Plato’s Embryology

James Wilberding
Ruhr-Universität Bochum
James.Wilberding@ruhr-uni-bochum.de*

Abstract
Embryology was a subject that inspired great cross-disciplinary discussion
in antiquity, and Plato’s Timaeus made an important contribution to this
discussion, though Plato’s precise views have remained a matter of
controversy, especially regarding three key questions pertaining to the
generation and nature of the seed: whether there is a female seed; what
the nature of seed is; and whether the seed contains a preformed human
being. In this paper I argue that Plato’s positions on these three issues can
be adequately determined, even if some other aspects of his theory
cannot. In particular, it is argued that (i) Plato subscribes to the encephalo-
myelogenic theory of seed, though he places particular emphasis on the
soul being the true seed; (ii) Plato is a two-seed theorist, yet the female
seed appears to make no contribution to reproduction; and (iii) Plato
cannot be an advocate of preformationism.

Keywords
Plato – embryology – seed – history of medicine – ancient Greek science

In antiquity embryology was a subject that inspired great cross-

disciplinary discussion. As even a brief look at two later doxographical
texts sufficiently illustrates, embryology generated interest not only among
dedicated physicians but also among philosophers – these doxographers
refer to Alcmaeon, Anaxagoras, Aristotle, Democritus, Diogenes,
Empedocles, Epicharmus, Epicurus, Hippon, Leucippus, Parmenides,
Plato, Pythagoras and Pythagoreans, the Stoics, Strato and Zeno of Citium
by name – and this interest was spread out over a wide-ranging collection
of questions.1 When viewed against this background, it should come as no

*
Ruhr-Universität Bochum; Institut für Philosophie II; GA 3/31;
Universitätsstraße 150; D-44780 Bochum (Germany). I would like to thank
the journal’s anonymous referees for their useful comments on the first
version of this article.
1
Censorinus, De dei nat., 5-11 and Aëtius, Plac. phil., 5.3-21. A new
reconstruction of Aëtius’ Plac. phil. currently being produced by Jaap

1
surprise that Plato, too, seeks to include embryology within the scope of
his dialogue on natural philosophy, the Timaeus. What is surprising,
however, is Plato’s rather selective engagement with the traditional issues
here. There are a number of classic and very central embryological
questions that Plato simply remains silent on, for example: how twins are
formed, how the offspring’s sex is determined, and how to account for
deformities and (lack of) resemblance. Moreover, on other issues, notably
on the three major issues in spermatogenesis – the number of seeds
involved in reproduction, the corporeal origin of the seed, the manner in
which the offspring is present in the seed – previous scholarship has not
been able to reach a consensus on Plato’s positions.2 As I shall show in
what follows, there are indeed difficulties here, but I shall argue that Plato’s
views can nevertheless be adequately determined on all three of these
issues.
Let us begin with a preliminary description of the three issues in
question. The first concerns the number of seeds involved in normal
biological reproduction. On one theory the male is the sole supplier of
seed. References to such a theory in Aeschylus and Euripides suggest that
it was widespread among early Greeks, and we have evidence that it was
advanced by a number of philosophers, including Anaxagoras, Diogenes
of Apollonia, Aristotle, and the Stoics.3 An obvious difficulty connected to

Mansfeld and David Runia. For book 5, however, the Stobaean material is
also lost, so that we are basically left with Pseudo-Plutarch’s Plac. phil. [=
Mor. 904C-911C], an abbreviated version of Aëtius’ lost work. I have been
working with Lachenaud’s edition of that text, though I have adopted the
more informative system of referring to the text by book, chapter and
lemma (as in H. Diels, Doxographi Graeci (Berlin, 1879), 415-44), even
though Lachenaud’s edition does not include the numbers for the lemmas. I
would like to thank an anonymous referee and David Runia for clarification
on the background and current state of affairs regarding this Pseudo-
Plutarchean epitome.
2
See E. Lesky, Die Zeugungs- und Vererbungslehren der Antike und ihr
Nachwirken (Wiesbaden, 1951); I.M. Lonie, The Hippocratic Treatises “On
Generation” “On the Nature of the Child” “Diseases IV” (Berlin and New
York, 1981), 99-110; and H. von Staden, Herophilus. The Art of Medicine in
Early Alexandria (Cambridge, 1989), 288-96.
3
Aeschylus, Eumen., 657-666 (cf. Sept., 754) and Euripides, Orest., 551-6
(and cf. Sophocles, Od. Tyr., 1211 and 1257). Anaxagoras is sometimes

2
the one-seed theory is how to account for maternal resemblance, which the
alternative two-seed theory could easily explain. The view that both the
male and the female emitted seed found a wide-ranging scope of
acceptance among philosophers – Alcmaeon, Hippon, and other
Pythagoreans, Parmenides, Empedocles, Democritus, Epicurus – and
especially among ancient physicians: the Hippocratics, Diocles,
Herophilus, Soranus, and Galen.4 One difficulty of the two-seed theory that

reported to have advanced a two-seed theory, e.g., Censorinus, De dei

nat., 5.4 and 6.8, and this is sometimes accepted by modern scholars, for
example, by G. Lachenaud, Plutarque. Oeuvres Morales. Tome XII. 2e Partie.
Opinions des Philosophes (Paris, 2003), 298, who claims to find this in
Aristotle, GA, 763b30-764a1 (= 59A107 DK), but Aristotle here clearly
attributes a one-seed theory to Anaxagoras. R. Joly, Recherches sur le traité
pseudo-hippocratique du regime (Paris, 1960), 78-80, prefers Censorinus’
testimony to Aristotle’s. For Diogenes of Apollonia, see Censorinus, De dei
nat., 5.4 (= 64A27 DK) and cf. 64A24 DK. For Aristotle, see GA, 1.17-23
(726a30-731b14). Concerning the Stoics, see Censorinus, De dei nat., 5.4;
SVF, 1.128 and cf. Aëtius, Plac. phil., 5.11.4 (on which see Lesky, Zeugung,
171). For Zeno, see Aëtius, Plac. phil., 5.5.2 (= SVF, 1.129); for Sphaerus,
see Diogenes Laertius, Vit. phil., 7.159 = SVF, 1.626).
4
For Alcmaeon, see Censorinus, De dei nat., 5.4 (= 24A13 DK) and 6.4 (=
24A14 DK). Hippon is sometimes described as having advanced a one-
seed theory on the basis of 38A14 DK and Censorinus, De dei nat., 5.4, but
Aëtius, Plac. phil. 5.5.3 (= 38A13 DK) attributes a theory of female seed to
Hippon, though this seed does not contribute to the embryo as it is not
conducted to the uterus, as in Herophilus. See Lesky, Zeugung, 28, and
below on the female seed in Plato. For other references to the
Pythagoreans, see Diogenes Laertius, Vit. phil., 8.28 (= 58B1 DK); Aëtius,
Plac. phil., 5.5.1; and L. Zhmud, Pythagoras and the Early Pythagoreans
(Oxford, 2012), 374-80. For Parmenides, see 28B18 DK; Aëtius, Plac. phil.,
5.11.2 (= 28A54 DK); Censorinus, De dei nat., 5.4; 6.5; and 6.8 (the latter
two passages are included in 28A54 DK). For Empedocles, see 31B63 DK.
Cf. 31A81 DK. For Democritus, see Aëtius, Plac. phil., 5.5.1 (= 68A142 DK);
Aristotle, GA, 764a6-11 Drossaart Lulofs (= 68A143 DK), and P.-M. Morel,
“Aristote contra Démocrite. Sur l’embryon,” in L. Brisson, M.-H.
Congourdeau and J.-L. Solère, eds., Porphyre. Sur la manière dont
l’embryon reçoit l’âme (Paris, 2008), 43-57, at 46ff. For Epicurus, see
Aëtius, Plac. phil., 5.5.1. For the Hippocratics, see especially Genit. and
Nat. Puer. and Lonie, Hippocratic Treatises, 119-122. Also Mul. I, 8 (8.34,9f.;
8.56,21f.; and 8.62,20f. Littré) and Vict. I, 27 (144,4-5 Joly = 6.500,8f. Littré).
For Diocles, see Fr. 42a/b in P.J. van der Eijk, Diocles of Carystus. A
Collection of the Fragments with Translation and Commentary, 2 vols
(Leiden, 2000-2001) = Fr. 172 in M. Wellmann, Die Fragmente der
sikelischen Ärzte Akron, Philistion and des Diokles von Karystos (Berlin,
1901). Cf. Lesky, Zeugung, 30. For Herophilus, see Galen, De sem., 146,20-

