in which the petals were green, and pro-
vided with stomata like leaves, whilst the rest of the
flower was in the natural state.

3d. Dumas and Keeper have both found a monstro-
sity of Campanula rapunculoides, which is of great im-
portance in the study of the structure of flowers. This
Campanula presents sometimes upon the same plant
flowers in the ordinary state, others where the petals
are transformed into leaves, and others also where the
petals and stamens, and even the carpels, are changed
into leaves.

I have observed a very analogous fact in Anemone
nemorosa. The flowers were disfigured by the transforma-
tion into leaves of most of their organs, but the anthers
still remaining here and there, clearly showed the primi-
tive character of these organs. M. Bridel has observed
a very similar fact in Erysimum officinale, where most
of the floral parts were transformed into leaves. Cassini

STRUCTURE OF FLOWERS. 123

has given a description of Scabiosa columbaria, the fila-
ments of which were thick and herbaceous, and the
anthers were changed into a small green leaf, of which
the filament was the petiole. Therefore all the floral
organs are only verticils of leaves in a particular state.

We shall presently revert to this theory and its con-
sequences ; let us confine ourselves at present to the
observation, that the leaves which surround or form the
flower may present themselves in a foliaceous or in a
petaloid state, and that, although each of them has a
greater tendency to one of the states, it can, notwith-
standing, pass into the other from causes unknown to
us. These two states seem rather physiological pheno-
mena than truly anatomical differences. The foliaceous
state is that in which these organs serve for the nutri-
tion ; the petaloid has a more or less energetical tendency
to approach the sexual organs. Let us observe in con-
clusion that the state of verticils, of which the flower, or
even the inflorescence is composed, is generally modified
in regular succession. Thus, the bracts only become
petaloid when the calyces are likewise so ; the stamens
become foliaceous only when the petals have passed into
that state, &c.

Section XVII.

Of the particular Analogy between the Male and Female
Organs of Floivers.

The facts contained in the preceding section have
already proved that there is a great analogy between the
different parts of the flowers ; if we pursue this kind of

124 VEGETABLE ORGANOGRAPHY.

examination, and observe in particular the sexual trans-
formations, we shall always be more struck with this
singular similarity of the organs.

The male parts of plants, or the stamens, may some-
times, from causes unknown to us, be changed into
female organs or carpels, and bear ovules instead of
pollen. In the cases of this kind which have been ob-
served, the filaments are found in the natural state, and
the anthers are transformed into carpels ; more frequently
the stamens, if numerous, remain partly in the male
state, and the inner rows only are changed into female
organs. Sometimes we find stamens with the anther
half filled with ovules and half with pollen. The first
observation of this extraordinary metamorphosis was
made by Du Petit-Thouars on Sempervivum tectorum,
in which this accident appeared frequent, at least in the
north of France and in England. My attention having
been roused by this beantiful observation, I found a
short time after the same metamorphosis in the inner
rows of stamens in Magnolia fuscata, and I have since
frequently seen the male catkins of different species of
Willow, where some of the stamens were transformed
into carpels, and most frequently the two stamens of the
same flower changed into carpels, formed a fruit re-
sembling the ordinary kind. Richard has found a
similar transformation in Erica Tetralix ; Mr. Brown
in Cheiranthus Cheiri ; Du Petit-Thouars and Defrance
in the Poppy ; Guillemin in Euphorbia esula ; Seringe
in Cucurbita Pepo ; Rceper in Campanula Rapuncu-
loides, &c.

In some of the last examples which I have mentioned,
the phenomenon was presented in a particular manner,
in being complicated with a case of union. Thus, Mr.
Brown remarked that the stamens of Cheiranthus Cheiri,
changed into carpels, were united together around an

STRUCTURE OF FLOWERS. 125

ordinary pistil, in such a manner as to form a kind of
sheath, so as to present on a transverse section, besides
the two central cells, as many others in an outer row as
there were anthers converted into cai-pels. The same
fact was observed by Roeper in Campanula Rapuncu-
loides, the fruit of which was found to present two rows
of carpels. It is not improbable but the small number
of cases where fruits with two rows of seminiferous cells
have been described may be phenomena of this class.

Another change, more rare than the preceding, is
that where a carpel is changed into a stamen : Roeper
has observed it in Euphorbia palustris and Gentiana
campestris. In these cases one of the carpels seemed
wanting, and was found under the form of an anther.
Not having had an opportunity of seeing this phenome-
non myself, I cannot describe it in detail. It is likely
that in several dioecious plants we might find that
the central stamens of the male flowers are carpels meta-
morphosed. Perhaps, also, in some flowers which have
an inner row of incomplete stamens, and the number of
carpels less than in the normal state, it might be found
that these incomplete stamens are transformed carpels.
We might thus have new means of knowing the organic
symmetry of beings.

Section XVIII.

General Conclusions and Considerations upon the Struc-
ture of Flowers.

It results from all the preceding Sections, that a flower,
considered anatomically, is composed of several verticils

12G VEGETABLE ORGANOGRAPHY.

of floral leaves, placed symmetrically one above or within
the other, of which some, as the calyx and sometimes
the ovary, are of a foliaceous or nutritive nature, and
the others, as the petals and stamens, of a petaloid or
sexual nature ; or, if we consider them in another point
of view, of which some serve as protecting organs (calyx,
corolla) and the others as sexual organs (stamens, pistil).

Each verticil may be formed of several rows of the
same nature, whence it results that the total number of
rows may vary from one to several. Thus, we find one
in the female flower of Euphorbia, and in naked uni-
sexual flowers ; two in the male flower of Euphorbia, and
most unisexual Monochlamydeae ; three in Cneorum,
and almost all hermaphrodite Monochlamydeas ; four in
isostemenous Dicotyledons ; five in diplostemenous
Dicotyledons and in Monocotyledons considered as being
isostemenous ; six in Dicotyledons with three rows of
stamens, and in diplostemenous Monocotyledons, &c. &c.

On observing flowers in this respect, we see that there
exist calyces formed of one or two rows of sepals ;
we cannot affirm that there are any which have a great
number, because of the difficulty of distinguishing with
precision the outer rows of the calyx from those of the
bracts, properly so called.

There exist corollas with one, two, or more rows of
petals.

There are also stamens disposed in one, two, or more
rows. It is in the verticil that this number is most
variable.

There exist carpels in one row. This is almost univer-
sally the case when the rows are numerous ; they are
then disposed upon an axis which is a prolongation of
the pedicil ; they are arranged spirally, sometimes pro-
vided with special bracts at the base, and thus these
flowers approach the structure of capitate ones. This

STRUCTURE OF FLOWERS. 127

axis is capable of being prolonged into a branch with
leaves, and the same disposition is met with also in
flowers where the axis is but little if at all prolonged,
provided that the carpels are not arranged in a regular
verticil. Thus, flowers with verticillate carpels are true
terminations of branches ; those with spirally arranged
carpels may be terminations of branches, but are so only
by the exhaustion of the central parts, and when these
are well nourished, or when the sexual parts are abortive,
the branch may be prolonged by its apex.

The number of parts in each row of a verticil is fixed
in each plant, often in each family : it is most frequently
quinary in Dicotyledons, and ternary in Monocotyledons.
We may believe that the number of parts in the verticils
or rows of the same flower is naturally the same in each ;
but it appears very different in several cases : — 1st, Be-
cause the number of rows of verticils is different : thus,
frequently there is one row of petals, and two, three,
four, &c. of stamens, &c. 2d, Because there is an abor-
tion, union, or metamorphosis of some parts.

The parts of each verticil or of each row are capable
of being united together by cohesion in every possible
degree, and the degree of union causes what are called
the divisions of the parts.

The parts of each row of a verticil or of each uni-
serial verticil are generally placed alternately with those
of the preceding row. When the verticils are multi-
serial, each row is likewise alternate with the preceding
and following one. The verticils and their rows may be
unequal, and not resemble each other, without the flower
ceasing to be regular ; it becomes irregular when one of
the parts of a verticil or row is different from others of
the same row.

The parts of each verticil are capable of being united
by adhesion with those of the neighbouring one ; thus,

128 VEGETABLE ORGANOGRAPHY.

the petals may be united to the sepals and stamens ; the
stamens are frequently so to the petals, but very rarely
to the carpels. The sepals and carpels may be united
by the interposition of the torus, which is the common
base of the petals and stamens, which organs then are
adherent to the calyx, and appear to arise from near its
summit, or from the point where it begins to be free.

The parts of each verticil are capable of being changed
into true leaves, like those of the plant. This pheno-
menon is most frequent in the parts already more folia-
ceous, as the sepals and petals.

The parts of each verticil are capable of taking a
petaloid appearance ; this phenomenon is constant in the
petal, frequent in the stamens, and more seldom found
in the carpels (except in the stylary prolongation), sepals,
and bracts.

The parts of each row or verticil are capable of
changing to the nature of the row which immediately
touches it. Thus, we find the sepals changed to a petal-
oid nature (Primula calycanthema), petals changed to
stamens (Capsella Bursa pastoris), stamens to carpels
{Magnolia fuscata) ; or quite the reverse, viz. — carpels
changed to stamens [Euphorbia palustris), stamens to
petals (all double flowers), or petals changed to the nature
of the calyx {Ranunculus abortivus). Goethe has very
happily designated the first of these series of transfor-
mations, by the name of Ascending or Direct Meta-
morphosis, and the second by that of Descending or
Inverse Metamorphosis.

All the floral verticils then are of a very analogous
nature as regards their tissue, but they differ much in
their physiological state. Those which are foliaceous,
as the bracts and calyx, serve for the nutrition, the
others for the sexual reproduction. In several verticils
we may distinguish the parts of the leaves which com-

STRUCTURE OF FLOWERS. 129

pose them and find more or less clearly the trace of the
petiole and limb : the former being developed in the
outer organs which serve as the integuments, and the
latter in the interior and reproductive organs. Thus, in
the calyx, the sepals represent usually dilated petioles,
more or less fibrous or foliaceous : sometimes the limb is
visible, as in the Rose. In the corolla, the petals appear
generally formed by the petiole dilated into a petaloid
limb ; sometimes they present a claw, which acts the part
of a petiole, and a lamina, which acts that of a limb. In
the stamens we may also distinguish the filament which
represents the petiole, and the anther which is formed of
the two margins of the limb rolled upon themselves,
thus forming two cells. Lastly, in the carpels it hap-
pens most frequently that the petiole is wanting ; the
limb forms the carpel and the ovules arise from the ex-
tremity of the lateral nerves.

The petiole sometimes exists in the carpels (for exam-
ple, in Sterculia, Phaca, &c.) and then they are pedicel-
late; but when there are several carpels with the petioles
united, we must take care not to confound this support,
which seems an axis (as in Helleborus), with the central
axis, which is the prolongation of the stem (as in Myosu-
rus). I do not know any general rule for distinguishing
them. The extremity of the carpels is prolonged into
a style, which arises from the point where in a great
number of leaves we see a bristle or mucro, or a terminal
tendril, which is therefore the rudiment of this organ.

The important differences which are observed between
the ordinary or nutritive leaves, and those which com-
pose the flower, are : —

1st. That the ordinary leaves bear a bud in their axil,
and rarely have germs developed at the extremity of their
nerves (except in Bryoplujllum) ; those, on the contrary,
which form or surround the flower have no axillary

VOL. II. K

130 VEGETABLE ORGANOGRAPHY.

buds, but, on the contrary, at least when they are formed
by the limb and not by the petiole, they have lateral
germs capable of being developed into grains of fecunda-
ting pollen, or ovules susceptible of fecundation. Per-
haps the little bulbs which are developed in the axils of
certain floral organs are the representatives of the axil-
lary buds of ordinary leaves, just as the lateral germs of
Bryophyllum are in ordinary leaves the representatives
of the ovules of carpellary leaves. We, however, find
examples of floral leaves furnished with buds more or
less developed. Roeper has mentioned examples derived
from Euphorbia, and has shown this fact in E .Cyparissias.
Choisy has observed, in the Botanic Garden of Geneva,
a monstrosity of the Rose, where, in the place of the
stamens, upon the inner border of the torus, was deve-
loped a verticil of floral buds irregularly formed but
capable of being recognised as such ; we may add to
these facts the proliferous Marigold, Daisy, and Scabious,
where the axils of the bracts of the involucrum bear
pedicellate floral buds.

2d. Ordinary leaves are almost always opposite or
spiral, and those which form the flower are almost always
verticillate. Among ordinary leaves there are but very
few examples of real verticils (in Hippuris, and Myrio-
phyllum), for, in most verticils, there are only two oppo-
site leaves which bear buds in their axils, and the others
are, consequently, kinds of stipules. In the leaves of
the flower there are no examples of spires except in
carpels, disposed upon a real axis, and we have seen that
this structure indicates perhaps an aggregation of flowers
and not a single one ; let us also add here that, even
when the leaves of the stem are verticillate, the number
of each of its verticils has no more connexion with that
of the parts of the flower than the number of leaves of
each spire can have.

STRUCTURE OF FLOWERS. 131

3d. Ordinary leaves, when they are atrophied or
coloured, can take the appearance of petals, but they
always differ much, and we never see them produce any
thing analogous to the sexual organs. The leaves of the
flower, on the contrary, are, in their ordinary state, very
different from the preceding ; but, in certain cases, they
completely take their characters, except the existence of
axillary buds. Can the position of these organs explain
this difference? Is there any means of joining the ver-
ticillate position of the leaves of the flower, with the
frequently very different one of the ordinary leaves of
the same plant ? This last point would be of great
importance in this, — that it would completely connect
the history of the reproductive organs with that of the
organs of vegetation ; but the efforts made to attain this
object are as yet too hypothetical and incomplete to be
mentioned.

A curious example tends to confirm the extreme
analogy of the leaves with the floral parts. There is
cultivated in gardens a monstrosity of the White Lily,
in which, in the place of the flowers, the extremity of
each branch bears an indefinite number of leaves dis-
posed spirally, or imbricated as the ordinary ones, but
which are distinguished because they are coloured, and
entirely petaloid ; they differ then from the parts of the
flower only in not being verticillate.

From all the examples and the analogies which I have
pointed out, we may conclude, as the illustrious poet
Goethe maintained; as several botanists of the German
school, and especially Rceper, have admitted; as Turpin
has partly developed in his Iconographie ; as Mr. Robert
Brown appears to admit in different passages scattered
throughout his works; as I have myself partially indi-
cated in several of mine ; we may, I say, conclude that
the leaves, or appendicular organs of the stem, modified

k2

132 VEGETABLE ORGANOGRAPHY.

by their position, compose all the parts of flowers. A
flower, then, is a kind of rosette, or terminal bud, the
leaves of which are verticillate, and take a less degree of
nutritive development than ordinarily, but acquire, in
return, new forms and functions. When the force of
vegetation is very great, a larger number of leaves take
a foliaceous state, and the branches bear fewer flowers ;
when the vegetative force is diminished, the upper leaves
have a greater tendency to be transformed into floral
parts : this is a law to be observed, practically, by
gardeners.

The extreme ease with which, by this theory, we can
explain all the anomalies and monstrosities of flowers, i
a sure pledge of its truth. Astronomers only regarded
their science as well proved, when they were able to
explain by its means the apparent aberrations of the
stars.

We may say, in a very widely extended sense, that
there only exists three organs in plants — the root, the
stem, and the leaves, and that the different modifications,
which the summits of the stems and the appendicular or
foliaceous organs present, constitute all the apparatus of
the flowers and fruit.

But because the floral parts may be considered as
modified leaves, we can by no means conclude that, in
changing form, they cannot take new functions ; and it
seems to me, we ought not to attach to this idea any
argument against the theory of the sexes. Every analogy
tends to prove, on the contrary, that the modified organs
often serve very different purposes from their primitive
ones, and in particular the sexual fecundation appears
to me to be demonstrated in plants, almost to the same
degree as in animals.

STRUCTURE OF FRUIT. 13#

CHAPTER III.

OF THE STRUCTURE OF THE FRUIT OF PHANEROGAMOUS

PLANTS.

Section I.
Of the Fruit in general.

When fecundation has taken place, the organs which
were destined to effect it, perish with more or less
rapidity ; the stamens wither and fall off, in the greatest
number of cases ; the petals follow the same fate ; all the
sexual portion of the carpels undergoes the same change ;
the stigma and style wither and usually fall off; the
pistillary cord, or the fibres which go from the style to
the ovules, wither likewise, — they disappear either by an
evident destruction in the small number of cases where,
as in Lychnis dioica, they are free from all adhesion,
and, consequently, visible, or by simple obliteration in
the cases, by far the most numerous, where they are
imbedded in the tissue of the carpels.

Whilst the truly sexual organs disappear after they
have fulfilled their office, the fecundated ovules have
taken a life of their own ; they attract the nourishing
juices, and are developed with their immediate covering
— the foliaceous part of the carpels. The name of Fruit
(fructus) is given to the body which results from the
ovules transformed into seeds, by the fecundation of the

134 VEGETABLE ORGANOGRAPHY.

carpels which surround, contain, and nourish them, and
from all the parts of the flower which, adhering to the
carpels, seem more or less to form an integrant part of
all this apparatus. The fecundated ovules bear the name
of Seeds (seminaj, and their envelopes have collectively
received that of Pericarp (pericarpium), a term which
is not exactly applicable to an organ which is not around
the fruit, but forms an integrant part of it.

The study of the fruit, considered collectively, bears
the name of Carpology.

It is to Gcertner that we owe the first exact descrip-
tions of the fruits and seeds of plants, and Bernard and
A. L. de Jussieu were the first to make us understand
the importance of the carpological characters in the clas-
sification of plants; since then several distinguished
botanists have directed their attention to Carpology.

Two causes, however, have principally contributed to
render the study of the pericarp more difficult and con-
fused than it ought to be, viz. — 1st, that for a long time
it was only studied when it had attained maturity, and,
consequently, the true character of the parts of which it
was composed could not be judged of, seeing that several
of them are obliterated or developed, or united together
during maturation ; 2d, that it, as well as most of the
floral organs, has been considered as a single organ
divisible into distinct parts, whilst it is always more true
and advantageous to consider first the elementary organs
themselves, and afterwards the results of their different
aggregations. We do not say then, with a modern
botanist (Mirbel), " that the flower never has more than
one ovary, and that the little distinct bodies, fixed upon
the same receptacle, are only portions of a single peri-
carp ;" but we affirm, on the contrary, that the flower
usually has several carpels, which are sometimes sepa-
rate, sometimes united into a single body. In order to

STRUCTURE OF FRUIT. 135

avoid these causes of doubt, after having examined the
pericarp, as we have already done, in the state of an
ovary, and in its connexions with the floral parts,
properly so called, let us consider here, in their isolated
and simple state, the elementary parts of which the
pericarp is composed, and afterwards let us examine the
consequence of their union, first with each other, and
then with the neighbouring organs.

I suppose throughout all the following description of
the pericarp, that all that I have said in the preceding
chapter upon the flower, and particularly upon the
pistil, is recollected. I beg those who would read the
chapter upon the fruit, not to do so until they have
read the preceding one.

Section II.

Of the Carpels considered in the state of separation from

one another.

The Carpels, as we have said in speaking of the pistil,
are the female organs of plants, most frequently verticil-
late in the centre of the flower, and which, sometimes
free, sometimes united, form the pistil during flowering,
and afterwards the fruit.

Each carpel may be considered as a leaf folded longi-
tudinally upon itself; if we examine its texture, we
shall find that, like the leaf, it is composed of three
parts, which really constitute a single envelope, viz. —

1st. Its external surface, which represents the lower
face of the leaf, is a kind of cuticle separable in a great
number of fruits, such as the Peach, and existing in all.
Richard has given it the name of Epicarp (epicar-

136 VEGETABLE ORGANOGRAPHY.

pium) ; like the lower surface of the leaf, it frequently
bears hairs, glands, and stomata, and presents in most
cases a decided analogy to the part of the leaf to which
it corresponds.

2d. The inner surface, which represents the upper
part of the leaf, is a membrane to which Richard has
given the name of Endocahp ( Endocarpium) ; its inter-
nal position, preventing its exposure to the air and light,
causes it to differ more than the epicarp from the
corresponding part of the leaf. Sometimes it appears
under the form of a fine membrane, foliaceous and even
greenish, as in the Pea ; sometimes under that of a fine,
pale, and as it were, etiolated membrane, as in Asclepias;
this membrane sometimes becomes thicker, and some-
times even hard and bony, as in the Peach, and then it
forms the stone of the carpel. All the intermediate
states between those which I have mentioned are found
in various fruits.

3d. Between the Epicarp and Endocarp is found the
plexus of fibres, vessels, and cellular tissue, which con-
stitutes the body of the leaf or carpel, representing the
mesophyllum of the former ; it has received the name of
Mesocarp (mesocarphcm). Sometimes it is very thick
and fleshy, as in the Peach, or Cherry, and then the
name of Sarcocarp, or more commonly, that of the
Flesh (caro) of the fruit is given to it. Sometimes it
is thick, but dry and fibrous, and then it is called the
Husk, as in the Almond. Sometimes it happens that
the epicarp and endocarp separate during their growth,
and leave between them an empty space, in which are
still observed traces of the fibres of the mesocarp ; this
rare and singular organization is seen well in Cysti-
capnos Africana. Sometimes it is also distinct, but less
thick and more foliaceous, as in the Bean and Pea ; at
times it is so thin, that it cannot be easily distinguished

STRUCTURE OF FRUIT. 137

from the epicarp and endocarp ; but however thin it
may be, we are always compelled to admit its existence,
since it is only by this plexus of vessels that the organ
is nourished.

In the same manner as we have seen, in speaking of
leaves, that the more stomata there are, the thinner will
this plexus be ; so, among carpels we shall find three
particular states of the mesocarp connected with the
structure of the epicarp. Thus, sometimes the carpellary
leaf is membranous, and almost scarious, and then the
epicarp has no stomata, and the endocarp little or no
flesh, as in Amaranthus ; sometimes the epicarp has
stomata, and the endocarp has the ordinary foliaceous
texture, as in the Pea; sometimes the epicarp has no
stomata, and the endocarp becomes thick and fleshy, as
in succulent leaves ; this is what we see in the Peach
and other fleshy fruits.

Fruits compared together when ripe, present different
kinds of adhesion or separation of the mesocarp ; thus it
adheres sometimes very strongly to the endocarp, as in
the Nectarine, or Beau ; at other times it easily separates,
as in the Peach, or husk of the Walnut, or in Entada, in
which it is detached naturally or easily, remaining free
from the epicarp. The adhesion of the mesocarp with
the epicarp is equally variable ; thus the latter is easily
detached in the Peach, with difficulty in the Cherry.

The leaves which form the carpels are presented
under several different systems ; thus, they may be : —

1st. Curved in the form of a cylinder, tapering to the
two extremities, having the two margins beveled off
upon one another, and the back round, as in Colchicum,
Delphinium, Sterculia, &c.

2d. Curved in the form of a horn, so that the margins
approach each other at the base, and the upper part re-
mains open, as in Astrocarpus, Helleborus, Isopyrum, &c.

138 VEGETABLE ORGANOGRAPHY.

3d. Folded upon their middle nerve, so that the
margins are applied to each other, forming a longitu-
dinal line, as in the Pea, Bean, &c.

4th. As the leaves are curved or folded, their two
seminiferous margins may be more or less folded in-
wards, so as to divide the carpel by semi-partitions, or
by longitudinal ones, as in Astragalus.

5th. The dorsal nerve of the carpel or the middle one
of the leaf, may be so pushed inwards, as to form a pro-
jection which also tends to divide the carpel into two
longitudinal cells, as in Oxytropis.

These different modes of folding or curving of the leaf,
correspond, as it is easy to ascertain, with the modes
already described, of the rolling or folding of the leaves
in the bud. When the carpels are formed of a leaf,
folded lengthways upon itself, the two lateral surfaces
may be either flat, as in Spartium Junceum, or more or
less convex, as in the Bean, or Crotolaria, or folded at a
more or less obtuse angle, a combination which in
general takes place only by the pressure of neighbour-
ing organs.

The line formed by the approximation of the two
margins of the leaf, and which represents a suture, is
called the Seminiferous Suture, because it is upon its
inner edge that the seeds are usually attached, or the
Ventral Suture, because it is opposite the back of
the carpel.

When the carpel is formed of a leaf curved longitudi-
nally, it only presents this suture, and receives then the
particular name of Follicle, (folliculus.) It is called
a Cocca when the suture opens with elasticity.

When the carpel is formed by a leaf folded upon its
middle nerve, it frequently happens that, at maturity,
the dehiscence takes place along this nerve, which
has for this reason received the name of the Dorsal

STRUCTURE OF FRUIT. 139

Suture ; and this is given to it by analogy, even when it
does not open, provided that the nerve be well-marked :
but it must be remarked that this suture is the rupture
of an organ by a natural dehiscence, whilst the first is
disunion of two portions cohering together ; the carpels
formed by leaves folded lengthways having consequently
two sutures, bear the generic name of Legume, (legu-
wen). There are some families, such as the Ranuncu-
laceae, where the existence of the dorsal suture is so
slightly perceptible, that we see in neighbouring genera
carpels which may be referred, almost at will, to follicles
or legumes.

When the carpels are long, with their valves flat, or
nearly so, and their seeds situated at small distances,
two phenomena frequently happen, which modify their
ordinary state. Sometimes the portions of the carpel
which are found between the seeds, are united together
by a kind of natural union, or by a development of the
cellular tissue, which forms kinds of false partitions be-
tween the seeds: this constitutes the legumes which
are said to be multilocular, or to have transverse cells,
as, for example, in Clitoria. At other times, the por-
tion of the carpel situated between the seeds is de-
veloped less than that which is around them, and then
the legume presents here and there swellings and
contractions. These different states of legumes with
transverse cells or joints are expressed collectively, by
calling them Lomentaceous carpels or legumes. Some-
times the two phenomena occur at once, as in Sopliora.

The carpels sometimes have a thecaphore or support,
which is to the carpellary leaf what the petiole is to the
ordinary one. This Thecaphore is very visible in
Phaca, Glottidium, Colutea, and several other Legumi-
nosas. It must be observed that this pedicel of the fruit
is often spirally twisted, whence it results that the

I'M) VEGETABLE ORGANOGRAPHY.

carpel is then presented in a position contrary to the
natural one ; the seminiferous suture is found situated
externally, and the dorsal internally.

The point whence the style takes its origin, whatever
be its position, is considered as the anatomical apex of
the carpellary ovary. In the greatest number of cases, it
is situated at the apparent apex of the fruit ; thus, for
example, in the Pea or Larkspur, the [style proceeds
from the tip of the ovary ; but there are plants where
the ventral suture is very short and the dorsal very much
swollen out, whence results a lateral position of the
anatomical apex, for example in the genera Rubus,
Fragaria, Potentilla, Sec.

The style most frequently dries up or falls off after
fecundation ; but sometimes the whole of it, or at least
its base remains, either without changing its form, or
becoming elongated, or hardening so as to form at the
top of the ovary a more or less decided point : for
example, in Dry as and Clematis it forms a long bearded
process ; in several species of Geum, one without a
beard ; in Trigonella and some species of Ranunculus, a
nearly spiny process, &c. Sometimes the two placen-
tary styles are separate, and then result ovaries termi-
nated by two points or bifid carpellary styles, as, for
example, in certain kinds of Euphorbia.

There are great differences between the carpels,
resulting from what takes place in them at maturity.
Some do not open naturally, and are said to be In de-
hiscent ; the others, which are called Dehiscent, open
in various ways.

Indehiscent carpels are of two kinds : —

1st. Those of a dry, scarious, bony or membranous
nature, have very little juice, few or no stomata, and
only contain two ovules, one of which is abortive, most
frequently before maturity. In these carpels the seed is

STRUCTURE OF FRUIT. 141

often united to the pericarp, or this organ is so well
moulded upon it without adhering, that the two bodies
seem confounded ; it is this which has caused the term,
Naked Seeds, to be applied to them, a very incorrect
one, which might be replaced by that of Pseudosper-
mous fruit or carpels ; the sutures, even the ventral one,
are here scarcely perceptible, being often reduced to a
simple nerve or fold, sometimes hardly visible. At
maturity these carpels, detached from the peduncle, are
sown without opening ; the seed, which is within, ger-
minates without coming out, and, as it is usually soli-
tary, this is performed without difficulty. It is to this
class of fruits that the following belong, viz. —

The Utricle (utricuhis), where the carpel is mem-
branous and does not adhere to the seed ; as in Ama-
ranthus.

The Nut (nux), where the carpel is osseous or stony,
and does not adhere to the seed, as in the Cashew-nut.

The Caryopsis, where the carpel adheres intimately
to the seed, as in Wheat.

2d. There are other indehiscent carpels, which have
the mesocarp more or less developed and fleshy. In
plants with carpels solitary by abortion, or isolated from
one another, we only find them fleshy in those which
have naturally one or two seeds. Some are of a fleshy
nature, with the endocarp membranous ; such are, for
example, the little round carpels, which, by being
collected on a common axis, form the fruit of the
Bramble and Raspberry : others have the mesocarp
fleshy, with the endocarp bony ; to these the name of
Drupe (drupa) is given ; such are the fruit of Detarium
and Geoffrcea among the Leguminosae ; Plums, Cherries,
and Peaches among the Amygdalaceae : finally, there are
others which have the mesocarp fibrous and the endocarp
bony, as the fruit of the Almond. It must be observed,

142 VEGETABLE ORGANOGRAPHY.

that in these two last classes, the endocarp retains the
original form peculiar to legumes; for whilst the
epicarp and mesocarp are perfectly continuous and inde-
hiscent, the stone presents two sutures and two valves
like legumes, and has a tendency to open, at least at the
period of germination, or sometimes sooner. In all
fleshy carpels, the epicarp and mesocarp are destroyed
by putrefaction or maceration ; and the seed, invested
with the bony or membranous endocarp, is sown and
germinates as in pseudo-spermous fruits.

We must not confound with the flesh, which is a
watery or fleshy development of the mesocarp, the Pulp
(pulpa) of the fruit ; this is only found in the interior
of the carpel : thus, for example, the legume of Cassia
fistula is dry and encloses a true pulp : this pulp is not
an organ, properly so called, but is a secretion of the
endocarp, placenta, umbilical cord, or of the surface of
the seed. It is probable, that we confound under this
name, various matters secreted by different organs ;
thus, I am inclined to believe that the sour and acid
pulp of Sophora proceeds from an organ different from
that from which the sweet and laxative pulp of Cassia,
&c. does.

Dehiscent carpels may open in different manners. The
most simple case is that where the two margins of the
carpellary leaf separate at the point of union and open
lengthways, as in follicles. Sometimes this dehiscence
takes place throughout the whole length, as in Asclepias ;
sometimes only at the top, as in TroUius.

A second very frequent case is that where the de-
hiscence takes place by the ventral suture and by the
dorsal nerve or suture, as in the legumes of most of the
Leguminosas.

But it sometimes happens that the two sutures so
cohere together, that they cannot open. Thus, if the

STRUCTURE OF FRUIT. 143

legume contain several seeds, the dehiscence takes place
in one of the two following ways : — when the fruit is uni-
locular, each of the valves splits lengthways along the
middle, forming two longitudinal ruptures ; this takes
place in the legume of Hcematoxylon : or when the
fruit is divided into transverse cells, as I have above
explained, a rupture is made transversely along the
partition or contraction which separates the cells, and
each of them (which then take the name of Joints)
is found separated from the others, and transformed, as
it wrere, into a pseudo-spermous fruit, which does not
open, but is sown with the seed enclosed. And here also
are presented two cases : sometimes, as in the He-
dysareae, each joint carries with it the two sutures
perfectly entire; sometimes, as in several Mimoseae, the
sutures remain, and the cells open at the same time that
they separate from one another.

In all these cases, it evidently results from the
dehiscence that the seeds, when they are numerous in
the same carpel, are permitted to be dispersed so as to
be sown separately.

We have already said that the seeds are attached
either to the inner margin of the carpellary leaf, and,
consequently, along the ventral suture, or at the base or
apex of the carpel, but always laterally, and in reality
near the top or bottom of the ventral suture. These
three positions, then, differ only in this, that in the first
case they arise all along the suture, in the second only
at its base, in the third only at the apex. In every case,
the portion of the carpellary leaf from which the seeds
arise, has received the name of Placenta.

The placenta is usually a kind of thick ridge, com-
posed of spongy cellular tissue, and pierced by two sets
of fibres : the first, which proceeds from the peduncle,
conducts the nourishing fluids ; the other, which comes

144 VEGETABLE ORGANOGRAPHY.

from the style, has borne the fecundating liquor to the
ovules, and has generally disappeared when the fruit is
formed. These two orders of vessels are subdivided
into as many filaments as there are ovules, and a
filament of each kind penetrates the seed, which is
connected with the placenta by a cord of variable form
and length, called the Umbilical Cord, Funiculus,
or Podosperm ; this cord, at the time of flowering, was
composed of one filament coming from the pistillary
cord, and of another from the nourishing one ; the first
usually disappears very soon after fecundation. There
are very rare cases where the two filaments, the com-
bination of which composes the ordinary funiculus, are
found completely distinct: thus, in Statice, the nourishing
filament proceeds from the base of the ovary, the
fecundating one from its summit, and they both reach
the seed distinct.

The placenta is but little evident at the period of
flowering ; it afterwards enlarges and becomes full of
juice, which the seed absorbs during its growth ; whence
it results, that at maturity, the placenta is dry and
flaccid, resembling old pith. The larger it is in proportion
to the seeds, the more does it serve to nourish them :
thus, it is remarked that when fruits have a large
placenta, as in Coba>a, we may cut them long before
they are ripe, without endangering the maturation of
the seeds, which absorb from this resorvoir the juices
which are necessary for them ; whilst in pericarps where
it is small, the seeds cannot ripen after the fruit is
detached from the plant. When the placentae are dry,
it sometimes happens that, at maturity, they naturally
separate from the carpellary leaf from which they
originated.

When the placenta? are placed along the ventral
suture, they are very evidently double ; it is the same

STRUCTURE OF FRUIT. 145

also when they are reduced so as to occupy only the
base or apex; for these are still the same two organs,
but much shorter than ordinarily. Since the placentas
of a carpel are necessarily double, and each of them has,
in the regular state, an equal right to bear seeds, it
results: — 1st, that the natural number of seeds in a
carpel ought to be always even when there is no
abortion ; but they are rarely placed exactly at an
equal height along each suture ; on the contrary, they
are situated alternately : this disposition is very evident
in legumes and long follicles ; but when the carpel is so
short as only to present one ovule on each placenta,
there are presented some cases which deserve notice : —

1st. The two ovules may arise, as in long carpels, one
above the other, at so great a distance that both arrive
at maturity : this is what happens in the legumes of the
dispermous Leguminosas, and then the two seeds are
clearly horizontal.

2d. These two alternate ovules are sometimes so close,
that one of them becomes abortive, the other only
arriving at maturity ; in this case, it happens either that
it is the upper one which is abortive, and then the
lower one finding more space towards the higher part of
the carpel, takes an erect position, or it is the lower
one which is abortive, and then the superior one finding
more space towards the base of the carpel, takes a
pendent position. It is very probable that it is to this
double cause that we must refer the diversity of direction
of the seed of the monospermous Ranunculaceae, which
are either ascending or pendent, whilst those of the
polyspermous ones are horizontal ; if one were to find a
Ranunculus or Clematis, the carpel of which presented
the two ovules arrived at maturity, they would be either
horizontal, or one ascending and the other pendent.

3d. The two ovules may be very near together either

VOL. II. L

146 VEGETABLE ORGANOGRAPHY.

at the base or apex of the fruit ; in this case one is very
frequently abortive, and in this manner the monosper-
mous carpels of the Compositae and Dipsaceae are formed.
But when we find, now and then, one of the Composites
with the fruit having two ovules, we see that they are
both ascending, and it is very likely that, if we should
find two in the Dipsaceae, they would be both pendent.

Whenever a carpel contains several seeds, they are
free and not united to the inner surface of it ; but when
it only contains a single seed, this one is sometimes free,
as in the utricles of the Amaranthaceas, or united by its
entire surface with the carpellary leaf, as in the fruit of
the Gramineae ; and then the carpel is so confounded
with the proper integument of the seed, that it does not
seem to exist ; it is in this case that seeds were called
naked, but there are none really devoid of the pericarp.
In fact, the style necessarily takes its origin from the
pericarp and not from the seed, and consequently, every
organ from which we see, at the period of flowering, a
style or stigma arise, is a true pericarp, whatever be its
appearance.

Seeds may appear naked from three causes : — either
by the intimate union of the seed with the carpel, as in
the Gramineae : — or because, as in certain species of
Leontice, or in Slateria, the seed, growing rapidly,
breaks the carpellary leaf and is found exposed : — or
because, as in Reseda, the carpellary leaves not being
completely folded upon themselves, leave their ex-
tremity open, and consequently the seeds naked. But
we know that neither of these causes answers exactly to
what was intended by the term of naked seeds, and that
the pericarp exists or always has existed.

The manner in which the seeds of a carpel ripen and
are dispersed, accords with the principles above declared.
In all carpels with a long placenta, i. e. when the seeds

STRUCTURE OF FRUIT. 147

are situated all along the ventral suture, the seeds
receive the fecundating matter by the branches of the
pistillary cord, which terminate there ; those at the top
receive it before the others, and their vital action imme-
diately commences; consequently, they ought to ripen
first : we see this in all polyspermous legumes and folli-
cles ; and as it is also at the top that the dehiscence of
the sutures commences, it follows that the seeds fall out
according as they ripen.

As soon as the seeds have fallen out, or a little after,
the Valves of the legumes (this name is given to the
two portions of the carpellary leaf separated by the de-
hiscence of the sutures), the valves, I say, twist, either
rolling up towards the outside or spirally upon them-
selves, or by irregular torsions ; they sometimes retain,
on separating, their primitive position.

The carpels frequently bear foliaceous or spiny tufts,
or tubercles, sometimes upon both margins of the
suture, sometimes upon their sides or valves ; these
peculiarities, which are sometimes of interest in order
to know such or such a fruit, are generally of but slight
importance in Carpology.

All that we have said in this section is applicable —
1st. To carpels which are naturally isolated from each
other in the same flower, and which constitute what is
called a Compound fruit : such are the two follicles of
the Apocyneae, the verticillate carpels of Alisma and
Delphinium, the carpels collected into a head or spike in
Ranunculus. On combining what is contained in this
Section and in the preceding Chapter, it seems to me
that we have their complete history presented to us.

2d. To carpels, which, originally resembling the pre-
ceding, have become solitary by the abortion of those
which ought, in the normal plan of the flower, to form a
complete verticil, as the solitary legume of most of the

l2

148 VEGETABLE ORGANOGRAPHY.

Leguminosne, the solitary carpels of the Larkspur.
This solitariness of the carpel, produced by the abortion
of the neighbouring ones, is perceived by the lateral
position of the seeds ; this has caused them to be con-
founded with fruits formed by the natural union of
several carpels. To these has been given, very impro-
perly, and solely on account of their external appear-
ance, the name of Simple fruits, although they are more
complicated, and this name is much more applicable to
those of which we are about to treat. Let us now
examine the results of the natural union of the carpels
of the same flower.

Section III.
Of the Carpels of the same Flower united together.

The carpels proceeding from the same flower may be
united together at two different periods : —

1st. There are some which are originally perfectly
free and distinct, but so near to each other, that, if they
become fleshy, they can unite together, on approaching
maturity, into a single body usually slightly irregular.
This late union of numerous and fleshy carpels is very
well seen in Dillenia and Anona; there results from
this aggregation a fruit marked with areolae, which are
the traces of the tops of the carpels ; the seeds appear
irregularly distributed in the mass, because the walls of
the carpel being fleshy and united, we cannot recognise
their primitive disposition.

2d. In a great number of flowers the carpels are
naturally united together from the first; this circum-

STRUCTURE OF FRUIT. 149

stance so modifies their form and appearance, that it is
necessary to describe it in detail. That fruits with
several cells placed horizontally are composed of carpels
united together, is what, I think, cannot appear doubt-
ful to any one who has attentively studied the article
upon the Pistil in the preceding Chapter. Some exam-
ples seem to render this fact more particularly evident ;
thus, among the Helleboreae, we find some with the
carpels perfectly free, such as Aconitum, whilst in certain
genera, as Nigella, we find species where the carpels are
united at the base only, as iV. orientalis : others, where
the union proceeds half-way, as N. saliva, and others
where they are united nearly to the apex, as in
N. Damasccena. It is the same in the Apocyneas and
Asclepiadeae, in which we find all the gradations, from
the perfectly free carpels of Asclepias, to those of Cer-
bera, Rauwolfia, &c. which are united into an apparently
simple fruit. Similar examples are found in a great
number of families.

We have already found, in these facts, a very simple
explanation of what was, or ought to be meant in
speaking of entire, divided, parted, and compound fruits.
Entire ones are those where the carpels are united
throughout their whole length ; divided ones, where the
union only proceeds half-way ; parted ones are those
where the carpels are only united at the base ; com-
pound ones, where the carpels are free from all cohesion.

In fruits with the carpels united throughout their
whole length, there may also be several cases : the
carpels either have the ventral suture prolonged more in
proportion than the dorsal, and then the whole fruit is
more or less pointed at the base, or vice versa, and then
the fruit is necessarily lobed at the summit: or, lastly,
the two sutures are perceptibly equal, and then the
fruit is obtuse or truncated. Thus, all the longitudinal

150 VEGETABLE ORGANOGRAPHY.

divisions observed in fruits are easily understood to
result from the union of the carpels.

When the carpels are verticillate, their approaching a
central axis (which is called the Columella when it is
real, and the Axis properly so called, if it be ideal,)
causes them to take a triangular form; their union
takes place by the two sides sloping inwards, and the
dorsal face of all the carpels forms the external part of
the fruit, which results from their union. When this
dorsal face is uniformly convex, the fruit is round, as in
Rhizophora ; elliptical, as in the Coffee ; or globular as
in the Grape, the Elder, &c. When the dorsal face is
more convex than the fruit taken collectively, the fruit
then presents as many furrows as there are sutures, or
points of union of the carpels, and as many round pro-
jecting sides as there are convex carpels, as for example,
in the Melon, Ricinus, &c. If the back of the carpel
be angular, or if the carpel be, as it were, folded upon
the middle nerve, the fruit then presents as many angular
sides as there are carpels ; the retreating angles indicate
the sutures, and the projecting ones the backs of the
carpels, as in Hibiscus esculenlus, Oxalis, &c. ; these
projecting angles are sometimes even prolonged into
wings, as in Dodoncea. Thus all the lateral depressions
or eminences which are observed on the surfaces of
fruits, are also easily understood in the theory of the
union of the carpels, and depend upon their elementary
forms. These forms are sometimes obscured by the
unusual development of the fleshy part of the mesocarp.

