Book - A History of Embryology 1959-4

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I have decided to take early retirement in September 2020. During the many years online I have received wonderful feedback from many readers, researchers and students interested in human embryology. I especially thank my research collaborators and contributors to the site. The good news is Embryology will remain online and I will continue my association with UNSW Australia. I look forward to updating and including the many exciting new discoveries in Embryology!

Needham J. A History of Embryology. (1959) Cambridge University Press, London.

1959 Needham: Chapter 1 | Chapter 2 | Chapter 3 | Chapter 4

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This historic 1959 embryology textbook by Needham was designed as an overview of embryology history. Currently only the text has been made available online, figures will be added at a later date. My thanks to the Internet Archive for making the original scanned book available.

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Chapter 4. Embryology in the Eighteenth Century

1. Theories of Foetal Nutrition

DURING the course of the seventeenth, and the first quarter of the eighteenth, century, many theories were propounded concerning foetal nutrition. It is convenient to classify them (Table I).

At this point the Emmenalogia of John Freiud' deserves special reference. This was a book which dealt with all aspects of menstruation.’ As has already been mentioned (p. 15o n. 2) be supposed that the blood passing through the placenta to the embryo was distinctively menstrual blood. This view he supported by an arithmetical argument. Calculating the amount of menstrual blood evacuated in nine months, he said,

The quantity of Blood which the Mother may bestow upon the nourishment of her Ofispring will be lib. 13 nzt.2§, which will outweigh the newborn Foetus with all its Integurnents, if they should be put into a balance; and leave no room to doubt, its being able to bestow very proper nourishment on the Embryo. For the mean weight of a new-bom Foetus is about 12 lx'b., sometirns it is found greater, and very often less.

This quantitative outlook forms a parallel to Harvey's approach in his famous calculation about the circulation of the blood.

Freind's view that the matemnl and foetal circulations were continuous was derived from the experiments of Rayger and Gayant, who had injected a blue dye into the foetal circulation and found it again in the maternal. Vl/orlt of this kind had begun as far back as about I555, when apparently Axuatus Lusitanus had made similar observations on a woman, and in the Cracolfia of J. F. I-lertodt, published in 1671, where under the heading “An erocus foetum tingat in utero ?" we find a description of the public dissection of a pregnant dog to which this dye had been given in the diet. The embryos were markedly yellow.

' Cf. the interesting recent resume of W. schopfer.

' Biographical detail: of Freind in an essay by Greenwood.

' Muller-lies: has written I monognph on the development of knowledge on menstruation from the sixteenth century ouwuds.


I. That the embryo was nourished directly by menstrual blood.

Beckher, 1633.

Plempxus, 1644. Plcmpius did not deny that the umbilical cord was functional, but insisted that the blood passing through it was menstrual. In 1651 Harvey’: vmrk was published.

smnemfi. 1554» F. Sylvius, I680.

Seger, 1660. Cyprizmus, 1700.

van Linde, 1672.

II. That the embryo was nourished through its mouth. (:1) By the amniotic liquid. (i) In addition to the umbilical blood.

Harvey, 1651. Linsing, 1701. W. Nctdhaxn, x667. Pauli, 1717;. de Gruf, 11977. Barthold, x717.

C. Bzrthulihus, 1679. S. Middlebezk, 1719. van Diemerbroeck, 1685. Teichmeyer, 17x9.

Ordob, X697. Gibson, 1726. D. Tauvry, 1700. (ii) Alane; the umbilical blood being regarded as unneces- mry or of minor importance.

Moellenbrcck, r672. P. Stalpartius, 1687. Everardus, I685. Bierling, 1690.

Case, 1696. Case thought the embryo arose entirely out of the amniotic liquid like a predpitate from 3 dm solution; see, p. 184.

Berger, 1702.

These writers assumed as their prindpal experi- mental basis reports of embryos born without umbilical cords, e.g. those of:

Rommelius, 1675 (in Velsch).

Valentinius, 1711.

(17) By the uterine milk or mu-um Iadzo-diylamm.