3
its proponents must address is why the male is required for reproduction,
seeing as the female already has a seed at hand. A common (but not
universal) response to this difficulty was to posit that the female seed is
either inferior or else completely inactive.5
The second issue relates to the corporeal origin of the seed, and
there were three standard responses to this issue. The oldest was the so-
called encephalo-myelogenic theory, which states that the seed comes
from the brain and/or the marrow.6 This was held by Pythagoreans such as
Alcmaeon and Hippon, and traces of this theory can still be found in the
Hippocratic corpus and in Diocles.7 This theory eventually gave way to the
theory of pangenesis, which has the seed being drawn from the entire
body in order to better account for the family resemblances, as was
advanced by Anaxagoras, Democritus, Hippocratic authors and Epicurus.8

148,24 De Lacy (= 4.596,4- 598,7 Kühn and T60 von Staden) but also note 5
below. For Soranus, see Gyn., 1.4.93-98 Burguière et al. Regarding Galen,
see especially De sem., 2.1 (144,4 - 160,23 De Lacy = 4.593,1-610,10 Kühn)
and D. Nickel, Untersuchungen zur Embryologie Galens (Berlin, 1989), 40-
49.
5
Herophilos held that the female seed simply does not contribute anything
to the embryo because his anatomical studies suggested to him that the
seed was conducted to the bladder and from there expelled. Soranus
(Gyn., 1.12.93-98 Burguière et al.) takes over this view from Herophilus.
This same view has also been attributed to Hippon (see 38A13 DK). More
on this below.
6
Cf. Galen’s reference to this view as tau/thß palaia◊ß do/xhß (In Tim.,
14.10 Schröder).
7
Regarding Alcmaeon, see Aëtius, Plac. phil., 5.5.3, and Censorinus, De dei
nat., 5.2 (both are included in 24A13 DK). Censorinus testifies that
Alcmaeon actually opposed the encephalo-myelogenic theory, but as
Lesky, Zeugung, 12, points out, this is simply due to his “ungenaue
Sammelberichterstattung.” For Hippon, see Censorinus, De dei nat., 5.2 (=
38A12 DK); Aëtius, Plac. phil., 5.3.3 (= 38A13 DK). For more general
evidence of Pythagoreans holding this view, see Diogenes Laertius, Vit.
phil., 8.28 (= 58B1 DK) and the note ad loc (905A) in Lachenaud, Plutarque.
There are traces of this view in the Hippocratic corpus at, e.g., Genit., 1.2
(44,10-20 Joly = 7.470,8-16 Littré). In general, see Lesky, Zeugung, 13-18;
Lonie, Hippocratic Treatises, 101-3; von Staden, Herophilus, 288-96. For
Diocles, see Fr. 41a-b van der Eijk.
8
Anaxagoras 59B10 DK; Democritus 68B32 DK and cf. Aetius, Plac. phil.,
5.3.6 (= 68A141 DK). Concerning the Hippocratics, see, e.g., Morb. Sacr., 5
(12,21-14,2 Jouanna = 6.368,10-370,11 Littré); Aer., 14 (58,8-26 Diller =
2.58,11-60,8 Littré); Genit., 1.1 (44,1-10 Joly = 7.470,1-8 Littré), and in

4
The third and final position to emerge was the hematogenous theory,
which derives seed from the concoction of blood. This appears to be
advanced by Parmenides and Diogenes of Apollonia and is taken up and
developed in much greater detail by Aristotle and later physicians such as
Erasistratus, Herophilus and Galen.9
The final issue concerned the manner of the offspring’s physical
presence in the seed. Preformationists held that the body of the offspring
exists pre-formed in the seed, whereas epigenesists (e.g., Aristotle and
Galen) argued that the parts are formed successively after conception.10
Some form of preformationism necessarily follows from pangenesis, where
the exact type of preformationism to follow will depend on the kind of parts
being supplied and whether they are pre-arranged into an organic unity
(though preformationism would seem to be at least conceptually possible
even in the absence of pangenesis). It is helpful to distinguish between
three varieties of preformationism, which I shall call homoiomerous
preformationism, anhomoiomerous preformationism and homuncular
preformationism. The first two maintain respectively that the
homoiomerous parts such as the humors or flesh and bone and the
anhomoiomerous parts such as the head, hands and organs pre-exist in the
seed but are not yet organized into a unified whole, while according to
homuncular preformationism the seed already contains a unified organic
living thing. It is not clear that anyone in antiquity actually intended to

general, Lesky, Zeugung, 76ff.; Lonie, Hippocratic Treatises, 99-110; von

Staden, Herophilus, 288-96. Epicurus, Ep. Hdt., 38 and 66; cf. Lucretius, De
rerum nat., 1037-57; Aetius, Plac. phil., 5.3.5.
9
Parmenides 28B18 DK. For Diogenes of Apollonia, see 64A24 and B6 DK;
Diocles, Fr. 40 van der Eijk (= Wellmann, Fragmente, 208ff. = Herophilus,
T191 van Staden), and see van der Eijk’s extended comments ad loc. For
Erasistratus and Herophilus, see Herophilus, T191 von Staden (with his
discussion on pp. 293-6). For Galen, see, e.g., De sem., 1.12-14 (106,14-
114,21 De Lacy = 4.555,5-563,13 Kühn); De usu part., 14.10 (2.316,5-319,22
Helmreich) and 16.10 (2.412,21-424,15 Helmreich). On Galen’s
hematogenous theory, see Lesky, Zeugung, 181, and Nickel,
Untersuchungen, 34 and 89.
10
Aristotle, GA, 734a16ff. For Galen’s views on preformationism and
epigenesis, see Nickel, Untersuchungen, 67-83, and M. Boylan, “Galen’s
Conception Theory,” Journal of the History of Biology 19.1 (1986), 47-77.

