THE 
VEGETABLE INDIVIDUAL, 
IN ITS 
RELATION TO SPECIES. 
BY 
Dr. ALEXANDER BRAUN, 
Professor of Botany in the University of Berlin, &c. &c. 
EXTRACTED FROM THE AMERICAN JOURNAL OF SCIENCE AND ARTS, VOLUME XIX, SECOND 
SERIES, MAY, 1855. 
NEW IIAVEN: 
PRINTED BY B. L. HAMLEN, 
Printer to Yale College. 
1855. The Vegetable Individual, in its relation to Species, (Das Indi- 
viduum der Pflanze in seinem Verhaltniss zur Species,—Gen- 
erationsfolge, Generationswechsel und Generationstheilung der 
Pflanze); by Dr. Alexander Braun, Professor of Botany in 
the University of Berlin, &c. &c. Berlin, 1853.* Translated 
by Chas. Francis Stone. 
Part I.—Introduction and History.f 
In Organic Nature the two principal phenomena, in which the 
shifting scenes of Life are unfolded, are individual development 
and individual propagation. Through them the intricate course 
of Nature, and its living chain of organized beings, are refreshed 
and renewed. Every new generation seems to bring back the 
old form ; still, to the investigator who looks deeper into the 
graves of the past, a slow, but certain, progress reveals itself even 
in this apparently identical succession. If nature is to be for us 
something more than a labyrinth of varied and intricate phenom- 
ena ; and if, in the apparent disorder, the hidden threads of the 
connection are to become visible, we must first of all separate 
and compare the different spheres of life, placing them higher or 
lower, according to their rank. The starting-points, which nature 
offers for such a purpose are, the Individual and the Species; 
whose reciprocal relations, however simple they may at first 
appear, when followed out to particulars lead to difficulties which 
* From the Transactions of the Royal Prussian Academy of Sciences, for 1853. 
f I have omitted the author’s brief introductory remarks.—Transl. 2 
The Vegetable Individual, in its relation to Species. 
demand an accurate examination.* From the Botanist such an 
examination is particularly demanded ; as the vegetable ideal pre- 
sented to us by the science in its earlier stages has been obscured 
by conceptions obtained from the animal kingdom having been 
transferred to Botany, though based upon the mistaken assumption 
that plants possess the same independent individuality as animals, 
the same organs with equally well-defined functions, and the same 
mutually dependent relations of the vital activities. And the 
investigations of late years, forsaking the old views more and 
more, have arrived at no well-defined conclusions, and, particu- 
larly as regards vegetable individuality, seem to lead more to 
negative than to positive results. After all, this should not surprise 
us; for even a superficial investigation shows relations in plants 
which will hardly harmonize with the common conceptions of 
individuality, and which require a careful review. 
In the whole realm of organic nature, we know of not a sin- 
gle species of which any one individual is a perfect representative : 
on the contrary we see each species adding generation to genera- 
tion, by multiplying the individuals in time and space, until its 
day has ended, whether from internal or external causes. In this 
particular, the species resembles the individual itself; having its 
allotted age, though measured by days of a higher order, and its 
appointed cycle of life,—in which the individuals appear as mem- 
bers occupying a certain time and place,—resembling the succes- 
sive relative forms through which the individual passes. For 
the organic individual does not manifest itself in one single per- 
manent form, but in a succession of forms, now gradually con- 
nected, now broadly interrupted; and these last, especially in 
plants, may attain to an independence, which gives them the 
character of a subordinate species. To this analogy between in- 
dividuals and species it may be objected, that, in most cases, a 
very remarkable metamorphosis is connected with the successive 
forms of the individual, while within the sphere of the species 
the consecutive members continue to have essentially the same 
* Should any one be inclined to doubt that the nature of the vegetable individual 
needs a futher discussion, I would beg him to turn to the latest works on Botany 
and compare the passages which treat of the plant’s individuality. I take KiUzwg's 
Grundzuge der phil. Botanik (2nd Part), as we have a right to demand from a work 
that lays claim to philosophical development, a fundamental discussion of this sub- 
ject, since it is the ground-work of the whole science. The first two paragraphs un- 
der the heading “ Das Pjlanzenindividuum als Organismus,” read as follows: “ By 
individual we here mean a single vegetable body not organically connected with a 
similar vegetable body. Vegetable individuals have the power of developing the 
general phenomena of vegetable life by themselves, unassisted by any other individ- 
ual of the same species. It is the nature of an organism to consist of members. . .. 
The possession of members is the first, as well as the most essential condition of the 
existence of the vegetable individual.” Not one of these assertions is true of veg- 
etable individuals, either in the broader or the narrower signification of the term. To 
say nothing of the connection, in which the individuals appear which are successively 
developed by shoot-formation, the coalesence of stocks which were originally sepa- The Vegetable Individual, in its relation to Species 
3 
character.* But, however important this fact may be, still we 
may assert of the individual as well as of the species, that it com- 
pletes the cycle of its existence in a succession of subordinate 
generations, while, on the other hand, we may affirm of the spe- 
cies, that like the individual, it exhibits a determinate cycle of 
development.! In comparing the processes of propagation with 
the process of the formation of the individual, cell-formation, 
which lies at the foundation of both, reveals the intimate con- 
nection which exists between the small and the great spheres of 
development; while the numerous cases which admit of a double 
explanation (since they may be ascribed with almost equal jus- 
tice to the inferior cycle of development of the individual, or to 
the superior one of the species) establish the close relationship of 
both. The above-mentioned circumstance, that the cycle of de- 
velopment does not present as graduated a progress in the species 
as it does in the individual, seems to suggest that the most relia- 
ble view of the analogy between the species and the individual 
is that in which the species is not compared with the whole cycle 
of the individual’s successive development, but with the single 
steps of the metamorphosis (which of course has its own sub- 
ordinate members), and in which the species itself is regarded 
as an inferior “ momentum” of a still more comprehensive cycle 
of development; but to determine this would lead us mo far from 
our subject.! In a word, the relation of the individual to the 
species is that of an inferior cycle of development to a superior: 
the individual is a member of the species. However, although 
they are under one and the same specific law, all the members of 
the species are not identical: a single member only represents the 
idea of the species more or less incompletely; and certain mem- 
bers, or series of members, are thus reciprocal complements. 
The regular relations here brought to view will form the princi- 
pal subject of the present investigation. But we must first care- 
fully determine the sphere of the individual. The individual 
shall not and may not be considered by itself: it must be viewed 
rate is no rarity. Are the pines of a pine-forest no individuals because, as Goppert 
has shown, they are connected with each other by their roots ? Do the filaments of 
Zygnemce cease to be individuals when they copulate. Are the cells of Hydrodic- 
tyon and Pediastruvi, originally separate, no longer individuals when they have joined 
themselves into a net, or a star ? To refute the second assertion, we may refer to 
dioecious plants; to refute the third, we refer to the one-celled Algae and Fungi, a part 
of which, at least, are of such a character that we can by no means ascribe to them 
an organisation in the usual acceptation of the term. However, we may regard it 
as an improvement, that Kulzing's Grundziige treats of the vegetable individual at 
all; for the earlier manuals do not even mention this important subject, but com- 
mence their account of plants with descriptions of the root, stem and other organs, 
or, as it has been preferred of late years, of the cells and vesicles. 
* Those of the forms and properties, which persist through the successive genera- 
tions, determine the species. Link: Grundlehren der Krciuterknnde, vi, p. 11. 
f The species is an individual of a higher rank (higher power). Link, 1. c., p. 11. 
+ Cf. the Author’s work on Verjungimg (1849), note to p. 344. 4 
The Vegetable Individual, in its relation to Species. 
in the successive generations to which it belongs. This succes- 
sion may be similar or dissimilar, simple or complicated by divi- 
sions, continuous or graduated by cyclical changes. It is by this 
that the phenomena of fissiparous and alternate generation may 
be explained. It is only by a consideration of these relations that 
the nature of the individual itself, as a subordinate sphere of the 
species’ development, can be rightly comprehended, and that the 
single individuals in their worth and importance, in their relations 
to each other and to the whole realized cycle of the species, can 
be understood. 
Preliminary remarks on Vegetable Individuality: different 
views in regard to it. 
We must determine what constitutes the vegetable individual, 
before we can investigate its relations to the whole cycle of gen- 
eration of the species. But it is this determination itself which 
presents so many difficulties; and these difficulties become the 
greater, the further we push our investigations. Individuality in 
plants seems as obscure and ambiguous, as in animals (at least 
in their higher orders,) it appears clear and simple; so that, as 
Steinheil remarks, it escapes us just when we are upon the point 
of seizing it ;* and investigators might even conclude that we can 
realize no other individuality than that which is manifested in the 
totality of the species. The first obstacle to our comprehending 
the vegetable individual as a single sphere of conformation, as a 
morphological whole, is the disconnected and separate character 
which obtains in the most heterogeneous modifications of vegeta- 
ble organisms. For no where in the vegetable kingdom do we 
perceive that indissoluble connection, and those pervading recipro- 
cal functions, which in the animal kingdom we are accustomed 
to associate with the idea of an individual organism. Neverthe- 
less, by starting from a comparison with animals we get an appo- 
site point of departure for a comprehension of the plant’s indi- 
viduality. Among the higher animals, the individual appears as 
a member of a race produced by sexual generation ; and this very 
test may be applied to plants, except in the very lowest forms, 
to which sexual generation does not apply at all, or not posi- 
tively. Without at present discussing the question whether the 
vegetable individual thus conceived is truly analogous to the an- 
imal individual, we may here state, that this conception car- 
ried out to its consequences, involves the assumption that all the 
plant-stocks produced, not by sexual generation, but by any mode 
of vegetable division, are not individuals, but only parts of the 
primary individual to which they owe their origin; as Gallesio has 
* “ Dans chacun de ces organes nous nous croyons au premier aspect sur le point 
de saisir l'individualite normale, et partout elle nous 6chappe.” Steinheil: De I'in- 
dividualite vegetate (1836), p. 9. The Vegetable Individual, in its relation to Species. 
5 
in fact contended.* Botanists have often asserted that it is the 
individual! alone, which is reproduced by slips (branches, buds, 
tubercles etc,), and their opinion coincides with this view. Still, 
how are we to distinguish plant-stocks of such an origin, from 
those derived from seeds ? The former take root, ramify, blos- 
som, ripen their fruit and seeds, just as the latter do, so that in a 
physiological sense they are complete For exam- 
ple, let us cast a glance at the weeping-willow (Salix Babylonica). 
It is well known that this tree, which was originally brought 
from the banks of the Euphrates, is always propagated by slips; 
for with us it never bears seeds—not because our climate is unfa- 
vorable, but because in our gardens there is no fructifying male 
tree.<§> According to Loudon (Arboret. the weeping-wil- 
low was sent to England in 1730, by a French merchant named 
Vernon. It was planted in Twickenham Park, whence it spread 
rapidly over England and the continent. The tree, from which 
the first slips that were brought to Europe were taken, was most 
probably a cultivated one itself, raised from a slip. However this 
may be, could the descent of all our weeping-willows be traced, 
it would undoubtedly lead us back to a willow, a female willow, 
grown in its native country from a seed. And so, on this ac- 
count, we are to regard all the beautiful weeping-willows of our 
gardens and our cemeteries—and surely they are perfect trees— 
not as individual stocks, but as the disjecta membra of a primary 
trunk, now hidden in mythical darkness ! In other cases this pri- 
* Gallesio: Teoria della riproduzione Vegetale (1816), a work, which T am sorry to 
say I have not been able to consult myself. Huxley (upon Animal Individuality, in 
the Ann. and Mag. of Nat. Hist., June 1852), holding corresponding views, regards 
all the animals which spring from an egg by non-sexual increase, as one individual, or, 
as he expresses it, as a representative of the individual by successive coexisting sep- 
arable forms ;—regards as such, for example, the sum total of all the Aphides, pro- 
duced in successive generations, by non-sexual increase, from the first “nurse” which 
sprung from the egg. If we assume with Bonnet that one nurse encloses one hun- 
dred yonng Aphides in the tenth generation (and according to Kyber they often 
reach even a higher number) the series would amount to much more than a billion 
(1,010,101,010,101,010,101). Those who regard sexual reproduction as the criterion 
of individuality must admit this as a perfectly legitimate consequence of their view. 
f “ Gemma; individuum continuant cum semina speciem propagent.” Link: Elem. 
phil. Botan., ed. II, v. i, p. 332. “ Continuant,” in antithesis to “ propagent,” cannot 
be mistaken. Again Endlicher und Unger: Grundzuge der Bot., p. 85, say, “ In these 
cases (i. e. when the buds drop off) the bud-formation is a true propagation, by which 
the individual is multiplied; though we must distinguish this mode of propagation 
from that of generation, by which the species is reproduced.” Here the meaning is 
obvious, though the expression is perfectly paradoxical; for how can we imagine 
that the individuals are multiplied without the species being reproduced ? I have 
elsewhere attempted to show what is here meant, by representing non-sexual propa- 
gation as a propagation subordinate to the cycle of sexual reproduction (cf. Ver- 
jungung, pp. 26, 27). 
;]; In many cases the experienced gardener can distinguish them, but certainly not 
in all; in some the difference is very remarkable: e. g. in Araucarice raised from 
branches. 
§ This has the advantage of avoiding the disagreeable seed-down. For the same 
reason, it is said, in China they cultivate the male tree only. 6 
Tlie Vegetable Individual, in its relation to Species. 
mary trunk is known with perfect certainty. It can be proved 
by history that many hybrids and varieties have been produced 
in one single exemplar; though they now ornament our gardens 
far and wide, having increased by means of slips, as they do not 
bear seeds. This was the case of the famous Cytisus Adami, 
which sprung, shortly before the year 1825, from the mingling 
of C. purpureus and C Laburnum. The single parent-stock, 
preserved in the garden of the celebrated Adam in Paris, has 
long since disappeared ; but its scions and scions-scions, have 
grown up into fine trees in half the gardens of Europe.* In 
the view just stated, they all form but one individual ! To 
support such a view, its partisans adduce the fact of certain 
individual particularities being preserved (in dioecious plants espe- 
cially the gender), when propagated by slips. In general this is 
true, and for practical gardening, e. g. for the cultivation of the 
finer kinds of fruit, of the greatest importance; but exceptions 
are not rare ; among which the well known re-division of Cytisus 
Adami into its two primary stocks is one of the most striking 
and remarkable. In our gardens the rule is that from slips the 
weeping-willow produces female trees; still some exceptions 
may be noted here. Napoleon’s grave in St. Helena is shaded 
by a weeping-willow, which has become the subject of scien- 
tific discussions. It was supposed to belong to a species (Sa- 
lix Napoleonis) indigenous to that Island ; but Loudon’s ex- 
haustive researches show that it is descended from our weeping- 
willows, one of which was carried from England to St. Helena 
in 1810. Branches of this iSalix Napoleonis were brought back 
to England, and to the astonishment of botanists they bore male 
flowers. Since up to that time no male weeping-willows had 
been seen in England, a change of gender must have been pro- 
duced through vegetative increase. A similar case has also oc- 
curred in Germany. In the-Grand-ducal Gardens at Schwetziu- 
gen there is a weeping-willow which, although a descendant 
from the common parent tree of all European weeping-willows, 
has changed its gender to such a degree that we not only find on 
it the most heterogeneous stages of transition from female flowers 
to male ones, but on many branches purely male catkins are pro- 
duced.! Besides these cases, a curled variety of weeping-willow 
Salix crispa or S. annularis of the gardens, is known ; which, 
as it is a mere garden plant, has probably been produced by slip- 
propagation. If it be true that we sometimes obtain varieties 
with hanging branches from several kinds of trees by grafting 
the slips inverted, we should have one of the most remarka- 
* Cf. Verjungung, pp. 337 and xi. In another place I shall communicate the his- 
tory of this hybrid, which has since been investigated. 
•f This tree was first observed by C. Schimper in 1827. Some remarks upon it 
may be found in Spenner’v Flora Friburgensis, vol. iii, p. 1061. The Vegetable Individual, in its relation to Species. 
7 
ble examples of the production of a singular peculiarity by 
non-sexual increase. But even if such exceptions did not exist, 
and if in every case a series of peculiarities which are extin- 
guished in seminal propagation were continued by grafting, yet we 
cannot perceive how we can seriously refuse an individual exist- 
ence to such stocks as these, produced, it is true, by non-sexual 
propagation, but still completely separated externally, developing 
in different places and under the most dissimilar relations, and ex- 
hibiting subordinate differences indefinitely, though with certain 
similar characteristics. But if we were to make any concessions 
on this point we should be carried irresistibly on to others. 
Most of the modes of non-sexual propagation thus far consid- 
ered agree in this particular ; that some shoot of the plant, whether 
it be undeveloped (eye, bud), or developed (branch, sucker, layer, 
&c.), is separated from the parent-stock by natural development 
itself, or by artificial means. As the nature of the separable part 
is not changed by the separation, it is no great step to attribute 
individuality to the shoot, (or as it is commonly called, the bud,) 
even when it is not separated from the stock. Each single plant- 
stock could then be no longer regarded as an individual in the 
usual meaning of the term, but as an united family of individual 
shoots;—a view which seems to be of high antiquity ; as passages 
are found in Aristotle* and Hippocratesf, which are interpreted 
in this sense. In later times, this view has been more or less, ad- 
vocated, especially by De la Hire,J Linnseus, Batsch, 
Goethe, Roper, Schleiden|| and others. 
But, even in this narrower view of vegetable individuality, the 
same difficulty meets us ; for the shoot itself is divisible, and new 
stocks may be produced by its parts: i. e. by the members of the 
* Cf. Wirnmer: Phytologice Aristotelicce Fragrnenta, §§23—28, 66 etll3. I cannot 
discover that explicit acknowledgment of the individuality of shoots or buds, which 
is said by Schultz (Anaphytose, p. 24) to be found in Aristotle, either in Schultz’s 
quotations, or even in Wimmer’s complete collection of the passages in Aristotle re- 
ferring to plants. It is true that Aristotle repeatedly speaks of the divisibility of 
plants; says that separated parts of plants may continue to exist; that on this account 
many trees may spring from a single source; that many plants are propagated by slips 
(cwi CTTrapayatittTtov cwo(puTtuojitvt*>v), and by lateral bud-formation (tw TrapafkaoTcivfiv) 
e. g. the bulbous plants; but he does not state his opinion of the parts which de- 
velop after such a separation, and explains the phenomena in general, by saying that 
the vegetable soul of plants (SpjTmxn is simple in actuality, (IvTttoxsO though 
multiple in capacity (SuvAao). 
f According to Moquin-Tandon: Teratologie, p. 5. 
| Hist, de 1’Acad. Roy. des Sciences, 1708, p. 233. De la Hire regards all the 
branches as new plants proceeding from hidden ovules. Myriads of these ovules, 
he thinks, exist between the bark and the wood; more or less of them come to ma- 
turity, according to circumstances. 
§ Darwin: Phytologia (1800), p. 1. “ If a bud be torn from the branch of a tree, or 
cut out and planted in the earth . . . ; or if it be inserted into the bark of 
another tree, it will grow and become a plant in every respect like its parent. This 
evinces, that every bud of a tree is an individual vegetable being, and the tree there- 
fore is a family or swarm of individual plants ...” 
H I shall consider the views of these authors more at large in the next section. 8 
7he Vegetable Individual, in its relation to Species. 
stem and its leaf or leaf-whorl.# Besides, the several members 
of the shoot are not contemporaneous creations, but, developing 
successively out of and over each other, they constitute a suc- 
cessive generation, composed of divisions each of which repeats 
essentially the same form, each of which may be compared to the 
embryonic plant originally developed in the seed, and consisting 
of its stemlet with one or two leaves (cotyledons). Thus the 
shoot itself came to be regarded as a succession of individual 
vegetable members, built up one above the other, like the stories 
of a house. The earliest traces of this view may be found in 
Darwin's Phytologia;f it was developed at a later period in vari- 
ous ways and with various modifications : e. g. by AgardhJ, En- 
gelmann, $ Steinheil|| and GaudichaudU—the last of whom calls 
the member of the shoot elevated to the rank of an individ- 
ual vegetable being, “thephyton,” and ascribes to it not only 
a stem and leaves, but even a root, by which he imagines it is 
connected with the preceding phytons, as the first phyton (the 
embryonic plant) is connected with the ground. Steenstrup** 
and Forbesff employ a similar view for their comparison of alter- 
nate generation in plants with that in the lower animals. 
But this restriction of vegetable individuality could not stop 
here ; for even the members of the shoot, the “ phyta” or “ sto- 
ries,” are themselves too complex organisms not to present subor- 
dinate divisions, which, like the whole member possess a certain 
independence, and under certain circumstances may even give 
birth to new stocks. Although botanists have attempted to 
view the petiole as the lower part of the leaf,or, vice versa, 
the leaf as the upper part of the petiole,(so as not to be com- 
pelled to divide the phytons of the structure themselves into rel- 
* I adduce this point in connection with the history of the views held by botanists 
in regard to vegetable individuality, in the terms in which it has been usually ex- 
pressed; further on I shall show that this view needs qualification. The individual 
members of the stem cannot expand into a new stock by direct development, like 
the separated shoot; they have this property only by being connected with a lateral 
sprout, by means of the eye which they bear, or have the power of producing. This 
view naturally brings us back to the shoot as the individual. 
f P. 9; where even the single well-defined stem-members of different herbaceous 
plants are described as so many buds, and hence as so many individuals. 
f Agardh : Essai de reduire la Physiologic vegetale d des principes fondamentaux, 
1829, (Ann. des Sci. Nat., tom. xvii, p. 86). 
§ Engelmann: de Antholysi, (1832) p. 12. 
| Steinheil: VIndividuality dans le regne vegetale. 1836. 
*|f Gaudicliaud: Recherches sur 1’ Organographie, la Physiologic et V Organogenic 
des Vegetaux. 1841. 
** li teen strap: On alternate Generation (1842), p. 128. As this important little 
work may be supposed to be in every one’s hands, I refrain from quoting this inter- 
esting passage. 
ft Forbes: On the Morphology of the reproductive system of Sertularian Zoophytes, 
Ac., Ann. and Mag. of Nat. Hist., v. xiv, (1844), p. 385. 
XX Ernst Mayer: die Metamorphose der Pflame und ihre Widersacher. Linrnea, 
1832, p. 401. 
§§ Hochstetter: Aufbau der Graspfanze. (Wurtemberger Jahreshefte, 184*7 and The Vegetable Individual, in its relation to Species. 
9 
atively independent members) this much at least is certain (and it is 
the important point here), that each of these two parts is capable 
of producing new growths by itself, yes, this capacity is enjoyed 
even by different determinate or casual parts of either member. 
It is well known that the leaf of Bryophyllum produces sprouts 
in every notch on its edges, while on the .other hand, caducous 
leaves of many bulbous plants (e. g. Eucomis regia, according 
to Hedwig, Ornithogalum thyrsoides according to Turpin)* pro- 
duce new plants in the form of bulblets on any portion of the 
whole of the upper surface. The petiole itself under certain cir- 
cumstances, has the power of producing the so-called adventi- 
tious buds, not only on the portions determined by the position 
of the leaf (leaf-axil), but sometimes on any other portions; a 
power enjoyed by the root in many cases. Hence parts of plants, 
otherwise most dissimilar, when they contain cambium, may 
have the power of reproducing the plant.f This is the founda- 
tion of the Schultz-Schultzetistein-ian doctrine of anaphytons; 
viz., those vegetable members “which, even when separated from 
the plant, continue to live, bud, and develop,and which are 
hence regarded as the individuals proper, as the true element- 
ary forms or morphological elements ; and it is by various com- 
binations of these that the organs (commonly so-called), root, 
stalk and leaf, are formed, by the repetition of which the whole 
plant is built up and indefinitely renewed. 
