Mnemosyne (2024) 1-23

brill.com/mnem

“Is the Embryo a Living Being?” (Aët. 5.15)

Embryology, Plants, and the Origin of Life in Presocratic Thought

Claudia Zatta
Department of Philosophy, University of Milan, Italy
[email protected]

Submitted 28 February 2023 | Accepted 25 August 2023 |

Published online 3 July 2024

Abstract

Building on previous studies, this essay discusses the use of embryological images and
analogies in Anaximander, Empedocles, Democritus, and Lucretius. It pursues their
intertextual connections arguing that in ancient philosophy embryology was not only
relevant for conceiving the early formation of the cosmos as has been claimed so far,
but that it also shaped the conception of the primeval rise of animal life and the living
processes of plants.

Keywords

embryology – life – animals – plants – Empedocles – Democritus – Lucretius

The veiled allusion to embryological doctrines in the Presocratics’ accounts

of the formation of the cosmos was at the center of scholarly attention many
decades ago. In a seminal study, titled “Embryological analogies in Pre-socratic
Cosmogony” published in 1932, Baldry identified in Anaximander’s cosmolog-
ical account elements of the conception of the embryo found in the initial
chapter of Hippocrates’ On the Nature of the Child, showing the hermeneutic
fruitfulness of the parallel between cosmogony and anthropogony and arguing
that knowledge about the embryo well precedes the work of the Hippocratic

Published with license by Koninklijke Brill bV | doi:10.1163/1568525X-bja10268

© Claudia Zatta, 2024 | ISSN: 0026-7074 (print) 1568-525X (online)
Downloaded from Brill.com 07/22/2024 12:39:01PM by [email protected]
via MRW Tester and BRILL
2 zatta

doctor.1 In line with Baldry, in the 1968 article “Embryological analogies in

Empedocles’ Cosmogony” Wilford has similarly identified traces of biological
thought in the early phases of Empedocles’ cosmogony. Wilford takes as a point
of reference for embryological doctrines the same chapter from Hippocrates’
On the Nature of the Child as Baldry but, differently from him, he focuses on
the phenomenon of embryonic respiration to argue that it was a model for
the Empedoclean cosmos. In Wilford’s reading, the dynamics of Love and
Strife at the cosmic level would reflect the phenomenon of respiration, which
is extensively documented in Empedocles’ fragment 100 and which, based on
the Hippocratic text, Wilford feels legitimized to assume also for Empedocles’
conception of the embryo.2 This is a problematic point to which I will return
later. Here let me stress that according to Wilford, in Empedocles’ doctrine, the
rhythmic exchange that enables the embryo to live would be applied to the cos-
mos to signify its alternating epochs, from the gradual aggregation of the four
elements into the Sphairos to their ultimate disaggregation under the domi-
nation of Strife. The parallels between embryology and cosmology discussed

1 Hippocrates’ text, quoted in the second section of this essay, belongs to an extensive discus-
sion on the embryo, from its conception to gestation and delivery, but embryology was a
shared item in the Presocratics’ agenda, and we can trace it back as early as Alcmaeon and
Parmenides. The first inquired about the formation of the embryo (fr. 24A14 Diels-Kranz /
Cens. 5.4), its nutrition (fr. 24A17 Diels-Kranz / Aët. 5.16.3) and sex determination (fr. 24A14
Diels-Kranz / Cens. 6.4), the latter, besides sex determination (fr. 28B17 Diels-Kranz / Gal.
In Hipp. Epid. 6.2.46, p. 119.12-15), discussed also intergenerational resemblances (fr. 26A54
Diels-Kranz / Aët. 5.11.2). Already Thales, however, may have held some interest in embryo-
logical questions if Aristotle’s testimony about the choice of water as arkhē is accurate. For
Thales would have held water as principle of all things based on the observation that the seed
of all things is moistened (fr. 11A12 Diels-Kranz / Arist. Met. 3.983b19-28).
2 In the words of Wilford 1968, 111: “Love and Strife are to the world what blood and air are to
the body, so that Love would expand from the centre and drive Strife out to the circumfer-
ence and beyond, just as blood does to air.” Besides Baldry and Wilford, another interpreta-
tive line that pursues the cosmological relevance of the embryological model involves the
revival of the theory of recapitulation according to which ontogeny recapitulates philogeny,
that is, the development of the embryo mirrors that of the species, see Wilderbing 2021,
131-134, who interprets the extant evidence regarding Anaximander and Empedocles in
these terms. We may certainly catch a glimpse of the recapitulation theory in Presocratic
doctrines, thus underscoring their relevance to modern biological thought, but I refrain
from pursuing this interpretive line because the Presocratic framework of inquiry was other
than that of modern biology. The interest, for instance, did not lie in the development of the
species per se, but in the rise of broad categories of living beings according to their habi-
tats. Even in Anaximander terrestrial life (human being included, see 12A30 Diels-Kranz /
Ps.-Plu. 5.9.14, discussed below) developed from the prōta zōa in water and the seemingly
evolutionary account of Empedocles’ zoogonies is in fact inserted in a larger cosmological
cycle that supersedes unidirectional species development.

10.1163/1568525X-bja10268 | Mnemosyne ( 2024) 1-23

Downloaded from Brill.com 07/22/2024 12:39:01PM by [email protected]
via MRW Tester and BRILL
“ is the embryo a living being? ” ( Aët. 5.15 ) 3

in these studies are not only interesting because they indicate that the world
itself is alive.3 They also document the inclination toward unifying the mode
of explanation for the origin, genesis, at the microcosmic and macrocosmic
level.4 What works in ordinary life for the birth of humans and other animals
must have been at work also for the rise of the cosmos qua living being. In
this essay I will expand on the discussion of the cosmogonic relevance of the
embryological model, showing, on the one hand, that this model lurks also,
within a cosmogonic framework, in Presocratic conceptions of the very incipit
of animal life, if only allusively in Anaximander, then, more explicitly in testi-
monies related to Democritus and in Lucretius.
On the other hand, however, I intend to pursue a new thread and discuss
how the embryological model is applied to understand the life of plants.
For this second aspect, I will focus mainly on Empedocles who, according to
Aëtius, investigated the origin of plants within a cosmogonic framework and
separately answered two similar, complementary questions, namely, whether
plants and embryos are living beings (zōa).5 True, these questions may well
result from the doxographic process and reflect the interests of later interpret-
ers who gauged the Presocratics’ ideas in light of successive doctrinal points,
particularly, in this case, Aristotle’s categorical distinction between plants and
animals and Plato’s idea that the embryo is a living being.6 Nevertheless, the
inquiry about whether embryo and plants are living beings creates a mean-
ingful network of references that help us reconstruct the nature of plants in
Empedocles, especially, with respect to the process of respiration. In the end,
I hope to show, whether implicitly or explicitly, that in Presocratic philosophy
embryological analogies were good for thinking not only about the early for-
mation of the cosmos as has been argued so far, but also about the primeval

3 In fragment 2, Anaximenes addressed explicitly the respiration of the world while more than
a century and a half later, in Timaeus, Plato responding to earlier studies of nature, will state
that the world is a living animal (zōon) endowed with a soul (30b). For a discussion of the
connection between biology and cosmology in these authors, see respectively Laks 2019,
12-14 and Sattler 2019, 29-45.
4 On the living being as the most important model for the cosmos, see Wright 1981, 56-74
(Chapter 4. Macrocosm and Microcosm); cf. Kahn 1960, 110.
5 These questions are brought up respectively in 5.26.4 and 5.15.3 and will be quoted in the
second section of this essay.
6 Aristotle and Plato are cited in Aëtius’ chapters dealing with the aforementioned questions.
For Aristotle’s denial of animal status to plants, see, for instance, Arist. de An. 2.2.413a20-413b1,
435a17-b3; cf. HA 7.588b7-11 on plants’ midway position between soulless (simple) bodies, on
the one hand, and the animals, on the other; for Plato’s belief that the embryo is alive, see
Ti. 91c-d where he calls the embryos ἀδιάπλαστα ζῷα.

Mnemosyne ( 2024) 1-23 | 10.1163/1568525X-bja10268

Downloaded from Brill.com 07/22/2024 12:39:01PM by [email protected]
via MRW Tester and BRILL
4 zatta

rise of animal life and the living processes of plants, conceived, themselves,
as the earliest forms of life.