5
defend homuncular preformationism, though some remarks come close to
suggesting it.11
Plato’s views on all three of these issues have been the subject of
various interpretations over the years. Remarks that give some indications
on Plato’s embryological views can be found scattered throughout the
dialogues, but it is in the Timaeus that we find Plato’s most considered
views on the issue, with the two most critical passages appearing at 73b1-
e1 and at the end of the dialogue in 91a4-91d5.
The first passage is concerned with the nature of the seed and its
origin in the body:

As for flesh and bones and things of that nature, this is

how it is. The starting point for all these was the
formation of marrow. For life's chains, as long the soul
remains bound to the body, are bound within the
marrow, giving roots for the mortal race. The marrow
itself came to be out of other things. For the god
isolated from their respective kinds those primary
triangles which were undistorted and smooth and
hence, owing to their exactness, were particularly well
suited to make up fire, water, air and earth. He mixed
them together in the right proportions, and from them
made the marrow, a panspermi÷a contrived for every
mortal kind. Next, he implanted in the marrow the
various types of soul and bound them fast in it. And in
making his initial distribution, he proceeded
immediately to divide the marrow into the number and
kinds of shapes that matched the number and kinds of
11
Notably, Aeschylus, Eumen., 657-666, and Euripides, Orest., 551-6.
Among philosophers, Democritus 68B32 DK strongly suggests the
homuncular variety, but when understood in the context of the rest of his
embryology, he seems to be advancing a weaker form of preformationism.
See the discussion of Plato below. Note one of Leon the Physician’s
etymological explanations of the Greek term embryon is “a mortal is
within” (endon broton) (Synopsis, 16,13 Renehan).

6
 shapes that the types of soul were to possess, type by
type. He then proceeded to mold the “field,” as it were,
that was to receive the divine seed, making it round,
and called this portion of the marrow, “brain.” Each
living thing was at its completion to have a head to
function as a container for this marrow. That, however,
which was to hold fast the remaining, mortal part of the
soul, he divided into shapes that were at once round
and elongated, all of which he named “marrow.” And
from these as from anchors he put out bonds to secure
the whole soul and so he proceeded to construct our
bodies all around this marrow, beginning with the
formation of solid bone as a covering for the whole of
it.12

This passage has been pointed to as evidence that Plato is an encephalo-

myelogenic theorist, and there is certainly a good deal of truth in this

12
Plato, Tim., 73b1-e1: To\ de« ojstw◊n kai« sarkw◊n kai« thvß toiau/thß fu/sewß
pe÷ri pa¿shß w—de e¶scen. tou/toiß su/mpasin aÓrch\ me«n hJ touv muelouv ge÷nesiß:
oi˚ ga»r touv bi÷ou desmoi÷, thvß yuchvß tw◊ø sw¿mati sundoume÷nhß, e˙n tou/twˆ
diadou/menoi katerri÷zoun to\ qnhto\n ge÷noß: aujto\ß de« oJ muelo\ß ge÷gonen e˙x
a‡llwn. tw◊n ga»r trigw¿nwn o¢sa prw◊ta aÓstrabhv kai« lei√a o¡nta puvr te kai«
u¢dwr kai« aÓe÷ra kai« ghvn di’ aÓkribei÷aß ma¿lista h™n parascei√n dunata¿,
tauvta oJ qeo\ß aÓpo\ tw◊n e˚autw◊n eºkasta genw◊n cwri«ß aÓpokri÷nwn, meignu\ß
de« aÓllh/loiß su/mmetra, panspermi÷an panti« qnhtw◊ø ge÷nei mhcanw¿menoß,
to\n muelo\n e˙x aujtw◊n aÓphrga¿sato, kai« meta» tauvta dh\ futeu/wn e˙n aujtw◊ø
kate÷dei ta» tw◊n yucw◊n ge÷nh, schma¿twn te o¢sa e¶mellen au™ sch/sein oi–a¿ te
kaq’ eºkasta ei¶dh, to\n muelo\n aujto\n tosauvta kai« toiauvta dihØrei√to
sch/mata eujqu\ß e˙n thvø dianomhvø thvø kat’ aÓrca¿ß. kai« th\n me«n to\ qei√on
spe÷rma oi–on a‡rouran me÷llousan eºxein e˙n auJthvø periferhv pantachvø
pla¿saß e˙pwno/masen touv muelouv tau/thn th\n moi√ran e˙gke÷falon, w˚ß
aÓpotelesqe÷ntoß e˚ka¿stou zw¿øou to\ peri« touvt’ aÓggei√on kefalh\n
genhso/menon: o§ d’ au™ to\ loipo\n kai« qnhto\n thvß yuchvß e¶melle kaqe÷xein,
a‚ma stroggu/la kai« promh/kh dihØrei√to sch/mata, muelo\n de« pa¿nta
e˙pefh/misen, kai« kaqa¿per e˙x aÓgkurw◊n ballo/menoß e˙k tou/twn pa¿shß
yuchvß desmou\ß peri« touvto su/mpan h¡dh to\ sw◊ma hJmw◊n aÓphrga¿zeto,
ste÷gasma me«n aujtw◊ø prw◊ton sumphgnu\ß peri« o¢lon ojste÷inon. The above
translation is drawn from D.J. Zeyl, Plato. Timaeus. Translated with
Introduction (Indianapolis, 2000), with some minor revisions.

7
attribution.13 For here Plato describes the marrow, which is constituted of
the most perfect triangles, as the starting point of the formation of all the
other parts. More importantly, he labels the marrow a panspermi÷a, which
usually gets translated as a kind of seed.14 But a closer look at the details of
this passage reveals that Plato appears to be employing the term
panspermi÷a in the sense of a receptacle of seed or a seedbed rather than a
seed itself.15 For he goes on to say that the Demiurge “implants” (futeu/wn)
the various types of soul in the marrow. Moreover, the concentration of
marrow that constitutes the brain is called a “field” (a‡rouran), and this
field is again said not to be a seed but to “receive the divine seed,” which
is the immortal (rational) soul.16 In short, the marrow appears to be
introduced not as a universal seed but as a universal seedbed, that is, a
receptacle for all kinds of seeds. The true seed appears to be the soul,
though Plato does not explain here or elsewhere in the Timaeus how
exactly the soul is supposed to execute the essential seminal function of
forming the embryo. Moreover, Plato explicitly refers only to the rational
soul as a seed, but the larger implication of the passage appears to be that
all three kinds of soul are in fact seeds, since otherwise the marrow could

13
E.g., Lesky, Zeugung, 18-20; E. Lesky and J.H. Waszink, “Embryologie,”
Reallexikon für Antike und Christentum 4 (1959), 1228-1244 at 1228; F.M.
Cornford, Plato’s Cosmology: The Timaeus of Plato translated with a running
Commentary (London, 1937), 295; van der Eijk, Diocles of Carystus, vol. 2,
92; M.-H. Congourdeau, L’embryon et son âme dans les sources grecques
(Paris, 2007), 197.
14
Zeyl, Timaeus, and Taylor (apud Cornford, Cosmology, 294): “universal
seed;” Cornford, Cosmology: “a mixture of seeds of every sort;” R.D.
Archer-Hind, The Timaeus of Plato (London, 1988): “a common seed;” T.
Paulsen and R. Rehn, Platon. Timaios (Stuttgart, 2003): “den gesamten
Samen;” L. Brisson, “La vivant, sa reproduction et sa nutrition selon
Platon,” in L. Brisson, G. Aubry, M.-H. Congourdeau and F. Hudry, eds.,
Porphyre. Sur la manière dont l’embryon reçoit l’âme (Paris, 2012), 31-46 at
40: “une semence universelle.” Cf. H. Müller, Platons sämmtliche Werke,
übersetzt von Hieronymus Müller, vol. 6 (Leipzig, 1857): “eine Verbindung
aller Samen.”
15
H.D. Rankin, Plato and the Individual (London, 1964), 33, comes close to
drawing this conclusion.
16
Cf. Theophilus Protospatharius, De corp. hum. fabr., 5.3.1 (189,5-8
Greenhill), where the brain is described as a “field” in which the rational
soul is planted, either directly or by means of the spinal marrow.