Mercklin, 1679. J. xvaldschmidt. 169:- Drelincurtius, 1685. Tauvry. I694- Bohnius, 1686. F73“? 1732' Zacnhias, x688. D3011”: 177-4-

III. That the embryo was nourished through the umbilical cord only. (a) By foetal blood (the circulations distinct).

Arantius, 1595. Snelle. 17o5. Harvey, 1651. Falconnet, 1711. W. Ncedham, 1667. F. Hoffman, 1718. Ruysch, 1701. Monro, 1734.

It is to be noted that Bicrling, P. Stzipnrtius, Berger, Barthold and Charleton, who supported the discontinuity theory of the circulations, were all upholders of the theory of foetal nourishment per 0:, so that their reasons for doing so were not those on account of which we agree with Hofimann and Needham at the presmt time.

(b) By maternal blood (the circulations continuous). Laurentiuz, 1600. Hamel, 17oo. de Marchette, 1656. de Craan, 17o_-4. Rallius, 1669. Lang. 1704. Muraltus, 1672. van Home, 17o7. Blasius, 1677. Freind, 1711. Vcslingius, 1677.

de Méry, 1711. De Méry comhated Faloonnet's view of the separate circulations. He said that he had not himself tried Falconnetla experiments, but that some students had, and could not repeat them.

Aubert, 1711. Narrative of a use in which the um- bilical cord had not been tied at the maternal end and the mother had nearly bled to death through it.

Nenterus, 1714. Wedel, 1717.

Bellinger, 1717. Bellinger believed that the maternal blood was transformed by the embryonic thymus gland into proper nourishment for itself, after which it was secreted into the mouth by the salivary ducts and so went to form meoonium without the necw sity for dcglutination. Heister's comments on this extraordinary theory are worth reading. Perhaps Bellinger was indebted to Tauvry for his idea of the importance of the thymus gland. Tauvry had dnwn attention in 17m to its diminution after birth.

dc Smidt, 1718. Dianis, 172.1.

(continued overleaf) 181 A A 1-nsronr or

new 1 (tonlirwed)

(t) by menstrual blood. Plempius, 1644.

(J) By uterine milk.

Em, x687.

Cameraxius. I714. (Opfnfo concilialrixl)

F. Hotfmann, 1718.

(t) By the amniotic fluid. Vicafills. 170°: Goelicke, 1723. IV. That the embryo was nourished through pans in is skin,

Deusingiua, 1660. Stockhamer, X682.

Nitzsch, 1671. This was suggested on the ground that in the earlier stages of development there is no umbilical cord. In 1684 de_St Romain argued against it on the ground 1-bit: if I! Went true, the embryo would dissolve in the amniotic liquid.

During this period also there were continued disputes about the origin of the arru-riatic liquid. Van Diemerhmeck and Verheycn con- sidered that it could not be the sweat of the embryo, for the embryo was always much too small to account for it, and, moreover, du Tertre had described cases where the secundincs had been formed with the mem- branes but in the absence of the embryo. Dionis aflirmed that whatever it was it could not be urine, for urine will not keep good for nine days, afnrtian‘ not for nine months. Drelincurtius put forward a theory that the embryo secreted it from its eyes and mouth by crying and salivating, while Bohn and Blancard derived it from the foetal breasts. Lang, Berger and de Gouey criticised this notion without bringing forward anything constructive, and de Gouey was in his turn annihilated by D. Hoffinzmn, who with Nenter and Kiinig supported the modem view, namely, that it was a transudation from the maternal blood-vuseis in the decidua. The question was complicated further by the alleged discovery by Bidloo in 1685 of glands in the umbilical cord, and by Vieussens in 1705 of glands on the amniotic membrane. J. M. Hoffmann and Nicholas Hoboken supported the view that these were the impor- tant structures. There the problem was left during the eighteenth century, various writers supporting different opinions from tune to time, and it is still not fully solved.