But where are the limits of the anaphytons ? How shall lines 
be drawn to include all the buds of the root, stalk and leaf, from 
which new formations may spring ? Aub. du Petit Thouars§ 
who had already developed doctrines similar to those of the ana- 
phyton-theory, attempts to draw the line between individuals by 
means of the cellular tissue, regarding every vascular bundle as 
an individual, since it has in itself, and independently of all oth- 
ers, the means of its growth, its preservation, and the reproduc- 
tion of new bundles. But it is difficult to perceive how, in such 
a view, the labyrinth of anastomosing bundles, (not less compli- 
cated in the majority of petioles, than in most reticulated leaves,) 
* Cf. Treviranus: Pflanzcnphysiologie, where several examples are adduced. 
f Aristotle himself says that plants possess the power of reproducing “ stalk and 
root” in every one of their parts. (TravTaxii 7a? *at pi£ctv xal xauAovi (Suv&iif. Yit. 
long, et brev. c. 6, p. 467). • 
\ Schultz: die Anaphytose (1848) and, System, der Morpliologie (1847). The 
passage quoted is taken from his later work, Verjiingung irn Pflanzenreich (1861). 
The remark made above, when treating of the members of the petiole, holds good 
here. The so-called Anaphyta can by no means grow into new plants themselves ; 
on the contrary, the new plant is produced as a germ, which is not identical with 
the anaphytons. 
§ Essais sur la vegetation considers dans le developpement des bourgeons (1809) cf. 
e. g., p. 174. “ C’est done le bourgeon en qui reside toute l’energie vegetale; aussi 
le regarde-t-on depuis longtemps comme un individu D’apres les principes que 
j’ai developpes dans mes precedens memoires, il faut aller plus loin, car je crois que 
chaque fibre vegetale est un Individu, puisqu’elle a en soi, independamment des 
autres, les moyens d’accroissement, de conservation et de reproduction.” 10 
The Vegetable Individual, in its relation to Species. 
can be disentangled and resolved into separate individuals and 
why the same independence and the same rank should not be 
allowed to the parts of the vascular bundles. And how shall 
we regard the lower plants, which have no fibres at all ? If 
our conclusions are to be anything more than mere arbitrary 
assumptions, we must go still farther; and we shall find no 
halting-place till we reach the cell, the true seat of every reno- 
vation in the plant, the starting-point of all non-sexual increase,* 
as it is of sexual propagation.! The cell has a better right to 
be considered as the vegetable individual than any other subor- 
dinate member of the plant; when connected with other cells 
it still continues to be an independent sphere of formation, 
sharply defined and, in youth at least, completely isolated.! Be- 
fore the universal law of cell-formation was known, and before 
botanists had succeeded in reducing all the elementary organs of 
plants to cells, Turpin hit upon the idea of seeking the vegetable 
individual in the cell ; though his views did not rest on as solid a 
foundation as Schleiden’s assertion that: “ in a scientific point of 
view, the cell is the vegetable individual. 
The most reliable authorities have agreed that new cells can 
never be formed externally to, but only within, other cells al- 
ready formed,H so that cell-multiplication must be regarded as a 
propagation, while all the cells of the mature plant must be re- 
garded as the progeny of the first embryonic cell. Besides, each 
and every plant is at first a cell; and there are single-celled plants 
in the strictest sense of the term, in which the first formation 
of new cells is that destined to reproduction ; i. e.: the germina- 
ting cells or spores.** Again, there are other plants in which the 
cell-generations contained between the first generation (which 
* Earlier investigations into the origin of adventitious buds had made it probable 
that, in its formation, each new shoot arises from a single cell. The first convincing 
proof of this fact, Avas given by llofmeister (Vergleichende Untersuchung u. s. av. 
der Coniferen, p. 94), in Equisetum. The propagating cells on the foliage and edges of 
the leaves of liverwort, which develop into new plants, have long been known. The 
spores of the Cryptogamiie belong here, as they are cells originating and developing 
non-sexually. 
f Pollen-cells, and the embryonic utricle and germinating cells,—as well as those 
of the archegonium of the higher Cryptogamiae. 
X Malpighi himself (Anatom. Plant., 1675), calls cells utriculi, or sacculi, though 
he distinguishes the wood and bast-cells as “fibres',' the vascular cells as “fistula:” 
and the cells containing milky sap as “ vasa spccialia.” As early as 1805, Link (Ro- 
mer’s Archiv., iii, p. 439), had expressed himself very explicitly in regard to the iso- 
lated position and the independence of cells : “ Quaevis cellula sistit organon pecul- 
iare, nullo hiatu nec poris conspicuis pneditum in vicina organa transeuntibus. Con- 
spicies non raro cellulam rubro colore tinctam inter reliquas virides.” 
§ Schleiden: Grundzuge, lte Aufl., 1842, vol. ii, p. 4, [Eng. trans. (1849), p. 127 T.] 
Cf. Schleiden: Grundzuge, i, p. 267 [Eng. trans. p. 103 T], “ The process of the 
propagation of cells, by the formation of new cells in their interior is an universal law 
in the vegetable kingdom.” Mold: Anat. und Phys. d. veg. Zelle, 1851, p. 63. “Cell- 
formation in plants takes place only in the cavities of older cells, not between or 
upon them.” Schacht: die Pfanzenzelle (1852), p. 47. “ The formation of new veg- 
etable cells always takes place in the interior of cells already formed.” 
** E. g.: Ascidium, Chytridium, Codiolium (a genus lately discovered in Heligo- 
land) Sciadium, Hydrodictyon (the last two AArith “ colonial formation”). The Vegetable Individual, in its relation to Species. 
11 
sprung from spores) and the last (itself returning into spores) sep- 
arate from each other, so that all the cells belonging to one cycle 
of vegetable development are segregated, and live completely 
independent of each other.* The importance of the cell as 
an individual seems to be decided by these facts; that of the 
entire plant, as a superior whole composed of individual cells, 
seems to be settled, and a firm foundation for the doctrine of veg- 
etable individuality to be gained. But let ns try to obtain a 
clearer view of some of the most important of these facts. The 
view which regards all cell-formation as a process of reproduction 
rests upon observations of the formation of free daughter-cells 
(blastidia) in the contents of the mother-cells (matrices),—the 
so-called free, or endogenous, cell-formation. Schleiden, who dis- 
covered this process, and Karstenf the most decided and original 
of his followers, regarded endogenous formation as the universal 
law of cell-formation. By this view the whole doctrine was 
turned in a wrong course, from which it could only be gradually 
recovered by the discovery, or rather the farther investigation, 
of another mode of cell-formation, which Nageli designated as 
“ Avandstandige,” Unger as “ merismatic,” and Mohl as “cell-for- 
mation by division of the primordial utricle.” But even at this 
day the misconception caused by generalizing the view that new 
cells are formed within old ones, has not been entirely removed. 
I have already! called attention to the fact that cells are divided 
which have no cell-wall, which is often the case among the 
Algse.$ In several genera in which numerous spores are formed 
in one mother-cell, its entire contents first divide into two parts 
(the so-called daughter-cells) which, without first secreting a 
cell-wall, immediately divide again into two; and this process 
may be repeated over and over,J| according to the number of 
spores which are to be formed (8, 16, 32, &c). In the second 
and subsequent divisions there is no formation of new cells in 
old ones, of daughter-cells in mother-cells, and hence no repro- 
duction, in the sense of one or more individuals being produced 
in an old one. The entire mother-cell is converted into two 
filial cells ; the filial cells are nothing but the mother-cell divided. 
And this is essentially the case in every cell-formation by division : 
for the wall of the mother-cell (within which the division gener- 
* Many Palmellacese, Desmidiacese, and Diatomeas. Cf. Braun; Verjungung, p. 132, 
et seqq. 
f H. Karsten, (De Celia Vitali, 1843,) emphatically rejects every mode of cell-for- 
mation by division and by sprouting, and asserts that every cell originates at its first 
appearance as a dot-like utriculus; regarding all formations found in the contents of 
the cell as cell-brood. 
\ Cf. Verjungung, p. 245. 
§ E. g.: Protococcus (viridis), Characium, Pediastrum, Ulothrix, Enteromorpha, 
Ulva, etc. during the process of spore-formation. 
| Nageli (Monocellular Algce, p. 28) calls such cell-generations “ transitory gener- 
ations.” 12 
The Vegetable Individual, in its relation to Species. 
ally takes place) certainly is not the living mother-cell, but merely 
its cast-off garment, its perishing shell. Cell-formation by divi- 
sion (called the “ merismatic” or “ wandstandige”) is that which 
obtains through the whole realm of vegetative development; 
while free cell-formation occurs only in fructification. Thus, the 
same phenomenon, which, regarded as endogenous cell-formation, 
seemed so favorable to the importance of the cell as the vegeta- 
ble individual, when more justly comprehended only brings us 
back to the divisibility of the vegetable organism, repeated in the 
most heterogeneous spheres. But still more : even the cell whose 
contents are not converted by division into new cells, but re- 
main simple, presents phenomena which can hardly be reconciled 
with their view by those who regard such a cell as an individual, 
isolated in space and independent in time. In the genera Vau- 
cheria, Bryopsis, Caulerpa, and other related Alga; in the fam- 
ily of SiphonicB, we find such cases, examples of the most extra- 
ordinary kind of cell-formation. The single cell, which forms 
the vegetable organism of these plants, has in fact a development 
which may continue indefinitely. Certain parts of the elonga- 
ted stem-like cell shoot forth into branches which lengthen by an 
independent terminal growth, without separating from the cavity 
of the maternal trunk by any partition. The principal trunk of 
the cell is either creeping, with an indefinite terminal growth 
though dying off from behind (Caulerpa-prolifer a)* or it is up- 
right and deciduous, while the sucker-like branches, club-shaped 
at the ends, and filled with a denser contents, are perennial ( Vau- 
cheria tubcrosa).-f In both cases the branches separate from the 
dying trunk, closing up at the bottom; and thousands of new 
trunks may thus be produced without any proper cell-formation. 
Thus the cell leads us back to the point from which we started 
at the tree; and, as we could not refuse individuality to the rami- 
fications of the tree, neither can we refuse it to the ramifications 
of the cell. Hence we cannot regard the cell as an absolutely 
single being, completely isolated and indivisible. Shall we pen- 
etrate still further into the anatomy of the cell itself, in the hope 
of possibly finding a valid vegetable individual ? All that we dis- 
cover here is, first, the vesicles, spherules and granules in the con- 
tents of the cell (amylum, chlorophyll and other pigment-vesicles, 
spherules of fat and, finally, the granules of the viscous cell-con- 
tents, whose chemical nature it is difficult to determine); and sec- 
ondly, the fibres, which compose the cell-membrane according to 
the old view advanced by Grew and lately revived by MeyenJ 
* Of. Kageli’s important paper on this plant (Zeitschrift fur wissen. Bot., i, p. 134), 
especially the exposition of the above-mentioned relations beginning p. 158. 
f A new species from the vicinity of Lake Neuenberg in Switzerland, remarkable 
for its purely furcated ramifications, with constrictions at the bottom of the branches, 
as well as for the club-shaped suckers at the ends. 
\ Meyen: Pfianzenphysiol., i, p.45 ; answered by Mohl, in his Vermischte Schrif- 
ten, p. 314. The Vegetable Individual, in its relation to Species. 
13 
and J. Agardh.* These parts, it is tru$, have often been regarded 
as the elementary formsf of plants, or their primary “ individual- 
ized” bodies the attempts, however, to represent them as the 
true and real vegetable individuals are not numerous; and they 
astonish us by their daring rather than entice to imitation. 
Turpin, who commenced by considering plants to be composed 
of different kinds of individual cells, which he compared with 
various lower plants (especially the Algae-genera Protococcus and 
Conferva), afterwards expanded his views, so as to regard the cells 
themselves as individuals of a second rank; while he consid- 
ered the true primary individuals to be the granules of the cell- 
contents, from which, in his opinion, the cell (cell-wall) is formed 
by agglomeration.<§> Mayer of Bonn, basing his theory upon 
molecular motions, considers the smallest granules of the cell-con- 
tents as individuals possessing animal life (biospheres) which 
buildup plants for their dwellings. “Like hamadryads these 
sensitive monads inhabit the secret halls of the bark-palaces we 
call plants, and here silently hold their dances and celebrate their 
orgies. ”|| 
Farther than this we cannot go : if we did we should have to 
leave specific vegetable life, and, instead of investigating its most 
minute spheres of formation, the visible cells, vesicles, granules 
or monads, turn to the invisible individual of brute matter, so 
as to consider plants as phenomena of appellant and repellant, co- 
herent and incoherent atoms. If we must understand by an in- 
* J. Agard: de Cell. Veg. fibrillis tenuissimis contexta (1852). Notwithstanding 
the importance of the author’s new investigations, they still need a more searching 
examination, as some points directly contradict well-ascertained facts, e. g.: the di- 
rect transition of the fibres from the outer to the inner layers of the cell-wall. The 
whole theory of the formation of cells by the uninterrupted growth of fibres cannot 
be admitted in view of the undoubted independence of the formation of tbe cell-wall 
from the contents. Mold is certainly right in regarding the fibrous division and di- 
visibility of many cellular tissues as a mere structural relation of the membrane 
(which in other parts is continuous); and he thinks it depends upon the peculiar 
mode of agglomeration of the molecules. As such molecules of the cell-wall are in- 
visible, I think it preferable to regard it as dependent upon a regular change of the 
relations of density. 
f Kutzing: Phil. Bot., i, p. 126, 129, does not regard the cell as the elementary 
form of plants, but as a complicated structure itself, and preceded by many other 
more simple forms, which he comprehends under the name of “ molecular tissue,” 
and which, he says, present in themselves many lower vegetable forms. Plants 
which are not even cells! 
\ Unger: Grundz. d. Anat. u. Phys. der Pjlanzen, p. 4. The cell is represented as 
the “ elementary vegetable organ but the vesicles, fibres and granules within it are 
further distinguished as very minute, “ individualized” bodies. 
§ Turpin: “ Sur le nombre deux,” (Mem. du Musee, xvi, 1827, p. 305): “Ainsi 
des individus globuleux, rapproches simplement contigus, foment la membrane de la 
vesicule Individu du tissu cellulaire, le filament Individu du tissu tigellulaire, et la 
membrane cuticulaire Individu. Des agglomerations de ces derniers constituent les 
Individuality, provenantes des bourgeons developpes, et enfin, celles-ci achevent 
VIndividualite composes d’un arbre.” 
|| Mayer: Supplements zum Lehre vom Kreislauf (1837), p. 49. I am acquainted 
with Mayer’s views through Meyen’s Pflanzenphys., ii, p. 256. 
*[ Cicero calls the atoms “ Individual 14 
The Vegetable Individual, in its relation to Species. 
dividual, a being perfectly,simple and indivisible, this is our last 
refuge, in which we may indeed reach an individual, but not a 
vegetable individual ; for this would then be identical with the 
material individual common to all corporeal existence. But, even 
if we could give up all hopes of a specific vegetable individual, 
doubt would still linger round these physical individuals; for even 
the existence of the universal primary particles of bodies,—the 
material individuals, the atoms,—is not conclusively established. 
No eye has seen them; we do not even think of considering them 
as objects of direct perception; we only accept them as an hy- 
pothesis, to eke out our theories of motion and of chemical affinity, 
and to enable us to compute their relations. The question might 
easily be asked, whether the same phenomena may not be as well 
explained by assuming the continuity, expansibility and penetra- 
bility of matter. However this may be, the question concerning 
the existence of atoms certainly lies beyond the limits of botan- 
ical investigation; and if the existence of vegetable individuals 
depends on this question, the botanist must despair of proving it. 
Thus the question at which we have now arrived is this: can we 
speak of individuals in botany? and this is identical with an- 
other : are plants mere products of the operations of matter (i. e., 
of a substance self-moving, uniting and separating by an innate 
force), and hence non-entities, or mere phenomena resulting from, 
or produced by, the blind forces of nature ; or may we ascribe 
to plants an independent existence in nature, notwithstanding 
their connection with the external world ? 
If what we call plants are nothing but complex chemical and 
physical processes, then we can no longer speak of their individ- 
uals and species in the sense the words usually bear; for the 
mere phenomena of the operations of the primary substance, 
which have no other efficient principle than the forces of this 
substance, cannot be regarded as self-existent beings, or as pe- 
culiar (specific) kinds of these beings, or as single (individual) 
modifications of them. This is, in fact, the result towards which 
the later physiological investigations are hastening, grounded on 
the positive results of investigations in the physical sciences. 
Even vegetable physiology cannot resist this tendency of science, 
although it struggles more or less against these conclusions.* The 
operations by which plants, and all organic beings, form and pre- 
serve their organisms, were formerly ascribed to peculiar vital 
forces; but the physiology of our day would recognize in the vital 
functions of the organism the same forces by which the processes 
of inorganic nature are perfomed. Thus physiology becomes 
* Even Schleiden, the most prominent and most decided of the representatives 
of this tendency, seeks to counterbalance the deadening effects of the purely mate- 
rialistic view by an aesthetic one (Die Pflanze und ihr Leben; last lecture: d. J5s- 
thetik der Pflanzenwelt). The Vegetable Individual, in its relation to Species. 
15 
physics and chemistry, or, according to the usual conception of 
the physical and chemical processes themselves, the “ mechan- 
ics” of organic nature in the most comprehensive meaning of the 
term mechanics. And thus the life of the enchanter is un- 
veiled, who had seemed to be the immediate cause of his own 
works; the lofty partition-wall between organic and inorganic 
nature falls, and one common foundation is laid for investiga- 
ting all material processes in every realm of nature. This impor- 
tant result is reached : the existence of the higher orders of nat- 
ural phenomena, which had been regarded as the peculiar realm 
of Life, is referred to the same natural causes (the same mate- 
rial substance and the same kind of forces) by which the lower 
orders, those of “inanimate” nature, have their being and per- 
form their functions. Still further conclusions may be attempted, 
and it is in the nature of scientific progress that these attempts 
should be made. As physical forces seem to be everywhere 
indissolubly connected with matter, and as a fixed regularity dis- 
plays itself in their operations, men were found bold enough to 
consider the totality of natural phenomena as the result of orig- 
inal primary substances, cooperating with determinate forces, 
according to the laws of a blind necessity a natural mechan- 
ism revolving in its endless orbit.* 
Though this view seems to explain all the phenomena of na- 
ture from one principle, in fact it precludes any real explana- 
tion of them, that is when exlusively applied to their solution. 
That which is eternally necessary can only be conceived as eter- 
nally carried out; and thus any real event becomes an absurdity. 
If the “mechanical” (physical and chemical) forces of nature are 
necessarily active, then if any motion is to take place, the first im- 
pulse, the proximate cause, cannot be explained by the nature of 
the motion ; it must be another principle above necessity ; and this 
is true not only of nature as a whole, but also of every particu- 
lar motion in nature as well. Thus not only the first impulse, but 
the universally apparent final cause, remains an inexplicable rid- 
dle in the doctrine of blind necessity. Hence the insufficiency 
pf the “physical” theory, compared with the “teleological”! is 
peculiarly obvious in the realms of organic nature, where the 
* As far as concerns Natural History these views are developed, e. g.: in both of 
Mohlschott’s works: d. Physiol, des Stoffwechsels in Pflanzen u. Thieren, (1851), 
and d. Kreislauf des Lebens (1852); in the last mentioned work we find such sen- 
tences as these: “ The miracle of nature is the interchange of matter, the first- 
cause of physical life,” p. 83. “ Creative omnipotence means the relations of mat- 
ter,” (p. 258). “ The hinge round which the wisdom of the present day is turning 
is the doctrine of the interchanges of matter,” (p. 363).—The doctrine, that the uni- 
verse is the play of attrahent and repellant atoms, belongs, after all, to the “ wis- 
dom” of the past, professed by Democritus and Epicurus. 
f Cf. Schwann: Microscopische Untersuchungen iiber die Uebereinstimmung in 
der Structur u. d. Wachsthum d. Thiere u. Pflanzen, (1839), especially p. 221-225 ; 
on the other side, Eschricht d. Physische Leben, (1852), in sections ii and iii. 16 
The Vegetable Individual, in its relation to Species. 
final cause of each particular life appears so distinctly. The ad- 
vocates of the physical view perceive this; but they explain the 
fitness of means to ends in nature as a whole, and in its individ- 
ual parts, by supposing matter, with its blind forces, to have been 
created by an intelligent being.* But we can regard this as a 
germ of an explanation only in proportion as it is also granted, 
that the intellect of the Creator lies not only behind and without 
nature and her processes of development, but that, as if incorpo- 
rated in nature, it is taken into the destiny of each created being, 
in proportion to its individuality. But this, again presupposes the 
admission of a substantiality of nature fit for such an hypothe- 
sis ;—a substantiality not grounded on mere matter, like a blind 
force; but which, on the contrary, must comprehend matter as 
subordinate to itself, and must realize itself through matter :—an 
assumption which modifies the physical view essentially, and 
would seem to be a modification of some ideal, or teleological 
theory. 
Without underrating the great importance, which the phys- 
ical view possesses for vegetable physiology, still we must confess 
that we cannot find in it the key to a conception of vegetable in- 
dividuality: for, after all, this must be sought for, not in the ex- 
ternal conformation, but in the essence of the plant, determined 
from within. This leads us from the last negative results to an 
historical view of the attempts at a positive explanation. 
It is evident from the foregoing review that, if we would not 
give up all hope of conceiving plants as beings, realized in indi- 
vidual conformations, we must not allow so great and decisive an 
importance to the external divisibility of their organism as has 
been usually done. We must seek :a decision in the essential 
concatenation of all the steps in the plant’s development forming 
one whole, according to one idea. This is the tendency of the 
concluding remark of Nageli, to which he is lead by the rela- 
tions of growth and propagation in Caulerpa ; when he says that 
indivisibility of form is not an element essential to individuality, 
—which, indeed must be constructed upon a new, and somewhat 
less material a basis. Link calls attention to this same unity, 
which is expressed in the whole development of the plant, and 
which forms the essence of its individuality, in the following true 
words: “ We cannot recognize an individual unless we are con- 
vinced that it remains the same in different periods of its exist- 
ence.”f Now the question is just this: how can we perceive 
* “ The fitness of means to ends, in every organism, even a superior degree of 
this individual fitness, cannot he denied; but in this (the physical) view, the cause 
of the fitness does not consist in the fact that every organism is produced by an in- 
dividual force tending towards a certain end, but, like the cause of the fitness of 
means to ends in the inorganic world, that matter is the creation of an intelligent 
being.” Schwann, 1. c., p. 221, and, in almost the same words, p. 224. 
•(■ Link : Eleni. Phil. Bot., Ed. ii, p. 11. The Vegetable Individual, in its relation to Species. 
17 
such a oneness of essence amid these changes of form and mate- 
rial ? How do we perceive that, with all its divisibility, the 
plant remains after all really one and the same individual ? 