8
not be called a universal seedbed “for every mortal kind.”17 To be sure,
Plato does subsequently refer to the marrow as a “seed” (spe÷rma), which
would seem to be in tension with what he says here, but the tension is
easily alleviated by acknowledging that in the remaining part of the
Timaeus Plato is conceiving of marrow that is already in possession of
soul.18
The remaining two issues explored in our overview are best
approached by considering the second and the major embryological
passage in the Timaeus:

For the fluids we consume there is a channel, and where it

receives the fluids going through the lungs over the
kidneys to the bladder and expels them under pressure
from air, they bored a channel to the compacted marrow
that runs from the head down the neck and over the spine
– the marrow, that is, that we called “seed” above. This
marrow, seeing that it is ensouled and has been given
vent [in the male member], created a vital desire for
emission in the [part] where the venting takes place and
so brought an e¶rwß of begetting to completion. This is the

17
There has been some disagreement on the sense in which the marrow
here is said to be a panspermi÷an panti« qnhtw◊ø ge÷nei (73c1-2). Cornford,
Cosmology, 294-5, following A. Rivaud, Platon. Tome x: Timée, Critias
(Paris, 1925), and subsequently followed by Rankin, Individual, 34, has
suggested that Plato is here alluding to the future degeneration of human
beings into all the other kinds of mortal animals. As the skeletal structures
of all these animals are degenerate forms of the human structure, the
marrow, it is claimed, is a suitable foundation for the souls of all mortal
kinds of animals. I believe the main motivation behind this interpretation
has now been sufficiently diffused by T.K. Johansen, Plato’s Natural
Philosophy. A Study of the Timaeus-Critias (Cambridge, 2004), 151-2, and
follow him (and others) in taking the rational, spirited and appetitive parts
of soul to be the proper residents of the marrow.
18
This is particularly clear in Tim., 91b1-2. All of these characterizations of
marrow as “seed” (74a3-4; 74b3; 86c3-4; and cf. 77d3-4) are subsequent
steps in the generation of the human body that depend on the crucial initial
procedure of implanting the soul into the marrow (73c3-4), as is made
clear, e.g., in 75a2-3 (cf. 74e1-2), 81d4-7 and 91b1-2.

9
 reason why the male genitals are disobedient and self-
willed, like a living thing that will not listen to reason and
on account of its raging desires tries to overpower
everything else. And in women the wombs and uteri are
said for these same reasons to be a living thing within
them that desires to give birth to children; whenever this
living thing remains unfruitful for too long beyond the due
season, it becomes irritated and difficult and wanders
throughout the entire body and blocks off the passages of
breath, and by restricting its respiration sends the body
into severe difficulties and provides for all sorts of
diseases,19 until the [female] desire and the [male’s] e¶rwß
bring [these male and female parts] together and, like
plucking a fruit from the trees, sow into the womb as if
into a tilled field living things that are too small to see and
unformed, and then after having separated them again,
they nourish them until they grow large inside [the
womb] and after this they bring them to the light of day,
completing the generation of living things.20

19
It has been often and correctly remarked that Plato here is taking up the
doctrine of the “wandering womb,” which figures prominently in
Hippocratic gynecology, on which see J. Longrigg, Greek Medicine From
the Heroic to the Hellenistic Age. A Source Book (London, 1998), 194-201.
Although Longrigg gives no examples here of the wandering womb
causing respiratory problems, this is an ailment described often enough in
Hippocratic medicine (e.g., Mul. II, 125-6 and 130 [8.268,9-272,8 and 278,7-
11 Littré). This is likely Plato’s intended meaning, though his use of pneuvma
(91a6), aÓnapnoh/n (91b2) and aÓne÷pneusen (91b3) in connection with the
male seed raises some questions about whether the ailment in question
(ta»ß touv pneu/matoß diexo/douß aÓpofra¿tton, aÓnapnei√n oujk e˙w◊n) is rather
the obstruction of the menstrual flow, which is still more frequently
associated with the wandering womb (see Longrigg, Greek Medicine, 194-
201).
20
Plato, Tim., 91a4-d5: th\n touv potouv die÷xodon, h∞ø dia» touv pleu/monoß
to\ pw◊ma uJpo\ tou\ß nefrou\ß ei˙ß th\n ku/stin e˙lqo\n kai« tw◊ø pneu/mati
qlifqe«n sunekpe÷mpei decome÷nh, sune÷trhsan ei˙ß to\n e˙k thvß kefalhvß
kata» to\n aujce÷na kai« dia» thvß rJa¿cewß muelo\n sumpephgo/ta, o§n dh\
spe÷rma e˙n toi√ß pro/sqen lo/goiß ei¶pomen: oJ de÷, a‚t’ e¶myucoß w·n kai«
labw»n aÓnapnoh/n, touvq’ h∞øper aÓne÷pneusen, thvß e˙krohvß zwtikh\n

10
Before examining some of the larger issues in this passage, a few words
are necessary on the phrase in italics. This is a translation of kai« pa¿lin
diakri÷nanteß (Tim. 91d3), and scholars have proposed a number of
plausible interpretations of this phrase. (i) One possible interpretation has
been put forth by Praechter, namely that the living things that were initially
sown into the flesh of the womb are then separated from the wall of the
womb for gestation.21 Such an interpretation might be behind Archer-
Hind’s (1888) translation: “and again separating them.”22 (ii) Praechter also
suggests another interpretation on behalf of Michael of Ephesus, though
Praechter distances himself from it.23 Working on the assumption that Plato
is a two-seed homuncular preformationist, one might take kai« pa¿lin
diakri÷nanteß to mean that each of the tiny human beings – the one
supplied by the male and the other by the female – must be divided again

e˙piqumi÷an e˙mpoih/saß aujtw◊ø, touv genna◊n e¶rwta aÓpete÷lesen. dio\ dh\

tw◊n me«n aÓndrw◊n to\ peri« th\n tw◊n ai˙doi÷wn fu/sin aÓpeiqe÷ß te kai«
aujtokrate«ß gegono/ß, oi–on zw◊øon aÓnuph/koon touv lo/gou, pa¿ntwn di’
e˙piqumi÷aß oi˙strw¿deiß e˙piceirei√ kratei√n: ai˚ d’ e˙n tai√ß gunaixi«n au™
mhvtrai÷ te kai« uJste÷rai lego/menai dia» ta» aujta» tauvta [I delete the
comma here. See below note 45] zw◊øon e˙piqumhtiko\n e˙no\n thvß
paidopoii÷aß, o¢tan a‡karpon para» th\n w‚ran cro/non polu\n gi÷gnhtai,
calepw◊ß aÓganaktouvn fe÷rei, kai« planw¿menon pa¿nthØ kata» to\ sw◊ma,
ta»ß touv pneu/matoß diexo/douß aÓpofra¿tton, aÓnapnei√n oujk e˙w◊n ei˙ß
aÓpori÷aß ta»ß e˙sca¿taß e˙mba¿llei kai« no/souß pantodapa»ß a‡llaß
pare÷cei, me÷criper a·n e˚kate÷rwn hJ e˙piqumi÷a kai« oJ e¶rwß sunagago/nteß,
oi–on aÓpo\ de÷ndrwn karpo\n katadre÷yanteß, w˚ß ei˙ß a‡rouran th\n
mh/tran aÓo/rata uJpo\ smikro/thtoß kai« aÓdia¿plasta zw◊øa
kataspei÷ranteß kai« pa¿lin diakri÷nanteß mega¿la e˙nto\ß e˙kqre÷ywntai
kai« meta» touvto ei˙ß fw◊ß aÓgago/nteß zw¿øwn aÓpotele÷swsi ge÷nesin.
21
K. Praechter, “Platon Präformist?,” Philologus 83 (1928), 18-3 at 29n23.
22
And cf. Paulsen and Rehn’s translation: “sie dann wieder von ihr
trennen.”
23
Michael of Ephesus was a Byzantine commentator working in the 12th
century, who is the author of the earliest known commentary on Aristotle’s
GA (which was wrongly preserved under Philoponus’ name). Within this
commentary Michael presents an interpretation of Plato’s embryology (see
passages in next note) that is hard to reconcile with the complete picture
offered by the Timaeus (see below note 48). Praechter, “Platon Präformist,”
28, has suggested that this is due to the influence of Neoplatonism, but
Michael’s understanding of Platonic embryology is in fact very far
removed from Neoplatonic embryology (see J. Wilberding, “The
Revolutionary Embryology of the Neoplatonists,” Oxford Studies in Ancient
Philosophy 49 [2015]: 321-361).

