LXXII
A.’s model of inheritance. For the absence of an heredity theory of hair, eye, skin colour and hair type, see GA V. Much of my account of A.’s genetics is modelled on HENRY (2006a)’s insightful analysis of GA IV. Nevertheless, my interpretation differs from his in several ways: see below. A. discusses the inheritance of deformities at HA 585b29, GA 724a3 and teratology at GA IV, 3. The basic phenomena of inheritance are given at GA 767b1. His attack on pangenesis is in GA I, 17–18, specifically as applied to the children of deformed people at GA 724a4, cf. GA 721b28 [trans. PECK (1943)], as applied to plants, GA 722a13. MORSINK (1982) pp. 46–7 argues that A.’s target is the Hippocratic author of On Generation rather than Democritus – see Littré VII, On Generation, 3, 8, 11 for crippling and LONIE (1981). Morsink is surely right to suppose that A.’s opponent is the Hippocratic author but at GA 769a7 A. discusses two flavours of the theory, one of which may be Democritus’ who may have held some such theory, DK 68B32, DK 68A141 and DK 68A143. DARWIN (1868) vol. II, ch. 27 gives his theory of pangenesis; PECK (1943), MORSINK (1982), HENRY (2006a) among others have applied C.D.’s term to A.’s theory. C.D. acknowledges ancient Ppangenesis in DARWIN (1875) 2nd edition, vol. II, p. 370, footnote. See MORSINK (1982) ch. III for an analysis of A.’s argument against pangenesis; HENRY (2006a) notes the plant example.
LXXIII
Dual-inheritance theory. This term is a small novelty of mine; it emerges from a solution to a problem in A.’s theory of inheritance. In A.’s standard theory of reproductive hylomorphism males supply the form and females the matter, see HENRY (2006b), but GA IV allows that maternal matter (the menses) can also encode hereditary information. One solution to this apparent conflict is to allow that when A. talks about indivisible forms he means individuals not species. This is the solution that HENRY (2006a, b), among others, adopt – and it implies that both parents transmit form. I, however, think that the weight of evidence favours the idea that form picks out the essential features of kinds and that only fathers supply it (see n. Chs XXXIII and LXX). If this is so, then we need another term for variation within an atomon eidos, hence ‘informal variation’. Since such informal variation can come from both mothers and fathers and is also encoded in movements in the seed, we have then a dual-inheritance system: one (paternal) that encodes essential, functional features; the other (bi-parental) that encodes non-essential features (snub noses, sex, etc.), both of which depend on seminal movements and are susceptible to mutations. GA 767b24 speaks of several levels of inheritance.
Sex determination. A. critiques existing sex-determination theories at GA IV, 1. His own theory is framed in terms of hot/cold, GA 766b8. It is important to remember that for A. ‘hot’ does not merely denote the presence of heat (thermal energy) and ‘cold’ its absence, rather ‘hot’ and ‘cold’ are opposing qualities that are more like forces – hence the language of conflict and conquest. The idea of a proportion in sperm and menses (a logos or symmetria) occurs at GA 767a16, cf. GA 723a29; later, A. will restate the hot/cold theory in terms of actual and potential movements and tie it to a theory of general inheritance in GA IV, 3. A. speaks of environmental sex determination at GA 767a28 and refers to the parts (heart) as principles at GA 766a28. PLATT (1910) n. GA 716b5 points out the distinction between primary and secondary sex determination; PECK (1943) n. GA 776a30 points out that A. often seems ambivalent about whether the sexual parts are ‘principles’ or not, but clarifies his position at GA 766a31 and identifies the heart, PECK (1943) n. GA 766b8. A. discusses castration and eunuchs at GA 716b4, GA 766a26. He does not explain how castration might affect the heart. Perhaps he didn’t recognize just how direct his analogy was, for post-natal castration affects only some secondary sexual characteristics such as balding and voice pitch, but not the genitalia. See LEROI (2003) ch. 7 for an account of Jost’s experiments and sex determination.
