THE

NATURAL PHILOSOPHY
OF PLANT FORM
BY
AGNES ARBER
M.A., D.Sc., F.R.s., F.L.S.
CAMBRIDGE
AT THE UNIVERSITY PRESS
ARISTOTLE AND THEOPHRASTUS
of parts that do so subdivide, for instance, hand is composed of
flesh, sinews and bones. "
1
In another great zoological work,
De partibus animalium, he carries the analysis a stage further back,
and traces 'similar parts' (Tex .601oepfl) to certain elementary
forces,
2
or primary substances,
3
from which they are compounded,
while the' dissimilar parts' (Tex &voo1oepfl) are correspondingly
compounded out of the similar parts. His description of the
. system of the living body is thus that it consists of elements, which
are the material for Tex 601oepfl, which are, in their turn,
material for Tex &voo1oepfl, which, in the final synthesis, make
up the organism as a whole. The division into similar and dis-
similar-or homogeneous and heterogeneous-parts, is roughly
equivalent to the more modern classification into tissues and
organs. In.the Oxford translation of De partibus animalium, it was
noted that this equivalence was incomplete, because Aristotle
included among homogeneous parts "much that we should not
call tissue, e.g. the blood ";
4
but in recent times there has been
. a return to Aristotle in this matter, for blood is now regarded as
a liquid tissue. 5
Aristotle held that the plant body was divisible on the same
principles as that of animals, but, as it was -only incidentally that
he touched upon plants, there is more content for us in the writings.
ofhis successor, Theophrastus (S70 B.c. to 285 B.c.), to whom it
seems possible that he may have handed over, deliberately, the
continuation of his work on the botanical aspect of biology.(>
- Theophrastus was n<?t more than fifteen years younger than
Aristotle, and, like him, had studied under Plato, so that he is
perhaps more justly described as a junior contemporary and
colleague of Aristotle, than merely as his pupil.
The technical terms, which Theophrastus adopts for the minor
parts of plants, which had not, like the principal parts, already
acquired primary folk names, illustrate one of the more charac-
1 Thompson, D'Arcy W. (1910), vol. IV [Hist. animal. I. 1. 486a (Oxford
trans.)J. Thompson points out that Aristotle's distinction of similar and dissimilar
parts is derived from Anaxagoras .
.
2
Ogle, W. (1912), vol. v [De part. anim. 11. 1. 1. 646a (Oxford trans.)].
,
3
Peck, A. L. (1937) ,.p. 12 [De part. anim.].
4
Ogle, W. (1912), vol. v [De part. anim. note to II. 1. 646a (Oxford trans.)].
6
Cf., for instance, Cabot, R. C. (1904), p. 4, and Roberts, M . . (1926), p. 180.
6
Cf. Stromberg, R. (1937), p. 25; for an important analysis of the relation which
, the biological work ofTheophrastus bears to that of Aristotle, see Senn, G. (1933).
I I
II. MORPHOLOGY OF THE ARISTOTELIAN SCHOOL
teristic features of the botany of the Aristotelian school-a con-
stant reference to animals; in consequence, our botanical language,
from classical times until to-day, has faithfully reflected zoological
terminology. As examples from Theophrastus we may mention:
(flesh), for plant pulp; pQ:x1s (backbone) for the midrib of
a leaf; Ka:p5ia: (heart) for heart-wood or the pith region; and is
(nerve or sinew) for a nerve in the leaf. The theory that the key
to the understanding of plants was to be found in zoology, led
Aristotle to suppose that the roots of plants correspond to the
mouths of animals, since both roots and mouths are members by
means of which food is absorbed; the plant, on this view, is com-
parable with an animal standing on its head.
1
This idea can be
traced back to Democritus,
2
while Plato
3
speaks of man as like a
plant whose roots are not in the earth, but in the heavens. The
notion that the root corresponds to the mouth was handed on by
Boethius, who, in the sixth century A.D., wrote of plants, "as it
were, thrusting their head into the ground ",
4
while, even in the
sixteen hundreds, Bacon quotes with approval, "Homo est planta
inversa. "
5
As late as the mid-nineteenth century, a well-known
text-book
6
sponsored the crudely conceived analogy between root
and mouth. The author writes that "many insects support
selves wholely by suction; and ... all plants do the same". He
goes on to explain that plants are provided, "not with a single
sucker, like the leech or the flea, but with many". These suckers
he describes as taking the form of spongelets at the extreme tips
of the rootlets, which absorb 'carbonised water'.
Theophrastus, with his wider botanical knowledge, tended to
be more cautious than Aristotle about analogies between the
animal and vegetable worlds: he says that "we must not assume
that in all respects there is complete correspondence between
plants and animals'' ;
7
but with this reservation, he often turns to
1
Peck, A. L. (1937), pp. 370-1 [Depart. anim. 1v. x. 686bJ; Hett, W. S. (1935),
pp. 69, 89 [On the Soul, 11. i. 412b; II. iv. 416c].