2. Growth and Differentiation; Stahl and Maitre-Jan

Very early in the eighteenth century (1708) there appeared a work by G. E. Stahl, van Helrnont’s most famous follower, which struck the keynote of the whole period. Stahl‘s Thearia Zlrledim Vera, divided as it was into physiological and pathological sections, belonged in essence to the a prion‘ school of Descarts and Gassendi. It differed from them profoundly, however; for instead of trying to explain all biological phenomena, including embryonic development, from mechanical first principles, it started out from first principles of a vitalistic order, and, having combined all the arthaei into one informing soul, it sought to show that the facts could be convincingly explained on this basis. The spiritual kinship of Stahl with Desmrtes and Gassendi is due to an atmosphere which can only be called doctrinaire, and which was common to them all. Like the methodist school of Hellenistic medicine, they subordinated the data to a preconceived theory.

Stahl is also interesting in that he represents a trend of thought which favoured what has been called "instantaneous generation followed by metamorphosis.” Cole calls this the “precipitation” theory! but I cannot altogether accept his account of it.

"Metamorphosis” was defined by Harvey (1653, pp. 222 if.) m the bringing into being of a formed object from a mass of material previously possessing no form; as opposed to “epigenesis," where assumption of form and increase of mass proceed simultaneously.

Artificial productions are perfected two several waies; one, when the artificer cuts and divide: the matter which is provided to his hands, and so by paring away the superfluous parts doth leave an Image remaining behinde, as the Statuary doth; the other, when the Potter forrnes the like Image of Clay, by adding more stuff, or augmenting, and so fashioning it, so that at one and the same time, he provides, prepars, fits, and applia his materials. Harvey's "metamorphosis" we should now call "differentiation without growth” and his “epigenesis” we should call "differentiation plus growth." It should be carefully noted that to epigenesis Harvey does not oppose “preiorrnation,” for he was writing some thirty years before Malpighi’s unfortunate surnmer-time experiments, and Joseph de Aromatari's seed had not yet sprouted. "Preformation” in modern temts would correspond to "growth without differentiation," all the complexity of the finished form being supposed to be present initially. “Metamorphosis" in Harvey’s mind was nothing more nor less than the Aristotelian blood-and-seed theory, and his description of :1 sculptor making a statue out of a pre-existent mass can best be understood in the light of Ruefl"s drawings (Fig. 10). Harvey opposed Fabricius precisely because the latter ventured to point to the chalazae in the hcn's eg as the homologue of the mammalian blood-and-sccd,

But the relations between growth and difl'erentiation were still open to different opinions in the eighteenth century and many believed rm: '1" former was by far the more impomm ofthe two. If the differentiation process was pushed far enough back in the life of the embryo the distinction between epigenesis and prefornation tended to disappear We have already had (p. 167) in Croone an example of a prefer-rnationisti Who believed illogically that “instantaneous generation” was followed by "metamorphosis,” and Stahl is an example of an epigenesist who thought that the governing artlmeur provided by the semen organised whatever it found in the uterus into the form of the body, after which there was pure growth and no further differentiation. The distinction thus became almost academic.

After the mysterious organising process the entire remaining business of generation [said Stahl‘] is taken up with the fomzation of the body, which from the first rudiment, so to speak, is nothing afterwards but nutrition; namely, such as is carried on from this time up to old age; while in the body what is once completely shaped is not merely preserved by a perpetual supply, nor, if it pen-hanoe fails, is it merely rebuilt, but in fact it continually grows until it is completely formed in all its pans. A perpetual assimilation everywhere accompanies apposition, or rather the apposition itself is set up immediately in such order and situation that assimilation is brought about and exists bemuse of that very position. . . . That Principle which unfolds its activity primarily in the brain and the nerves prmides over the formation of the body, and those parts which constitute the only immediate instrument of in action: being formed first of all, make it probable for that reason that something ought to, or at least an, be provided by themselves alone. . . . As for how the blood is generated, it’ we are to be- lieve the theories commonly held today, it is thought to be born from a spontaneormalgllidhtg-together and chance meeting of particles uniting them- selves mutu y.