LXXIV
A general theory of inheritance. A. explains his model of inheritance at GA IV, 3 and sex-associated features at GA 768a24. The woman of Elis features at HA 586a4 and GA 722a8. A. argues that the Hippocratic theory cannot explain ancestral similarities of this sort at GA 769a24; see HENRY (2006a). Failure of the semen’s heat as the cause of atavism, GA 768a9. Littré VII, On Generation, 8, shows that the Hippocratic theory is a blending theory since the author states: ‘If from any part of the father’s body a greater quantity of seed is derived than from the corresponding part in the mother’s body the child will, in that part, bear a closer resemblance to its father; and vice versa’ [trans. LONIE (1981), modified, italics mine]. Thus any trait has a continuous, rather than a discrete distribution, and depends on proportionate contribution; see also GA 769a7 where A. reports, much less precisely, of this (or a related) theory that if ‘the same amount comes from each of the two [parents], then, they say, the offspring formed resembles neither’. That probably means that the offspring is a blend of the two parents, but, admittedly, it could also mean that it’s something completely different. Monsters are said not to be hybrids at GA 769b11. A. gives his reversion theory of monstrosity at GA 767b1. For early modern theories of genetics see GLASS (1947) on Maupertuis, DARWIN (1868) vol. 2 pp. 399–401 and ch. XIII and MAYR (1982) ch. 14 for the dismal record of early theories of genetics. PA 642a29 tells how Democritus was brought to the theory of substantial definition by the facts.
LXXV
Shellfish of the lagoon. The biology of the ostrakoderma, HA IV, 4–7; the porphyra (murex), HA 528b36, HA 546b18 and PA 679b2; see THOMPSON (1910) n. HA 547a3 on the royal purple industry. On oyster gonads see GA 763b5; cf. HA 607b2.
LXXVI
Spontaneous generators. Some animals are generated from animals, HA 539a21. Cockles, clams, razorfish, scallops, oysters, fan mussel, ascidians, limpets, barnacles, murex, other snails, hermit crabs are said to be spontaneous generators at HA V, 15; sea anemones and sponges at HA V, 16; fish lice at HA 557a21; worms at HA 551a8; cockchafers, scarabs, flies, horseflies, pseudoscorpions, clothes moths at HA V, 19 and fish fry and Cnidian mullet at HA VI, 15–16. Oysters are provided as evidence for spontaneous generation at GA 763a26; cf. for mullets and eels HA 569a10 and HA 570a3. The recipe for an oyster is given at GA 762a19, GA 763a25; cf. HA 569a10. A. discusses eel reproduction at HA 538a3, HA 570a3 and GA 762b27. The gēs entera appears at HA 570a15, GA 762b22; see PLATT (1910) and PECK (1943) n. GA 762b22; THOMPSON (1947) p. 59 for varying ideas as to what it might be. THOMPSON (1910) n. HA 538a12, BERTIN (1956) and PROMAN and REYNOLDS (2000) discuss eel-head shape. On not removing the foundations of a science without replacing them, DC 299a5.
LXXVII
The fate of A.’s theory of spontaneous generation. A.’s theory of spontaneous generation and early modern science are discussed by FARLEY (1977), RUESTOW (1984) and ROGER (1997). Oyster’s gonads and larvae first observed by Brach in 1690; Leeuwenhoek independently described them in the following letters: 151 (1695), 157 (1695), 170 (1696) in LEEUWENHOEK (1931–99). Sea-urchin pluteus larvae were identified by Müller in 1846, barnacle nauplius larvae by Thompson in 1835, ascidian tadpole larvae by Kowalevsky in 1866. See WINSOR (1969) and WINSOR (1976) for accounts of their significance. Leeuwenhoek discusses his observations on eels and contemporary theories of eel reproduction in the following letters: 33 (1677), 15 (1691), 123 (1693), 169 (1696) in LEEUWENHOEK (1931–99). Leeuwenhoek initially observed putative eel progeny in the intestines of eels, but later identified them as parasites; he remained convinced, however, that he had identified the womb and progeny of the eel. For the discovery of the eel’s gonads see BERTIN (1956).