The most usual structure of the carpels is when their
two retreating faces reach into the interior of the fruit,
as far as the axis, and then the fruit presents as many
Cells (loculi) as there are carpels entering into its
formation ; we then say that it is Multilocular, or
Bi-, Tri-, Quadri-, Quinque- &c. locular, when we

r

STRUCTURE OF FRUIT. 151

wish to designate the number. These cells are sepa-
rated by vertical Partitions (septa), formed by the
more or less intimate union of the retreating faces of the
two contiguous carpels. These retreating faces appear
composed only of the endocarp and a very thin expan-
sion of the mesocarp ; as for the epicarp, it is not pro-
longed, or at least it is not visible upon the partitions.
The seeds are then placed at the central angle of each
cell, attached to the extremity of the retreating face of
the carpel, and consequently, (except in cases of abor-
tion,) to the number of two at least in each cell, or
always in equal numbers. All that I have said above of
their position in isolated carpels, is applicable to the
cells of fruits with cohering ones.

When the carpels of which the fruit is composed are
lomentaceous, or divided by diaphragms, as transverse
partitions, then each principal cell is subdivided bv
these cellular diaphragms into smaller ones situated one
above the other ; we see this in Amaioua among the
Rubiacea?, in the lomentaceous Cruciferae, Sec.

We reserve the name of False Cells, or of Cavities,
for certain spaces which are found in some fruits, and
which do not contain seeds, not from abortion, but from
their peculiar nature. The most remarkable example of
these cavities is observed in Nigella Damascccna, so
common in gardens ; its fruit, cut transversely, seems to
have ten cells, of which the five inner ones which con-
tain the seeds at the internal angle, are the true cells,
and the five outer ones, devoid of seeds, are the cavities;
these result from the epicarp swelling up during matu-
ration, so as to break the mesocarp, and form in its
place a space or air-cavity. In several fruits we find
cavities more or less decided, either in the axis of the
fruit, when the carpels, instead of reaching the centre,
leave there a small empty space ; or between the cells,

152 VEGETABLE ORGANOGRAPHY.

when the retreating faces of the carpels are not inti-
mately united together ; or on the sides of the valves
when they are puffed up, as in Myogram ; or at the top
of the pedicel, or in the axis, when it is fistular ; or,
lastly, at the base of the style, when this base is, in like
manner, fistular. These last cavities are remarkable for
sometimes containing a seed, as is seen in Brassica
cheiranthos, and in Trianthema monogyna ; this kind of
seminiferous cavity or stylary cell is of rare occurrence,
and inexplicable by any carpological theory ; its detailed
observation merits the attention of anatomists.

We have seen how the cells are formed in fruits by
the folding back of the margins of the carpels as far as
the axis; let us now examine what happens when the
retreating parts do not reach the centre. Three cases
are here presented to us : they either extend half-way in
from the top to the bottom, or they are so short as to
seem absent, or they nearly reach to the centre at the
base of the fruits, but are absent at the upper part.

When the retreating parts of the carpels are prolonged
into the interior without reaching to the axis, there
results a fruit, the centre of which is empty and the
circumference presents as many cells, open on the inside,
as there are carpels ; these cells bear the name of Semi-
cells. The partitions, which are then called Semi-
partitions, bear the seeds on their inner margins, as
usual ; this is seen in certain Poppies and Hypericineae :
on comparing together the species of these two groups,
we meet with almost every degree, from partitions which
reach very nearly to the centre, to those which are so
short as hardly to have their edges visible.

When the partitions are so short as to be hardly
visible, the placentse are then, as it were, applied to the
sides of the fruit, and each carpel is reduced to its
dorsal face. The fruit is then said to be Unilocular,

>'

STRUCTURE OF TRUIT. 153

and the seeds are Parietal ; we see this in Viola,
Helianthemum, Passiflora, Reseda, Argemone, the
Capparideae, &c.

Lastly, if the retreating parts only bear the seeds at
their base, and as it is this portion which is extended
towards the centre, the seeds are found placed at the
centre of the base of the fruit, and then the two fol-
lowing cases may happen : —

1st. Sometimes the partitions are prolonged towards
the centre as far as the top, and then, not bearing seeds,
they are usually thin and membranous ; in this case, the
fruit has still several cells with the seeds at the bottom
of each, as in some multilocular Caryophylleae.

2d, Sometimes the upper part of the partitions appears
to be wanting at maturity, because the carpels, which,
at the period of fecundation, were of the same length
as the placenta, afterwards elongated so as to rupture
the upper part of the partitions, and, more or less, com-
pletely isolate the placenta. This appears to take place
in several Caryophylleae. In all these cases, the fruits
are said to be Unilocular, and the seeds attached to a
Central placenta, although in reality the fruit is
always formed of united carpels, the retreating parts of
which bear the placentae on the inner margins.

We have above seen, that each carpellary placenta is
prolonged into a style, that the union of the two placen-
tary styles forms the carpellary one, and that the union
of the carpellary ones forms the style properly so called.
This organization never presents any difficulty when the
placentae occupy the whole length of the fruit ; thus,
whether the placentae reach the centre, whether they
stop half-way, or whether they hardly exceed the margin,
we know that they have a direct communication with
the style. But what takes place when the placenta is
central, and does not reach the apex of the fruit ?

131 VEGETABLE ORGANOGRAPHY.

There may happen two cases which correspond to those
mentioned above.

Sometimes the partitions may originally be in a state
of great tenuity, or the carpels, at the time of flowering,
may not be longer than the placentae, and then the
filament which proceeds from the placenta can reach the
base of the style and transmit the fecundating matter
to the ovules. This filament is destroyed after fecunda-
tion, either by the destruction of the partitions, or by
the elongation of the carpels; and then at maturity we
do not find it, and can only conceive how the fecundation
could take place by having recourse to the anatomy of
the ovary at the period of flowering. We see this in
all the Caryophylleae with a central placenta; sometimes
the filaments, which at the time of fecundation arise
from the placenta, are distinct, as in Lychnis dioica,
where there are five ; in Stellaria, where we see three
(PI. 20, fig. 1) ; sometimes they are all united into one,
as in Arenaria (PL 20, fig. 3, 4). An analogous organ-
ization is met with in the Portulaceae (PI. 20, fig. 7, 8),
where we find three distinct filaments; in Primula, where
the placentae are all united into a nearly globular body,
the filaments also being united into a point which
reaches the base of the style. In all these examples,
the filaments are wholly or in part destroyed after fe-
cundation, and the placenta seems isolated from the style.

It may also happen that the branch of this pistillary
cord may go along the margin of the carpel which does
not retreat, as in fruits with parietal placentae, and thus
it arrives at the base of the fruit and to the seeds which
are there situated. It is this, probably, that takes place
in the fruit of Luzula, for example, where we find the
seeds attached to the base of the valve.

The placenta is usually placed at the inner angle of
the more or less retreating part of the carpel, either

STRUCTURE OF FRUIT. 155

throughout its whole length, or only at the base ; but
its form and dimensions present some remarkable
differences, which modify the structure of the fruit.
Most frequently, it presents an elongated ridge bearing
one or two rows of seeds ; sometimes becoming very
large and thick, it extends into the interior of the cell,
forming a large projection, as is seen in Datura,
Solarium, Nicotiana, &c. ; sometimes it spreads out and
lines the whole of the retreating part of the carpel, as
is seen in the Poppy and Nymplicea; and at other
times, it is expanded into a kind of net-work, applied
to all the inner walls of the carpel, and bearing here
and there the seeds, as is seen in the Flacourtianeae and
Butomeae.

The Cruciferae present, in this respect, an organization
which is peculiar to them ; the two carpels, which
compose the Siliqua (this is the name given to this
kind of fruit), have their retreating margins reduced to
an extremely thin and delicate membrane, which may
be regarded as an internal prolongation of the epicarp
alone, and the placentae are situated upon the margins
of the endocarp, which is not prolonged into the interior,
so that the seeds are parietal, although the fruit is
bilocular.

All these various combinations are often rendered
obscure in fruits which open at maturity, by the different
modes of dehiscence ; and in indehiscent fruits by the
development of the pulp or flesh, which confounds their
different parts in an almost indistinct mass : these two
causes of obscurity, as well as those which proceed from
abortions or from the state of the central axis, deserve
to be analyzed.

All the modes of dehiscence which we have found in
isolated carpels, may be met with in cohering ones, but
modified and multiplied by this cohesion.

156 VEGETABLE ORGANOGRAPHY.

The most simple, but not most frequent case is that
which is called Septicidal dehiscence, because it takes
place in the partitions which seem to split in two : there
is a particular case of this, which T name Dehiscence
by Separation ; it consists in the carpels being so
slightly joined together, that they separate from one
another at maturity, forming so many distinct bodies, at
first closed, afterwards opening, by one of the systems
mentioned above, in solitary carpels ; thus, the carpels of
the Colchicaceas separate at maturity, and open in the
manner of follicles by a fissure along the ventral suture,
which in the entire fruit was central. Thus the carpels
of Hermannia laevigata separate at maturity, and each
of them opens by its two sutures, as most legumes.

This dehiscence is also modified by the existence or
non-existence of a central axis : when this does not
exist, it may happen, either, — 1st, that the carpels
detach themselves entirely from one another, leaving the
centre of the fruit void, as we see in Colchicum ; or, 2d,
that the extreme portions of the partitions, which bear
the placenta), may be so united together as not to
separate, then the rupture takes place along the placenta;
the carpels open by leaving in the centre a false semini-
ferous axis, formed by the intimate union of the inner
margins and placentas ; this takes place in the Balsam.
When the axis exists, the same two cases may happen :
sometimes the carpels, on detaching, carry with them
the placenta?, and leave the axis bare, as in the Malva-
ceae, Euphorbiaceae, &c. Sometimes the placentas may
remain united to the axis, and the rupture may take
place along the partitions; but I do not know an ex-
ample of this mode of dehiscence, and, in general, it is
not always easy to distinguish the case where the
apparent axis is formed by the placentas alone, or by the
placentas united to the axis.

STRUCTURE OF FRUIT. J 57

Let us suppose, now (and of this there are numberless
examples), that the two retreating faces of the carpels
are so united together that they cannot separate, and
that, notwithstanding, the fruit must open, and this
always happens when, if not fleshy, it is filled with
seeds. There takes place then a Dehiscence by
Rupture, and this may be presented under six different
forms, viz. : —

1st. (And this is the most frequent case), the de-
hiscence takes place along the dorsal nerve, or middle
line at the back of the carpel ; this is what we call
Loculicidal dehiscence, because it occurs in the
middle of the cells ; in this case, one is always inclined,
at first sight (and I myself, with most botanists, have
for a long time committed this error), to take for the
primitive elements of the fruit, not the carpels properly
so called, but the Mediastins, that is to say, the
bodies formed by the halves of two carpels united
together by their retreating faces ; it is in this sense,
founded only upon appearance, that we have called
Valve the outer part of the mediastin, although really
formed of two semi-valves, and that we have said that it
bears the partition upon the middle of its inner part,
although this partition, really double, arises from the two
margins of neighbouring valves. This organization is
found in Liliaceae, Ericinese, Tiliaceae, See. &c. It is
modified, like septicidal dehiscence, by the existence or
non-existence of the central axis, and by the more or
less great degree of adhesion of the placentae, either
with each other or with the axis. Thus, in the Irideae,
all of which have loculicidal dehiscence, the placenta?
remain united, forming a false axis in Belamcanda,
whilst they accompany the partitions in most of the
others, and especially in Iris.

2d. It happens in some families, such as the Cruciferas.

158 VEGETABLE ORGANOGRAPHY.

Capparideae, Fumariaceae, and some Papaveraceae, that
the margins of the carpels which do not extend into the
interior, or, if they do, by a very delicate lamina, are
however so united together that they cannot separate at
maturity. These united margins, together with the
placenta, form kinds of thick, firm nerves : the rupture
in this case takes place on each side along this nerve,
the whole of the intermediate part of the carpel is
detached and receives the name of Valve, and the
filament composed of the two placentae, united to the
margins of the carpels, is called the Intervalvular
placenta. An analogous phenomenon takes place in the
Orchideae.

3d. It happens in some genera, with what are called
central placentae, that the carpels, after fecundation,
have a tendency to elongate beyond the placenta, the
partitions being, at the same time, very delicate and
easily broken, whilst the outer parts are strongly united
together both at the base and apex ; in these com-
plicated circumstances, the union of which is con-
sequently rare, the rupture takes place transversely
across the middle of the carpels: we call this Trans-
verse (circumscissa) dehiscence, of which Portulaca,
AnagalUs, &c. present examples. It is met with in the
Lecythideae with a peculiar combination.

4th. Among genera with central placentae, the capsule
of which elongates after fecundation, and by this means
seems unilocular, at least at the upper part, it also fre-
quently happens that the outer portions of the carpels
remain united together the greater part of their length,
but at the upper extremity, they have a tendency either
to separate from one another or to split open along the
middle nerve ; this constitutes Apicilar dehiscence.
It is observed in a great number of the Caryophylleae :
the number of the teeth is equal to that of the carpels

STRUCTURE OF FRUIT. 159

when each of them remains entire, and double this number
when there is a fissure of the middle nerve. The same
kind of dehiscence is produced in the Poppy by a
different cause — the existence of the torus in a mem-
branous state surrounding the carpels.

5th. The contrary, also, sometimes takes place, as in
Cuscuta, for example, where the carpels are more
united at the apex than at the base, and separate at
maturity at the lower extremity; this constitutes
Basal dehiscence, which is almost always slightly
irregular, and is nearly confounded with transverse de-
hiscence.

Gth. Lastly, it sometimes happens, even in dry poly-
spermous fruits, that the carpels are so united together
that they cannot separate or split regularly by any part
of their surface ; there generally result, then, towards
the top of each carpel, kinds of pores, or irregular
ruptures, which give passage to the seeds, but which can
only be placed among dehiscences called Irregular;
we see it in Linaria and several other Scrophularinese.

In indehiscent fruits, the true nature of the carpels is
obscured by causes differing from the preceding ; some-
times the fruits do not open, because the pericarps are
membranous and dried up, and then there is usually an
abortion of several parts ; sometimes they are indehis-
cent because they are fleshy : and here we find the same
distinctions as among fruits with isolated carpels ; the
flesh, which is only a development of the mesocarp, is
found outside the cells, and the pulp is found inside
them : there are fruits, as the Quince, which have flesh
and pulp at the same time. Generally, in fleshy fruits,
we cannot easily recognise the original position and
place of the carpels, because the unions here are much
more intimate.

A frequent cause of error in the manner of appreci-

160 VEGETABLE ORGANOGRAPHY.

ating the symmetry of fruits with united carpels, is the
abortion of some of them, either wholly or in part.
Thus, a fruit which, as Lodoicea, ought to have six
lobes, is found to have not more than two or three, by
the constant abortion of the others. A fruit of the
Rubiaceae, which ought to have two equal cells and a style
springing from the centre, is found to have a single cell
and a lateral style, as Pleurogaster. A fruit which ought
to have three complete cells, is found, as the Pistachio,
to have a single fertile cell, and two others, half or
completely abortive, &c. &c. The number of similar
examples is immense, for there are few families in which
it is not met with.

The manner in which the carpels are placed with regard
to the axis, deserves also to occupy our attention for a
short time ; the axis which supports the carpels, such
as that which is observed in Magnoliaceae or Anonacese,
only becomes an integral part of the fruit when the
carpels are united after flowering ; the axis of the
Malvaceae, which is generally very visible, bears the
carpels adhering by their inner margins, and the carpel-
lary styles are either free or applied to it : we see this
also in the Geraniaceae, and, in general, in all fruits
which have a true axis. But it sometimes happens that
the carpels are articulated with a body which makes an
integrant part of the style, and through which the
vessels which carry the fecundating matter must ne-
cessarily pass ; this is observed in the Ochnaceae, for
example, and it is the swelling of the base of the style
which I have called the Gynobase. Some naturalists
have confounded it with the axis properly so called ;
but there is this important difference between these two
organs — the pistillary cord does not pass through the
axis, which I consider an elongation of the pedicel des-
tined to support the carpels, whilst it does pass through

STRUCTURE OF FRUIT. 101

the gynobase, which is nothing but an extraordinary
swelling of the base of the carpellary styles united
together. The true axis is usually elongated ; but it is
nearly globular in several Anonaceas ; it also takes this
form and a fleshy consistence in the Strawberry, where it
presents, moreover, the singularity of detaching itself
from a kind of more solid axis in the centre ; the
carpels of the Strawberry are little granular styliferous
bodies, dispersed over the surface of a fleshy body
which serves for their nourishment, and is nothing but a
round axis, to which several authors have given the
name of Polyphore. These axes must not be con-
founded with the thecaphores, which form part of the
carpels, of which they are, as it were, the petioles : the
axes, on the contrary, are prolongations of the pedicel
of the flower.

Hitherto 1 have always spoken of the carpels as being
leaves folded inwards, or upon their upper surface ; but
it would seem that the inverse organization takes place
in some Cucurbitacece ; when the young fruits of this
family are cut transversely, we find the carpels with
their backs opposite the centre of the fruit, and the
ovules are directed towards the side of the adherent
calyx. Are these carpels folded in a contrary direction
to all other plants, or have they twisted upon them-
selves before their development, so as to have the upper
part of the carpellary leaf directed towards the outer
side of the fruit ? I am ignorant of this. I would ven-
ture to add here an observation which is at least sing-u-
lar : — M. Seringe found flowers of the Gourd, the
anthers of which accidentally bore ovules, which were
directed outwards, because the anthers are extrorse. Is
there any relation between the extrorse direction of the
anthers and of the carpels of the Cucurbitaceas ? Does
this relation exist in other families ? These are ques-

VOL. II. M

162 VEGETABLE ORGANOGRAPHY.

tions which I leave to botanists accustomed to the study
of analogies, but upon which I cannot venture as yet to
hazard an opinion.

Section IV.

Of Carpels considered with regard to their relation to
the other parts of the flower which are persistent, or
united around them.

We have seen in the preceding Section what results
from the natural union of the carpels together, but that
is not sufficient to form a complete idea of the modifica-
tions of the fruit ; we must also study the pieces of the
flower which form, or seem to form, part of the fruit at
its maturity, viz. — the torus, the calyx, or the perigone.

The torus, as we have said, is the base of the male
and corolline parts of flowers. It is sometimes pro-
longed around the fruit, either under the form of
distinct petaloid scales, as in the Columbine ; or of pili-
forrn filaments, as in several Cyperaceee, and then it
cannot produce any illusion : or under the form of a
membranous cup which surrounds the carpels without
adhering to them ; thus in Pcsonia Moutan, var. papave-
racea it is thin and membranous ; it surrounds the
carpels without adhering there; it opens at its extremity
to give passage to the stigmata, and as it does not de-
hisce it seems to make part of the fruit, from which
however it is distinct. In Carex a similar cup is found,
open at the top, and enclosing the solitary carpel with-
out adhering to it, although it presses closely upon it.

In Nuphar, or the yellow-flowered Water-lily, we find
a thick cup, green and shining externally, closed at the

STRUCTURE OF FRUIT. ] 03

top, and surrounding the membranous, polyspermous,
verticillate carpels, which form the true fruit- During1
flowering and maturation, it seems to adhere strictly to
the carpels, but at maturity it becomes detached at the
base, and then we clearly see the distinction between
these organs.

It is nearly the same in the Poppy : here the torus
appears under the form of a thin lamina, which sur-
rounds the carpels and completely adheres to them, ex-
cept at the top of the ovary ; the valves of the fruit,
when they open at the top, are retained in their place
by this adherent sheath of the torus, and it is this which
causes the dehiscence of the Poppy under the form of
teeth or very short valvules, and not throughout the
whole length of the valves, as in the other Papaveraceae.

The Orange only seems to differ from the precedino-
examples, in that the torus, which is thick and glandular
externally, completely surrounds the carpels to the
origin of the style, and adheres to them by means of
very lax cellular tissue ; remove this continuous torus,
and you find the carpels verticillate around an imaginary
axis, separable without tearing, of a membranous tex-
ture, and etiolated like all shaded organs ; filled inter-
nally with a peculiar kind of pulp, which differs from
that of all other fruits, in its being enclosed in kinds of
utricles which arise from the walls of the carpels.

In the Capparideae, Passifloreaa, and some Legumi-
nosae, the torus only adheres to the thecaphore, and the
fruit itself is completely naked.

Such are the principal examples where we see the
torus adhering to or surrounding the fruit, without the
calyx or perigone following the same course. In Nym-
phcea, or white-flowered Water-lily, the stamens and
petals are united at their base with the torus, whence it
results that they seem adherent to the ovary ; they are

m CZ

164 VEGETABLE ORGANOGRAPHY.

destroyed after flowering, and the torus, which sur-
rounds the fruit, is found marked with their cicatrices.
I do not know any other example where we can find
these organs adherent to the fruit. But frequently they
remain without falling, and surround the base, as we see
in the Campanulaceas, Ericineae, several Leguminosae,
&c. But these persistent stamens or petals, do not
cause any remarkable differences in the history of the
fruit.

Let us now see what takes place when the torus, and
the calyx or perigone, united, adhere to the carpels, and
form what is called an adherent ovary or calyx. Through-
out the following, I only speak of the calyx for the sake
of abbreviation, but the whole of the section is equally
applicable to the perigone. This phenomenon necessa-
rily supposes, — 1st, that the pieces of the calyx or
perigone are united together, so as to form a tube more
or less prolonged ; 2d, that the torus is united to this
tube, and consequently the stamens and petals are peri-
gynous; 3d, that the carpels either adhere together or
are reduced to one. All these conditions are frequently
found united in the calyciflorous or perigynous families,
in which alone the phenomenon can be met with. We
frequently find in the same family every intermediate
degree between the free and adherent calyx. Thus, we
observe in the Rosaceae, genera with the calyces free
and expanded, and with the carpels distinct, as in Poten-
tilla and Spircea: others with the calyx free, and more
or less cup-shaped, enclosing sometimes several carpels,
sometimes a solitary one, without adhering to them, as
Alchemilla and Rosa ; and there are others (Pomaceae)
where the carpels cohere together, and are surrounded
by the fleshy calyx, as the Pear and Medlar. Analo-
gous transitions are observed in the Ficoids, Saxifragese,
Caprifoliaceae, &c. ; on the contrary, the adhesion of the

STRUCTURE OF FRUIT. 165

calyx with the ovary is constant in Myrtaceae, Cucurbi-
tacese, Umbelliferae, &c. ; it never takes place in Cras-
sulacese, Salicariae, and, perhaps, in Leguminosae. This
last family, however, presents in some cases a commence-
ment of adhesion : thus, in Arachis, Jonesia, and some
species of Bauhinia, the thecaphore or proper pedicel of
the carpel, is laterally united with the calyx. From this
fact, it is not impossible, perhaps, but that one day a
leguminous plant may be found with an adherent ovary.

The adhesion of the calyx with the ovary, only takes
place at the part where the torus is itself united to the
tube of the calyx ; consequently, if the tube be shorter
than the ovary, the adhesion can only take place to a
certain height, the stamens and petals will arise from the
margin of the tube around the ovary, the upper part of
which will be free, as in several Ficoids.

If the tube be as long as the ovary, which is most
frequently the case, the adhesion will take place through-
out the whole length of the two organs, the stamens and
petals will arise at the point of separation, and only the
limb of the calyx will be free ; lastly, if the tube be
prolonged beyond the ovary, and the torus also beyond
it, the ovary is then entirely adherent, and surmounted
by a tube, at the top of which arise the stamens and
petals, as in Oenothera : in almost every case we remark
at the top of the adherent ovary and around the style, a
little space usually round, or with as many angles as
there are sepals : this is the upper portion of the ovary,
which is not covered by the calyx ; sometimes it
increases after flowering, and then forms a very evident
mark upon the fruit ; it is very large in several Cucur-
bitacese, and especially in Cucurbita ^felopejoo ; it is also
very remarkable in the Medlar, in several Rubiaceae,
and, when carefully observed, wre find it in almost all, or
perhaps in all adherent fruits. This exposed portion of

1G6 VEGETABLE ORGANOGRAPHY.

the ovary is usually very smooth, and is thus distin-
guished from the calyx. Outside this disc formed by
the ovary, we find a small circular zone, which is the
trace of the point where the torus terminates. This
zone is very easily perceived when, as in the Pomaceae,
the stamens are more or less persistent on the fruit, or,
as in the Campanulaceae, where the corolla itself is per-
sistent ; it is also very visible when, as in several Cucur-
bitaceae, it increases after flowering : I suspect that it is
the torus, perhaps, which is prolonged a little after the
flowering, and forms, in the Rubiacea?, the little cup
which is found between the limb of the calyx and the
base of the style. In most adherent fruits this zone,
produced by the torus, is effaced at maturity.

The disc, formed by the naked part of the ovary, the
zone, produced by the torus, and especially the rest of the
free part of the calyx which remains, or leaves, at least,
some trace at the top of the fruit, form by their union
what is called the Eye, visible in this class of fruits, for
example, in the Pear.

The tube of the calyx, united with the ovary, may,
according to its texture, be moulded to the form of the
fruit, or compel the fruit to take its form; but most
usually the two bodies are modified a little in their
general form. Its texture is also variable : sometimes
it remains foliaceous or membranous, and then the fruit
is dry ; at other times it becomes fleshy with the ovaries,
and sometimes grows to a considerable size ; most fre-
quently it is impossible, in adherent fleshy fruits, to dis-
tinguish which is the part which is transformed into
flesh ; thus, in the Pear, for example, it may be either
the development of the sarcocarp of the carpels, or of
the torus, or of the calyx, or, what is most probable, of
all the parts at once.

The adhesion of the calycinal tube with the ovary is

STRUCTURE OF FRUIT. 167

usually intimate and lasting ; but it happens in some
cases, such as Cosmibuena, a genus of the Rubiacese,
near Quinquina, that at maturity it becomes detached
from the ovary, and only covers it over without exactly
adhering.

The free part of the calyx is presented under very
different forms, which influence the appearance of the
fruit and frequently its history ; it is sometimes scarious,
sometimes membranous, and sometimes completely ab-
sent, either from the time of flowering or at maturity.

When the entire tube of the calyx is united with the
ovary, and its lobes do not undergo any change, they
remain at the top of the fruit under the form of teeth,
as in CEnanthe or Conium, or they form a kind of eye,
as in the Pear or Apple.

If the tube be prolonged beyond the ovary and harden
after flowering, it results that the fruit is crowned with
a kind of peculiar collar, as in the Pomegranate and
some species of Gardenia.

Sometimes these lobes enlarge, remaining foliaceous,
or becoming slightly fleshy after flowering.

We have seen, in speaking of the modifications of
flowers, that it constantly happens in those which are
collected into a compact head, and sometimes in others,
that the limb of the calyx has a membranous and sca-
rious texture : in this case it remains at the top of the
fruit, and the name of Pappus is given to it.

Its function only begins to be important at the time
of the dissemination of the seeds.

These scarious calyces sometimes have their lobes
united into a single entire or toothed body, which causes
the fruit to be crowned with a scarious cup, as in Favo-
nium, Chrysogonum, Scabiosa stellata, &c. At other
times each lobe remains distinct, and takes either the
form of a little scale, as in Apuleia, Centaurea Crupina,

1G8 VEGETABLE ORGANOGRAPHY.

&c, or that of a long process, as in Pedis. Most fre-
quently each lobe is, as it were, replaced by a greater or
less number of scales in the form of hairs, which are
called the hairs of the pappus : they are sometimes
simple and free, and then the pappus is said to be
Hairy, as in Sonchus ; sometimes irregularly united
together, and then the pappus is Branched, for exam-
ple, in Stcuhelina ; sometimes toothed on the margins,
as in Hieracium and Chondrilla ; sometimes bearded
laterally, as in Scorzonera, and then the pappus is
Feathery.

The pappus, which is called Stipitate, is produced
because the calyx, and perhaps an elongation of the
pericarp, is prolonged perceptibly above the point where
the seed terminates ; as this portion is empty it remains
thin, filiform, and appears, at first sight, rather a sup-
port of the pappus than a part of the fruit ; we see it in
Tragopogon, &c.

It sometimes happens that the pappus is in two rows,
which do not resemble each other. In this case, the
outer row is certainly the limb of the calyx ; but I should
not be surprised to find it proved that the inner row is
a prolongation either of the torus, or of the pericarp ;
this is observed in some species of Centaurea.

The limb of the calyx of the Valerianeas, during
flowering, is rolled inwards, so as only to present a small
circular limb ; it afterwards unrolls, and the fruit is
crowned with a feathery pappus ; the Proteaceas have a
kind of pappus which is formed by the limb of the
perigone.

Finally, the limb of the calyx is sometimes com-
pletely absent ; this phenomenon may take place even
during the time of flowering, when the whole of the
calyx is united with the ovary, as happens in most of
the Umbellifera? ; still even in this case the lobes almost

STRUCTURE OF FRUIT. 169

always exist, but reduced to very small teeth. The
entire absence of the limb is more visible in those Com-
positas without a pappus, as the Daisy, &c. ; the limb is
here indicated by a small circular rim, entire or un-
equally toothed.

In other cases the limb is visible at the period of
flowering, but it is destroyed or detached naturally at
maturity ; we observe this in Epilobium, &e.

Nyctago and the Marvel of Peru present, in this
respect, a phenomenon worth mentioning : the base of
the perigone, united with the ovary, forms a kind of oval
nut, and the upper part separates immediately above it,
after flowering, and falls off, the nut remaining within
an involucrum which has the form of a calyx.

It is not necessary that the calyx should be, strictly
speaking, adherent to the ovary so as to form an in-
tegrant or apparent part of the fruit ; thus, for example,
in the Rose, the carpels are dispersed in a kind of cup,
which forms the tube of the calyx ; they adhere to it only
by their bases ; after flowering, the calyx and torus unite
together, increase in size, and become very fleshy, prin-
cipally in the inner part. The internal cellular tissue
penetrates between the bony, indehiscent, monospermous
carpels, which appear to be simple seeds contained in a
pulpy pericarp, whilst they are caryopses embedded in a
calyx become fleshy.

In a great number of plants, and especially in the
Monochlamydea?, the calyx or perigone, without adher-
ing to the ovary, covers it over so closely that it abso-
lutely seems to form part of the fruit ; in this case it
sometimes remains membranous, as in Atriplex, and at
other times it becomes fleshy, as in Blitum.

When the calyx, without adhering to the ovary,
remains around the fruit in a looser manner than in the
preceding case, we are contented to say that the fruit is

170 VEGETABLE ORGANOGRAPHY.

Covered, when, as in Phy salts, the calyx has a ten-
dency to be closed at the top, and thus entirely surrounds
the fruit ; we say that it is Concealed, when the per-
sistent calyx only surrounds it in part, as in Nicandra
or Hyosciamus. The calyces of the Labiatae are tubular,
persistent, and contain four monospermous caryopses ;
after flowering, in certain genera, their lobes close in
together, and the fruits may be said to be covered ; in
others they remain more or less open, and the fruits may
be said to be concealed ; in this last case, it ahnost
always happens that little hairs, which are not perceived
before on the inner face of the calyx, are developed
after the fall of the sexual and corolline parts ; they
close the entrance of the tube, and protect the young
fruit from rain and insects.

Section V.

Of the Organs situated outside the Flower, and which
sometimes seem to form Part of the Fruit.

Not only can the organs of the flower, in certain
cases, become integrant or apparent parts of the fruit,
but the same thing may happen with the bracts or
involucra as well as the peduncles and receptacles.

All that I have said of the calyx or perigone, in their
connexion with the fruit, may almost apply to the
bracts or involucra, observing only that the examples
are much less numerous ; thus, we sometimes find
bracts which adhere to the calvx, or cover it so inti-
mately, that they seem part of the fruit : in Scohjmus

STRUCTURE OF FRUIT. 171

Hispanicus, the bracts, from the axils of which the
flowers are developed, surround the ovary so closely and
are united to it in such a manner as to seem an integrant
part of the fruit ; it is tin's which caused Gsertner to
give it the name of Scolymus angiospermus.

In Echinops, the bracteoles, which collectively form
the involucellum, perform the part of a calyx towards the
ovary, being united with it and forming a kind of false
scaly pappus.

In Lagasca, the involucellum surrounds the achenium
without adhering to it, and seems to be a cup-like calyx
around the pericarp.

In all the Compositae and Dipsaceae with a double
involucrum, the bracts, which form the involucellum or
proper involucrum of the flowers, present, more or less
distinctly, analogous phenomena.

Lastly, to mention other families, the bracts of
Pollichia become fleshy after flowering, and are easily
taken for an integrant part of the fruit, which they
cover ; the foliaceous involucrum of the Hazel seems to
form part of the fruit ; the cup of the Acorn is a true
involucrum formed by the union of a great number of
little bracts, and the Acorn, like the Nut, is a fruit
formed by an ovary adherent to the calyx. These two
examples present a rare peculiarity in the vegetable
kingdom, viz. : — a fruit which adheres by a large portion
of the base, which at the time of separation caused a great
cicatrice like that which is most commonly seen in
seeds. This cicatrice of the fruit, or Carpal Cicatrice,
ought to be distinguished from that of seeds, or the
hilum, of which we shall shortly speak.

The peduncles themselves sometimes seem to form
part of the fruit; thus, in Semecarpus and Anacardium,
they enlarge after flowering, become fleshy, and take
the form of a Pear, whilst the true fruit, which is dry,

172 VEGETABLE ORGANOGRAPHY.

is situated at their apex, seeming to be a kind of ex-
crescence.

In Hovenia dulcis, they also become fleshy after
flowering, and seem to form the true fruit.

Section VI.

Of the Aggregation of Fruits which proceed from
differe7it Flowers.

The facts mentioned in the preceding section lead us
to the study of Aggregated fruits, or those formed
by the intimate or apparent union of fruits proceeding
in reality from different flowers. This phenomenon
never occurs but in plants where the carpels are solitary
and most frequently monospermous by abortion ; it
likewise almost always supposes, as necessary condi-
tions, on the one hand that the solitary carpel is united
to the calyx, and on the other that the flowers are
placed very near together. I shall explain this by
examples derived first from capitate or umbellate flowers,
and afterwards from those in spikes.

Honeysuckles have naturally two flowers which arise
from the same axil ; their pedicels are frequently united
into one, which consequently bears two flowers and
two berries ; but it happens in several species, as for
example, Lonicera Xylosteon, that the two fruits are
more or less united into a single one, bilobed or almost
entire ; in this last case, the union is perceived, either
because, during flowering, we saw two corollas upon an
apparently single ovary, or because, after this period,
we recognise the two eyes which indicate the fall of the

STRUCTURE OF FRUIT. 173

sexual parts and appendages of the adherent ovaries.
In Symphoricarpos, a genus so near Lonicera that it was
for a long time united with it, instead of there being only
two flowers, there *are several united by the ovaries,
whence there results a fruit composed of several joined
together, each of which still presents its own eye : the same
occurs in Morinda ; it is also met with in Opercularia, with
only this difference, that the flowers, which by their near
approach form a compact head, have not a fleshy fruit,
but their calyces and bracts are all united together : at
flowering, we see all the corollas distinct — at maturity,
the fruit is slightly irregular, and composed of all the
partial ones joined in a head. The same thing takes
place in the Composite, in Gundelia ; here the bracte-
oles united together surround the partial fruits, so that
at maturity there results a mass composed of the recep-
tacle, bracteoles, and achenia of all the flowers of which
the head was composed.

The fruit known under the name of Fig, is a remark-
able example of aggregation, analogous to the preceding
cases : it is either a hollow pedicel, or rather (if attention
be paid to those exotic species which have scales
externally), a kind of fleshy involucrum, formed of a
great number of thick bracts intimately united at the
base either with each other, or with the top of the
peduncle, and very slightly free at their extreme apex.
The flowers are very numerous within this involucrum,
the top of which is scarcely open ; the female ones,
which are the most numerous and more central, are trans-
formed into as many little caryopses as there seem to be
seeds, and which, at maturity, are as nuts in the centre
of this fleshy or pulpy involucrum : it may be said, then,
that there is no other difference between the fruit of the
Fig and that of the Rose, except that the pulpy part of
the Fig is an involucrum, and that of the Rose a calyx ;
that, consequently, the seeds of the former are caryopses

174 VEGETABLE ORGANOGRAPHY.

proceeding from different flowers, and those of the latter
caryopses proceeding from but one flower.

What I have said of the Fig becomes still clearer
when we compare it with the neighbouring genera
Amhora and Dorstenia, in which the receptacle is open.

Flowers disposed in spikes sometimes present all the
phenomena which I have above mentioned ; thus, when
we follow the Mulberry from its flowering to maturity,
we see that the flowers are sessile on an axis which is
articulated at its base ; after flowering, the ovary is
covered by the perigone, and is transformed into a small
pulpy fruit; all these fruits are united incompletely
together, and appear the more readily to form a single
one, as the general peduncle disarticulates at its base, so
that the Mulberry is detached from the tree like simple
fruits. All that I have said of this fruit is strictly
applicable to the Bread-fruit, except that in the latter
the partial fruits are more completely united, and the
whole fruit which results from the aggregation is larger,
and the flesh more farinaceous ; and as for the culti-
vated Bread-fruit, we may add that the seeds are almost
always abortive, leaving their places empty, which forms
the irregular cavities in the centre of the mass.

The history of the Pine-apple slightly differs from the
preceding examples : the flowers are disposed in a dense
spike along the stem, almost as in Eucomis; after
flowering, the flowers which have the perigone adherent
to the ovary, are each transformed into a fleshy fruit,
originally trilocular; these are united first with the
bracts situated at their base, afterwards with each other;
the development of the fleshy part, and the intensity of
the union, are greater in proportion as the number of
abortive seeds is great ; and when they are all abortive,
as in the cultivated ones, there results a compact oval
head, in the centre of which we see, as in the Bread-

STRUCTURE OF FRUIT. 175

fruit, empty cells which indicate the abortion of the
seeds, and externally kinds of scale, which are the per-
sistent remains of the bracts and lobes of the perigone ;
the whole is crowned with a tuft of leaves, which are
nothing but foliaceous bracts devoid of flowers, which
spread out at the top of the spike, as in Eucomis, and
the development of which is favoured by the abortion of
the seeds of the lower flowers.

The fruits of the Coniferae present phenomena very
analogous to the preceding. If we examine the female
cone of a Fir, we find small flowers sessile in the axils
of the bracts, and disposed in a spike along an axis ;
after flowering, the flowers which have the perigone
adherent to the ovary ai'e each transformed into a kind
of nut, or samara, and the bract, which grows much,
completely covers the fruit ; this assemblage has re-
ceived the name of Cone; its axis is sometimes acci-
dentally prolonged into a leafy branch (PI. 16, fig. 4),
which is similar to what constantly takes place in the
Pine-apple. The cones of the Proteaceae, and the
follicular heads of the Hop, present an analogous organ-
ization; these kinds of cones differ from those of
Magnolia, or the Tulip- tree, in proceeding from the
aggregation of the carpels of several flowers in a spike,
whilst in the Magnoliaceae, they are formed by the
aggregation of several carpels in a spike proceeding from
a single flower.

But there are some Coniferae where the phenomenon
is complicated in consequence of the form or texture of
the organs. Thus, in the Pine, we find the same
general disposition ; but the bracts, after flowering,
enlarge and become very thick at the top, so as to form
a close mass, which only opens shortly before their
separation ; the Cypress and Thvja have these same
bracts less numerous, and so dilated at the top that they

176 VEGETABLE ORGANOGRAPHY.

form kinds of convex discs and pedicels; the cone, which
has then very improperly received the name of nut, has
a globular appearance ; it is close and semi-fleshy in its
young state, but at maturity it becomes dry, and the
scales separate by kinds of slits which give passage to
the caryopses or achenia which they enclose, and are
usually falsely called seeds. The Juniper differs from
the Cypress only in the bracts, thick at the top, being
fleshy and much more united, whence it results that the
fruit at maturity presents the appearance of a globular
berry, which name it has improperly received : the traces
of the union of the bracts are hardly perceptible, and
the enclosed caryopses have still more the appearance of
simple seeds. Thus the apparently simple berry of the
Juniper is formed by the natural union of fruits pro-
ceeding from several flowers, nearly as the berry of
several species of Anona and of Dillenia is formed by
the natural union of the carpels proceeding from one
flower.

These apparent affinities between the fruit of different
classes, have frequently caused analogous terms to be
used in popular nomenclature. The fruits of the Chest-
nut (JEsculus) and those of the Horse-Chestnut have
a very great external resemblance. The Chestnut,
seen at the time of flowering, presents several female
flowers, collected in an involucrum, which enlarges and
becomes very spiny ; each flower has an ovary surrounded
by an adherent calyx ; this ovary is formed of three
united carpels, each containing two ovules : during and
after flowering, several of the seeds become abortive ; and
there sometimes remains but one.

In the Horse-Chestnut, on the contrary, the flowers
are perfectly distinct, and the calyx is not adherent ; the
ovary is formed of three carpels united into a body,
bristly externally, each of which encloses two ovules ;

STRUCTURE OF FRUIT. 177

but during and after flowering, several of them become
abortive, so that the capsule frequently has but two
cells and two or three seeds.

Thus, the spiny husk of the Chestnut is an involucrum,
that of the Horse-Chestnut a capsule. The brown,
shining, round bodies of the Chestnut are achenia,
furnished at the base with a large carpal cicatrice; those
of the Horse-Chestnut are seeds with a large spermal
cicatrice. The bodies enclosed in the brown envelope of
the Chestnut are distinct seeds ; those within the brown
skin of the Horse-Chestnut are the cotyledons, or por-
tions of the seed. Although this example is trivial for
botanists, I thought that I ought to mention it in detail
for beginners, because, better than all reasoning, it
proves the necessity of referring to the period of flower-
ing in order to comprehend the structure of fruit.

Section VII.

Of the Umbilical Cord and its Expansions.

We have already said, that the Funiculus or Um-
bilical Cord proceeds from the placenta, and supports
the seed ; that it is composed, during flowering, of the
filament proceeding from the style and bearing the
fecundating fluid, and of a fibre coming from the pedicel
and carrying the nourishment ; that after this period,
the pistillary filament is obliterated, and the funiculus
remains formed of the nourishing fibre alone : we may
consider it as making part of the pericarp, either on
account of its texture analogous to the placenta, or
because that, at maturity, it usually happens that it
remains adherent to the placenta when the seed is

VOL. II. , N

178 VEGETABLE ORGANOGRAPHY.

detached ; but this last character is subject to several
exceptions, and we shall presently see that it is often
difficult to fix the precise line of demarcation between
the pericarp and the seed.

The funiculus is usually presented under the form
of a short and scarcely visible filament ; it is very long
either in those fruits where the cells are large, as certain
Mimoseae, or where it is curved or folded, as in the same
plants, in some Cruciferas, &c, or where it is destined
to support the seed when out of the cell : thus, in
Magnolia, the free carpels, of which the fruit is com-
posed, open along their dorsal suture, and the one or
two seeds which they contain hang out, supported by a
long, slender, silvery-white, and flexible funiculus. It
has been remarked, that it is a bundle of tracheae ; I do
not know that any similar observation has been made
upon the funiculi, which, in almost all other plants, are
not capable of being extended.