(A) EPIC!-‘N515 E Diflerentiation + Growth Harvey

(B) PRErDR.\tA‘I10. Growth alone Malpighi. Sw:rnmcrdzm.€t€- (C) luzrluuoxrflosts Differentiation alone Aristotle, Fabriciu:

(D) PRECIPITATION E (A) in the very early Sub]. C359

stages, followed by (B) (E) (B) in the very early stages, followed Croone by (C) (F) (C) in the very early stagm, followed Butfon ‘by (B) Thewid. on 425 S-

In 1722 Antoine Maitre-] an published his book on the embryology of the chick, the only one on this subject between Malpighi and Heller. It was an admirable treatise, illustrated with many drawings which, though not very beautiful, were as accurate as could be expected at the time. Perhaps its most remarkable characteristic is its almost complete freedom from all theory — Maitre-Jan says hardly a word about generation in general, and is far from putting forward a "system" in the usual eighteenth-century manner. He contents himself with the recital of the known facts including those added by his own observations. He gives no references, and writes in an extremely modem and unaffected style.

The only traces of theoretical presupposition which can be found in him are Cartesian, for he speaks of the activity of fen-nents in blood- fomzation. He is an epigenesist, and long before Brooks, he gives the right explanation of Malpighi’s error, affirming that the hot Italian summer was responsible for some development in Malpighi’s eggs before Malpighi examined them. Although Maitre-Jan’s book must have been accessible both to Buffon and Haller, they perpetuated Malpighi’s mistake till nearly the end of the oentury.

In technique, Maitre-Ian was pre-eminent. He was the first embryologist to make practical use of Boyle’s suggestion regarding “distilled spirits of vinegar" for hardening the embryo so that it could be better dissected.‘ He also used “weak spirits of vitriol"; after treating blaste- derms with it, he said, “I saw with pleasure an infinity of little capillary vessels which had not appeared to be there before” (see Plate XVII, facing page r86). He made a few chemical experiments also, noting that vinegar would coagulate egg-white, and estimating quantitatively the difference in oil-content of different yolks—though for this he gives no figures.

His theory he relegated to an appendix entitled Objettimz: :10‘ la génératian dc: animaux par de petilr em‘. There were sixteen of them, but the most cogent one was that, as little worms had been found under the microscope in pond-water, vinegar and all kinds of liquids, there was no reason to suppose that those in the semen were in any essential way connected with generation. For his time, this argument was an

‘ His Ietuzl words are as follows:

p. nu. "Si l'on verse dan: cet zzuf (70 hours) nu dans un autre de pareil tems de oouvée, du vinaigre distillé, on verrn tn peu de tam: le fetus blanehir ex devenir plus solid: at xi opnque, qu'on no | uroit plus dixtinguer nu lntvers lea vésiculu du cczur, m mam: lea vusseaux qui y s utissent, hon celui qui régne le lung du Carpl."

p. 148. :']e du-ni enfin que quoiqu‘on n: décourre quhssez ebseurément ls plflpnt ties pnnapales parties intérieureu, hon le ctzur ct quelque: vnisseaux (144 houn), I cause d:_ lcur trop dc moluse et de la viseoaité de tout le fetus, elles ne laisxent pas que d'A_vmr sen quelque farme, comm: on 1: oonnolrn en fnisanr infuser le ftztux dun le vinugre dxsulé."

excellent one, and was . exiebrimen: which hflli)xX’1el)[ity‘:;:I?):leer:n!1!l]:§:VE3S:;I;l‘1)c §:oIr:1!1:)of filtration out I 's time ' . ‘ _ blood, the fwamm ézgfélisiiilrgeezgtrgvexrsrygter the circulation of “"1 d° M57 “"5 mgflgtd in a pnlcmi<?ori this stil7)?c:i° I7.i‘:'1,Tauwy also corrwponded with Duvemey, snvcstm and ear: . e latter “'°V°’5}’ “'hi°l\ refills that of Laurcntius and Petreus a hunrirn L: can. before. Nichnlls wrote later on the same sulfect Daniel Ta e yyurs i"‘°’°51i“8. h°W€V€1'. for other reasons. He an epi Wag

agtinst the view that the soul constricted diuiinn cmbryvgeny a suitable home for itself. 3

Nine years later two books appeared which form very notable land-

in the history of embryology. One was Martin Schurig’s Embryo- IUEW. and the other the Elementa Clxemiae of Hermann Boerhaave