LXXVIII
Flies. Flies copulate and produce larvae, HA 539b10, cf. HA 542a6, GA 721a8; flies are produced from larvae, HA 552a20; flies are spontaneously generated, HA 552a20 and GA 721a8. The same confusion applies to fleas and lice: e.g. HA 556b21. A. also considers what would happen if maggots were to reproduce. He says they can’t since, if they did, they would necessarily produce a third kind of animal – some sort of ‘nondescrip
t’ – whose progeny, in turn, would be yet another kind of animal, and so on to infinity. They would engender an endlessly mutating lineage of living things, and that cannot be, for as he says, ‘nature flies from infinity’, HA 539b7 and GA 715b14.
Spontaneous generation recipe v. sexual reproduction. For the comparison see GA 762b1; specificity of the spontaneous generation recipe, GA 762a25. Many scholars have noted the tension between A.’s theory of spontaneous generation and his metaphysical commitments, though they agree neither on the exact nature of the problem nor on its solution, see PECK (1943) pp. 583–5, BALME (1962b), LLOYD (1996), ch. 5; LENNOX (2001b), ch. 10; GOTTHELF (2012) ch. 6; ZWIER (in prep.). Why believe in spontaneous generation. ZWIER (in prep.) argues that A. is investigating how spontaneous putative spontaneous generators actually are. My solution differs from hers only in the relative emphasis placed on the influence of A.’s predecessors on his thought and the degree to which ‘spontaneous’ generation and ‘spontaneous’ events sensu Physics II are intended in the same way. Following BALME (1962b) and LLOYD (1996) ch. 5, I think they’re being used in quite different ways. Theophrastus discusses spontaneous generation at CP I, 5.1–4; cf. CP I, 1.2, HP III, 1.3–6 and among the physiologoi HP III, 1.4. On origin-of-life theories and spontanous generation, see Prob X, 13; cf. GA 762b28. On traditional beliefs about spontaneous generation in cicadas see CAMPBELL (2003) p. 72. A.’s empiricism is evident in his discussion of spontaneous generation in mullets, HA 569a23, and muricids, HA V, 15 and GA 762a34. He gives the life cycle of the cicada at HA 556a25.
LXXIX
Life cycles. On the need for life cycles see Ch. XCVI and KING (2010). A. describes the natural history of the tuna at HA 537a19, HA 543b32, HA 543a9, HA 543a12, HA 571a8, HA 597a23, HA 598a18, HA 598a27, HA 599b9, HA 602a26, HA 607b28 and HA 610b4. He speaks of the regulation of monthly menstrual cycles in women at HA 582a34 and how most animals mate in spring in HA 542a20. A. talks of the alkyōn at HA 542b1, cf. HA 616a14; see PECK (1970) n. pp. 368–72 and ARNOTT (2007) for its identity and mythological associations. Most of A.’s information on the seasonal habits of animals, other than reproductive, is in HA VII, 12–30. He gives fish spawning times at HA VI, 17, cf. HA V, 9–11 and elsewhere, speaks of the hibernation of bees at HA 599a21 and bears at HA 600b28, the migration of cranes at HA 597a4, cf. HA 597b30, and the reasons why fish migrate at HA 598a30. Animals adjust their habits to the season at HA 596b20 and have certain thermal tolerances at HA 597a14. He discusses the relationship of life cycles to celestial cycles at GA 778a5 and GA IV, 10, GC 336b16 and LBV 465b26.