1 Embryological Models and the Incipit of Animal Life

We can start considering the embryological model in the conception of the

origin of life by looking first at its resonances in Anaximander’s cosmogony.
A key passage accounts for the birth of the cosmos with a charged language:

φησὶ δὲ τὸ ἐκ τοῦ ἀιδίου γόνιμον θερμοῦ τε καὶ ψυχροῦ κατὰ τὴν γένεσιν τοῦδε
τοῦ κόσμου ἀποκριθῆναι καί τινα ἐκ τούτου φλογὸς σφαῖραν περιφυῆναι τῷ
περὶ τὴν γῆν ἀέρι ὡς τῷ δένδρῳ φλοιόν· ἧστινος ἀπορραγείσης καὶ εἴς τινας ἀπο-
κλεισθείσης κύκλους ὑποστῆναι τὸν ἥλιον καὶ τὴν σελήνην καὶ τοὺς ἀστέρας.

He (Anaximander) says that the seed of the warm and the cold, coming
from the eternal, was detached at the birth of this world and that a cer-
tain sphere of fire coming from this grew around the air surrounding the
earth like the bark around a tree. When this was torn away and enclosed
within certain circles, the sun, the moon, and the stars were formed.7

Whether they reflect or not Anaximander’s words, the reference to the seed
of hot and cold, the idea of separation expressed by the verb ἀποκριθῆναι, a
passive form generally adopted for the separation of the seed at the microcos-
mic scale,8 the use of περιφυῆναι to describe the formation of a sphere of fire
around the air and of φλοιός, here translated as ‘bark’, to qualify it—all these
terms signal a parallel with embryological doctrines.9 Φλοιός indicates the ten-
der bark of a tree (and hence of a growing tree)10 and by extension “the rind,
skin of fruit, pellicle of a leaf or egg”.11 In Aristotle, for instance, it is the mem-
brane enclosing the eggs of vipers which break (περιρρήγνυνται) after a short
period of time, letting the young out12 while the related form φλόος is used in

7 Fr. 12A10 Diels-Kranz / Ps.-Plu. Strom. 2. All text and translations of the Presocratics are
from the Loeb edition by Laks and Most.
8 In History of Animals, for instance, Aristotle uses it to indicate the distinct formation of
the body of the embryo in the female’s uterus (6.561a17).
9 See Baldry 1932; cf. Classen 1970 who further inquires after the overall nature of the bio-
logical parallel and whether it constituted a theory of change.
10 See, for instance, Hom. Il. 1.237, Thphr. HP 1.5.2.
11 See Beekes’ Etymological Dictionary of Greek s.v. φλοιός.
12 See Arist. HA 6.558a28; at 552a10 it refers to animals that break out from larvae.

10.1163/1568525X-bja10268 | Mnemosyne ( 2024) 1-23

Downloaded from Brill.com 07/22/2024 12:39:01PM by [email protected]
via MRW Tester and BRILL
“ is the embryo a living being? ” ( Aët. 5.15 ) 5

Nicander to designate the skin of humans and snakes alike.13 Given this range
of applications, it is not surprising that φλοιός appears in another testimony
about Anaximander to indicate the prickly skin enveloping the first living
beings, the prōta zōa, born in moistened earth heated by the sun:

Ἀναξίμανδρος ἐν ὑγρῷ γεννηθῆναι τὰ πρῶτα ζῷα φλοιοῖς περιεχόμενα ἀκανθώ-

δεσι, προβαινούσης δὲ τῆς ἡλικίας ἀποβαίνειν ἐπὶ τὸ ξηρότερον καὶ περιρρη-
γνυμένου τοῦ φλοιοῦ ἐπ᾽ ὀλίγον χρόνον μεταβιῶναι.

Anaximander states that the first animals were born in moisture, sur-
rounded by thorny bark, but as they increased in age they moved to
where it was drier, and when the bark burst open they changed their way
of life in a short time.14

The bark surrounding the first animals is analogous to the bark-like sphere of
fire surrounding the air in the nascent cosmos—the analogy is functional to
capture a similar embryological stage during the process of formation of both
living beings and the world,15 but the outcomes are different. If the sphere of
fire around the air breaks, giving rise, in the torn parts, to the sun, moon and
stars, the bark around the first animals will break completely and disappear
leading to the development of living beings suitable to live on earth—a diver-
gence that shows the circumscribed functionality of the embryological refer-
ence. Now if in Anaximander’s testimony regarding the rise of the first animals
the image of the embryo is conjured up merely through linguistic allusions,
namely, the terms φλοιός for the membrane and περιρρήγνυμι for its breaking,
in the tradition attributed to Democritus the embryological model is apparent:
at a specific cosmogonic stage, the earth produces embryo-like formations.
The relevant testimony is found in two different texts, i.e., Diodorus Siculus’

13 See, respectively, Nic. Alex. 302 (in which φλόος is applied to the skin of Marsyas, flayed by
Apollo) and Ther. 355, 392.
14 Fr. 12A30 Diels-Kranz / Ps.-Plu. 5.9.14. Gregory 2016, 29-36 advances the hypothesis that
this doctrinal tenet is based on the observation of the life-cycle of the Caddis fly and
indicates a process of metamorphosis rather than adaptation to a drier environment, thus
disconnecting the emergence of life from cosmogonic dynamics. Observation of the nat-
ural world may have well contributed to Anaximander and other Presocratics’ doctrinal
tenets but their reflections on the origin of life are essentially inscribed in the cosmologi-
cal framework and explained in terms of the arkhē, whether in monistic or pluralist terms
(see in this respect Aristotle’s description of the Presocratics’ inquiry in PA 1.640b5-13).
15 Kirk, Raven, and Schofield 1983, 142 remark on the similarities of the general principles
informing the two phenomena of birth: in both cases, the bark-like covering contains
moisture, which expands under the effect of the heat causing an explosion.

Mnemosyne ( 2024) 1-23 | 10.1163/1568525X-bja10268

Downloaded from Brill.com 07/22/2024 12:39:01PM by [email protected]
via MRW Tester and BRILL
6 zatta

History and Hermippus’ On Astrology.16 These texts provide a similar picture

for the origin of life in a cosmogonic framework, but they also differ in that
Diodorus focuses strictly on animal life, humans included, while Hermippus
comprises the origin of plants, hence offering a testimony that seems more
adherent to Democritus’ (and other Presocratics’) doctrines inasmuch as we
know that they investigated the origin of animals and plants from a cosmo-
gonic perspective.17 After the four elements have taken their place in the lay-
ered structure of the cosmos, with fire at the top, above air and water, and the
earth at the bottom, attention is directed to the effects of the sun on the recent
stratification of water and earth. Hermippus writes,

ἀλλ᾽ ἐπεὶ τὸ ὕδωρ ἐπὶ γῆς τὴν οἰκείαν χώραν ἐπέλαβεν, αὕτη δὲ διάβροχος
οὖσα τὴν οἰκείαν μορφὴν ὑπὸ τοῦ ἡλίου προσβάλλοντος καὶ πρὸς τὸ ξηρότε-
ρον μεθιστάντος κατὰ μικρὸν ἀπελάμβανε, φύονται δὴ πρῶτον οὕτω δένδρα τε
καὶ φυτὰ καί τινες ὑμένες ἐοικότες πομφόλυξιν, αἳ δὴ μεθ᾽ ἡμέραν μὲν ὑπὸ τοῦ
ἡλίου διαθερμαινόμεναι, νύκτωρ δὲ ὑπὸ τῆς σελήνης καὶ τῶν ἄλλων ἀστέρων
περιθαλπόμεναι χρόνῳ διαρραγεῖσαι τὰ ζῷα ἀπέτεκον.

Once the water had found its proper place on the earth, and that place,
being thoroughly moist, had got its own form from the sun which shone
on it and gradually dried it, first there came into being trees and some
membranes resembling vesicles; they were dried by the sun during the
day and warmed at night by the moon and the other heavenly bodies, and
they eventually burst and gave birth to the animals.18

16 These texts are included in the Diels-Kranz edition of the Presocratic philosophers
(respectively fr. 68B5 and 68B5.20) and paired together in Luria’s edition of Democritus
(515). Some scholars, however, have disputed the Democritean filiation of Diodorus’
account (see, for instance Cole 1990, 11 who prefers to speak about them in terms of tradi-
tional material rather than ultimate attribution). Similarly, Guthrie warns against a direct
attribution to Democritus noting many common points with Empedocles’ zoogonies
but claiming that even if the attribution is uncertain Diodorus still preserves Presocratic
material (1965, 210 n. 1). For Luria, however, the Democritean filiation of the account is
clear from Diodorus’ claim that the view on the origin he presents belongs to ‘those who
think that the world comes to be and perishes’ (D.S. 1.6.3; Luria ad 515), and this was in
fact a tenet held by Democritus (see, for instance, fr. 68A40 Diels-Kranz / Hippol. Haer.
1.13.2-3 and Guthrie 1965, 408).
17 See Arist. PA 1.640b5-13 where Aristotle summarizes the earlier physiologoi’s project into
nature referring to the doctrines of Empedocles, Anaxagoras, and Democritus. Note,
however, that Diodorus’ testimony could be inclusive of plants qua belonging to the
terrestrial animals.
18 Fr. 68B5.2 Diels-Kranz / Luria 515, 534 / Hermippus, On Astrology, 2.16-12. This and the fol-
lowing translation are by C.C.W. Taylor.