The funiculus is usually free from all adhesion ; but
there are plants in which, being very near together, they
are constantly united ; this is observed among the Cruci-
feras in the genus Eunomia. It more frequently happens
that it is found naturally united with the walls of the
cells ; thus, for example, in some Cruciferaa, such as
Lunaria or Petrocallis, the funiculus is united, through-
out its whole length, with the partition in the middle of
the fruit. In some Mimoseaa, it adheres to the valve
from which it originates ; in these cases, the seed,
although arising from the margin of the carpel, seems
to proceed from the middle of the partitions or valves.
It is possible that it may be from analogous adhesions of
the umbilical cord and not of the placenta, that arises
the position of the seeds of the Flacourtianeae and
Butomeas, scattered over the inner valves of the fruit.

When the funiculus of a free carpel or of a cell of

STRUCTURE OF FRUIT. 179

the fruit arises from near the base, if it be short, the
seed is necessarily erect, for example, in all the Com-
posite ; if it be long enough to reach the top of the
cell and then curve at its extremity, the seed, although
originating from the base, is found pendent ; as is seen
in the upper cell of the fruit of Crambe, in the fruit of
Paronychia, &c.

Let us suppose, now, that the nourishing cord is long,
ascending, and united to the wall of the cell, the seed is
attached to its extremity, and appears pendent from the
top of the cell, as, for example, in the Dipsaceae ; in
this case, as in the preceding, one of the margins of the
fruit presents a small nerve ; in the first, this nerve,
which is very delicate, is produced by the pistillary cord,
in the second, by the nourishing cord.

When the seeds arise from the margins of the carpels,
or from the inner angle of the cells, they are naturally
horizontal ; but when the funiculus is long, and especi-
ally in pulpy fruits, it happens that they take a pendent
or uncertain position according to the development or
particular position of the fruit, or according to their own
weight. Thus the length, adhesions, and inflexions of
the umbilical cords, or of the pistillary and nourishing
cords, determine the general position of the seeds in
the cells of the fruit or in the carpels, combining
these characters with those above mentioned, with
regard to the position of the placentae and the number
of seeds.

The umbilical cord always bears the seed at its ex-
tremity, and the part of the seed to which it adheres is
that which is called the Umbilicus, Hilum, or Cicatri-
cula ; but as this cord has a tendency, in several fruits,
to expand a little before reaching the seed, these expan-
sions have received the name of Arillus ; and their
history is the more important, as, in certain cases, one

N 2

ISO VEGETABLE ORGANOGRAPHY.

is inclined to confound them sometimes with parts of the
pericarp, at other times with those of the seed.

The most simple cases are those where the funiculus
expands laterally, so as to form an appendage upon the
seed ; thus, in several Polygaleae, we find a lateral
arillus which evidently arises from the funiculus. In
this case, it is usually fleshy or membranous; it is,
perhaps, to this order of unilateral arilli, that the carun-
culi which are found in some species of Dolichos and in
Chelidonium ought to be referred. In the Nutmeg the
arillus is large, fleshy, and ramified, forming a kind of
incomplete envelope at the base of the seed ; it is what
is commonly called Mace : in Blighia it is so large and
•fleshy as to be worth the trouble of collecting for food.
The same phenomenon occurs in the Passifloreae, where
the inside of the arillary coat is full of a pulp secreted
apparently by the walls of the arillus ; when abundant,
it causes some capsules of the Passion-Flowers to be
classed with edible fruits.

In all the examples which I have mentioned, the
arillus forms an incomplete envelope around the seed,
and this is what ought to be considered the distinctive
character of this kind of expansion of the funiculus.

We give, on the contrary, with Gaertner, the name of
Epidermis, to a dry, thin, membranous sac which en-
tirely covers the seed ; this organ is very visible in the
Malvaceae, Bombaceae, &c.

It is to be remarked that the arillus, whether it be
fleshy, membranous, or pulpy, never bears hairs ; on the
contrary, the epidermis is sometimes smooth, as in the
Gourd, but more frequently furnished with hairs ; and
as the skin of the seed, properly so called, never bears
hairs, whenever a seed appears so covered, it is because it
is invested with a very adherent hairy epidermis ; these
hairs are either very short, as in most Mallows, or very

STRUCTURE OF FRUIT. 181

long, as ill Gossypium, where they form the substance so
celebrated and useful, called Cotton ; sometimes they are
found scattered over the whole surfaces of the epidermis,
as in Ochroma ; sometimes only in certain places, as in
several varieties of the Cotton Plant : sometimes in a
tuft at the extremity of the seed, as in several Apo-
eyneae. These tufts, which have been called Crests,
{comae) so resemble pappi, that they have frequently
been confounded with these organs ; but they present
this essential difference — that the pappus, which is a
degeneration of the limb of the calyx, is on the outside
of the pericarp, and the crest, which is an extension of the
epidermis, is within the cells of the fruit, and upon the
seed itself. Notwithstanding this important anatomical
difference, their nature and properties have a great
analogy. These two kinds of tufts are formed of mem-
branous and very hygroscopic hairs, endowed with the
faculty of approximating when moist, and diverging
when dry ; whence it results, that as long as the ma-
turation has not been completed, these hairs, being
moist, remain close together ; but becoming dry at ma-
turity, they diverge, and thus tend to facilitate the
bodies to which they are attached in coming out of their
envelopes. The pappus draws the achenium out of the
involucrum ; the crest carries the seed out of the peri-
carp ; both of them, being expanded, permit the least
wind to bear to a distance these little bodies, to which
they perform the office of wings, or rather of parachutes.
I shall now revert to the modifications of the epidermis.
It frequently happens that this membrane is ex-
panded around the seed, and, instead of bearing hairs,
is dilated into a wing frequently well developed and very
delicate. It is thus that in several Apocyneae, Malvaceae,
&c. the seed is terminated, surrounded or enclosed by a
membranous wing, which, like the crest, contributes to

182 VEGETABLE ORGANOGRAPHY.

facilitate its passage and dissemination. But it must be
observed here, that this wing resembles several very
different organs, or rather that analogous expansions
may be developed upon almost all the organs of the
fruit: thus, although I am inclined to believe that the
greatest number of winged seeds owe this organization
to the epidermis, it is nevertheless possible, that some-
times the skin itself of the seed may be expanded into a
wing ; this seems to take place in the Bignonias with
winged seeds ; and I confess that, seeing the adhesion
and fineness of certain epidermes, I scarcely know any
means (except analogy) to ascertain if the wing of a seed
results from its own covering, or from its epidermis. The
carpels themselves may expand into wings, as in the soli-
tary ones of JYissolia, Sec, or in those united into a single
fruit in the Elm ; calyces adherent to the ovary and
becoming part of the fruit, form membranous wings,
either by the expansion of their limb, as in several
Dipsaceas and Compositas, or by the expansion of their
angles, as in several Umbelliferse ; and what is remarkable
in this degeneration, as well as in the preceding, is that
the physiological function of these expansions is ab-
solutely the same in every case, whatever be their
anatomical origin. The wings always serve for the
dissemination either of the seeds properly so called,
or of the carpels or fruits which contain but one or
two seeds ; for they are hardly ever formed upon poly-
spermous fruits. Thus they always serve in the end,
wherever they may be placed, to separate the seeds
from one another for their natural dissemination.

I believe that it is also to the presence of a very
delicate, but hygroscopic epidermis, that we must
attribute a curious phenomenon, viz. — the faculty of
certain seeds of absorbing moisture, and of being found,
when placed in water or wet earth, surrounded by

STRUCTURE OF SEED. 183

an aqueous pulp, retained around them by a very
delicate membranous net-work ; Lepidium sativum, the
common Flax, and several other seeds, exhibit this
phenomenon, which must tend to facilitate their ger-
mination.

Among the different examples of the accessory integu-
ments of seeds which I have mentioned, the origin of
the arillus, as a prolongation of the funiculus, is very
evident ; but the origin of the epidermis is much less so ;
it may be considered as proceeding also from the funi-
culus on account of its position around the seed, and
because it is evidently an organ in addition to those
which essentially compose the seed.

CHAPTER IV.

OF THE STRUCTURE OF THE SEED OF PHANEROGAMOUS

PLANTS.

Section I.

Of the Seed in general.

A Seed (semen) considered with regard to the flower,
is a fecundated ovule ; considered individually, it is a
cavity closed on all sides, containing the rudiment of a
plant. It is composed of the embryo or germ, which
has received the fecundation, and of its different appen-
dages, some of which serve as nourishing organs, and
the others as protecting integuments.

184« VEGETABLE ORGANOGRAPHY.

The seed properly so called, such as I have defined it,
must be distinguished from monospermous fruits and
tubercules ; thus, it may be confounded, as it frequently
has been in botanical, and as it constantly is in ordinary
language, either with a monospermous pericarp adherent
to the seed, as the caiwopsis, such as the seed of the
Wheat, or with the body which proceeds from the
union of a solitary seed with the pericarp and calyx, as
the achenium of the Compositse ; or with this same
achenium also united with the involucellum, as in Sco-
lymus. In all these cases the seed forms part of the
body to which it gives its name, but it is not isolated,
and unless one take care to separate it, either really or
in imagination, from the organs to which it is joined, it
will be impossible to understand its description.

On the other hand, one is often inclined to take for
seeds, the tubercules or bulbs which are produced in
certain parts of plants, but which are germs developed
without fecundation. The distinction of the seeds from
these bodies is often very difficult, sometimes impossi-
ble ; therefore, to avoid all uncertainty, I shall derive
all that I have to say from the seed of plants in which
this doubt does not exist, and I shall leave for the
following chapters the examination of the doubtful
cases.

A seed may be considered as a germ which is de-
veloped in the axil of a leaf, curved upon itself in the
form of a closed envelope. This fecundated germ takes
the name of Embryo ; the leaf which surrounds it, that
ofSpERMODERM ( sjoermodermis J , or skin of the seed:
these are the only two organs essential to the ripe seed.
We sometimes find in the spermoderm another body,
which is called Albumen, and which deserves special
attention ; the spermoderm, albumen, and embryo, will
be, then, the three parts which we shall have to study.

STRUCTURE OF SEED. 185

The umbilical cord bears the seed at its extremity;
the trace which it leaves upon it after it is detached, or,
in other terms, the place by which it adhered to the
funiculus, is its Cicatricule (cicatricula), also called
Hilum or Umbilicus : this place is always considered
as the base of the seed ; the apex is not determined
anatomically, as in the fruit, where the trace of the style
clearly indicates it; whilst the seed does not give origin
to any other organ, but it is found convenient to call the
ideal Axis of the seed, the straight or curved line, which,
arising from the base, proceeds at an equal distance from
the margins ; and we name the Apex the extremity of
this line. It evidently follows from these definitions —
1st, that the base of a seed is nearest the pedicel of the
fruit in erect seeds, nearest the axis or walls of the fruit
in horizontal ones, and nearest the style in pendent
ones ; 2d, that the position of the seed is only con-
sidered with regard to the pericarp, and not to the rest
of the plant ; thus, when a fruit is pendent, we say that
the seed is erect, when its apex is directed towards the
earth ; and that it is pendent if its apex point towards
the sky.

The abortion of the ovules or seeds, either before
fecundation, during flowering or at maturity, is a phe-
nomenon so frequent, that it might be said, without
exaggeration, that it is rare to find fruits, all the ovules
of which have arrived at the state of ripe seeds. It may
be caused by the slightest derangement, either in the
fecundating or nourishing apparatus of the ovules ; and
even when these two systems of organs are in a perfect
state, and when no external accident deranges them,
there are still two frequent causes which produce these
abortions : —

1st. The more or less lateral position of the flowers
with regard to the axis either of the spike, branch, or

186 VEGETABLE ORGANOGRAPHY.

stem itself, causes an inequality in the facility with
which the sap penetrates the different sides of the
flower or fruit, and the less favoured sides often present
abortions.

#d. The fecundation cannot take place upon all the
stigmata at once, and the fecundating vessels which go
from the stigmata to the ovules, do not carry the fovilla
to all of them at the same time. When the ovules do
not grow rapidly after fecundation, this inequality in the
period of fecundation does not cause any abortion ; but
if one or more of them grow rapidly, then they tend to
render the others abortive, either by attracting all the
nourishing sap, or by compressing or obliterating the
filaments of the pistillary or nourishing cord of the
other ovule. As these causes are connected with the
original structure of each species, the abortions which
result are nearly constant, as we clearly see in the Oak,
Horse-Chestnut, Lodoicea, &c.

The unions of the seed with parts of the pericarp,
have already occupied our attention ; but I ought to
mention here the accidental union of them with one
another, a rare phenomenon, and of which I have as yet
only seen a positive example, shown to me by M. Hey-
land: it is that of two seeds of the Horse-Chestnut,
which were united together half-way. I mention this
fact, not only on account of its rarity, but because it
may lead to the explanation of another, more important
and less rare, viz. — the plurality of embryos in one seed.
This fact is frequent in different species of the Auranti-
aceae ; thus, the Orange has usually three or four, and
it is accidentally observed in some other plants, as in
Ardisia coriacea. Richard does not hesitate to regard
this plurality as monstrous. I should be inclined to
believe that it results from the incomplete union of two
or more ovules, the spermoderms of which united to-

STRUCTURE OF SEED. 187

gether do not seem to make more than one, and the
embryos of which are developed simultaneously. What-
ever be the cause, this plurality of embryos does exist
in some seeds, and they are sometimes isolated from one
another, and at other times united together. This last
case has been observed by my son : having remarked a
plant of Euphorbia Helioscopia, (PL 22, fig. 1,) which
came up with four cotyledons, he perceived that this
number was owing to there being two embryos united
together throughout their whole length; he has since
observed in Lepidium sativum, and Sinapis ramosa, this
same monstrosity, which bears the same relation among
plants to the monstrous animals formed by the union of
two young ones. We know that in these animal mon-
strosities, it frequently happens that a part of the organs
of one or both of them disappears ; it is in this way that
calves with two heads, or six legs, &c, are formed.
The same happens in the union of embryos ; some
instead of four cotyledons, have but three ; this is what
is observed in the Euphorbia and Lepidium of which
I have spoken, and in Ranunculus, Solanum, the Bean,
&c. &c. In order to complete what relates to the plu-
rality of embryos, I have slightly deviated from the
natural order of facts, to which I now return.

The seeds of the same plant are not all exactly of the
same size ; in most cases this difference is of little im-
portance, and we generally select the largest seeds for
sowing, because it is remarked that the plants proceed-
ing from them are more vigorous. But in some cases
this difference of size has a peculiar interest ; thus,
M. Autenrieth has remarked that among the seeds of
the Hemp, which is a plant constantly dioecious, the
longest, largest, and heaviest, produce male plants, whilst
those which are rounder and less heavy, produce female
ones ; the former have a longer radicle, and germinate

188 VEGETABLE ORGANOGRAPHY.

more quickly than the latter. Similar observations have
not as yet been made upon other dioecious plants, so
that it would be imprudent, in the present state of our
knowledge, to affirm if these laws be more or less
general.

The weight of seeds is much more capable of being
appreciated : in general, ripe and fecundated ones are
heavier than water, and this law appears to be universal.
Seeds which have not attained maturity, or the embryo
of which has not been fecundated, are almost always
lighter than water, a practical character which enables
all gardeners to distinguish good seeds from bad : it
must be remarked that good ones may swim when they
retain a stratum of air, confined around them by means
of hairs, wings, or cavities which may surround them.

Section II.

Of the Spermoderm, or Skin of the Seed.

The proper skin, envelope, or coat of the seed, is an
organ so distinct, that it was very right to give it a
name. Richard, from analogy with the word pericarp,
proposed that of Perisperm, and afterwards that of
Episperm; but these terms do not seem to me to be
admissible ; the first, because Jussieu used it in another
sense ; the second, because, having no similarity with
the sense of the word epicarp, it would produce con-
fusion. I have replaced them by the term Spermoderm,
which expresses, in one word, the skin or coat of the
seed; this envelope exists in every seed, and I cannot
admit what Mirbel says of its absence in some ; Gaertner
considered it as formed of two membranes, which he

STRUCTURE OF SEED. 189

called the testa and the inner coat; but if I differ from
the nomenclature of Richard, I entirely adopt his
opinion upon the nature of this envelope. It is, like all
foliaceous organs, composed of two membranes and of an
intermediate tissue ; the external one bears the name of
Testa, the internal one that of Endopleura, and the
intermediate plexus that of Mesosperm ; these three
parts form, by their union, the close coat without valves
or sutures which surrounds the nucleus.

The testa, when deprived of all the accessory integu-
ments, which the arillus, pericarp, calyx, or even the
involucrum may furnish it with, appears most frequently
under the form of a shining membrane ; sometimes,
however, it is rough as in the Tulip, or marked with
little tubercles or furrows, as in Oxalis. But in general,
it is smooth, even glossy, dry, scarious, osseous, or
almost petrous, as in Guilandina Bonduc.

Notwithstanding this appearance, it is eminently en-
dowed with the faculty of absorbing water, and performs,
in this respect, an important office in germination. It
also presents this singularity — that although the micro-
scope can discover no kind of pores, it not only absorbs
water, but even the coloured particles of this fluid,
tinted, for example, with cochineal ; this progress of
absorption is entirely analogous to what takes place at
the extremities of the radicle and in the stigma, which
obliges me to consider the testa formed of seminal
spongioles ; its colour presents much variety, and affords
here and there, especially in the Leguminosae, the most
vivid and glaring tints ; for example, in Abrus, Ery-
t/irina, Beans, &c.

The testa is interrupted at the point where the um-
bilical cord is attached to the seed; this forms the hilum,
to which I shall presently revert : this interruption
seems to indicate that this organ, like the epidermis of

190 VEGETABLE ORGANOGRAPHY.

leaves, owes its existence to its being more exposed to
the air than the other parts of the spermoderm : what
confirms this opinion is, that sometimes the testa cannot
be distinguished, or at least, is not of its usual texture
in all seeds which are invested with an epidermis, and
especially in those which are united to the pericarp.

The endopleura, or inner coat of the spermoderm, is
exactly in the seed what the endocarp is in the fruit,
that is to say, the upper face of the leaf folded upon
itself; this membrane never has either the shining ap-
pearance or solidity of the testa ; it is almost always of
an uniform white colour, and composed of a cellular
tissue, which, one would think, ought readily to absorb
water; but the case is quite the contrary, for it contains
the aqueous juices of the young seeds without absorbing
them, and at the period of germination it prevents the
water from passing directly to the embryo. The endo-
pleura seems moulded upon the Nucleus <of the seed
(this name is given to the whole of what is contained
within the spermoderm); but, in reality, it is the nucleus
that is originally moulded, as it were, in the empty
space within the spermoderm, which, when it begins to
grow, it distends.

The endopleura, at a certain point which is called
the Chalaza, gives passage to the vessels which bear
the nourishing or fecundating juices to the embryo.

The mesosperm is in the seed, what the mesocarp is in
the fruit ; that is to say, the plexus of fibrous vessels
and of cellular tissue which is found between the two
membranes ; in fact, it is most frequently a very delicate
and scarcely apparent fibrous plexus ; it takes a fleshy
or pulpy consistence in but a very small number of
cases, as, for example, in Magnolia, Irisfcetidissima, &c.
The seeds which have this peculiarity, are named
Semina saccata in descriptive works ; the dry and

STRUCTURE OF SEED. 191

brittle nature of the testa causes it to adhere less to the
mesosperm than this latter does to the endopleura ; it is
on this account, that those carpologists who say that
there are but two membranes in the spermoderm, have
united under the name of inner coat or hilofer, the en-
dopleura and mesosperm.

The fibres which form the mesosperm generally arise
from the hilum, and expand between the two membranes
of the spermoderm ; they fulfil two offices, and are, per-
haps, of two different natures ; the one kind, which may
proceed from the umbilical cord, may bear nourishment
to the embryo and spermoderm during maturation, and
when maturity has arrived, may be obliterated; the
other kind, which may be directed from all parts of the
surface, towards the point of the endopleura where the
embryo is situated, may be for the purpose of bearing
thither the water absorbed during germination. These
two orders of fibres have not as yet been accurately dis-
tinguished ; but those which are employed during ger-
mination may be observed, by causing large seeds to
germinate in coloured water. I have employed those of
the Broad Bean, and have very well seen the fibres of
the mesosperm become gradually coloured, when intro-
duced into cochineal water; I have even seen the colour
reach the embryo. In selecting any large seed where
the position of the chalaza may be different from that of
the hilum, we might decide, by direct experiment, the
as yet obscure problem of the nature and direction of
the fibres of the mesosperm.

The portion of the seed where the testa is absent is,
as we have said, the hilum ; here we may distinguish
two parts : the one, situated near the margin, is a little
depression which Turpin has called the Micropyle,
which is, according to him, the trace of the place where
the branch of the pistillary cord entered ; the other,

192 VEGETABLE ORGANOGRAPHY.

which the same naturalist names Omphalode, and
which occupies almost all the rest of the hilum, is
slightly convex in the centre, and appears to be the
trace of the cicatrice of the nourishing cord.

When the embryo is directed towards the hilum, the
vessels go directly from it to the chalaza, which is then
confounded with the hilum ; but when the embryo is
directed in the contrary way, the chalaza is very dis-
tinct from the hilum, and the umbilical cord is prolonged
through the mesosperm from the hilum to the chalaza;
this course of the fibres is called the Raphe, and appears
externally as a kind of little nerve. The umbilical cord
is a prolongation of the carpellary fibre, which bears the
seed, and is itself prolonged into the raphe ; the cha-
laza is the true hilum, that is to say, the point where
the embryo draws its nourishment from the mother
plant; but it is often difficult to determine its position,
both on account of its small size, and of the change of
position of the embryo during the growth of the seed.

There are some Monocotyledonous seeds, in which the
radicle causes by its position a little projection on a given
part of the spermoderm ; and at the period of germina-
tion it shoots out from this part ; it is doubtful whether
it be an organ properly so called, seeing the small num-
ber of plants in which it is found.

Section III.

Of the Albumen.

If we examine an ovule at the period of flowering,
we find its spermoderm already well-formed, and its
canty filled with a mucilaginous fluid, to which the

STRUCTURE OF SEED. 193

name of Amnios is given. This liquid is, or may be,
transmitted into the cavity of the seed by the umbilical
cord, or more probably is secreted by the endopleura.

As soon as fecundation has taken place, the embryo,
which swims in the amnios, begins to be developed ; it
occupies more space, and the amnios consequently di-
minishes; it is probably absorbed by the embryo, to
which it may serve for nourishment, or it may be re-
absorbed by the neighbouring organs : whatever becomes
of it, it happens after a certain time, that, in some
plants, the whole of it disappears, and the embryo alone
occupies the cavity ; in others, the more fluid part of it
alone disappears, but its solid particles are deposited,
and solidify into a particular body, to which many
different names have been given, but I prefer that of
Albumen, both because it makes allusion to the albumen
of eggs, and because the name is applicable to this
substance in its being of a white colour in all seeds.

The albumen is less an organ, properly so called, than
the residue of one, or a deposit formed in cellular
tissue ; it does not present, at least at the maturity of
the seed, any organic connexions either with the endo-
pleura or embryo ; we do not perceive in it any vascular
organization, but only a mass composed of cellular
tissue, and not adhering to any of the neighbouring
organs : if this be not the case perhaps in some of the
Coniferae and Cycadeae, where there is a slight adhesion
between it and the radicle. What has often induced
persons to speak of the albumen adhering to the spermo-
derm, is, that several naturalists have described as
albumen, the endopleura when it is thick and fleshy : it
is this that happens in those Leguminosae in which the
existence of albumen has been admitted. The seed in
which the nature of this body can be followed with the
greatest facility is the immense one of the Cocoa-Nut

VOL. II. o

194 VEGETABLE ORGANOGRAPHY.

Palm, of which the albumen occupies so large a portion.
The Cocoa-Nut, in its young state, is filled with a watery
fluid, which soon becomes of the consistence of an
emulsion, and then takes the name of milk ; it is at this
period that it is made use of as a beverage ; soon, the
solid part suspended in the emulsion is deposited, and
concretes upon the walls of the seed in a state which
does not bear a bad resemblance, in consistence, to our
Almonds, and which is edible as they are. Finally,
this amygdalaceous deposit hardens and forms an albu-
men with a slightly oily flesh which lines the walls of
the seed; the centre, at first occupied by a watery fluid,
is converted into an empty cavity by the evaporation and
absorption of this water. What I have described in the
Cocoa-Nut, takes place in all other seeds where albumen
is formed, except that this substance usually fills the
whole cavity without leaving any space, and its size,
form, nature, and position vary in different plants.

In a seed of a given size, the volume of the albumen
is essentially in an inverse ratio to that of the embryo.
The families in which the albumen is larger in proportion
than the embryo, are the Palmas, Liliaceae and neigh-
bouring families, Euphorbiaceae, Nyctagineae, Rubiaceas,
Umbelliferae, Ranunculacena, &c. ; it is met with, but in
much less proportion, in the Convolvulaceae, Violarieas,
&c. ; it exists only in certain genera in the Labiatae ;
lastly, it is completely wanting in the Cruciferae, Legumi-
nosae, Compositae, &c. The families which have no
albumen, have the embryo large ; in those furnished
with it, the embryo is sometimes of extraordinaiy
minuteness ; thus, in Ranunculaceae and Umbelliferae,
it frequently presents only a small niche in the albumen,
near the base of the seed. In general, we must remark
that it is hardly ever absent in any family of Monocoty-
ledons ; the Alismaceas alone appear devoid of it, whilst

STRUCTURE OF SEED. 195

its entire absence is frequent in Dicotyledons ; of the
families of this class, about a third are constantly devoid
of it.

The nature of the albumen presents great variety in
the different families, and a remarkable constancy in
each of them.

1st. One of its most ordinary states is that of being
fleshy, as is seen in the Cinchonaceae. This fleshy state
degenerates into a firm and almost woody texture in
some families, as in the Umbelliferae.

2d. It is often oily, as is seen in several Palms, and
especially in the Euphorbiaceae, in which we remark
that their embryo is also impregnated with oil, but that
the nature of these two fixed oils is different; that of the
embryo is acrid, as is the entire plant, of which the em-
bryo is a miniature representation ; that of the albumen,
which is a particular secretion of an organ, is in general
bland and wholesome, although more or less laxative.

3d. It is frequently feculent or farinaceous, as is
observed in the Caryophylleae, Nyctagineae, and espe-
cially in the Gramineae ; for it is the albumen of the
Cereal Gramineae which serves for the principal nourish-
ment of the human species.

4th. We find horny albumens, such as that of the
Asparageae, the Coffeaceao, &c.

All known albumens are of a wholesome nature, to
whatever family they belong ; those of the Euphorbiaceae
are laxative ; their properties are also similar in all those
which have an analogous consistence. Thus, all farina-
ceous albumens are sensibly of the same nature ; the
farinaceous albumen of the Polygoneae, for example,
may be substituted for that of the Gramineae : all horny
ones present some analogy with that of the Coffee ; thus
those of Galium and Ruscus, when roasted, have the
odour of Coffee.

o2

190 VEGETABLE ORGANOGRAPHY.

The general form of the albumen is moulded upon
the inner cavity of the endopleura, and is modified by
that of the embryo. In general, it is all in one mass ;
but there are some genera of the Rubiaceae, such as
Rutidea and Grumilea, where it is presented under the
appearance of little portions detached from one another.
It presents in some plants, and especially in all the
Anonaceae, a character which is very remarkable, viz. —
the endopleura is wrinkled or prolonged into projecting
laminae, so that when the albumen is cut lengthways, it
seems furnished upon its margins with little transverse
lamellae, a very remarkable character which Mr. Robert
Brown has met with in the anomalous genus Eupomatia,
which he discovered in New Holland.

The position of the albumen is always in the empty
space left by the embryo, which is in general more or
less central, and then the albumen surrounds every part
of it ; the embryo is sometimes lateral, or situated at the
base near the chalaza, and then the albumen occupies
the rest of the cavity, as in Portulaca, the Marvel of
Peru, &c. : lastly, when the embryo is peripherical, then
the albumen is found in the centre, as, for example, in
several Polygoneae and Chenopodeaa. There are seeds
among the Malvaceae and Bombaceae, in which the albu-
men is reduced to a small farinaceous deposit, lying
between the cotyledons.

The use of the albumen has not yet been perfectly
studied ; it is evident that the amnios, especially when it
is absorbed, must serve to nourish the embryo, and that
the albumen must serve to nourish the young plant at
the period of germination. We see, in fact, that most,
if not all albumens are transformed at this period, by the
addition of the water which they absorb, into an emulsion-
like matter, which is absorbed by the embryo, and serves
to develop it ; but the details of this phenomenon have

STRUCTURE OF SEED. 197

not been sufficiently observed, and I ought the more to
abstain from mentioning them here, since they belong
entirely to Physiology.

Section IV.
Of the Embryo.

The Embryo is the object and end of the whole func-
tion of sexual reproduction. It is a young plant in
miniature, already provided with every organ essential
for nutrition, — with a root, which at this age bears the
name of Radicle : with a stem, which receives from
analogy the name of Cauliculus, or more usually that
of Plumule ; and lastly, with leaves themselves, to
which, seeing that their appearance differs much from
others, the name of Cotyledons has been given. Let
us examine the embryo at its three ages — 1st, whilst it
is in the state of an unfecundated germ ; 2d, during its
state of torpor in the fecundated seed ; and 3d, during
the changes it receives by the act of germination.

The first of these articles will be short ; for the em-
bryo is hardly visible before fecundation : as soon as it
can bo perceived it appears very small, floating in the
water of the amnios. It is possible that the radicle
draws its nourishment, either from the hilum or from
the amnios, without having any organic connexion with
either, and by simple absorption, analogous to that by
which the roots draw the nourishing fluids from the
earth. It is possible that at this period of its life, the
extremity of the radicle communicates by a vascular
filament with the umbilical cord, and by this filament it
may receive its fecundation and nourishment; but it has

198 VEGETABLE ORGANOGRAPHY,

not as yet been seen by anatomists in a positive manner ;
if it exist, it is probably destroyed before maturity. Is
not the filament which has been seen by Richard going
from the hilum to the radicle in the Cycadeee and Coni-
ferae, that of which I have been speaking, which in these
families may be more persistent than in others ?

When the embryo has not been fecundated, or when,
having been so, some particular cause has arrested its
development, we may suppose two cases possible — either
the whole ovule may become abortive on account of the
abortion of the embryo, and then the seed is wanting in
the place where it ought to be found ; this is most fre-
quently the case ; or the integuments of the seed may
continue to be developed, so that the seed appears per-
fectly formed externally, but internally it is found
empty : when it has albumen, this is often formed then
as usual ; but the place of the embryo is vacant. Thus
it is not rare to find seeds of the Coffee well-formed as
to the spermoderm and albumen, but having the cavity
of the embryo empty. There are some cases in which it
is difficult to affirm which of these abortions has taken
place ; thus, for example, if we examine the fruit of
Ranunculus lacerus, a hybrid plant, we shall find the
seeds well-formed externally, but empty within; is it the
seed which is abortive in the carpel, or the embryo in
the seed ? In this particular case I believe that the seed
is entirely abortive, because the albumen is absent ; but
if it took place in a plant without albumen the question
would be insoluble ; this happens in Centaurea hybrida,
the achenia of which are empty, without our being able
to affirm whether the spermoderm exists or not.

The fecundated embryo generally grows very rapidly ;
the water of the amnios gradually disappears, either
wholly, or, its fluid parts being absorbed, the solid resi-
due concretes into albumen. In the first case, the seed

STRUCTURE OF SEED. 199

or embryo is said to be Exalbuminous, and the embryo
is sometimes called Naked or Epispermic; in the con-
trary case, the seed or embryo is said to be Albumi-
nous or Endospermic. It must be observed that the
word Perispermic signifies, in the nomenclature of
Jussieu, that there is albumen, and in that of Richard,
that there is none ; an example which, in the midst of
numbers of others, proves the inconvenience of changes
of names.

When the radicle is directed towards the hilum,
which, as we know, is the base of the seed, we say that
the embryo is Erect, or that the radicle is Inferior,
or directed to the base ; when it does not point to the
hilum, it may be either lateral as in the Coffee, or
directed upwards, it is then Superior, and the embryo
Inverted, as in the Doom-Palm; this is the only
exact sense in which the position of the embryo can be
designated. But carpologists have often extended these
terms in another sense; they have frequently referred
the direction of the embryo, not to the seed, but to the
fruit ; so that when a seed is pendent in the fruit, all
the terms which designate the position of the embryo
must be generally taken in a contrary sense.

Whether the embryo be inferior or superior, it may
be erect, curved, or folded upon itself. In the first case,
if it be long, it occupies the axis of the seed, and is
named Axile, as in Sjwndias, Empctrum, See. ; if it be
short, it only occupies a small portion of the axis ; and
it is said to be Basal if at the base, as in Ranunculus;
and Apicilar if at the apex, as in Clematis. Similar
differences take place in curved ones ; they are usually
lateral, and situated at the side of the seed ; if they be
equal in length to it, or shorter, they proceed from their
origin to the other extremity, and are then said to be
Curved, as in different species of Polygonum, ; if their

200 VEGETABLE ORGANOGRAPHY.

length exceeds that of the seed, they return by the
other side toward the base, and are then termed Peri-
pherical, as in the Spinach ; if they be still longer, they
may describe one and a half, two, or three turns, and
are then called Spiral, as in Dodorup.a. As to those
which are folded upon themselves, I cannot explain their
structure before having spoken of the parts of the
embryo.

The Radicle (radicula) is that part which represents
the root ; in most Dicotyledons, it is presented in a
conical form, very like that of ordinary roots ; it goes
on gradually tapering from the neck to its extremity,
which is pointed ; at the period of germination it
elongates by its extremity, as other roots do during the
whole course of their lives, and makes lateral shoots,
but very slowly. To plants thus organized Richard
has given the name of Exorhize/e, because their
radicle is, as it were, projecting and developed : on the
contrary, in all Monocotyledons and some Dicotyledons,
as Berberis, Nuphar, &c, the radicle is thick and
rounded at the extremity ; it scarcely elongates at the
period of germination, but gives origin, either laterally,
or from its apex, to some usually simple little roots,
which act the part of radicles, and sometimes seem to
issue out of the round radicle by particular slits ; this
structure has caused Richard to give to plants, thus
organized, the name of Endorhize^;. This has been
for a long time observed in Wheat, Rye, and Barley,
and more attentive observation has proved that a great
number of other plants partake of the same structure.

It frequently happens in the embryos of the Endor-
hizeEe, either that the radicle, from which the little
roots ought to proceed, is very thick, and, as it were,
capitate, and the\| the embryo is said to be Macro-
podous, as in Pekea ; or, that one of the lateral parts

STRUCTURE OF SEED. 201

of the radicle takes some unusual development, and
then this kind of tubercule has been confounded with
several others under the name of Vitellus ; or that
the radicular extremity, being arrested in its elongation,
becomes reflected upon itself, and forms a kind of
perfectly close sac, which surrounds all the embryo, this
has received the name of Saccule, and is seen in the
Nymphaeaceaa. Thus, this distinction of plants into
Exorhizeas and Endorhizeae, which would seem to
promise a new confirmation of the natural division of
the two great classes of Phanerogamese, is found reduced
to a remarkable phenomenon it is true, but which
cannot serve for the purposes of classification.

When at the germination of the Exorhizeae we
cut the extremity of the radicle at the moment it pro-
trudes from the seed, it is transformed, as it were,
artificially into an endorhizeous one ; that is to say,
we oblige it to produce lateral radicles, much more
than it was naturally intended to do. The distinction
of Exorhizeae and Endorhizeao, which seems so decided
at first sight, becomes less pronounced when we
examine the intermediate cases ; thus, the common
Radish presents below the neck two laminae applied
upon the root, which are kinds of Coleorhizaa, for they
are the remains of a kind of sheath which the radish
has pierced through or torn, so then we might say that
the Radish is an Endorhizeous plant, which shoots out
but one radicle.

Radicles, whatever be their form, are often furnished
at the moment of their development with particular
hairs which are of a silvery-white colour, long, bristly,
but of a very soft texture, and short duration ; they
arise principally from near the neck, and always in the
parts exposed to the air ; their particular use is not yet
well known, but their existence is especially remarkable

20.2 VEGETABLE ORGANOGRAPHY.

in the roots being in general devoid of true hairs, to
which, in order to designate their situation, I have given
the name of Radical Hairs. — (Book i. ch. 10, sect. 7.)

The radicle of the embryo, whatever be its form, is
perceived — 1st, in the seed before germination, because
it is always directed outwards — a very important charac-
ter to observe in Monocotyledonous plants, where it is
sometimes the only one by which we can readily dis-
tinguish the two extremities of the embryo ; 2d, after
germination ; because, with a very small number of
exceptions, as the Misletoe, the radicle is directed
towards the centre of the earth. This direction is so
decided that it presents itself under the most different
circumstances. We observe in particular, that whatever
be the position of the germinating seeds, the radicle is
always directed downwards, and if one more or less
developed be turned to the zenith, it always tends to
return of itself, so as to resume its natural position.
The cause of this phenomenon is an object of delicate
physiology, which will be discussed elsewhere ; I only
mention the fact here, as a distinctive character of ger-
minating radicles.

The Plumule (plumula) is, as we have said, the stem
of the embryo or young plant, already present in the
seed or at germination ; it is distinguished, whatever be
its form, by characters opposite to the preceding, viz.
because in the seed it is directed inwards ; at germina-
tion, because it has a tendency to rise towards the
zenith, becoming green when exposed to the light, and
presenting all the other characters of stems. It ma}*
be divided into two parts, which Richard has called the
cauliculus and the gemmule.

The Cauliculus is that part of the plumule which is
between the neck and the cotyledons ; the Gemmule
(gemmula) that which is above the cotyledons; in

STRUCTURE OF SEED. 203

embryos which have no apparent cotyledons, as the
Dodder, these parts are confounded.

The existence of the cauliculus has often been denied,
because this organ is sometimes so short, as to be scarcely
distinguishable ; but as the cotyledons always spring
from the stem, the great or small distance from their
point of origin to the neck, can always be perceived.
This length of the cauliculus also appears to be of little
importance in the symmetry of the young plant ; thus, in
the Papillionaceae, the French-bean presents it nearly
two inches long, and in the Pea, it is so short as to be
hardly visible. Its length influences, at the period of
germination, the situation of the cotyledons — whether
they be elevated above ground, or on a level with the
surface or subterranean ; these three manners are met
with in the Leguminosae, and the example is of itself
sufficient, independently of all other reasoning, to prove
that we cannot derive the primary divisions of plants
from these characters, as Willdenow has proposed. The
cauliculus is always simple, even in plants which become
the most branched ; almost always devoid of leaves, even
when they are very numerous near the root ; the branches
and leaves only begin to be developed above the cotyle-
dons. I only know one exception to these rules —
Euphorbia, which has sometimes buds upon the cauli-
culus below the cotyledons.

The existence of the cauliculus, which is so evident
at germination, tends to prove that we ought not, as
some naturalists have done, to confound the neck, pro-
perly so called, i. e. the plane of separation between
the root and stem, with the point of origin of the coty-
ledons. These two points in the French bean are
nearly two inches apart, and never exactly coincide.

The gemmule, or the part of the stem which is ele-
vated above the cotyledons, is truly the first bud of the

204 VEGETABLE ORGANOGRAPHY.

plant ; there are seeds where it is presented under the
appearance of a little sharp point, hardly visible ; there
are others where it already has small leaves visible, to
which the name of Primordial Leaves is given. In
general, the gemmule, on developing at the period of
germination, presents all the appearances of a young
branch which proceeds from a bud furnished with leaves.
In some cases, such as Cactus Melocactus, Euphorbia
Canariensis, and in general, in succulent plants with
very small leaves, it is very thick, fleshy, round, and
devoid of leaves : in this case it has sometimes been
taken for a single cotyledon ; but on observing it more
closely, we find the two cotyledons very small it is true,
and, as it were, concealed in the mass of the gemmule.
Embryos, with this kind of gemmule, have received the
name of Macrocephalous ones.

The name of Cotyledons, as we have said above, is
given to the first leaves of the plant already formed, and
visible in the seed. That they are leaves is easily proved :

1st. By their usual transformation into leaves, and
becoming green at germination.

2d. By their respective position, similar, or analogous
to that of the plant already developed.

3d. By their structure, exactly resembling that of
leaves ; and because they are generally furnished with
vessels, and stomata distributed in the same manner.

4th. Because, in plants where the leaves present
peculiar phenomena, such as the mobility of those of
the Sensitive-plant, or the presence of glands in those
of Hypericum, or Gossypium, (PI. 23, fig. 1), &c,
the cotyledons present the same characters.

5th. Because their development, death, and fall, are
analogous to what takes place in ordinary leaves.

6th. Because the cotyledons are absent in plants
devoid of leaves, as Cuscuta.

STRUCTURE OF SEED. 205

7th. Because, when they are opposite they bear in
their axils as opposite leaves, sometimes a single ter-
minal gemmule, at other times, three small ones ; a
central one, which is the prolongation of the cauliculus,
and two axillary ones.

8 th. Finally, because the analogy of the radicle to
the root, and of the plumule to the stem, having been
proved, that of the cotyledon to the leaves, is an evident
consequence.

The principal difference that plants present, as to
their cotyledons, results from the respective position of
those organs which are distributed in two manners :
the first takes place when two or more are situated
upon the same horizontal plane, and then they are
opposite or verticillate ; as the first case is by far the
most frequent, we give to the whole class of plants,
with two or more cotyledons, the name of Dicotyle-
dons. When it is necessary to express positively, that
the cotyledons are verticillate, we call the plants Poly-
cotyledons, but they cannot be considered as a class.
In fact, 1st very similar plants present both these
manners ; thus, Pinus (PI. 23, fig. 2) and Abies among
the Coniferae are polycotyledonous, and the other genera
of the family dicotyledonous ; 2d. also in polycotyledo-
nous genera or species, the number of cotyledons is not
regular ; 3d. in several, they are disposed in two opposite
parcels, which seem to refer to the primitive type of the
class ; 4th. in all verticillate cotyledons, as in all, or
nearly all verticillate leaves, we only find two opposite
buds, besides the central prolongation of the stem ;
5th, some species with two cotyledons accidentally pre-
sent three or four, as I have seen in the French bean,
Ranunculus, and Cabbage ; 6th, lastly, I have above
said (Book II. chap. iii. sect. 7,) that the distinction be-
tween opposite and verticillate leaves is very uncertain ;

206 VEGETABLE ORGANOGRAPHY.

therefore it is with just reason that all plants with oppo-
site or verticillate cotyledons have been classed together :
perhaps a more applicable name than that of Dicotyle-
dons might have been given to them ; but it must be
observed that it is not the number which is essential,
but their respective position. Although I do not attach,
as one sees, as great importance to the number as the
term Dicotyledon would lead one to believe, I ought to
notice, however, notwithstanding what has been said,
that I have never seen Ranunculus, Fumaria, Sfc. ger-
minate with one cotyledon, the binary number peculiar
to the class being always met with.