Every development presents a succession of phenomena, which, 
while they present themselves in a regular order, also show un- 
mistakably a point of departure, an end, and a course between 
the two advancing after a fixed plan, and which indicate a com- 
mon internal principle.* They point to an internal vital principle,! 
common to the whole succession ;—to a principle which must be 
conceived of, not only as an idea which guides the whole pro- 
cess, or as a force determining the specific type of this plastic 
succession, but also as a living essence, comprehending the idea 
as its internal determination, and the force as the means of its 
realization ;—an essence which precedes and shapes the external 
existence ; as intentions precede and determine acts-! If, in ac- 
* Du Petit-Thouars, 1. c., p. 284 : “ L’individu est un etre dont toutes les parties 
sont subordonnees a un principe unique d’existence.” Link, Elim. Phil. Dot., ed. ii, 
p. 3. “ Nos individuum vocamus, quod ab uno eodemque principio interno deter- 
minatum est, ad idealem potius quam ad realem respicientes divisionem.” 
f Spring: Ueber d. Begriffe v. Gattung, Art u. Abart, (1838), p. 65. “ It is this in- 
dwelling principle which makes the individual; and in natural history, every body is 
an individual in as far as it really exists as a single being, whose existence is deter- 
mined by a peculiar indwelling vital principle.” Spring afterwards distinguishes 
the systematical and the physiological individual: in the former one moment of the 
development is comprehended, in the latter the whole metamorphosis. The physi- 
ological individual comprehends an assemblage of forms, which might be regarded 
by a casual observer as so many systematical individuals.—Still, a true systematick 
must protest against such a purely subjective distinction of systematical and physi- 
ological individuals. However much the embryos of mosses resemble Conferva?, or 
the larva of an insect resembles a worm, a true systematick will not separate the 
young individual from the developed one ; and genera which are founded upon our 
ignorance of their successive development, as Protonema, Lepra, Sclerotium, etc., 
must be given up by the systematist himself. True, we shall be called upon at a 
later point in this inquiry to decide, whether a sphere of development which really 
belongs to the individual can present itself to us so divided that the divisions them- 
selves attain to the importance of subordinate individuals. 
\ Aristotle describes the internal essence of plants as a “plastic soul,” (HpirrixTi 
■4uxn, tou £uvtos cwgaTos am'a. xal apx’i)* Cf.Wimmer: Phytol. Arist.frag., c. iii, de pi. 
vita atque anima. The charge of anthropomorphism has been made against such a 
view, which attempts to conceive of nature as a chain of essences, both in the recip- 
rocal relations of her forces, and in her internal developments ; but, if man himself 
is a member of nature, if he is the highest member in the order of natural beings, 
that member which presents the most complete unison of all the phases of life in 
nature,—then all his knowledge of nature must be connected with his knowledge of 
himself. However meanly we may estimate this knowledge at the present stage of 
psychological science, still it is sufficient to assure man of his own “ ego. And if 
man is justified in regarding himself as a human being, by analogy he is justified in 
regarding his relations, the animals, in the same manner, as animal beings; plants as 
vegetable beings ; and every single animal, every single plant, as an individual being 
(even though included in a higher entity). To attain a unity of idea in Natural 
History, man must apply this idea farther down in the scale of nature, and must re- 
gard minerals, even the elements themselves, as beings of their own kind. But the 
materialist will reply: individual beings are only the elementary substance: all other 
beings, so called, are formed by a temporary composition and cooperation of these. 
But who has seen these elements of chemical combinations, as elements,. or has 
proved their existence in any way ? But even if they should exist as such, is it not 18 
The Vegetable Individual, in its relation to Species. 
cordance with this idea, we regard external development as the 
revelation of the internal essence, which exhibits its purport in 
the processes it undergoes in connexion with the world without 
it, and whose realization is thus produced by a determinate sphere 
of activities, necessary for such a realization, then, vice versa, we 
may infer the essential unity of each particular sphere of devel- 
opment, from the complete unity of the functions relating to this 
realization. This leads us to the attempts made at a physiolog- 
ical determination of the vegetable individual. The usual defi- 
nition, and one entirely in accordance with the physiological 
point of view, is that an individual is a perfect representative 
of the character of the species, possessing all the functions neces- 
sary to the continuance of the species. Now if we would con- 
ceive of a physiological individual, in the broadest meaning of 
the term, we should certainly be compelled to demand that our 
conception should be such as to exhibit not only single phases, 
but all the phases of the specific life during its entire develop- 
conceivable that a higher being should include the lower beings? We say: hydrogen 
and oxygen form water ; but it would do as well to say, water forms itself out of 
hydrogen and oxygen. The elements do not form the plant; the plant forms its 
body out of the elements. We may declare both these views to be hypotheses; but 
of hypotheses that is preferable which is nearest to man,—I would almost say, most 
necessary to man’s nature, when he proceeds from the data of his own existence. Shall 
the elements have a stronger claim to be acknowledged as real existences than man 
himself? Or will any one say that it is a more daring hypothesis to assume that 
man thinks ; that brutes move themselves; that plants themselves produce the deter- 
minate form of their organism, than to suppose that elementary substances in their 
connexions and cooperations produce the phenomena of thought, voluntary motion 
and typical conformation ? But after all, is it not true that the elementary sub- 
stance is everywhere present ? that without it none of the phenomena just mentioned 
can occur ? Certainly, this is so; the higher stages cannot be realized without the 
lower, which enable them to exist; but these higher stages can never be explained 
by, and comprehended in, the lower. No one, as yet, has shown even the shadow 
of a possibility of explaining, from the things themselves merely, why the ele- 
mentary particles form a mineral kingdom, a vegetable kingdom, an animal king- 
dom, and man. And why do they not fulfill their task after an eternal immutable 
manner, since such a fulfillment is one of their necessary, eternal, and immutable prop- 
erties ? Why have they succeeded in composing man only in the most recent geo- 
ogical epoch ? Why have they not from eternity produced in man’s brain the the- 
ory of their actions, and thus, in accordance with their eternity, eternally manifested 
and glorified themselves ? The most industrious investigations into the relations of 
the physical world promise us a deeper insight into the regular connexions of all 
the parts of nature ; into the cunning mechanism, which carries on and upholds all 
natural life. Still a key to the interior of this structure, and an admission to the es.- 
sence of plastic nature in her operations, cannot be found by our investigations if, 
by presumptuous hypotheses, they debar us from the higher realms of development, 
especially those of organic nature and of human life. Flesh and blood are hypoth- 
eses ; but mind is truth, says a well known writer; and Des Cartes could find a 
proof ol his own existence and of that of the world around him in his mind alone. 
It would be a strange contradiction, if the investigation of the most distant realms 
into which the human mind can penetrate should rob us of what is nearest and 
surest, the intellectual ego itself, the starting point of all investigations. But he 
who has not recognized the foundations of the spiritual world in nature itself, must 
of consequence deny their existence in man, if he would not lose, in an inexplicable 
dualism, the hope of obtaining coherent views of nature. The Vegetable Individual, in its relation to Species. 
19 
ment; that it should realize all the capabilities of the specific be- 
ing, and thus present to us the whole plan, the whole destiny of 
the species. If we examine the preceding conclusions from this 
point of view, it will be evident that single cells cannot be such 
individuals ; for, although the whole construction of the plant 
and all the functions of its life are carried on by means of the 
cells, still, viewed as a connected whole, the cells are only single 
stones, single elements, in the great mechanism of the organism. 
Any single member of a plant (as the internode and leaf) corres- 
ponds no more to such a physiological individual than does the 
cell; for plants undergo their metamorphoses in their successive 
members; and the various processes of their preservation, repro- 
duction and propagation are connected with the various steps of 
these metamorphoses. Nor can it be the shoot; for that usually 
does not embrace all the steps of the metamorphosis; besides, 
the functions are variously distributed in the shoot; and in many 
cases, this takes place for the reciprocal completion of the func- 
tions. Besides, whatever is characteristic in ramification and in 
growth depends upon the combined shoots, and without these it 
is impossible to conceive of trees, for instance. Then we come 
back to the whole plant-stock ? Nay, farther ; we cannot stop at 
the plant-stock ; for the single stocks are far from being perfect 
representatives of all the phases, and tendencies of the specific life. 
I would refer to the division according to gender, or the modes 
of fructification, which is often made in botany ; the dioecious 
and trioecious,* relations, and farther, to the varieties, especially 
to those which do not possess essential organs and functions, 
which belong to the species as such ; e. g.: those varieties which 
never bear blossoms (Ball-acacias), or which never produce fruit 
(congested blossoms), or which never perfect seeds (currant- 
grape, cultivated bananas and bread-fruit trees). Besides, no 
stock is exactly similar to another: we ascertain only the limits 
of the possible relations of the specific form by a comparison of 
many stocks. As in animal physiology the solution of the prob- 
lem of the life of many animals depends upon their social relations 
(societies composed of couples or of flocks, or of self-governing 
states), so in vegetable physiology it depends upon characteristic 
physiological traits whether plants live single and dispersed, or in 
societies. For example, in considering the life of turf-mosses we 
must determine whether they grow in great sods or in carpets; 
and of grasses, whether they form meadows; or of trees, forests. 
Even the relations of geographical distribution, which are discov- 
ered by a comparison of all the stocks, depend upon the physio- 
* Trioecious plants are exceedingly rare among Phanerogamic (Ceratonia, some 
kinds of Rhus), but are more common among the Cryptogamic; perhaps we may 
add the Floridic. In Polysyphanio violacea I have found three kinds of stocks mixed, 
and in the same stage of development in the same place : (upon the same thread in 
Chorda Filurri). 20 
The Vegetable Individual, in its relation to Species. 
logical character of the plants: plants of sensitive and inflexible 
constitutions are found only within narrow limits ; while plants of 
adaptive and pliant constitutions are more widely distributed, be- 
come migratory plants, and by degrees spread over almost all parts 
of the earth, if their seeds possess the necessary properties. From 
these considerations, and many others which might be adduced, 
it is obvious that there are no determinate limits to a purely 
physiological conception of the vegetable individual; and that 
we may expand the definition of the individual until it coincides 
with that of the species itself. 
How then can we steer a middle course, between the mor- 
phological view, which results in indefinite subdivision, and the 
physiological, which ends in indefinite expansion ? The physio- 
logical view has shown that none of the divisions or spheres of 
formation, which have been regarded as the individual ones, fully 
realizes the idea of the species; and that each needs the others 
to render this idea complete. The morphological view has shown, 
in the same manner, that there are subordinate and comprehen- 
sive spheres of development, none of which exhibits complete 
independence, since all appear in unequal degrees, as more or less 
perfect members of the entire succession of the specific develop- 
ment. If we would discover the individual under such circum- 
stances, we must not demand of it all that belongs to the species ; 
for this is completely represented only in the totality of the indi- 
viduals, not in any single individual. We must answer this ques- 
tion : Which member of the graduated potential series in the 
sphere of development subordinate to that of the species deserves 
preeminently the title of individual ? And we shall be compelled 
to reply : That which exhibits the most complete independence 
and definiteness. Good use has decided in regard to man (and 
the higher animals), and it justifies itself by the fact, that what is 
usually termed an individual undoubtedly possesses great organic 
independence: and this is true both of its subordinate spheres (i. e. 
the members of the organism, down to the cells) and of those by 
which the individual is comprehended (family, state, race, etc.). 
By means of comparison and analogy, the signification of the 
more doubtful spheres of development among the lower animals 
and plants may receive some new light from such a view. I pro- 
pose to attempt this in the second part of this Investigation, but 
now I will only subjoin a few general remarks. 
In the conception of individuality, there are two elements ; that 
of multiplicity, and that of unity. Each development exhibits 
multiplicity; but this multiplicity is not equally subordinate to 
the unity in every development. The more complete this sub- 
ordination, the more perfect is the individuality; for it is only 
this subordination to the unity which binds up the multiplicity 
of the conformation into an indivisible organism. The less com- The Vegetable Individual, in its relation to Species. 
21 
plete the subordination, the more perfect will be the independence 
of the parts, and the more indefinite will be the individuality of 
the whole. If we apply this view to plants, whatever is dubious 
in our conception of vegetable individuality will be explained. 
Successive development, we may say, is the peculiar nature of 
plants, which, beyond the power exhibited in the process of for- 
mation and propagation, possess no higher vital power; while in 
animals the process of the formation of the body appears only as 
an operation preparatory to its connection with a higher vital ac- 
tivity. For animals, in addition to their powers of external man- 
ifestations, have a power of internal vital comprehension, which 
expresses itself in the life of the soul (by which animals possess 
an internal centre, from which the organism is governed and reg- 
ulated). It is the soul alone which connects in indivisible unity, 
and for reciprocal services the products of the plastic power, and 
gives to the organism of animals the character of a definite indi- 
viduality. Among plants the case is different: plants in their ope- 
rations are active solely in one direction, externally—are split up 
so to say in the process of external conformation, so that the parts 
appear less connected, as compared with the plant as a whole more 
independent, and more divisible among themselves. Thus the 
vegetable organism is a dividual, rather than an individual ; a 
multiplicity* rather than an unity ; i. e. a whole whose parts hold 
the same relation to each other as individuals to each other, but 
which present spheres as indivisible as the whole itself. This is 
the doctrine of the relativef individuality of plants, which Stein- 
heil has especially noticed. According to this doctrine, different 
orders of vegetable individuals, as it were different powers of indi- 
viduality, are distinguished. In the same manner 
distinguishes the cell-individual (Vindividu cellulaire, in which 
he has been preceded by Turpin); the bud-individual (Vindividu 
bourgeon, after Darwin); the slip-individual (Vindividu bouture); 
the stock-individual, or the vegetable individual (Vindividu vege- 
tal penes quern est jus et norma loquendi); and the embryo-indi- 
vidual (Vindividu embryon), which, in accordance with the mean- 
ing in which Gallesio used the term, comprehends all that pro- 
ceeds from one germ, even if multiplied by division. Since the 
* “ Planta est multitudo.” Engelmann: de Antholysi, p. 12. 
f Steinheil: 1. c., especially p. 4 and p. 17 : “Les vegetaux ne peuvent arriver a, 
l’iudividualite absolue; ils se presentent a nous dans un etat, qu’on peut designer par 
le nom d’individualite relative; ce qui distingue cette partie de la creation du regne 
mineral, ou. l’individualite est nulle, et du regne animal, ou elle est presque toujours 
absolue.” 
+ BeCnndolle: Physiologic Veget., p. 957. The author does not attach much im- 
portance to his division, as he says he has assumed it for convenience of expression, 
and to avoid the usual confusion of language. His son Alphonse DeCandolle consid- 
ers it quite an arbitrary matter -which part of the plant we call the individual: “ Les 
vegetaux sont evidemment des etres composes: mais jusqu’ ou veut-on les decom- 
poser, pour que les el6mens s’appellent des individus ? C’est une chose arbitraire, 
qui depend de l’id6e par laquelle on se laisse dominer” (after Steinheil, p. 6). The Vegetable Individual, in its relation to Species. 
22 
slip-individual is essentially the same as the bud-individual (i. e. 
shoot-individual), we have four degrees of individuality, in which 
at least one more might have been easily inserted, between the 
cell and the shoot-individual, i. e.: the member or “ story”-indi- 
vidual (Gaudichaud’s phyton). With this view Schleiden’s divi- 
sion is connected: he distinguishes the cell as the plant of the 
first order ; the shoot as that of the second, which he calls the 
simple plant (a term borrowed from C. F. Wolf, who used it in 
the same sense); the whole stock as that of the third order, which 
he designates as the composite plant By a searching investigation 
into the shoot, I shall endeavor to decide whether all these rela- 
tive individuals can be considered individuals with the same jus- 
tice ; or whether, after all, one of them does not deserve the title 
preeminently, corresponding to the animal individual. In either 
case Goethe’s words may be applied with perfect justice to plants 
and their individuality : 
Freuet euch des wahren Scheins, 
Euch des ernsten Spieles; 
Rein Lebendiges ist Eins 
Immer ist’s ein Vieles. 
Herder, in speaking of the works of the Creator, says : “ Every 
one of Thy works Thou makest one and perfect, and like itself 
alone.” 
This sentence presents the other aspect of existence, by which 
the multiform is one ; and every unity in the one-sidedness and 
incompleteness of all single manifestations, is after all a perfect 
whole. These words lead us to the internal essence of things, re- 
ferring us at the same time to the primary ideas, which Nature 
comprehends and realizes in Life. 'From the American Journal of Science and Arts, 2nd Series, Vol. XX, Sept. 1855.] 
THE VEGETABLE INDIVIDUAL 
IN ITS 
RELATION TO SPECIES. 
The Vegetable Individual in its relation to Species ;* by Dr. 
Alexander Braun.—Translated from the German by Charles 
Francis Stone. 
Part II. f 
As I attempted to show in Part I, whatever seems arbitrary 
and indefinite in the existing views of what constitutes the Veg- 
etable Individual has its ground in the nature of plants them- 
selves, which, in their realization are resolved into a plurality 
which they are not capable of reducing to as complete an unity 
as animals are. As we ascend in the natural kingdoms, individ- 
uals increase in importance, until they reach their most perfect 
independence in'Man. Hence, if we would appreciate them justly 
in the lower departments, in which their character is less defin- 
ite, we must try to comprehend the less perfect structures by 
starting from the more perfect ones: to appreciate vegetable in- 
dividuals we must start from a comparison of animal individuals. 
* Das Individuum der Pflanze in seinem Verhaltniss zur Species,—Generations- 
folge, Genrationswechsel und Generationstheilung der Pflanze, by Dr. A. Braun, Pro- 
fessor of Botany in tbe University of Berlin, Ac. Ac. 
f For Part I, see Number for May, 1855. 2 
The Vegetable Individual in its relation to Species. 
From this point of view we perceive at once that the cell can- 
not be regarded as the proper individual in plants, otherwise it 
would have to be considered in the same manner in animals. 
Cell-formation is a property common to plants and animals: but 
in animals it appears far more obviously as a subordinate element 
in the organization of the whole body, than it does in plants ; 
since the animal cell, in most cases, is not so independent, nor so 
determinate, nor so permanently isolated as the vegetable cell. 
For this reason, too, it is rarer to find the animal cell considered 
as the proper animal individual, although Schwann has shown 
that animal cells are analogous to vegetable cells and may be as 
justly considered individual organisms as they. Yet as rnerecw- 
riosa we might adduce the somewhat similar assertion of Gaillon, 
that “ men and animals are properly masses of Infusoriaand 
Oken’s doctrine of generation, “ a synthesis of Infusoria,” might, 
perhaps, be interpreted in the same sense. The “ stories” of the 
axes, the internodes with their leaves, might claim to be com- 
pared with the animal individual with more justice than the cell, 
especially if leaf-formation really took place as the defenders of 
such doctrines have represented : that is, if every successive leaf 
were produced as a new structure out of the old one (out of its 
base which becomes the internode), and if the whole stem were 
thus merely a concatenation of leaves shooting out of and grow- 
ing above each other. But this is not so ; the rudiment of the 
stem as an uninterrupted growth (“ continuance”) is formed be- 
fore the leaves, while the latter, emerging as developments of the 
upper surface of the stem, are evidently members dependent upon 
and belonging to the axis, and forming with it one whole. Hence 
the structure of the internodes may be more aptly compared with 
the lateral structure of the animal body, and that of the leaves 
with its terminal structure. Thus we arrive at the shoot; and 
we must investigate the question, whether it should be considered 
as what corresponds best with the animal individual, or whether 
we must ascend still farther, up to the whole plant-stock. 
The Shoot as the Vegetable Individual. 
The first and most common view is that which considers the 
individual in plants, as in animals, to be merely each single spe- 
cimen, i. e., each representative of the species which appears to 
be one whole from the connexion of its parts. To some extent 
this view is correct, for in a forest of trees of the same genus and 
species, in a meadow, or in a cornfield, each single tree, each 
stock of grass or of grain, appears as a single member of its spe- 
cies, as each single beast does in a flock of animals forming a The Vegetable Individual in its relation to species. 
3 
community. But the question arises whether these individual 
beings, regarded as such in this superficial way, can each be con- 
sidered individuals in the same sense. When the flocks or socie- 
ties of animals are numerous, as in an apiary, each hive or swarm 
will appear as an individual member of its species, and the more 
so in proportion to the closeness of the connexion between the 
members of such a community. Many flocks of animals whose 
members are organically connected during life, have until lately 
been considered to be individual animals ; and even when the 
separation of the individuals is more complete, such conceptions 
are to a certain extent justified as long as the community is really 
a natural growth—when in fact it consists of members of one 
single stock—and we are not surprised to find that the oldest his- 
tory of the human race describes the family itself, and the tribe 
which springs from it, as one person, named after its patri- 
arch. As regards the plant-stock, even a superficial examination 
shows us peculiarities which will hardly allow us to consider it 
as an individual in the precise meaning of the term, and which 
calls upon us carefully to consider whether it is to be regarded 
as such an individual, or merely as an individual in the broader 
sense,—as one united family. Even our feelings aroused by the 
sight of the most ramified plant-stocks,—especially by a tree with 
its numerous branches, with the thousands of blossoms and fruits 
which it bears, and the numberless buds through which it will 
deck itself again in the following year with leaves and flowers— 
excite the presentiment that this is not one single being, one sin- 
gle life, comparable with the animal or the human individual, but 
rather a world of united individuals which have sprung from each 
other in a succession of generations, and although they do not 
separate, going through their particular cycles of existence,— 
here dying off, there reproduced, and thus building themselves up 
in uninterrupted succession into a family-tree, perennially laden 
with an increasing posterity. That such a view, so consistent 
with our healthy natural feelings, is corroborated by scientific in- 
vestigation, I hope to show in the following observations. 
Comparing plants with animal individuals, it is at once evident 
that the tree loses annually flowers and fruit,—the highest and 
noblest structures which vegetable life produces,—to generate 
them again in the following period of vegetation. Even the 
whole dress of the tree, even its foliage when compared with the 
trunk and branches, is only a superficial growth periodically 
dying off, and reproduced by the succeeding generation : in 
the paradoxical words of Schleiden :* “ No tree has leaves.” 
* Beitr., p. 152, where the following view of the arboraceous stem, as a common 
ground bearing many individuals is developed ; but this whole view, after all, needs 
to be corrected by a precise limitation of its meaning by what follows it. 4 
The Vegetable Individual in its relation to Species. 
The leaves, in fact, never grow out of the woody portions of the 
tree, but only on its herbaceous extremities, which grow upon 
the woody stem as upon a ground formed by the process of veg- 
ation. This common ground, namely the woody stem, which is 
almost lifeless in comparison with the herbaceous parts engaged 
in active growth, is annually covered with a vigorous sheath un- 
der the protecting bark, and this sheath is the ground of the nour- 
ishment of all the vegetating herbaceous extremities. This 
sheath is the so-called cambium, a layer of active, living tissue 
which, contemporaneously with the lignification of the herba- 
ceous extremities of the branches, becomes a new woody layer, 
united to the old trunk in the form of an annual ring—to be cov- 
ered in its turn in the following period of vegetation with a new 
layer, which, again, will be the immediate supporter of the new 
generations. The history of the grand development of nature 
on the surface of our globe presents an analogy which may per- 
haps serve to set this relation in a clearer light. The successive 
geological formations superposed during the course of countless 
ages, present, buried in their depths, the traces of as many forma- 
tions of the organic world, each of which carpeted the then su- 
perior stratum of the earth with a new life, until it found its own 
grave in the succeeding formation, when a new uprising of or- 
ganic life took its place. In the same way the stem of a tree is 
a multistratified ground, in whose layers the history of earlier 
growths are legibly preserved. The number of the woody layers 
indicates the number of the generations which have perished, i. e., 
the age of the whole tree ; a distinct annual ring is the monu- 
ment of a vigorous season, an indistinct one of a bad season, a 
sickly one (which is often found among healthy ones) indicates 
the unhealthiness of the foliage of that particular year. The 
practised woodman can decipher many facts of the past in the 
layers of the trunk, e. g., a good season for foliage or for seed, 
damage by frost or by insects, etc. 