10.1163/1568525X-bja10268 | Mnemosyne ( 2024) 1-23

Downloaded from Brill.com 07/22/2024 12:39:01PM by [email protected]
via MRW Tester and BRILL
“ is the embryo a living being? ” ( Aët. 5.15 ) 7

Not all living beings come into being at the same time, but trees emerge first
and animals later. Both require a period of gestation. Trees are born after the
sun had gradually dried the moistened earth while animals require a longer
and differentiated period. They are born, Hermippus tells us, ‘from mem-
branes (hymenes) resembling vesicles (pompholyxin)’ that had formed in the
earth. The reference to embryological doctrines is evident: the term hymenes,
which may be considered analogous to Anaximander’s φλοιός, is used in medi-
cal literature for the membrane enveloping the embryo.19 Note also that the
similitude of the membranes with vesicles points at once to the physical pro-
cess that thickens the moistened earth and to the swollen membranes includ-
ing their contents20 and, further, that animals’ longer period of gestation
promotes their growth at the embryological stage. Two external forces are
distinguished: if the sun dries the embryos up, moon and stars warm them
up. These effects are complementary. The prefixes dia before thermainomenai
and peri before thalpomenai seem to suggest a difference in impact: the heat
of the sun affects the membranes with their content throughout while sun,
moon and sun affects them all around. Diodorus’ testimony fittingly integrates
Hermippus’ narrative by explaining, first, that the effects of the sun on the
moistened earth product pockets of fermentation (σηπεδόνες)21 covered by
thin membranes and, then, that the sun solidifies them while the nocturnal
mist nourishes them, thus pointing to the phenomenon of respiration and its
nutritive role which I will soon discuss.22 Finally, as for Anaximander’s first
animals, in the case of this testimony too, the enveloping surfaces break (περιρ-
ρήγνυμι) and the animals are born.

19 See, for instance, the discussion of the embryo in On the Nature of the Child (3 and pas-
sim). Further, for the atomists too we note an overlapping of terms and hence of con-
ceptions for the origin of the cosmos and the origin of life, particularly animal life. If
hymēn is used in the plural to account for the generation of animals from the earth in
Diodorus’ account, it also indicates in the singular the membrane forming around the
nascent cosmos and growing as it incorporates inside more and more atoms (D.L. 9.32;
see Guthrie 1965, 407-408, who however stresses that the atomists’ view was purely mech-
anistic: they assimilated animal formation to the nascent cosmos and not vice versa).
20 Significantly, the term pompholyx is used in the plural by Aristotle in On the Generation
of Animals within an analogy between the thickening of fluids composed of water and
earthy matter under the agency of heat and the foam qua composed of water and hot air
(pneuma) (Arist. GA 2.735b). In Hippocrates, the related πομφός indicates the blister on
the skin (Mul. 2.118; Morb. 2.70); LSJ s.v. πομφόλυξ and πομφός.
21 Cf. Pl. Phd. 96b in which Socrates, recalling a similar theory, refers to a sort of fermenta-
tion (σηπεδόνα τινά) as the phenomenon that brings about the formation of animals.
22 See D.S. 1.7.4 / Fr. 68B5 Diels-Kranz. In other words, the nourishment by the mist implies
that the embryonic organisms breathe, see below the discussion of the nutritive role
of respiration.

Mnemosyne ( 2024) 1-23 | 10.1163/1568525X-bja10268

Downloaded from Brill.com 07/22/2024 12:39:01PM by [email protected]
via MRW Tester and BRILL
8 zatta

Three and a half centuries after Democritus, the embryological model will
dominate Lucretius’ account of the origin of animal life in De Rerum Natura, a
poem based on atomistic theories and influenced by Empedocles.23 In book V,
after dealing with the origin of the cosmos, Lucretius accounts for the emer-
gence, first, of plants, and then, of animals, thus reflecting earlier chronologies
for the rise of life. He announces this novel phenomenon with a suggestive
expression, that is, novus fetus. We read,

Nunc redeo ad mundi novitatem et mollia terrae

Arva, novo fetu quid primum in luminis oras
Tollere et incertis crerint committere ventis.

Now I return to the youth of the world and the earth’s soft fields, and to
what in new birth (novo fetu) they decided first to bring into the shores of
light, and entrust to the changeable winds.24

Here fetus, or—to use the later and now common spelling—foetus has been
translated as ‘birth.’ In other editions it is rendered as ‘parturition,’ ‘parto’, and
‘productions nouvelles’25 but, literally, it means ‘embryo.’ As Campbell notes,
in Origines Isidorus defines foetus as ‘what is nourished in the womb’.26 And as
in Empedocles brephos, one of the corresponding Greek terms for foetus (the
other being embryon), may mean either the organism inside the mother’s body
or the new-born baby,27 so in Lucretius foetus seems to preserve an intrinsic
ambiguity. For, on the one hand, it may be taken as, so to speak, resultative, thus
foreshadowing the accomplished emergence of all the life-forms (plants and
animals qua fully-formed bodies)28 but, on the other, it fittingly harmonizes
with the very mode of animals’ emergence accounted for by Lucretius, that is,
from wombs rooted in the earth. At one stage of their formation, in Lucretius’
doctrine animals were indeed literally embryos enclosed in the earth-wombs.
Thus, in this parallel reading, the expression ‘new foetus’ would hypostasize

23 For Empedocles’ philosophical influence on Lucretius, see Furley 1970.

24 Lucr. 5.780-782; all translations of the De Rerum Natura are by G. Campbell.
25 See, respectively, the translations by H.J. Munro in the Loeb edition, Marchetti in
Sonzogno, and Ernout in the Belles Lettres. Bailey translates novus fetus even in more
abstract terms as ‘with new power of creation.’
26 Isid. Orig. 11.1.136 (Barney, Lewis, Beach and Berghof 240); see Campbell 2003, 45.
27 See Zatta 2023, 256 and n. 18.
28 In this respect, Campbell notes that in Lucretius and elsewhere foetus is a poetic meta-
phor for the fruit of the earth (2003, 45), hence holding an extensive meaning that applies
to plants and animals alike.

10.1163/1568525X-bja10268 | Mnemosyne ( 2024) 1-23

Downloaded from Brill.com 07/22/2024 12:39:01PM by [email protected]
via MRW Tester and BRILL
“ is the embryo a living being? ” ( Aët. 5.15 ) 9

the process of emergence of life in its different forms assimilating the sponta-
neous growth of plants with that of the animals. Significantly, in the lines that
follow, Lucretius insists repetitiously on the origin of plants and animals from
the earth hence stressing its role of mother while excluding competing physi-
cal theories, namely that animals fell from the sky or rose from salt pools.29
By these theories he may be referring respectively to Anaxagoras who claimed
that the air contains the seeds of all things30 and to Anaximander, who con-
sidered the first animals to be ‘fishes or similar to fishes.’31 By contrast, for
Lucretius plants grew from the body of the earth in the same way as ‘feath-
ers, hairs and bristles grow first on the limbs of four-footed creatures and the
birds’, while the first animals emerged from it in the same way as they do in
present-day phenomena of spontaneous generation.32 Regarding the earlier
formation of plants, Waszink remarks that, on the one hand, Lucretius echoes
Empedocles’ doctrine in which plants are the first living beings to have been
born from the earth, but that, on the other, he follows Epicurus33 for whom
plants are not empsykha and hence not members of the animal club34—hence,
we may add, the difference in their emergence with respect to Hermippus’
account quoted earlier.35 For while both plants and animals originate from the
earth, in Lucretius plants emerge spontaneously (i.e., like feathers and hairs)
while animals require a complex gestation. He writes,

tum tibi terra dedit primum mortalia saecla.

multus enim calor atque umor superabat in arvis.
Hoc ubi quaeque loci regio opportuna dabatur,
crescebant uteri terram radicibus apti;
quos ubi tempore maturo patefecerat aetas
infantum, fugiens umorem aurasque petessens,

29 Lucr. 5.793-794.
30 See fr. 59A117 Diels-Kranz / Thphr. HP 3.1.4.
31 Fr. 12A30 Diels-Kranz / Hippol. Haer. 1.61.6. 6-7. For Anaximander’s conception of the
origin of life from sea water, see, for instance, Cornford 1952, 170, Loenen 1954, 222, and
Kahn 1960, 113, although, based on the Hippocratic De Victu, this author seems to trace a
line of influence in explanation from the cosmos to the living being rather than vice versa.
32 See Lucr. 5.783-791 and 797-798. These different proofs account for the reliability of Lucre­
tius’ account targeting at the same the creationist model for the origin of life as that found
in Plato’s Timaeus and rebutted by Epicurus; see Campbell 2003, 41.
33 Waszink 1964, 49.
34 See Aët. 5.26.3.
35 Recall that in Hermippus’ testimony, plants still required a period of gestation (even if
shorter than animals) while, as will we see in the second section of this essay, Empedocles
considered plants ‘embryos in the earth.’