The second system of arrangement of the cotyledons
is, that where the leaves being naturally and essentially
alternate, the lower one or cotyledon is solitary, and
consequently lateral. The plants or embryos in which
this disposition takes place, bear the name of Monoco-
tyledons ; this term is more correct, in general, than
the preceding, but still anomalies are met with here and
there. In most monocotyledonous embryos the cotyle-
don or lower leaf is alone so large and developed as to
be visible in the seed ; but it frequently happens that
along the gemmule we observe other little bodies analo-
gous to the cotyledons, and situated alternately : these
are, properly speaking, primordial leaves ; if this name
be given to them, the plant might be said, in a strict
sense, to be monocotyledonous; but if they be called se-
condary cotyledons, which their appearance and nature
might authorize, it must then be said that they are
Monocotyledons with two or more alternate cotyledons ;
thus here also it is the position and not the number
which serves for the character of classification. Cycas
is the only really doubtful case which may be mentioned
as tending to impair the separation of the two great
classes of Vasculares ; we here find two cotyledons more

STRUCTURE OF SEED. 207

or less unequal, but they are not strictly opposite, the
smaller one arises a little above the larger, and, conse-
quently, the embryo, although having two cotyledons,
belongs to the class of Monocotyledons, which con-
firms the whole structure of the stem and fructifying
apparatus.

We call those plants Acotyledons, which are devoid
of cotyledons ; but under this name one may comprehend
two very different organizations ; — 1st, The Nemea of
Fries, or cellular plants, which are all considered as
Acotyledons, although in most the germination is ill
understood ; — 2d, Among vascular plants, those which
are devoid of cotyledons, and which Fries names exclu-
sively Acotyledons, are generally also devoid of leaves:
such are Cuscuta, and Orobanche, which, notwithstand-
ing, we are obliged to class among Dicotyledons, and
perhaps some leafless Orchideae, which do not less belong
to Monocotyledons. The only known example of a vas-
cular plant furnished with leaves, and apparently devoid
of cotyledons, is Lecythis, the singular germination of
which Du Petit Thouars has described. We must con-
sider this seed as formed of two fleshy cotyledons, united
with each other and with the plumule, and which do
not separate at germination. In several succulent plants
the cotyledons are so small, as in Cactus Melocactus,
&c. (PI. 23, fig. 3,) or so adherent to the cauliculus, as
in Stapelia, that we might think them absent, from a
superficial observation.

Next to their position, that which most distinguishes
cotyledons from each other, and which causes the great-
est difference in their history, is the presence or absence
of stomata, which is always connected with their tex-
ture. All those which, when developed, present stomata
on their surface, are more or less foliaceous, and of a

208 VEGETABLE ORGANOGRAPHY.

green colour; they then take the exclusive name of
Seminal Leaves.

All those which have no stomata, remain in a fleshy
or farinaceous state, and do not become green; these are
usually only called fleshy cotyledons.

Foliaceous ones are very frequent among Dicotyle-
dons, and are hardly ever found in Monocotyledons, but
they exist in Ferns. Fleshy ones are common in Mono-
cotyledons ; they are also met with among the Dicoty-
ledons in the tribes Phaseoleae, Viciae, &c. of the Legu-
minosae, in Hippocastaneas, Trapa, the Chestnut, &c. ;
foliaceous cotyledons are, in general, more frequent in
albuminous Dicotyledons ; and fleshy ones in those which
are exalbuminous ; the reverse would seem to occur in
Monocotyledons.

Foliaceous cotyledons being furnished with stomata,
can, from the moment they are exposed to the air,
elaborate the sap which is transmitted to them by the
radicle, and, consequently, it is not necessary that they
should be provided with a store of nutriment prepared
beforehand for the young plant ; fleshy ones, having no
stomata, cannot elaborate the sap, and there would be
no action if they were not filled with a quantity of fecula
or mucilage, which, diluted by the water absorbed by the
radicle, is thus transformed into nourishment. We may
say, then, that the cotyledons nourish the young plant,
when they are foliaceous by elaborating the sap in the
same manner as leaves, and when they are fleshy by fur-
nishing nutriment prepared beforehand, as in the albu-
men or tubercules ; whence it results that every organ
which has no stomata, and is not fleshy or filled with
fecula, is not a cotyledon, an important character which,
in certain doubtful cases, elucidates the nature of
organs ; for example, it has enabled me to deter-

STRUCTURE OF SEED. 209

mine the character of different parts of the seed of
Nuphar.

It must be observed, that as germination has mos"
frequently been studied in the French-bean, Wheat, and
other plants with fleshy cotyledons, that which is only
true as regards this class, has, in general, been too
readily extended to all.

Foliaceous cotyledons are, on account of their nature,
always destined to come out of their envelope, and also
above ground, at germination ; but it is not the same
with fleshy ones ; the first come out of their integu-
ments as the French-bean, the second continue in them,
and remain underground, such as the Pea, Vetch, Horse-
chestnut, &c. As most monocotyledons have the coty-
ledon fleshy, it is conformable to analogy that it should
be subterranean, which in reality is the case.

A curious consequence which results from the nature
of the two kinds of cotyledons which I have mentioned,
is, that the fleshy ones alone are those which man makes
use of for food ; he takes for his own use the deposit of
nourishment which the mother-plant had prepared for
its offspring, in the same manner as in the eggs of birds :
it is thus that the seeds of the Leguminosa? with fleshy
cotyledons, such as Beans, Peas, Lentils, Cajanus, &c.
serve for the food of man, whilst those with foliaceous
ones are useless or dangerous. There are no apparent
exceptions to this rule but in albuminous seeds : it is
then the albumen, which is itself a deposit of nourish-
ment, that man makes use of; thus Buckwheat (Poly-
gonum) is edible on account of its farinaceous albumen ;
the Gramineae have at the same time a fleshy cotyledon
and farinaceous albumen, a double circumstance which
contributes to place them in the first rank among nutri-
tive seeds.

The forms of foliaceous cotyledons are as variable as

VOL. II. P

210 VEGETABLE ORGANOGRAPHY.

those of leaves; most, however, are entire; but some
are lobed either at the apex, as in Helicteres, Convol-
vulus, the Radish, &c, or at the base, as in Polygonum,
&c. ; some are divided, being palmate, as in the Lime-
tree, or pinnatifid as in the Erodium Pimpinellifolium.
But the principal difference which they present in this
respect consists in the lateral cotyledons of phaneroga-
mous Monocotyledons being almost all sheathing at the
base, whilst this form is rare in Dicotyledons ; this cor-
responds very well with the ordinary form of the leaves
of the two classes.

Among Dicotyledons, it happens from time to time
that the two cotyledons are found united ; this union
takes place laterally, unequally, and purely accidentally,
in a great number of plants — for example, in Ebenus
Cretica or Tithonia; it is constant and regular, in some
united at the base, so that they seem to form a kind of
disc pierced by the stem, as is seen in several Ficoids.

Fleshy cotyledons are generally of a more irregular
form, and frequently united together by their whole
inner surface ; this is seen in the seeds of the Horse-
chestnut, Nasturtium, Eugenia, See, which at first sight
might appear Monocotyledons, because their two coty-
ledons are united into a single mass.

The inequality of the cotyledons, which is rare and
entirely accidental in Dicotyledons, where they are foli-
aceous, is not very rare in those where these organs are
fleshy ; the seeds which I have mentioned present good
examples: but in this respect the seed of Trapa deserves
mention. In Trapa natans, which is so well known in
Europe under the name of Water-chestnut, the two
cotyledons are extraordinarily disproportionate; they are
exactly opposite, as in all Dicotyledons, but one is so
small, that it must be carefully sought for before it can
be .perceived, and it comes out without difficulty -with

STRUCTURE OF SEED. 211

the rest of the embryo, by a small circular hole, from
the envelope formed by the spermoderm invested with
the pericarp and calyx ; the other is very large, fari-
naceous, borne upon a long petiole, and remaining in the
spermoderm, the whole of which it fills ; the first, which
is rudimentary, is almost useless, and does not furnish
any nourishment ; the second provides the radicle with
all the nourishment which serves for its development ;
whence it results that the side of the root which cor-
responds to the great cotyledon increases much, and
gives rise to a great number of little roots ; whilst the
opposite side, which corresponds to the small cotyledon,
does not produce any little roots, and, as it remains
very short, draws towards it the whole body of the
root.

The cotyledons, considered as to the manner in which
they are folded or rolled up in the seed, present as many
varieties as leaves in the bud ; considered as to the
position of the radicle, they are sometimes continuous
with it, as in erect embryos, and sometimes curved or
folded upon it, — a character which is more frequently,
but perhaps less correctly expressed, by saying that the
radicle is curved or folded upon the lobes. Dicotyledons
present two very evident varieties among folded em-
bryos ; thus, sometimes the radicle is folded back upon
the cotyledons, so as to he in the fissure between them ;
this is seen in all the Papilionaceous Leguminosa? and
Cruciferao Pleurorhizeae ; in this case we say that the
radicle is Lateral, which is indicated by the sign o=;
or the radicle is folded upon the back of one of the
cotyledons, and it is then said to be Dorsal, which is
expressed by the sign o|| : this is seen in the Cruciferao
Notorhizeae.

If we consider the respective position of the cotyledons
themselves, we shall find several varieties : —

p 2

2\2 VEGETABLE ORGANOGRAPHY.

1st. The greatest number of plants present the coty-
ledons absolutely flat, that is to say, without any curve
or folding ; such are those of Cytisus, Ricinus, Arabis,
&c. ; this form is compatible with every position of the
radicle.

2d. There are some which are folded longitudinally
upon their middle nerve ; these always have the radicle
dorsal ; such are, for example, the Cruciferse Ortho-
ploceaa ; these cotyledons are said to be Conduplicate,
and are designated by the sign o^.

3d. There are some curved or rolled spirally, in a
longitudinal direction, as, for example, those of the
Combretaceae — Puyiica, Helicteres, &c.

4th. Some are bent double, as in the Cruciferae Di-
plicolobiae, such as Heliophila.

5th. We find cotyledons flat, but rolled crossways one
upon the other, as, for example, in the Cruciferae Spiro-
lobeae, such as Bunias.

6th. Lastly, there are some which are irregularly
plaited or crumpled upon one another ; such are those
of the Mullow.

This kind of character does not appear connected in
an intimate manner with the symmetry of plants, since
there are families where we find united several of these
different foldings of the cotyledons ; the Cruciferaa, in
particular, present five different systems.

As leaves, some cotyledons are furnished with petioles,
others are sessile ; the former, as well as several of the
latter are, as it were, articulated at the base, and fall off
soon after germination ; there are, however, annual
plants, where they remain until the flowering, as some
species of Veronica and Galium, &c. Sheathing coty-
ledons, or even simply sessile ones, are more permanent,
or only destroyed in part. Those of several succulent
plants are particularly remarkable for their permanence;

STRUCTURE OF SEED. 213

thus, Euphorbia Canariensis still presents remains of
them at the end of one, or even two years.

We have not as yet any certain proof of cotyledons
furnished with stipules, if this do not happen in Trapa
natans, where the ascending twin filaments, which are
observed near the base of the stem as far as the origin
of the cotyledons, appear to be true stipules.

The primordial leaves which we see sometimes fully
developed in the seed with the cotyledons, as in the
French-bean, or which are developed immediately after
them, are always of an analogous nature to the true leaves
of the plant ; but they often differ — 1st, in form ; thus,
those of the French-bean are simple and heart-shaped,
instead of being trifoliate with oval leaflets ; the differ-
ence, however, rarely proceeds as far as this ; 2dly, in
size, which is usually less ; 3dly, in position, which, in
Dicotyledons, is opposite, or nearly so, even in species
where the leaves become alternate as the plant advances
in age ; sometimes the change takes place suddenly, as
in the French-bean, which has the first two leaves oppo-
site, and all the others alternate ; at other times gra-
dually, so as to show that the alternate position is a
simple degeneration, owing to the mode of development.
The contrary takes place in Monocotyledons, which have
their leaves alternate. It results from this, that when
we see a plant with the lower leaves opposite, we may
be pretty certain that it belongs to Dicotyledons ; and if
we find one with them alternate, there is great pro-
bability that it is a Monocotyledon.

214 VEGETABLE ORGANOGRAPHY.

CHAPTER V.

OF THE ORGANS OF REPRODUCTION" WITHOUT FE(
DATION IN PHANEROGAMOUS PLANTS.

I said, on commencing- to treat of the organs of
reproduction, that all organized beings appeared to be
reproduced by the development of pre-existing germs.
Are these germs, as Bonnet maintains, bodies existing
in infinite numbers from the commencement of the
species, inserted into one another and destined to be
successively developed under favourable ciicumstances ?
Or, are they productions successively formed by vital
action, or, as has been said, by the plastic forces of
individuals, so as to pre-exist only a short and definite
time before the period when their development is visible?
It is hardly necessary to discuss this question as regards
the subject upon which we are at present engaged. It
will be sufficient to admit that there exist in different
parts of plants, germs which are developed after two
manners : one kind requires the particular act of fe-
cundation, and forms the seeds of which we have been
examining the structure, &c. ; the other kind, in order
to be developed, only requires the concurrence of certain
circumstances purely relating to nutrition.

Among these last, there are some, which, without
any preparatory apparatus, are developed when the
nourishment becomes more abundant in a given place :

ORGANS OF REPRODUCTION'. 215

this phenomenon is purely physiological, and can hardly
be placed in Organography; thus, when we make a
notch in the bark of a tree, a swelling, that is to say a
deposit of juices, is caused there by the stagnation of
the sap, and the latent germs are developed with fa-
cility at this point; this fact is only connected with
Organography in these respects : —

1st. That every species presents fixed points where
certain developments of germs readily take place.

2d. That certain species present particular points
where there is naturally a stagnation of the juices and a
deposit of nourishment, and where, consequently, the
germs are already visible in the natural state or more
easily developed.

As to the first respect, we remark that the axils of the
leaves are the principal of these fixed points in all
plants where, by the ordinary progress of vegetation,
there is facility for the development of the germs of
branches : this is what happens in the natural course of
things, forming ordinary buds.

As to the second, there are plants which present
naturally, here and there, articulations or transverse
swellings, which retain the sap, form deposits of nou-
rishment, and consequently favour the development of
germs ; such are the articulations of Pinks, Vines, and
Geraniums, the nodes of the Graminea?, &c. There
are others which form, here and there, kinds of ex-
ostoses or tubercules filled with a quantity of fecula, and
which have a tendency to cause the germs situated upon
their surface to be developed ; such are Potatoes, &c. :
the germs appear upon these tubercules as opaque slightly
fleshy points, to which the name of Eyes is frequently
given. Every one knows, at least from the popular
example of the Potatoe, that these eyes or germs, when
separated from the feculent part of the tubercule and

216 VEGETABLE ORGANOGRAPHY.

placed in favourable circumstances, can develope and
produce a new individual ; but we also know that this
development is more ready and vigorous, when we leave
around each the whole or part at least of the nourish-
ment which had been previously collected for it. Thus,
developments of this kind are favoured by the nourish-
ment accumulated in the tubercules, but may take place
by the forces peculiar to the germ which attracts the
surrounding water. There are, in fact, other tubercules
where we find the germ furnished with a very slight
provision of nourishment ; such are those which arise
from the roots of Saxifraya granulata, and the little
bulbs which are accidentally or constantly developed in
the axils of the leaves of several Liliaceas, and even in
the axils of their spathes, and which ma}r indifferently
be considered either as buds or tubercules.

There are cases where the germs exist almost without
any provision, appearing under the form of dots, but
ready to be developed under favourable circumstances;
such are the points visible in the notches of the leaf of
Bryophyilum Calycinum, and which are developed
when this leaf, having become old, happens to touch the
moist earth.

When the tubercules, which bear germs, become
detached from the plant which has given birth to them,
we can easily imagine that the double circumstance of
their being isolated bodies, closed on every part, and
capable of producing a new individual when sown, has
caused them to be taken for seeds ; this happens in
Ficaria, for example, where the development of these
bulbs has been described as a true germination.

This error is the more excusable, as there are cases
where it is really difficult to elucidate the truth, and
where we observe remarkable affinities between seeds
and tubercules.

ORGANS OF REPRODUCTION. 217

Thus, we find several species of Crinum and Ama-
ryllis, in which the cells of the fruit, instead of enclosing
seeds in the usual state, each contain only a thick,
round, fleshy body, on which we remark a small eye ;
this body is detached from the pericarp at maturity, and
when sown produces a new individual. Is this a tubercule
or bulb, as it is generally said to be ? Is it a seed,
modified in texture, as some modern botanists think ?
In order to embrace with any confidence either of these
opinions, we must first of all know what essential
difference is found between seeds and tubercules. Cannot
the same germ, according to the state of its developments,
either require fecundation, which is the ordinary case,
or not require it, and then be developed under the form
of a tubercule or bulb ? This supposition seems to
acquire some weight if we reflect that the germs of
Bryophyllum are placed in the leaf precisely as the
ovules in the pericarp, and, consequently, appear to be
of the same nature. A second example, as curious
though less clear, is presented to us in Lemna; the ordi-
nary mode of reproduction in this plant, is the develop-
ment of a lateral germ situated upon the margin of the
foliaceous disc which composes the whole plant; this
germ, on developing, forms a second foliaceous disc,
united to the first, but which afterwards separates and
forms an entire plant. When these plants happen to
flower, which is very rarely the case, the flowers are
found precisely in the same point where the germ
usually is ; whence we may suppose that this germ can
be developed, according to circumstances, with or without
fecundation.

Lastly, we shall see in the following Chapter, that
there are Cryptogamous plants, in which it is perfectly
impossible to affirm whether their development results
from a true fecundation, or from circumstances purely

218 VEGETABLE ORGANOGRAPHY.

regarding the nutrition. If the identity of nature of
these germs, developed with and without fecundation,
could be completely demonstrated, it would be a power-
ful argument against the system of Epigenesis. I must
explain myself: — among the naturalists who have studied
the theory of animal generation, there are formed two
opposite classes : — some, -such as Haller, Bonnet, and
Spallanzani, maintain that the germ exists perfectly
formed, before fecundation, in the female organ, and
only receives from the male organ its vital action. The
others, as Needham, &c, have thought that the germ
exists in the male organ, which transmits it to the
female, which only serves as a matrix for it. The recent
observations of Prevost and Dumas seem to give force to
this last opinion, although, in fact, all that is known can
be explained in both theories. When we wish to apply
these considerations to the vegetable kingdom, we are
asked if the little granules, which are perceived in some
fovillas, do not enjoy an analogous function to what is
attributed to the spermatic animalculas ; but notwith-
standing the numberless facts which prove the existence
of the ovules before fecundation, and the continuity of
the embryo with the mother-plant, it is evident that if
the unfecundated germs are developed in the same
manner as fecundated ovules, we must conclude that
they are produced by the female organ, and only owe
their vital action to the male.

The reproduction of plants by simple division, or,
which is to say the same thing, by unfecundated germs,
is an universal phenomenon, and all plants appear
capable of this mode of multiplication. Vegetable
fecundation, some philosophers have said, is a useless
phenomenon, since all plants have another mode of
reproduction, and consequently, we ought not to admit
it. ¥c may respond to this kind of argument : — 1st,

CRVPTOGAMOUS PLANTS. 219

that the fecundations of animals capable of division must
also be denied, whilst there are several where the two
modes of reproduction are very certain ; 2d, that it is
very true that all plants can be reproduced without
fecundation, but in most it requires the hand of man to
cause this phenomenon ; that therefore all phaneroga-
mous plants, which are not either creeping, rooting, or
furnished with tubercules, i. e. three-fourths of all known
plants, would be deprived of all natural reproduction, if
the germs of their flowers were not vivified by fecun-
dation.

It remains then from this observation, that reproduction
by unfecundated germs is common to the whole vegetable
kingdom, a very remarkable circumstance when com-
pared with the animal kingdom ; but that this form of
reproduction requires a concurrence of physiological
circumstances which are rarely found in the state of
nature, and that fecundation is the natural form which
replaces it, and thus insures the perpetuation of the
species.

CHAPTER VI.

OF THE ORGANS OF REPRODUCTION IN CRVPTOGAMOUS

PLANTS.

Section I.

General Considerations.

As soon as we began to study with any care the
structure of the flower and fruit of plants, we imme-
diately divided them into two great classes : — Phanero-

2L20 VEGETABLE ORGANOGRAPHY.

gamous ones, of which we have been speaking, and
Cryptogamous ones, which we are about to study in
this Chapter.

Some naturalists, struck with the extreme difference
of these two classes, and believing that all plants, which
did not present a flower formed as in ordinary ones, had
really no flower, but were reproduced by simple unfe-
cundated germs, have given collectively to these plants
the names of Agamm or Inembryone^e ; others, struck
with the fact that their reproductive bodies were formed
without apparent cotyledons, have designated them by
the name of Acotyledons. Some, admitting the ex-
istence, in these plants, of fecundating organs, but per-
ceiving their difference from those of phanerogamous
ones, have named them iETHEOGAMyE. Lastly, there are
some, as Gaertner and Borckhausen, who have desig-
nated them by the name of Aphrodites, in order to
make it understood that they have, it is true, fecundated
seeds, but that the fecundating fluid has no peculiar ap-
paratus, and is secreted by the same organs, or in the
same cavities, as those in which the ovules are found.

But all these terms, although admitted by distin-
guished naturalists, are less generally used than that of
Cryptogamous plants, which Linnaeus very felicitously
gave to this class of plants, and which is peculiarly
applicable to them. This term indicates that the organs
of fructification are not visible to the naked eye.

The name of Agamaa, which affirms the non-existence
of fructifying organs, and the absence of all fecundation,
probably expresses that which is not exactly true.

Perhaps it will one day be necessary to divide these
plants into two classes: — 1st. Cryptogamae, properly so
called, where fecundation is performed, although by
organs scarcely if at all visible to the naked eye ; and
2d, true Agamae, which have no fecundation : but if, in

CRYPTOGAMOUS PLANTS. 99 [

the present state of the science, we can affirm that the
first of these classes does really exist, it would be as yet
imprudent to declare that there are true Agamaa. We
can easily understand, in fact, 1st, that the fecundating
organ may have escaped, and still continue to escape
detection by our microscopes, and it may afterwards be
discovered ; 2d, that if the same cavity contain the germ
to be fecundated and the fecundating fluid, we might
not be able to see the sexual apparatus, and, notwith-
standing, fecundation may exist ; 3d, that as in phane-
rogamous plants there are some which are reproduced
with and without fecundation, it may happen in Crypto-
gamous ones, that these two modes of reproduction may
also exist, but that reproduction without fecundation
may be the most frequent.

The details into which we shall enter upon the dif-
ferent families of the Cryptogamae, will tend to prove
that these different reasons of doubt exist in several
cases; I regard it impossible, in the present state of
things, either to affirm that there are plants absolutely
devoid of fecundation, or that all are endowed with it.
I admit then the word Cryptogamous, in this sense, that
it designates plants in which the fructification is obscure,
or perhaps wanting.

The circumstance, which has most retarded the dis-
covery of the sexual organs of the Cryptogamae is, that
for a long time, these plants were only studied at the
period of maturity ; for it is clear that then we should
be no more able to find the male organs in them, than
the stamens in phanerogamous plants when their seeds
are ripe. It was the celebrated Hedwig who first made
this remark, and who has succeeded in finding the male
organs of several Cryptogamae, by observing them at the
period when they must be perceived, i. e. a long time
before maturity.

222 VEGETABLE ORGANOGRAPHY.

A second difficulty which also contributes to cast
doubt upon the structure of these plants is, that several
of them appear furnished at the same time with both
modes of reproduction ; thus, several Mosses and Hepa-
ticae present both reproductive bodies, which, proceeding
from a fecundating apparatus, must be considered as
seeds, and others which seem to be true bulbs. If the
distinction between these two classes of bodies is so dif-
ficult in certain phanerogamous plants, we can compre-
hend how the difficulty must go on increasing here,
seeing the smallness of the organs, and the almost
impossibility of applying the laws of analogy.

In fact, that which is most remarkable among Crypto-
gamous plants is, that the families of this class compared
together differ much more than the phanerogamous
families, and that the most nearly related present diver-
sities which would seem to declare a totally different
nature ; the laws of symmetry, which have so powerfully
aided us in discovering the true nature of the organs of
phanerogamous plants, can only be applied here in rare
and uncertain cases; this obliges us to study each family
in particular, without being able to derive from this ex-
amination any general laws for the entire class.

All Cryptogamas are furnished with bodies which
serve to reproduce the species in the same manner as
seeds. The name of Spores has been given to them, a
term which ought to be considered as provisional ; for,
when it shall have been proved that these bodies have
been fecundated, we must call them seeds, and if it be
demonstrated that they are not fecundated, they would
take the name of bulbs.

In most Cryptogamous plants, perhaps in all, the
spores are contained in a vesicle, or membranous cap-
sule, to which the name of Sporangium is often given ;
this organization is met with from the Ferns to the

CRYPTOGAMOUS PLANTS. 223

Algae, and it seems to be one of the constant characters
of these plants; the Sporangia are sometimes so small,
as to be taken for simple seeds, and the error is the
more easy, as they seem to germinate when placed upon
earth ; at other times they have been confounded with
the grains of pollen.

The most certain means of avoiding this last error, is
to observe the series of phenomena as well as their form.
The function of male organs is limited to the period of
fecundation, and it is observed in all known plants, that
the stamens perish after the emission of the pollen, and
usually fall off in a short time ; the female organs, on
the contrary, which have been fecundated, then com-
mence a new series of phenomena ; they increase in size,
become firm and opaque, and thus declare their true
nature. This system, simple, and founded at once upon
observation and reason, will enable us to recognise the
nature of the different powders which are observed in
the Cryptogamae: the more fugacious will be considered
as the male organs, the more lasting as female. We now
proceed to examine, in a general maimer, the structure
of the different families.

Section II.

The EquisetacecE.

We have seen, (Vol. I. p. 200,) on speaking of the
structure of the Equi^etaeea?, that their branches and
the scales, the union of which formed their sheaths, are
verticillate around the axis. This disposition is also met
with in their organs of fructification. Their stems, and
often their principal branches, terminate in an oval or

224 VEGETABLE ORGANOGRAPHY.

conical spike, composed of verticillate scales, each of
which is a disc with five, six, or seven angles, borne at
the centre upon a nearly cylindrical support. From the
lower borders of the disc are prolonged downwards
from five to seven whitish horns, which open by a longi-
tudinal fissure on the inner side, that is to say, nearest
the pedicel. There proceed from this slit, at the period
of maturity, globules which, received upon paper and
examined by the naked eye, present a kind of singular
spontaneous motion.

When placed under the microscope, we perceive that
each globule is formed: — 1st, of a green central body,
globular and compact ; 2d, of two laminae, dilated at
their two extremities into small club-shaped bodies,
placed crossways at the middle, at the base of a green
body, and rolled spirally around it ; these two laminae,
or four half-ones, are covered, especially at their swollen
extremities, with small red or browu corpuscules. They
are endowed with a very decided hygroscopic motion ;
they are rolled around the green body when moist, un-
rolled when they are ^dry, and evidently seem to serve
for the purpose of dispersing the green bodies out of the
horns which enclose them. What is the nature of these
different organs ?

Hedwig thinks that the green globule is an ovary, and
that the elastic laminaa are stamens, the pollen of which
is represented by the powder which adheres to their
surface. That the green globule is a true pistil, is what
Hedwig appears to affirm, asserting that in its young
state it presents a small point, which afterwards disap-
pears, and which he takes for the stigma. But is this
ovary furnished with a cavity containing several seeds,
as he appears to believe ? Agarclh and Vaucher invali-
date this opinion by the following observation : they
have seen that if these globules be plunged into water,

. CRYPTOGAMOUS PLANTS. 225

and if, after they have thus been made to swell up, they be
placed upon moist earth, they elongate and ramify, and
produce a young plant. This globule first gives birth to
articulated and confervoid filaments, analogous to those
which are observed on the development of the seeds of
Mosses.

The green globule then is a reproductive organ, but it
may be either a monospermous fruit, or a simple tuber-
cule analogous to a bulb. This last opinion would seem
confirmed by the fact, that in its development the foli-
aceous part does not appear to proceed out of an integu-
ment as in seeds, but that the seed itself appears to
dilate.

As to the elastic laminae, we cannot positively demon-
strate their nature ; on comparing them with stamens,
which their general position seems to authorize, it must
be confessed that several circumstances have been nesr-
lected : — 1st, We have no example of elastic stamens
which remain, without being obliterated, till the matu-
rity of the fruit ; 2d, We have no example of pollen in
the form of globules placed on the external surface of
the filaments ; 3d, If the laminae be filaments and
their inflated parts anthers, it is at least singular that
of four of these anthers two are placed on the side of the
globule, not at the point where the stigma is supposed
to be. Tn this state of circumstances I have great doubts
upon the reality of the character assigned to these
organs, and I am almost disposed to consider them as
simple elaters analogous to those of the Hepaticae, and
intended only to promote the dispersion of the green
globules ; and if these latter bodies be true fruits and
not tubercules, we might suppose that the fecundating
matter is contained with them in the horn or follicle
from which we see them issue.

VOL. II. Q

226 VEGETABLE ORGANOGRAPHY.

Section III.

Of the Marsileacea or Rhizospermce.

The Marsileacegg are of all Cryptogamous plants those
in which we most easily distinguish the sexual organs.
Most of them have their parts of fructification enclosed
in a kind of close involucrum, which appears divided
into several cells. We may count four in Pilularia, and
one in most species of Marsilea ; in each of these cells
or distinct cavities, we find unilocular sessile anthers,
which contain yellow globular pollen, and pistils also
sessile, formed of an oval ovary surmounted by a small
stigma ; these ovaries change into a monospermous inde-
hiscent fruit. At germination, the seed gives birth,
first to a radicle, and a leaf, the number of each of which
afterwards increases, and they finish by forming a small
bundle of roots and leaves. Bernard de Jussieu does
not hesitate to consider them as Monocotyledons,
near Ferns, on account of the circinate vernation of
their leaves. Salvinia, also presents a closed involucrum,
which encloses the male filiform organs surrounding a
solitary ovary, surmounted by a sessile stigma, and in-
closing several seeds. These three genera are specially
organized to live in water and inundated places, since
the organs of the two sexes are contained in the same
closed envelope, so that the pollen may fall imme-
diately upon the stigma. Azolla, which Mr. Brown
refers to this family, differs from the other genera in the
male and female flowers being contained in different
involucra.

CRYPTOGAMOUS PLANTS. 221

Section IV.
Of Ferns.

The Ferns have sometimes been called by the name
of Dorsiferous or Epiphyllospermous plants, making
allusion to one of their most striking characters, viz :
that their fructification arises in general upon the back
of their foliaceous organs, which may be considered
either as true leaves, bearing, by a structure peculiar to
this family, the organs of fructification, or as peduncles
bordered with foliaceous limbs.

In favour of the first opinion, we may allege that these
foliaceous organs are not always fructiferous, and that
some Ferns bear their fructifications in spikes which seem
distinct from the leaves, that they present absolutely the
use and structure of true leaves, and are provided with
stomata ; that, lastly, there are some phanerogamous
plants, as Polycardia, where we find a somewhat similar
structure.

We may say, in favour of the second opinion, that the
leaves without fructifying organs owe this state to an
abortion analogous to that of peduncles transformed into
tendrils or spines ; that the pretended spikes of certain
Ferns are only peduncles which are not bordered ; that
certain phanerogamous plants, as Urtica membranacea
and Paspalum membranaceum, present peduncles bordered
in a similar manner ; that if petioles can be bordered
with a foliaceous limb furnished with stomata, pedun-
cles can also present the same singularity ; and, lastly,
that the example of phanerogamous plants with epi-
phyllous flowers are all doubtful when closely examined.

It is, perhaps, in order not to decide this question, that

q 2

228 VEGETABLE ORGANOGRAPHY.

several botanists have given to the leaves of Ferns the
vague name of Fronds.

Whatever term may be preferred for these peduncu-
late-leaves, or foliaceous-peduncles, we shall observe
that their position upon the stem is similar to that of
leaves, and that they may in like manner be divided into
petioles and limbs furnished with nerves and parenchyma.
Although the limb is often very much divided, there is
never any articulation between its parts, and it must
always be compared with simple leaves.

In general, those fronds which do not bear the fruc-
tification, as those of Osmunda, are large and foliaceous
throughout their whole extent. This appearance is met
with in all those which bear a moderate number of fruc-
tifying organs, for example, Polypodium, Pteris, Sec.
But when the number of these organs is very great, then
the foliaceous limb diminishes and seems to disappear,
being covered or hidden by the development of fruits ;
this is seen clearly in several species of Acrostichum;
and following analogies which leave but little doubt, we
come to understand that Ferns said to be spiked, as
Ophioglossum, only owe this appearance to a more com-
plete and constant abortion of the foliaceous limb. Let
us also remark that in the cases where the organs of
fructification are moderately dispersed upon the fronds,
these latter still being able to perform the physiological
functions of true leaves, may be all fertile, as Polypodium,
Pteris, Sec. ; whilst, if the fructifying organs are accu-
mulated in great numbers upon certain fronds, so as to
deprive them of all the functions of leaves, it then hap-
pens that there are upon the same stem other fronds,
said to be sterile, which perfectly enjoy the functions of
leaves ; this is seen in Osmunda, Ophioglossum, Sec. The
vernation of the fronds of all or nearly all Ferns is cir-
cinate, or in other terms, they are rolled up from the

CRYPTOGAMOUS PLANTS. 229

apex to the base ; this disposition, analogous to what is
observed hi the Droseraceae and Cycadese, is remarked
not only to affect the middle lobe of the leaf, but also
each of its partial lobes. It results that at the moment
of expansion the upper surface is entirely external, and
the lower is protected by being thus rolled up.

When we examine Ferns under the microscope, at
this period, we find, along the middle, oval, pedicellate,
naked, scattered little bodies, which Hedwig considered
to be stamens; and upon the part of the limb which is
rolled up, we observe other bodies, more numerous, and
concealed under a proper membrane. These last are,
doubtless, the rudiments of young fruits, for we can fol-
low their development on to their maturity.

As to the first, the opinion of Hedwig is founded
upon the following reasons: 1st, they are only found at
a period long before maturity, and disappear soon after-
wards ; this is the property of the male organs of plants:
2d, their form and appearance are analogous to [those
of ordinary male organs : 3d, if this character be not
assigned to them, we shall be puzzled to attribute any
other to them.

In answer to these arguments it may be objected : —
1st, that these bodies have as yet been seen in only a
small number of Ferns ; 2d, that their position is uncer-
tain, and very different from that of the female organs ;
two circumstances which are contradictory with regard
to the character assigned to them ; 3d, that the female
organs being covered over by a membrane, we do not
see in what way fecundation can be performed ; 4th, that
observers have not as yet perceived the dehiscence of the
organs which Hedwig considers to be males ; 5th, that
nothing appears to act the part of a style, or stigma, in
those which he regards as females.

In opposition to the opinion of Hedwig, some natura-

230 VEGETABLE ORGANOGR Al'HY.

lists have thought that the capsules of Ferns were kinds
of bisexual flowers. Muratti was the first who main-
tained the hermaphroditism of the flowers of these
plants. Hill and CEder have thought that the ring of
the sporangium was the organ which contained the
fecundating fluid. Gartner and Mirbel have main-
tained that each of the globules contains, in its young
state, the fecundating fluid and the ovules. This opinion
is founded, not upon direct observation, which would be
impossible, but upon the analogy of the Ferns with the
Marsileaceffi, and upon the presumed necessity of fecun-
dation.

Lastly, Bernhardi has expressed a new opinion upon
the nature of the sexual organs of these plants : he
thinks that the male organs are little bodies of a glan-
dular appearance, sessile upon the small scales which are
observed upon the upper surface of the leaves ; that the
ovaries of the female organs, situated in clusters on the
lower surface, have kinds of styles which pierce the
tissue of the leaf, and terminate in the pores on the
upper side, performing the part of stigmata. In support
of this opinion, he remarks that the points which he
supposes to be globules of pollen, are situated at the ex-
tremity of vessels which appear stronger, and conse-
quently more important than the neighbouring ones;
that the scales are first of a yellowish brown colour,
afterwards pale, and fall off, as anthers do ; that the
globules of pollen may pass along the upper surface of
the leaf, as far as the points which appear to perforin
the function of stigmata. Bernhardi found species where
the organs which he regards as males, are situated upon
different leaves ; such as, for example, Onoclea Stru-
thiopteris and 0. crispa, the sterile leaves of which are,
according to him, the males; but in the greatest number
of observed genera, as Poly podium, Polystichum, Cyathca,

CRYPTOGAMOUS PLANTS. 231

Davallia, Asplenium, &c, the male organs are upon
the same leaves as the capsules.

The theories of Hedwig and Bernhardi are at present
the only ones which deserve attention, but I will not
venture to say any thing upon the preference which
ought to be given to either of them : that of Bernhardi
presents, it is true, less objections ; but the facts upon
which he rests are known in so small a number of
species, and have been observed so little in detail in
those where they have been described, that it appears to
me premature to adopt them without new examinations.

As soon as the leaf is developed, the male organs
(I here speak collectively in the sense of the two hypo-
theses) disappear, and the female organs begin to in-
crease. We see them gradually become elevated, and
afterwards, at maturity, they break the pellicle which
covers them ; the clusters bear the name of Sori, and
their integument that of Indusium; the disposition of
the clusters upon the limb or margins of the leaf, and the
existence or form of the indusium, are the principal cha-
racters from which the classification of Ferns is derived :
let us examine now the bodies of which the clusters are
composed.

These bodies, at maturity, are of a brown or reddish
colour, round or reniform, and furnished with a short
pedicel. The name of Capsules, or more correctly that
of Sporangia, is given to them; they are most fre-
quently bordered with an elastic ring, which opens from
within outwards, and causes the dehiscence of the cavity ;
in some anomalous Ferns the ring is absent, and the
dehiscence takes place by a transverse rupture ; almost
all have the capsule unilocular ; but in Myriotheca it is
divided into several cells.

From this cavity there proceeds, at dehiscence, a small
cloud of powder, which is composed of seeds or spores ;

232 VEGETABLE ORGANOGRAPHY.

these are little corpuscules, usually round, and reddish-
brown ; when sprinkled upon a sponge or moist earth we
see them evidently germinate, and reproduce the species
which gave birth to them.

The seed or spore produces laterally a green body, at
first nearly cylindrical, but which afterwards expands
into a foliaceous limb, devoid of nerves, very similar to
those of certain Hepaticae, and which may be considered
as the cotyledon ; it sometimes becomes lobed at the
apex, sometimes it surrounds the base of the plant, so
that the following fronds appear to proceed from its
centre. It frequently shoots out radicles, either from
its margin or lower surface, and sooner or later is
destroyed, as is the case with the cotyledons of phanero-
gamous plants. It only remains, for the complete assi-
milation of these organs with cotyledons, to be assured
whether the foliaceous parts proceed from an integument,
or whether they are a simple prolongation of the globule.
The extreme minuteness of this body has not as yet
permitted this to be accurately observed, but the ana-
logy of this germination with that of Mosses, in which
Hedwig asserts that he has seen the rupture of the inte-
gument, ought to make us believe that it will also be
observed in Ferns.

There are ferns which are called viviparous, because
young individuals are seen to arise from the margins of
their leaves, or from the centre of their clusters of fruc-
tifications. This phenomenon may be compared, either
to the development of embryos in Bryophyllum, or to
the germination in the pericarp which is observed in
certain species of Cuscuta. The Ferns in which this
takes place are Darea, Asplenium bulbiferum, A. ramosum,
Cyathea bulbifera, &c.

CRYPTOGAMOUS PLANTS. 233

Section V.
Of the Lycopodiacece.

The family of the Lycopodiaceas, although not nu-
merous in species, is one of those, the structure of which
is the most difficult to be understood. The diversity of
organs which are found, either collected together, or
separated in the different groups of the family, is the
principal difficulty met with in this study.

The only species which may be regarded as sufficiently
known, is Lycopodium denticulatum, very well described
by Mr. Brotero, and figured by Mr. Salisbury in the
Transactions of the Linnean Society; this species joined
to L. Helveticum, forms a genus or particular section, to
which the name Diplostachyum, proposed by Beauvois,
may be retained, although the character is hardly cor-
rect. These species present two kinds of spikes upon
the same plant, or a single spike, which encloses two sets
of organs in the axil of the bracts. We find in the upper
part of these spikes, slightly crustaceous, reniform, bi-
valve bodies, full of an angular, yellowish or orange
powder. Mr. Brotero thinks that this organ is an anther
full of pollen, and he affirms that he has sown it without
his ever having seen it germinate. Beauvois adopts the
same opinion. At the base of the spikes or on the
shorter ones, borne upon the same plant, we find in the
axils of the bracts other bodies, which are also crusta-
ceous, and which open by four lobes, and contain four
yellowish globules, marked at their base with three pro-
jecting ribs ; these globules are seeds, for, in the midst
of a great number which were abortive, Messrs. Brotero
and Salisbury have seen some of them germinate ; con-
sequently the shell with four lobes which contains them

234 VEGETABLE ORGANOGRAPHY.

is an ovary ; according to the former, the stigma of this
ovary is represented by a pellucid and transverse line,
placed at the apex ; the same character is attributed to
the little central protuberance winch surmounts this
line.

At the period of germination, we see the young plant
come out of the seed by the side ; its radicle is simple
and perpendicular; its plumule rises vertically, and is
terminated by two opposite leaves, from the axils of
which spring two branches. Inside the seed is an oily
body adherent' to the embryo, which Mr. Brotero calls
the Vitellus, and which appears to me to be the true
cotyledon ; the two opposite leaves, which these ob-
servers call Cotyledons, represent, in my opinion, the
primordial leaves. The change of character, assigned to
these organs, appears to me sufficiently authorized, both
because the vitellus is an almost imaginary organ, and
because it is impossible to compare Lycopodium with
Dicotyledons.

From well known facts with regard to Lycopodium
dcnticulatum, we may easily conclude: — 1st, that in L,
Selaginoides, or the section Selaginella, the bivalve
shells which Hedwig described as the female organ are
the male, and the quadrivalve and reniform ones which
he described as males are the female organ : — 2d, that
it is the same in the sections Gymnogynum and Stachy-
gynandrum of Beauvois, although their structure has not
been as well studied ; but what is the character of the
shells which are observed in the Lycopodiums compos-
ing the sections Plananthes, Lycopodium, and Psilotum
of Beauvois ; that is to say, in all the Lycopodiaceae,
where but one class of organs is known ? Beauvois con-
siders them always as males, and regards the females as
unknown. Linneus also regarded the powder of these
shells as analogous to pollen, on account of its inflam-

CRYPTOGAMOUS PLANTS. 235

mable nature. This opinion would seem confirmed by
the extreme analogy which is observed between the bi-
valve shells of the section Plananthes, compared with
those of the section Selaginella, which analogy obliges
us to consider as males ; the contrary opinion has been
maintained by Hedwig, but to support it he has been
compelled to admit for the male organs of these plants
kinds of foliaceous buds which do not resemble any
known male flower. I admit then, with little doubt,
that the bivalve shells of Plananthes and Lycopodium of
Beauvois are male organs, of which we do not know the
females ; but I am much more uncertain with regard to
the nature of the shells with three valves in Psilotum,
although the globules contained in them appear to in-
close fovilla rather than an embryo.