11
into their organs so that a single composite human being may be created
out some male parts and some female parts. In fact, there is good reason to
doubt that Michael’s Plato was a homuncular preformationist; he appears
rather to be a two-seed vital anhomoiomerous preformationist.24 More to
the point, there are very good reasons for not ascribing any variety of
preformationism to Plato, as we shall see below. (iii) The current scholarly
consensus favors a third interpretation that takes this phrase together with
mega¿la […] e˙kqre÷ywntai, with both picking up on the description of the
seed as aÓo/rata uJpo\ smikro/thtoß kai« aÓdia¿plasta: since the seed is still
small and unformed, it must be given form and made larger. Thus, kai«
pa¿lin diakri÷nanteß is given the technical embryological sense of
articulating the embryo.25 It is certainly true that it was common in
embryological contexts to discuss when the articulation of the limbs took
place – the so-called point of first articulation, and that diakri÷nein and
dia¿krisiß are terms commonly used to refer to this process.26
When confronted with the details of the Greek text, however, the
consensus interpretation runs into serious problems. First, there is the
problem of the pa¿lin at 91d3. For what, on the consensus interpretation
(iii), would it mean for the embryo to be articulated again? But the real
problem concerns the subjects of the participle diakri÷nanteß: the male
e¶rwß and the female e˙piqumi÷a remain the subject throughout 91c7-d5.27 But
how can Plato think that the male e¶rwß and the female e˙piqumi÷a are
responsible for articulating the embryo’s features? This is a problem that
has already been brought out with great force by Sarah Broadie.28 (iv)

24
See again Praechter, “Platon Präformist,” 29n23. Cf. Michael, In GA,
25,20-31 and 33,19-34,6, and In PA, 36,35-37,3. And see below note 48.
25
E.g., A.E. Taylor, A Commentary on Plato’s Timaeus (Oxford, 1928);
Cornford, Plato’s Cosmology; Zeyl, Timaeus; Brisson, “La vivant.”
26
See, e.g., the Hippocratic Nat. Puer., 17.1-18.1 [59,9-61,7 Joly].
27
contra Rankin, Individual, 35.
28
See Broadie, Nature and Divinity in Plato’s Timaeus (Cambridge, 2012),
270-1. Broadie does not even consider alternative senses of diakri÷nanteß,
but she does an excellent job drawing out the absurd consequences of the
consensus interpretation. If the male e¶rwß is responsible for the formation,
then “Plato is on the way to admitting a third sort of principle into his
cosmology, something that is neither Intelligence nor material Necessity

12
These problems disappear if we simply do not insist on importing the
technical embryological sense of “articulation” and instead translate the
participle diakri÷nanteß straightforwardly as the complement to
sunagago/nteß: first, the male e¶rwß and the female e˙piqumi÷a bring the two
reproductive parts together, and then they separate (diakri÷nanteß) them
again (pa¿lin). While the suggestion that love is responsible not just for
bringing the male and female together but also for separating them might
sound surprising, it is reasonable if we bear in mind that the desire at issue
is a desire not for union but for creation. In other words, on the Timaeus’
account, there is no desire for sexual intercourse per se; there is only a
desire for procreation. This desire for procreation leads to sexual
intercourse in the first instance, but afterwards it promotes other, non-
sexual activities, such as caring for and nourishing the embryo during its
full period of gestation (cf. Tim. 91d4-5).29 This should be compared to
Plato Symposium 207a-b, where again e¶rwß and e˙piqumi÷a (207a7) are
described as causing certain ailments in living things that account not only
for sexual intercourse (summighvnai aÓllh/loiß b1-2) but also for their
nourishing of the offspring (th\n trofh\n touv genome÷nou b2; e˙ktre÷fein b5).
Now back to the larger issues. As has been remarked, Plato is not
delivering a fully worked-out embryology here, and the concision of his
account has led to some diverging opinions on where Plato stands with
respect to the existence of maternal seed and preformationism.30 This
passage initially appears to provide strong evidence for Plato’s

nor the combined effect of the two of them […] its mode of causation is sui
generis.”
29
This is analogous to the effects of the basic desire to restore the natural
state of the body. When my body is cold, this desire will lead me to
approach the fire, but once my body becomes too warm, it will also lead
me away from the fire (cf. Philebus 32a-b). This is possible because the
motivating desire is not simply a desire to be warm. Perhaps Plato even
followed the Hippocratic author of Superf. in believing intercourse during
pregnancy to be harmful to the child (see Superf., 13 [78,15-16 Lienau] and
26 [82,14-15 Lienau]; cf. Soranus, Gyn., 1.46.64-67 Burguière et al.), though
interpretation (iv) hardly requires us to assume this.
30
E.g., Taylor, Commentary, 639, and Lesky, Zeugung, 20.

13
subscribing to the one-seed theory.31 There is no explicit mention of a
female seed here or anywhere else in the Timaeus. The focus of Timaeus
91a4-b7, where the creation of the channel that leads the seed to the
reproductive organ is described, is exclusively on men, and when Timaeus
turns to discuss female reproductive organs no such channel is mentioned.
Moreover, the language of plucking a fruit (karpo/n – singular) and sowing
it in the womb, and describing the womb as a field further suggests that the
female is simply receiving the male seed. This would also seem to be
corroborated by Plato’s likening the receptacle to a mother and the source
of this reception to the father.32 Finally, a one-seed theory would also
harmonize well with several remarks regarding the male and female
contributions to generation that Plato makes elsewhere in the corpus. The
Republic, for example, already appears to draw a strong distinction
between the male and female contributions to reproduction: the male
begets (ojceu/ein) children, while the female bears (ti÷ktein) them.33 And the
same goes for Diotima’s account of physical pregnancy in the Symposium,
where Plato appears to buy in to the traditional view that the male actually
gives birth to the child when he emits the seed, where the female’s role
appears limited to providing a beautiful receptacle.34
Nevertheless, a strong case might be made for saying that Plato is in
fact a two-seed theorist and is simply not giving voice to the role of the
female seed in this passage.35 For the identification of the seed with the