Mnemosyne ( 2024) 1-23 | 10.1163/1568525X-bja10268

Downloaded from Brill.com 07/22/2024 12:39:01PM by [email protected]
via MRW Tester and BRILL
10 zatta

convertebat ibi natura foramina terrae

et sucum venis cogebat fundere apertis
consimilem lactis, sicut nunc femina quaque
cum peperit, dulci repletur lacte.

It was at that time, you see, that the earth gave birth first to mortal races,
for great heat and moisture abounded in the field. So, wherever a suitable
place was given, wombs grew fastened to the earth by roots, and when in
due time the age of the infants had broken these open, fleeing the mois-
ture and seeking the breezes, nature redirected there pores of the earth
and forced a juice like milk to flow from open veins, just as now each
woman when she has given birth is filled with sweet milk.36

Earth, moisture, and heat conjure up the traditional scenario favorable for the
origin of life. And as in Diodorus’ account, only certain spots provide an opti-
mal condition for the formation of animals, which are born from wombs rooted
in the earth. Crescebant uteri terram radicibus apti, Lucretius tells us, resorting
to a “strange idea” as Bailey defined it. Wright finds an antecedent to this imag-
ery in Empedocles’ οὐλοφυεῖς (the whole-natured).37 In fragment 62, to which
I will return in the second part of this essay, we learn indeed about a zoogony
of males and females, which sprung from the earth driven up by fire desiring
to reach the celestial fire and possessed an inarticulate body, without repro-
ductive organs and the capacity to speak. Empedocles calls these living beings
‘nocturnal saplings’ (ἐννύχιοι ὅρπεκες), literally ‘saplings in the night’ alluding
with such an adjective to the fact that they sprung up in a nightlike environ-
ment without sun. Hence this generation of living beings, the οὐλοφυεῖς, lived
in conditions very similar to plants, which, we learn from Aëtius in a chapter
dedicated to the question whether plants are animals, were for Empedocles the
first living beings to spring up from the earth at a time when the sun had not
started yet its rotation and day and night were not yet separated.38 The com-
monality between Empedocles’ conception of the whole-natured and plants,
on the one hand, and Lucretius’ rooted wombs, on the other, lies in their vis-
ceral connection to the earth from which they are born and from which they
derive their nourishment through the roots whereby they grow.39 At the same

36 Lucr. 5.805-814, transl. by G. Campbell.

37 Wright 1981, 217.
38 Fr. 31A70 Diels-Kranz / Aët. 5.26.4.
39 Regarding Lucretius’ rooted wombs, Schrijvers 1974, 256-260 speaks of a fusion between
vegetal life (roots) and animal and human life (wombs), stressing the connection of
Lucretius’ idea with Greek biological thought and further pursuing the analogy with

10.1163/1568525X-bja10268 | Mnemosyne ( 2024) 1-23

Downloaded from Brill.com 07/22/2024 12:39:01PM by [email protected]
via MRW Tester and BRILL
“ is the embryo a living being? ” ( Aët. 5.15 ) 11

time, Empedocles spoke of plants as embryos in the earth40—a claim that will
be discussed extensively in the next section. That plants and embryos shared a
similar mode of life was a Greek scientific idea. The botanic excursus in On the
Nature of the Child (22-27) articulates a set of parallels between the nutrition
and growth of plants at the early stage of formation and those of the human
embryo while in On the Generation of Animals Aristotle will explicitly equate
umbilical cord and roots based on their respective roles in the nutrition of the
embryo and the trees.41 Lucretius’ rooted wombs are built on similar grounds
although from the perspective of the earth as the mother (and, we may add,
endorsing the ontological distinction between plants and animals).42 For his
attention lies on the numerous wombs that the earth qua mother provides
rather than the embryos the wombs contain. The wombs’ roots, then, are anal-
ogous to that part in the female body that allows the nourishment to reach the
embryo. Interestingly, we may find here an echo of Democritus’ theory on the
prolific nature of pigs and dogs, which he explained in terms of the presence of
many wombs passible of receiving seed.43 Be that as it may, while Empedocles’
whole-natured and actual plants continue to nourish themselves through their
roots during their silent existence, Lucretius’ rooted wombs break out giving
birth to various species of mammals which find a suitable form of nourish-
ment in the juice-like milk springing from the earth.

2 Embryos and the Life of Plants

I turn now to the second part of my essay addressing the reference to the embryo
as a tool to understand the life of plants and particularly the phenomenon

plants also when accounting for the nutrition of the newborns by means of the milk-like
juice flowing from the earth. Still, if the generation of animals is fused with that of
plants, at the same time Lucretius separates animal and vegetal life, hence holding an
approach contrary to Presocratic doctrines which considered animals and plants in terms
of continuity.
40 Fr. 31A70 Diels-Kranz / Aët. 5.26.4.
41 See for instance Arist. GA 745b24-30. For Democritus, we may add, the navel was the first
part to form and served as ‘an anchorage against tossing and wandering, a cable and a rope
for the fruit, engendered and future’ (fr. 68B148 Diels-Kranz). Thus, Democritus seem-
ingly excludes that the navel may have provided an organ of nutrition since Democritus
also believed that the embryo got its nourishment by suckling (fr. 68A144 Diels-Kranz /
Aet. 5.16.1).
42 See Waszink 1964, 53 who, based on a discussion of the sources, argues that the rooted
wombs are Lucretius’ own visual device to account for an invisible phenomenon.
43 Fr. 68A151 Diels-Kranz / Ael. Nat. An. 12.16.

Mnemosyne ( 2024) 1-23 | 10.1163/1568525X-bja10268

Downloaded from Brill.com 07/22/2024 12:39:01PM by [email protected]
via MRW Tester and BRILL
12 zatta

of vegetal respiration.44 The focus will be on Empedocles but I will start with
remarking on the Presocratics’ pervasive interest in plants, their bodies, and
their lives. Whether they are simply alluded to or discussed explicitly, plants
are often referred to in a framework that considers them in analogy and conti-
nuity with other forms of life. Already in Anaximander we saw how the emer-
gence of the world and the first animals involved the formation of a bark-like
membrane covering a growing body. Empedocles, as mentioned earlier, con-
ceived a generation of males and females emerging from the earth and stretch-
ing upwards similarly to trees. They are offshoots (ὅρπηκες):

νῦν δ᾽ ἄγ᾽, ὅπως ἀνδρῶν τε πολυκλαύτων τε γυναικῶν

ἐννυχίους ὅρπηκας ἀνήγαγε κρινόμενον πῦρ,
τῶνδε κλύ᾽· οὐ γὰρ μῦθος ἀπόσκοπος οὐδ᾽ ἀδαήμων.
οὐλοφυεῖς μὲν πρῶτα τύποι χθονὸς ἐξανέτελλον,
ἀμφοτέρων ὕδατός τε καὶ εἴδεος αἶσαν ἔχοντες·
τοὺς μὲν πῦρ ἀνέπεμπε θέλον πρὸς ὁμοῖον ἱκέσθαι,
οὔτε τί πω μελέων ἐρατὸν δέμας ἐμφαίνοντας
οὔτ᾽ ἐνοπὴν οἷον τ᾽ ἐπιχώριον ἀνδράσι γυῖον.