Essentially the same relations as those seen in the tree, or the 
shrub, are to be found in the subterranean perennial growth of 
plantas rediviv® (herbaceous perennials) whose subterranean stem 
(rhizoma), like the stem above the surface, emits annually a new 
generation of herbaceous growths ; whose stalks however, unlike 
those of the tree, do not lignify and form a part of the common 
supporter, but die off wholly, or mostly, at the close of the sea- 
son of vegetation. 
The relations indicated above compel us to recognise a suc- 
cession of generations in trees, shrubs, and perennial herbs; and 
thus our first idea of them as individuals is necessarily modified. 
Another remark may be made here which confirms our idea thus The Vegetable Individual in its relation to Species. 
5 
modified. Natural death closes the life of the individual.* The 
development of the life of individuals in organic nature has a 
goal, an acme ; after it has attained this goal its course draws to 
an end. This is not the case in the tree and the perennial herb. 
True the tree is destroyed by time ; but this seems to result more 
from external, and in part mechanical causes, than from any in- 
ternal decrepitude. The more numerous the generations which 
the tree builds up, one above the other, the greater is the distance 
of the growing extremities from the source of their nourishment; 
the thicker the supporting trunk the thinner is the layer of cam- 
bium which connects the new shoots with the extremities of the 
root by which the nourishment is absorbed. This increased dif- 
ficulty of communication between the upper and lower extremi- 
ties is probably the cause of the decrease of vigorous growth af- 
ter the plant has arrived at a certain age. But in most cases ex- 
ternal casualties are superinduced, which accelerate the termina- 
tion of the tree’s life. It is injured by wind and weather, the de- 
cay of the injured part spreads through the whole organism, vari- 
ous fungi fix themselves upon the tree, and are especially fatal 
when they attack the roots. Oftentimes the tree breaks down 
under the weight of the productions of its own vital powers, the 
luxuriance of its fruit. These statements are corroborated by 
the cases of trees of unusual age, now so well known through 
De Candolle’s investigations. One of the examples adduced by 
him shows in particular that those trees whose branches have 
been prevented from breaking down by props or supports attain 
to a great age. I refer to the celebrated Linden in Neustadt on 
the Kocher, which, as early as 1229, was the cause of the town’s 
being called “Neustadt an der grossen Linde” (Neustadt of the 
great Linden), whose wide-spreading branches were supported al- 
ready in 1408 by sixty-seven stone pillars, and this number was af- 
terwards increased up to more than one hundred.f The hoary tree 
still flourishes, having survived its many scientific admirers, among 
whom was my predecessor, to whom Botany is so greatly indebted, 
who visited and described it a few years ago, (in I849).J Natural 
supports are more efficacious in preserving trees than even artifi- 
* Cf. Schleiden: Beitr., p. 151. “ The idea of individual life necessarily implies as 
its distinguishing characteristic individual death, preconditioned in the organization it- 
self.” Although this remark is not universally true in many respects, yet I have 
adopted it for the light it is calculated to throw on the nature of the tree. For the 
very reason that natural death is the result of a determinate conclusion of the de- 
velopment, those shoots (vegetable individuals) which have no such conclusion fre- 
quently undergo no death at all except that of some of their parts : but this is a con- 
comitant of animal life itself, (casting the skin, moulting, and the organic changes in 
the body). Cf. on this point Roeper: Linnrca, 1826, p. 439, and the following remarks 
on Paris, Lysimachia nummularia, Adoxa, etc., and the preceding ones on Caulerpa, 
f JJe Candolle: Physiol. Veg. II, p. 988. 
\IAnk: ErinneruDgen an die grosse Linde bei Neustadt am Kocher (Flora, 1850, 
No. 8). 6 
The Vegetable Individual in its relation to Species. 
cial ones ; since they not only prop the branches, but conduct 
nourishment to them by a shorter road, as is actually found to be 
the case in Rhizopltora Mangle, in various species of figs, [Ban- 
yan, &c.], and other tropical trees, whose branches high in air 
send down strong roots into the earth. A similar example nearer 
home, though indeed on a much smaller scale, is found in the Ju- 
niperus iSabiua. Its branches, which spring from a low stem, 
curve down to the earth, strike numerous roots, and raise them- 
selves again, so that the comparatively feeble stem may carry a 
creeping crown of considerable extent, like a thick wood contin- 
ually spreading, and which may continue to flourish in its parts, 
even when the communication between the original supporter 
and nourisher of the whole colony and the succeeding new 
growths, which are constantly receding from it, has finally ceased. 
A remarkable specimen of this tree stands in the lloyal Botanical 
Garden at Schoneberg, which, if not as old as the Garden itself, 
which was laid out in 1679 under the great Elector, Frederic 
William, certainly dates as far back as Gleditsch’s time, and his 
directorship commenced in 1744. The main stem is not more than 
33 inches in circumference at eight inches above the ground, 
close under the place where the first branches originate ; the cen- 
tre-piece of the crown which belongs immediately to the stem, 
is only nine feet high, and has been dying off' during several 
years, while the maximum diameter, from S. W. to N. E., of the 
hundred-rooted crown, which has spread out over the ground by 
the declination of the branches, measures 35 feet; the entire cir- 
cumference of the crown, which amounts to about 100 feet, 
would be still more considerable if it had been permitted to spread 
on every side, and if the branches on the N. E. side had not been 
removed at an early day. 
What has just been said of trees admits of no doubt as regards 
perennial herbs (plantae redivivae) with subterranean creeping 
stems or stolons. Such plant-stocks as those of the well-known 
Paris, Anemone nemorosa, Convallaria majalis, Asperula odo- 
rata, are undoubtedly exposed to none but a casual death.* All 
plants which renew the cycle of vegetative life repeatedly and 
without any determinate limits to their existence, and which I 
would hence call anabiotic, cannot therefore be considered sim- 
ple individuals.! 
* The same relations of great unlimited age are found in polyps -which form 
stocks. Cf. Ehrenberg: Abh. d. Acad, for 1832, p. 382, 420 , where among others, 
stocks of Mseandrise and Faviaj are referred to, larger than a cord of wood—which 
may readily be supposed to have been seen by Pharaoh. 
f I pass over the further question, intimately connected with this subject, whether 
the composite plant-stock itself, with all its subordinate generations, with all its pos- 
sible divisions,—viz., the individual in the most comprehensive sense (in which Galle- 
sio conceived it) has not a determinate term of life, though not easy to be ascertained, 
on account of the narrow space of time accessible to our direct experience. The Vegetable Individual in its relation to Species. 
7 
At first sight the case seems to be different in the haplobiotic* 
plants, which terminate their existence at the end of the simple 
process of development, with the formation of flowers and fruit; 
and this they do whether they exist one year, as Adotiis cestivalis 
and auiurnnalis, Nigella, Papaver Rhoeas, Erigeron 
or for two years, as Oenothera and Verbascum, or for many years, j 
as Agave [Century-plant], the East Indian Corypha, and the Mex- 
ican Fourcroya.<§> which suddenly puts forth its flowers only after 
400 years of extremely slow growth, and ends its life with the 
formation of its first and long-deferred fruit. The development of 
these plants, when compared with that of the first mentioned ana- 
biotic plants, seems at first to comprise only one generation, and to 
depend upon the development of one individual. But here, too, a 
closer examination shows conditions incompatible with the nature 
of the simple plant (the individual). One constituent element in 
the idea of an individual is, that the parts of the organism are es- 
sentially connected] yet the stock of annuals themselves pre- 
sents a multitude of parts which bear no essential relation to the 
whole plant. This is true of a large part of the ramifications, of 
branches which may exist in one case and not in others, and 
* De Candolle calls anabiotic growths polycarpic and haplobiotic growths mono- 
carpic, terms which are useless from their ambiguity. With an equally inap- 
propriate choice of terms he divides the first (Phys. Veg., II, p. 73) into caulo- 
earpic and rhizocarpic, according as the stem which produces the fruit is perma- 
nent, or dies off down to the root; but the latter in fact never takes place in per- 
ennial growths; for in such cases the life of the plant-stock is preserved not by 
the mere root alone, but by a subterranean portion of the stem. It is one of the 
most remarkable confusions which a want of true biological ideas has engendered, 
that De Candolle should have regarded the simplest and most natural circumstance 
in the plant’s life,—its death after having attained the goal of its development,—as an 
unnatural and to some extent casual occurrence, as a kind of sickness comparable to 
the succumbing of the mother in childbed, which he accounts for by the rapacious- 
yiess of the flowers and seeds. Roper, however, in a note to his translation of the 
above work, justly remarks that there are annuals with double flowers which die off to 
the ground although they produce no seeds. We may convince ourselves beyond a 
doubt that the flowers, on the contrary, are much less rapacious than the vegetative 
parts of the plant, that they even shut themselves off from the afflux of too copious 
nourishment; for many plants develop vegetative branches close under the terminal 
flower, as e. g., Stellaria media, Datura, Mirabilis, etc. In such cases the flower- 
stalk, which cuts itself off from almost all farther afflux of nourishment, remains slen- 
der, while the portions of the stem directly beneath, and the branches which spring 
from it, gorged with succulent matter, enlarge more and more, and attain a most 
disproportionate size. 
f These plants, like other annuals which germinate in the autumn, are usually reck- 
oned among biennials ; but this is a mistake, for like our winter corn they are plantce 
annuce hiemales. So, too, many vernal plants, as Teesdalia, Erophila, Cardamine hir- 
suta, Spergula Morisonii, and many weeds of the winter corn, e. g., several species 
of tares, Bromus secalinns et aff. 
j C'orypha umbraculifera. Cf. Phcede, Hort. Mai., iii, pi. 1-12. This is also the 
case in the palm-genera Metroxylon and Eugeissona, according to Martius (Hist. 
Palm, I, p. 108). 
§ On Fourcroya longccva, cf. Zuccarini in the Roy. act. nat. cur., xvi, 2, p. 666 and 
pi. 48. 8 
The Vegetable Individual in its relation to Species. 
which are proved to be unessential by the plant’s losing no essen- 
tial function when deprived of them. For even when the plant 
does not produce them, it can fully consummate the object of its 
individual life ; it can produce flowers and fruit. A glance at the 
examples just now adduced, Nigella, Papciver Rhoeas, Adonis, 
etc., will make these statements obvious. The branches of these 
plants, each of which, like the stem, is crowned with flowers and 
fruit, are evidently only unessential repetitions of the simple 
plant, absolutely identical with the main stem, and hence to be 
ranked as equal to it in importance, i. e., equally to be viewed as 
particular individuals, and with as much reason as in Zoology we 
concede individuality to the branches of the coral-stock (polypi- 
dom), which are now universally acknowledged to be individuals, 
and which offer an analogy of decisive importance for ascertaining 
the nature of the branch in vegetables. In view of this analogy 
Ehrenberg regarded plants as aggregations of individuals.* 
We can now turn back, and apply what has been shown to be 
the case in the annual herb to the shrub and the tree, each of 
whose annual generations now appears more distinctly than before 
to be, in their peculiar connexion, not one individual, but a world 
of individuals developing in the same period of vegetation and 
upon the same stem. To this intent many of the early botanists 
have expressed themselves, as I stated in the Introduction. Thus, 
B. Batsch, e. g., says of branches, that they shoot forth from the 
stem “ as if they were so many plants rooted in it ;”j* and Goethe 
“ lateral branches may be regarded as particular plantlets which 
are rooted upon the maternal stem, just as this stem is upon the 
earth.” Among moderns, Unger, at the close of his investigations 
into dicotyledonous stems, says . . “ Buds and the branches 
they develop are individual plants, which live by preying upon 
the maternal stem.”<§> Similar expressions are used by Schlei-. 
* Abli. d. Akad., 1835, p. 247 “Hence a polyp-stock is a mass of ani- 
mals. We have no satisfactory comprehensive expression for our idea of a plant. 
What an individual is remains still unknown; most of them are evidently aggregates 
of individuals which may be compared with coral-stocks.” . . . The origin of coral- 
stocks is minutely described by Ehrenberg in the Abhandl. for 1832, where he makes 
the following remarks : “ The coral structure is neither a mere structure composed 
of many animals arbitrarily conjoined, as Ellis supposed; nor one single animal with 
many heads, or with simple furcations, as Cavolini maintained ; nor a vegetable stem 
with animal flowers, as Linnaeus expressed it; it is a body of families, a living tree 
of consanguinity, the single animals belonging to it, and continually developing upon 
the primary ancestor, are entirely isolated within themselves and capable of complete 
independence although unable to achieve it.” 
f Bot. fur Frauenzirnmer, p. 15-16. 
\ Versuch d.Metam. d. Pfl. zu erkldren, p. 59. The words “just as” in the passage 
quoted imply too much, and remind us of Du Petit-Thouars’ unfounded doctrine 
of the formation of the woody layers of the stem by the ‘ roots’ of the buds which 
penetrate it. 
§ Ueber d. Bau u. Wachstlium des Dicotyledonenstammes, p. 177. Here, too, 
“ preying” is too strong a term. The Vegetable Individual in its relation to Species. 
9 
den ;* they are most definite in Roper’s works.f Linnmus ex- 
pressed the same thought in the words u gemma} totidem lierba3.” 
And I am thus led to make a particular remark, which is intended 
at the same time to modify in some degree what I said before in 
relation to the annually renewed generations of trees. It is in- 
deed true that branches of trees and perennial herbs, especially in 
temperate climates, first appear as buds ; and in a more extended 
sense we call in general every young branch a bud, even if its 
parts are not, as they usually are, compactly arranged and folded 
together ; still, all buds are not the rudiments of branches. Lat- 
eral buds are the only ones from which branches originate, and 
therefore they alone are to be regarded as new lines of develop- 
ment,—as individuals. Terminal buds, on the contrary, are noth- 
ing but still undeveloped parts of the (relative) principal axis: 
they are mere continuations and augmentations of the individual 
already existing, and are not to be regarded as commencements 
of a new one.jj: Hence, only those trees which produce no ter- 
minal buds, as the Linden, Willow and Elm develop new indi- 
viduals and nothing else at each renewal of vegetation ; while, 
on the contrary, those which do produce terminal buds also, as 
for example the Oak and Poplar, bear a mixed annual generation, 
which consists partly of new individuals, partly of old ones re- 
awakening and continuing their development with renewed vigor. 
I have already remarked how unessential the presence of 
branches is in many plants. A comparison of stocks grown on a 
rich soil with those of a poor one shows what license is given to 
plants in regard to producing branches, and how different the ap- 
pearance of specimens of the same species thus becomes. Plants 
grown on a poor soil are often called dwarfs ; but unjustly, for 
they present the most normal development of all essential parts, 
dispensing with everything that is unessential, and are much less 
* Grundz., ii, p. 4. “ New identical individuals develop upon the maternal stem 
by continuing the growth,” etc. Here the expression “ continuing the growth,” is 
improper, for the shoot does not “continue” the growth at all, but is a new growth 
fro m a new rudiment. 
f “ Omnis gemma solitaria aut ejusdem continuatio immediata et perpendicularis 
(caulis, ramus, ramulus, flos) individuum vegetabile vocatur.” This is the most defi- 
nite description I know of ; for in this passage not only the branches so-called, but 
also every arbitrary shoot, even when it is merely a flower, is acknowledged to be a 
particular individual. Besides what I have stated in the text in regard to the ap- 
pearance of terminal buds, I have only to remark, against the word “ gemma,” that 
in its growth every shoot does not enjoy a perceptible state of gemmation, i. e., a 
state of rest in which its parts are folded together. The term bad is applicable to 
but one state of a shoot or of its parts, and therefore cannot be a suitable expression 
for what is to be regarded as the vegetable individual. 
\ Kutzing, {Phil. Bot., ii, p. 146,) aptly expresses these relations by calling the 
terminal bud the continuation of the “ series of formations,” lateral buds begin- 
nings of a new “ series of generations.” In contradiction with these terms, however, 
he calls the bud an “ organ” as long as it is connected with the natural individual— 
a term inapplicable to the bud as it is to the developed branch, of which it is the 
adolescent state. 10 
The Vegetable Individual in its relation to Species. 
inclined to malformations than the lusty giants of the rich soils. 
Not unfrequently we find diminutive speciments of Eryihrcea 
pulchella s. ramosissima which are branchless and perfectly sim- 
ple, as they terminate with a flower after four or five pairs of 
leaves. More vigorous specimens produce two branches out of 
the axils of the highest pair of leaves, which after a single pair of 
leaves terminate in the same manner with a flower ; and branches 
of the second order may be also emitted from the axils of the two 
leaves preceding this flower; and so on. In the first order of ram- 
ification the number of flowers amounts to three, in the second 
to seven, and so on ; in the seventh, which is not unfrequently at- 
tained, it amounts to 127 ! Here, if we would consider the stock 
or specimen as the individual, and the flower as the superior 
termination of the vegetable organism, comparable, say, to the 
head of the animal, this variation in the number of the flowers 
would be as astounding as if we were to learn that an animal 
might have 3, 7, 15, 31, 73, or 127 heads, according to circum- 
stances. The same thing occurs in Radiola, linoldes. Erigeron 
Canadensis, which often grows to the height of a man and bears 
as many branches as a tree, presents dwarfed specimens scarcely 
two inches high and of a perfectly simple form.* After devel- 
oping two early deciduous cotyledons it presents about 13 leaves 
on the stem, which are followed by a terminal capitulum of 21 
involucral bracts and about 34 flowers. One middle-sized speci- 
men about three feet high presented nearly 100 branches of the 
first order, out of which branches of the succeeding orders pro- 
ceeded, together bearing about 2000 heads, and hence (reckoning 
the head at 34 flowers) 6S000 flowers.f 
I may here remark, that such unessential branches maybe sep- 
arated and reared independent of their parent stem ; on which fact 
depends propagation by artificial divisions, which is so variously 
employed in horticulture. The most remarkable case of this artifi- 
cial division is recorded by Miller: in the year 1766-67, he ob- 
tained 500 stocks of winter Rye, by dividing one stock and re- 
peating the operation three times; these 500 stocks emitted 
21,109 spikes, bearing together 576,840 grains. Nature herself, 
as well as art, in various ways may effect such an independent 
separation of developed branches or of undeveloped buds, and 
this too either above or beneath the ground. Propagation of the 
Strawberry by its runners; of the Potatoe and the Helianthus 
tuberosus by their tubers ; of bulbous plants by their bulbs; of the 
* Not counting the florets, which also are properly so many branches, 
f Similar cases occur in most annuals. The forms of Bromus mollis and racemosus 
with simple spikelets instead of rich panicles, are well-known ; less known and less 
remarkable are the depauperate specimens of Umbelliferce with one single unifloral 
umbel, some of which of Scandix Pecten are in my possession. I have also speci- 
mens of Solanum nigrum, one and a half inches high, with a solitary terminal flower. The Vegetable Individual in its relation to Species. 
11 
Garlic by the bulblets formed in the process of flowering, and fall- 
ing off like seeds ; of the varieties of the beautiful Achimenes 
by the amentaceous or the strobiliaceous deciduous shootlets, are 
well-known examples of this process; and thousands of others 
might be adduced.* 
The gardener can not only separate individuals, but unite them 
upon one stem. This is true not only of individuals of the same 
species, but even of those of different species ; sometimes even 
of different genera of the same family. The Lilac is not unfre- 
quently grafted upon the Privet (Ligustrum), the Pear upon 
the Mountain Ash (Soj-bus Aucuparia), ihe Peach upon the Al- 
mond. Ely the insertion of a bud (inoculation), or of a developed 
sproulet (grafting), we are thus enabled to pluck different kinds 
of roses from the same bush, to gather different kinds of fruit 
from the same tree. It would evidently be a contradiction in this 
case to consider the whole tree, or the whole bush, as the indi- 
vidual ; for we should then give the name to a compound of sev- 
eral species, or even of several genera. 
In attempting to comprehend the vegetable individual in its 
simplest form, we have thus far spoken of unessential branches 
only, and have endeavored to show that they cannot be regarded 
as mere parts of the individual. But there is another kind of 
branches, those which are essentially requisite for the attainment 
of the end of vegetation,—for the formation of flowers and fruit. 
These occur in all plants which possess no terminal buds, and 
which must hence necessarily have some branches in order to 
attain the end of their existence. This is the case with the Eve- 
ning-Primrose, Larkspur, Orchid.ece,etc., whose lateral flowers are 
just such essential branches. If we demand that the individual 
should be a complete representative of the characters of the spe- 
cies, as is implied in the usual view, then we must add to the 
principal axis such branches as these,—without which the process 
of vegetation is not concluded, and on which, in fact, the most 
essential and characteristic parts of the plant make their appear- 
* I will only adduce a few more of these examples, which might be multiplied 
indefinitely. Besides the Garlic (Allium sativum) in many other species of Allium. 
e. g., A. olcraceum, carinatumj vineale, Lilium bulbiferum, tigrinum, humile, and 
other species, Gagea fstulosa, Ficaria ranunculoides, Dentaria bulbifera, Saxi.fraga 
bulbifera and cernua, Cicuta bulbifera, Polygonum viviparum, Begonia bulbifera, 
diversifolia and other species, Remusatia vivipara, Cystopteris bulbifera—buds fall off 
above the ground (as bulblets). In Stratiotes aloides rosette-like developed axillary 
shoots separate close to the base. The separation of lateral shoots in Lemna is well- 
known ; and it occurs in a similar manner in Pistia, by the separation of thin-stalked 
lateral rosettes, and in Hydrocharis in the separation of peculiar winter buds. When 
the inferior leaf-formation is gorged with sap, bulblet-like buds form from the axils 
of the root-leaves (frondes fundi) in Saxifraga granulata and many exotic spe- 
cies of Oxalis, in the same way as the bulb-brood of monocotyledonous bulbous 
plants. Inferior leaf buds which are placed on the ends of their stolons become free 
by the death of the runners in Epilobiumpalustre, Lycopus Virginicus, etc., and swell 
out and form little lumps. Cf. on this subject Wydler (Flora, 1853, p. II—24). 12 
The Vegetable Individual in its relation to Species. 
ance,—and call these parts of the same individual. In this sense 
Schleiden’s view of the simple plant might perhaps be justified, 
although, as he starts from different premises, he does not con- 
sider mere floral branches as particular individuals. He says : “ If 
nothing but organs of reproduction, or flowers, spring from the 
bud we still call the plant a simple one.”* 
Here, however, we arrive at a contradiction, which shows us 
that we cannot carry out the idea of the vegetable individual with 
the requisite definiteness in this way, since we thus regard essen- 
tially similar branches, now as individuals in themselves, now as 
mere parts of individuals. As I have already remarked, Schlei- 
den allows individual importance to branches which are identicalf 
with the main axis ; those on the contrary which produce flow- 
ers alone, and in this respect differ from the main axis, he regards 
as mere parts of the simple individual. This distinction when 
analyzed is perfectly nugatory ; since it only lays down two ex- 
tremes, between which there are an infinite number of gradations. 