I have met with two organs analogous to those of Ly-
copodium in the genus Isoetes, which might be defined
by the name of Lycopodium aquaticum. Having had an
opportunity during my stay at Montpellier to see a living
plant, I attentively examined it : the leaves spring
from a kind of fleshy subterranean stem, slightly analo-
gous to those of bulbous plants. Each of them bears in
its axil a fructifying organ, or a flower which is adherent
to it ; in those which may be considered as the lower
ones, we find a membranous indehiscent body concealed
by a small foliaceous lamina, surmounted by a filament,
divided internally into three compartments by small
transverse columns, and enclosing about fifty spherical
globules, marked at their base with three projecting
ribs, as the seeds of Lycopodium denticulatum. In the
axil of the central leaves we find other bodies very simi-
lar to the preceding, but which are divided internally
into more numerous compartments, which contain an
impalpable powder, at first white, afterwards black.

If I had caused either of the two powders which I

236 VEGETABLE ORGANOGRAPHY.

have described, to germinate, the history oflsoetes would
have been completely elucidated ; but having quitted
Montpellier before I could do so, I am still in doubt
with regard to the nature of these two organs. On the
one hand, the extreme similitude of the three-ribbed
globules with the bodies which Brotero has seen ger-
minating, leads me to consider them as seeds ; but I have
always found them empty at the period of maturity,
which would seem to indicate that they are of a male
nature ; moreover, the powder of the central capsules,
which becomes brown and opaque at maturity, would
seem more to represent seeds.

Gaertner and some other naturalists have considered
the Lycopodiaceae as devoid of sex, and furnished with
two kinds of seeds ; but this hardly probable opinion
must be confirmed or destroyed by experiments upon
the germination of the two kinds of powder.

Section VI.
Of Mosses.

Mosses are more distant from phanerogamous plants
than the preceding families, since they are devoid of ves-
sels and stomata ; they present, however, various affini-
ties with these plants in their organs of fructification,
which are better known than in any other cryptogamic
family. Hedwig has so extended the field of our know-
ledge with regard to Mosses, that, neglecting former opi-
nions, I shall limit myself to the exposition of his, and
only examine the later doubts and objections to the
works of this philosopher.

CRYPTOGAMOUS PLANTS. 237

The fructifying organs of Mosses are contained in
kinds of buds, placed sometimes at the tops of shoots,
sometimes laterally, and at other times even at the base
of these shoots : those which are really terminal some-
times seem lateral on account of the elongation of the
shoot after flowering. These buds, either star-like or
capitate (for their appearance may cause these different
names to be given to them), are formed of leaves imbri-
cated without any order, and the number of which does
not appear fixed. This envelope has received the name
of Perichcetium when it is found at the base of the
pedicellate fruits of Mosses, or, in other terms, around
the female organs ; and the name of Perigone, when it
surrounds the male organs. These two terms, although
generally admitted, appear to me to rest upon ideas
hardly correct ; in fact, it seems evident that these leaves
constitute the same organ, and the frequent cases where
they cover at the same time both the male and female
organs, are sufficient to demonstrate it ; moreover, when
the organs of the two sexes are always separate, which is
not the case here, one would not be more authorized to
give two names to their integuments than to the calyx
or involucrum of the two sexes of dioecious phaneroga-
mous plants. If we ought to admit but a single name
for identical organs, that of perichcetium ought to be
rejected ; that of perigone is less incorrect, but it also
presents a great objection.

This term, already admitted for phanerogamous
plants, supposes that the floral bud of Mosses is a sim-
ple flower, and this, in fact, is the opinion of Hedwig,
and of almost all Muscologists. Others have thought
on the contrary, that the floral bud of Mosses is a
true capitulum formed of several flowers. I would wil-
lingly compare it with the compound flower of Euphor-
bia. The leaves which surround the fructifying organs

238 VEGETABLE ORGANOGRAPHY.

appear to me to be a true involucrum with several leaf-
lets, inclosing sometimes the male, sometimes the female
flowers, and more rarely the two sexes together.

The leaves of the involucrum, or the bracts of Mosses,
differ from the ordinary leaves nearly as the bracts of
phanerogamous plants, either in size, form, or even in
colour; frequently the middle nerve is absent when the
other leaves are furnished with it. At other times they
are prolonged into a long bristle, which is wanting in the
ordinary leaves ; sometimes those of the two sexes or
those of different rows differ from one another ; but
these leaflets are never verticillate as in the perigone or
calyx, but always imbricated as in involucra.

In these capitula, whatever be the sex of the organs
they contain, are found an indefinite number of simple
divided filaments. Hedwig has given them the name of
Paraphyses ; they are most frequently cylindrical, and
longer than the sexual organs ; we find some which are
gradually thickened towards the apex, and others abruptly
dilated into an oval or globular kind of club. They
usually spring from very near the base of the sexual
organs. They have been compared to the nectaries of
flowers, but I have already shown in what a vague sense
this word was used ; I should be inclined to compare
them with the little scales found in the involucra of
Euphorbia, and to regard them either as bracteoles, or as
the rudiments of a true perigone. They continue for a
long time without changing their form, and their func-
tion, with regard to the fructification, is entirely un-
known.

The male organs of Mosses are found scattered among
the paraphyses in an indefinite number : they entirely
compose the male capitula of the monoecious or dioecious
species, and surround the female organs in the herma-
phrodite capitula. Each of them is composed of a very

CRYPTOGAMOUS PLANTS. 239

short and sometimes scarcely visible filament, and of an
oval or oblong sac or anther : this sac presents no trace
of a suture, and is unilocular ; the apex presents a glan-
dular point, by which, at maturity, the cell opens, and
we see it emit, in intermittent jets, a viscid fluid. Does
this represent the fovilla contained in the grains of pol-
len ? or, do the pollinic globules here swim in a parti-
cular liquid? This liquid, as well as all the male or-
gans, is of a greenish colour ; after it has been emptied,
the half-dried sac takes a yellowish or brownish colour,
and the cellular organization is visible to the microscope
under the form of a net-work. In a small number of
Mosses, Sphagnum palustre, for example, the anther is
large, oval, and borne upon a long filament.

The female organs of Mosses, considered at the time
of flowering, entirely compose certain capitula said to be
female flowers, and are found in the centre of those
called hermaphrodite ones. In both cases their number
varies from three to four, or even eight or ten ; but,
whatever be the number, there is hardly ever more than
one which arrives at the state of fruit; the others become
abortive, and are thrown oif under the form of scales
intermixed with the paraphyses.

Each organ is sessile or nearly so ; at the time of
flowering, we distinguish an ovary, usually oval and of
a reddish brown colour, a filiform style of the same co-
lour, and a slightly expanded stigma opening at fecun-
dation. All this apparatus is surrounded by a mem-
branous integument, which, after flowering, is elevated
by the elongation of the pedicel of the fruit, breaks at
the base, and receives the name of Calyptra, on ac-
count of its hood-like appearance. Hedwig considers it
as the corolla of Mosses, and this comparison is tenable,
provided that we only consider it as a short way of in-
dicating that it is an intimate integument of the flower;

240 VEGETABLE ORGANOGRAPHY.

but in the opinion which this author had that the capi-
tulum of Mosses is a simple flower, this term was con-
tradictory with regard to his own theory ; for each
female organ has its own calyptra ; and in that which he
called hermaphrodite flowers, he said that the corolla was
situated within the stamens. These difficulties do not
exist when the capitula of Mosses are considered as
aggregated flowers. The calyptra is the integument of
each of the female flowers, and may be compared to the
perigone of monochlamydeous flowers.

After flowering, the calyptra, elevated by the elon-
gation of the pedicel, is broken across near the base j
sometimes, as in Sphagnum j)alustre, the lower part
remains at the base of the fruit, but most frequently this
is not visible, and the calyptra remains placed as a small
cup at the top of the fruit, and falls off at the approach
of maturity. Sometimes the enlargement of the fruit
causes it to break laterally, at other times the pedicel
curves at the apex, so that the fruit is pendent, and the
calyptra drops off, which organ, at this period, is always
membranous or half-withered, resulting from its having
no longer any organic connexion with the plant ; it is
almost always smooth, but it sometimes bears hairs,
which appear to be the remains of paraphyses united
with it ; these hairs are directed upwards in Oligotr'i-
ckum and Orthotrichum, and downwards in Polytrichum.

The pedicel, which was so short at the period of flow-
ering as to be hardly visible, afterwards elongates so as
often to be longer than the stem ; it is a true thecaphore,
and is slender, simple, cylindrical, firm, and composed of
compact elongated cellular tissue. The name of Pedi-
cel or Bristle, (seta,) is given to it.

The Theca, which terminates the pedicel, is the true
pericarp ; its form is most frequently oval, sometimes
either tapering or swollen at the base, or slightly pro-

CRYPTOGAMOUS PLANTS. 241

truding laterally. It opens at maturity by a true circu-
lar dehiscence, which takes place near its apex; the
upper part, which resembles the cover of a vessel, has
received the name of Operculum ; it is slightly depressed
at the margins, and conically elevated at the centre.

After the fall of the operculum, we see that the inner
border of the theca is furnished with one or two mem-
branes terminating in regular teeth ; these membranes
bear the name of Peristome (peristoma), because they
surround the opening of the theca. The peristome,
when there is but one, or the outer one when there are
two, is very remarkable for its diversities of form and its
irregularity ; in a small number of cases it bears no
teeth, as in Gymnostomon ; most frequently it is bor-
dered with teeth or cilia, which are always equal and
of the number four, or one of its multiples, four in
Tetraphis, eight in Splachnum, sixteen in Grimmia,
thirty-two, forty-eight, or sixty-four, in different species
of Polytriclium. In several cases each tooth is half-
divided by a fissure, as in Dicranum, and in the same
case where it is not found, we perceive traces of it under
the form of longitudinal lines. We might believe that
the number of teeth is as great in the normal state, and
that they are united two and two, three and three, four
and four, &c. The inner peristome only exists in a part
of the Mosses ; it is more membranous ; its margin is
divided into 8, 16, or 32 teeth, which are often more
\ unequal and irregular than those of the outer peristome.

In some genera, as Polytriclium, the tops of the teeth
of the peristome are all united together by a transverse
membrane stretched over the entrance of the theca ; this
membrane bears the name of Epiphragm ; when it
exists, the seeds can only come out between the teeth.
In almost all the other genera the teeth are free, and are
endowed with a very decided hygroscopical motion ; they

VOL. II. R

242 VEGETABLE ORGANOGRAPHY.

are curved inwards when they are moist, and outwards
when dry ; by means of this motion they raise up the
operculum, and facilitate the dispersion of the seeds.

The centre of the theca is occupied by a vertical axis
called the Columella, which proceeds from the base
and reaches the top of the operculum, to which probably
it bears nourishment ; it is sometimes cylindrical, at
other times slightly swollen in the middle ; its apex is
obliterated at the fall of the operculum.

The seeds or spores are very numerous, attached,
according to Hedwig, to the walls of the theca and to
the columella ; they are very small, reddish or brown at
maturity, and of a globular or round form. Hedwig has
seen those of several species germinate; from his account,
the integument breaks, and the young plant presents at
its birth a descending filament, which might be taken
for a radicle, and a cylindrical divided body, which ap-
pears to be a kind of cotyledon; there are afterwards
developed kinds of cylindrical and branching primordial
leaves, the number of which is uncertain ; they remain
for a long time in certain species, as Phascum Confer-
voides. Mr. Drummond, who has observed the germina-
tion of Mosses since Hedwig, asserts that these confervoid
filaments penetrate the earth and form roots.

This theory of the reproduction of Mosses, although
hitherto universally admitted, has not been free from
contradiction : some have begun to deny facts upon
which there is at present the most accordance ; others,
admitting the structure of the organs, have denied the
use assigned to them, more, it appears to me, from
general opinions upon the absence of sexes in Crypto-
gamse, than from the real examination of facts. The
principal positive objection has been, that it is difficult
to conceive how the female flower, invested with its
calyptra, can be reached by the matter emitted bv the

CRYPTOGAMOUS PLANTS. 243

anthers, and especially in the dioecious capitula; but
nothing prevents our admitting (and several say that
they have seen it) that at this period the calyptra is
slightly open at the apex, or we might believe at least
that there is some direct communication with the stiarma.
It has also been said, that the mode of fecundation
described above is impossible in aquatic Mosses; but
Hedwig has remarked that when they flower, which is
very seldom, the tops are then raised above the water.

Beauvois has maintained, that the whole reproduction
of Mosses is performed in the theca alone, partly found-
ing his opinion upon the very small number in which
male organs have as yet been perceived. He thinks that
the seeds of Hedwig were the pollen, and that the true
seeds were inclosed in the columella. As an objection to
this theory, has been urged the improbabihty of finding
the pollen at the same time as the ripe seeds, so per-
fectly resembling them in size and form, and in so great
a quantity; it has been especially answered by the germi-
nation of the pretended grains of pollen. Finally,
Mr. Robert Brown appears to have found the cause of
the illusion of Beauvois; when the theca is cut trans-
versely, the knife carries with it into the columella some
of the seeds, which are those which he believes are
lodged there ; but when the columella is either cut
lengthways, or after having completely isolated it, we
find nothing but cellular tissue devoid of seeds.

Besides the sexual reproduction which we have de-
scribed, Mosses are also propagated by shoots which
proceed from the trunk, and taking root, form new indi-
viduals. This mode is common in aquatic Mosses, and
those of very damp places, which rarely flower.

B -'

244 VEGETABLE ORGANOGRAPHY.

Section VII.
Of the Uepaticce .

The family of Hepaticaa, though very natural, presents
forms too dissimilar for it to be convenient to describe it
in a collective manner ; it will be more clear to speak
successively of the small number of genera which com-
pose it, beginning with those which have most affinity
with Mosses.

The Jungermannia?, which form the most numerous
genus of this family, have been unfortunately described
by Linneus, who designated their fruit by the name of
anther, and confounded under the name of female flowers
the true male flowers, and the gemmules. Schmidel was
the first to clear up this difficult subject ; Hedwig has
confirmed and extended his observations, and Dr. Hooker
has thrown a new light upon it in his excellent Mo-
nograph.

The Jungermanniae are all monoecious ; the male
flowers are presented under the form of whitish an-
thers, solitary, sessile or nearly so, oval or ovate, com-
posed of a fine reticulated membrane, full of pollen, and
situated along the nerves of the leaves, or more rarely
scattered upon the disc. Dr. Hooker has made them
known in more than forty species of this genus. These
anthers are usually naked, sometimes surrounded by
leaves analogous to an involucrum or perigone. The
female flowers are produced in very different situations;
they are almost always surrounded by a foliaceous or
membranous calyx, or perigone, sessile upon the stem or
the leaf, and most frequently of a single piece, tubular,
or slightly dentated at the apex ; it is only absent in a
very small number, as Jungermannia concinnata, and

CRYPTOGAMOUS PLANTS. 245

J. Hookeri; it is found double in /. Lyellii and /. Hi-
bernica. Each calyx incloses from three to four, or ten
linear pistils, very like those of Mosses, and covered in
the same manner with a calyptra, which differs from that
of Mosses in its being broken at the apex ; consequently
it is not raised with the fruit, and it forms a kind of
membranous cup at the base of the peduncle, which, as
in Mosses, is but little if at all visible at flowering, and
elongates much and very rapidly at the approach of ma-
turity ; this peduncle is almost always of a whitish
colour, delicate texture, and formed of very elongated
cellules ; the theca or capsule is globular, brown, always
devoid of an operculum, and opens at maturity by four
spreading valves; it contains a great number of seeds
attached to filaments or linear lamellae, elastic, very hy-
groscopic, rolled up as a snail-shell, and most frequently
of a brown colour : they are called Elaters ; they
appear intended for the dispersion of the seeds, which
are spherical, brown, and opaque. Hedwig has seen
those of Jungermannia epiphylla shoot out, at germi-
nation, a simple radicle, and dilate at their upper part
into a leaf.

Besides these seeds, they have likewise almost every
kind of gemmae or bulbs which serve to reproduce them :
it also appears that the bodies collected into a compact
head, at the top of the leaves of some species, as Jun-
germannia nemorosa, and which Hedwig considered to
be male flowers, are nothing but collections of bulbs.

Marchantia does not differ from Jungermannia as
regards its fructification, except in the following circum-
stances:— 1st. The anthers, though resembling in form
those of Jungermannia, are collected together, and, as it
were, regularly inserted into an orbicular disc, nearly
flat, slightly wavy, and borne upon a long peduncle.
2d. The female flowers, organized as those of the Jun-

246 VEGETABLE ORGANOGRAPHY.

crermanniae, are sessile at the lower face of a star-like
pedunculate disc, and directed downwards ; the capsules
only open at the apex, by scarcely distinguishable teeth,
and the elaters are more slender. 3d. The bulbs are
more frequent, collected in kinds of sessile cups.

In Anthoceros the male organs are, according to
Hedwig, oval anthers, slightly pedicellate, collected three
or four together, in points scattered upon the disc of the
leaf, at first concealed under a pellicle which breaks, and
forms around them a kind of perigone. The female
flowers also grow upon the disc of the leaf; they are at
first presented under the form of a cone, they pierce it
through at the apex, and retain the remains at their
base, under the form of a sheath ; they afterwards have
an elongated bivalve capsule, which opens longitudinally,
and then presents an isolated filament situated in the
axis of the fruit. The seeds are spherical, slightly
bristly, and furnished with compressed laminae, which
appear to perform the part of elaters.

Targionia only presents a globular capsule surrounded
by a perigone ; the seeds are devoid of elaters. Sprengel
thinks that the male organs are corpuscules scattered
upon the membrane situated around the female flower,
and which perish before the maturity of the fruit.

Finally, Riccia only presents for the fruit, according
to Hedwig, kinds of univalve capsules, buried or sunk
in the leaf, surmounted by a small filament which seems
to be a style, and containing several ovules devoid of
elaters. The male organs are small whitish points,
sessile, and scattered upon the leaf near the margins of
its expansions. But the mode of reproduction of these
two last genera deserves to be studied.

Here ends the series of cryptogamic plants, where we
can recognise the sexes with any degree of precision.
In the following families we shall find no organs which

CRYPTOGAMOUS PLANTS. 247

can, with any likelihood, be considered as males, and if
fecundation take place, it is probable that the fecun-
dating fluid is contained in the same cavities as the
ovules, without having any apparatus of its own.

Section VIII.
Of Lichens.

Lichens, considered with regard to their fructification,
present kinds of discs or tubercules, which have received
the common name of Apothecia, and to which that of
Shields (scutellce), or Lyres (lyrellce), are given
when it is wished to designate their particular forms.
All these apothecia contain, at maturity, what appears
to be the true fruit, in which we find oval or globular,
opaque, blackish bodies, which appear to be the repro-
ductive corpuscules ; they have never, however, been
seen to germinate, but it is only by analogy, and not by
direct observation, that they are compared to seeds or
spores.

This point being almost universally admitted, one is
asked if these bodies have been fecundated, and if the
function of a male organ can be attributed to any
known part of Lichens. Some have thought that certain
farinaceous efflorescences which are observed on different
parts, were masses of pollen ; others have believed that the
nearly globular parcels of pulverulent matter, collected
at the extremity of certain lobes of the thallus, replaced
this function. Others have attributed it to kinds of
cavities hollowed out in the thallus, and where it is said
that a poliniform matter is found. Neither of these
opinions is founded upon proof, or even upon sufficient

248 VEGETABLE ORGANOGRAPHY.

probability; and we may object to all of them, because
the greatest number of Lichens are devoid of the organs
to which so important a function was attributed, and
consequently it is more probable that these efflorescences,
or warts, peculiar to certain species, are connected with
more general uses. Cassini has shown that the globules
collected at the extremities of the leaf of Physcia tenella
are capable of reproducing a new individual, and it is
likely that we ought to consider them as bulbs, and that
the globules analogous to these in other species are of
the same nature.

Those who, knowing the truth of these observations,
persist to admit fecundation in Lichens, have been com-
pelled to suppose, either that the pollen was produced
by the margin of the shields, which is usually rolled in-
wards at the period of the first development of these
organs, which must be supposed to be that of the
flowering, or that the fecundating matter is contained in
the cavities with the ovules. It is evident that nothing
can show either the truth or falsity of these opinions,
founded, not upon observation, but upon the theory of
analogy ; if they be true, the supposed reproductive
corpuscules of Lichens are real seeds.

Others, not willing to admit this, which it is possible to
see, and denying, perhaps imprudently, the existence of
that which does not fall under our senses, have unhesi-
tatingly decided that Lichens are devoid of male organs,
and of fecundation ; consequently some have called them
Agamae, others Anandrae, some Inembryones, and others
Acotyledons ; but all have been evidently guided by
an hypothesis — the non-existence of fecundation. If
this opinion be true, the reproductive corpuscules are
bulbs.

As I do not know any reason for admitting or reject-
ing either of the two theories which I have mentioned,

CRYPTOGAMOUS PLANTS. 249

it is convenient to give these corpuscules a name which,
if I may thus speak, will be entirely neutral ; those of
Spores or Gongyles have been proposed, as not affirm-
ing any thing beyond what appears known.

Section IX.

Of Fungi.

The immense family of Fungi presents forms so varied,
that I should greatly exceed the limits of this work if I
were to attempt to describe them. I shall confine my-
self to stating that all Fungi, taking this term in the
more extended sense that botanists have assigned to it,
present, at their maturity, globular coloured bodies,
which arc regarded as reproductive corpuscules, or in
other terms, spores or gongyles ; they are placed very
differently in the various tribes : — sometimes inclosed in
the body itself of the Fungus, as in the Truffle, &c. ;
sometimes situated on the surface, as in C Lav aria ; ;\t
other times between the lamellae, as in Agaricus, Bole-
tus, &c. &c. These are considered the reproductive
bodies, although they have never been seen to germinate.
We observe them in two states: — sometimes, as in Aga-
ricus, they appear in the form of an impalpable powder,
which separates at maturity from the membrane which
produces it, and which is called the Hymenium; in this
case they appear to be naked spores : sometimes, as in
Sphceria, we see them contained in a membranous en-
velope, which is a Sporangium, most frequently in the
form of a globe or oblong spindle. In both cases, the
spores or sporangia may be dry in cavities or upon sur-
faces which are not mucous ; sometimes, as it were,

250 VEGETABLE ORGANOGRAPHY.

immersed in cavities or upon surfaces which secrete a
particular mucus ; the sporangia, when they are not
imbedded in mucus, frequently adhere to divided fila-
ments.

As to the fecundating organs, there is still the great-
est doubt upon their existence. Bulliard has remarked
that in some species of Sphccria there exists, indepen-
dently of the sporangia of which we have spoken, a white
fugaceous inflorescence, which he supposes analogous to
pollen. Hedwig has believed that he has perceived,
both in the cells of Sphcsria and upon the margin of the
cap of Jgaricus, kinds of bodies filled with pulverulent
matter, and which he thinks are the male organs. But
neither of these assertions is founded upon observations
sufficiently positive, or extended to a sufficient number
of species, for it to be possible to repose any confidence
in them.

It may be said that the fecundating fluid is inclosed
with the spores in the sporangia, or around them in the
cavities which contain them. This may be so ; but those
who affirm it do not know more about it than those who
deny it. It would be premature, then, to give the least
importance to theories which are not founded upon any
positive facts. Until these are well known, if ever they
can be, we shall call spores or gongyles the fecundated
or unfecundated corpuscules, which we suppose, from
analogy, to reproduce these plants.

The practical multiplication of the mushroom ( Aga-
ricus campestris ) , which is performed by means of the
remains of old layers, or of what gardeners call mush-
room spawn, does not serve to elucidate the theory of
the reproduction of these plants ; in fact, in this rude
operation, they heap up without any order both the
remains of the roots as well as the caps of old mush-
rooms, and we may as well believe that reproduction

CRYPTOGAMOUS PLANTS. ^51

takes place by shoots or suckers, as by germination of
seed ; even admitting this last hypothesis, the fact has
not yet been observed in detail.

Section X.
OfAlgce.

In the same manner as in the preceding article I have
spoken, under the collective name of Fungi, of all the
( ryptogamae of a fungous nature ; here also I shall
combine under the name of Algae all cellular aquatic
plants. I do not pretend to give this definition as a strict
classification, but the sense in which it is here taken,
though very vague, is sufficient for my present purpose.
Considering these plants collectively, with regard to
fecundation, we see, that if they are provided with a
fructifying apparatus, they must have it organized in a
particular manner ; their pollen, or at least the fovilla,
must be able to reach the ovules through the water ; this,
perhaps, happens in the Characeae : or the' fecundating
matter is inclosed in the same cavities as the ovules, or
transmitted to these cavities by particular canals, which
perhaps is the case in the Thalassiophytae and Confervas ;
and even admitting that several groups of Algae are fur-
nished with fructifying organs variously formed, there
are others in which the most scrupulous investigation
has not been able to discover any vestige of sexual or-
gans, and which only appear to be reproduced by simple
division : such are the Batrachospermeae and Dia-
tomeae.

We shall notice these different groups without

252 VEGETABLE ORGANOGRAPHY.

pretending to give here a methodical arrangement of
the Algas.

Those who desire to study the subject more in detail
should consult the works of Muller, Hedwig, and espe-
cially those of Bory St. Vincent, Agardh, Lymgbye,
and Fries.

§ 1 . — Of the Characese.

The Characeae, though composed of the single genus
Char a, present an organization so remarkable that we
cannot as yet affirm what is their true place in the na-
tural system. I have elsewhere (Book II. Ch. IV. Sect. 6)
spoken of the structure of their stem ; it only remains
for me to make known that of their flowers and fruit.

Upon the inner side of the verticillate branches of
Char a there arise from each node two little bodies
which appear to be the sexual organs ; one of them,
situated a little below the other, and laterally, seems to
be the male apparatus, for it disappears very soon ; the
other, surrounded at its base with three or four little
branches, appears to be the female organ, for it remains
long after the other, and produces a new individual.

The male apparatus or anther is a red reticulated
disc, bordered by a white transparent membrane formed
of cellules, the partitions of which ars distinct. When
this body is cut across, we see it filled : 1st, with divided
transparent filaments ; 2d, with oval corpuscules filled
with the red matter which colours the disc ; this matter
comes out when the membrane which contains it is
pressed. Hedwig and Vaucher consider these bodies as
grains of pollen, and the matter which they inclose as
the fovilla.

The objections made to this opinion are — 1st, that no
one has seen the anther open to give passage to the

CR.YPTOGAMOUS PLANTS. 253

globules ; but Hedvvig answers, that this may take place,
or at least that the fovilla may come out, by imper-
ceptible pores ; 2d, that the flowering taking place in
water, it is not known how the fovilla reaches the female
organ: but Vaucher explains this anomaly by the nature
of this fovilla, which is resinous and does not mix with
water.

Others have thought : —some, that this apparatus or
red disc was a kind of swimming bladder ; others, that
it was an apparatus containing a peculiar kind of seeds ;
but its fugacity, its appearance at the moment when the
other is developed, have caused almost all naturalists to
regard it as a true anther, in which it remains to be dis-
covered in what manner the fovilla can escape.

The female apparatus is composed — 1st, of three or
four very short branches which surround it at the base,
forming for it a kind of involucrum ; 2d, of an oval
body, marked with five or six lines disposed in regular
spires, enclosing a green opaque body surmounted with
five or six lobes, each of which is situated at the top of
one of the lines. Vaucher considers these as stigmata.
Hedwig considers them as prolongations of an adherent
calyx, and says that he has remarked near their centre
a projecting point, which he considers as the true stigma.
This last opinion appears more likely, because it accords
on the one hand with the position of the lines, and on
the other with the solitariness of the central body.

This body is filled with a multitude of little globules
of different sizes, which Hedwig and Martius consider
as spores or seeds, but which Vaucher denies to be so,
without affirming anything upon their real nature.

This observer has shown me. that when the whole ap-
paratus above described is placed in water, it opens at
the top into five teeth, and there comes out of it a
cylindrical filament, which is the stem of a new individual,

254 VEGETABLE ORGANOGRAPHY.

and from the base of which is prolonged a radicle sur-
rounded with little radical fibrils ; it is not then doubt-
ful that the central body of this apparatus is a repro-
ductive body. But is it a monospermous fruit, as
Vaucher seems to believe ? Is it a fruit containing
several seeds, one only of which is developed at germi-
nation ?

This remains to be investigated.

§ 2. — Thalassiophytse.

I designate with Lamouroux under this name all the
marine Algae described under the names of Fucus, Cera-
mium, and Ulva. In order to make their reproductive
system known, I shall select some examples from the
different groups.

Fucus vesiculosns presents at the extremity of its ra-
mifications kinds of oval swellings ; these are not pro-
jecting tubercles, but simple dilatations of the tissue: the
surface of this dilated part presents kinds of round pores
disposed regularly. When, at the period of fructifica-
tion, this dilated part of the frond is cut, we see that it
is formed internally of a cellular tissue, very much swol-
len up with an abundant watery mucus differing in con-
sistence from the juice of the rest of the plant. Under
each of the pores on the surface is found a round mass,
which seems formed of divided filaments, transparent,
and interlacing one another : these masses, seen by the
naked eye, do not bear a bad miniature resemblance to
the prickly husk of the Chestnut. If they be cut across,
we find in their interior a grand number of oval mem-
branous bodies, which, at maturity, appear isolated from
the rest of the tissue, and destined to come out of the

CRYPTOGAMOUS PLANTS. 255

husk by the pores of the surface. The oval bodies,
placed under the microscope, appear dotted, and this
results from their containing a great number of small
globules ; on observing a recent specimen under the mi-
croscope, I have several times seen these sporangia open
by one of their extremities, and there comes out a viscid
mucus, heavier than water, which falls to the bottom of
the glass on which it is placed, carrying with it the seeds
swimming with it, and which, on account of their
opacity, were visible in the sporangium before its
dehiscence. It is evident that these granules are the
spores or seeds, and that the young plants proceed
from them.

But have these reproductive grains been fecundated ?
This point is very obscure. Reaumur has taken for the
stamens of Fucus, certain divided diaphanous filaments,
which grow in little tufts upon Fucus serratus, F. vesi-
culosus, &c. ; but these appear to me to be simple pro-
ductions analogous to hairs : in fact — 1st, their texture
indicates nothing which calls to mind the structure of an
anther, and there cannot be perceived either pollen or
fovilla; — 2d, they are only found upon a small num-
ber of species; — and, 3d, in those even which are fur-
nished with them, they are scattered over nearly the
whole surface, and remain all the year — two circum-
stances which are incompatible with the idea of consider-
ing them as stamens. Correa has given a much more
likely opinion, in admitting that the fecundation of these
plants is performed by the viscid mucus which surrounds
the bristly masses ; but this opinion is, perhaps, impos-
sible to be demonstrated in a direct manner.

W hatever it be, every Fucus, which has the frond
dilated at the period of fructification, comes, with slight
differences, under the description which I have given.

There is another order of Fucus, in which lateral

256 VEGETABLE ORGANOGRAPHY.

tubercles are produced at the period of fructification ;
these are pierced at their apex by a round pore ; and I
have seen in F. Confervoides, in a fresh state, the spo-
rangia come out through it in intermittent jets ; these
sporangia do not differ from those of F. vesiculosus ; but
we find in the tubercle, where they are lodged, nothing
like the bristly masses or even the viscid mucus of which
I have spoken above, This is my principal objection to
the theory of Correa ; and if one wished to admit the
existence of a fecundating fluid, it must be said that it
is that which is found with the spores in the sporangia;
for it is this alone which is observed in all the Thalas-
siophytse. F. pinnatifidus only differs from those which
I have mentioned in the sporangia being pear-shaped
instead of oval. The fructification of Ceramium presents
little difference from that of the Fuci with lateral tuber-
cles ; but I have not sufficiently studied it to describe it.
As for the marine Ulvae, they only appear to differ from
the Fuci in their sporangia, which are oval, and perfectly
like those of.F. vesiculosus and F. Confervoides, growing
in parcels in the tissue of the frond, and, as this is not
pierced, they can only come out by the destruction of
the tissue ; it is this which causes the regular holes, often
found in old species of Ulva.

It results from these descriptions, that all the Thalas-
siophytae have spores contained in a membranous spo-
rangium, floating in a viscid fluid which, at maturity,
sinks to the bottom of the water, and serves probably to
attach them to rocks. At germination, these spores are
developed by forming a small cup, more or less regular,
which disappears in most, but is met with in Fucus
Loreus, for example, at an advanced age. The spor-
angia are collected several together, or lodged in the tissue
of the leaf, as in Ulva and certain Fuci, or in lateral
tubercles, as in the other Fuci and Ceramium ; they come

CRYPTOGAMOUS PLANTS. 257

out either by the destruction of the tissue ( Ulva), or by
regular pre-existing pores ( Fucus).

In all these plants it frequently happens that the
spores germinate in the sporangia, or in the cavities
which contain these bodies ; this we can easily see by
the naked eye.

§ 3. — Confervas.

The Conferva) (I use this term here in the sense in
which Vaucher has employed it) all grow in fresh water.
Notwithstanding their external resemblances, they pre-
sent great differences in their reproduction, which I
shall rapidly point out, taking for my guide the excellent
work of Vaucher (ll'tstoire des Conferves d'eau douce).

The Vaucheriae, or Ectospermae of Vaucher, present,
at the period of fructification, sessile pedicellate tuber-
cles, solitary or in pairs, or sometimes collected several
together upon a peduncle. These bodies separate natu-
rally from the plant, and Vaucher has seen them ger-
minate : at this period they usually shoot out a green
filament like the plant which has given birth to them ;
more seldom another filament proceeds from the oppo-
site side. The body is not seen to open for the de-
velopment of these filaments, so that we may as well
consider them to be bulbs as seeds. Vaucher has also
observed in most species of this genus little clubs or
hooks, from which he has seen a fine greenish powder
proceed : he considers them as male organs.

The Zygnemse present a much more complicated
organization. At the period of their fructification, their
filaments approach one another by pairs ; from one
filament to the other there are kinds of hollow tubercles,

vol. II. s

258 VEGETABLE ORGANOGRAPHY.

forming transverse passages. A fine green-coloured
matter, disposed in a radiated or spiral manner, or in a
mass, passes from the cellules of one of the filaments
into those of the other ; we then see either this matter
collect in each cell into a globule, or, what is more
probable, a globule, hitherto imperceptible, enlarges
after this (perhaps prolific) operation, and is transformed
into an oval body, which comes out of the cell by the
rupture of its partitions. This body opens at germina-
tion into two valves, and there proceeds from it a
filament very like the plant that gave birth to it. It is
difficult not to believe that these reproductive bodies
are true seeds, and that the green matter performs the
part of pollen. This coupling of the filaments is also
so remarkable, that, although no one has yet been able
to discover any movement in these beings, one would
readily be induced to place them in the animal kingdom.
The Chan transit, or Polyspermy of Vaucher, present
a third mode of fructification. Their internodes slightly
swell up at the period of fructification ; and by the
destruction of the tissue, a multitude of oval globules
comes out of each. Vaucher has seen them shoot out,
either by one of their extremities, or more rarely by
both, a divided filament resembling the mother plant.
This kind of germination sometimes takes place without
the globules coming out of the cell ; and it is this that
makes me think that the Proliferae of Vaucher do not
essentially differ from his Polyspermas.

These three modes of reproduction are the only ones
in which we can perceive an apparatus analogous to a
true fructification. In the other genera we only see,
it appears to me, a simple division, but which is pre-
sented under very different forms.

Thus, in Hydrodyction, each partial filament, which
forms one of the sides of the pentagonal areolae of whicli

CRYPTOGAMOUS PLANTS. 259

the whole sac is composed, separates, swells up, and
forms a sac resembling that of which it formed a part,
without our being able to distinguish any thing that can
be compared to a seed.

In the Batrachospermas little buds detach themselves,
which are developed and produce a new individual in a
manner more analogous to reproduction by bulbs than
any other.

In the Diatomeae the filament is continually broken
across by rectilinear dehiscences, and each fragment,
which at first seems simple, appears double, &c, and is
itself subdivided by transverse ruptures.

The Oscillatoria? do not differ perhaps from this
mode of division, and as they present a kind of motion,
apparently independent of external causes, they are
placed by most naturalists in the animal kingdom.
We here touch the limits of two organized kingdoms,
and find here no other mode of reproduction than that
of simple division.

260 VEGETABLE ORGANOGRAPHY.

BOOK IV.

OF THE ACCESSORY ORGANS ; OR, OF THE MODIFICA-
TIONS OF THE FUNDAMENTAL ORGANS, WHICH
RENDER THEM CAPABLE OF SERVING AS THE
MEANS OF PROTECTION FOR THE OTHER ORGANS,
OR OF FULFILLING OTHER ACCESSORY USES.

In Book I. we have described the elementary organs
common to all plants ; in Book II. we treated of the
fundamental organs of plants, that is to say, of those
which constitute their framework, serve for their nu-
trition, &c. ; and in Book III. we have followed the
numerous modifications of the fundamental organs as
regards their transformation into reproductive ones.
It now remains for us to point out other modifications
of the fundamental organs both of those destined for
the nutrition, properly so called, and of those which
seem intended for reproduction : from these modifications
results their transformation into organs very different
from those from which they are derived ; they become
suited for new uses, which are almost all related either
to the support, defence, or protection of the organs
essentially destined to nourish or multiply the individual.
It is to indicate this less degree of importance, and this,
as it were, subordinate function, that I have given them
collectively the name of Accessory Organs. I have
already made some slight mention of them when speaking
of the organs from which they are derived ; I shall now
describe their origin, form, and use. It is in these
different points of view that I shall speak of Thorns,
Tendrils, and of the different foliaceous, fleshy, petaloid,
and scaly expansions of plants.

OF THORNS. 261

CHAPTER I.

OF THORNS.

I here designate under the general name of Thorns
{Armd) all the organs, or parts of organs, which de-
generate into a hard and more or less sharp point, and
which become kinds of defensive arms for the plants
furnished with them. It has been usual to distinguish
them into spines and prickles; but this distinction is less
easy than was at first thought. It has for a long time
been said, that spines proceed from the wood and
prickles from the bark ; but from this definition it must
be admitted, that only one of the organs exists in
Monocotyledons, where the wood and bark cannot be
distinguished, and one would even be puzzled to say
whether they belonged to spines or prickles ; in Di-
cotyledons themselves, there has been uncertainty in
knowing to which class ought to be referred the thorns
of several leaves, and those which grow upon the
flower and fruit.

In this state of the science, I have remarked that
all the organs of plants are capable of taking, at their
extremities, a degree of induration which transforms
them into thorns ; and it has been easy to see that
those, which, until then, had received the name of
Prickles (Aculei), rather from some vague analogy
than from any strict definition, were organs of the
nature of hairs, indurated and larger than ordinary, and
that all the organs transformed into prickles came to be,
and have been, generally considered as Spines (Spinas).

202 VEGETABLE ORGANOGRAPHY.

Some examples will place these principles beyond
doubt, and will serve at the same time to make known
the varieties and origin of spines and prickles.

The circumstance which most frequently produces
spines is the defective development of the branches of
certain trees which become indurated, and thus are
transformed into thorny points : thus, for example,
the spines of Prunus spinosa, the common Sloe, are
evidently only indurated branches ; they spring from
the axils of the leaves like branches, and frequently
bear leaves ; their structure is absolutely that of
branches; moreover, when one of these plants is found
in a very dry soil, there are more spines, or, in other
terms, many more abortive branches ; if, on the contrary,
it grow in fertile land, it loses its spines — that is to say,
all its shoots, instead of being abortive, are prolonged
into true branches. It is on account of this circumstance,
common to several trees and shrubs, and especially
those of the Rosacea?, Amygdalacese, &c, that we often
observe that spiny plants lose their thorns by culture,
as I have observed in the wild Medlar, in which all
the spines disappeared in two years. The spines of
Gleditsia, which are so enormous and branching, those
of Genista, Cytisus, and a multitude of others, are only
abortive and indurated branches. We might say that
these are ramal spines.

The petioles of some species of Astragalus, of Hali-
modendron and Ammodendron, present an analogous
phenomenon ; they harden at the end of the life
of the leaflets, and when these fall off, or are ready
to do so, they change into very hard and sharp true
petiolary spines : from the nature of their origin, they
are always simple ; they become almost as hard as the
stem itself; for all petioles which have this spinescent
tendency are continuous, and not articulated at the base.

OF THORNS. 263

The stipules of several plants become so indurated as
to present the appearance of true stipulary spines: such
are those of Pictetia. But it must be observed, that
the coussinets, or protuberances of the branch upon
which the petioles are placed, sometimes acquire so
large a lateral development as to form true spines,
which have often been taken for stipulary ones : they
are very well seen in certain species of Acacia, where
they coexist with the true stipules, in A. Hcematomma
for example ; but when only one of these two organs is
met with, it is almost impossible to affirm whether the
spines on both sides of the leaf are indurated stipules
or lateral expansions of the coussinet. Analogy with
neighbouring plants can alone remove the doubt.

It happens in a small number of cases that the
leaflets are wholly or partially abortive, and the petiole
is changed into a spine, which is simple when all the
leaflets are abortive ; trifid, when the two stipules
adhering to the base of the petiole, or the two lower
leaflets reduced to the middle nerve, harden, forming the
two lateral branches of the spines ; quinquefid, when
the stipules and leaflets are present at the same time.
It is in this manner that the spines of different species
of Berberis appear to be formed, being evidently
nothing but leaves reduced to their middle nerve, the
bundles of axillary leaves replacing their physiological
functions.

The leaf itself may be transformed into a spine in
two ways : sometimes it is found reduced to a foliaceous
petiole, more or less dilated, and terminating in a spiny
point, as appears to take place in Littcea, Yucca, &c. ;
sometimes the limb itself is prolonged into a spine
formed by the prolongation of the middle nerve, as in
Chuquiraga. What I have said of leaves is equally
applicable to leaflets, the middle nerve of which is

264 VEGETABLE ORGANOGRAPHY.

prolonged into a spine, as in Coulteria; to the lobes of
leaves which have their nerves prolonged into spines, as
is seen in Thistles ; and to spiny teeth, which are only
lobes smaller than the preceding. The points of the
leaves of the Holly come under the class of foliary
spines ; but those of Aloe and Agave are analogous to
the lateral ones of petioles.

Leaves, reduced to the state of scales, involucra, or
bracts, present similar phenomena, and approach, for
the most part, petioles devoid of limbs and prolonged
into spines ; this is readily seen on examining the
involucra of the Thistle and other spiny Compositae.

Peduncles, like all the other organs, may harden at
the point so as to become spines. This takes place after
flowering, and is presented under two remarkable
forms : — sometimes the floral branches, more or less
ramified, remain after the fall of the flowers and fruits,
and form kinds of spines, usually branched, and ap-
parently terminal, as, for example, in Alyssum spinosum,
Mesembryanthemum spinosum, &c. ;— sometimes the
axis of the spike becomes indurated after flowering,
and is terminated, at maturity, by a hard point, which
in certain plants, as Trifolium subterraneum, serves, by
the curving of the peduncle, to penetrate the earth and
bury the seeds there. The pedicels, when they do not
bear flowers, are sometimes changed into spines ; this
appears to take place in Nauclea, Sec.

The parts of the flower even, though more fugitive
than the others, and having consequently less time to
harden, occasionally present spiny degenerations.

Thus, the sepals so partake of the nature of leaves
that they frequently terminate in spines, as in Stachys.
The spiny pappi of certain Composite come under this

class.