Come then: how fire, separating off, drew upward the nocturnal sap-
lings of much-weeping men and women—Hear this. For my tale is not
aimless nor ignorant. First, whole-natured outlines sprang up from the
earth, possessing a share of both, of water as of heat. These, fire sent
upward, wishing to reach what was similar to it; as yet they displayed
neither the lovely framework of limbs, nor the voice and the organ that is
native to men.45

The nocturnal existence and wholeness of these living beings, combined with
their lack of sexual differentiation, are aspects common with plants, since,
on the one hand, in Empedocles’ doctrine, plants too were born at night46
while, on the other, in combining the male and female sexes, they too were
whole beings.47 Democritus considered plants upside-down animals holding
their heads underground, thus crafting a metaphor implying that plants’ roots

44 For plants’ respiration in Presocratic doctrines (Empedocles, Menestor of Sybaris and

Democritus) see Zatta 2023, 251-272. While building on this study, the line of inquiry
I pursue here is different in that I focus on embryological analogies in ancient Greek phi-
losophy and engage with the Hippocratic On the Nature of the Child.
45 (Fr. 31B62 Diels-Kranz / Simp. in Ph. 381.31-382.3 (and passim).
46 See n. 40 above.
47 See fr. 31A70 Diels-Kranz / Aët. 5.26.4, quoted below.

10.1163/1568525X-bja10268 | Mnemosyne ( 2024) 1-23

Downloaded from Brill.com 07/22/2024 12:39:01PM by [email protected]
via MRW Tester and BRILL
“ is the embryo a living being? ” ( Aët. 5.15 ) 13

were equal to animals’ heads and pointing to the function of nutrition of both
these parts.48 Plato called the plant a living being rooted in the earth, endowed
with the appetitive soul and hence of sensations and desire, although he also
claimed that the sensations of plants are ‘other’, allai, with respect to those
of animals.49 Given this network of body analogies and shared functions, it
is logical that the Presocratics may have found a common ground also for
the reproduction of plants and animals prior to Hippocrates and Aristotle
cited earlier. For instance, Alcmaeon, a doctor, traced a firm correspondence
between plants and human beings’ bodies with their respective reproductive
systems. As a tree’s blooming indicates its readiness to produce seed, so does
the growth of hairs in the adolescent.50 But it is Empedocles that is particularly
relevant to our discussion. In the same chapter about the question whether
plants are animals and right after reporting their primeval origin before the
sun had come into existence, Aëtius adds that,

διὰ δὲ συμμετρίαν τῆς κράσεως τὸν τοῦ ἄρρενος καὶ τοῦ θήλεος περιέχειν
λόγον· αὔξεσθαι δ᾽ ἀπὸ τοῦ ἐν τῇ γῇ θερμοῦ διαιρομένου, ὥστε γῆς εἶναι μέρη,
καθάπερ καὶ τὰ ἔμβρυα τὰ ἐν τῇ γαστρὶ τῆς μήτρας μέρη.

because of the equilibria of their mixture, they [i.e. trees] include the
principle of the male and that of the female. They grow from the heat
in the earth that separates, so that they are parts of the earth, just as
embryos in the belly are parts of the womb.51

Just as at the outset of the world, so now, plants emerge and grow because
of the heat in the earth that separates and thus becomes an integral part of
their bodies. But plants will never separate from their mother to which they
remain attached, holding throughout their lives a visceral connection with it.
They are like embryos in the mother’s belly. The difference with the other ani-
mals (of the fourth zoogony)52 is that the latter have developed from an embry-
onic stage and fully separated from their mothers while plants live thanks to
that connection: they are like permanent embryos in the mother’s belly. The
Empedoclean model for conceiving plants may be fruitfully discussed in terms

48 Fr. 68B5.2 Diels-Kranz / Hermip. De astrol. 2.1.11.

49 Pl. Ti. 76e-77c.
50 Fr. 24A15 Diels-Kranz / Arist. HA 1.581a14-16.
51 Fr. 31A70 Diels-Kranz / Aët. 5.26.4.
52 Empedocles conceives four different zoogonies of plants and animals, among which only
the creatures of the fourth one stop being born from earth and water and reproduce sexu-
ally; see fr. 31A72 Diels-Kranz / Aët. 5.19.5.

Mnemosyne ( 2024) 1-23 | 10.1163/1568525X-bja10268

Downloaded from Brill.com 07/22/2024 12:39:01PM by [email protected]
via MRW Tester and BRILL
14 zatta

of the passage from the Hippocratic On the Nature of the Child on which Baldry
and Wilford have based their parallels between embryology and cosmogony in
Presocratic doctrines, discussed at the beginning of this essay, and to which it
is finally time to revert. The Hippocratic doctor writes,

Ἢν ἡ γονὴ μείνῃ ἀπ᾽ἀμφοῖν ἐν τῇσι μήτρῃσι τῆς γυναικός, πρῶτον μὲν μίσγεται
ὁμοῦ, ἅτε τῆς γυναικὸς οὐκ ἀτρεμεούσης, καὶ ἀθροίζεται καὶ παχύνεται θερμαι-
νομένη. ἔπειτα πνεῦμα ἴσχει, ἅτε ἐν θερμῷ ἐοῦσα, τῆς μητρὸς πνεούσης, ἔπειτα
δὲ τοῦ πνεύματος ὅταν πλησθῇ, ὁδόν οἱ αὐτὸ ἑωυτῷ ἔξω ποιέει καὶ κατὰ μέσον
τῆς γονῆς τὸ πνεῦμα ἔξεισιν· ὅταν δὲ ὁδὸς γένηται τῷ πνεύματι ἔξω θερμῷ ἐόντι,
αὖτις ἕτερον ψυχρὸν εἰσπνέει ἀπὸ τῆς μητρὸς καὶ τοῦτο διὰ παντὸς τοῦ χρόνου.
θερμαίνεται μὲν γὰρ ἅτε ἐν θερμῷ ἐοῦσα· ψυχρὸν δ’ ἴσχει ἀπὸ τῆς μητρὸς πνε-
ούσης· πάντα δὲ ὁκόσα θερμαίνεται πνεῦμα ἴσχει. τὸ δὲ πνεῦμα ῥήγνυσι καὶ
ποιέει οἱ ὁδὸν αὐτὸ ἑωυτῷ καὶ χωρέει ἔξω· αὐτὸ δὲ τὸ θερμαινόμενον ἕλκει ἐς
ἑωυτὸ αὖτις ἕτερον πνεῦμα ψυχρὸν διὰ τῆς ῥαγῆς, ἀφ᾽οὗ τρέφεται.

If seed from both (sc. parents) remains in a woman’s uterus, first this is
mixed together—since the woman does not remain still—and then on
being warmed it aggregates and becomes thicker. Then, owing to the fact
that it is a warm place, it takes in breath when the mother breathes, and
then, when it is filled with breath, the breath forms a passageway for itself
to the outside, and passes out through the middle of the seed. When the
passageway out for the warm breath has been formed, the seed draws
in fresh, cold breath from the mother, and goes on doing this through
time. For the seed is warmed because it is in a warm place, and it acquires
cold breath from the mother’s breathing. In fact, all things that are
warmed take in breath: and the breath causes a tear, forms a path for
itself and then passes out again, and the thing that is warmed draws
fresh, cold breath to itself through the tear and from this it is nourished.53

This passage aims at explaining how the embryo begins to grow by identifying
the key mechanism by which it introduces some nourishment. Such mecha-
nism consists in the process of respiration as the first food is air, the nutritive
properties of which are stressed also in the Hippocratic treatise On Nutrition,
although in this context the respiration at stake takes place through the lungs
(48). On the Nature of the Child focuses, instead, on the first intake of air by the
embryo at a time when the lungs are not formed yet and claims that the succes-
sive exit of air articulates the embryo by opening a passageway, thus enabling