Strictly speaking, there are no branches which are perfectly iden- 
tical with the main stem, as is evident from the fact that no 
branch begins with cotyledons, as the main axis does.j: Besides, 
the foliaceous leaves on the branch are ajmost always fewer than 
those on the main axis, and generally fewer in proportion as the 
point is higher where the branch originates. The arrangement 
of the leaves on the branches, also, often differs from the arrange- 
ment on the main axis, as e. g., in most of our broad-leaved trees, 
—in the Elm, Hazle, Chestnut, Linden, etc., in which the phyl- 
lotaxis on the main axis, and often at a later period in the so- 
called ‘‘water-shoots” (Wasserschossen), is spiral or decussate, 
while on the branches, it is, on the contrary, distichous. In Al- 
nns viridis the phyllotaxis is tristichous on the main axis, and dis- 
tichous on the branches. On the main axis of Cypresses and Thuja 
there are 3-4-leaved whorls; on the branches the pairs of leaves 
are nearly decussate; this is also the case in Lysimachia vulgaris. 
In the same way in Equiselum the number of the rameal verti- 
cillate leaves is always inferior to that of the cauline ones. While 
thus on the one hand the vegetative branches are nowhere en- 
tirely similar to the stem from which they spring, on the other 
hand it appears that those branchlets which seem to bear flow- 
ers only are usually more numerous than they seem to be ; since 
in most cases one, two, or even more small leaves (bractlets), are 
present beneath the flower, which may easily escape notice on 
account of their diminutive size, although their existence may 
* Qrundz., ii, p. 4. f Grundz., ii, p. 4. 
\ The basillary cotyledons of the branches, indeed, have been compared to cotyl- 
edons. This comparison is partly justified |in view of the commencement of phyllo- 
taxis on the branch ; which often resembles that on the main axis, whilein regard to 
form and consistency almost all resemblance disappears. The Vegetable Individual in its relation to Species. 
13 
be often ascertained with certainty even in those cases in which 
they are not visible when the flower has reached its complete de- 
velopment.* If we are to deny individuality to those buds 
(branches) only which are composed of a flower alone, as a strict 
interpretation of Schleiden’s language demands, we should have 
to draw a most unnatural and often impracticable line of demar- 
cation between branches which, physiologically speaking, are per- 
fectly homologous (floral branchlets which really have no bracts), 
and those which bear imperceptible or even suppressed (abortive) 
bracts. If on the other hand we would reckon the latter also 
among the branches which are not individuals, then it may be 
contended that there is such a series of gradations in regard to 
number and vigor in the leaves which precede the rameal flower, 
that it is impossible to draw a dividing line even in this manner. 
The above-mentioned distinction between unessential and es- 
sential branches seems to afford a better stopping-place, no matter 
* In fact, all the constant lateral flowers of Primulacece, Cruciferce, Uapparidece, 
Resedacece, Balmminece, Orchidece, never have any bractlets. Among moUocotyle- 
vlonous plants in many cases there is only one bractlet; among the dicotyledonous 
there are generally two. Gesneriacece have generally three ; Empetrum and Santa- 
lum have four, Eriostemon five, Polemoniecece, Cuscutece and other plants with pani- 
cled inflorescence an indeterminate number. We possess the following means of 
showing the existence of suppressed bractlets : 1. The position of the parts of the 
flower relatively to the axis of origination from which the lateral flowers spring. 2. 
Analogy. 3. The study of malformations. 4. Observations of the flower’s devel- 
opment. The first criterion can be applied only where we can determine the succes- 
sion of the parts of the flower. The position of the parts of a lateral flower depends 
in fact upon determinate laws of rameal origination; when they do not harmonize 
with these laws we must conclude that preceding leaves have been suppressed. In 
this way, e.g., we can explain the very common position in the 2-5th arrangement 
of the calyx with the second sepal posterior, by supposing two bractlets according 
to the fixed law, while it cannot be explained without these bractlets. Analogy aids 
us most by confirming our conclusions, as e. g., in the families Scrophularinece, Labi* 
atcce, etc., in which many genera present distinct bractlets, while others appear to be 
without them. In monstrous flowers (in cases of (ant holy sis and chlorysis), sometimes 
without any other malformation, bractlets otherwise imperceptible appear in an ab- 
normal growth. Not unfrequently in Digitalis purpurea, which in its normal state 
presents no bractlets, but in which we inferred their original existence from aestiva- 
tion and the position of the calyx relatively to the axis, I have found bractlets de- 
veloped in the most heteregeneous degrees, especially on the lowest flowers of the 
raceme of cultivated specimens. C. Schimper and myself have both observed the 
same fact in Tropceolum majus, which, like most species of this genus, presents no 
trace of bractlets in the normal state. We have seen them in the form of very 
small, white, subulate leaves about in the middle of the flower-stalk, while the flower 
remained unchanged in all other respects. Their existence, however, was already in- 
dicated by the position of the quincuncial calyx relatively to the axis, as well as 
confirmed by analogy, for Tropceolum ciliatum E. et. P. (Popp, et Endl. Nov. Gen. 
t. 38) in its normal development has two round and prettily ciliated bractlets on the 
the flower-stalk. I have mentioned the history of development last, not to disparage 
study, but because the morphology must be rightly understood beforehand by means 
of comparisons of developed structures, and because in its present stage the devel- 
opment is incapable of giving us reliable information in regard to all the leaves which 
are present in the germ, though they may not develop. To know what parts then 
exist we should have to be able to distinguish the leaf as a cell or a group of cells 
before it rises to view above the surface of the stem. 14 
The Vegetable Individual in its relation to Species. 
whether the branch hears nothing but a flower or not. We might 
say, all essential branches must be regarded as individuals since 
they repeat the process of specific development laterally, and can 
become independent plants, as layers, whether natural or artificial,, 
Those branches, on the contrary, which appear as necessary mem- 
bers in the line of development which is advancing towards flower 
and fruit, and which therefore complete the series of formations 
belonging to the species, and without which the plant is either 
unable to eke out its vegetable life or to accomplish propagation, 
must be regarded as members of one and the same history of de- 
velopment. Let us take a case where the main stem bears only 
proper leaves, branches of the first order only bracts, and those 
of the second order only flowers and fruit, as is really the case 
in Pla?itago, Melilotus, Veronica officinalis and Chamcedrys ; 
here it is evident that these three divisions cannot be isolated ; 
that all three must necessarily be present in order that the specific 
life may attain a complete representation in one individual.* 
Notwithstanding the importance of this discrimination between 
essential and unessential branches, it cannot, when analyzed, es- 
tablish a distinction which will enable us to decide upon their im- 
portance as individuals ; for even those branches which appear 
unessential, in relation to the formation of flowers and fruit, may 
yet be essential to the plant in other relations : as when they ap- 
pear as characteristic elements of the vegetable structure, or when 
they play any important part in the economy of the plant, as I 
have shown in extenso elsewhere.f Nay, more ; one and the 
same branch, as to whose nature there seems to be no doubt, may 
appear either as essential or as unessential, according to circum- 
stances. When those branches which conduct the structure to a 
higher stage of its development appear in great numbers on a 
principal axis, as e. g., in indefinite racemose or spicate inflores- 
cence, the lateral branchlets appearing as flowers are then in- 
deed, generally speaking, necessary to the plant’s full completion 
of the series of formations, and in this sense essential ; but their 
number is immaterial as regards this completion ; and this the 
plant itself shows in producing either a larger or a smaller num- 
ber of them ; sometimes the number is reduced to one.J There- 
fore, properly speaking, only one lateral flower is essential ; and 
we may arbitrarily consider any one of the number to be this es- 
sential one. Hence each of them may be regarded indifferently 
as essential or unessential. This is not the case in those racemes 
and spikes which possess a terminal flower, as is the case in many 
* [But why assume (as here and supra) that the species must attain a complete 
representation in a single individual in vegetables ?—since this is by no means the case 
in the higher (unisexual) animals, where there is no doubt as to what corporeally con- 
stitutes the individual,—that is, in the very cases whence we derive our idea of in- 
dividuality, and the standard of comparison which our author is endeavoring to apply 
to the case of plants, a. g.] 
| v. Yerjiingung, p. 41, et. seq. 
\ E. g. not unfrcquently in the raceme of Lalhyrus odoratus. The Vegetable Individual in its relation to Species. 
15 
Campanulacce, e. g. in Campanula rapunculoides. Here all 
the lateral flowers are unessential: yet if the terminal flower is 
cut off, the lateral branchlets which bear the flowers at once be- 
come essential. Such a change is not always artificial, for it often 
happens naturally, as there are plants in which the terminal flower 
may be either present or absent. Agrimonia Enpatoria, and 
Campanula rapunculoides, are examples of this variability.* 
We can cut this Gordian knot only by deciding to consider every 
branch as an individual, however appearances may be against 
it, provided that we have other grounds sufficient to regard 
branches as individuals. The genesis of branches justifies us in 
so doing ; for each branch is not a direct continuation laterally, is 
not a development belonging to the stem (like the leaf), but is a 
new formation ; like the main axis itself, it has its own centre of 
formation, with its peculiar development. Branch and stem, main 
axis and lateral axis, differ therefore only in their origin and rela- 
tive position ; but they are essentially of the same nature ; they 
are united in the idea of the shoot. The stem is the primary 
and principal shoot of the whole plant, the branch is a lateral 
shoot in reference to the main shoot; but it can itself become a 
relatively main shoot, and the stem of a succeeding generation of 
shoots in its turn. As far, then, as we are justified in speaking 
of vegetable individuality at all, we must hold fast to the indi- 
viduality of the shoot: the shoot is the morphological vegetable 
individual; is that form or that part of its specific realization 
which is analogous to the animal individual, if any part is. 
In zoology we give the name of individual to every whole 
which is controlled and bound together from one vital centre. 
Since such an internal domination of the organism as that which 
characterises animal life is wanting in plants, whose existence is 
a process of growth directed externally alone, we can only de- 
mand, as the criterion of vegetable individuality, that the indi- 
vidual shall be formed in direct continued development from one 
centre, and thus in accordance with its origin, shall, in all its 
parts, belong to one centre. Now this is the character of the 
shoot. Its centre of formation has been known since C. F. 
Wolff’s celebrated “ Theoria Generationis” (1759) under the 
name of “punctum vegetationis it is about what is called in 
* Agrimonia Enpatoria bears usually one spike without any terminal flower; in 
weak specimens, a terminal flower not unfreqnently makes its appearance which 
opens before the upper lateral flowers. This has been observed by Wydler (Bot. 
Zeit. 1844, p. 642). In Campanula rapunculoides the case is just the contrary : its 
looser spikes are usually terminated with a flower, while denser ones end in a coma 
of bracteal leaves, without any terminal flower. Dictamnus resembles Agrimo- 
nia; while Triglochin (especially Tr. maritimum) on the other hand imitates Cam- 
panula. Even in plants in which the essentiality of the lateral position of the 
flower is expressed by their zygoraorphic development, terminal flowers make their 
appearance in some cases; they then resemble Pelorioe. This is the case in Linaria, 
Orobanche, and a Digitalis purpurea monstrosa (described by Yrolik, Flora, 1844, Ho. 
1), which propagates by seeds, and is now widely disseminated in our gardens. 16 
The Vegetable Individual in its relation to Species. 
common life the “ heart,” of the plant, or, at the first appearance 
of the lateral shoot, the “ eye.” The whole future of the plant 
slumbers unseen within it; leaf after leaf arises out of it, step by 
step, at a measured pace, prescribed by law, until (in case the shoot 
is destined to conduct the development thus far) the series con- 
cludes with the last formation, that of the carpels, which close 
over the dying point of vegetation and form the fruit. In this 
progress the centre, always keeping the lead, is ever advancing, 
rising more and more, and leaving behind it an axis arrayed with 
the organs already formed. Hence we may designate the vegeta- 
ble individual as the sum of the parts belonging to one axis. Just 
as the body of the animal has only one trunk and one head, the 
shoot has but one axis and one apex. As the trunk of the ani- 
mal has a second extremity opposite to the terminating head, and 
gradually dwindling down till it forms the tail, so the perfect 
shoot has a second extremity opposite to that which terminates 
with the most perfect structure (the fruit), and dwindling down 
to an indeterminate end, the root, by means of a punctum vege- 
tationis turned downward.* 
But it will be objected : is not the vegetable shoot indefinitely 
divisible, can we not cut it up into an arbitrary number of pieces, 
each of which is capable of reproducing the whole plant in 
its turn? Were this the case the phenomenon would not be 
without its parallel among the lower animals. But this is not the 
case. The supposed divisibility of the vegetable shoot, at least 
in perfect plants (the Phanerogamia) to which I am now alluding, 
is a delusion, which rests simply upon the fact that the formation 
of new shoots has been confounded with a reproduction of the 
shoot as such. As the injured shoot has the faculty of producing 
new shoots, so the parts of the divided shoot have also this faculty 
in many cases; but this is no re-completion of the shoot itself; 
the fragment of the old shoot can continue to develop in one sin- 
gle case only : when, in fact, it bears the apex of the axis with 
the point of vegetation. Let us examine this case more closely. 
If a shoot is divided transversely, under certain circumstances the 
upper part, on which the punctum vegetatioiiis (“ the heart”) is 
* Aristotle, on the contrary, considered that the root, being the imbibing organ, 
was the part of the plant which corresponds to the upper part, to the head and 
mouth of the animal; and he regarded the stem as the inferior part. He found the 
cause of this topsy-turvy position of plants in the necessity under which they labor 
of drawing their nourishment from the earth, as they are incapable of moving from 
place to place. In this respect he compares plants to muscles ((6<maxo5fjua). 
which also have their heads turned downwards. Cf. Wimmer: Pliyt. iris. Frag. 
56-65. This comparison of the root with the animal’s head is however, morpholog- 
ically speaking, inverted; for as the highest stratum of the spinal chord (the sen- 
sorial portion) attains its maximum state of development in the head of animal?, it 
can only be compared to that extremity of the plant’s axis in which the highest and 
noblest part of the plant is exhibited. Besides, the peculiar and striking character- 
istic of the animal’s head, its involved structure terminating the organism, is by no 
means to bo found in the root end of the plant; but it is seen in the opposite end 
which terminates with flower and fruit. The Vegetable Individual in its relation to Species. 
17 
still remaining, may continue the development; but the lower 
part is nothing but a stump, aud continues to be a stump which 
can never complete itself by a terminal shoot, and which never 
fails to die if it is not nourished by lateral sprouts formed before, 
or sometimes after, the division took place, and thus kept alive by 
its posterity. This cannot be called divisibility, in the usual 
meaning of the term ; the whole phenomenon, on the contrary, 
strongly reminds us of the capacity animals possess of losing the 
less essential caudal extremity without any cessation of life. In 
favor of this view the fact may be adduced that a similar phe- 
nomenon occurs in the normal process of development of plants 
and animals. As there are animals which may spontaneously 
lose the posterior extremity of their body during the course of 
their development, as e. g., Cercaria, Comatula, frogs, etc., so 
there are also numerous plants in which the posterior extremity 
gradually dies off, and is cast aside, during the course of growth, 
while the anterior end of the shoot, which bears the punctum veg- 
etationis continues to unfold; as is seen in the growth of many 
mosses, especially of Peat-mosses, in the creeping and climbing 
rootstocks of Ferns and Aroidece, in the long creeping stems of 
Lysimachia nummularia, the little subterranean creeping root- 
stocks of Paris, in most plants which possess a radix prcemorsci, 
as e. g., Siiccisa pratensis, the perennial species of Plantago, in 
Tormentilla, etc., with which the perennial bulbs of monocoty- 
ledonous plants agree in all essential respects; and finally, this is 
especially remarkable in Utricularia, and in Selaginella incres- 
centifolia, whose apices only form close buds, and last through 
the winter, while all the remaining parts of the shoots perish. If 
the shoot is indivisible transversely, it is still less so longitudin- 
ally. There is not a single case to prove that a shoot longitu- 
dinally divided can as such continue to develop ; nor do we know 
of a single case where such a longitudinal division takes place 
spontaneously. What has been usually described as a bifurca- 
tion of the stalk depends in the Phanerogamia in every case upon 
a true ramification which takes its rise laterally close under the 
apex, as I have already described it in the case of Erythrcea pul- 
chella. As a normal formation no immediate division of the 
stalk occurs among Phanerogamia; for the phenomenon known 
as “ fasciation,” which might be adduced here, is always a mon- 
strosity.* The stalk, or axis of the shoot, is hence indivisible in 
* Fasciation depends upon a real division of the punctum vegetationis into two 
parts of equal importance ; in the simplest case it produces a simple division into 
two parts. Here neither of the two parts can be regarded as a branch of the other. 
If repeated bifurcations follow each other in the same plane, and in unbroken con- 
nexion the well known “ribbon and fan” like forms arise which however usually end 
at last in single apices. Very rarely more than two parts lying in different planes 
are produced by the division of the punctum vegetationis, a case which I have no- 
ticed in the capitula of Compositae. The rarest phenomenon which bears upon our sub- 
ject is the annular fasciation, in which an annular border arises from the simple 
point of vegetation, of which I shall speak more at large in the following Part, when 18 
The Vegetable Individual in its relation to Species. 
the higher plants, in the same sense that the body of the higher 
animals is indivisible.* The only phenomenon which might be 
described as a division of the stalk is leaf-formation. This, how- 
ever, is not a division into new stalks, but a formation of subor- 
dinate parts belonging essentially to the stalk, as it were an eradi- 
ation of the stalk itself, which may be aptly compared to the for- 
mation of the extremities in the animal body. We may there- 
fore justly describe the shoot, or the vegetable individual, as an 
indivisible axis,—as an axis with its appertinent radii which are 
inseparable from, and regularly arranged by, its own development. 
With the first appearance of the branch a new axis is formed, and 
a new system of subordinate radii appears. However completely 
the branch may contrive to interweave itself with the trunk dur- 
ing the course of its development, it always owes its origin to an 
accessory point of vegetation which develops into a particular 
axis. The vegetable individual thus presents in its nature a cer- 
tain analogy to the mineral individual,—the crystal,—as well as 
to the animal individual; for the crystal is determined by the re- 
lation of its parts to one and the same system of axes. As soon 
as this system of axes holds another position there results another 
individual, which may be distinguished even when two or more 
individual crystals intersect, so as to form twin crystals, or stel- 
late crystals. 
In the preceding considerations on the indivisibility of the axis 
I described the leaves as its radiations,—as members of the 
stalk, and belonging essentially to it,—and I attempted to distin- 
guish the leaves from the branches, by considering the latter as 
new axes. But how are leaves and branches distinguished in 
their genesis ? Are not the branches as much radiations or lat- 
eral members of the stalk as the leaves? It would lead me too 
far from my subject to make a fundamental critical investigation 
into this question, and to examine the existing views of the mode 
of formation of leaves and branches, especially as investigations 
into this subject have not been complete enough to enable us to 
obtain reliable results. 1 can therefore only allow myself a few 
hints in this place. The leaf originates in the earliest period of 
the formation of the stalk ; and its rudiment is contemporaneous 
with the first stages of the formation of tissue in the punctum 
I compare the relations of growth in the Cryptogamia. A division of the individual 
corresponding to fasciation in phanerogams and to dicthotomy, its homologue, in many 
cryptogams also occurs in the animal kingdom, as appears especially in many genera 
of corals, e. g. Caryophylliece whose stocks are formed in this manner exclusively, and 
in Antrim and Favice in which it appears in conjunction with shoot-formation (gem- 
mation), as was shown by Ehrenberg, (Beitriige, etc., Abb. der Acad., 1832, p. 242). 
Ehrenberg explains the form of Bccdalince as a result of incomplete termination of the 
individuals in gemmation; in appearance it resembles the coxcomb-like forms of fas- 
ciation as they occur in a remarkable way in some monstrous Cacti of the genera 
MammilUiria and Fchinocactus, as well as in Celonia cristata, well-known as an orna- 
mental plant. 
* [Some criticisms upon this may be given at the close of the whole memoir, a. g.] The Vegetable Individual in its relation to Species. 
19 
vegetationis. A leaf can never be formed at a later period from 
the developed axis. It is a necessary consequence of the manner 
in which the leaf originates, that an absolute dividing line cannot 
be drawn between leaf and axis ; for the subsequent position of 
the leaves upon the organism affords no standard of appreciation, 
especially as most of them do not mark the basis of the leaf, which 
loses itself in the axis. Earlier, before the extension of the axis 
begins, the rudiments of the leaves are always closely pressed to- 
gether, so that they appear as a peripherical development of the 
axis itself, occupying the whole upper surface, and dividing it 
into clearly defined planes, which maybe recognised even in the 
developed state, in those plants whose foliaceous pulvini are dis- 
tinctly marked, as e. g. in many Ferns, most acerose plants, in 
Cacti, and particularly in Nymphcea and Victoria where the pul- 
vini may be distinguished even in the interior of the axis. The 
primitive vascular system of the axis enters directly into the 
leaves, and ramifies there ; while the woody layers of the stem, 
which are found later, have no connexion with the leaves. With 
branches the case is totally different. In their origin and devel- 
opment they always succeed the leaves ; and even at a much later 
period, when the leaves have been long cast off, shoots may ori- 
ginate in places where, at an earlier period, no trace of a rameal 
rudiment, or of an eye, was to be found. If we now consider 
the axillary shoots,—i. e. those branches whose position is pre- 
determined by the situation of the leaves,—at an early period we 
shall find their rudiments, even though they develop very late or 
not at all, in the form of a circular and slightly prominent gibbos- 
ity, which may be compared with the apex of the axis; or rather, 
it is an accessory punctum vegetationis forming near the apex. 
The circumstance of the epidermis of the axillary shoot’s being 
a continuation of that of the stem, is explained by the early date 
at which it originates ; for this takes place at a time when the 
surface of the axis has not yet lost its flexibility. The eye is 
shown to be an independent centre of vegetation by its subse- 
quent internal and external conformation ; for it not only devel- 
ops leaves upon its surface, and this too with an independent com- 
mencement of its phyllotaxis, but even in its interior the first sys- 
tem of vascular fibres seems to be formed independently of that 
of the main axis ; as originally it lies upon it, and afterwards be- 
comes intimately blended with it by later layers of tissue. Not- 
withstanding the intimacy with which later formations of woody 
tissue unite branch and stem, still, according1 to Unger’s investi- 
gations, no immediate influence is exerted by the branch upon 
the conformation of the stem, since the stem owes none of its es- 
sential parts to the branches.* This independence of the branches 
is shown still more decisively in adventitious shoots, whose posi- 
* Unger : Ueber den Bau dcs Dicotyledonen-Stammes (1840), p. 65, et 66. 20 
The Vegetable Individual in its relation to Species. 
tion is not predetermined by the leaves. Originating at a later 
period, they take their rise not from the surface but from the cam- 
bium layer,—the internal tissue which preserves the faculty of 
producing new growths. Hence if they would come to the light 
of day they must break through the bark. Their origin has been 
particularly described by Trecul.* W. Hofmeister, however, as 
1 have already remarked, succeeded in tracing it in Equisetum 
back to the first cell, a cell in the interior of the stem. As is the 
case with axillary buds, such adventitious buds sometimes remain 
undeveloped for a long time (ten years and more) without losing 
their vital activity ; a fact to which attention has lately been 
called by C. Schimper.f in a report on exostoses. When this is 
the case they not unfrequently develop into spherical or conical 
wood-kernels, which continue to exist without any connexion 
with the ligneous body of the maternal stem; this is especially the 
case in Beeches and Poplars. 