LXXX
Theory of elemental movement and transformation. On the natural movements of the elements see Phys 225a28, Phys 255b14 and DC 297a30. My account rests on claims in Physics VIII and DM that the elements are not, strictly speaking, self-movers. It follows COHEN (1996) ch. II and FALCON (2005) p. 11; see WATERLOW (1982) pp. 167–8 and GILL (1989) p. 238 for different accounts. On the transmutation of the elements see GC II, 1–5. On seasons and elemental transformation see GC 336a13, GC 336b16, GC 337a4 and GC 338b1; FALCON (2005) p. 11. LEUNISSEN (2010a) ch. 5.2–3 discusses the teleological connection between the theory of elemental formation and the celestial movements. The following passages in the Meteorology contain A.’s theory of winds and rains: Meteor I, 9, Meteor II, 4–6; but the wind has a life cycle in GA 778a2; on rivers Meteor 347a2 and geological cycles Meteor I, 14; see WILSON (2013). Many scholars have discussed Phys II, 8 198b16ff. on the winter rain; see JOHNSON (2005) ch. 5.5 and WILSON (2013) ch. 5. Wilson rightly weighs the ambiguities of this passage against the complete lack of teleological explanation in the Meteorology. See WILSON (2013) ch. 5 for a rich discussion of the use of biological metaphors in the Meteorology. He also offers the intriguing suggestion that meteorological phenomena should be viewed as dualizing between elements and spontaneous generators; and spontaneous generators as dualizing between meteorological phenomena and sexually reproducing animals.
LXXXI
Figs. A. on figs, HA 557b25; T. on figs, HP II, 8.1–3, CP II, 9.5–15. T. on seasonal flowers, HP VI, 8.1–5; on flower structures, HP I, 12. The quotes are respectively from HA 557b25 and GA 715b21; cf. GA 755b10. T. on the date palm, HP II, 6.6, HP II, 8.4, CP II, 9.15; AMIGUES (1988–2006) vol. I, p. xxiii, discusses the source of T.’s information about date palms; cf. Herod I, 193. See LLOYD (1983) ch. III, 2 on T.’s background. A. on plant sexes, GA 715b16 and GA 731a21; T. on plant sexes, CP II, 10; NEGBI (1995) discusses Theophrastus’ concept of male and female though gives him more assurance in distinguishing plant sexes than I think is his due. For the identity of fig-related insects see DAVIES and KATHIRITHAMY (1986) pp. 81–2, 92 and figs on Lesbos CANDARGY (1899) p. 29. I thank Charles Godfray, University of Oxford, for suggesting the identification of the kentrinēs, and Filios Akriotis and Theodora Petanidou, both of the University of the Aegean, Mytilene, for telling me respectively about fig varietal names and fig culture; I also thank Dimitrios Karidis, an Erresos fig farmer, for further information about the last at Erresos. For the history of the study of caprification see Gasparrini quoted in LELONG (1891). See KJELLBERG et al. (1987) and WEIBLEN (2002) for fig wasp life cycles.
LXXXII
Bees. For the origin of honey see HA VIII, 40 and Theophrastus HP VI, 11.2–4; SHARPLES (1995) pp. 208–10 for the missing Theophrastan work on honey. A. discusses the generation of bees at GA III, 10; MAYHEW (2004) ch. 2 defends A. from sexism on bees. A. speaks of his uncertainties about bees at GA 760b27; cf. DC 287b28 for a similar look to the future possible resolution of explanatory difficulties. MADERSPACHER (2007) gives a brief history of the elucidation of bee life cycles.
LXXXIII
Life history. The swallow winds, HP VII, 15. For the migratory and nesting habits of swallows see HA VII, 16 and HA VIII, 8. Eye regeneration in swallows, HA 508b4, HA 563a15, GA 774b31; and in chicks, DEL RIO-TSONIS and TSONIS (2003). On the altricial cubs of bears see HA 579a20 and PECK (1970) pp. 376–8.