53 Hp. Nat. Puer. 1, transl. by P. Potter.

10.1163/1568525X-bja10268 | Mnemosyne ( 2024) 1-23

Downloaded from Brill.com 07/22/2024 12:39:01PM by [email protected]
via MRW Tester and BRILL
“ is the embryo a living being? ” ( Aët. 5.15 ) 15

a continuous exchange with breath coming in and breath going out. It is the
embryo’s heat that triggers the first intake of air supporting throughout the
process of breathing. This heat is not intrinsic to the embryo but depends on its
position inside the mother’s body. In other words, the embryo possesses heat
because of its warm location. And it breathes taking in air when and because
the mother breathes. In his study of this treatise Lonie has remarked how it is
liable to contain traces of Presocratic doctrines, namely of those of Anaxagoras,
Empedocles and Democritus,54 who, incidentally, were also among those most
interested in plants.55 And while for the passage just quoted, we do not have
explicit references to any of these authors, it is legitimate to ask whether it may
reflect Empedocles’ doctrine and illuminate his conception of plants, particu-
larly with reference to whether they were living beings that breathed.56 For if
plants are like embryos in the earth and the embryo is breathing, do plants,
also, breathe? And, furthermore, if the embryo breathes through the mother’s
breathing as we read in On the Nature of the Child, does this happen to plants
too? Do they breathe in unison with the earth?
In the pages that follow, I will attempt to answer the first of these questions.
I will leave aside the second because it would imply to argue for the respiration
of the earth, which lies out of the scope of this essay, but regarding which let
me remark that in fragment 100 Empedocles claimed that ‘all things breathe’
(πάντα ἀναπνεῖ) and that in fragment 55 ‘the sea is the sweat of the earth,’ thus
alluding to a form of terrestrial perspiration. As for the question of plants’ res-
piration based on the analogy with the embryo, we find a seemingly negative
piece of evidence because Empedocles denied that the embryo had the capac-
ity to breathe. The relevant source comes from Aëtius in a chapter dedicated
to the question of whether the embryo is a living being (zōon). Aëtius writes,

Ἐμπεδοκλῆς μὴ εἷναι μὲν ζῷον τὸ ἔμβρυον ἀλλ᾽ἄπνουν ὑπάρχειν ἐν τῇ γαστρί·

πρώτην δ᾽ ἀναπνοὴν τοῦ ζῷου γίνεσθαι κατὰ τὴν ἀποκύησιν, τῆς μὲν ἐν τοῖς
βρέφεσιν ὑγρασίας ἀποχώρησιν λαμβανούσης, πρὸς δὲ τὸ παρακενωθὲν ἐπεισ­
όδου τοῦ ἐκτὸς ἀερώδους γινομέμης εἰς τὰ παρανοιχθέντα τῶν ἀγγείων.

Empedocles (says that) the embryo is not a living being but exists without
breathing in the womb. The living being’s first breath occurs (he says) at

54 Lonie 1969, 392.

55 See Arist. PA 1.640b12-14.
56 This question is even more legitimate if we think that one of the a priori principles of
On the Nature of the Child is that like is attracted to like, which was also at the core of
Empedocles’ doctrine. Recall the explanation of the οὐλοφυεῖς’ stretching toward the sky
as due to the terrestrial fire longing to reach the celestial one.

Mnemosyne ( 2024) 1-23 | 10.1163/1568525X-bja10268

Downloaded from Brill.com 07/22/2024 12:39:01PM by [email protected]
via MRW Tester and BRILL
16 zatta

the time of birth, when the moisture in the new-born babies is excreted
and in the space (thus) vacated entry of the external air occurs in the ves-
sels that have opened.57

From this passage respiration emerges as the hallmark of life. Empedocles

denies that the embryo is a living being because it does not breathe inside the
womb but starts doing so at the moment of its delivery as a newborn. We do
not possess further explanation for this claim. Perhaps, differently from the
Hippocratic doctor of the On the Nature of the Child, Empedocles did not see any
connection between the embryo and the mother’s breathing and merely con-
ceived it constrained in her body. In this respect, in Parts of Animals Aristotle
notes that for Empedocles ‘many of the characteristics which animals have are
due to some accident in the process of formation.’ For instance, the vertebrae
of the backbone originate when ‘the fetus gets twisted and so the backbone
is broken into pieces.’58 Presumably we may add, due to the pressure on the
embryo from the mother’s body, a process which seems to indicate an environ-
ment unfavorable to respiration. Jaeger tried to salvage the life of the embryo
in Empedocles’ doctrine by claiming that Empedocles denied that respira-
tion through the lungs was possible for the embryo but postulated respiration
through the skin pores instead, and, as we saw earlier, Wilford endorsed this
interpretation to argue for the cosmogonic dynamics in Empedocles as reflect-
ing his presumed doctrine of embryonic respiration.59 To support Empedocles
conception that the embryo breathes, however, seems problematic and not
only because it is made ex silentio. For in another chapter dedicated to the
phenomenon of respiration, still from Aëtius, Empedocles describes the first
breath of the first living being (πρώτη ἀναπνοὴ πρώτου ζώου) offering a state-
ment which seems to harmonize with his denial of life to the embryo. We read,

Ἐμπεδοκλῆς τὴν πρώτην ἀναπνοὴν τοῦ πρώτου ζῴου γενέσθαι τῆς ⟨μὲν⟩ ἐν
τοῖς βρέφεσιν ὑγρασίας άποχώρησιν λαμβανούσης, πρὸς δὲ τὸ παρακενωθὲν
ἐπεισόδου τοῦ ἐκτὸς ἀερώδους γινομένης εἰς τὰ παραχνοιθέντα τῶν ἀγγείων·
τὸ δὲ μετὰ τοῦτο ἤδη τοῦ ἐμφύτου θερμοῦ τῇ τὸ ἐκτὸς ὁρμῇ τὸ ἀερῶδες ὑπα-
ναθλίβοντος, τὴν ἐκπνοήν, τῇ δ᾽εἰς τὸ ἐντὸς ἀνθυποχωρήσει τῷ ἀερώδει τὴν
ἀντεπείσοδον παρεχομένου, τὴν εἰσπνοήν.

57 See Aët. 5.15.3; all translations of Aëtius are from the Mansfeld and Runia edition.
58 Arist. PA 1 640a20-24 and fr. 31B97 Diels-Kranz (Ἐμπεδοκλῆς οὐκ ὀρθώς εἴρηκε λέγων ὑπάρ-
χειν πολλὰ τοῖς ζῴοις διὰ τὸ συμβῆναι οὕτως ἐν τῇ γενέσει, οἷον καὶ τὴν ῥάχιν τοιαύτην ἔχειν ὅτι
στραφέντος καταχθῆναι συνέβη).
59 Jaeger 1963, 216 n. 1.

10.1163/1568525X-bja10268 | Mnemosyne ( 2024) 1-23

Downloaded from Brill.com 07/22/2024 12:39:01PM by [email protected]
via MRW Tester and BRILL
“ is the embryo a living being? ” ( Aët. 5.15 ) 17

The first inhalation of the first living being came about when the liquid in
the brephesin60 withdrew and the external airy element entered from the
outside via the opening of the vessels into the empty spaces that resulted.
Afterwards, exhalation [scil. came about] when the innate heat expelled
the airy element by its impulse outward, inhalation when it provided the
airy element with a reverse passage inward by withdrawing in the oppo-
site direction.61

If indeed the embryo were a living being and breathing, then, it seems,
Empedocles would not have spoken of the ‘first breath’ of the newborn, for the
latter was breathing also in a prior stage. And if one could object, along with
Jaeger, that in this passage Empedocles is referring to the respiration through
the lungs, and not the skin pores, then it should be remarked that the respira-
tion at stake is that of the first living beings, the prōta zōa, which in Empedocles’
doctrine were the plants. For we have seen earlier that plants were the first liv-
ing beings to have emerged when the sun had not started its rotation and no
alternation of day and night existed yet. Aëtius’ testimony must refer to the
respiration through the skin pores in line with the phenomenon described in
fragment 100 in which respiration is presented as pertaining to the surface of
the body. We read,

ὧδε δ᾽ ἀναπνεῖ πάντα καὶ ἐκπνεῖ· πᾶσι λίφαιμοι

σαρκῶν σύριγγες πύματον κατὰ σῶμα τέτανται,
καί σφιν ἐπὶ στομίοις πυκναῖς τέτρηνται ἄλοξιν
ῥινῶν ἔσχατα τέρθρα διαμπερές, ὥστε φόνον μέν
κεύθειν, αἰθέρι δ᾽ εὐπορίην διόδοισι τετμῆσθαι. 5
ἔνθεν ἔπειθ᾽ ὁπόταν μὲν ἀπαΐξῃ τέρεν αἷμα,
αἰθὴρ παφλάζων καταΐσσεται οἴδματι μάργῳ,
εὖτε δ᾽ ἀναθρῴσκῃ, πάλιν ἐκπνέει.

It is in this way that all [scil. probably: living beings] inhale and exhale:
for all
Channels of flesh, which the blood leaves, extend to the surface of their
bodies;
And at the openings, the furthest limits of their skin (rhinôn)
Are perforated through and through with dense furrows, so that the
blood

60 Given the ambiguity of the term βρέφος, discussed below, I leave it transliterated.
61 Fr. 31A74 Diels-Kranz / Aët. 4.22.1.