Th: {°m‘"v h°W¢V€|'. gave to the world no new experiment‘; of °b5C1'V3"'0fl5; it Was the first of What we should now call the typical “'°"i°“"' kmd °f Plllflimtion. Schurig saw that he was living at the end of :1 great scientific movement following the Renaissance. and set him. self accordingly for many years to Compile large treatises on specific physiological subjects, taking care to give all references with meticulous accul'-“»'}'- and to omit no work, significant or insignificant. His Spam. tologfa was the first to appear (in 1720), and it was followed in 1723 by Sialologia (on the saliva), Chylologia (1725), Muliebria (1729), Parxlmm. 1031}! (1719). Gyflllttnlogia (1731) and Hazmatologia (1744). His Embryo- Iogm was the last but one of the series. In it he treated compendiously of all the theories which had been advanced about embryology during the immediately preceding two centuries, and his chapters on foetal nutrition and foetal rcspiration throwa flood of light on the "intellec- tual climate" in which Harvey and Mayow worked. Schurig‘s biblio- graphy is a very striking part of his book, extending to sixteen pages, and inlfll-‘l.‘l:.i-llg five hundred and sixty references, it was the first attempt of its ‘ .

3. Chemical and Quantitative Approaches to the Origin of Organisation; Boerhaave, Hamberger and Mazin Hermann Boerhaave was a more prominent figure, a professor at Leiden for many years, and renowned for his cncyclopaedic learning on all subjects remotely connected with medicine. His Elemmta C/zhniae, which became the standard chemical book of the whole period, demonstrates throughout the exceedingly wide outlook of its and contains in the second vmlume what must be regarded as the author, I reproduce here the first detailed account of chemical embryology. 186 PLATE XVII

llluxlralmmfram Antoine fllailn-]n1|‘x Obsenutions sur la Iormznon du poulcl, oh la: dnen changtmms qui arriu-nx i l’<:ufi mcsun: qu'|l 2:: com é, mm exacxemem cxpliqufi ct rcprésnntéa en F: ns D‘IIour_v. Parix, 172:.

A. Drlvungs olcmbryos of rum. zoc»15o hours inzubznon.

n. The am dnlung u! lh: fillosines on uh: inlumr of the yoli-sac.

tp. 2,2) “um: pike .1. 1. “mad: membrane de_1:-me sur 1. ...,msc.e mttnmre at hqurlle on wit ..x..,.=un nnglu d: m penis vusiuux cmnrnllées dc dnusn grlndeurs ti dxlhrement amass" L179 hrs 1 c. Photaznph at the .-.11:-mm. from Rcrnoui. They puy m impomm pan m mg Ibsotpllnn n( the nu.

relevant passages in full because of their great interest. It will be noted that they are cast in the form of lecture addresses, as if they had been taken down direct from the lectures of the professor, a fact which gives them a peculiar charm when it is remembered how many great men must have listened to them, among them Albrecht Von Heller and Julien de la Mettrie. In considering what follows, it should be noted that Boer- haave's interest is biological all the time, and that he does not trmt the liquids of the eg. as nearly all the chemists before him had done, as substances of curious properties indeed, but quite remote from any question relating to the development of the embryo. Another interesting point is that he deals only with the white, and hardly mentions the yolk; this is perhaps to be explained by the Aristotelian theory‘ that the embryo was formed out of the white, and only nourished by the yolk (ex albofieri, ex luteo nutn'n'), a theory which was still alive, in spite of Harvey, in the first half of the eighteenth century. If this was what was at the bottom of Boerhaave's mind, then it is obvious that the egg-white would be to him the liquid inhabited more particularly by the plastic force. This, then, is what he has to say about the biochemistry of the egg.