The individual nature of the shoot is confirmed not only by 
the mode, but by the place, of its origin. While the organs of 
the individual organism,—the leaves of the plant,—occupy a posi- 
tion determined with geometrical accuracy, shoots on the con- 
trary can arise out ot almost any part of the plant, wherever in- 
deed any cambium exists; and they may be even enticed by art 
out of places where they do not usually appear. There are 
shoots from the stem, the root and the leaves. In herbaceous 
stems they appear in situations determined by the leaves (in the 
axils of the leaves), while they may be found anywhere on 
old woody as adventitious buds, or on any part of the lig- 
nified roots of most dicotyledonous woody growths, and even 
on some monocotyledonous ones, as in vmbraculifercE.§ Shoots 
appear less frequently on the roots of herbaceous plants. || Shoot- 
formation from leaves has often been discussed and described in 
regard to many plants, especially Bryophyllvni, Cardamine pra- 
tensis, Drosera, Malaxis paludosa, etc. A fine example of this 
is shown by a Chelidonium majus var. laciniatum reared by 
Bernhardi in the Botanical Garden at Erfurt, from whose leaves 
* Trecul: Recherches sur I’orig. des bourg. adv. Ann. des sc. nat., viii, (1847) p. 
268. 
f In Sept., 1852, in the Yersammlung der Naturforscher in Wiesbaden. 
\ Rarely scattered shoots appear on the herbaceous stem, and especially on the first 
internode under the cotyledons, as Roper (Enum. Euphorb. 1824) first showed in Eu- 
phorbia, and Bernhardi in the germ of Linarice. A specimen of Begonia manicata, 
dipetala cultivated in our [Berlin] Botanical Garden, which is probably the same 
species as the B. phyllomaniaca of Martius, presents the case of a plant which pro- 
duces a multitude of shootlets in the whole leaf-region; they arise from the sappy 
stem which is not yet hardened, soon after the fall of the leaves. 
§ According to Rheede, Corpypha urnbraculifera sends forth root-shoots when the 
stem dies off after the fruit has ripened. 
| I have often observed them in Linaria vulgaris, llelichrymm arenarium, Rum ex 
Acetosella, Ajuga Genevcnsis, Jurinea Bollichii, Nasturtium sylvestre et Pyrcnaimm. 
According to Wydler, they often appear in Viola sylvatica. The Vegetable Individual in its relation to Species. 
21 
floral bractlets arose, partly unifloral, partly multifloral, without 
any preceding leaves.* Shoots may be allured by the gardener 
out of most leaves which do not wither too soon.f Finally, 
the little budlets in whose bosom the germ of the new plant is 
formed and developed, and which we call seeds, are a kind of 
shoots, which in most cases owe their origin to leaves, (carpels) 
out of which they spring (on the margins, which unite to form 
the placenta) or more rarely, out of their whole inner surface. 
While thus, on the one hand, all the facts seem to unite in 
establishing the individual nature of the shoot, on comparing 
shoots in their qualitative relations, phenomena are brought to 
view which seem to contradict such a view of its individuality. 
The higher departments of the animal kingdom usually present 
as individuals representatives of the specific type agreeing in all 
essential respects, though, perhaps, not perfectly identical. The 
fact of the separation of the sexes was all that modified this 
view; and here, indeed, the essence of the species does seem to 
be divided between two different individuals. Attempts have 
not been wanting to obviate this contradiction by the Platonic 
doctrine of the original unity of the sexes, by the assertion of 
Paracelsus that, in fact, the two together must be regarded as 
the one real individual,—and such like. 
This contradiction to the usual view of what constitutes the 
individual is shown in a far higher degree by qualitative compari- 
sons of vegetable shoots, not merely of the same species, but also 
of the same stock. Thus we see, e. g., in Equisetum arvense 
(Field-Horsetail) shoots totally diflerent in aspect proceeding from 
the same root-stock; in early spring they are pale, discolored, 
unbranched, terminating with a strobilaceous-like fructification; 
later, green and foliaceous ones appear, verticilfately ramified. In- 
vestigations into subterranean vegetation show even other varie- 
ties of shoot-formation, viz., offsets dwindling down to a point, 
and club-shaped buds which, at a later period, drop off of them- 
selves. The Colt’s-foot (Tussilago Farfara) presents similar 
phenomena, in early spring putting forth leafless shoots, with as- 
paragus-like scales terminating with yellow capitula, which in 
summer are followed by others bearing leaves. The flowers in 
the little capitula of the first present a third variety of shoots in 
their lateral branchlets. Even in common life we distinguish 
* I may add to the examples I have given of shoot-formation taking place out of 
the leaves, one which I observed in June, 1853, in Levisticum officinale. I found, in 
fact, in several species of this Umbellifera, one or more, frequently two, shoots in 
the points of division of the leaves, which, after producing a few weak leaves bore 
a small umbel. (Later note.) 
f Kirschleger (Flora 1844, No. 2) notices a line example of this in Gloxinia spe- 
ciosa. 
\ “For this ye must know: man without woman is not a whole; only with wo- 
man is he a' whole. That is as much as to say: both together make man, and nei- 
ther alone.” 22 
The Vegetable Individual in its relation to Species. 
leaf-buds from flower-buds, on many trees. Let us consider this 
relation in the Cherry tree, for example. On the same branch 
we find, on the one hand, buds which develop into branches 
bearing leaves, without producing flowers; on the other hand 
some bearing only little squamate leaves on the shortened axis, 
from whose axils the flowers rise and form a third kind of 
shoot. 
On examining closer into the real origin of these differences, 
we find their ground to be a partition of the different steps of the 
metamorphosis (of the formations) among different shoots. True 
there are many plants which go through the whole series of form- 
ations, from the inferior* and the foliaceons formations up to 
flower and fruit; but the cases are quite numerous in which this 
does not take place, in which the single shoot is not able to pro- 
duce all the formations. Thus there are shoots which are only 
able to realize the lower steps, and never attain to flowers and 
fruit; while others overleap all the inferior degrees and com- 
mence immediately with the formation of flowers. Hence, on 
the one hand, we see the metamorphosis interrupted, a stop- 
page taking place at a determinate step; on the other, the meta- 
morphosis attained by passing over the intermediate steps. Still 
more remarkable are the cases in which the retardation is not 
merely an interruption at a determinate step, but. appears as a real 
retrogression in the metamorphosis, whereby an alternate rise 
and fall,—an oscillation,—usually takes place, which may at last 
pass over in victorious progress to the formation of flower and 
fruit; though in most instances it prevents the shoot in question 
from ever attaining its end. Helleborus viger is an example of 
the first case ; for after many years of inferior- and foliaceous- 
leaf formation, at last it attains superior leaves and fruit by over- 
leaping the formation of foliaceous-leaves which until then had 
prevented its farther progress.f Many of our trees with true fo- 
liage present examples of the second case. Their branches com- 
mence with bud-scales (inferior-leaves), the succeeding foliaceons 
branch ends with a terminal bud, (thus falling back to inferior- 
leaf formation,) and in the next period of vegetation they rise 
* On the terminology of the leaf-formations, see Wydler: Bot. Zeit., 1844, 36tes. 
Stuck., and A. Braun Verjiingung, p. 86. (Henfrey’s Transl Ray Soc., 1853, p 62, T.) 
f Analogous cases occur in the branches in AEsculus and many Maples which at- 
tain to flowers. Among herbaceous plants Anemone nemorosa and Asarum Euro- 
pcenm also belong here, and especially remarkable is the Tulip, the plants of which, 
not yet ripe for flowering, annually develop one single foliaceons leaf, followed by a 
central-bud hidden in the middle of the bulb and composed of several inferior-leaves. 
This bud preserves this position in bulbs deep in the ground, but in those nearer the 
surface it is, as it were, led out of the centre of the bulb, and sinks deeper into the 
earth, causing an indentation of the surrounding base of the preceding leaf in form 
like a spur, boring through the old bulb and penetrating vertically into the ground, 
and at the same time sinking itself into a deeper stratum with the spur;—an ar- 
rangement explained, but not with sufficient clearness by Henry in Nov. Act. Nat. 
Cur., vol. xxi, p. 275, 1.16 et 17. The Vegetable Individual in its relation to Species. 
23 
again to foliaceous-leaf formation,*—as in the Oak, Beech, and 
Poplar. A similar oscillation between inferior-leaf formation and 
foliaceous-leaf formation, keeping pace with the change of sea- 
son, is seen in the creeping main-shoot of Adoxa, and in the 
stock of Hepatica vobilis, creeping close to the soil, with its 
short internodes, and which in so far deserves its French name 
(la fille avant la mere) as its flowers, which unfold before the fo- 
liage, do not belong to the same individual as the foliage, but are 
produced laterally as a “ daughter generation” from the axils of 
the inferior-leaves of the maternal stem.f A similar phenome- 
non only in a higher degree, (a rising and falling between folia- 
ceous- and superior-leaf formation.) is presented by those plants 
whose inflorescence ends in a foliaceous coma, as is remarkably 
the case in the Pine-Apple, and also in the New Holland species 
of Melaleuca, and Callislemou, whose crowded, brush-like inflo- 
rescence (i. e. the region covered with superior-leaves and bear- 
ing the flowers in the axils of these) returns and forms foliace- 
ous-leaves, and in the following year again attains an inflorescence. 
While every leaf-formation may bring the progress of the met- 
amorphosis on a single shoot to a consummation, it is conceivable 
that one shoot may be allowed to each step for itself alone. Thus, 
there are shoots which represent inferior-leaf formation alone; e. g. 
the root-stock of Paris quadrifolia, the tuberiferous branches of 
the rhizoma of the Potatoe,| and there are some which are en- 
dowed with the foliaceous-leaf formation only, as the primary 
axis of many species of Veronica, the sterile leafy branches of 
several E upltorbice, as well as the leafy branches of those woody 
growths which have no bud-scales and no terminal inflorescence, 
(e. g. Rhamnus Fra.ngula'). Cases of pure superior-leaved shoots 
may be seen in the peduncles of Veronica Chamcedrys, ojjicina- 
* In such librations, of course, the formation of the flower can only be attained 
by particular branches, deviating in character from the rest,—the catkins which pass 
over leaf-formation advancing from the inferior-leaves immediately to the superior- 
lsaves out of whose axils the flowers are emitted. 
\ The same obtains in Galantlms nivalis, in which every annual generation con- 
sists of one inferior-leaf, one foliaceous-leal'with a vagina, and one without a vagina, 
which follow each other in simple alternation, in a distichous arrangement. The 
flower, as a branch, is emitted from the axil of the second foliaceous-leaf while the 
direct continuation of the shoot returns again to inferior-leaf formation. In striking 
contrast to the extremely simple relations of this plant we find Oxa/is tetraphylla 
and other species of that genus, in which the subterraneous main-stem also presents 
an alternation of inferior leaf-formation and foliaceous-leaf formation, advancing with 
the change of season, but conjoined with a rare abundance of leaves and a compli- 
cated phyllotaxis. The number of the inferior-leaves amounts to several hundreds; 
and transverse sections of the bulbs, which last through the winter and are formed 
by the close approximation of these leaves, form some of the prettiest specimens of 
phyllotaxis, showing 21-15 arrangement through easily computable 8-, 13- and 21- 
ranked oblique spirals. The number of the foliaceous-leaves is not so large; they 
develop in the summer, and form an 8 to 13 leaved rosette, out of which the axil- 
lary inflorescences issue, with their long peduncles. 
\ In case (as sometimes occurs) the tuber does not pass through this formation 
and advance to foliaceous-leaf formation. The'tuber is the thickened apex of the 
inferior-leaf shoot. Cf, the figure by Turpin: Mem. du Mus. d’Hist. Nat., 1.19, pi. 2. 24 
The Vegetable Individual in its relation to Species. 
Us, etc., in the (always lateral) spike-bearing scapes of Plantago, 
and the racemes of Covvallaria majalis, which shoot out of the 
axil of the highest lower-leaf as branches. Even the leaf-form- 
ation belonging to the flower can be divided among different 
shoots, and thus the flowers may be produced piecemeal, so to 
say ; as is the case in all dioecious plants, where the two most 
essential formations of the flower (the stamens and pistils) are 
found, not in the same flower, but in two separate ones. Even 
the less essential parts of the flower, the sepals and the petals, 
may occur separated from the other particular shootlets; as may 
be seen in the neutral flowers in the coma of the spike of Mus- 
cnri comosum and in the ray-flowers of the cyme of Viburnum 
Opulus. The destitution of the shoot may be carried so far as to 
cause it to produce but one single leaf, or one single formation 
(whether from the sphere of the plant-stock, or from that of the 
leaves); in which case the individual represents only one single 
organ ; as, for instance, in the branches which form the axis of the 
inflorescence in Vida, monanthos and other Leguminosse with ra- 
cemes reduced to one flower, bearing one single superior-leaf, from 
whose axil the flower proceeds. The male flower of Euphorbia 
is a peduncle whose flower consists of one single stamen.* Must 
we, now, still regard as individuals, these shoots, so partially en- 
dowed, and the last-named so destitute? Certainly! For if the 
individual can fall short, though ever so little, of the perfect real- 
ization of the specific idea, then there are no limits to its imper- 
fection and destitution ; for, after all, the realization of this vegeta- 
* The genuine eases will be of rare occurrence if we look at the cases which be- 
long here rigorously, that is. if we take into account the dwarfed foliaceous forma- 
tions which may possibly exist, suppressed or scarcely discernible. The male flower 
of Euphorbia itself properly belongs here only in appearance, as two small scales 
(inferior-leaves) occur, more or less developed, at the base of the peduncle. The 
small involucre of the male flower proceeds to develop itself out of one of these 
scales. (Cf. Wydlcr: Linnsea, 1843, p. 409.) Another example of a one-leaved 
shoot (though a spurious one) is presented in the Californian Pinus monophyllos 
[Fremont), whose lateral branchlets bear a fascicle of needle-shaped leaves reduced 
to one single needle: but this, as well as the pair of such leaves of our ordinary 
pines, is preceded by a vagina composed of several bud-scales. Perhaps another 
deception is played upon us in this case, for the perfectly round form of this needle 
excites the suspicion that it may be composed of two which have grown together 
through their whole length. The seed-bearing fruit-scales of the cone of Abietiuce, 
which are placed in the axils of the scales, also appear to be one-leaved shoots; but 
the series of changes which these scales present in cones of Pinus Larix which 
have completed their growth, proves that these fruit-scales are composed of two 
concrete leaves. The spurious axis of the Grape is a concatenation of alternating 
one- and two-leaved leaf-shoots, if we do not count the one or two little dwarfed su- 
perior-leaves, which in most cases are perceptible on the apex of the single shoot 
which finally forms a cirrhus. Ophioglossum presents a genuine case of a one-leaved 
shoot. The spike of this plant is a single fertile leaf, standing in the axil of the 
sterile one and hence belonging to a lateral axis, of which however nothing is per- 
ceptible but this leaf. (Cf. Schnizlein: Icon. fam. riat.. Heft II, t. 32.) The utricu- 
lus of Carex is the solitary leaf of an axis which in its normal condition develops 
no farther, and out of which, as the axillary formation of the utrieulus, the female 
flower is emitted. And the so-called neutral flower of Panicum., and the related 
grasses, is a shootlet which develops nothing but one leaf (the bract of the flower). The Vegetable Individual in its relation to Species. 
25 
ble Idea by the different members of the vegetable kingdom 
is precisely similar to the realization of the species by its single 
individuals. To be sure our idea of a plant implies that it shall 
manifest its life in a series of successive formations, that it 
shall put forth its leaves, flowers and fruit by successive steps; 
and yet there are plants which produce no leaves and no fruit 
(the Cryptogamia); again, there are others which hasten on to 
form flower and fruit with various intermissions of the regular 
steps, as is especially the case with the ugly parasites destitute of 
that green foliage which elsewhere is so characteristic a product 
ofthe vegetable world * One of these (the Hydnora,\ which preys 
upon the root of the South African Euphorbia) seems entirely 
devoid of all the foliage which is usually formed before the flower. 
Hence, therefore, in general we cannot necessarily regard indi- 
viduals as perfect representatives of the specific idea, and hence, 
too, we cannot regard them as representations invariably identi- 
cal in their realizations. Individuals appear rather as living at- 
tempts, by which the Idea is more or less attained, and is thus 
realized with various modifications. From this point of view 
even the differences in individuals, as pointed out by the doctrine 
of shoots, within the limits of vegetable species will no longer 
surprise us; on the contrary it will open to us a deeper insight 
into that independence presented to us even in the life of nature, 
in the realization of the internal problems of the creation. 
But here, too, as is so variously the case in nature, the regula- 
tive law is admirably united to the free configuration; for what 
gives a peculiar interest to the differences among shoots in the 
same species is the regular reciprocal relation among the shoots, 
as they reciprocally complete each other by their very one-sided- 
ness, and thus form a higher whole. In this respect the qualita- 
tive difference of shoots bears a certain relation to their origin, 
that is, to the order of ramification to which they belong.. And 
as the formation of shoots, as was shown, is a process of propa- 
gation, we see here, in the history of the development of the 
species, propagation taking the place of individual development. 
A second individual takes up the thread of reproduction which 
the preceding one was unable to carry any farther. Thus, what 
we are accustomed to see elsewhere attained in the individual, is 
here reached by the generation in a more or less strictly deter- 
* Orobnnche, Lathrcea, Monotropa, Gynomorium, all of -which agree in the infe- 
rior-leaf formation passing immediately into superior-leaf formation, and thus the 
formation of foliaceous-leaves is omitted. In the celebrated Ra.fflesia the immense 
flower is preceded by bud-scales only, which must be considered as the inferior-leaf 
formation. The same occurs in Frostia, which preys upon the branches of arbores- 
cent Leguminosce, and which resembles a mere flower so much that one might doubt 
whether it is merely a monstrous papilionaceous flower or a real parasite. (Cf. End- 
licher: Gen. plant., p. 16, and Guillemin: Nouv. Ann. des. sc. nat., II, t. 1, and as 
to parasites in general, Uvger : Annalen d. Wiener Museums, part II.) 
f E. Meyer: Nov. act. acad. L. C. nat. cur. XVI, 2, p. 771, t. 58 et 59, and R. 
Brown: On the female flower and fruit of Raffiesia and Hydnora, 1844, pi. 6-9. 26 
The Vegetable Individual in its relation to Species. 
mined cycle,—in other words, where the single shoot is incapa- 
ble, a determinate succession of shoot-series arises to bring the 
internal problem of its existence to a consummation,—to complete 
the metamorphosis into flower and fruit. This remarkable phe- 
nomenon,—which is a very frequent one in the vegetable king- 
dom, and is one of the essential characteristics of many of the 
most important families of plants, e. g., the grasses, Synanthe- 
rea>, Labiatiflorece, Crucifer cc, Leguminosce, etc.,—is the same 
as that which in the animal kingdom (in whose lower orders it 
re-appears) was, we cannot say discovered, but brought to a 
clearer comprehension not long since by the Norwegian natural- 
ist Sars,# completed and confirmed by von Siebold's investiga- 
tions into the history of the development of Medusa auriia.f 
and soon after substantiated in its universality by the Dane, 
Steenstrup, under the name of “alternation of generation,” or 
propagation and development by alternate series of 
Single cases of alternation of generation had been already care- 
fully observed but they were too much in opposition to the 
usual mode of reproduction to be understood in their true mean- 
ing. It was attempted to reconcile them with the customary mode 
by an unnatural interpretation, which regarded them as subver- 
sive exceptions to the general rule; while on the contrary almost 
all later works|| bring to light a multitude of unexpected facts 
* In Wiegmann’s Arcliiv, 1844, where the observations published in the author’s 
earlier works, on the adolescent states of Medusa are completed and concluded. 
f Beitrage zur Naturgesch. der wirbellosen Thiere. Danzig, 1839. 
Ueber d. Generationswechsel, iibersetzt von Lorenzen, Copenhagen, 1S42. 
§ Bonnet's industrious observations, the first that were made, of the alternating 
mode of reproduction of Aphis, published in his Trait6 de l’lnsectologie in 1745, 
though made in 1740, belong here. Also Chamisso’s correct observations of alter- 
nation of generation in Salpce described in his Memoir: de Animalibus quibusdam e 
classe vermium Linnaeana, Fasc. I, 1 SI9. Fragments in regard to the alternation of 
generation of Trematodce were known, (but as such they did seem very enigmatical,) 
by Bojwuus's Beschreibung d. konigsgelben Wfirmer (the “nurses” of Trematodce ac- 
cording to Steenstrup) aus welchen Cercarien (the larva; of the final generation) 
herauskommen (Isis, 1818), and by v. Bauer's important work on Cercarice and the 
related Bucephalus, (Beitrage zur Kenntniss d. uiederen Thiere. Act. nat. cur., Vol. 
XIII, 1827). 
|| Of the later works, by which the field of alternation of generation has been 
extended, I will adduce in particular; Sars: Fauna litoralis Norwegia;, 1846, in 
which the sections especially important in relation to alternation of generation are 
those on Sgncorgna, Podncoryna, Perigonimus, Cytais, as well as on Aga/mopsis, 
Biphyes and Scdpa.— Van Beneden: Recherelies sur l’embryogenie des Tubulaires 
(1814); Mem. sur les Campamdaires de la cote d’Ostende (1845 in the mem.de 
l’Acad. roy. de Bruxelles, T. XVII); Recherches sur l’anat., la physiol., et le (level, 
des Bryozoaires (Mem. de l’Acad. roy. de Br. T. XVIII).—Dujardin: Sur le devel. 
des Meduses et des Polypes hydraires (Ann. des sc. nat., Nov. 1845).— Krohn: Be- 
merkungen fiber die Geschlechtsverhaltnisse de Sertularinen (in Muller’s Arcliiv, 
1843, p. 174); Ueber d. Fortpfl. u. Entw. der Biplioren (Froriep’s neue Notizen, No. 
868, 1846).— Busch: Beob. fiber Anat. u. Entw. d. Infusorien (Arch. f. Naturgesch. 
XV, p. 92). How great an importance must be attributed to the discovery of alter- 
nation of generation in dispelling the darkness which until then settled on the his- 
tory of the life and development of Entozoa, may be seen in particular in v. Sie- 
bold's pregnant communications in B. Wagner's Handwbrterbuch d. Physiologie, p. 
640 (Article: Farasiten). The Vegetable Individual in its relation to Species. 
27 
which take their places naturally under the law of alternation of 
generation as now known, and substantiate the pertinent words 
of Goethe with which Steenstrup opens his Memoir: “Nature 
keeps on her course, and what seems an exception is in rule.” 