Mnemosyne ( 2024) 1-23 | 10.1163/1568525X-bja10268

Downloaded from Brill.com 07/22/2024 12:39:01PM by [email protected]
via MRW Tester and BRILL
18 zatta

Lies hidden, while easy access is cut by these passages for

the air (aithêr) 5
When from here the delicate blood then rushes backward,
The air (aithêr), boiling, rushes after it in a raging surge.62

In this fragment Empedocles illustrates the phenomenon of respiration, which

involves all things, by describing the process in the blooded animals. As the
blood recedes from the surface of the body inwards, the air enters through the
pores which the receded blood had left empty, while, subsequently, the air
is expelled as the blood advances to the surface of the body. This oscillatory
movement is emblematic also for the process of respiration of the living beings
which do not possess blood. For since all, he tells us, inhale and exhale, those
creatures who do not have blood must possess an analogous part playing the
same role which blood plays in blooded animals. Significantly, in the testimony
about the first breath of the first living being, Empedocles explains the respira-
tory process in similar terms as in fragment 100, that is, as an oscillatory move-
ment but introduces, also, a revealing difference. Now Empedocles does not
speak of blood but more generally of liquid, ὑγρασία, thus mentioning the part
of the vegetal bodies that is equivalent to blood. In this respect, Theophrastus’
History of Plants tells us that the ancient Greeks used different terms to iden-
tify the moisture (ὑγρόν) of trees: if some called it juice (ὀπός) and others tear
(δάκρυον), still others, presumably including Empedocles, referred to it with-
out a specific name.63 Besides, Aëtius’ testimony also offers us another piece
of information which is missing from fragment 100 and which contributes to
our reconstruction of the phenomenon of respiration in terms of what trig-
gers it. For if fragment 100 simply describes this phenomenon as an oscillatory
movement between internal blood and external air focusing on the necessary
condition of a vacuum in the body (i.e., in the channels through which the
air is taken in when the blood recedes and is, then, expelled when the blood
advances), now we are told that what triggers the expulsion of air, for the living
being’s first breath and arguably the successive ones, is the body’s internal heat
(θερμόν). This seems an important clue for the comprehension of plants’ respi-
ratory process in Empedocles’ doctrine. Note that for this author plants had a
contrasting elemental constitution. The upper part, the one stretching toward
the celestial fire, was composed by fire while the part below, stretching into
the earth, was composed by earth. It was by resorting to this composite nature

62 Fr. 31B100 Diels-Kranz / Arist. Resp. 7.473b9-474b16.

63 See fr. 32A2 Diels-Kranz / Thphr. HP 1.2.3. Empedocles, however, did use ὀπός, but in a
technical way as referring to the thick sap of the fig tree (fr. 31B30 Diels-Kranz).

10.1163/1568525X-bja10268 | Mnemosyne ( 2024) 1-23

Downloaded from Brill.com 07/22/2024 12:39:01PM by [email protected]
via MRW Tester and BRILL
“ is the embryo a living being? ” ( Aët. 5.15 ) 19

that Empedocles, we learn from a passage in Aristotle’s On the Soul, was able
to explain the peculiar phenomenon of vegetal growth extending at the same
time toward opposite directions, upwards and downwards.64 The fire of plants’
upper part was stretching towards its like in the sky while the earth in the lower
parts stretched toward its like in the earth.65 So, bringing together the pieces of
evidence discussed so far, if for Empedocles all things inhale and exhale, and if
plants are like embryos in the earth, which do not breathe, but still contain fire
in the upper part of their body, and if fire is what triggers the expulsion of air,
then plants must breathe with the upper part of their body, which is exposed
to the aerial environment, and not with the part underground. As for the pre-
cise moment in which plants start to take in air and breathe, it can be grasped
by considering Empedocles’ discussion of the embryo in Aëtius’ testimony,
quoted earlier. Embryos, we have seen, start breathing and thus living only at
the moment of their delivery as newborns and analogously to them, plants
must start breathing as soon as they start emerging from the earth (although
they will never be delivered).
In claiming that the embryo does not breathe, Empedocles seems to have
endorsed a different model than the one presented by the Hippocratic doctor
in On the Nature of the Child. In his doctrine, plants’ partial embodiment in
the earth did not constitute a condition for the breathing of their lower parts,
the earth did not confer heat to the roots because its fire became constitutive
of their upper bodies. Still, the analogy between the embryo and the body of
plants shows that Empedocles thought about plants in relation to the embryo.
The same analogy might probably have also shaped his understanding of
plants’ nutrition through the roots. Plants derived their food from the earth
like embryos do from the body of their mothers—as later in Lucretius’ rooted
wombs. For Empedocles, however, we have evidence that discusses the nutri-
tion of plants from the earth in relation to plants’ different internal anatomy
but not in explicit terms of analogy with the embryo.66 Be that as it may, in this
discussion of the Hippocratic embryological model and Empedocles’ concep-
tion of plants’ life there is one more point to bring forward which highlights the
role of respiration in Presocratic thought and more specifically in Empedocles.
In On the Nature of the Child the process of respiration is tied to the process
of nutrition: ‘the thing that is warmed draws fresh, cold breath to itself … and
from this is nourished’, the Hippocratic doctor writes in the passage quoted

64 Fr. 31A30 Diels-Kranz / Arist. DA 2.415b28.

65 Vegetal growth was rigorously explained according to the doctrine of likes, which
informed at another level also Empedocles’ theory of sensation.
66 See, for instance, Plu. Quaest. conv. 3.2.2 649D / fr. 31 ad B77 Diels-Kranz.

Mnemosyne ( 2024) 1-23 | 10.1163/1568525X-bja10268

Downloaded from Brill.com 07/22/2024 12:39:01PM by [email protected]
via MRW Tester and BRILL
20 zatta

above, likely extending a doctrinal principle from the earlier physiologoi.67 To

the embryo air is food, the first type of nourishment, besides being the tool
that articulates the body’s internal anatomy. Now for Empedocles too the air
contributed to the process of plants’ nutrition and ensuing fructification. We
can gather it from a couple of fragments, 77.78, part of which has been trans-
mitted by Theophrastus in On the Causes of Plants and emended by Diels in the
1903 edition of the Presocratic philosophers. Let us look at these fragments:

⟨δένδρα δ᾽⟩ ἐμπεδόφυλλα καὶ ἐμπεδόκαρπα τέθηλεν

καρπῶν ἀφθονίῃσι κατ᾽ ἠέρα πάντ᾽ ἐνιαυτόν.

Evergreen and ever-fruiting ⟨trees⟩

with abundant fruit all the year due to the air.

The subject is plants’ capacity to produce leaves and fruits throughout the
year rather than at a specific period. In the passage that follows the quota-
tion of Empedocles’ words, Theophrastus explains that Empedocles sup-
posed that ‘a certain blend of air, the springtime air, is temperate.’68 In other
words, for Empedocles (as well as for Theophrastus) the aerial environment
has an impact on trees’ fructification, which is due to the optimal mixture of
springtime air. But while Theophrastus implied a mere contact between air
and tree,69 Empedocles believed that, along with all other living beings, also
plants breathed and that they breathed with the upper part of their body.
And it was by ‘inhaling’ the optimal mixture from the environment that the
trees contributed to the formation of the surplus complementing the process
of nutrition through the roots.70 In a similar way, in the Hippocratic On the
Nature of the Child the embryo is nourished both by the air which it inhales
in unison with its mother and, eventually, by the blood that reaches it inside
the mother’s belly.

67 I’m referring here to the conception of air as nourishment. Regarding the mechanism of
respiration, there is a slight discrepancy with the Empedoclean model. In Aëtius’ testi-
mony about the first breath of the first living being we are told that the expulsion of air is
due to the internal heat (see fr. 31A74 Diels-Kranz / Aët. 4.22.1) rather than its intake as in
the Hippocratic treatise.
68 τινα τοῦ ἀέρος κρᾶσιν, τὴν ἐαρινήν, κοινήν (Thphr. CP 1.13.2).
69 At CP 13.2.1 Theophrastus considers the lack of food (ἔνδεια τροφῆς) and the coldness of
air two distinct factors that influence plants’ production of fruit, thus showing that the
quality of the air is uninvolved with the process of nutrition.
70 For an extended discussion of fragments 77 and 78 in Theophrastus and Empedocles’ doc-
trines, see Zatta 2023.