The petals themselves, notwithstanding their usually

OF THORNS. 265

tender and fugacious nature, are sometimes terminated
by spiny points, as in Cuviera.

The persistent or sterile stamens of someByttneriaceae
acquire so firm a texture, that they may take the name
of spines.

The styles often remain after flowering, and form at
the apex of the fruit spines often very hard and long:
as in Martynia.

Thus all the organs of plants, except the roots and
seeds, are capable of hardening or being prolonged into
spines ; so that it is impossible to say that a spine is an
organ properly so called, but that it ought to be con-
sidered a particular state of vegetation.

All thorns which do not result either from the in-
duration or prolongation of any of the organs which I
have mentioned, bear the name of Prickles, and may
be considered as kinds of hairs larger, stronger, and
harder than usual. There are cases where the transition
from hairs to spines is so gradual as to show their iden-
tity of nature : thus, in the bundles of hairs which spring
from the axils of the leaves of Opuntia, we see some
larger than the others, and transformed into very long
and hard prickles. It is nearly the same in Roses : we
often see the glandular hairs of their peduncles and
calyces harden into true prickles. There are cases,
then, where there can be no doubt that prickles are
analogous to hairs. I know that it cannot be affirmed
in a general manner, except by means of analogy.

Prickles are distinguished from spines in their never
taking the place of any of the great organs of the plant ;
they are usually found upon the stems, branches, pe-
duncles, and petioles, and upon the nerves of the leaves,
calyx, or even of the petals ; but they never terminate
either the fibres or nerves ; whilst spines, being indura-
tions of organs, are always placed at their ends. This

26() VEGETABLE ORGANOGRAPHY.

appears to be the more certain way of distinguishing
them, especially in Monocotyledons.

From the prickles replacing hairs, and spines replacing
all the other organs, it follows, consequently, that the
former are superficial, and the latter intimately connected
with the tissue.

It is remarked that most prickles of stems or petioles
are curved with the point directed downwards, as in the
Rose ; but this rule ought not to be taken as a general
one, for there exist perfectly straight prickles in several
Mimosea3.

Thorns, whatever be their origin, are generally the
defensive weapons of certain plants, serving to protect
them from the teeth of large animals, &c. Some,
perhaps, may serve either to penetrate the earth to
favour the natural sowing of seeds, as in Trifolium sub-
terraneum; or to hook the fruits or seeds of certain
plants to the wool of animals, in order to transport them
to a distance, as in the Burdock.

The existence, and consequently the use of spines, is a
fact entirely peculiar to certain species, sometimes to cer-
tain varieties, and is but very slightly connected with the
general symmetry, and consequently with the funda-
mental laws of their organization. We frequently observe
in the same families and the same genera, species, some of
which are furnished with, and others devoid of thorns.

Let us observe, on concluding this chapter (and this
remark will be equally applicable to the following),
that the analogy which is observed in the manner in
which the sepals, petals, stamens, and carpels can, like
true leaves, change into spines or tendrils, tends to
confirm the identity of origin of these organs.

OF TENDRILS. 2G7

CHAPTER II.

OF TENDRILS.

Under the name of Tendrils (cirrhi), are designated
those soft, cylindrical prolongations, which are capable
of twisting or twining round bodies they meet with ; they
are found in different parts of plants, and generally
serve to support those which are unable to do so of them-
selves.

The origin of tendrils is perfectly analogous to that
of spines ; they are not organs properly so called, but
degenerations or modes of prolongation of which almost
all the organs are susceptible, and they only differ from
spines in their softness, flexibility, and in their usually
greater length.

We call those Petiolary which are produced by the
prolongation of common petioles into flexible filaments ;
this is frequent in the simply winged leaves of the Le-
guminosge, among the Vicieac, and is met with, though
more rarely, in bipinnate leaves, as in Entada. In this
last case, the common petiole is prolonged into an elon-
gated tendril, and the partial ones either do not lengthen
at all, or only present a small, scarcely apparent point.
We also find tendrils analogous to simple leaves, but
with the segments so distinct that they resemble winged
leaves, as in Mutisia, and especially in Cobcea. Petiolary
tendrils are sometimes simple, sometimes branched : —
simple, when they are formed by the unbranching pro-

268 VEGETABLE ORGANOGRAPHY.

longation of the petiole, as in Lathyrus Aphaca; —
branched, when this prolongation bears lateral branches,
which probably represent the middle nerves of the unde-
veloped leaflets or lateral segments ; this structure is
frequent in Vicia. When the tendrils are long they
readily twine around neighbouring bodies, and serve to
support the plants ; but sometimes they are so short, as to
be unable to perform this office, and only exist as indi-
cations of the tendency of certain petioles to be thus
prolonged ; we see this in Orobus. Lastly, the petioles
of several Fumariaceae, although terminated by foliaceous
limbs, are frequently twisted at the point so as to act the
part, and have the appearance of tendrils. Those of
Clematis cirrhosa, and some other species, remain after
the destruction of the segments of their limb, and form
kinds of tendrils.

We know but a very small number of examples of
Foliary tendrils, that is to say, of leaves thus pro-
longed ; and these examples also are not of true limbs,
but of foliaceous petioles devoid of the limb, and with
the nerves, straight or parallel at the base, converging at
the apex into a flexible filament; as in Flag ellaria Indica,
Gloriosa superba, and the upper leaves of Fritillaria
verticillata. If the filament, which proceeds from the
middle nerve of Nepenthes, and is afterwards prolonged
into a cup, can be considered a true tendril, it will ap-
proach this class.

Stipulary tendrils are very rare, and even doubtful ;
we ought to refer, perhaps, to this class: — 1st. The
filaments which arise from the axils of the cotyledons
of Trapa, and along the base of its stem. 2d. The
tendrils of the Cucurbitaceas, which occupy the place of
a stipule, but are only on one side of the leaf.

Petiolary glands are usually sessile, or nearly so, and
but little prolonged ; but it sometimes happens that they

OF TENDRILS. 269

are prolonged into slender filaments more or less ana-
logous to tendrils ; thus, the glands of the petiole of
Passiflora ligularis are lengthened into long and nearly
cirrhiform filaments.

'The petiolary tendrils of Smilax are difficult to be
understood with regard to their anatomical origin.
These climbing shrubs generally have the petiole
dilated at the base into a kind of sheath, which might
readily be taken for a stipule adhering to the petiole, if
the analogy with other Monocotyledons were not opposed
to it ; above this sheath there proceed from the petiole
two opposite, simple, filiform, or elongated tendrils ; are
they prolonged petiolary glands, as in Passiflora ligu-
laris ? Their position would appear to make us believe
it ; but as no Smilax bears petiolary glands, this hypo-
thesis is not admissible. Are they prickles prolonged
into a filiform appearance? The irregularity of the
position of the prickles of Smilax, compared with the
regular position of the tendrils, ought to clear up this
opinion. Are they prolongations of the petiolary
sheath ? This idea is founded upon the circumstance
that they arise from the extremity of it, and it is con-
firmed by the fact that in *S. herbacea, which has no
sheath, there are no tendrils. Lastly, the opinion
which appears to me to be the most likely is, that the
leaf has originally three segments, and that the tendrils
represent the two lateral ones, which are abortive or
transformed. When very young leaves of Smilax aspera
are examined, it is very difficult not to admit this opi-
nion ; it is confirmed by the analogy of the Smilaceas
with the Aroideae, which often have leaves with several
segments.

Pedunculary tendrils are more frequent in nature,
and clearer as to their origin, than the preceding ; they
are, as the name is intended to indicate, produced by

270 VEGETABLE ORGANOGRAPHY.

the prolongation of the peduncles ; this supposes the
total or partial abortion of the flowers which they ought
to bear. Thus, for example, it is easy to satisfy oneself
that the tendrils of the Vine and all the Ampelideae are
peduncles ; in fact, we always see that they are opposite
the leaves as the bunches are, and it is not rare to find
some which are half furnished with flowers and half with
tendrils. The little bundles which are found at the upper
part of the Vine, frequently present states intermediate
between absolutely fertile bunches and those which, by
the abortion of the flowers, are changed into tendrils.
Analogous facts tend to prove that the tendrils of Pas-
sion-flowers are likewise only abortive peduncles, for
they occupy their places in the axils of leaves ; and in
some species, as in Passlflora clrrhiflora^ the peduncle,
which is branched, is partly changed into tendrils and
partly furnished with flowers. The pedicels at the
base of the scapes of Cardiospermum, and of some
other Sapindaceae, are almost always transformed into
tendrils. In a species of Smilax, several of the axillary
peduncles are transformed either usually or accidentally
into tendrils, which must not be confounded with those
which spring from the petiole.

Bracts and sepals so resemble leaves in their nature,
that it is difficult to believe that they are not capable of
being transformed into tendrils ; the examples, however,
of this transformation are rare and doubtful. The floral
leaves of Fritillaria verticillata change into tendrils
very analogous to those of Gloriosa superba; the sepals
of Calytrix are prolonged into a very slender filament,
which seems to be the rudiment of a tendril analogous
in form to that of Orobus. The awns of the glumes of
the Gramineae also appear to be an analogous degenera-
tion; according to Rceper, the glume represents the
sheath, and the awn is the abortive limb. They

OF TENDRILS. 271

frequently have a tendency to twist spirally as true
tendrils.

Corollas themselves, notwithstanding their fugacity,
sometimes take the appearance of a tendril; thus, in
the genus Strophanthus the lobes of the corolla are
prolonged into a fine filament, from one to twTo inches
long in most of the species, and even attaining seven
inches in S. hispidus of Sierra Leone : the five filaments
proceeding from the five lobes are twisted together
before the expansion of the flower, thus forming a kind
of floral tendril which twines around the neighbouring
branches. The tops of the anthers of Nerium Oleander
are prolonged into kinds of apparently corolline ten-
drils, which are sometimes twisted together, as those of
Strophanthus. Thus all the organs capable of changing
into spines appear endowed, in other plants, with the
faculty of changing into tendrils.

Stems put on this appearance in a great number of
cases ; and it is usual to call them simply by the name
of twining or climbing stems. The annual shoots are
the parts which most frequently present this tendency ;
whence it results, that when the plant is an annual, it is
twining throughout its whole life. In perennial plants,
two cases happen : — the stem either constantly remains in
the same twisted state as it was in during the first year,
as in most Passion-flowers ; or, the lower part becomes
so firm as to be able to support itself, and then we have
a shrub with an erect stem and twining branches ; this
is observed in several species of Convolvulus. The
reverse takes place, according to Vaucher, in Periploca
Grceca, which scarcely twines the first year, but after-
wards very strongly encircles trees which it meets with.

The ti'ansformation of organs into spines supposes in
general the existence of a hard and solid fibrous tissue ;
therefore this texture is more or less remarkable in all

212 VEGETABLE ORGANOGRAPHY'.

spiny plants. The transformation into tendrils supposes,
on the contrary, a soft, flexible, fibrous tissue, capable of
elongation; therefore we remark in each family, that
those plants, in which the stem has a tendency to be
prostrate or climbing, have at the same time and for the
same reasons some organs transformed into tendrils.
The Vicieae, Mimoseae, PassifloreaD, Sapindaceae, and
Smilaceae, with weak stems, have well-developed ten-
drils ; whilst in the same groups we find no tendrils, or
only rudimentary ones, where the stem is stronger ;
thus Orobus and Faba, which have a firm stem, are the
only Vicieae which are almost devoid of them ; none of
the Mimoseae with strong stems have any, whilst
Entada, which has twining stems, is furnished with
them ; the arborescent Passifloreae are alone devoid of
them ; the weak and climbing Sapindaceae only, as Car-
diospermum, Urvillea, Paullinia, are furnished with
them ; Smilax herbacea, which has an erect stem, is
devoid of them, whilst all the other species are provided
with them. Thus it is generally found that tendrils are
only developed in plants too weak to support themselves.
The existence of this kind of support enables them to
twine round trees or shrubs; therefore most plants of
this kind live, in preference, in forests when they are
very large, and among bushes and hedges when small.
The farmer imitates this natural phenomenon when he
sows Vetches among Oats, which serve to support them.
The twisting of twining stems and tendrils takes
place in each species in a determined manner — either
crossways or spirally ; the former is performed upon the
same plant, and only takes place in tendrils which find
no other body to lay hold of: this is frequently seen in
Sapindaceae. The twining in a spiral manner, properly
so called, always takes place in stems or tendrils which
are twisted around a long body. What is most remark-

OF FASCICULATED EXPANSIONS. 273

able is, that in each species the direction appears to be
strictly fixed, viz. — from right to left in the French
Bean, from left to right in the Hop. Bryonia presents
in this respect a phenomenon of which I know no other
example ; its tendril suddenly changes its direction in
the middle of its course, so that the upper half twines in
a contrary direction from the lower. The causes which
determine the twining in general, and the particular
direction, are very imperfectly known, and purely
physiological.

CHAPTER III.

OF FASCICULATED EXPANSIONS.

All the caulinary organs, which are not expanded into
foliaceous or petaloid limbs, have a tendency, in certain
constant or accidental cases, to form kinds of expansions
of a singular nature, and which I call Fasciculated
Expansions, extending a little the ordinary use of this
term. In these expansions, the branches, peduncles, or
petioles, instead of being cylindrical, become spread out,
and, as it were, semi-foliaceous, the fibres or nerves re-
maining either nearly parallel or converging, or diverging
towards the apex, but nearly simple, and not expanded
as in the limb of leaves. It might be said that in this
mode of vegetation, the fibrous bundles, usually arranged

VOL. II. T

274 VEGETABLE ORGANOGRAPHY.

in such a manner as to form a nearly cylindrical body,
spread out at the base, arranged side by side so as to
form an expanded disc ; and in certain cases, one would
be inclined to believe that the fasciculated expansions
are produced by the natural union of several small
branches which arose, side by side, from the same
point. There is no doubt that this happens sometimes,
but it would be imprudent to affirm that it is the only
cause of these expansions.

Branches are very subject to this unusual kind of de-
velopment ; the fasciculated branch or stem is nearly
cylindrical, afterwards it becomes flattened, more or less
striated or channelled longitudinally; near the extre-
mity the small portions separate by the striae having a
tendency to separate from each other, and frequently
form as many small branches situated nearly upon the
same plane ; when they do not separate, they frequently
present the appearance of nerves united by cellular tis-
sue. Almost all vascular plants may accidentally pre-
sent this phenomenon ; thus I have observed it among
herbaceous Dicotyledons in Ranunculus, Euphorbia,
Cyparlssia, Chicorium, Jasione, the Cockscomb, some
species of Stapelia, See. ; among woody ones in Daphne
Mezereum, the Jessamine, Ash, Broom, (PI. 4, fig. 1,)
&c. ; and among Monocotyledons in Asparagus, and also
in some Ferns.

If it were always accidental, it would possess little
organographical interest ; but it is sometimes presented
under an appearance so constant, that it seems to form
part of the ordinary state of plants ; this appears to be
the case in Xylophylla (PI. 16, fig. 1.)

On examining the nature of plants capable of forming
these expansions, we see them either much branched, or
furnished with a very abundant cortical cellular tissue —
two circumstances which tend to confirm the hypothesis

OF DEPOSITS OF NOURISHMENT. 275

that they result from the union of several neighbouring
branches upon the same plane.

Fasciculated stems must not be confounded with
stems or branches which have the cortical parenchyma
so extended upon their two opposite sides as to give
them the expanded appearance of a foliaceous limb ;
thus, several species of Cactus, as C. Pkyllanthus and
Opuntia, have the branches expanded laterally into a
limb which has a leaf-like appearance. Ruscus aculeatus
(PI. 16, fig. 3,) also presentsw inged branches, which have
exactly the appearance of a true leaf. In these different
cases, we recognise their true nature either by studying
the origin of the organs, or by following their develop-
ment ; when they begin to enlarge, the woody body, by
distending the bark, gradually obliterates these foliaceous
appendages, and the winged branches are changed into
cylindrical stems.

CHAPTER IV.

OF DEPOSITS OF NOURISHMENT, OR THE FLESHY, FE-
CULENT, ETC. DEGENERATIONS WHICH MODIFY THE
TEXTURE OF ORGANS.

In the same manner as some organs, taking a more
ligneous texture than ordinarily, may be changed into
thorns, and thus become the defensive arms of the
plant ; so there are other parts which, acquiring a con-
siderable thickness, receive into their tissue a large
quantity of watery, mucilaginous, feculent, or oily

t 2

27G VEGETABLE ORGANOGRAPHY.

matters, and are then found to become deposits of nutri-
ment for a given period or a given part of plants. The
organs thus modified are the more important to study,
since they perform a considerable part in the nutrition
of plants, and often produce phenomena apparently con-
tradictory to the progress of the sap. In fact, if it be
true, as vegetable physiology appears to demonstrate,
that the sap is only elaborated in the foliaceous parts,
and only becomes a nourishing juice after this elabora-
tion, how can we understand the nutrition of a great
number of organs which can receive no action from the
foliaceous parts, and which are evidently nourished by
the ascending sap ?

The cellular tissue of several very different organs is
capable of being dilated, and of receiving a much larger
quantity of water than usual ; it is this which constitutes
the ordinary state of the leaves of succulent plants, that
of several fleshy roots, that of succulent pericarps called
fleshy fruits, that of fleshy spermoderms, &c.

The nature of the water accumulated in the tissue
presents differences, both between one organ and an-
other, and one plant and another : thus, the water which
swells up the leaves of several Ficoids contains earthy
and alkaline salts in solution ; that which is in most
fruits contains mucilaginous or saccharine matters, &c.

The tendency of each organ to a state of anasarca is
sometimes constant in the species, sometimes accidental.
Thus, the leaves of Ficoids, Crassulaceae, PortulaceaB,
Aloes, &c. the stems of the Cacteas, Stapelia, &c, and
the perigones oiBUtum, are constantly fleshy. The peri-
carps of a great number of plants also present this dis-
position in a permanent manner. In all these cases we
remark that this state is connected with the total ab-
sence of the stomata when it affects fruits, or with their
small number when it affects leaves.

OF DEPOSITS OF NOUR1SHMKNT. 277

But this state, constant in certain plants, is met with
accidentally in others evidently caused by external cir-
cumstances ; thus Lotus corniculatus, Plantains, and
several other plants, have leaves more fleshy than usual
when they grow near the sea.

It results from this disposition in leaves, that they
become receptacles of water, and the plants, thus organ-
ized, can consequently support drought much better
than others, since they then reabsorb the water of their
leaves. Thus eminently succulent plants, as the Ficoids,
can support the long drought of the African deserts
by a phenomenon nearly analogous to that which
enables camels to travel for a long time in the same
regions.

As to fleshy pericarps, it is not easy to determine the
use of this particular state to the plant. Does this de-
posit of juices, gradually absorbed by the plant, serve
to continue the nutrition of the seeds until their matu-
rity ? Does it serve, by decomposing it, to favour their
coming out of the pericarp, which in fleshy fruits is
always indehiscent ? Does it serve, at this period, as a
kind of manure to nourish the germinating seeds ? All
these opinions are evidently true in certain cases.

It is rare to see pericarps pass accidentally from a dry
to a fleshy state, or vice versa. We can mention a small
number of examples ; such as that singular kind of the
Almond-peach which sometimes bears upon the same
tree fruit with a fibrous pericarp, and others with a
fleshy one. But we know a host of examples where
plants, very similar in structure, differ in the dry or
fleshy nature of their pericarp ; such are the Almond
and Peach, Silene and Cucubalus, Hypericum and Andro-
soemum, &c.

The deposits of mucilaginous and feculent matters are
as frequent as the preceding ; they may be found in all

278 VEGETABLE ORGANOGRAPHY.

the organs of plants, and their presence determines the
possibility of the development of certain parts. In fact,
without this deposit prepared before-hand, it would be
impossible to understand how certain ascending parts are
nourished until the period when their own nourishing
organs are developed, or how certain parts are developed
although apparently devoid of proper organs to elaborate
the ascending sap.

If I form a just idea of this phenomenon, which is in
itself very remarkable, this is how I understand it : the
watery sap or lymph absorbed by the roots passes
through the cellular tissue essentially by the intercel-
lular passages, as Kieser and others appear to have
clearly demonstrated ; when it passes into the canals or
passages which separate the very long cellules, which
are usually empty, it follows its course without any
alteration; when it traverses the organs abundantly
supplied with round cellules, its motion nearly ceases,
or is very slow, and then another phenomenon may take
place : if the progress of the vegetation of the preceding
year has accumulated in these cellules a certain quan-
tity of mucilage, this is partly or wholly dissolved in the
lymph which surrounds the cellules, and when, by the
development of the upper parts, this lymph is attracted
there, it does not arrive in the state of pure water, but
it contains a certain quantity of mucilage in solution.

I think that the same takes place with regard to fecu-
lent or oily matters, although we do not possess, espe-
cially as concerns the former, any correct ideas of the
manner in which they can be affected by the water so as
to become soluble in it. Although we do not know
how to render fecuia or starch soluble except by pro-
cesses hardly likely to be met with in vegetation, it is,
however, certain that it becomes so by the force alone
of vegetable life, and the history of the germination of

OF DEPOSITS OF NOURISHMENT. 279

Wheat is an example. It appears to me that an analo-
gous phenomenon evidently takes place when the lymph
traverses a feculent or oily deposit.

If we now apply this general idea to all the cases
where certain organs are nourished without the possi-
bility of their being so by the descending sap, we shall
see that they owe it to deposits prepared before hand in
the course of the ascending sap.

Thus, in all perennial plants, mucilaginous or fecu-
lent matters are deposited, towards the end of summer,
in the upper parts of their roots : when the new stems
shoot up in spring, they are nourished by the ascending
sap, which, in passing through these nutritious deposits,
dilute them, and, becoming charged with them, carry
them to the parts destined to be developed until the
production of leaves allows them to prepare their own
nourishment. Tuberous roots present, in this respect,
special organs for the deposit of these matters, and we
see that they are destroyed after the development of the
young shoots: this is also applicable to subterranean
stems and tubercules, as well as to the nodes of ordinary
stems from which we see the young branches produced.

In dicotyledonous trees the pith is a true deposit of
nourishment as regards the young shoot, and we see it
dry up and perish after it has fulfilled this office.

The receptacles of several flowers serve the same pur-
pose ; before flowering, they are found loaded with
mucilaginous and feculent juices, which are carried up
by the ascending sap during flowering, and serve to
nourish the flowers and fruits ; after this period, they
become empty and dry up, as may be seen in the Arti-
choke, &c.

What we have clearly seen in those receptacles,
which are very large, and nourish a great number of
flowers, takes place in all peduncles, but in a more or

£80 VEGETABLE ORGANOGRAPHY.

less visible manner : thus, for example, the centre of the
rays of umbels is a point where a deposit of nourishment
is effected, and from which the flowers draw their sup-
ply. In all cases flowers are developed by the ascending-
sap, which, in its passage, meets with deposits prepared
beforehand by the action of the foliaceous organs.

The placentas of fruits enjoy the same function with
the greater energy as they are thicker or more fleshy :
thus, in several Solaneae, Rubiaceae, Primulacese, &c. in
Cobcea and a number of others, they are true deposits
of fecula, which serve to nourish the seeds.

Finally, cotyledons themselves are often fleshy, and
then become true deposits of nourishment prepared by
the mother-plant, and absorbed by the embryo at ger-
mination.

All that I have said of the deposits of mucilage and
fecula is equally applicable to those of fixed oil situated
either in the pericarp, as in the Olive, in the albumen,
as in the Euphorbiaceae, or in the cotyledons, as in the
Poppy.

Thus, all the organs of plants are, in certain cases,
capable of performing a particular physiological function,
in becoming depositories of nutriment prepared for dif-
ferent growing organs ; but this circumstance, which
alters their use, ought only to be considered with regard
to Organography as a particular modification or dege-
neration. It is in this point of view that I have here
spoken of the subject, which will be treated of more in
detail in the Physiology.

OF SCALES. 281

CHAPTER V.

OF SCALES.

We generally designate under the name of Scales
(Squamce) the little flat pointed bodies which are found
upon different parts of the surface of plants ; but there
are few terms under which more dissimilar objects have
been confounded ; the simple mention of them will be
sufficient to make their differences understood, and to
show how we must distrust, in natural history, those
vague denominations which are not founded upon
anatomy.

The organs confounded under this name may be
referred to three principal classes : — appendages analo-
gous to hairs ; — excrescences of certain organs ; — and
foliaceous organs more or less abortive and reduced to a
rudimentary state.

Scales analogous to hairs are : — either kinds of radi-
ating and peltate discs, as those of Elceaguns (PI. 3,
Fig. I.) which seem formed by the constant union of
several hairs radiating upon the same plane ; or of en-
larged hairs, scarious and dilated at the base, as those
on the petioles of Ferns. These appear, at first sight, to
differ much from hairs ; but if they be studied in the
whole family, every degree of size is met with, from
those which are entirely in the form of hairs, to those
which are large and dilated into scales. The membra-
nous expansions which crown the fruits of several Com-

282 VEGETABLE ORGANOGRAPHY.

posita? and some Dipsaceas, may be considered either
as the hairs of pappi united together, or as membranes
formed by a greater abortion of the limb of the calyx
than ordinary.

The second class of scales are the expansions peculiar
to certain organs ; thus, the calyx of Salsola bears-upon
its back membranous appendages which form part of
that organ, the throat of Nerium Oleander, Silene, &c.
is prolonged into petaloid scales which collectively form
a kind of crown ; these different bodies, whatever their
origin, are not special organs, but simple forms peculiar
to such or such a part.

Lastly, the most frequent sense of the word scale is
to designate small fiat bodies which are the rudiments of
abortive leaves or analogous organs, such as stipules,
bracts, or sepals, or even of the other floral organs
reduced to very small dimensions. This degeneration
changes their form and appearance, and causes them not
to be recognised by those who are not acquainted with
this kind of change. Examples taken from the different
organs will suffice, I think, to make me understood.

It is customary to say, that the calyces of Pinks are
furnished with four scales at their base ; but whoever
will examine them with care, will perceive that they are
only the upper leaves or bracts, which from being near
the flowers, remain very small, and have taken the
appearance designated in other cases by the term scales.

The branches of most of the Erythroxyleas, of
Pictetia squamata, and several other plants, are often
invested with little imbricated and scarious scales, which
are persistent and very close stipules, the leaves of
which are wanting.

The bracts which form the involucra of the flowers
of the Compositae and Dipsaceas are leaves reduced to
small dimensions, and for this reason have received the

OF BUDS. 283

name of scales. This name is also given in the same
plants both to the abortive bracts situated between the
flowers, and more usually called Yaleje, and to the
pieces of the calyx, reduced to the state of pappus,
when their form differs from that of hairs, and their
texture gives them a rude resemblance to scales ; it is
also in this sense that this name has been given to the
bracts of cones, to the glumes of several Graminese and
Cyperaceae, &c. &c. Lastly, the scales of buds come
evidently under the same class, and are only the rudi-
ments or abortions of leaves, petioles and stipules ; but
their history is so important that it deserves particular
mention, and therefore I dedicate the next chapter
to it.

CHAPTER VI.

OF BUDS.

The term Bud is taken, in the French language, in
two very different senses. 1 st, the young productions or
branches of perennial plants are commonly meant by it.
Botanists call these Young Shoots or Scions (turiones).
When these shoots are found covered or protected in
their infancy by particular scales, the plant is said to
have scaly buds ; in the contrary case, they are said to
be naked. 2d, Botanists, on the contrary, designate by
the name of Bud {Gemma), not the young shoot, but
the collection of scales or coats which surround and

284 VEGETABLE ORGANOGRAPHY.

protect it in its infancy ; in this sense, which we shall
adopt here, they say that a young shoot is naked, or
without buds, when at its development it lias no
particular integument ; they say that it has a scaly bud,
when it presents an integument formed of pieces
analogous in texture to scales ; and they may also say
that it has a bud with membranous coats or fleshv
scales, &c.

Gardeners are also accustomed to call by the name of
bud both the flowers, as yet undeveloped, and the buds
(in the sense of botanists) before their expansion ; but
botanists have admitted the name of Alabastrum to
designate the uuexpanded flower : but let it be remarked
that, throughout this chapter, we restrict the name
Bud to the integuments of the young shoots at whatever
age we examine them.

Buds present very different appearances according to
the place they occupy, and according to the nature of
the plant: we shall distinguish in this respect two
classes, viz. 1st, caulinary buds, which grow upon the
stems of trees and shrubs, or the buds, properly so
called ; 2d, those which are formed at the neck of
perennial plants, on a level with the surface, as scions,
or underground, as bulbs properly so called.

The origin of these two kinds of buds is always a
semi-abortion or degeneration of the foliaceous parts,
but their position produces such great changes in their
appearance, that it will be better to study them sepa-
rately at first, and afterwards to show their points of
affinity.

All Dicotyledonous trees have not their young shoots
covered with special integuments ; and these integu-
ments themselves, when they exist, proceed always from
external leaves or stipules, which, on account of their
premature exposure to the action of air and light, suffer

oi ih;ds. 285

in their development, and are more or less completely
changed into scales. When we examine the structure
of a bud of the Ash or Sycamore in spring, we see the
external scales short, hard, of a reddish brown colour,
and slightly downy, and the inner rows gradually be-
come more membranous, paler, and longer; afterwards
they bear at their extremity the rudiments of leaflets ;
then they become small leaves, so that it is impossible
to doubt that the external pieces are of the same nature
as the internal.

Buds have received particular names, according as
they are formed by different portions of the foliaceous
organs, and according to the degrees of their degenera-
tions and adhesions.

1st. We say that buds are Foliaceous when the
leaves being sessile, their limb, reduced to the form of a
scale, forms the buds, as in Daphne Mezerewn.

2d. Buds are said to be Petiolaceous when the
bases of the petioles, dilated into scales, form the co-
vering of the young shoot; this is seen in exstipulate
petiolate leaves, as the Walnut, the Ash, the Horse-chest-
nut. (PI. 12.)

.'id. Stipulaceous buds are those which are formed,
not by the leaves, but by the stipules, which are not
united to the petiole. We distinguish two kinds of
stipulaceous buds: 1st, those which are formed by
the superposition of a great number of stipules which
together inclose an entire young shoot ; this hap-
pens in almost all the Amentaceae, as the Oak, Willow,
Elm, &c. ; 2d, those, the stipules of which, free
or united together by their outer margin, form the
peculiar envelope of each leaf, and are gradually de-
veloped with the branch itself; this is seen in Ficus,
(PI. 6) and the Magnoliaceae ; these kinds of mono-
pbyllous buds can be recognised at first sight by their

286 VEGETABLE ORGANOGRAPHY.

terminating the branch in the form of a very sharp
cone.

4th. When the stipules adhere to the petiole, these
two organs, united into one, form the scales of the buds,
which are then called Fulcraceous ; this happens in
most Rosaceae ; these kinds of scales frequently have
three lobes or teeth, which indicate the origin of the
scale, formed by the petiole and the two stipules
united together. (PL 15, figs. 4, 5, 6.)

Monocotyledonous trees, or palms, have buds exactly
resembling the preceding as to their origin ; they are of
the petiolaceous class : it may be said that the tops of
Draccena, and other trees of this kind, have foliaceous
buds ; but these are the only classes of buds which can
be found among Monocotyledons, because the stipules
are wanting here, and, consequently, stipulaceous and
fulcraceous ones are impossible to be met with.

In perennial herbs the shoots perish each year, or at
least after each flowering, and new ones are developed,
which spring from the permanent part of the stem un-
derground, or on a level with the surface, and which is
usually confounded with the root. These new shoots
often proceed from buds which are named Turiones,
which, considered as to the origin of their scales, pre-
sent the same varieties as the buds of trees. Thus, in
Dicotyledons, we may say that foliaceous turiones are
met with in Asters, petiolaceous ones in Paeonies, (PI.
15, figs. 1, 2, 3,) and fulcraceous ones in Potentilla. I
do not know any example of purely stipulaceous turi-
ones, for all of the families furnished with them have
only herbaceous species ; but their existence is not im-
probable, and it might be said that Salix herbacea has
them when its stem is subterranean.

Among Monocotyledons it may also be said that the
scales of the bulb of the Lily are simple leaves, which

OF BUDS. 287

from being underground are etiolated and fleshy, and
come under the class of foliaceous buds ; whilst the radical
buds, produced by the dilated petioles of Hemerocallis,
are examples of petiolaceous ones. We know that in
this class we can find none of those which suppose the
existence of stipules.

Although aerial and subterranean buds have the same
origin, the difference of their position produces diver-
sities in their nature, which deserve to be analyzed.

Aerial buds owe to their position the peculiar texture
of their scales, which, exposed to the action of air and
light, evaporate much, and are found reduced, as it
were, to their fibrous tissue. These characters go
on diminishing in the inner scales, which from the
same circumstance evaporate less, and retain the juices
more.

The buds have eminently two uses to perform — to
defend the young shoots from damp and cold.

In the first respect the scales are generally so nume-
rous and applied to one another so exactly, that the
rain cannot penetrate their interstices before their
expansion. Several buds present, moreover, a parti-
cular protection against moisture, in having their scales
covered over with a varnish or viscid layer of a resinous
or waxy nature, which is not miscible with water, and
thus prevents its introduction. The buds of the Horse-
chestnut present this in a high degree ; those of the
Alder and Black Poplar are also invested with a
resinous matter, which defends the young shoots from
moisture.

With regard to temperature, the superposition of the
scales is a means cf protection from cold, because each
of them incloses a certain quantity of air ; moreover,
several buds are invested externally with a close down,
and some have the inner cavity full of a soft thick down

288 VEGETABLE ORGANOGRAPHY.

resembling cotton, which surrounds the young shoots,
and protects them, like fur, against frost, and also a
little against moisture. The buds of the Horse-chest-
nut are also examples of this structure. There are
some shrubs, as a species of Viburnum, where, accord-
ing to Kceler, the scales are wanting, and are replaced
by a cloth -like down.

It is on seeing this important character of buds, and
comparing their existence in different plants, that we
have been led to conclude that trees devoid of scaly
buds cannot live in cold climates, and that those of
warm countries can only be acclimatized in the north,
which are furnished with them. These two rules are
generally true, but they are subordinate to the peculiar
nature of the leaves of each species, and both present
exceptions. Thus, Viburnum Lantana and Rhamnus
Frangula, although natives of cold countries, have no
scaly buds; and Palms, though furnished with petio-
laceous buds, cannot exist in the north.

It would be curious to follow the vegetation of trees
of species like our own in very different climates, in
order to know — 1st, if the outer leaves of the young
shoots continue to be changed into scales, when the
trees are placed in a warm and very fertile climate ? 2d,
if certain trees, which in such climates do not present
this transformation, would not be liable to undergo it
in less warm climates, and if they could not be induced
to do so by culture ? If these two questions could be
answered in the affirmative, the field of naturalization
would be found greatly extended.

The buds of trees usually proceed from the axils of
the leaves, and consequently the disposition of the
young shoots is determined by that of the leaves; but of
the buds which are developed, a great number sooner or
later become abortive, whence it results that the branches

OF BUDS. 289

of trees are frequently irregular, although they were
originally disposed in regular order.

Besides the buds evidently axillary, certain trees
present terminal ones, which are generally larger and
more forward than the others ; they are met with in
trees both with opposite and alternate leaves. In the
first case three buds grow at the top of the branch, viz.
the terminal one, and the two which are produced in
the axils of the upper leaves : these three are rarely
developed together ; the two lateral ones are sometimes
abortive, and the terminal one alone is developed, as in
the Horse-chestnut, Pavia, the Sycamore, &c. Some-
times the terminal one is abortive, and the lateral ones
are developed, whence a bifurcation results, as in the
Lilac. The same differences take place in trees with
alternate leaves. Thus, the terminal bud continues the
branch in the Holly, Oak, Peach, &c. ; it is absent or
abortive, and the branch is continued by the buds of the
upper axils in the Apricot, Rose, Hazel, &c.

The development of the buds of a branch in spring
commences almost always at the apex, and proceeds
downwards, so that the lower buds are the last to shoot,
and sometimes are not developed. This appears to
result from the upper part being more herbaceous, and
consequently more sensible to the action of the warmth
of the atmosphere ; whence it results that an equal
degree of warmth applied to a whole branch has a
greater action upon each bud in proportion as it is
nearer the top. The exceptions even confirm this rule ;
for in the trees where the branches are of an equal
degree of hardness, or, as gardeners say, equally ripened
throughout, the buds follow the contrary order of de-
velopment, being influenced by the ascending sap ; as in
the Larch, Gincko, &c.

The buds of Dicotyledonous trees differ from one

VOL. II. v

290 VEGETABLE ORGANOGRAPHY.

another according to the nature of the young shoots they
are destined to protect : some inclose branches furnished
only with leaves and devoid of flowers ; these are called
Leaf-buds : the others contain only bunches, umbels,
or heads of flowers, and are called Flower-buds ;
some contain, at the same time, leaves and flowers, and
for this reason are named Mixed Buds. The first are
generally recognized by their long pointed form; the
second, by being round ; and the last, by an intermediate
form. It is evident that the distinction between leaf
and flower buds is only possible in trees where the
flowers arise independently of the leaves, as in the
Cherry, Apple, &c, and that the mixed buds are the
only ones which can be found in the trees where the
flowers grow upon the same branches as the leaves. In
the first, the position of the two kinds of buds is
determined beforehand, and the period of development
of each frequently disturbs the usual order of their
evolution from the top to the bottom.

As to the young shoots which proceed from each bud,
the development takes place from the base upwards, to
whichever class they belong : the scales of flower-buds
ought to be considered as the rudiments of bracts, and
in many respects these buds might be compared to
involucra; there are really no other differences than
that the floral buds are usually caducous, whilst invo-
lucra are ordinarily persistent: but there are many
intermediate cases in duration and appearance ; thus the
envelope which surrounds the flowers of Cornus mascula
and Aucuba is sometimes called bud, at other times in-
volucrum, and both terms are, in fact, equally applicable
to it. We might very well say, that the head of flowers
in the Compositse and Dipsaceae is a kind of bud.

When the petioles of trees are dilated at their base
into a sheath, this surrounds the young shoots and often

OF BUDS. 291

acts the part of a "bud ; sometimes it surrounds it so
completely, that no bud is perceived in the axil, but it
appears lodged in a cavity of the petiole formed by the
two edges of the sheath folded upon one another. This
is seen very clearly in Smilax aspera, where the peti-
olary sheath surrounds the young shoot, and remains
upon it until the spring. We meet with an analogous
and still more singular fact in the Plane ; here the leaves
fall off in the Autumn, so that the bud is not protected
as at its first development : the margins of the petiolary
sheath are completely united together so as to seem to
inclose the bud ; but if we make an examination just
before the fall of the leaves, we see that the base of the
petiole is split longitudinally on the upper side, exactly
at the place where the theory indicates that the two
margins of the sheath ought to be found. Phenomena
analogous to the preceding are met with, with slight
variations, in Negundo, Pkiladelphus, Robinia, and some
species of Sumach (Rhus).

The buds which are developed in perennial herbs,
either on a level with the surface or under ground, differ
the more from aerial ones, in proportion as they are
more decidedly subterranean. The more a vegetable
surface is deprived of the action of air and light, the
paler will it be, the less will it evaporate, and conse-
quently it will take, according to the texture of its
organ and the period of its vegetation, either the appear-
ance of a simple membrane, or that of an etiolated body,
but, however, full of juice.

If we compare the aerial buds of the tree Poeony
with those at the neck of the herbaceous ones, (PL 15,
Fig. 1, 2, 3,) it will be impossible to perceive any other
differences between them than those which result from
their position ; and all the turiones, or buds on a level
with the ground, in perennial non-bulbous plants, hardly

u 2

292 VEGETABLE ORGANOGRAPHY.

present any other differences ; but they usually take the
name of Bulbs (bulbi) when they present certain pecu-
liarities which deserve to be studied.

1st. A small number of Dicotyledons, which are
called bulbous, have received this term from a double
peculiarity of their organization, viz., that their leaves
have a petiole flattened at the base, more or less sheath-
ing, and their stem is swollen up above the neck into a
kind of tubercule ; it results from this, that this tuber-
cule, covered by the petiolary sheaths, resembles the
bulbs of several Monocotyledons ; such is the structure
of Ranunculus bulbosm, Fumaria bulbosa, &c.

2d. Several Monocotyledons present an analogous
disposition, that is to say, they have the leaves sheathing
at the base (which is frequent in this class), and at the
same time the base of the stem is swollen into a tuber-
cule, as in several Irideae, whence results a kind of bulb,
which many naturalists call a Bulbo-tuber.

3d. True bulbs present a very short subterranean
stem, reduced almost to a simple plate ; from it the
leaves arise in great numbers, overlaying one another,
and thus forming an oval or round body ; the external
leaves are reduced to the state either of fleshy scales, as
in the Lily, and then the bulb is said to be Scaly ; or
of short truncated membranous sheaths, as in the
Hyacinth, when it is said to be Tun icated. In these last
the base of the sheaths, and especially that of the inner
ones, is fleshy, as in the scales of the Lily, and approaches
them in texture although distinct in form. The inner
sheaths have a tendency to elongate into true leaves,
and all the radical leaves of the Liliaceee are prolonga-
tions of the inner pieces of this bulb. An Onion of the
first year is nothing but a terminal bud, situated at the
top of an extremely short subterranean stem.

When this organization is compared with that of a

OF BUDS. 293

Palm, we shall find, in this respect, that there is no
other difference but that the stem of the Palm is very
long, and, consequently, bears its bud very high, whilst
that of the Tulip is very short and leaves its bulb to be
developed under ground. Every intermediate state is
met with in species of the same class ; thus, we see the
stem take a greater elongation in certain species of
Allium, Crinum, Yucca, Draccena, &c. ; and we thus
arrive by insensible degrees from the scarcely visible
stem of bulbs to the long ones of Yucca, from the sub-
terranean buds of the Liliacese to the aerial ones of
Palms : we understand, then, how it happens that in the
same class there are sometimes very visible stems with-
out true bulbs, and sometimes bulbs without there
seeming to be any stem.

What are called Offsets are nothing but the axillary
buds of bulbs, or in other terms, the young branches
which are developed in the axils of the leaves; they pre-
sent this peculiarity, which probably results from their
position, that they are only attached to the stem by a
slender filament which is easily broken. As they have
their scales fleshy and full of nutriment, they can be de-
veloped by themselves as tubercules ; and this frequently
happens when they are separated, either artificially or
naturally, from the mother plant. The buds of Dicoty-
ledons, detached from the tree, may vegetate, provided
they are placed, by the operation of budding, in a
similar situation ; those of bulbous Monocotyledons
carry with them sufficient nutriment stored up to enable
them to continue to grow, especially where they find
heat and moisture.

We may distinguish in bulbous plants, as in trees,
leaf-buds, flower-buds, and mixed-buds ; thus, most
species of Amaryllis have at the same time the two first,
and the Tulip the last.