It was Sars, however, who first gave the answer to the riddle, 
the key to the newly opened domain, when he said of the course 
of development of Medusa, that here “it was not the individual, 
but the generation, which underwent the metamorphosis.”* This 
was the true point of view; for Steenstrup d welt too exclusively 
on the physiological side, the functional relations, of the alter- 
nating generations. Steenstrup, in fact, considered that the sig- 
nificance of alternation of generation consisted in its being an 
organic nursing of the brood connected with particular genera- 
tions, for which reason he termed the individuals of these genera- 
tions “ nurses —a mode of viewing the subject, which, with all 
Steenstrup’s pregnant elaboration of his idea, and with all the 
analogies he pointed out between it and the well-known phe- 
nomena of nursing the brood by particular individuals among 
bees, wasps, ants and termites, does not seize the essential point 
of the phenomenon of alternation of generations! R. Leuck- 
hardtj conceives alternation of generation from a more compre- 
hensive physiological point of view, in connection with the to- 
tality of all the other phenomena of the formation of different 
individuals, whether it occurs in a different or in the same gene- 
ration; regarding all these phenomena from the point of view of 
a division, not merely of the generic task, but of the vital task in 
general, among certain individuals; considering it as a polymorph- 
ism determined by a division of labor. But even this view must 
lead to the morphological one; for the division of labor is deter- 
* Sars: 1. c., p. 29. This assertion, of course, must not be understood as if the 
particular generation did not come in for its part of a metamorphosis. Sars’ vieAv 
is most beautifully corroborated by a comparison Avith plants; as in plants the meta- 
morphosis of the individual itself is connected Avith the formation Avhich leads to the 
completion of new parts, Avhich in their turn have their OAvn subordinate meta- 
morphosis. 
f Steenstrup’s explanation is most correct in regard to the history of the devel- 
opment of Distomce, whose nurses and grand-nurses are at last utricles entirely 
filled with the brood, and forming mere receptacles of the brood. Its application 
is less happy to those cases where the transition from the preparatory generations to 
the final generation takes place through external shoot- or bud-formation, as in Ser- 
tularice, Campanularicc, and Corynce, whose nurses forming the polypstem can con- 
tinue to live even after the concluding generations, comparable to the floAver in plants, 
separate or wither off. Hence the vital activity of the preparatory generations is 
not exhausted in the production of the brood. Steenstrup’s view, accordingly, 
would only be correct if non-sexual brood-production (by internal or external shoot- 
formation or by division) and alternation of generation were correlative conditions 
of each other. But this is not the case, as reproduction by shoots takes place Avith- 
out any alternation of generation in a great number of animals [Ascidia, Bryozoa, 
Madrepora), and by division as Avell [Astrcea. Annulate, Infusoria). These cases are 
comparable to the occurrence of unessential branches in plants; while alternation 
of generation represents the succession of essential shoots. 
\ Ueber d. Polymorphisms d. Indiv. od. d. Ersch. der Arbeitstheilung in d. Natur 
Ein Beitrag z. Lehre v. Generationsw. (1851.) 28 
The Vegetable Individual in its relation to Species. 
mined by the organic development, while this itself obtains its 
peculiar character from the determinate step of the metamorpho- 
sis at which the development ceases;—and this is just what is 
so unmistakable in the phenomena of alternation of generation 
in plants. Hence as a typical phenomenon of development, as a 
metamorphosis of generation, alternation of generation (as well 
as the metamorphosis of the individual) presents analogies with 
the graduated series in the animal and vegetable kingdoms, and 
the organic scale of the creation, in general ;—a point to which 
Y. Cams* called attention, and Reichert, his predecessor, as well. 
The difficulties which the qualitative differences of shoots of 
one and the same species seem to present to our conception of 
shoots as individuals, will be entirely obviated if we can demon- 
strate that a partial outfit and equipment of individuals, perfectly 
analogous to those found among plants, are likewise found in the 
animal kingdom, where in most cases there is less doubt as to 
what is an individual,—if we can show that in both kingdoms, 
and in a similar maimer, a polymorphism of individuals occurs 
which depends upon a division of the steps of development and 
of the vital problem of the species among individual members, 
whether of the same generation (divisions of generation), or of 
different generations cyclically succeeding each other (alternation 
of generation). 
Let us first compare the phenomena of alternation of generation 
(or, as it should be called, cyclical succession of generations) in 
both kingdoms.! As is the case in the alternation of generation 
* Zur naheren Kenntniss d. (1849); and, Einige Worte lib. Metam. 
u. Generationsw. (v. Seibo/d u. Kolliker: Zeitscbr. f. wiss. Zool. Ill, 1851, p. K59). 
f These remarks on alternation of generation in plants, do not depend, as one 
might perhaps be disposed to think, upon a zoological doctrine fancifully applied to 
plants. But I recognized the phenomenon as the same, and I treated of it in my 
papers, if not under the same name, still in the same meaning, before my attention 
was called to the occurrence of this phenomenon in the animal kingdom by Steen- 
strup’s work. As soon as the doctrine of the shoot as the vegetable individual was 
assumed in all its consequences, a determinate succession of generations emitted 
one from the other necessarily appeared to be the ground of the flower’s first making 
its appearance in many plants in a determinate degree of ramification, and of the 
occurrence of a determinate succession of steps in the series of axis up to this goal, 
caused by a peculiar partition of the leaf-formations. Hereby the essential shoot- 
succession, which is the one which represents alternation of generation, was accurately 
distinguished from the unessential one. Twenty years ago, or more, C. Schimper 
distinguished between essential and unessential shoots, denominating the first (in a 
wider sense of the word) “ Ableger” [off-setts], the latter “ Ausleger” [out-sets]. In 
the Versammlung d. Naturforscher in Mayence in the autumn of 1842, I made a 
communication on this subject, and at the same time in particular I called attention 
to the frequent importance of the characteristics involved in these relations when ap- 
plied to improving the differentiation and grouping of species. Of this commu- 
nication a report appeared in the Flora for 1842, p. 962, though, indeed, somewhat 
distorted by inaccuracies. Wpdler treated the same subject in the Bot. Zeit. 1844, 
St. 37, under the heading “ Achsenzahl der Gewachse,” and gives a compendium of 
examples, in which, however, much appears which needs qualification. As Wydler 
informs us, Aug. de St. Hilaire is said to have turned his attention to ascertaining 
the number of essential axes in plants; however I find nothing in the place referred The Vegetable Individual in its relation to Species. 
29 
of animals, a twofold reproduction appears in plants: sexual and 
non-sexual. Disregarding for the present the various relations of 
alternation of generation among the Cryptogamia, we find sexual 
reproduction (in animals by fertilized ova,—in plants by fertil- 
ized seeds) always vested in the generation which concludes the 
cycle of generations. That the consideration of this generation 
as the concluding one is not arbitrary, is shown by comparing it 
with the usual course of the metamorphosis; for the concluding 
generation is invested with the concluding formations of the met- 
amorphosis (flower and fruit), in the same way in fact as in the 
animal the complete development of the organs of generation 
occurs at the summit of the individual metamorphosis. The pre- 
ceding (preparatory) generations, which Steenstrup calls “nurses,” 
on the contrary invariably produce their brood by non-sexual re- 
production ; in the animal kingdom this takes place, now through 
germ-granules which develop in the interior of the body (as 
the nurses of Distoma:), now by a process of division in the pos- 
terior part of the body (the nurse of the Medusce, the Tapeworm), 
or finally by external, persistent or deciduous, shoot-formations, 
(Corynce, Campanularice, Sertularice, etc.). Among Phanero- 
gamia the last is the only kind occurring subservient to alterna- 
tions of generation. 
In animals, as in plants, the number of the generations in 
which the cycle of alternation of generation is completed, is for 
the most part a determinate one. Medusce, Salpce, Corynce, Tu- 
bularice conclude this cycle in the second generation ; according 
to Steenstrup’s showing, Distoma pacificum has a trimembral al- 
ternation of generation, and the family stock of Pennat.ula seems 
also to be formed by a trimembral succession of shoots. Cam- 
panularia has a quadrimembral cycle, in which however the two 
first generations are of the same character. Among Sertularice 
cycles of still more numerous members appear to occur; eight 
to ten generations form the annual cycle of generation of Aphides, 
though, excepting the last one, they are all similar and not even 
determinate as to number. 
To these examples from the animal kingdom much more nu- 
merous ones from the vegetable kingdom might be added, though 
to in the Legons de Bot unique but the distinction between determinate and indeter- 
minate growth, which has been known since Joachim Jung’s time, and was brought 
forward especially by Roeper and applied by him to classifying infloresences. It 
is exemplified, in that place, by creeping stems, upright root-stocks, and by bulbs; and 
the section on indeterminate stems is unluckily exemplified by wrong cases, viz., 
Scirpus palustris, Primula officinalis and Menyanthes, to which indeterminate main- 
shoots are falsely ascribed.—Steenstrup, also lays down an alternation of generation 
in plants, in the concluding remarks in his work quoted above, as well as in his later 
book, “Ueber das Vorkommeu des Hermaphroditismus in der Natur,” (On the phe- 
nomenon of hermaphroditism in Nature), though in an entirely different manner from 
mine as here given, for he compares the single leaves of the plant with the individ- 
ual in animals,—a mode of viewing the subject in regard to which I have already 
expressed my opinion in the Introduction. 30 
The Vegetable Individual in its relation to Species. 
I will only adduce a few of them here. Most Labiatiflorce, 
iSynantherecc, Grasses, Polygalece, Primulacece, the Dictamnus, 
Iris, Galanthus nivalis, etc., have a bimembral alternation of 
generation in different ways, according to the partition of the 
formations. In Paris, for example, the first generation takes 
the lowest grade: it presents a subterranean inferior-leaf shoot, 
(rhizoma) which never leaves the darkness of the earth, only 
reaching the world of light, towards which all plants strive, in 
its posterity, viz., in the quadrifoliate and unifloral lateral shoots 
which it sends up. The first generation of Viola odorata and 
related species forms foliage proper; still, the main axis tarries 
close to the earth, and the second generations (the lateral flowers) 
scarcely rise above the foliage. In Lysimachia numrnularia, 
the main-shoot, a rooting leaf-stem, creeps along the surface of 
the ground, growing indefinitely, and terminating only in the 
(essential) lateral branches by its golden-yellow flowers. The 
main shoot rises perpendicularly, forms foliage proper, and passes 
on to superior-leaf formation in many species of Veronica, e. g., 
F. acinifolici, producing its flowers as a second generation out of 
the axils of the leaves. The same holds good in regard to Oro- 
banche ramosa, which fixes itself and preys upon the root of hemp, 
though its main-shoot has no green leaves. A very remarkable bi- 
membral alternation of generation is shown by Adoxa, now so fa- 
mous, its name to the contrary notwithstanding.* The main-shoot 
creeps along the ground, oscillating with the seasons between leaf- 
and inferior-leaf formation,—at every return of the latter stretching 
out like a runner and boring into the earth. Flowers and fruit, 
frustrated by the invariable retrogression of the main-shoot, are 
produced by the aspiring perpendicular branches, after a pair of 
small leaves on the scape, and several insignificant superior-leaves, 
out of whose axils the lateral flowers are emitted as unessential 
shoots of the third degree. Hepatica presents a similar division of 
the formations among the two generations of shoots; but the 
main-shoot, rejuvenated from year to year and alternating be- 
tween inferior-leaf and leaf-formation, is short and upright. The 
branches with their single flowers, forming the second generation 
arise in the axils of the scale-like inferior leaves. A bimembral suc- 
cession of shoots occurs in Convallaria, Polygonatiim, the genus 
Aloe, all species of Plantago, Veronica officinalis, Chamcedrys, 
etc., Viola sylvatica, Lysimachia thyrsifolia, Alyssum saxatile 
and some other Cruci/erce, Echeveria coccinea, all the species 
of Melilotus, Medicago, Galega, in Pisum, and many other 
leguminous plants, and in Succisa pratensis, Anacyclus, Pyre- 
thrum, Polygonum Bistorta, etc. A familiar example occurs in 
\ 
* E. g. Adoxa moschatellina, 'which derives its name from Soifa (fame). The rela- 
tions of growth in this plant have been correctly described by Wydler: Bot. Zeit 
1844, p. 667. The Vegetable Individual in its relation to Species. 
31 
Secale. Its spiciferous culm forms the shoot of the first degree, 
the lateral spikelets which compose the spike itself are those of 
the second,* and the florets in the axils of the superior leaves 
(paleas) of these spikelets are the shoots of the third degree, i. e., 
the third generation of the cycle. A quadrimembral succession 
of shoots occurs in Trifolhim montanum, Hedysarnm corona- 
rium, and in several of the New Holland phyllodineous Acacice. 
Several species of Carex, e. g., C. maxima and leptostachys, have 
a trimemhral succession of shoots up to the male flower and a 
five-membral one up to the female. 
If we were to reckon the similar generations which are reared 
one above the other until the tree gains strength enough to per- 
fect its flowers, in many trees without terminal buds, as, in the 
Willow, the Linden,! we might find a number of generations 
equal or even much superior to that presented by Aphis. 
Besides the generation essential to itself, and by which it gives 
existence to the next grade in the cycle, every generation can 
have still another unessential reproduction, which only extends 
the same grade. As above we distinguished between essential 
and unessential shoots, so here accordingly we must distinguish 
an essential succession of generations,—the true alternation of 
generation,—and an unessential one. Very often both occur in 
the same species of plants. A fine example of this is shown in 
Lysimachia mimmularia, from whose creeping and rooting leaf- 
axis are emitted not only peduncles, but here and there new creep- 
ing leaf-axis exactly repeating the original one (except as to the 
two early-lost cotyledons): and from the undetermined leaf-bear- 
ing main-axis of Tropceolum minus are emitted in regular alter- 
nation three lateral flowers at a time, and then again one (unes- 
sential) leaf-shoot. In Cardamine amara the first generation (the 
stem bearing foliaceous and superior-leaves) is repeated in a two- 
fold manner, by lateral branches from the cauline leaves, and by 
creepers from axils of the root-leaves. Similar relations obtain 
in Mentha and a large number of other plants. This same phe- 
nomenon is repeated in the animal kingdom. The polyp-like 
nurses of the Medusa increase as such (according to Sars and 
von Siebold) by lateral buds and runners. Syncorynce are spadix- 
polypi, which represent trees by their formation of unessential 
branches, emitting finally from every branch and from the middle 
stock a whorl of individuals of the second (and last) degree. 
Campannlaricc and Sertularirc put forth runners from the bases 
of the main-individual, which again shoot up and become new 
main-stems, or new stems emerge out of them ; and perhaps the 
* Secale, in fact, has no terminal spicule; neither has Trilicum monococcum, 'while 
the other cultivated species of Trilicum have. 
f I have described the grape in reference to this subject in another place, (Ver- 
jiingung, p. 49,) [Henfrey's Transl. op. cit. p. 46. T.]. 32 
The Vegetable Individual in its relation to Species. 
ramifications of Bucephalus (which according to Steenstrup’s 
supposition is the larva of Aspidogaster conchila) as represented 
by Baer in Nov. Act. Nat. Cur., xiii, 2 belong here. 
In our qualitative comparison of shoots, it was shown how the 
shoot can be limited to a few leaves, or even to a single one ; in 
like manner the animal individual, in the division of role which 
occurs in alternation of generation, can become the representa- 
tive of one single organ, of one single function. Thus the fe- 
males of Coryne squamata are hardly anything more than egg- 
stocks, and the males than seed-stocks.# The members of the 
tapeworm, which are so many individuals of the final generation, 
hardly represent anything more than hemaphrodite sexual appa- 
ratus. As an analogous example in the vegetable kingdom per- 
haps the Willow\ may be compared to the Coryne; here too the 
shoots of the last degree are nothing but naked unisexual appa- 
ratus of reproduction. In Potamogetonon the contrary, they 
are hermaphrodite, as in the tapeworm. The construction of 
many of the lower animals, which when considered as individ- 
ual animals seem to be the strangest monsters, becomes more 
intelligible as soon as they are regarded from this point of view, 
—as soon as we make up our minds to regard the supposed indi- 
viduals as a family stock, and its parts (formerly held to be mere 
organs, and which, physiologically considered, are really nothing 
more) as individuals. In particular this is true of Physophora 
Stephanomia and Agalmopsis. 
In many cases we find alternation of generation connected 
with division of generation, that is, the appearance of heteroge- 
neous individuals in one and the same generation. Just as is the 
case in animal and vegetable forms without alternation of gene- 
ration, so where it is connected with alternation of generation, 
division of generation relates principally to the sexual functions; 
and a glance at the animal kingdom shows us relations of alter- 
nation of generation complicated by division perfectly similar to 
those which occur in the vegetable kingdom. In animals which 
go through an alternation of generation the individuals of the 
preparatory generations are non-sexual; still they may neverthe- 
less have a determinate importance in relation to the completion 
of the race which is to form their posterity. When in fact the 
final generation does not consist of hermaphrodite individuals, as 
obtains, for instance, in the tapeworm, various alternations are 
conceivable : the final individuals of both sexes can be nourished 
by the same nurse, and hence the sexual division will first take 
place in the second, or generally speaking, in the last generation ; 
* Hence Rathhe regards the male individuals as mere testicles. Cf. Wiegm. Ar- 
chiv., 1844, p. 155, and Steenstrup: Hermaph., tab. I, f. 17-20. 
f The two stamens in the Willow, and the floriferous bud as well, is preceded by 
only two very small bracts, which grow together and form a little scale. 
| The flowers of Potamogeton are branches which bear only stamens and carpels. The Vegetable Individual in its relation to Species. 
33 
or, different nurses may nourish the two sexes so that a division 
of generation will occur even at the degree of nurse-formation. 
If in the last case the nurses are not single ones, but even then 
form perse a family stock, then on the same stock we may either 
have male-bearing and female-bearing nurses together, or these 
two kinds of nurses may be divided among different stocks, ac- 
cording as the division of generation occurs in a determinate 
later generation, or is present already in the first. Although as 
yet the observations of these relations by no means form an un- 
broken chain,* still this much is certain, that in animals, in the 
same way as in plants, both monoecious and doecious forms occur; 
and hence there are families partly bisexual, partly unisexual. 
Cory nee, Tubularice, Campanulariae, and probably all Seriula- 
rine (hence, doubtless, the greater part of Hydroids), also Vere- 
tillum, Cynomorium, according to Steenstrup, Krolm and other 
observers, are dioecious,—whether they form small simple stocks 
as Coryne sqnarnala, or small ramified trees, as Syncorynce, 
Campanularice,\ etc. On the other hand Siphonophorice, ac- 
cording to Milne Edwards’ description of Stephanom,ia% (and 
judging from Sars’ description of Agalmopsis), are monoecious 
family stocks; Hydras are also To enter any fur- 
ther into these relations as they occur in the loweranimals would 
lead us too far from our subject; but it may be in place to give 
some details as to the manifold relations under which sexual di- 
vision of generation occurs in plants. 
Dioecious relations may occur without alternation of genera- 
tion when, in fact, the flower has a terminal inflorescence and no 
branches, or only unessential ones,—when, therefore, as it is usu- 
ally expressed, it is “ uniaxial,” as e. g., in Rubus, Chamoemorus, 
Lychnis, and Viscum. Much more frequently, however, divis- 
ion of the sexes occurs in plants which at the same time have a 
cyclical succession of shoots (alternation of generation),—a suc- 
cession which each of the two heterogeneous stocks passes 
through independently, and not always pari passu. This is a 
circumstance which must not be neglected in considering the 
differences of habitus in male and female flowers. Thus, in Mer- 
* Thus e. g., as far as I know, it remains to be shown whether the single nurses 
of Medusae produce Medusae of both sexes, or, as is most probable, only those of 
the same same sex. In Aphis also this point still needs to be more accurately de- 
termined. 
\ Steenstrup: Hermaph., pp. 66, 6*7, 72, 
% Ann. des Sc. Nat., 1841, p. 217, pi. 7-10. 
§ The later investigations into Siphonophorice by Huxley: Edin. Phil. Journ., 
1852,Kdlliker: Zeitschr. f. wiss. Zool., 1852, and Leuckardt: Zool. Untersuch., lstes 
Heft, 1853, corroborate the monoecious relations of these wonderful creatures as re- 
gards most of their genera, e. g., Agalma, Agalmopsis, Stephanomia (Apolemia), 
Physophora, and the other closely related genera; Busch's researches into the group 
of JDiphyidce have proved them to be dicecious(i and the same obtains in the related 
genus Epibulia. (Later note.) 34 
The Vegetable Individual in its relation to Species. 
scuriali the female plant bears flowers even on the second axis ; 
in the male plant, however,—if I do not misunderstand the in- 
florescence (a spike composed of small glomerules)—this first 
occurs on the third. In Carex dioica, vice versa, the male plant, 
flowers in the second line and the female in the third.* In other 
dioecious plants on the other hand, the male and female flowers 
appear in the corresponding generation, e. g., in the second: 
Slratiotes, Empetrum and Taxmen in the third: Salix, Popu- 
jus, Myrica, Cannabis; in the fourth : Phoenix. In Hemp the 
extremely heterogeneous appearance of the inflorescence of the 
male and female plants does not depend upon a division of the 
flowers of the two sexes among different axes, but upon the pro- 
duction of numerous unessential peduncles in the male inflores- 
cence.! 
Monoecism necessarily presupposes a succession of shoots (alter- 
nation of generation); in the simplest case at least for one of the 
two sexes, as both cannot be united in the same terminal flower : 
but vice versa, both may easily appear in determinate (equal or 
unequal) degrees of ramification. The most important circum- 
stance to be considered in monoecious relations, consists in both 
the sexes (i. e., the shoots which bear them) occurring either sub- 
ordinately or coordinately,| for one either arises out of the other, 
or they both spring from a common mother-stem. In the first case, 
the female flower usually belongs to the earlier, the male to the 
later (subordinate) generation ; the male flower-shoot springing 
from the female,$ as e. g., in Euphorbia, Ricinus and Poterium, 
in which the female, flower terminates the main axis, and the 
male occurs as a lateral shoot.|| In Buxus the female flower oc- 
curs as the second, the male as the third axis ; in many species 
of Phyllanthus (e. g., Ph. niruri) the female as the third, the 
* The second axis, which is a complete dwarf or a mere bristly spine bears the so- 
called Urceolus, in the axil of which the female flower is placed, as the third mem- 
ber of the succession of generations. 
f The female flowers are placed at the sides of the primary branches as branches 
of the second degree. In the same place where one single flower occurs in the fe- 
male plant, a furcately ramified inflorescence is found in the male, produced by 
branching out of the two bracts of the original flower. 
\ Both these cases doubtless occur in the animal kingdom, the first probably in A l- 
cyonella, where the stock is said to be composed partly of males and partly of females. 
As the stock is here formed by individuals continually shooting out of each other, 
one sex must shoot out of the other. The second case occurs in Agalmopsis (accord- 
ing to Sars,) where partly female (seminal vesicles,) and partly male individuals 
grow out of the same main-stem. 
§ The opposite case seems to occur very rarely or not at all. A monstrosity, 
which, for some reasons might be adduced here, is found in Larix JEuropcea and Picea 
alba, in which transitions of the amentaceous male flowers into female cones occur, 
where the fruit scales are emitted from the axils of stamens which are often only 
slightly abnormal. 
|| As in all the examples adduced, the unessential aggrandizement of the inflores- 
cence must be disregarded, which occurs in Ricimcs and Poterium in the form of lat- 
eral female flowers emitted beneath the terminal female flower. The Vegetable Individual in its relation to Species. 