31
E.g., Lesky, Zeugung, 30n1; Congourdeau, L’embryon, with caution;
Taylor, Commentary, 638-9 appears to lean towards the one-seed theory,
but is uncertain.
32
Tim., 50d2-3.
33
Plato, Rep., 454d10-e1. As K. Dover, Plato Symposium (Cambridge,
1980), 147, notes, the verb ti÷ktein can be used of both male begetting and
female bearing, but the contrast here is clear.
34
Plato, Symp., 206b-e (esp. 206d7: i¶scon to\ ku/hma), on which see E.E.
Pender, “Spiritual Pregnancy in Plato’s Symposium,” The Classical
Quarterly N.S., 42.1 (1992), 72-86 at 73-76.
35
A two-seed theory has been attributed to Plato by H. Cherniss, Aristotle’s
Criticism of Presocratic Philosophy (Baltimore, 1935), 284n243; W. Gerlach,
“Das Problem des ‘weiblichen Samens’ in der antiken und mittelalterlichen
Medizin,” Sudhoffs Archiv für Geschichte der Medizin und der
Naturwissenschaften 30:4-5 (1938), 177-193 at 182; Geurts (apud Lesky,

14
(ensouled) matter of the brain or marrow would seem to demand a two-
seed theory, since brain and marrow are just as much a part of female
anatomy as they are of male anatomy. Perhaps this is also why, according
to the little evidence we possess, all Presocratic encephalo-myelogenic
theorists were also two-seed theorists.36 When viewed from this
perspective, then, Plato’s endorsement of a version of the encephalo-
myelogenic theory would already seem to commit him to a two-seed
theory. Further, certain details in the present passage supply some
additional support for a two-seed theory. He conspicuously says that the
female reproductive organs are living things “for these same reasons” (dia»
ta» aujta» tauvta) that explained the male case, which could be taken to
refer to a single cause of life in the male and female reproductive organs:
the life-status of the female reproductive organs is owed to the presence of
an ensouled marrow-seed.37 Likewise, what is said to be sown into the
womb are “living things” (zw◊øa – plural), which could be taken to mean that
two seeds are involved. Finally, denying that Plato allowed for a female
seed would seem to raise certain difficulties, such as how to account for
maternal resemblance.
Given the considerable amount of evidence pointing in each
direction, the truth would seem to lie near some intermediate position. In
light of Plato’s identification of marrow and seed, it is very difficult to deny
the existence of a female seed,38 but he might well have thought that this
seed made no direct contribution to the formation of the embryo. Such a

Zeugung, 30); and H.D. Rankin, “On ΑΔΙΑΠΛΑΣΤΑ ΖΩΙΑ (Plato, Timaeus
91d3),” Philologus 107 (1963), 138-145 at 141, and Individual, 36.
36
For Alcmaeon, see 24A13-14 DK; for the Pythagoreans, see Diogenes
Laertius, Vit. phil, 8.28 (= 58B1 DK), Aëtius, Plac. phil., 5.5.1; for Hippon,
see Censorinus, De dei nat., 5.2 (= 38A12 DK); Aëtius, Plac. phil., 5.5.3 (=
38A13 DK).
37
Cf. Taylor, Commentary, 639.
38
Cf. Cherniss, Aristotle’s Criticism, 284n243: “I think anyone who reads
Timaeus 91A-D will admit that Plato could not consistently have otherwise
supposed [viz. that there is also a female seed]. Spinal marrow which is
seed exists in both sexes, and he expressly says both sow the unformed
animals into the womb.” As will become clear, I agree with Cherniss only
regarding the former claim.

15
theory was advanced by Herophilus, the famous physician living
approximately a century after Plato, whose detailed anatomical studies led
him to conclude that although the female produces a seed, there are no
spermatic ducts leading the seed to her uterus so that the seed was
expelled from the body without making any contribution, and this view was
subsequently taken over wholesale by Soranus.39 A similar theory has
been ascribed to the Presocratic Hippon, who lived in the century prior to
Plato. Presumably it was theoretical rather than anatomical considerations
that led Hippon to this conclusion: the encephalo-myelogenic theory
demands the existence of the female seed, but perhaps Hippon denied it
any role in embryology in order to avoid the problem of
parthenogenesis.40 We may do justice to the present passage, then, by
allowing for a female seed but then taking Plato seriously when he
describes the seminal duct as being constructed exclusively in the male.41
The other considerations in favor of a more efficacious female seed
are not compelling. First, Plato does not seem particularly interested in
addressing the problem of maternal resemblance, and even if he were
concerned about this, the remarks he makes elsewhere suggest that he
would have again agreed with Hippon that the nourishment provided by
the mother is sufficient to the task.42 For example, when Socrates in the
Republic complains about the current Athenian constitution’s ability to
corrupt the nature of its citizens, he draws a comparison to how differences
in soil can alter and pervert the natures of plants: “just as a foreign seed
sown in alien ground: when it is overcome, it fades and likes to go over

39
Herophilus, T61 von Staden, with the comments by von Staden on 230ff.
Soranus, Gyn., 1.12.96-98 Burguière et al.
40
Aetius, Plac. phil., 5.5.3 (= 38A13 DK) and see above note 5. According to
Aetius, Plac. phil., 5.7.7 (= 38A14 DK) Hippon viewed the female
contribution to consist only in trofh/. See Lesky, Zeugung, 27-28.
41
Taylor, Commentary, 637, also appears to restrict the seminal duct’s
creation to the male. Note that Aristotle considered the uterus to be the
female counterpart to male seminal passages (GA, 720a12-14).
42
For Hippon, see Aetius, Plac. phil., 5.7.7 (= 38A14 DK) with Lesky,
Zeugung, 27-28.

16
into the native species.”43 Given the widely accepted analogy between the
earth-plant relationship and the mother-embryo relationship, the
implication would seem to be that the nourishment supplied by the mother
is sufficient to account for significant formative changes in the embryo.44 In
addition, Plato’s description of the living things in the seed still being
“unformed” (aÓdia¿plasta) also leaves the door open for the female to
exercise some formative influence during the period of gestation, a point
to which we shall return below. Second, when the womb is said to be a
living thing “for these same reasons” (dia» ta» aujta» tauvta), Plato is
making an epistemological point rather than giving a causal account.45 He
is drawing our attention to that fact that it is possible to witness behavior of
both male and female reproductive organs from which it may be inferred
that they are independent living things. Just as we may infer that the male
member is an independent living thing on account of its unruliness in
being aroused, so too may the womb be inferred to be a living thing on
account of its own brand of unruliness, namely its proclivity to wander. In
both cases the parts behave contrary to the interests of the whole on
account of their individual agendas – the male’s desire (e˙piqumi÷a) for
emission and e¶rwß of procreation (genna◊n), and the female’s desire
(e˙piqumi÷a) for child-bearing (paidopoii÷a). The dia» ta» aujta» tauvta is,
therefore, sufficiently accounted for by this parallel. There remains the
question of how to account for the womb’s desire for child-bearing, but it is
hard to see how transferring the account given for the male to the female

43
Plato, Rep., 497b4-5: w‚sper xeniko\n spe÷rma e˙n ghvø a‡llhØ speiro/menon
e˙xi÷thlon ei˙ß to\ e˙picw¿rion filei√ kratou/menon i˙e÷nai. Cf. Plato, Rep., 491d1-
4; Theat., 149e3-4; Menex., 237e1-238a5.
44
Interestingly, Aristotle himself seems to accept some version of this
theory in GA, 2.4 (at 738b27-36), for which Galen later criticizes him for
effectively turning nourishment into a seed instead of acknowledging a
female seed (De sem., 154,1-11 De Lacy [= 4.602,10-603,15 Kühn] and
156,1-7 De Lacy [= 4.604,12-18 Kühn]). Lesky, Zeugung, 173-7, takes
Galen’s criticism to be directed at both Aristotle and Athenaeus of Attaleia.
45
I read the dia» ta» aujta» tauvta as modifying lego/menai and delete the
comma after tauvta: “are said for these same reasons to be a living thing
within them.”