2
Cornford, F. M. (1937), p. 357.
s Cornford, F. M. (1937), p. 353 [Timaeus, 90J.
4
Boethius, A. M. T. S. (1609), bk. II, xi, opp. p. 80; for a revised version of this
translation, see Stewart, H.F. and Rand, E. K. (1918), The Consolation of Philosophy,
pp. 280-1.
5
Bacon, F. (1631), p. 151; this edition quoted as the first is not accessible to the
writer.
6
Rennie, J. (1849). pp. 88, 92.
7
Hort, A. (1916), vol. 1, pp. 6, 7 [1. i. 4J.
12
THEOPHRASTUS
analogy for help, and he is indeed right in doing so. For human
thought can proceed only by passing from the known to the
unknown, on the assumption that there is some degree of real
analogy between the two.
1
It has been a certain handicap to the
study of plants, that, owing to man's primary interest in the
animal world, to which he belongs, his attention was naturally
first focused upon this, and plants were thus left to be interpreted
subsequently through zoological comparisons; the reverse
process might at times have been more illuminating.
The botanical work of Theophrastus is of such importance, as
the basis on which even to-day we still build, that it is necessary
to consider it in some detail, at least as regards those aspects
which bear definitely upon morphology. Our knowledge of his
output is derived from two books, De causis plantarum ( Tiepl q>vTwv
ah1wv), and De historia plantarum ( TTEpi q>VTWV icrTopio:). It has
been suggested
2
on internal evidence that De causis is relatively
early, while the Historia, in which the vocabulary shows greater
richness, precision, and specialisation, represents a group of later
works. The influence of Aristotle is conspicuous in De causis,
while the Historia shows more independence.
3
Both treatises
have some of the characteristics of lecture notes.
4
The fact that
neither of them is a fully integrated individual work, representing
a final written version of the author's views, does not add to the
ease of interpretation; but, even so, a gre(:lt part of their content
is lucid and of profound interest.
The Historia opens with an account of the general scope of
botany. Theophrastus tells us that we must take into considera-
tion: ( 1) the parts of plants; ( 2) their qualities; ( S) the way in
which their life originates; and ( 4) the course which their life
follows. He adds that behaviour and activities, such as we witness
in animals, are not to be found in plants. This classification, which
is remarkable for its comprehensiveness, might be restated in
modern terms as representing the division into ( 1) morphology;
(2) physiology and biochemistry; (s) the study of reproduction
1
Cf. Hort, A. (1916), vol. 1, pp. 18, 19 [1. ii. 4J; on analogy in the history of
science, see Arber, A. (1946b).
2
Stromberg, R. (1937), p. 136; for a careful chronological study of the botany of
Theophrastus, see Senn, G. (1933).
8
Senn, G. (1930), p. 113.
4
'Vorlesungsmanuskripten', Stromberg, R. (1937), pp. 69-70.
lS
II. MORPHOLOGY OF THE ARISTOTELIAN SCHOOL
and development; ( 4) the study of life-histories. Theophrastus
goes on to say that, of these subjects, that dealing with the parts
of plants (morphology) presents the greatest difficulty. Writing,
as he did, more than 2000 years ago, he was naturally not in
a position to make a critical comparison between the obstacles to
be overcome in these various regions of research, but his apprecia-
tion of the complexity of morphology indicates that this was the
field into which he had the clearest insight; it is indeed the field in
which his influence has been most lasting, while that of Aristotle
has been more deeply felt in the study of behaviour, and of
'purposeful' structure.
1
The difference of emphasis in the work of
these two pioneers possibly represents a natural cleavage between
the standpoint of the botanist and zoologist-the botanist inclin-
ing to the consideration of form, and the zoologist to that of
function.
We have already mentioned Aristotle's realisation of the
existence in the animal body of homogeneous parts (tissues),
which are the components of heterogeneous parts (organs).
Among plants, correspondingly, he recognised organs, but he
says that they are of a simpler type; he instances leaf and seed-
vessel.2 Theophrastus carries the analysis to a further point, and
classifies plant organs under two categories: the main parts, such
as root, stem, bough, and twig; and the annual parts, such as
flower, leaf, fruit, and 'new shoot'. On comparing the parts of
plants with those of animals, he is struck by the impermanence of
the plant members of the second category, and with the fact that
trees make fresh shoots every year, so that the number of parts
is indeterminate and continually changing, whereas in the higher
animals the number of organs is fixed and definite. He evidently
feels that this is a divergence which must be accepted as such,
without trying to force our conception of the plant into the frame-
work derived from zoology. He holds that "it is waste of time
to take great pains to make comparisons where that is impos-
sible" ;
3
this may seem to be a glimpse of the obvious, but in those
days the overstrained animal analogy had a stranglehold which
it is hard now to realise.
1
Cf. Thompson, D'Arcy W. (1913).
2
Hett, W. S. (1935), p. 69 [On the Soul, 11. i. 412bJ.
3
Hort, A. (1916), vol. 1, pp. 6, 7 [1. i. 4J.
14