294 VEGETABLE ORGANOGRAPHY.

"What is remarkable in bulbs compared with buds is,
that their coats remain for many years : so that a bulb
is not only formed of the buds of the year, but of the
sheathing coats of preceding years, which are then ex-
hausted of all nourishment, but remain under the form
of membranes, and thus serve to protect the young offsets
either from cold, because they contain several strata of
air between their coats, or from wet, because their epi-
dermis is silicious and scarcely affected by moisture.
There are some bulbs which, like the buds of trees, pre-
sent a cottony layer between or outside their coats, as
in the Tulip.

It results from all that I have set forth in this chap-
ter, 1st, that buds are the integuments of the young
shoots formed by the external foliaceous organs, some-
times in their natural state, as the stipules of Ficus and
Magnolia, but most frequently converted into scales by a
kind of degeneration or semi-abortion caused by their
position ; 2d, that the buds of trees, exposed to the air,
and those which spring on a level with the surface, or
the turiones of perennial plants, and subterranean buds,
or the bulbs of the Liliacese, &c. only differ from one
another in what must necessarily result from their
position, and the stems which bear them.

CONCLUSION". 295

BOOK V.

CONCLUSION, AND GENERAL CONSIDERATIONS.

After having described all the organs of plants, and
having nearly come to the conclusion of this Work, I
ought now to give some general considerations, which
might have appeared either too hypothetical if I had
treated of them at the commencement, or too misplaced
if I had dwelt upon them occasionally. I shall proceed
to point out those general subjects which are as much
connected with Physiology as Organography.

296 VEGETABLE ORGANOGRAPHY.

CHAPTER I.

OF THE INDIVIDUAL PLANT.

What in the vegetable kingdom ought to be con-
siclered as an Individual ?

Illiterate, and even educated persons, accustomed to
see all the higher animals endowed with a life of their
own, have with difficulty believed that all which is pre-
sented under an analogous appearance can present
really different phenomena ; and have had much greater
difficulty in forming an idea of beings, in appearance
simple, but in reality an assemblage of individuals. They
have expressed great surprise when zoologists have
shewn that there exist apparently simple animals, but
which, in reality, are composed of several beings collected
together, living, however, with a common life : such are
the Botryllae, Pyrosomse, Polyclinums, and probably the
fresh-water Hydras and Polypes. Passing to the
vegetable kingdom, the question is whether plants are
single individuals, as vertebrate animals, or individuals
collected together, as the Polyclinums.

The common opinion is, that a Willow, Cherry, Cab-
bage, &c. are so many single individuals ; but when we
examine them, we find that they are singularly divisible :
almost all their parts are capable of being separated from
the whole, and of forming a new individual. This division
can go on ad infinitum; and there are examples, such
as the first Weeping Willow introduced into Europe,
(I select this example, because we only possess one

OF THE INDIVIDUAL PLANT. 297

of the sexes, and that does not seed,) from which, by
simple division, all the Weeping Willows existing in
Europe have been produced : all of these, then, are por-
tions of a single individual. The word individual, taken
in this sense, will be still more incorrect than if we con-
sidered a mountain of granite as a mineralogical indivi-
dual, divisible, at the will of man, into as many fragments
as he might l'educe it to by breaking the rocks.

Do we say that we only admit as distinct individuals
those plants which proceed from a seed ? This would
be an approach towards correctness ; for it is certain
that plants produced by simple division retain all the
peculiarities of the one of which they made part; whilst
those proceeding from seeds may present varieties or
differences, and seem to maintain in preference that
which forms the type of the species.

But how can trees which proceed from division, or from
seed, be distinguished when they are alike ? How can
this line of demarcation be drawn between this multitude
of beings, when we cannot distinguish seeds from bulbs
or spores ? How can this possibility of division, ad infi-
nitum, of an individual, supposed to be single, be ad-
mitted ? How can this definition be reconciled with
analogies elsewhere so remarkable as we have observed,
in the course of this work, between germs capable of
being developed either w7ith or without fecundation ?

All these difficulties vanish, on admitting that plants,
such as they appear to our eyes, are almost all aggre-
gations of individuals, and not single individuals.
Although allusions to this opinion are met with in several
authors, and particularly in the writings of Goethe, Mr.
Darwyn, on commencing his Phytologia with a chapter
on the individuality of buds, appears to me to be the
first who expressed it in general terms.

We consider, then, as an individual every developed

:298 VEGETABLE ORGANOGRAPHY.

germ, viz. ; 1st, sometimes a seed, supposing that, as
takes place in some annual plants, it produces a stem
without branches ; 2d, sometimes a branch considered
as a developed germ. Thus, in this sense, a tree is an
aggregation of the primitive individual proceeding from
the seed, and of all the individuals proceeding from un-
fecundated germs which are developed one upon another,
and have formed the prolongations or ramifications of
the primitive individual.

Cassini opposes this idea, and persists in the opinion
of the unity of the plant, founding it upon the continuity
of the fibres of the branches and of the trunk ; but this
continuity only proves, which no one can deny, that the
germs spring from the extremity of the fibres : in other
cases we find a continuity equally as great, at least from
our means of investigation, when we dissect a branch
proceeding from a grafted bud ; for, in this case, we
know perfectly well that there is a plurality of indi-
viduals and continuity notwithstanding. I do not think,
then, that the observations of this learned botanist can
modify the theory of Darwyn.

Each branch, or partial individual, presents great con-
formity of development with the primitive individual ;
its pith, full of juice, acts the part of a reservoir of nu-
triment ; and in Dicotyledons the two first leaves of
each branch are almost always opposite like the cotyle-
dons which they seem to represent.

Each partial individual, whether it originate from a
seed or from an unfecundated germ, is susceptible of
two kinds of terminations: — it is sometimes terminated
by a flower ; sometimes it is prolonged without flowering,
and only seems to stop from defect of nourishment. The
indefinite development of a branch requires more vege-
tative force, and is more frequent in young plants, and
in those growing in very damp situations. The termi-

OF THE INDIVIDUAL PLANT. 299

nation of a branch by a flower is more frequent in old
individuals and in those which have but little watery
nutriment. The indefinite development of branches
which do not flower favours the production and growth
of a great number of nutritive leaves, which tend to in-
crease the force of the aggregate, and to deposit here
and there stores of nourishment adapted to favour new
developments of germs or flowers. The termination of
branches by one or more flowers tends to deprive the
branches or trunks of the development of nutritive
organs, and to consume the deposits of nutriment which
may exist in the branches, stems, or roots.

When the flower only consumes the nutriment con-
tained in its peduncle or immediate support, these dry
up after flowering in the male flowers, and after the
maturity of the fruit in the females. But as the rest of
the plant has not been exhausted, it continues to vege-
tate, maintained by the branches which have produced
nutritive leaves, and the following year new germs are
developed. In this manner are formed trees, shrubs,
and under shrubs, or, in one word, all Caulocarpient
plants.

When the flowers are more numerous in proportion
to the force of the stem which bears them, they exhaust,
during the maturation of the seeds, not only the nou-
rishment deposited in their peduncles, but also all that
of the stem, which then perishes nearly down to the
neck ; and the following year the new buds arise from
the persistent part or stock ; this happens in perennial
herbs, or Rhizocarpient plants.

Lastly, if the flowers be still more numerous, or require
a greater quantity of nutriment in proportion to the
force of the stem which bears them, they exhaust, during
the maturation of their seeds, not only their peduncles
and stem, but also the root ; then, after the maturity of

300 VEGETABLE ORGANOGRAPHY.

the pollen in the male plants, or the seeds in the females,
the whole plant being exhausted dries up and perishes :
this forms plants said to be Monocarpient ; that is to
say, those which fructify but once at the end of one
year (annuals), of two years (biennials), or of several
years, as for example in Agave, &c.

These differences, although constant in each species,
since they are determined by causes inherent in its
structure, are nevertheless subject to modifications from
external circumstances. We can transform an annual
plant into a perennial one by preventing it from bearing
seeds ; thus, the Mignionette has been changed into an
under-shrub which, when once it has a woody stem,
flowers each year, without the exhaustion produced by
the flowering causing the stem to perish ; thus the dou-
ble Nasturtium has become perennial, because its
flowers, devoid of the faculty of producing seeds, do not
exhaust the stem ; and it is probable that every annual
plant, which may be rendered double by culture, will
become perennial.

We may likewise transform, in an analogous manner,
a perennial plant into an under-shrub ; this is frequently
the case in double pinks. Zizyphus presents a curious
phenomenon which renders it, as it were, intermediate
between Rhizocarpient and Caulocarpient plants ; we
see upon its old trunk kinds of exostoses, from which
proceed several simple branches, of which those which
bear a great number of flowers disarticulate, and fall after
flowering, exactly as the common petioles of pinnate
leaves ; whilst those which do not flower, lengthen,
remain upon the tree, and form true permanent
branches.

These details tend to prove that the differences in the
duration of plants only result very indirectly from their
anatomical structure, and serve to explain how, in the

OF THE INDIVIDUAL PLANT. 301

same natural families, we so frequently find plants of
various durations.

The individual plants proceeding from fecundated
germs (seeds), or unfecundated ones (bulbs/tubercules, or
young shoots), are some of them endowed with the faculty
of drawing up the sap by their own roots, and others
are devoid of this faculty, but are capable of receiving
the sap drawn up by the others ; thus, the individuals
proceeding from seeds are almost always provided with
roots to nourish them. The Misseltoe is an example of
a plant, which, although proceeding from a seed, has no
true root ; and its neck, implanted upon another plant, is
nourished at its expense, exactly as a bud inserted by
grafting. The individuals proceeding from bulbs or
tubercules are similar to those from seeds, as regards
the existence of roots.

Individuals produced in the manner of buds are con-
stantly devoid of roots, and are nourished by the sap
which is transmitted to them through the woody body
of the plant upon which they grow ; but if by any cause
the development of adventitious roots from the lenticils
be favoured there, then these individuals can live when
separated from the one which gave birth to them. The
processes by which we obtain new individuals are known
by the names of making Cuttings or Layers ; the graft
is nothing but the transplanting of a young shoot. The
laws relating to the duration of plants, or, rather, the
manners of expressing these laws, are subordinate to the
ideas adopted with regard to vegetable individuality; but
as this subject is entirely physiological, I shall here
confine myself to stating, from the preceding considera-
tions, that, with a small number of exceptions perhaps
even doubtful, plants are aggregations of as many indi-
viduals as there have been seeds or buds developed to
concur in their formation ; and that the plant is conse-

302 VEGETABLE ORGANOGRAPHY.

quently a compound, being analogous to the Polypes or
Botryllse in the animal kingdom.

This formation of new individuals, naturally grafted
upon that which gives birth to them, is not limited ; and
in this sense it is true to say, that if a tree be considered
as a single individual, its duration is infinite, and that it
only dies by accident. This proposition, which might
appear strange without reflection, is not in reality more
singular than if it were said that an aggregation of
animals which multiply without cessation, could not last
for an infinite period.

CHAPTER II.

OF VEGETABLE SYMMETRY.

When the kingdom of inanimate nature began to be
studied, there were only seen, as it were, irregularities, in-
termixed, however, here and there with more or less
evident symptoms of a regular order. How was it with
regard to astronomy ? The indications of a regular
order were evident ; but anomalies apparently inexplic-
able, such, for example, as the retrogradations of stars,
made persons believe that the laws would never be dis-
covered. These laws have been discovered, and the
apparent irregularities have become the most beautiful
confirmations.

How was it with respect to mineralogy ? The irre-
gularities were so numerous, and the cases of regular

OF VEGETABLE SYMMETRY. 303

forms so rare, that it seemed almost impossible to dis-
cover any general law ; by degrees it was perceived that
almost all, and probably all amorphous bodies might be
reduced to the form of crystals, and, consequently, that
regularity existed in their nature. Among crystals
themselves, it was perceived that a number of very
different forms were simple modifications of a few primi-
tive ones ; these primitive forms have not only been
reduced to a very small number, but the principal cir-
cumstances which determine the secondary forms have
been arranged ; and here, as in a great number of dy-
namic phenomena, it has been seen that the irregularities
result from the simultaneous action of several regular
causes which are increased and complicated in the
results.

If we examine more closely the progress of Crystal-
lography, we shall see that Rome-de-lTsle, considering
crystals as single bodies, explained their anomalies by
truncations, whilst Haiiy, going back in theory to the
primitive molecules, although they do not come under
our observation, was enabled to explain, in the most
happy manner, the most complicated forms, by referring
them to the different modes in which the molecules are
united together. The first reasoned as those botanists
do, who consider a leaf or corolla as a single whole,
notched upon its margins from some unknown cause ;
the second has served me as a guide, when I attempted
to show that the different indentations, or divisions of the
organs of plants, were essentially connected with the
various modes and different degrees of their aggregation.

There has been, then, a similarity in the progress of
these two sciences ; let us now see if there be any in
their nature.

Does this regularity exist in organized bodies, and
can the anomalies, so frequent in thenij be owing, as in

301 VEGETABLE ORGANOGRAPHY.

minerals, to complications of causes, each of which,
considered individually, would produce a regular effect ?
Those who believe most in the normal regularity of
organized bodies have perceived that it cannot be sub-
mitted to the same laws as that of minerals, &c. ; that
in particular no truly geometrical regularity is met with
in them ; but, although it is, perhaps, impossible to find
a flower with all its petals geometrically equal, or a leaf
with its two sides mathematically alike, it cannot be
denied that, even upon the most superficial examination,
we are not struck with the kind of regularity of these
organs. The name of Symmetry has been given to this
ungeometrical regularity of organized bodies. That
these, particularly vegetables, present a multitude of
examples of exact symmetry, is what no one can deny ;
and to these the name of regular bodies has been given,
to point out the fact, without pretending to compare
this regular symmetry or this regularity with the geo-
metrical order of crystals. But it cannot be denied that
in certain rather numerous cases the symmetry appears
deranged; does this symmetry any longer exist in the
cases of irregularity? or can this apparent derange-
ment of it be produced by regular causes ?

Even to the present day, the first of these opinions
has been held ; all the irregulai'ities of plants and ani-
mals have been described, without its being thought that
they conceal an order submitted to general laws. Each
unusual form of an organ received a new name, and it
became impossible to recognize their analogy with one
another. Each unusual form of a body was considered
either as a monstrosity if it were rare, (and the old bota-
nists were contented with this unmeaning word to dis-
pense with the study of it,) or as a distinct species, if the
phenomenon were frequent, and thus all correct means
of distinguishing organs were lost. They could not even

OF VEGETABLE SYMMETRY. 305

be classed with any method, for the least anomaly ob-
served between any two individuals or groups would
prevent all their other analogies from being recognised.

But the more the number of known beings increased,
and the more carefully they were studied, the more were
naturalists convinced of the principle which I have been
the first, or one of the first, to declare in general terms,
viz., that it is almost certain that organized beings are
symmetrical or regular when considered in their type,
and that the apparent irregularities of plants are con-
nected with phenomena constant within certain limits,
and capable of existing either separately or collectively,
such as the abortion or degeneration of certain organs,
their union with one another, or with others, and their
multiplication after regular laws.

I shall here confine myself to giving a few conside-
rations upon the importance and utility of this method
in studying plants, both as regards their organization
and classification.

The number of truly distinct organs is found to be
prodigiously reduced when we analyze their nature ; we
see that several to which an important character has been
attributed are only simple modifications of others ; and
we can recognise the same organ under different ap-
pearances, and consecpiently follow a true comparative
Organography.

All the numerous class of facts, known under the
name of monstrosities, which it was impossible to under-
stand by the old system, and were therefore left unstu-
died— all this class, I say, has been elucidated, and taken
a new interest, since they have been examined in their
true point of view, — as indications for recognising the
normal symmetry of beings. Monstrosities are, as it
were, experiments which nature makes for the profit of
the observer : here we see what are the organs, when

VOL. II. x

306 VEGETABLE ORGANOGRAPHY.

they are not united together, and we perceive what they
really are when an accidental cause has not prevented
them from enlarging. Proceeding thus from the opinion
that the primitive nature is symmetrical, that irregularity
is produced by different causes which alter this sym-
metry, we understand that monstrosities are owing to
certain variations of these causes, and that, consequently,
they may sometimes enable us to know the causes of
derangement when their action has been increased or
freed from all complication ; at other times they may
show us the symmetrical state when the causes which
change it have been weakened or destroyed.

The whole theory of natural classification evidently
rests upon the intimate knowledge of the organs and
their modifications. The arrangement of plants in na-
tural orders supposes, in my opinion, that we may one
day be able to establish the characters of these orders
upon what forms the base of their symmetry, and to
refer the various forms of the species or genera to the
action of causes which tend to alter the primitive sym-
metry. Thus, each leaf, like each class of crystals, may
be represented by a regular state, sometimes visible,
sometimes imaginary ; this is called its Type : unions,
abortions, degenerations, or multiplications, separate or
combined, modify this primitive type so as to produce
the constant characters of the beings they compose.
These modifications are constant within certain limits,
like the secondary forms of crystals. But each genus,
each species, is more or less submitted to the causes
which determine them, for plants which have the same
type are not more identical than the crystals which have
similar primitive molecules. If botany is much behind
mineralogy in this respect, it results, on the one hand,
from the much greater multiplicity of forms and causes
of action ; and, on the other, from all these facts being

OF VEGETABLE SYMMETRY. 307

submitted to a peculiar force (the vital force), the laws of
which are much more obscure and difficult to study than
those of affinity and attraction.

The simple description of vegetable facts and forms
has been singularly improved since the knowledge of
some general laws has caused describers to reflect upon
what they see. Those who refuse to believe these laws,
may, without doubt, describe the aberrations as the
natural state, because nothing causes them to doubt that
what they see is contrary to order ; they may easily
neglect minute organs, because nothing indicates their
existence, and they may take much trouble in describing
in detail certain peculiarities, which a few words,
founded upon analogy, would have expressed with more
clearness. Lastly, when two persons have described
the same object in a contradictory manner, which unfor-
tunately is not seldom the case, we have evidently no
other means of discerning the truth than by the greater
or less analogy of the descriptions with the laws of sym-
metry. For, in order to place plants in a rational order,
we most constantly decide it upon more or less incor-
rect descriptions, for the period no longer exists when
one man can see for himself all known plants.

Thus, in proportion as science makes new progress,
we more and more perceive the necessity of knowing
the general laws of organic symmetry. This necessity,
perceived by all those who love general truths, has
caused two divisions or schools to arise.

Some have endeavoured to establish laws upon the
structure of beings from general considerations, and, as
it is usually said, a priori. Others have attentively
observed the facts which would seem to remove laws
from regularity ; they have seen that by separating
almost all after uniform principles, and thus grouping
the apparent irregularities, they referred them again to

x 2

308 VEGETABLE ORCxANOGRAPHY.

regular laws, and ascending from partial to general
facts, they were enabled to recognise the laws of sym-
metry a posteriori.

Those who are attached to the second of these two
methods perform two parts in the general economy of
the science : on the one hand, they collect with care all
the facts in detail in order to derive general laws which,
gradually compared together, may lead to other laws a
little more general ; on the other hand, they examine
as simple hypotheses, to be verified or rejected, opinions
conceived a priori, and try to perceive in what points
the partial laws which they have recognised approach,
or from what they depart. This course appears to me
the same as that which is followed in all the physical
sciences, the only one which can lead to general truths.
If there still exist botanists who can believe, either
that there are no general laws in the structure of orga-
nized beings, or that it is not worth while to search
them out, I am persuaded that it can only result either
from their being frightened by the multitude of facts in
detail, or from their having studied only a small number
of objects selected without method from those found
within their reach.

I shall now proceed to give a summary of this work
under an aphoristical form, which may be able to
give some idea of these principles of symmetry ; this
will be the object of the following chapter.

GENERAL SUMMARY. 309

CHAPTER III.

GENERAL SUMMARY OF THE STRUCTURE OF PLANT-.

1st. A plant is an organized and living being, devoid
of voluntary motion, having neither nerves, muscles,
nor a central cavity resembling a stomach, and always,
or nearly always, attached to the soil from which it
draws its nourishment.

;3d. Plants are either wholly, or in a great measure,
composed of membranous Cellules, closed on all sides,
more or less united together, and enclosed, at least in
their young state, in a membranous cuticle. Those
which are entirely thus formed bear the name of
Cellular Plants (Book i. Ch. 2 and 16).

3d. Those which are thus formed in part, and which
are called Vascular Plants, present, besides the
cellules, cylindrical tubes which are called Vessels ;
these are never naked, but always surrounded by
cellules (Book i. Ch. o and 16).

4th. In vascular plants we observe moreover: — 1st,
that the cellules and vessels are united in very different
degrees, so as frequently to leave between them empty
spaces, called Intercellular Passages ; 2d, that
besides the purely membranous vessels, there are bodies
rolled spirally, and endowed with great elasticity, which
are called Tracheae ; «Sd, that their cuticle is pierced
(at least, in almost every part exposed to the air) with

310 VEGETABLE ORGANOGRAPHY.

pores or Stomata, which appear to be evaporating
organs (Book i. Ch. 2, 3, 5, G).

5th. The cellules are endowed with the faculty of
uniting together, of absorbing the moisture around
them, and probably of contracting and dilating. They
are round, or more or less elongated ; the former
inclose the feculent, mucilaginous, or resinous matters
which they have elaborated, of which the latter contain
little or none. The round ones form the parenchyma ;
the long ones (by themselves in cellular plants, united
with the vessels in vascular ones) compose the fibres or
nerves (Book i. Ch. ti, 4).

6th. The passages between the elongated cellules, or
the vessels, appear eminently to serve for carrying the
lymph, i. e. the as yet unelaborated watery juices.
Those which are formed among the round cellules con-
tain the more stagnant juices (Book i. Ch. 2).

7th. The vessels, whatever their form, seem eminently
intended to contain air or gas, and are true aerial canals,
at least in the ordinary course of vegetation (Book i.
Ch. 3).

8th. Certain particular points of the surface of plants,
and especially of vascular ones, are more eminently
endowed with the faculty of absorbing water. They are
called Spongioles, and arc situated at the extremity of
roots, at the top of the style, and on the surface of seeds
(Book i. Ch. 7).

9th. Dilatations of the intercellular passages, or, in
certain cases, ruptures of the cellules, cause irregular
cavities in the interior of the tissue. These receive the
name of Air-cavities when filled with air, or of Re-
ceptacles of proper Juice when they contain an
elaborated juice (Book i. Ch. 11, 12).

10th. Glands or glandular surfaces are some of them
composed only of cellular tissue, others of cellular tissue

GENERAL SUMMARY. 311

and vessels ; both secrete special juices, but the first appear
(at least in certain floral organs) to be excrementitial,
and the second recrementitial (Book i. Ch. 9).

11th. The surface of plants exposed to the air is often
invested with Hairs, which are prolongations formed of
projecting cellules. Some of these hairs are protecting
organs for the surfaces ; the others the supports or
canals of excrementitial glands. They are always situ-
ated upon the nerves, whilst the stomata are always
upon the parenchyma (Book i. Ch. 6 and 10).

12th. A vascular plant, considered lengthways, is
composed of two bodies opposed by their bases (stem
and root), and which grow in a contrary direction to one
another. Their point of junction is called the Neck
(Bookii. Ch. 1, 2).

13th. The body which descends, or the Root, elongates
indefinitely by its extremity alone; does not become
green by the action of the sun, except at its extremity ;
bears neither leaves nor flowers, and serves to fix the
plant in the ground and to draw up its nourishment
(Book ii. Ch. 2).

14th. The body which rises upwards, or the Stem,
elongates throughout its whole length till the period
when it ceases to grow, unless by the development of a
body resembling itself (branch) and which is grafted
upon it. It becomes green on exposure to the light
throughout its whole length, at least in its young state,
bears leaves and flowers, and transmits to them the
nutriment absorbed by the roots (Book ii. Ch. 1).

loth. The stem of vascular plants is sometimes
cylindrical, composed of a single system (the Woody
Body), which increases by the development of new
fibres internally ; sometimes conical and composed of
two systems (the Woody Body and Bark), winch
increase in diameter by means of layers which are

312 VEGETABLE ORGANOGRAPHY.

developed upon the surface of each of these systems
which is in contact with the other system. To the first
the name of Endogens is given, to the latter that of
Exogens. The structure of the root of each class is
similar to that of the stem (Book ii. Ch. 1, 2).

16th. The stem of vascular plants is furnished late-
rall}r with appendicular organs, which seem formed by
the expansion of one or more fibres (Book ii. Ch. 3).

17th. These appendicular organs, although very dif-
ferent from one another in their appearances and uses,
seem however entirely identical in their original nature
(Bookiii. Ch. 2, sec. 18).

Those which are already formed in the embryo, bear
the name of Cotyledons, or Seed-leaves; those
which are produced immediately afterwards, Primordial
Leaves. The following bear simply the name of
Leaves. Those which immediately surround the flower
receive the name of Bracts, and the flower itself is
composed of several verticils of appendicular organs,
much modified (Book ii. Ch. 3; Book iii. Ch. 1, 2).

18th. The appendicular organs perform, according to
their position and mode of development, several different
functions, of which the principal are : —

1st, That of nourishing organs, as the cotyledons and
leaves ;

2d, That of protecting organs, as the scales of buds,
bracts, sepals, petals, carpels in their last stage ;

3d, That of fructifying organs, as the stamens, and
the carpels during the first stage of their existence.
Several partake of both of these functions (Book ii.
Ch. 3; Bookiii. Ch. 1,2, 3).

19th. The nourishing appendicular organs are, at
their origin, alternate in endogenous plants, called also
for this reason Monocotyledons ; opposite or verticil-
late in Exogens, called also Dicotyledons. In the

GENERAL SUMMARY. 313

course of their development, those of Endogens always
remain alternate or spiral, those of Exogens may either
remain in their primitive state or take a spiral dis-
position (Book ii. Ch. 3).

20th. The appendicular organs which compose the
flowers are, in both classes, disposed in concentric ver-
ticils ; the innermost are sometimes spiral (Book iii.
Ch. 2).

21st. The protecting appendicular organs hold a middle
station, in form, size, colour, and often also in position,
between the two other classes ; and we frequently see
them metamorphosed, either into organs decidedly nou-
rishing, or more rarely into fructifying ones (Book iii.
Ch. 1).

22d. The appendicular organs are generally composed
of a petiole and limb, but one of them may be wanting.
The Petiole, which is the bundle of fibres not as yet
disunited, has its fibres longitudinal ; the Limb, which
is the part formed by the expansion of the fibres, has
them more or less diverging. These fibres of the limb,
or Nerves of leaves, are generally curved in Endogens,
and separate at angles more or less acute in Exogens
(Book ii. Ch. 3).

23d. The nerves of curvi-nerved leaves converge
towards the apex, or diverge from a middle bundle.
Those of anguli-nerved ones arc pinnate, palmate, or
pedate ; but the portions of the limb of the three last
classes are penni-nerved, so that this form seems essential
to the leaves of Dicotyledons (Book ii. Ch. 3).

24th. The leaves of Dicotyledons are the only ones
which have been seen, either composed of joints or
leaflets, or furnished with lateral stipules.

25th. Germs, or the undeveloped rudiments of new
individuals, appear able to arise from all parts of the
surface; but there are certain points where they are

314 VEGETABLE ORGANOGRAPHY.

developed in preference, such as the axils of the
appendicular organs and the extremities of the fibres of
their limbs (Book iii. Ch. 1, 5).

26th. The germs which are placed in the axils of the
appendicular organs, along the stem or petiole, may be
developed by the action of the nutritive forces alone.
Those which are situated at the extremity of the lateral
fibres of the limb almost always require (except in
Bryophyllum), in order to be developed, a particular
operation called Fecundation (Book iii. Ch. 5).

27th. The germs which are developed without fecun-
dation most frequently arise united to the mother plant
without having proper envelopes, and without shooting
out roots : they then form branches. Some separate
when they are furnished with a tubercule or store of
nutriment : they then form separate individuals and
produce roots (Book iii. Ch. 5).

28th. Every stem or branch can shoot out adventitious
roots. In Dicotyledonous trees, these spring from the
lenticels ; every branch, furnished with them or capable
of producing them, may easily be separated from the
mother plant and form a distinct being (Book iii. Ch. 5),

29th. The germs which are developed by fecundation
are always contained in a closed envelope, furnished
with the rudiments of a root and appendicular organs.
They receive the name of Embryos (Book iii. Ch. 4).

30th. The unfecundated germs perpetuate the varieties
of the mother plant; the embryos only retain the
characters of races or species.

31st. The appendicular organs which immediately
surround the flowers, or the bracts, hardly ever have
leaf-buds developed in their axils ; this is still more
seldom the case in the appendicular organs which com-
pose the flowers (Book iii. Ch. 1 , 2).

32d. The buds, or germs, which are developed into

GENERAL SUMMARY. 315

branches, are often protected in their young state by
scales which are nothing but the outermost appendicular
organs of the young branch, modified by their position
(Book iv. Ch. 6).

33d. The flower, in which is the apparatus destined
for the fecundation, is a kind of terminal bud formed of
verticillate appendicular organs, the outermost of which
act the part of protecting organs, the innermost of
sexual ones ; but they are capable of changing their
office by being transformed either into leaves, or from
one into another (Book iii. Ch. 2).

34th. In the modifications or transformations of the
appendicular organs, each is only usually converted into
the nature of the verticil which follows or precedes it in
the order of development or position. The first phe-
nomenon, which is the most frequent, has received the
name of Ascending or Direct Metamorphosis, and
the second that of Descending or Retrograde Meta-
morphosis (Book iii. Ch. 2).

35th. The flower being formed of verticillate organs
is necessarily terminal with regard to the pedicel, at
least when the pedicel is not prolonged beyond it, as
happens accidentally in certain proliferous flowers
(Book iii. Ch. 1).

36th. Pedicels near one another, and composing the
same inflorescence, are disposed after three systems : —
1st, the outer or lateral ones are developed first, and the
flowering proceeds indefinitely in a centripetal order;
2d, the central one, which is necessarily terminal, flowers
first, and the flowering proceeds in a centrifugal order;
3d, these two laws are combined, the one affecting the
general axis, the other the lateral branches (Book iii.
Ch. 1).

37th. The number of verticils in phanerogamous
flowers, is usually four ; but it may vary, being either

316 VEGETABLE ORGANOGRAPHY.

less when one is absent or united to the neighbouring
one, or more when one is composed of several verticils or
similar rows (Book iii. Ch. 2).

08th. The almost universal disposition of the pieces
of each verticil or row, is that of being alternate with
those of the preceding verticil or row (Book iii. Ch. 2).

39th. The number of pieces of each floral verticil is
generally three in Monocotyledons, and five in Dico-
tyledons (Book iii. Ch. 2).

40th. All the caulinary, and especially the appendi-
cular parts of plants, are capable of being united
together, especially during their infancy ; the union is a
distinct phenomenon from the graft ; it is the more easy
in proportion as the nature of the organs is more
analogous; it takes the name of Cohesion when it
comes between similar organs, and Adhesion when
they are different. The different degrees of adhesion of
similar organs, or of the parts of the same organ,
determine either its integrity, or the divisions or in-
dentations of most organs.

41st. All the caulinary or appendicular parts are
capable, when they are filiform, of expanding into
limbs ; and when naturally in the form of a limb, of
presenting a cylindrical appearance. They may also
put on, within certain limits, forms, sizes, texture,
colours, and even functions and positions, varying in
different points of the same individual or analogous ones ;
this constitutes the Degenerations or metamorphoses
of organs.

42d. All the appendicular organs, verticillate or spiral,
are capable of presenting multiplications of number,
both in the increase in the number of the verticils or
spires, or in the increase in that number of the pieces in
each of the systems.

43d. All the organs of plants are susceptible of being

GENERAL SUMMARY. 817

abortive either wholly or in part, and, consequently, of
presenting simple rudiments or leaving empty spaces.

44th. All the irregularities observed in the symmetry
of verticillate organs, and especially in that of flowers
and fruits, appear to result from one of the causes
mentioned in the four preceding paragraphs, or from the
combination of several of them.

45th. In particular, the unity or solitariness of the
verticillate organs can only exist by the abortion of
those which ought to complete the verticil or spire, or
by the union of several.

46th. The fruit is formed by the Carpels, which may
be free, or cohere together, or adhere to neighbouring
parts (Book iii. Ch. 4).

47th. As the two margins of each carpellary leaf can
bear ovules, the solitariness of the seed in a carpel, free
or united to others, can only result from an abortion
(Book iii. Ch. 3).

48th. The embryo must be considered as the de-
velopment, by fecundation, of a germ situated at the
extremity of one of the lateral fibres of the carpellarv
leaf (Book iii. Ch. 4).

49th. Cryptogamous plants present, in their organi-
zation, only partial indications of symmetry, which, in
the present state of the science, are not sufficient to
enable us to recognise the laws. We cannot affirm
particularly whether there is fecundation in all Crypto-
gamous plants, or whether several are not reproduced
by unfecundated germs.

ERRATA.

Vol. I. p. 133, line 20, for " annular," read " annual."

144, line 7, for " Alburum," read "Alburnum."

267, line 5 from bottom, for " two pair," read " one pair."

278, the last line, for " Euphorbium," read " Euphorbia."

279, line 10, for " Chicorium," read " Cichorium."
309, line 14, /or " Polytricha," read " Polytrichum."

Vol. II. p. 7, line 18 from bottom,/or " Veronica hederacea," read " V.
hederifolia."
59, the last line, for " Berberidse," read " Berberidese."
176, line 23,/or " ^sculus," read " Castanea."
212, line 21, for " Mullow," read " Mallow."

ALPHABETICAL TABLE

OF THE

GENERA AND FAMILIES MENTIONED AS EXAMPLES IN THE
COURSE OF THIS WORK.

ABIES (Fir), i. 36, 138, 175, 207,

228, 234, 287, 296, 301 ; ii. 11, 18,

175, 205.
Abrus, ii. 189.

Acacia, i. 242, 243, 247, 2G8, 283.
Acer, ii. 285.

Aconitum, i. 297; ii. 68, 149.
Acorns, i. 191.
Acro.sticlanii, ii. 228.
Adansonia (Baobab), i. 169.
Adenanthcra, ii. 58, 117.
sEschinomene, ii. 64.
AZsculus (Horse-chestnut), i. 176, 177,

264; ii. 176, 186, 209, 210.
Agaricus,i. 321; ii. 21".
Agave, i. 56, 58 ; ii. 264..
Agdestis, ii. 83.
Agrostis, ii. 12.
Ailantus, i. 150, 228.
Ajuga, i. 277.
Alchemilla, ii. 70, 164.
Alct., i. 11, 15,35, 51, 307, 322, 326 ;

ii. 251.
Alisma, i. 244; ii. 68, 1 17.
Alismagp.e, ii. 88, 194.
Alkekenge ; see Physalis.
Allium (Onion), i. 128, 196, 224, 273,

ii. 196.
Almond; see Amygdalus.
Alnus (Alder), ii. 287.
Aloe, i. 114, 128, 196, 246, 247, 279.
Alsine, i. 42.
Althaa, i. 250.
Alyssum, ii. 39.
Amaioua, ii. 151.
Amaranthacee, ii. 64, 88, 146.
Amaranthus, ii. 59, 120, 137, 141.
Amaryllis, ii. 217.
Ambora, ii. 37, 174.
Amentace*:, i. 282, 2S4, 287 ; ii. 3S,

83.

Ammannia, ii. 23.

Ammodendron, ii. 262.

Amome.e, i. 271, 297; ii. 89.

Amomum, i. 33, 196.

Ampelide.e, i. 251.

Amvgdalace.e, ii. 104, 141.

Amygdalus, i. 180 ; ii. 136, 141.

Anacardium, ii. 39, 171.

Anagallis, ii. 15S.

Anemone, i. 274; ii. 83, 98, 119, 122.

Anona, ii. 68, 148, 17'i.

Anon.ue.e, ii. 160, 161, 196.

Anthemis, i. 104 ; ii. 16.

Anthericum, i. 196.

Anthoceros, i. 316; ii. 246.

Anthyllis, i. 265, 292.

Antirrhinum, ii. 103.

Apocyne.*:, i. 103, 105, 110; ii. 107,

147, 149, 181.
Apricot ; see Armcniaca.
Apuleia, ii. 167.
Aquilegia (Columbine), ii. 68, 75, 84,

99, 119, 162.
Arahis, ii. 212.
Arachis, ii. 165.
AraliacejE, ii. 106.
Ardisia, ii. 186.
Arenaria, ii. \~> 1.
Argemone, ii. 153.
Armeniaca (Apricot), i. 295.
Aroide/E, i. 246, 252, 259 ; ii. 46.
Artichoke ; see Cinara.
Artocarpe;e, i. 110.
Artocarpus (Bread-fruit), ii. 174.
Arum, i. 129 ; ii. 12, 120.
Asclepiade.e, ii. G2, 73, 149.
Asclepias, ii. 73, 136, 142, 149.
Ash ; see Fraiinus.
Aspalathus, i. 267, 281, 282.
Asparageje, i. 193, 194, 196, 282;

ii. 195.

320

ALPHABETICAL TABLE OF GENERA, ETC.

Asparagus, i. 206, 207. 248.

Asperula, ii. 1 13.

Asphodele^:, i. 193, 197, 207.

Asphodelus. i. 220.

Asplenium, ii. 231, 232.

Astartea, ii. 92.

Aster, I 197; ii. 99.

Astragalus, i. 236, 244, 261, 268, 285 ;

ii. 70, 138.
Astrocarpus. ii. 137.
Atriplex, ii. 120, 169.
Aurantiace/e, i. 266; ii. 76, 186.
Avena (Oat), i. 200.
Azolla, ii. 226.
Balsamina (Balsam), i. 11, 38, 41,

114, 116; ii. 156.
Ba?nbusa (Bamboo), ii. 91.
Banana ; see Musa.
Baobab ; see Adansonia.
Barberry ; see Berberis.
Bar/eria, ii. 42.
Barnadesia, ii. 65.
Batrochospermese, i. 323.
Bauhinia, i. 192, 250, 261, 269, 270 ;

ii. 43, 165.
Beech ; see Fagus.
Begonia, i. 291 \ ii. 120.
Bellis (Daisy), ii. 38, 130, 169.
Berberide.e, ii. 51, 59, 94.
Berberis (Barberry), i. 207 ; ii. 5'J,

92, 200.
Betula (Birch), i. 172, 176.
Bignonia, i. 240; ii. 182.
Birch ; see Betula.
Bistort ; see Polygonum.
Blighia, ii. 180.
Blitum, ii. 169.
Boletus, i. 321 ; ii. 249.
Bombace/e, ii. 180, 196.
Bombax, i. 103, 155.

BoRRAGINEiE, ii. 20.

Borrago (Borage), i. 210; ii. 57.

Box ; see Burns.

Brassiea (Cabbage, Turnip, &c), i.

219, 272; ii. 152, 205.
Bromelia (Pine-Apple), i. 196 ; ii. 8,

174.
Broom ; see Spartium.
Bryonia (Bryony), i. 132.
Bruophillum, i. 238; ii. 69, 129.

216.
Bryum, i. 309, 311.
Buck-wheat ; see Polygonum.
Bunias, ii. 212.
Bupleurum, i. 241, 243, 247, 289 ; ii.

43.

Eutome.i, ii. 155, 178.

Butomus, i. 34.

Buxbaumia, i. 309, 312.

Buxus (Box), i. 158.

Byttneriace^e, ii. 97.

Cabbage ; see Brassiea.

Cacalia, i. 147.

Cactus, i. 134, 136, 215, 305 ; ii. 204,

207.
Cajamis, ii. 209.
Caladium, i. 203; ii. 12, 120.
Calamus, i. 193, 200, 205.
Calendula (Marygold), ii. 130.
Calytrix, ii. 270.
Calla, i. 114, 118; ii. 12.
Callistemon, i. 176 ; ii. 8.
Campanula, ii. 53, 57, 73, 94, 122,

124.
Campanulace.e, ii. 66, 72, 164,

166.
Candytuft ; see Iberis.
Candle-plant ; see Cactus.
Cane, i. 131.
Carina, i. 33.
Cannabis (Hemp), i. 56, 58, 110, 170 ;

ii. 83, 1S7.
Capparide^:, i. 293 ; ii. 69, 76, 78,

87, 153, 158, 163.
I <r i iris, i. 293 ; ii. 30.
Caprifoliace.e, ii. 78, 164.
Capsella, ii. 85, 128.
Caragana, i. 42.
Cardiospenuum, ii. 270.
Cardopatum, ii. 21, 26.
Carduus (Thistle), i. 128, 147, 148;

ii. 16.
Carex, i. 222 ; ii. 17, 162.
Carlina, i. 128.
Carpesium, i. 104.
Carrot ; see Daucus.
CARYOPHYLLE^.i. 132, 178, 280, 283 ;

ii. 19, 66, 82, 87, 88, 153, 154, 158,

195.
Cassia, i. 265 ; ii. 142.
Cassytha, i. 305.
Castanea (Chestnut), i. 147, 148,150,

155,249; ii. 43, 176,208.
Casuarina, i. 135, 305.
Catalpa, i. 147.
Catananehe, ii. 81.
Cedar, i. 279, 282.
Celastrine^e, ii. 78.
Centaurea,u. 81,167, 168, 19S.
Centranthus, ii. 110.
Cephalanthus, ii. 16.
Cephalotus, i. 272.

ALPHABETICAL TABLE OF GENERA, ETC.

32]

Ceramium, ii. 254.
Cerastium, ii. 333.
Cerasus (Cherry), i. 138 ; ii. 14, 70,

104,111, 121, 136, 141.
Ceratophyllum, i. 36, 74.
Cerberus, ii. 149.
Cercis, i. 155.
Ceropegia, i. 305.
Chailletia, ii. 31.
Chantransije, ii. 258.
Chura, i. 35, 322 ; ii. 252.
Cheiranthus (Wallflower), i. 11, 90 ; ii.

124.
Chelidonium, ii. 180.
Chenopodeje, ii. 196.
Chenopodium, i. 106 ; ii. 86.
Cherry ; see Cerasus.
Chestnut ; see Castanea.
Chondrilla, ii. 168.
Chrysanthemum, i. 104.
Chrysogonum, ii. 167.
Chuquiraga, ii. 263.
Cicer (Chick-pea, &c), i. 90, 93.
Cic Horace.?;, i. 110, 186.
Cichorium (Chicory), i. 186, 210, 279.
("niara (Artichoke), ii. 16.

ClNAROCEPHAL^E, ii. 62.

ClNCHONACEiE, ii. 195.

Cineraria, i. 86.

Cistine;e, ii. 104, 109.

Citrus (Orange, &c), i. 225, 234,240,

266; ii. 163, 186.
Cladonia,\. 318.
Clavaria, i. 321 ; ii. 249.
Claytonia, ii. 333.
Clematis, i. 185 ; ii. 87, 106, 119, 140,

145, 199.
Cleome, ii. 69.
Cliffortia, ii. 41.
Clinopodium, ii. 23.
Clitoria, ii. 139.
Clusia, i. 224.
Cneorum, ii. 126.
Cnicus, ii. 81.

Cobaa, i. 138; ii. 62, 116, 144.
Cocos (Cocoa-nut), ii. 193.
Coffea (Coffee), ii. 150, 195, 198, 199.
ColchicacejE, i. 120 ; ii. 156.
Colchicum, ii. 70, 96, 137, 156.
Columbine ; see Aquilegia.
Colutea, ii. 62, 64, 139.
Combketaceje, ii. 88, 212.
Commeline.e, ii. 89.
Composite, i. 280, 296; ii. 16, 21,

26, 37, 53, 55, 65, 72, 81, 82, 99,
VOL. II.