Life-history patterns. A.’s life-history data on mammals and birds are mostly in GA IV, 4–10. Important passages telling of particular associations are: GA 771a17ff. (litter size and body size); GA 773b5 (adult body size and neonate body size); GA 774b5 (neonate perfection, litter size and gestation time); GA 774b30 (neonate perfection, gestation time); GA 777a32 (gestation time, longevity, neonate size). All this material is interwoven with explanations of abnormalities. Besides these passages, for the predicted longevity from gestation time of deer see HA 578b23; and LBV 466 b7 (longevity and fecundity); see n. Ch. LXXXV; and on birds see GA 749a35 (also below). SUNDEVALL (1835) coined the modern terms altricial and precocial; see STARCK and RICKLEFS (1998) for some of this history.
Explaining life-history associations. A. argues that the negative association between body size and fecundity is causal at GA 771b8, and that the positive association between gestation time and longevity is not causal at GA 777a35. On confounding variables in the comparative method see LEROI et al. (1994). For weakness after sex see GA 725b6. On the infertility of fat people see GA 725b32 and PA 651b12. For the effect of castration on longevity and growth see HA 575a31, HA 578a33 and HA 631b19; also LEROI (2010) and Chs LXXXV and XCVII. A. discusses the Adrianic fowl at HA 558b16, GA 749b25; Aldrovandi does too, LIND (1963) pp. 27–9. For the connection between feet, wings and way of life in birds see Ch. XLV; for their connection, in turn, to life history see GA 749a30, cf. GA 771a17. I emphasize the allocative aspect of his argument, but A. also argues that some raptors acquire less nutrition than other birds. See Appendix V for further discussion of the way that life-history features covary in mammals.
LXXXIV
Fish life history. T. lists the summer flowers at HP VI, 8.1–5. For A.’s observations on fish life history see HA VI, 10–17 and GA III, 3–6. That it is the function of (egg-laying) fishes and plants to be prolific is asserted at GA 718b8 and that this is, in the case of egg-laying fishes, due to high embryonic mortality is explained at GA 755a30, cf. HA
570b30. The features of egg-laying fishes that permit high fecundity are: (i) reverse sexual dimorphism, GA 720a16; (ii) small eggs, GA 755a30; (iii) external ‘perfection’ (fertilization? – see below) to avoid uterine space constraints, GA 718b8å, cf. GA 755a26; (iv) rapid growth of the embryos, GA 755a26; (v) parental care in the glanis and its explanation, HA 568b15. A. describes brooding in the beloně at HA 567b22, HA 571a2, GA 755a30. On the contrast between the fecundity of viviparous selacheans and oviparous scaly fishes see HA 570b29.
Perfect v. imperfect eggs. When speaking of the relative perfection of birds and mammal progeny, it’s clear that A. means something like altricial v. precocial. When he speaks of perfect and imperfect eggs (e.g. GA 718b8, GA 732b1, GA 754a22 and GA 755a11), he means something related but rather different. Once again, his technical vocabulary is seriously underdetermined. Among the reproductive products of fishes, A. thinks that those that are live born (those of most selacheans) are the most perfect, then eggs with a hard shell-like case (other elasmobranchs, e.g. the rays and skates) are less perfect; soft eggs (e.g. most scaly fishes) are the least perfect of all. The distinction lies in how much development the reproductive product (i.e. the thing that emerges from the mother) has to undergo before it becomes a functional creature (little, some, lots). This difference in egg morphology is, in fact, closely associated with fertilization mode: elasmobranchs have internal fertilization while most bony fishes have external fertilization, and it’s probable that A. recognizes this, but it’s hard to be sure since he’s very vague about how fish copulate. Even so, it’s likely that he views fertilization itself as a ‘perfecting’ of the female matter; and the stage at which this ‘perfecting’ occurs (early, internal v. late, external) partially determines how perfect the progeny are at birth. For the expansion of the jelly coat in fish eggs that occurs at fertilization see COWARD et al. (2002).
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