35 
male as the fourth ; in Xylophylla, the female (on the margins of 
the spurious leaves) as the fourth, the male arising from the bracts 
of the female flower, (as in Phyllanthus) as the fifth. In Mo- 
mordica, Ecbalium, Cephalanthera and some other Cucurbitacece, 
the female flower, placed in the axils of the foliaceous leaves of 
the main stem, belongs to the third axis, and the male to the 
fourth ; for the third axis, which here arises from the base of the 
peduncle of the female flower as main axis of the racemose male 
inflorescence, is a su perior leaf-shoot. In the other cases,—in which 
the succession of shoots, in order to arrive at the two kinds of 
flowers, separates into two coordinate lines,—both kinds of flow- 
ers can appear either immediately in the first generation after this 
separation, or, since here again preparatory generations are inter- 
calated, in a later one. Further, the number of the generations 
(axes) in the two lines arising from the division, may be either 
equal or unequal. A few examples may serve to explain the man- 
ifold cases which thus occur. In Musa, Myriophyllum and Sag- 
itaria the coordinate male and female flowers appear in the first 
generation after the separation, and in the whole as a second 
system of axes. Here the female flowers stand in the lower, the 
male in the upper part of the spicate or racemose inflorescence. 
The contrary holds true of Cncurbita and the monoecious Bry- 
onice •* for here the earlier flowers, which appear in the axils of 
the foliaceous leaves, are male ; while the later ones which appear 
on the farther continuations of the stems are female. Arumf 
has below female, in the middle male, and above again female 
flowers, though these last are dwarfed and sterile. Likewise in 
the first generation after the separation, but in the whole as the 
third system of axes, we find both kinds of flowers in Pachysan- 
dra and Acalypha, and here again, as is usually the case in inde- 
terminate spicate inflorescences of mixed sexes, the female flower 
is in the lower, the male in the upper part of the inflorescence. 
The same obtains in monoecious Palms with axillary spadices ; 
though here the flowers appear in ramified spikes from the fourth 
system of axes. When the flowers make their appearance in the 
second generation after the division, they cannot easily be united 
in the same inflorescence, and special male and female inflores- 
cences will arise. Thus, e. g., in Platanus, Liquidambar and 
Sparganium, in which the female inflorescences occur on the 
lower part of the main shoot, and the male in the upper; like- 
wise in Quei'cus and Fagus, though here, vice versa, the male 
inflorescences are the lower, and the female the upper. Finally, 
* Bryonia has apparently axillary racemes, but a more careful investigation shows 
that they do not spring immediately out of the axil of the foliaceous leaf, but (as 
secondary branches) out of the peduncle of a single flower standing directly in the 
axil of the leaf which exactly corresponds to the flower in Cncurbita. 
f The inflorescence in Arum is terminal, as well as that in Call a. 36 
The Vegetable Individual in its relation to Spedes. 
if the division of the succession of shoots is an unequal one in 
the separated lines of generation leading to the two kinds of 
flowers; i. e., if the number of essential axes is unequal, it is 
greater sometimes for one sex and sometimes for the other. In 
the Walnut (Juglans) it is the male flower which attains the 
higher degree of ramification; in Xanthium and the species of 
Carex with separated male and female spikes it is, on the con- 
trary, the female flower.* 
Other dimorphisms or even polymorphisms of the flowers, more 
or less independent of sex, occur when the sexes appear in the 
two different lines of generation ; for even among flowers of the 
same sex, whether hermaphrodite, male, or female, differences 
often reveal themselves of a very striking character, which are 
generally coordinate according to fixed laws of division of gen- 
eration. Thus, in all Primulce, and in several Labiatce, two 
kinds of hermaphrodite flowers occur, in a state of dioecious sepa- 
ration : one with a large corolla and strongly developed stamens 
(forma brevistyla), the other with a small corolla and strongly 
developed pistils (forma longistyla). Acccording to C. Schim- 
per’s observationsf both forms occur at times in Labiatce even on 
the same stock and in the same inflorescence, e. g., in Dracoce- 
phalum Moldavica. Many species of Viola also produce two 
kinds of hermaphrodite flowers on the same stock: early ones of 
the usual form, and late ones without petals. In Viola mirabilis 
the first arise directly out of the main stem (as branches of the 
first degree) and are mostly sterile, while the latter spring from 
the foliaceous branches (as branches of the second degree) and 
are fertile. In Tmpatiens sterile flowers with perfect corollas and 
apetalous fertile ones occur in the same raceme. The cases in 
which normally formed above-ground and abnormally formed 
underground flowers appear belong here; the latter have their 
corolla developed slightly or not at all, and are merely female, 
and, par excellence, fertile. If both kinds of flowers are fertile, 
the subterranean fruit differs from that borne above the soil; such 
cases are found especially in the family of Leguminosce, e. g., in 
several species of Lathyrus and of Vicia, in Amphicarpcea, and 
in Arachis;% and also in the very remarkable Abyssinian Con- 
* In species of Carex with terminal male and lateral female spikes, the male 
flower belongs to the first generation after the division, and the female to the third. 
In most of the species where the shootlet which bears the inflorescences is a contin- 
uation of the main axis of the plant, the male flowers represent in general the sec- 
ond generation and the female the fourth; in those species, on the other hand, ■which 
have a shortened main axis, which forms a mere rosette of leaves whence the shootlets 
bearing the inflorescences proceed as branches,—in these species the male flower is 
the third system of axes, and the female the fifth ; as e. g., in Carex maxima, lepto- 
stachys and pilosa. 
f Communicated in the Yersam. d. Natur. zu Wiesb. in Sept. 1852. 
For details, vid. Treviramts: Bot. Zeit., 1853, p. 393. The Vegetable Individual in its relation to Species. 
37 
volvulacea, Hygrocharis Abyssinica.* Among the most striking 
cases of dimorphous flower-formation are those described by Jus- 
sieuf in Gaudichaudia, Camarea, and other Malpighiacece. 
Here, besides the flowers conjoined in racemes or in corymbs, 
and formed according to the common type of the family, other 
apetalons flowers occur, standing alone and hid in the axils of 
the leaves. Besides the normally formed glandulose corolla, 
they have only one stamen and two carpels. In several cases 
the dimorphism of the flowers is confined to the formation of 
the fruit alone, as, e. g., in some species of JEthionema, (espe- 
cially AE. heterocarpum, Gay,) which in the same raceme bear 
partly dehiscent silicles with two cells and several seeds, and 
partly one-celled and one-seeded indehiscent silicles. Ceratocap- 
nos,% a North African genus of FumariaccB, bears in the lower 
part of the spike oval, ribbed, one-seeded nutlets, and in the 
upper part, lanceolate two-valved and two-seeded siliques. Poly- 
morphism of flowers and fruit occurs in the most heterogeneous 
manner in the family of Composites; I will only refer to Zinnia, 
Dimorphotheca, Heterotheca, Thrincia, Geropogon, Crupina ; 
and especially to Calendula, where the hermaphrodite blossoms 
of the ray produce three different forms of fruit, so that, includ- 
ing the male flowers of the disc, the capitulum presents four dif- 
ferent forms of flower-shoots (belonging to the same generation). 
As somewhat similar cases in the animal kingdom, the instances 
of dimorphal insects, of which there are several, might be ad- 
duced.$ 
A separation of the series of generations into several distinct 
lines occurs in fact not only as regards the flower, but also, though 
less frequently, even among the inferior formations of the plant; 
this is especially the case where a particular lateral line is allotted 
to the leaf as well as to the flower. The true Pines afford the 
best known example of this. Their fascicles of needle-shaped 
leaves are nothing but foliaceous branches of circumscribed 
growth,j| which lie outside of the line which leads to the two 
kinds of flowers, while they are essential as the leaf-formation 
* Jlochstetter: in Scliimp. Iter Abyss., Ho. 572 et 1701. The same plant is called 
Nephrophyllum Abyssinicum by Richard: Tent. Flor. Abyss., and figured in pi. 76. 
The two kinds of flowers are emitted from the axils of the foliaceous leaves of the 
same creeping stem; those provided with corolla, stamens and pistil stand upright; 
the others without corolla and stamens, bend down to the ground on their long pe- 
duncles. 
j Adr. de Jussieu: Monographic des Malpighiacees. (1843.) 
| Durieu: Explor. scient. de l’Algerie, pi. 7 8. Endlicher: Gen. plant., Suppl. 
IV, p. 32. 
§ The first in several species of Dycticus (D. marginalis, circumcinctus, Lapponi- 
cus, Rceselii, according to Erichson : Gen. Dyticeorum, 1832, p. 31; the last in Aphis 
Quercus according to Bonnet. 
[| That the fascicles of leaves in Pinus are branches, is proved by the phenome- 
non of percrescence, which is not unfrequent, especially in young pines. 38 
The Vegetable Individual in its relation to Species. 
appears on them alone.* Here the generation splits up into 
three kinds of essential and coordinate shoots: 1st, the small 
leaf-shoots which after some few inferior-leaves forming the 
vagina, bear two, three, or five foliaceous leaves; 2d, the male 
flowers, or small shoots, which are provided with stamens 
only; 3d, female inflorescence, shoots with superior-leaves (the 
integumentary scales of the strobile) in whose axils the fruit- 
scales of the cone are formed, belonging to a farther system of 
axes. In the animal kingdom cases analogous to these occur 
in monoecious Siphonophorai, especially in Stephanomia and 
Agalmopsis, where even more than three kinds of coordinate in- 
dividuals are emitted from the main axis: in particular motory 
individuals (the so-called swimming-bells), nurses, the proboscis- 
like formations or imbibing tubes, and as already mentioned, two 
kinds of sexual individuals. 
The differences of shoots thus far considered depend princi- 
pally upon this: one portion represents exclusively the vegeta- 
tive formation, or a certain part thereof; the others represent the 
degrees of formation which belong exclusively or principally 
to the sphere of fructification. Hence, in regard to the division 
of functions, to one portion the functions of nutrition are allotted, 
to the others those of generation. For this reason the different 
kinds of shoots of such a partial character must unite in,a deter- 
minate succession, and complete each other; and even those 
which we have designated as unessential are of importance in 
enriching, preserving, and increasing the plant-stock. Finally, 
we have still to consider those shoot-formations which properly 
do not belong either to the essential or the unessential succession 
of shoots, but rather to an aberrant formation; as they neither 
conduce to the perfection of any of the common steps of the 
metamorphosis, nor perform any essential physiological function 
in the plant, but at the best are only of some service as organs 
of defence, support or adherence. These are the shoots which 
take the form of thorns, bristles, hooks and tendrils, which for 
the most part owe their peculiar abnormal character to an entire 
suppression of the leaf-formation, and a final induration of the 
point of vegetation: these seem to be the last, terminal or lateral 
members of the generation, abortive in every respect. Not un- 
frequently they form the last ramification of paniculate and dich- 
otomous inflorescences, like terminal flowerless peduncles, as, 
e. g., in Teloxys (Chenopodium aristatum, L.), Acroglochin, and 
in a very peculiar form, branching and complicated by aculeate 
or setiform leaf-formations, in Pupalia, Desmochceta, Digera and 
* The main-stem, as ■well as all the elongated branches essentially resembling the 
stem, bear only leaf-scales, which may be best compared to bud-scales, and ascribed 
to the inferior-leaf formation. It is only in early youth (in the first and second 
years) that the main-stem itself bears needle-shaped leaves. The Vegetable Individual in its relation to Species. 
39 
Cometes ;* also, in Scleropus, where they take the form of short, 
thick, cartilaginous stalks, with two converging leaf-apicules. 
Among the grasses they are known under the form of bristles in 
Setaria. In many Rhamnaceous and Sapindaceous plants (Heli- 
nus, Cardiospermum) they appear as small cirrhi, not as the last 
sterile ramifications of the inflorescence, but on the contrary as 
the first, followed by other fertile peduncles. They often occur 
in the axils of foliaceous leaves; and wherever they make their 
appearance they naturally arrest the farther succession of shoots, 
when they have neither of the two leaves at their origin, out of 
whose axil an additional shoot may be developed. This is the 
case in Pcissijlora, whose flower arises from the axil of a leaf 
situated at the side of the base of the tendril. The thorns of 
Ononis, Elceagnus and Macluraf present the same phenome- 
non. In other cases the succession of generation thus arrested 
by the aculeate shoot is restored by secondary formations ; when, 
with the thorn, a second shoot follows out of the axil, which in 
some cases may form a leaf-shoot, and in others a flower-shoot. 
This happens in Gleditschia, in several Acacice (e. g., A. pul- 
chella), in Prinsepia utilis,% the Lemon, the Egyptian Balan- 
ites, Duranta, Bouganvillea and Randia, in which the second- 
ary shoot arises close under the spine; while in Celasirus pyr- 
rhacantha§ and Europeans, as well as Pisonia aculeata,|| the 
secondary shoot occurs above the thorn. In Uncaria and 
Slrycknos spinosa, pairs of leaves with axillary thorns alternate 
with pairs which have peduncles in their axils. 
Have even these phenomena of extreme alienation of the in- 
dividual (as they occur in the thorns and hardened shoots of plants) 
analogous forms in the animal kingdom ? Yes, I believe they 
have ! I believe I may assert that in the animal kingdom itself 
there are individuals which occur as mere fixed claws, pincers, 
scourges, tactual and predial filaments, etc.,—individuals which 
perform neither functions of nutrition nor of reproduction in the 
society to which they belong, but which probably merely assist 
in seizing the food, or lend a helping hand in defending the 
community. The cases which I have here in mind are of fre- 
quent occurrence among Bryozoa, and especially in the group 
* The plumose tails which form the “envelope” of Cometes, are the last branches 
of the dichotomous inflorescence, accompanied by similar accessory (secondary and 
tertiary) branclilets. All these numerous sterile branchlets are elongated and beset 
with setiform leaflets arranged in spiral order (.|), commencing with two similar an- 
terior leaves. The direction of the phyllotaxis in all these branchlets follows the 
law of furcate inflorescence. 
\ Here belongs also the curious hook of Uncinia, which is also visible, though 
less developed, in many species of Carex. The utriculus is a leaf at the base of 
this spine. 
\ Boyle: Illustr. of the Bot. of Himal., pi. 38, fig. 1. 
8 Bolster: Voy. bot. en Espagne, t. 38. 
| Rheede: Hort. Malab., vii, t. 17. Wallich: Plant. As. rar., t. V70. 40 
The Vegetable Individual in its relation to Species. 
of Celia? 'ice. Individuals in the form of horns (which usually 
conclude the series of complete cell-inhabiting individuals) occur, 
e. g., in Eucrabea cornuta ,* and Cordierii;f in another form 
(reminding us of Teloxys,) as forked terminal spines, in Vesicu- 
laria spinosa.% Moveable individuals, representing mere weap- 
ons, in form like a bird’s beak, a crab’s claw or a pincers, appear in 
Acamarchis avicularia<§> and Jlustroides. || Retepora cellulosa 
iScrupoceUaria and many others. In the last named 
Cellarice, besides the claw-individuals, there are also scourge-in- 
dividuals, which Yan Beneden himself compared to the cirrhi in 
plants, and which even Leuckardt** acknowledges to be indi- 
viduals. Beside the ‘ Swimming-bells’ evidenily resembling Me- 
dusce, the peculiar retractile predial filaments of the Siphono- 
phorce doubtless belong here also ; they are remarkable for a pur- 
plish-red swelling on or under the apex, and they shoot out sin- 
gly as branches from the stalk of the nutritive individual (irn- 
bibing-tubes), and themselves bear a series of similarly formed 
filaments as secondary branches. They are found with unimport- 
ant departures from this form, especially in Physophora,-\f Di- 
phyesXX and Agalmopsis. In the last named genus, according to 
they have even three modifications : the spadiciferous ter- 
minal piece ends in a long simple filament, or in a short two- 
parted one, or without any filament at all. In Stephanomia\\\\ 
numerous filaments, called tentacles, arise out of the stalk of the 
nutritive animals (the so-called proboscis-formed organs) without 
such colored swellings, which in the same manner may also be 
regarded merely as individuals with a very incomplete outfit of 
organs.HY 
* Ellis: op. cit. pi. 21, f. 10. (Cellaria cornuta); M. Edw.: Ann. d. Sc. Nat., 
(1838) t. 8, f. 2 (C'risidia cornuta). 
f Descrip, de l’Eypte: Polypes, t. 13, f. 3. 
\ Van Beneden: Recli. sur les Bryozoaires, t. 4, f. c. 
§ Van Beneden : 1. c.,t. 6, f. 1-8 (Ce/lularia avicularia Pall. Crisia avicularia 
Lamx.) || Ellis : op. cit., pi. 38, f. 7. 
®;’ Van Beneden: 1. c., t. 5, f. 8-16 (Cellaria scruposa Auct.) 
** Leuckardt: Polymorphism, p. 17. f f Philippi: Muller’s Archiv, 1843, taf. 6. 
X% Sars: Fauna lit. Norw. tab. 7. §§ lb. tab. 6. 
|| | Milne Edwards : Ann. d. Sc. Nat., 1841, pi. 7-10. 
Since Sars observed the separation of the Medusa-like sexual individuals in 
Agalmopsis. the view that Siphonophorce are composite animal stocks has gained 
ground more and more among zoologists. But this mode of viewing the subject was 
for the first time carried out (after a fashion,), consequently in Leuckhardt’s latest work 
on strange animal forms (Zool. Pinters, erstes Deft: Siphonophoren, 1853); and 
this idea had forced itself upon me as early as 1847, when I compared the descrip- 
tion JDiphyes with Agalmopsis, in Sars’ Fauna lit. Norw. In the above named work, 
Leueklmrdt extends the view which allows individual importance to the parts of the 
stock of Siphonophorce not only to the tentacles and predial filaments, but also to the 
covercles, which in most of the genera are placed close above the nutritive individ- 
ual as protective envelopes; these formations, like all the other appendages of indi- 
vidual importance, being emitted from the stem as shootlets, and in the first stages 
of their formation, resemble the tentacles in particular. Accordingly Siphonophorce 
have not less than eight different forms under which the individual may appear on 
the whole stock. . . . (Later note.) [I have omitted the enumeration of these 
forms.—t.J The Vegetable Individual in its relation to Species. 
41 
After having in the foregoing review regarded all lateral shoots 
which spring from the main axis of the plant as real individuals, 
however 'unimportant a fraction of the total specific character 
they may realize, it will hardly be deemed surprising if we 
finally apply this mode of view to the branches of the root and 
to adventitious shoots. It is only possible for the main-shoot to 
develop freely both the points of vegetation of the axis ; yet 
even here the lower point remains undeveloped. On the con- 
trary, the lateral shoots, thus far considered, have no lower point 
of vegetation ; for their base is united to the maternal shoot, and 
hence they are mere developments of the upper point of vegeta- 
tion. Opposed to these, there are, however, other shoots by 
which the lower point of vegetation is represented, and which on 
the other hand have no upper point of vegetation. Among these 
may be reckoned not only the root-branches which take their rise 
from the main root, but also all adventitious roots which spring 
from the stem at determinate or indeterminate places. I must, 
however, content myself with this general hint, as any attempt 
to particularize these relations could after all only show the defi- 
ciency of the investigations into this subject, and how desirable 
a more comprehensive work is on root-formation in the vegetable 
kingdom. 
The few points which I have selected out of the inexhausti- 
ble field of shoot-formation in the vegetable kingdom may in the 
mean time suffice to show that the comparison of the vegetable 
shoot with the animal individual is not far-fetched or arbitrary, 
but is presented to us by Nature herself. The solution of the 
difficulties which this mode of conceiving the vegetable individ- 
ual encounters in the lowest grades of the vegetable kingdom, I 
must defer to a later day. These difficulties are founded upon the 
less complete organization of the inferior plants, and at all events, 
cannot invalidate the results gained in considering the higher or- 
ganizations. We may therefore consider it settled, that although 
the individual has not exactly the same importance in the veget- 
able kingdom as in the animal, plants still realize their vital cycle 
in sections which are not only comparable to the animal individ- 
ual, but are in fact its complete analogues. What distinguishes 
plants is the formation of family-stocks, (a formation manifested 
in the highest vegetable representations, and here in the richest 
fullness),—as ancestral trees organically connected, variously dis- 
posed in their ramifications, and comprising numerous genera- 
tions, rendered reciprocally complete through individuals vari- 
ously endowed. And this leads us back again to the tree from 
which we set out; in which even our natural perceptions seemed 
to discern something more than one common individual, and 
whose high import scientific research must confirm. Just what 
at the outset appeared to be an obstacle to our allowing the single 42 
The Vegetable Individual in its relation to Species. 
shoots of the tree their true significance,—now that we have com- 
pared them with alternation of generation in animals at length 
proves to be the most conclusive demonstration of the correctness 
of our first conception. The conception of these so heterogeneous 
shoots as individuals of one and the same species has led us, in 
fact, to a more profound and more pregnant conception of indi- 
viduality, which will no longer seem paradoxical when we per- 
ceive it is confirmed even in the highest realms of life—in the 
sphere of the mental development of the individual. Or are the 
differences of human individuals in mental endowment and de- 
velopment less important than those which we have seen in the 
morphological and physiological endowment and development of 
shoots ? Do we not meet with a similar reciprocal completion, a 
similar division of labor among the individuals of the family, of 
the state and of nations, and cannot even the human individual 
become likewise a mere organ ? Do we not see the development 
of the human race itself bound up with a succession, in which 
the later generations continue the edifice their predecessors be- 
gan, like branches depending upon the earlier stocks and nour- 
ished by them ;—in which generation is added to generation, 
and cycles to cycles; so that thus by the ever-renewed labor of 
the individual the problem of human life may be ceaselessly as- 
pired to, and at last reach its final accomplishment ?* 
* The preceding pages were almost all printed when I was fortunately enabled to 
read Reichert’s memoir (die monogene Fortpflanzung, Dorpat, 1852,) upon a subject 
closely allied to the one here discussed. His work is full of new views of the 
subject, elaborated with great acuteness. The vegetable individual itself is con- 
sidered in detail, and the author is thus led to a mode of viewing this subject similar 
to the Schultz-Schultzenstein-ian doctrine of anaphyta—regarding not only the shoot, 
but even its single parts, the internodes, with their leaves, as series of individuals 
shooting out of each other, or intimately connected by continuable bud-formation. 
Since, however, it is implied in the idea of an individual, that it shall somehow be 
limited by, and distinguishable from, (notwithstanding it is connected with) others ; 
it seems to me that even from this point of view Reichert’s idea can by no means be 
carried out. I will not deny that there are still other considerations in the nature 
of the shoot which it is difficult to reconcile with the idea of the simple individual, 
and I can only find the ground of this phenomenon in the fact, that the individual 
appears in its full import in the higher steps of the series of created beings, while 
in the lower it loses more and more its reality, if I may so say. I must reserve far- 
ther remarks on this subject until I treat of the individuality of the lower plants. 
[We cannot but think, after all, that this view of Reichert’s, &c., which our au- 
thor rejects, is the legitimate conclusion, to which the very line of argument so com- 
pletely and ably presented in the preceding pages, when fully carried out, naturally 
leads. It is merely a question of degree of individuality. As yet, perhaps, no sure 
middle ground has been secured between the two extreme views,—one of which re- 
gards all the vegetative offspring of a seed, however numerously multiplied, as phil- 
osophically the individual; while the other views the phyton, or in the simplest 
lower plants the cell, as philosophically representing the individual,—real individu- 
ality being incompletely realized (and with various grades of incompleteness) in 
all vegetables, and in many animals. The mind is reluctant to accept either of 
these conclusions, and seeks—thus far in vain—for some stable intermediate view. 
Of the two extreme views, if forced to the choice, we should incline to prefer the 
latter.—a. g.]