17
would be of any help here.46 For that account can only explain the male’s
desire to emit seed and procreate but not the female’s unique desire to
bear children, which is perhaps better explained by assuming that her
reproductive parts lack the seed that they naturally long for. Finally, Plato’s
use of the plural zw◊øa to describe the seed cannot be decisive, as he has
been loose with his use of singulars and plurals throughout the passage,
and the repetition of the plural in his account of birth – “completing the
generation of living things” – suggests that Plato has switched to the plural
because he is discussing all cases of reproduction collectively and not
because he means to suggest that there is more than one seed.47
Plato’s position on preformationism has equally caused a great deal
of confusion. A number of scholars have seen Plato as a preformationist –
even as a homuncular preformationist – but what he gives us in the Timaeus
is really just a puzzle. This puzzle is concentrated in his description of the
seed in Timaeus 91d2-3. On the one hand he calls the seed a zw◊øon (or even
a plurality of zw◊øa) that is simply too small to see, which suggests a strong
form of preformationism, but on the other he describes these same zw◊øa as
being “unformed” (aÓdia¿plasta), which would seem to speak against
preformationism.48 What has gone unnoticed is that this same puzzle is

46
Taylor, Commentary, 639, points to this question: “As we have had no
description of the formation in the female of a counterpart to the e¶rwß touv
genna◊n, one may perhaps suppose that T. regards the passion as due in
both sexes to the same cause, the attempt of the muelo/ß to exit […] Yet the
very next sentence suggests the cruder view that the whole of the child’s
body comes from the father.”
47
As noted above, Plato gives the analogy of plucking a single fruit
(karpo\n) at Tim., 91d1, and at Tim., 91b7-c2 Plato turns to the womb by
using the plural (ai˚ d’ e˙n tai√ß gunaixi«n au™ mhvtrai÷ te kai« uJste÷rai) but
then switches to the singular zw◊øon, which remains the subject of 91c2-7.
48
Taylor, Commentary, 640, attributes homuncular preformationism to
Plato: “The zw◊øon spoken of here is supposed to be the future infant itself in
minature, as we see from the words mega¿la … ei˙ß fw◊ß aÓgago/nteß.” H.
Balss, “Präformation und Epigenese in der griechischen Philosophie,”
Archivo di storia della scienza 4 (1923), 319-325 at 320, and M. Wellmann,
“Spuren Demokrits von Abdera im Corpus Hippocraticum,” Archeion 11
(1929), 297-330 at 307-8, also attribute some form of preformation to Plato,
though both put a great deal of weight on Michael of Ephesus’ account of
Plato’s embryology (on Michael of Ephesus, see above notes 23 and 24).
Lesky, Zeugung, 20, credits Praechter and Geurts with having refuted

18
already to be found in Democritus, whose embryology appears to have
had a significant influence on Plato’s own.49 This is certainly not to say that
Plato took over Democritus’ embryological theory wholesale, as there are
many features of the Democritean embryology that are incompatible with
the bits of theory to which Plato does clearly subscribe. Democritus, for
example, advances a two-seed theory of pangenesis in which the female
seed makes a significant contribution to the embryo, and his atomism
figures prominently into his embryology. Yet Plato does appear to help
himself to bits and pieces of Democritus, which is perhaps only to be
expected given Democritus’ contributions to the field.50 This is most
striking in Plato’s possible appropriation of the Democritean term
panspermi÷a (67A15 and 28 DK) as well as in the readiness of both
philosophers to infer the existence of organic entities that are “too small to
be seen” on the basis of rational inquiry.51 Yet it also appears in a more

Michael’s relevance, and she herself thinks that the aÓdia¿plasta decides
the matter: “denn der Ausdruck ‘nicht durchgeformte Lebewesen’ spricht
durchaus gegen die Annahme von Präformation.” The conclusion reached
by Brisson, “La vivant,” 41, may be viewed as a recapitulation of the
puzzle: “Quoi qu’il en soit, Platon peut être considéré comme un
préformiste, dans la mesure où les êtres vivants sont déjà formés dans le
sperme même s’ils sont invisibles et informes.”
49
This parallel between Plato and Democritus was noted by Wellmann,
“Spuren Demokrits,” though Wellmann depends too much on Michael’s
exegesis (see previous note), but is oddly not to be found in S.M. Nikolaou,
Die Atomlehre Demokrits und Platons Timaios (Stuttgart, 1998).
50
Democritus appears to have developed at least one of – if not the – most
comprehensive embryological theories of the Presocratic philosophers. He
is often regarded as having exercised a considerable influence on
Hippocratic embryology. See, e.g., Wellmann, “Spuren Demokrits;” Lesky,
Zeugung, 70-76; A. Stückelberger, Vestigia Democritea. Die Rezeption der
Lehre von den Atomen in der antiken Naturwissenschaft und Medizin (Basel,
1984), 49-87; Lonie, Hippocratic Treatises, 62-71 (and 176-86 and passim).
His influence has been called into question by L. Perilli, “Democritus,
Zoology and the Physicians,” in A. Brancacci and P.-M. Morel, eds.,
Democritus: Science, the Arts, and the Care of the Soul (Leiden and Boston,
2007), 143-179 at 162-172: “There are two radically opposing parties:
philosophers as a rule see an influence of Democritus on medical treatises
concerning specific aspects, while historians of medicine usually tend to
exclude it. I subscribe to the latter view” (167).
51
In PA, 665a31-33, (= Democritus, 68A148 DK) Aristotle, who maintains
that only blooded animals have viscera, criticizes Democritus for

19
subtle form in connection with Democritus’ commitment to
preformationism. A commitment on his part of some kind necessarily
follows from his pangenesis, though which kind exactly he subscribes to is
obscured somewhat by mixed messages in our evidence. One fragment
appears to provide strong evidence for Democritus’ commitment to
homuncular pangenesis: “Sexual intercourse is a minor stroke: for a human
being bursts forth from a human being, and is torn off, separating, by a
blow.”52 Yet homuncular pangenesis would seem to be irreconcilable with
two of Aristotle’s reports about the formation of the embryo in the womb.
He criticizes Democritus for maintaining that the embryo remains in the
womb “in order that the parts may be formed (diapla¿tthtai) after the
fashion of the parts of the mother,” whose articulation of the embryo
begins on the outside and works its way inwards.53 So just as Democritus
describes the seed as an a‡nqrwpoß that still needs to be diapla¿ttesqai,