105, 107, 146, 169, 171, 179, 182,

184, 194.
Conferva, i. 324; ii. 257.
Conifer/E, i. 36, 86, 110, 111 ; ii. 11,

83, 175, 193, 198, 205.
Coniocarpon, i. 317.
Conium, ii. 167.
Conocarpus, ii. 16.
Convallaria, i. 281.
Convolvulace/e, ii. 194.
Convolvulus, i. 137, 140; ii. 210.
Cork-tree ; see Quercus.
Cornus, ii. 120.
Correa, i. 120.

Corylus (Hazel), i. 30; ii. 11,43, 171.
Corymbium, ii. 21.
Cosmibuena, ii. 167.
Cotton ; see Gossypium.
Cotyledon, ii. 92.
Coulteria, ii. 264.
Cowslip ; see Primula.
Crambe, ii. 179.

Crassula, i. 215, 288, 289 ; ii. 92.
&IASSULACE2E, ] 136; ii. 68, 70,73,

92 ; ii. 116, 165.
Crataegus (Thorn), i. 147, 150.
Crinum, i. 114, 115, 196; ii. 217.

daria, ii. 138.
Croton, i. 94.
CruciancUa, ii. 12.
Crucifer^, ii. 28, 40, 64, 97, 116,

155, 157, 178, 194, 211,212.
Cryptogam/e, ii. 3.
Cucumis, ii. 150.
Cururbita, (Gourd), i. 11, 47; ii. 60,

65, 124, 161, 165, 180.
Cucurbitace.e, i. 284, 293; ii. 62,

78, 83, 161, 165, 166.
Cupressus (Cypress), i. 36 ; ii. 175.
Cuscuta (Dodder), i. 225, 305 ; ii. 159,

203, 204, 207, 232.
Cuviera, ii. 2d5.
Cyatheea, i, 201 ; ii. 230, 232.
Cycade^:, i. 193, 297; ii. 193, 198.
Cycas, i. 194; ii. 206.
Cyclamen, 5. 128, 242 ; ii. 38, 39.
Cydonia (Quince), ii. 159.
Cynomorium, i. 134.
Cyperace^:, i. 105, 222, 245, 273 ;

ii. 46, 162.
Cypress ; see Cupressus.
Cysticapnos, ii. 136.
Cytisus, i. 267; ii. 64, 212.
Daisy ; see Bellis.
Dalbergia, ii. 64.
Y

322

ALPHABETICAL TABLE OF GENERA, ETC.

Dahlia, i. 220.

Dandelion ; see Taraxacum.

Daphne, ii. 86.

Darea, ii. 232.

Datura, ii. 94, 155.

Daucus (Carrot), i. 210, 219, 226.

Davallia, ii. 231.

Deal ; see Abies.

Delphinium (Larkspur), ii. 119, 137,

140, 147.
Desmodium, i. 240, 267.
Detarium, ii. 141.
Dianthus (Pink), i. 215; ii. 20, 42,

55, 64, 93, 107, 215.
Diatoma, i. 326.
Dicksonia, i. 129, 201.
Dicrdnum, ii. 241.
Dictamnus, ii. 122.
Digitalis (Foxglove), ii. 9, 102.
Dillenia, ii. 148, 176.
Dioneea, i. 240.
DioscoRE«,i. 196,253, 281.
Diospyros (Ebony), i. 147, 155, 158;

ii. 82.
Diplolana, ii. 80.
Diulostachyum, ii. 233.
Dipsace^e, ii. 16, 17, 37, 81, 82, 146,

171, 179, 182.
Dipsacus, i. 186 ; ii. 37.
Dodder ; see Cuscuta.
Dodoncea, ii. 150, 200.
Dolichos, ii. 180.
Dorstenia, ii. 37, 174.
Dracana,i. 179, 180, 193, 194, 196,

205, 278.
Dracontium, i. 261.
Drosera, ii. 20.
Droseraceje, i. 297.
Dry as, ii. 140.
Durio, ii. 60.
EbENACEjE, ii. 82.
Ebemis, ii. 210.
Ebony : see Diospyros.
Echinophora, i. 223.
Erhinops, ii. 26, 44, 171.
Echium, i. 185; ii. 20, 25.
El.eagne^e, ii. 88.
El&agnus, ii. 88,281.
Elder ; see Sambucus.
Elm ; see Ulmus.
Empetrnm, ii. 199.
Entada, ii. 137, 267.
Ephedra, i. 135, 178, 305.
Epidendrum, i. 246.
Epilobium, ii. 169.

Epimedium, ii. 108.

EquisetacejE, ii. 223 — 225.

Equisetum, i. 138, 200, 205, 305.

Erica (Heath), ii. 124.

Ericaceae, ii. 94, 157, 164.

Erineam, i. 320.

Eriodendron, ii. 60.

Eriophorum, i. 103.

Erodium, ii. 210.

Ervum, i. 293.

Eryngium (Eryngo), i. 131, 186, 223,

228 ; ii. 15, 16, 37.
Erysimum, ii. 122.
Erythrcea, ii. 19.
Erythrina, ii. 189.
ErythroxylEjE, i. 286 ; ii. 282.
Erythroxylon, i. 285.
Eucalyptus, ii. 35, 50, 51.
Eucomis, ii. 8, 23, 174.
Eugenia, ii. 23, 24, 35, 210.
Eunomia, ii. 178.
Euonymus, i. 295.
Euphorbia, i. 278, 279, 305 ; ii. 19, 21,

43, 124, 125, 126, 128, 130, 140,

156, 187, 203, 204, 213.
EuphorbiacevE, i. 110; ii. 67, 71,83,

88, 156, 194, 195.
Eupomatia, ii. 75, 196,
Exoacantha, ii. 42.
Faba (Bean), ii. 191, 209.
Fagara, i. 240.
Fagus (Beech), i. 150, 163, 166,

295.
Favonium, ii. 81, 167.
Ferns ; see Filices.
Ferula, i. 146, 147.
Festuca, i. 237.
Ficaria, ii. 216.

Ficoid ; see Mesembryanthemum.
FicoidEjE, ii. 87.
Ficus (Fig), i. 136, 137, 147, 224,

285 ; ii. 37, 39, 173.
Filices (Ferns), i. 137, 201, 208,

222, 224, 263, 297; ii. 208, 227,

232.
Fir ; see Abies.
Fissidens, i. 313.
Flacourtiane^:, ii. 155, 178.
Flagellaria, i. 247.
Flax ; see Linum.
Floride;e, i. 323, 324.
Fontinalis, i. 311.
Fragaria (Strawberry), i. 136; ii. 68,

140, 161.
Fraxinus (Ash), i. 141, 147; ii. 85.

ALPHABETICAL TABLE OF GENERA, ETC.

323

French-bean ; see Phaseolus.

Fritillaria, i. 196; ii. 116.

FucACEiE, i. 323, 324.

Fucus, i. 113, 225, 323, 324; ii. 254.

Fumaria, ii. 65, 206.

Fumariace;e, ii. 65, 158.

Fungi, i. 11, 35, 51, 55, 107, 307,

319—322 ; ii. 249.
Galeopsis, ii. 103, 111.
Galium, i. 137, 276, 286; ii. 195,

212.
Gardenia, ii. 79, 167.
Genista, i. 267.
Gentiana, ii. 95, 125.
Geoffraa, ii. 111.
Geraniace^e, i. 132, 286: ii. 67,

160.
Geranium, i. 140; ii. 71, 215.
Geum, ii. 140.
Gincko,\. 141, 252.
Gladiolus, ii. 12.

Gleditsia, i. 263, 268 ; ii. 82, 104.
Globulea, i. 277.
Glottidium, ii. 139.
Glyceria, ii. 91.
Gomphia, i. 286.
Gossypium (Cotton Plant), i. 103 ; ii.

181, 204.
Gourd; see Cucurbita.
Gramine.e (Grass Tribe), i. 25, 105,

193, 198, 205. 219, 245, 246, 247,

253, 271, 273, 289; ii. 11, 17,46,

91, 115, 146, 195, 209,215.
Grewia, i. 291.
Grimmia, ii. 241.
Grumilia, ii. 196.
Guiacum, i. 147.
Guilandina, ii. 189.
Gundelia,n. 45, 112, 173.

GuTTIFER/E, i. 110.

Gymnostomum, i. 313 ; ii. 241.
Gymnostyles, ii. 85.
Gynandropsis, ii. 76.
Gyrostemon, ii. 47.
Hcematoxylon, ii. 142.
Halimodendron, ii. 262.
Hazel ; see Corylus.
Hedera (Ivy), i. 137, 225.
HedysarejE. i. 287 ; ii. 143.
HeUanlhemum, ii. 104, 153.
Heliantheje, ii. 62.
Helianthus, i. 186, 250.
Helicteres, ii. 110, 210, 212.
Hcliophila, ii. 212.
Helleborus, i. 252; ii. 129, 137.

Hemerocallis, i. 244, 247, 253, 261.

Hemp ; see Cannabis.

Hepatic^, i. 11, 15, 35, 36, 55, 72,

307, 313, 314—316; ii. 244.
Heracleum, i. 33.
Hermannia, ii. 156.
Hesperis, ii. 122.
Hibiscus, ii. 31, 150.
Hieracium, ii. 168.
Hippocastane^e, ii. 208.
Hippuris, i. 36; ii. 130.
Holly ; see Ilex,
Homalineje, ii. 92.
Hop ; see Humulus.
Hordeum (Barley), i. 199; ii. 200.
Horse-chestnut ; see JEsculus.
Hortensia, ii. 41, 87, 98, 120.
Hovenia, ii. 172.
Hoya, ii. 9.
Humulus (Hop), i. 139; ii. 11, 18,

175.
Hura, ii. 18.
Hyacinthus, i. 128, 131, 196, 210, 211,

220, 246, 247, 274, 297 ; ii. 13, 40,

116.
Hydrangea, i. 138.
Hydrodyction, i. 323, 325 ; ii. 258.
Ili/menaa, i. 269.
Hymenopappus, ii. 81.
Hyosciamus (Henbane), ii. 170.
Hypericineje, i. 110; ii. 152.
Hi/pericum (St. John's Wort), ii. 64,

97, 204.
Hyphtene (Doom Palm), ii. 199.
Hypnum, i. 309, 314.
Hypoxylon, i. 11.
Tberis, ii. 13, 15.
Ilex (Holly), ii. 264.
Illicium, ii. 68.
Impatiens, ii. 59, 97.
Indigo/era (Indigo Plant), i. 244.
Inga, i. 240; ii. 111.
IridevE, ii. 157.
Iris, i. 139, 197, 238, 247, 273,

296, 297 ; ii. 60, 71, 83, 114, 157,

190.
Isidium, i. 317.

Isoetes, i. 203, 273, 274 ; ii. 235.
Isopyrum, ii. 137.
Ivy ; see Hedera.
Jacksonia, i. 98.
Jasione, ii. 274.

Jasminum (Jessamine), i. 112, 148.
Jonesia, ii. 165.
Juglande^e, ii. 88.

y2

c3^4

ALPHABETICAL TABLE OF GENERA, ETC.

Juglans (Walnut), i. 112, 148, 268 ;

ii. 137.
Juncus (Rush), ii. 13, 86
Jungermannia, i. 314, 315 ; ii. 244.
Juniperus (Juniper), i. 36, 163 ; ii. 176.
Justicia, i. 288 ; ii. 62.
Kcempferia, i. 33.
Kalanchce, ii. 19.
Koelreuteria, ii. 13.
Labiat.e. i. 173,280; ii. 22, 102, 103,

117, 110, 170, 194.
Lagasca, ii. 44, 171.
Lagerttrocmia, i. 278 ; ii. 97.
Larix (Larch), i. 36, 282, 296.
Lathreea, i. 242, 306.
Lathyrus, i. 222, 240, 241, 244, 247,

284, 304; ii. 41,69, 121.
Laurine.e, ii. 59.
Laums (Laurel), i. 288 ; ii. 59, 96.
Lavandula (Lavander), ii. 23.
Lebeckia, i. 244.
Lecythide.e, ii. 158.
Lecythis, i. 153.
Leguminos.e, i. 266, 269, 280, 282,

283, 284 ; ii. 24, 28, 38, 69, 71, 76,

77, 78, 82, 104, 111, 119, 141,

142 ,145, 148, 163, 164, 165, 1S9,

193, 203, 208, 209.
Lemna, i. 36 ; ii. 217.
Leon/ice, ii. 108, 146.
Lepidium, i. 241 ; ii. 183, 187.
Lepra, i 317.
Leptospermum, i. 176.
Lichenes, i. 11, 35, 307, 316—319;

ii. 247.
Lignum-vitse ; see Guiacum.
Ligustrum (Privet), i. 295.
Lilac ; see Syringa.
LiliacevE, i. 193—197, 199, 206, 208,

220, 247, 296; ii. 86, 107, 119.

157, 193, 216.
Lilium (Lily), i. 196, 247; ii. 94, 131.
Lime ; see Tilia.
Limodorum, i. 306.
Linaria, ii. 103, 117, 159.
LINE.E, ii. 109.
Linum (Flax), i. 56, 58, 170; ii. 107.

183.
Liriodendron (Tulip-tree), i. 297 ; ii.

68, 175.
Littcea, i. 114, 115, 205, 246.
Lobaria, i. 316, 319.
Lobelia, i. 273 ; ii. 64.
Lobeliace.e, ii. 65.
Lodoirca, ii. 160, 106.

Lonicera (Honeysuckle), i. 288; ii.

34, 111, 172."
Lorantltu.-t. i. 75.
Lotus, ii. 277.
Lucerne; see Medicago.
Lunaria, ii. 178.
Lupinus (Lupine), i. 264.
Luxula, ii. 151.
Lychnis, ii. 82, 133, 154.
Lycoperdon, i. 307.
Lycopersicum (Tomatbe), ii. 112.
LYCoroDiACE.E, ii. 233 — 236.
Lycopodium, i. 203 ; ii. 233.
Lysimachia, i. 137,276.
Lithrarie.e, ii. 23, 77, 165.
Lyihrum, ii. 23.
Magnolia, i. 285; ii. 68, 124, 128, 175,

178, 190.
Magnoliace.e. ii. 60, 160, 175.
Mahemia, ii. 333.
Mais, (Indian Corn), ii. 18.
Malpighia, i. 94.
MalpighiacecB, ii. 117.
Ma/us (Apple), ii 111, 164, 167.
Malva (Mallow;, i. 250; ii. 180, 212.
Malyace.e, i. 170, 251, 282 ; ii. 43,

64, 66, 67, 107, 109, 156, 160, 180,

181, 196.
Mandragora, ii. 29.
Maranta, i. 33.
Marchantia, i. 316.
Marrubium, i. 72.
Marsilea, i. 265 ; ii. 226.
Martiusia, ii. 76.
Martynia, ii. 265.
Marvel of Peru; see Mirabilis.
Mayanthemum, i. 212; ii. 114.
Medicago, i. 228, 267.
Medlar ; see Mespilus.
Melaleuca, ii. 64, 92, 97.
Melampyrum, ii. 41.
Melastoma, i. 250.
Mei.astomaceje, ii. 110.
Meliace.e, ii. 64.
Melianthus, i. 286.
Melon ; see Cucumis.
Melothria, ii. 83.

Menispfiime.e, i. 106, 252; ii. 83.
Menispermum, i. 252.
Mentha, i. 138.
Menyanthes, i. 102.
Mesembryanthemum (Ficoid), i. 214,

234, 237, 248, 289 ; ii. 39, 50, 87,
95, 164, 165, 210.
Mespilus (Medlar), ii. 165.

ALPHABETICAL TABLE OF GENERA, ETC,

325

Methonica, i. 247.

Michauxia,\. 73.

Millet, ii. 12.

Mimosa, i. 217, 283, 292; ii. 104, 142,

178, 204.
Mirabilis (Marvel of Peru), i. 146 ;

ii. 43, 88, 169, 196.
Misseltoe ; see Viscum.
Monarda, ii. 119.
Monotropa, i. 242, 306.
Moroca, ii. 33.
Morina, ii. (>6.
Morinda, ii. 173.
Morus (Mulberry), ii. 39, 174.
Mosses ; see Musci.
Muracuja, ii. 78.
Musa (Plantain; Banana), i. 33, 3.5,

114, 191, 197,254, 261.
Musci (Mosses), i. 11, 15, 35, 55, 72,

78, 82,308—314; ii. 236—243.
Musscenda, ii 102, 110.
Mu/isia, ii. 267.
Myogram, ii. 152.
Myosurus, ii. 68, 129.
Myriophyllum, i. 36, 74; ii. 12, 130.
Myriotheca, ii. 231.
Myristica (Nutmeg), ii. 180.
Myrsine^e, ii. 92.
Myrtace;e, i. 1 10, 283 ; ii. 8, 38, 92,

114, 165.
Myrtus (Myrtle).
Najas, i. 36.
Narcissus, ii. 86.
Nasturtium ; see Tropeeolum.
Nauclea, ii. 264.
Neckera, i. 313.
Negundo, ii. 292.
Nemopanlhes, ii. 80, 87.
Nepenthes, i. 240, 272.
Nerium (Oleander), ii. 58, 271.
Nettle ; see Urtica.
Neurocarpum, ii. 76.
Nicandra, ii. 170.
Nicotiana (Tobacco), i. 210, 234; ii.

55, 155.
Nigella, ii. 73, 149, 151.
Nissolia, ii. 182.
Nostoc, i. 326.

Nuphar, i. 117 ; ii. 76, 162, 200,209.
Nyctagineje, ii. 194, 195.
Nyctago, i. 114, 116 ; ii. 43, 169.
Nymphaa (Water-lily), i. 222, 224,

225, 236, 295 ; ii. 30, 76, 9o, loo.
Nymphceace*:, ii. 107, 163, 201.
Oak ; sec Quercus.

Oat ; see Avena.

Ochnace^l, ii. 160.

Ochroma, ii. 181.

Oco/ea, i. 233.

CEnanthe, ii. 16, 80, 167.

(Enothera, i. 274 ; ii. 165.

(Ethionema, ii. 64.

Oligotrichum, ii. 210.

Onagrari/e, ii. 61, 62.

Onoclea, ii. 230.

Ononis, i. 241.

Opercularia, ii. 45, 112, 173.

Ophioglossum, ii. 228.

Opuntia, i. 305 ; ii. 32, 265.

Orange ; see Citrus.

Orchide/e, i. 134, 137, 220, 246; ii.

62, 102, 158,207.
Orchis, i. 306; ii. 103.
Ornilhogalum, i. 273; ii. 13, 14, 121.
Orniihopus, i. 220.
Orohanche (Broom-rape), i. 140, 242,

305, 306; ii. 207.
Orobus, i. 244, 268.
Orthotrichum, ii. 240.
Osmunda, ii. 228.
Othonna, ii. 43, 106.
Oxalis, i. 243, 251 ; ii. 150, 189.
Oxytropis, ii. 138.

Pceonia (Preony), i. 197 ; ii. 75,76,162.
Palm;e (Palms), i. 128, 188, 196,

208, 229, 244, 246, 247, 259 ; ii.

12,46, 194, 195.
Pancratium, ii. 33.'5.

PANDANE.E, i. 193 — 197.

Pandanus, i. 194, 278.

Panicum, i. 199, 222.

Papaver (Poppy), i. 106; ii. 49, 76,

95, 110, 124, 152, 155, 159, 163.
Papaverace^e, ii. 158, 163.
Papilliosace^;, i. 220 ; ii. 54, 64,

102, 109, 203, 211.
Papyrus, ii. 17.
Paris, i. 247; ii. 115.
Parnassia, ii 116, 117.
Paronychia, ii. 179.
Passerina, i. 236.
Pass/flora (Passion-flower), i. 252,

261 ; ii. 78, 153
Passifloreje, ii. 78, 163, 180.
Paspalum, ii. 227.
Patellaria, i. 317-
Pavia, ii. 289.
Pea ; see Pisum.
Peach ; see Persica.
Pear ; see Pyrus.

326

ALPHABETICAL TABLE OF GENERA, ETC.

Pedis, ii. 168.

Peganum, i. 73.

Pekea, ii. 200.

Pelargonium, i. 271.

Peraltea, ii. 76.

Periploca, i. 139, 165.

Persica (Peach), ii. 135, 136, 141.

Petrocallis, ii. 178.

Peziza, i. 320.

Phaca, ii. 129, 139.

Phascum, i. 309, 312, 315 ; ii. 242.

Phaseolus (French Bean), i. 49, 139,

211, 212 ; ii. 70, 85, 138, 187, 189,

203, 205, 208, 209, 213.
Philadelphus, ii. 96.
Pldeum, i. 199.
Phlomis, i. 148.
Phlox, ii. 53.
Phormitun, i. 56, 57, 58.
Phyllanthus, ii. 31, 32.
Phyllirea, i. 155.
Physalis, ii. 30, 170.
Physcia, i. 318 ; ii. 248.
Phyteuma, ii. 16, 54.
Phytolacca, i. 146.
Pictetia, i. 283 ; ii. 263.
Pilularia, ii. 226.
Pine-apple ; see Bromelia.
Pink; see Dianthus.
Pinkneya, ii. 102, 119.
Pinus (Pine), i. 36, 109, 278, 279,

288; ii. 175, 205.
Piper (Pepper), i. 114, 116.
Pisum (Pea), i. 137, 268 ; ii. 70, 136,

138, 140, 203, 209.
Pitcarnia, ii. 25.
Plane ; see Platanus.
Plantago (Plantain), ii. 11, 12, 16,

38, 40.
Platanus (Plane-tree), i. 172, 181 ;

ii. 16.
Pleurogastcr, ii. 160.
Plum ; see Primus.
Podospermum, ii. 81.
Pollichia, ii. 171.
Polycardia, ii. 32, 227.

POLYGALE.-E, ii. 180.

Polygonalum, i. 119.

Polygone^, i. 242, 286 ; ii. 195, 196.

Polygonum, i. 219; ii. 199, 209, 210.

Polypodium, i. 201 ; ii. 228, 230.

Polystichum, ii. 230.

Polytrichia)!, i. 309; ii. 240.

Pomaces, ii. 164, 166.

Poppy ; see Papavcr.

Populus (Poplar), i. 141, 155, 158,

239; ii. 83.
Portulaca, ii. 196.
Portulace^:, ii. 78, 154, 158.
Potamogeton, i. 222, 245, 246. 247,

297.
Potatoe ; see Solanum.
Potentilla, ii. 140, 164.
Poterium, ii. 333.
Pothos, i. 202.
Prenanthes, i. 200.
Primula (Primrose), ii. 14, 97, 98,

119, 128, 154.
Primulace^e, ii. 92.
Privet ; see Ligustrum.
Prosopis, ii. 117.
Proteaceje, ii. 62, 168, 175.
Protococcus, i. 324.
Prunus (Plum), i. 180 ; ii. 104, 141.
Psilotum, ii. 235.
Psora, i. 317.
Pteris, i. 202 ; ii. 228.
Pterospermum, i. 291.
Puccinia, i. 320.

Punica, (Pomegranate), ii. 167, 212.
Pyrola, i. 242.
Pyrus (Pear), i. 140, 147, 236, 247 ;

ii. 5, 164, 166, 167.
Quercus (Oak), i. 49, 134, 147, 148,

150, 156, 160, 164, 166, 172, 185,

284; ii. 11, 171,186.
Quinales, i. 67.
Quince ; see Cydonia.
Quinquina, ii. 167.
Radish ; see Raphanus.
Ramnct-lace.e, i. 241 ; ii. 60, 68,

84, 95, 98, 145.
Ranunculus, i. 210, 214, 220, 243, 258 ;

ii. 49, 68, 122, 128, 140, 145, 147,

187, 194, 198, 199, 205.
Raphanus (Radish), i. 210, 219; ii.

201, 210.
Rauwolfia, ii. 149.

Reseda (Mignionette), ii. 146, 153.
Rhamne.e, ii. 78, 79, SS.
Rhamnus, ii. 288.

Rhhophora, i. 136, 210, 224; ii. 150.
Rhodora, ii. 53.
Rhodorace.e, i. 120.
Rhus (Sumach), i. 147, 153, 224. 228.
Riccia, i. 316; ii. 246.
Ricinus, i. 252, 285 ; ii. 150, 212.
Robinia, i. 221, 22S, 247, 264.
Rocella, i. 318.
Rochea, ii. 92.

ALPHABETICAL TABLE OF GENERA, FTC.

32^

Rosa (Rose), i. 30, 34, 86, 285 ; ii.

5, 49, 68, 98, 109, 129, 130, 164,

169, 173.
Rosacea, i. 269, 282, 287 ; ii. 77,87,

95, 104, 164.
Rosmarinus (Rosemary), i. 295.
Rubia, i. 286.
Rubiace.e, i. 280, 282, 285, 286 ; ii.

79, 107, 114, 160, 165, 166, 194,

196.
Rubus (Raspberry), ii. 140, 141.
Rudbeckia, ii. 16.
Ruellia, L 293.
Ruscus, i. 206, 207, 248 ; ii. 33, 35,

195.
Ruta (Rue), ii. 96, 113, 114.
Rutaces, ii. 68, 81, 87.
Rut idea, ii. 196.
Sabinea, ii. 111.
Sage ; see Salvia.
Sagina, ii. 84.
Sagittaria, i. 245, 247.
Salicomia, i. 289.
Salix (Willow), i. 129, 141, 155, 181,

212, 215, 223, 287; ii. 11, 59, 61.

66, 83, 124.
Sahola, ii. 282.
Salvia (Sage), i. 134; ii. 23, 41, 58,

87,119.
Salvinia, ii. 226.
Sambucus (Elder), i. 145, 147, 148,

150; ii. 13, 150.
Samyde.e, i. 109.
Sapindace.e, ii. 272.
Saponaria, i. 295.
Sarcophyllum, i. 240, 266.
Sarracenia, i. 271.
Sassafras, i. 49.
Saxifraga, ii. 216.
Saxifrages, ii. 164.
Scabiosa, i. 30; ii. 123, 130, 167.
Schinus, i. 109.
Scilla, ii. 121.

SciTAMINE.E, i. 297.

Scolopendrium, i. 262.
Scolymus, ii. 45, 170, 184.
Scorsonera, i. 219; ii. 81, 168.
Sckophulari.nes, ii. 102, 103, 117,

159.
Scutellaria, ii. 50.
Secale (Rye), ii. 200.
Sedum (House-leek), i. 137, 215, 223;

ii. 20, 82, 92.
Semecarpus, ii. 171.
Sempervivum,\. 209; ii. 92, 124.

Sesemum,\\. 103.

Seseli, i. 289 ; ii, 43.

Silene, ii. 20.

Sitiapis (Mustard), ii. 187.

Slateria, ii. 146.

Smilace.e, i. 196.

Smilax, i. 253 ; ii. 269.

Solandra, i. 137.

Solane.e, ii. 112.

Solarium, i. 220, 222, 276, 290 ; ii. 30,

59, 155, 187, 215.
Solomon's Seal ; see Poly-gonatum.
Sonchus (Thistle), ii. 81, 168.
Sophora, i. 265; ii. 139, 142.
Sorocea, ii. 211.
Sparganium, ii. 16.
Sparmannia, i. 333.
Spartium (Broom), i. 135; ii. 50, 138.
Sphoeranthus, ii. 16.
Sphoeria, ii. 249.
Sphagnum, ii. 239, 240.
Spinacia (Spinach), ii. 200.
Spira-a, ii. 32, 110, 164.
Surfaces, ii. 10 1.
Splachnum, i. 35, 309.
Spondias, ii. 199.
Squammaria, i. 317.
Stachys, ii. 264.
Stcchelina, ii. 81, 168.
Stapelia, i. 130, 133, 134, 305 ; ii. 65,

73, 207.
Statice, ii. 11, 144.

a, ii. 67.
Stellaria, ii. 15 1.

Sterculia, i. 265 ; ii. 69, 129, 137.
Sterigma, ii. 6 I .
Strawberry ; see Fragaria.
Strelitzia, i. 246, 254.
Strophanthus, ii. 271.
Sumach ; see Rhus.
Sweet- William; see Dianthus.
Sylphium, i. 288.
Sympiioxie.e, ii. 65.
Symphoricarpos, ii. 173.
Syncarpha, ii. 45.
Syringa (Lilac), i. 134, 177; ii. 24,

114.
Tagetes (African Marigold), ii. 53, 99.
Tapura, ii. 31.

Taraxacum (Dandelion), i. 274.
Targionia, ii. 246.
Taxodium, i. 200.
Taxus (Yew), ii. 59, 66.
Telephium, ii. 333.
Terebinthace.e, ii. 77, 88.

321

ALPHABETICAL TABLE OF GENERA, ETC.

Tetraphis, ii. 241.

Thalassiophytje, ii. 254.

Thesium, ii. 31.

Thistle ; see Sonchus ; Carduus.

Thorn ; see Crataegus.

Thuja, i. 36 ; ii. 175.

Thymel^eje, ii. 82, 88.

Tilia (Lime-tree), i. 271, 298; ii.

106, 210.
Tiliace/E, ii. 157.
Tithonia, ii. 210.
Tobacco ; see Nicotiana.
Tolpis, ii. 26.

Tomatoe ; see Lycopersicuni.
Tradescaniia, ii. 89, 106, 109, 120.
Tragopogon (Salsify), ii. 168.
Trapa, i. 113, 284 ; ii. 208, 210, 213.
Trianthema, ii. 152.
Trifolium, i. 267, 285 ; ii. 264.
Trigonella, ii. 140.
Trillium, i. 247; ii. 115.
Triosteum, i. 289.
Triticum (Wheat), i. 199, 222; ii. 12,

141, 200, 209.
Tritoma, i. 63.
Trollius, ii. 142.
Tropteolum (Nasturtium, or Indian

Cress), i. 41, 252, 210.
Tuber (Truffle), i. 307 ; ii. 249.
Tulipa, i. 128, 196 ; ii. 86, 189.
Tulip-tree ; see Liriodendron.
Turnip ; see Brassica.
Vgena, i. 129, 201.
Ulmus (Elm), i. 150, 159, 221, 225,

228 ; ii. 83, 182.
Ulva, i. 262, 323, 324 ; ii. 254.
Umbellifer*:, i. 109, 241, 296; ii.

14, 17,28,36, 79, 80,81,107, 119,

165, 168, 182, 194, 195.
Uredo, i. 320.

Urtica (Nettle), i. 170 ; ii. 35, 227.

Urtice.e, ii. 83.

Urvillea, ii. 272.

Utricularia, i. 113; ii. 30.

Usnea, i. 318.

Valeriana, ii. 110.

Valeriane*:, ii. 81, 82, 168.

Vallisneria, ii. 39.

Vaucheria, ii. 257.

Verbascum, i. 274.

Verbenace^e, ii. 119.

Veronica, ii. 7, 212.

Verrucaria, i. 318.

Viburnum, ii. 13,99, 288.

Vicia (Vetch), i. 222, 244, 268, 272,

285; ii. 208, 209.
Vinca (Periwinkle,) ii. 7, 29.
Viola (Violet), ii. 103, 117, 153.
ViolariEjE, ii. 194.
Viscum (Misseltoe), i. 164, 166, 171 ;

ii. 202.
Vitace^e, i. 132.
litis (Vine), i. 147, 215, 250, 295,

300; ii. 24, 54, 106, 150, 215.
Wall-flower; see Cheiranthus.
Walnut; see Juglans.
Water-lily ; see Nymphcea.
Webera, i. 309.
JVeissia, i. 309.
Wheat ; see Triticum.
Willow ; see Salix.
Wisteria, i. 139, 165.
Xanthorrhcea, i. 195.
Xylophylla, i. 305 ; ii. 32, 35.
Yew ; see Taxus.
Yucca, i. 128, 193, 195, 196, 205,

206.
Zizyphus, i. 250; ii. 31.
Zostera, i. 36 ; ii. 33.
Zygnema, i. 323.

DESCRIPTION OF THE PLATES.

PLATE J.

Cellular Tissue.
Fig.

1. Round Cellular Tissue.

2. Hexahedral ditto ditto.

3 Ditto ditto ditto, showing the Intercellular Passages.

4. A Section of a portion highly magnified, from Kieser.

5, 6. Cellules separated from the others.

7. Elongated cellules of the wood and bark.

8. Elongated cellules which surround the vessels.

9. 10. Muriform cellular tissue from the Medullary Rays.

PLATE II.

The I 'esse Is.
Fig.

1. A longitudinal section of a fibre of the leaf of Tritoma Uvaria, showing in a. a.

the Trachea?, and b. b. the striped vessels.

2. u.a. the striped, and b.b. the dotted, vessels, from the stem of Lycopodium denti-

culatum.

3. A Trachea of Musa Paradisiaca, showing the band to be composed of several

spiral fibres.

4. A Trachea, from Dutrochet.

5. a. a. striped, and b. b. dotted, vessels, from the stem of Smilax aspera.

6. A strangulated vessel.

7. a. a reticulated ditto ; b. an example, from Mirbel, showing a trachea changing

into a striped or annular vessel.

PLATE III.

Cuticle and Stomuta.
Fig.

1. Cutic'e and Stomata of Patonia.

2. Ditto ditto of Iris Germanica.

3. Ditto ditto from the lower surface of the leaf of Lycopodium denti-

culalum.

4. Portion of a longitudinal and vertical section of a leaf of Tritoma Uvaria,

showing the cellules of the cuticle and those of the parenchyma.

5. A longitudinal section of the same leaf, made parallel to the surfaces, showing the

longitudinal fibres of the nerves, the parenchyma, and bundles of Raphides.

6. The Raphides more highly magnified.

7. Portion of a transverse section of a leaf of Yucca aloifolia, showing the dif-

ference between the compact cellules of the cuticle and those of the parenchyma.

8. A vertical section of Fig. 2.

o30 DESCRIPTION OF THE PLATES.

Hairs.

A. A cupulate hair.

B. A capitate ditto from Petunia.

C. An awl- shaped ditto, or sting from the Nettle.

D. A hair of Malpighia.

E. A simple hair.

F. Divided ditto.

G. Moniliform ditto.
II. A hair from Malm.

I. A shield-shaped ditto from the leaf of El&agnus.

PLATE IV.

Stems.

Fig.

1. A fasciculated branch of Spartium junccum.

2. A transverse section of the stem of a Juniper, half the natural size, showing a

gelivure, or frozen layer. (See vol. i. p. 1C3).

3. Section of the annual stem of Ferula communis.

PLATE V.

Stems.

Fig.

1. Bifurcation of a hranch of the Horse-chestnut.

2. A transverse section of a branch of the Ebony-tree (Diospyros Ebenum), showing

the sudden difference of colour between the Duramen and Alburnum.

3. Transverse section of Quercus Tanza, showing the large medullary rays, and the

very thick cellular layer of the bark.

PLATE VI.

Ficus Elastica.

Portion of a shoot of Ficus elastica, showing how the stipules inclose the young
leaves at their infancy; also a fragment of the bark with lenticils, from one of
which an adventitious root is seen proceeding.

PLATE VII.

Leaves.

Fig.

1. Pinnatiftd leaf of Comptonia asplemfolia.

2. A leaf of Acacia heterophylla. with the common petiole very slightly dilated,

and bearing two pair of partial petioles furnished with leaflets.

3. Another leaf of the same plant, with the common petiole more dilated, but still

bearing several pairs of partial ones.

4. Another leaf, with the petiole still more dilated, and bearing very few partial ones.

5. Example of the same plant, with the petiole entirely dilated into a phyllodium

and no longer bearing leaflets. The flowers in compact heads, pedicellate and
axillary.

PLATE VIII.

Root.

Extremity of one of the branches of the root of Pandanus odoratissimus, of the
natural size, showing the large scaly spongioles which terminate the principal
branches.

DESCRIPTION OF THE PLATES. 331

PLATE IX.

Leaves.
Fig.

1. Leaf or portion of the stem of an Indian Arum, showing the sheathing petiole

and pedalinerved limb.

2. Leaf of Liquidambar Styraciflua ; an example of a paimatifid, palminerved limb,

with serrated lobes.

PLATE X.

Leaves.
Fig.

1. Leaf of Ocotia Guianensis, showing the false nerves or marks caused by the

pressure of neighbouring nerves during its infancy; an example of an ohlong
leaf, pointed at both ends, slightly penninerved, with reticulated lateral nerves,
and entire margins.

2. Leaf of Begonia Arlicula/ri, the joints being formed by the dilation of the

winged petioles, the leaflets which ought to spring from the contractions being
entirely wanting.

3. Leaf of Desmodium triquetrum, with two stipules and a winged petiole, termi-

nated by a single leaflet.

4. Leaf of Sarcophyllum carnnsum, composed of a petiole and terminal leaflet.

5. Different states of the leaves of Lebeckia nuda, showing the petiole furnished

with rudiments of leaflets.

PLATE XL

Leaves.
Fig.

1. Leaf of Bauhinia purpurea, probably formed by the constant union of two

leaflets.

2. Leaf of Passifttra perfoliata, emarginated at the base.

3. Leaf and stipules of Lardizabala irilernata, which had better be called biternala.

An example of foliaceous, caulinary, caducous stipules.

PLATE XII.

Buds of the Horse-chestnut.

Fig.

1. A young branch with the buds as yet undeveloped, and with lenticils and cica-

trices of the old leaves.

2. 3, 4 and 5. The buds more developed, showing the progression from the scales

to the leaves, proving the scales to be petiolaceous.

PLATE XIII.

Leaves.

Fig.
1. Leaf of Desmodium gyrans, showing the three leaflets and the stipels.
2 and 2*. Leaves of Mimosa sensiliva.

3. Leaf of Melianthus cosmous, showing the stipules.

4. Leaf of Melianthus major, showing the great interpetiolary stipules, with a longi-

tudinal line through their centre, proving that they are formed by the union of
two.

332 DESCRIPTION OF THE PLATES.

PLATE XIV.

Union of Leaves.
Fig.

1. Two leaves of Laurus nobilis united toeether.

2. A similar example from Juslicia oxypkylla.

PLATE XV.

Btids.

Fig.
I, 2, and 3. Radical buds of Pceonia nfficinalis, in different states of development.

4, 5, and 6. Buds of Amelanchier in different degrees of development. We see the

stipules and bracteoles resembling each other.
7 and 8. Buds of Pyrus hi/brida at different ages.

5. Progress of the scale into leaves, showing that these buds are of a fulcraceous

nature.

PLATE XVI.

Varieties.

Fig.

1. Top of the winged branch of Xylophylla, enlarged; showing the origin of the

flowers from each little notch.

2. Portion of a branch of Capparis quadriflora, with spiny stipules on each side of

the leaf, and the pedicellate flowers springing in a vertical series above the
axil.

3. Ruscus aculeatus. The alternate sheaths may either be considered as true coty-

ledons, or as primordial leaves if the cotyledons be concealed in the seed.
The leaves resemble scales at the bases of the leaf, like compressed branches.

4. The axis of the cone of Larix Eurnp&a prolonged into a branch.

PLATE XVII.

Proliferous Rose.

Fig.

1. A rose with a flower bud in nearly the ordinary state, and with a proliferous

flower: in which the calyx is changed into leaves, and does not adhere to the
ovary ; the petals and stamens are nearly in the state of a semi-double rose ;
the axis is prolonged in the centre of the flower bearing a second.

2. The above flower seen from below.

3. The same, cut through the centre, the sepals and petals being removed.

4. An ordinary petal.

5. An ordinary stamen.

6 and 7. Stamens changing into petals.

8. A petal of the supernumerary flower.

9. A stamen of ditto ditto.
10. A carpel of ditto ditto.

PLATE XVIII.

Varieties.

Fig.

] and 2. Two flowers of Podospermum laciniatum ; one in the ordinary state with
the limb changed into the pappus : and the other with a five-lobed calyx.

3 and 4. Two flowers of Capsella Bursa-pastoris with ten stamens : the four petals
being changed into stamens in addition to the six ordinary ones.

DESCRIPTION OF THE PLATES. 333

Fig.

5. A flower of Datura fasttiosa, showing a triple corolla.

6 and 7. A monstrosity of Gentiana purpurea, in which the ovarium is formed of
two rows of carpels, all bearing ovules ; the outer row has four, the inner two
carpels.

8, 9, 10, and 11. Different states of the flowers of Phlox amaina from the same plant,
showing the different combinations of the petals, from their being nearly per-
fectly free, to their usual gamopetalous state.

PLATE XIX.
JEstivations.

In these figures, j designates the sepal?, p the petals, / the tepals, and b the bracts.

Fig.

1. Papaver Rhevas ; petals rumpled, sepals imbricate.

2. Muliernia pinnata; petals contorted, sepals valvate.

3. Tradescantia Virginica; petals imbricate, sepals valvato.

4. Philadelplius coronurius ; petals contorted, sepals valvate.

5. Cistus albidus; petals contorted, sepals quincunxially imbricate.

6. Clematis florida ; sepals induplicate; s. one seen from the inner side.

7. Sparmannia Africana ; petals half induplicate, half imbricate, sepals valvate.

8. Spartium junceum; petals in vexillary aestivation; v. (vexilla), the standard ;

a. a. (alae). the wings ; c. (carina), the keel.

9. Daphne alpina ; peritjone imbricate.

10. M. Irosideras lanceolata ; sepals and petals quincunxially imbricate.

11. Plantago media ; tepals imbricate.

12. ' iota arvensis ; sepals and petals quincunxially imbricate.

13. Poterium Sanguisorba ; sepals in regular aestivation.

14. Pancratium maritimum ; the outer row of the perigone formed of curved pieces,

the inner, of flat and valvate enes.

15. Hmja carnusa ; petals valvate.

PLATE XX.

Structure of Oraries with a central placenta.

Fig.

1. Stellaria graminea.

2. Cerastium arvense.

3. Arenaria tenuifolia.

4. Arenaria marina.

5. Silene, (a species very near .■?. conica.)

6. Silene nutans.

7. Claytonia perfoliata.

8. Telephium Impcrati.

PLATE XXI.

Monstrosities.

Monstrosity of an apple, produced by two (lowers united by their peduncles. Cen-
taurea with the peduncles united.

PLATE XXII.

Germinations.

Fig.

I. Euphorbia Hclioscopia ; a monstrosity with four cotyledons, formed by the. union
of two embryos.

SSI DESCRIPTION OF THE PLATES.

Fig.

2. Cubepa scandens, with its stem, cots'ledons, and primordial leaves.

3, 3*, and 3**. TroptEolum peregrinum, with its radicle; 3. the seed containing

the cotyledons ; 3**. the seed beginning to open at the base ; 3* the cotyle-
dons are seen, their covering being removed.

PLATE XXIII.

Germinations.

Fig.

1. A species of Bombax.

2. Quercus Suber.

3. Cactus Melncactus.

THE END.

LONDON :
PRINTED BY RICHARD CLAY, BREAD STREET HILL.

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UNIVERSITY OF TORONTO LIBRARY

QK

641

C213

1841
v.2

BioMed

Candolle, Augustin
Pyramus de

Vegetable organography

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