10.1163/1568525X-bja10268 | Mnemosyne ( 2024) 1-23

Downloaded from Brill.com 07/22/2024 12:39:01PM by [email protected]
via MRW Tester and BRILL
“ is the embryo a living being? ” ( Aët. 5.15 ) 21

To conclude, in his commentary of the initial chapter of On the Nature of

the Child Lonie remarks on its forceful style, including repetitions, details
and proofs, by which the author boldly asserts the basic, essential principle
‘that the embryo acquires breath.’ Lonie further notes a close parallel with
Empedocles’ fr. 17 in which the ancient physiologos accounts for the elemen-
tal metaphysics that sustains his interpretation of the phenomena of life and
death.71 But I would argue that the engagement with Empedocles in this chap-
ter goes deeper than the mere enactment of an incisive style. For the insistence
with which the Hippocratic doctor claims that the embryo breathes and the
several proofs he adduces to support it seem also to constitute a firm critique
of Empedocles’ belief that the embryo does not breathe. Significantly, among
the proofs the doctor provides for the respiration of the embryo there is also
the strange (at least to us!) evidence that, similarly to the embryo, also leaves
and wood take in air when they are warmed up. This argumentative strategy
seems to apply Empedocles’ very conception of vegetal respiration to the
nature of the embryo foreshadowing the analogy between the development of
the embryo and that of the plant which will soon appear in chapter 22 of the
botanic excursus of the treatise. If the embryo develops like a plant, and plants
breathe, then, by analogy also the embryo must breathe. Empedocles, however,
denied that the embryo had this capacity. Perhaps as the lower parts of plants
in the earth were earthy, so the embryo kept a similarly earthy constitution in
the mother’s body to be considered to breathe while, in Empedocles’ embry-
ology, fire was relevant only to describe the uterine environment that deter-
mined the embryo’s sex.72 What is certain, however, is that for Empedocles
the embryo became a model for thinking about plants and their lives in a
similar way as it did for the nascent cosmos and the very origin of animal life
in other sources. Permanently attached to the earth and thus being perma-
nent embryos, plants did not breathe with their body underground but started
breathing as soon as they emerged from the earth. They were hybrid, archaic
beings who perpetuated in contemporary times the rise of animal life from
the earth at the dawn of the world. So conceived, plants were not categori-
cally differentiated from the other animals but constituted a form of animal
life. Their embryological nature in Empedocles holds a diversity of resonances,
both within Empedocles’ and other Presocratics’ doctrines. On the one hand,
in tune with the lack of categorical separation between plants and animals, it
directly relates to an earlier generation of living beings, the whole-natured, at

71 Lonie 1981.
72 See fr. 31B65 Diels-Kranz / Arist. GA 1.723a24-26 in which the temperature of the uterus
is considered the factor determining the sex of the embryo.

Mnemosyne ( 2024) 1-23 | 10.1163/1568525X-bja10268

Downloaded from Brill.com 07/22/2024 12:39:01PM by [email protected]
via MRW Tester and BRILL
22 zatta

a time of Strife’s ascending power over Love in a cyclical view of their dynam-
ics. Further, also the zoogonies involving parts of animals’ bodies under the
period of ascending Love were rising from the earth,73 a phenomenon which,
we may add, might also imply some forms of embryological processes since for
the rise of these living beings too Empedocles uses a botanical verb, βλαστάνειν
(to sprout, line 1). On the other hand, however, the embryological nature of
plants in Empedocles resonates also with those accounts from Anaximander
to Lucretius, discussed in this essay, in which the origin of animal life is inter-
preted in embryological terms—and in sharing their origins with the other
forms of animals’ life, plants may well have shared similar capacities.

Bibliography

Bailey, C. (1948, repr.). Lucretius on the Nature of Things. Oxford.

Baldry, H.C. (1932). Embryological Analogies in Pre-socratic Cosmogony. CQ 26.1
pp. 27-34.
Barney, S.A., Lewis, W.J., Beach, J.A. and Berghof, O. (2016). The Etymologies of Isidore of
Seville. Cambridge (UK).
Beekes, R. (2010). Etymological Dictionary of Greek. Leiden/Boston.
Campbell, G. (2003). Lucretius on Creation and Evolution. Oxford.
Classen, C.J. (1977). Anaximander and Anaximenes. The Earliest Theories of Change?
Phronesis 22, pp. 89-102.
Cornford, F. (1952). Principium Sapientiae. The Origin of Greek Philosophical Thought.
Cambridge (UK).
Diels, H. (1903). Die Fragmente der Vorsokratiker. Berlin.
Diels, H. and Kranz, W. (1951-1952). Die Fragmente der Vorsokratiker, 3 vols., 6th ed.
Berlin.
Ernout, A. (1924). Lucrèce. De la nature. Paris.
Furley, D.J. (1970). Variations of Themes from Empedocles in Lucretius’ Proem. BICS
17, pp. 53-64.
Gregory, A. (2016). Anaximander: A Reassessment. London.
Guthrie, W.K.C. (1957). In the Beginning: Some Greek Views on the Origin of Life and the
State of Man. London.
Guthrie, W.K.C. (1965). A History of Greek Philosophy. The Presocratic Tradition from
Parmenides to Democritus, vol. II. London.
Jaeger, W. (1963, 2nd ed.). Diokles von Karystos. Berlin.

73 See fr. 31B57 Diels-Kranz and Campbell’s commentary on Lucretius’ rooted wombs,
2003, 75.

10.1163/1568525X-bja10268 | Mnemosyne ( 2024) 1-23

Downloaded from Brill.com 07/22/2024 12:39:01PM by [email protected]
via MRW Tester and BRILL
“ is the embryo a living being? ” ( Aët. 5.15 ) 23

Kahn, C.H. (1960). Anaximander and the Origins of Greek Cosmology. New York.
Kirk, J.S., Raven, J.E. and Schofield, M. (1983). The Presocratic Philosophers. Cambridge.
Laks, A. (2019). Soul and Cosmos before Plato. Five Short Doxographical Studies. In:
R. Salles, ed., Cosmology and Biology in Ancient Philosophy, Cambridge, pp. 8-28.
Loenen, J.H.M.M. (1954). Was Anaximander an Evolutionist? Mnemosyne 7, pp. 215-232.
Lonie, I.M. (1969). On the Botanical Excursus in de Natura Pueri 22-27. Hermes 97,
pp. 391-411.
Lonie, M.I. (1981). The Hippocratic Treatises “On Generation” “On the Nature of the Child”
“Disease IV”. Berlin.
Luria, S., Krivushina, A. and Fusaro, D., eds. (2007). Democrito. Raccolta dei frammenti,
interpretazione e commentario. Milan.
Mansfeld, J. and Runia, D.T. (2020). Aëtiana V. An Edition of the Reconstructed Text of the
Placita with a Commentary and a Collection of Related Texts. Leiden/Boston. Part 3.
Marchetti, A. (1909). Della natura delle cose. Milan.
Monte Ransome, J. (2013). Nature, Spontaneity, and Voluntary Action in Lucretius.
In: D. Lehoux, A.D. Morrison and A. Sharrock, eds., Lucretius: Poetry, Philosophy,
Science, Oxford, pp. 99-130.
Sattler, B. (2021). The Ensouled Cosmos in Plato’s Timaeus: Biological Science as a
Guide to Cosmology? In: R. Salles, ed., Cosmology and Biology in Ancient Philosophy,
Cambridge (UK), pp. 29-45.
Schrijvers, P.H. (1974). La pensée de Lucrèce sur l’origine de la vie (« De rerum natura »
V 780-820). Mnemosyne 27.3, pp. 245-261.
Waszink, J.H. (1964). La création des animaux dans Lucrèce. RBPh 42.1, pp. 48-56.
Wilberding, J. (2021). Recapitulation Theory and Transcendental Morphology in Anti­
quity. In: R. Salles, ed., Cosmology and Biology in Ancient Philosophy, Cambridge
(UK), pp. 131-151.
Wright, M.R. (1981). Empedocles: The Extant Fragments. New Haven and London.
Zatta, C. (2023). Plants and Vegetal Respiration in Early Greek Philosophy. AncPhil 43.1,
pp. 251-272.
Zatta, C. (forthcoming). Empedocles on Plants’ Fructification. In: M. Protopapa, ed.,
Empedocles from the Beginning to the 2oth century. Athens.

Mnemosyne ( 2024) 1-23 | 10.1163/1568525X-bja10268

Downloaded from Brill.com 07/22/2024 12:39:01PM by [email protected]
via MRW Tester and BRILL