overturning the empirical evidence and positing viscera that are too small
to see (dia» mikro/thta a‡dhla) in bloodless creatures. Plato appears to
make a similar inference about the seed in Tim., 91d2 (aÓo/rata uJpo\
smikro/thtoß). Cf. Tim., 43a3.
52
68B32 DK: xunousi÷h aÓpoplhxi÷h smikrh/: e˙xe÷ssutai ga»r a‡nqrwpoß e˙x
aÓnqrw¿pou kai« aÓpospa◊tai plhghvi tini merizo/menoß. Lesky, Zeugung, 72,
takes this to imply homuncular preformationism: “Der real-morphologishe
Zusammenhang zwischen den elterlichen Organen und Organteilen und
denen des Keims, die dieser in präformiertem Zustand – denn als Mensch
stürzt er schon aus dem Menschen heraus – vererbt erhält, ist […] zum
Ausdruck gebracht.” Lonie, Hippocratic Treatises, 180, with somewhat
more caution, agrees: “It is exceedingly tempting to see a form of the
homunculus theory here […] Clearly we cannot be certain that Democritus
did think in this way; but we can at least say that of existing theories, that of
Democritus was the best suited to explain organic structure.” As M.L.
Gemelli Marciano, “Le Démocrite technicien. Remarques sur la reception
de Démocrite dans la littérature technique,” in A. Brancacci and P.-M.
Morel, eds., Democritus: Science, the Arts, and the Care of the Soul (Leiden
and Boston, 2007), 207-237 at 215, shows, 68B32 DK “n’est toutefois attestée
nulle part sous sa forme complète.” Cf. also John of Alexandria’s use of a
similar phrase in what is clearly not a case of homuncular preformationism
(In Hipp. Nat. Puer., 134,16 Bell et al.).
53
Aristotle, GA, 740a35-7 (= 68A144 DK): tou/tou ga»r ca¿rin e˙n tai√ß
uJste÷raiß me÷nei to\ zw◊ion, aÓll’ oujc w˚ß Dhmo/krito/ß fhsin, iºna
diapla¿tthtai ta» mo/ria kata» ta» mo/ria thvß e˙cou/shß. And 740a13-15 (=
68A145 DK): o¢soi le÷gousin, w‚sper Dhmo/kritoß, ta» e¶xw prw◊ton
diakri÷nesqai tw◊n zw¿iwn, u¢steron de« ta» e˙nto/ß, oujk ojrqw◊ß le÷gousin.

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Plato describes it as zw◊øa that are still aÓdia¿plasta. Even Plato’s use of the
plural zw◊øa might have some connection to the enigmatic Democritean
fragment 68B124: oJ me«n Dhmo/kritoß le÷gwn a‡nqrwpoi ei–ß e¶stai kai«
a‡nqrwpoß pa¿nteß, if it has not been corrupted in transmission.54 For
Democritus, this paradox is presumably to be resolved by a more liberal
reading of 68B32 that credits him with a softer version of pangenesis. This,
in any case, would also seem to be the implication of 68A141: “Democritus
says that [the seed derives] from the entire body and the most important
parts such as its bones, flesh and sinews.”55 If this is right, then Democritus’
theory would be preformationist to the extent that the seed contains a
human being in the sense of containing all of the relevant parts of a human
being (certainly bones, flesh and sinews, and perhaps even
anhomoiomerous parts), though these parts would still need to be
assembled together and formed in the womb.56

54
Diels-Kranz label the fragment unintelligible and do not even translate it.
Diels conjectures that 68B124 DK is a corruption and suggests the original
read a‡nqrwpoß e˙xe÷ssutai e˙x aÓnqrw¿pou panto/ß (cf. 68B32 DK), which
might well be right. As it stands, the sense might be that every portion of
the seed may be counted as a human and that all of these “humans” go
together to form a single human offspring. The phenomenon of polygony
might be in the background here.
55
Aëtius, Plac. phil. 5.3.6 (= 68A141 DK): Dhmo/kritoß aÓf’ o¢lwn tw◊n
swma¿twn kai« tw◊n kuriwta¿twn merw◊n oi–on ojstw◊n sarkw◊n kai« i˙nw◊n [to\
spe÷rma ei•nai]. There is some disagreement about whether the first kai/ is
meant to be epexegetic (thus Perilli, “Democritus,” 171) or co-ordinative
(H. De Ley, “Pangenesis versus panspermia. Democritean Notes on
Aristotle’s Generation of Animals,” Hermes 108 [1980], 130-153 at 135-6). In
the former case we have homoiomerous pangenesis, the results for the
latter interpretation will depend on how one understands aÓf’ o¢lwn tw◊n
swma¿twn (which De Ley takes to refer to the organs).
56
As P.-M. Morel, “Démocrite et l’objet de la philosophie naturelle. A
propos des sens de φύσις chez Démocrite,” in A. Brancacci and P.-M.
Morel, eds., Democritus: Science, the Arts, and the Care of the Soul (Leiden
and Boston, 2007), 105-124 at 110-11, and “Aristote contra Démocrite,” 52-
53, notes, there remains some ambiguity about how to reconcile
Democritus’ two-seed theory, which is already supposed to account for
paternal and maternal resemblances at conception (68A143 DK), with his
claim (68A144-5 DK) that the mother forms the embryo according to her
own parts during gestation. On the former aspect of Democritus’ theory,
see Lesky, Zeugung, 73-4, and De Ley, “Pangenesis,” 142-3.

21
 The puzzle, however, that Plato presents to us in the Timaeus, is
more resistant to solution, though preformation, in any of its three forms,
would seem to be difficult to square with the theory of the Timaeus. Some
might look to buttress the case for preformationism by appealing to
Diotima’s account of pregnancy in the Symposium, and perhaps the
absence of any discussion in the Timaeus of the standard question of the
order of formation of the offspring’s parts could be interpreted as further
evidence that Plato is simply assuming that the parts are already formed.57
Yet the fact stands that in the Timaeus the seed consists only of soul plus
marrow, while even the weakest variety of preformationism,
homoiomerous preformationism, demands that all of the fundamental
homoiomerous parts be present in the seed.58
None of the above is meant to overturn our opening observation that
Plato is not attempting to deliver a fully worked-out embryology. Indeed,
there are many major issues of ancient embryology that appear simply not
to have fit into the Timaeus’ philosophical program. But we can clearly
distinguish the contours of his theory on the three central issues of
spermatogenesis, which may now be summarized as follows. Plato
subscribes to the encephalo-myelogenic theory of seed, though he places
particular emphasis on the soul being the true seed. From this it also
follows that Plato is a two-seed theorist, yet the female seed appears to

57
In the Symp., 206c-e, Plato appropriates the female experience of
pregnancy for the male. Ejaculation of the seed is recast as giving birth.
Plato is silent on the woman’s birth pains. Birth, moreover, is recast as an
entirely pleasant affair that no longer maps onto a female’s experience.
The only pain worth mentioning is that of the male, when he cannot off-load
his “offspring” (τὸ κύημα). See F. Sheffield, “Psychic Pregnancy and
Platonic Epistemology,” Oxford Studies in Ancient Philosophy 20 (2001), 1-
35 at 13-15.
58
Nor may one claim that for Plato marrow is the only fundamental
homoiomerous part. At least when Plato describes the creation of flesh,
there is no indication that it is being created out of marrow (Tim., 74c5-d2).
Bones, likewise, are soaked in marrow but are not described as being
generated out of marrow (Tim., 73e1-74a1). Additional (minimal) support
against preformationism might be drawn from Leg., 872e5-10, where Plato
advocates equal punishments for matricide and patricide, in stark contrast
to the preformationism-based defense of Orestes in Aeschylus, Eumen.,
602-10 and 652-66.

22
make no contribution to reproduction. Finally, given his commitment to the
above encephalo-myelogenic theory, Plato cannot be an advocate of
preformationism. Rather, his view must be that the soul in the seed is
responsible for forming the embryo, though he offers us no further
explanation of this aspect of this theory.

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