0]. Chem. in Am'nmIz'a. (Processus log.) The albumen of A fmh egg it not add, nor alluzline, nor doe: it contain afzrmenled .tj:x'n'l. The white of a fresh egg, separated from the shell, the membranes and the yolk, I enclose in clean glass vessels, and into each of these I pour different acids, and shake them up, mixing them, and no sign of ebullitiou appears however I treat them. Thero- fore I lay these vessels aside. Now in these other two vessels I have two fresh portions of albumen, and I mix with them in one case alkaline salt and in the other volatile alkali. You see they are quiet without any sign of effervescenoe. Now behold a remarkable thing, in this tall cylindrical vessel is half an ounce of the albumen of an egg and two drains of spirits of nitre, in this other vessel is half an ounce of egg-white, together with four and a half ounces of oil of tartar per deliquium both heated up to 92 degrees. Pray observe and behold, with one movement I pour the alkaline albumen into the acid albumen, with what fury they boil up, into what space they rarefy the mass, so that they stream out of the vessel although it is ten pints in size (decupli eapace). They have scarcely changed their oolour. But when the eifcrvescenoe has abated how suddenly they return to the limits of space occupied before. But now if more egg-white is heated to 100 degrees in a retort (nu-urbihz) an insipid water containing no spirit is given oil’. If egg-white is applied to the nalted eye or naked nerve it does not give the smallest sense of pain, and scarcely affects the smell; nothing more inert and more insipid can be put on the tongue. It appears muoous and viscid to the touch, not at all penetrable. Hence in the fresh white of an egg there is no alkali or acid, or both together. It is indeed

3 ‘hick, Slick ‘. incfl. and insi id li uor, e ‘ ' ‘ - heat of 93 degrees within the sgaee oci 2t dz): ;::.i,'§§:,'§,ff,§ egg from a tiny mass hardly weighing a hundredth of a grain into the perfect b°‘:1Y Qf 311 31151113]. Weighing an ounce or more. We have learnt therefore of a liquid distinct from all others, from which by inscruublc mum mm, membranes, vessels, entrails, muscles, bones, mrfilaggg’ and 311 lb, cum: parts, tendons, ligaments, the beak, the claws, the feathers, and all the humaurs can be produced-and yet in this liquid we find softness inertia absence of Mid. alkali, and spirit, and no tendency to eflervesce. Indeled if there were the slightest efl'ervescenee in it, it would certainly break the eggshell; therefore ne see from how slow and inactive a mass all the solid and fluid parts of the chick are constructed. And yet this iself is rendered absolutely useless for forming the chick by greater best. It smrcely bears no degrees with good effect but at a la: temperature never brings forth a chiclr, lorunder 80 degrees will not sutfioe. But by a heat kept between these limits, there is brought about so marvellous an attenuation of the mucous inactivity that it can exhale a great part through the shell of the egg and the two membranes, the yolk and chalazae alone remaining slang with the amniotic sac. For the yolk, the uterine placenta of the chick, takes little part in the nourishment. Meanwhile Malpigbius has shown that this albumen is not a liquid of a homogeneous kind, as the blood-serum flowing through the vital vessels is, but that it is a structure composed of numerous membrane-like and distinct small sacculcs, filled with a liquid of their own, in the same way as in the vitreous humour of e eye.

(Prooesus xxx.) Eajblonzlicm of the egg-white u-ill: almlrol. In this transparent vessel is the albumen of an egg, and into it, as you perceive, I gently pour the purest alcohol, so that it descends down the sides of the vessel and reaches the albumen. I do this deliberately and with such solidtude that you may see the surface of the albumen which, touching the alcohol, holds it up, being immediately coagulated, while the lower part remains liquid and transparent. As I now gently shake them together, it appears evident that wheret er the alcohol touches the albumen a onncretian is formed. Behold now, while I shake them up thoroughly together, all the egg-white is coagulated. If alcohol previously warmed is employed in this experiment, the same result is brought about but more rapidly. It appears therefore that the purest vegetable spirits immediately coagulate the plastic and nutrient material.

(Prooessus :12.) Thefresh albumen afar: egg is broken up by di'm'!hzti0n. These {rah egg have been cooked in pure water till they become hard. I now take the shining white sepanting off all the other things and break it up into small pieces. I put these, as you see, into a clean glass retort (nu-mbiln) and I duly cover it by fitting an an alembic and add a receiver. By the rules of the [chemical] art I place the whole retort in I bath of water and I apply to it successive degrees of fire until the u hole bath is boiling. No vnporous strealzs (mine) of spirits are given off but simple water detvy drops and this in incredible quantity, more than nine-tenths. I eonunue so with patience until by the heat of boiling water no more drops of thn humour are given 05.

Then this water shows no trace of oil, salt, or spirit; it is perfectly transparent and tasteless, except that it eventually grows rather sour. It is odourless, save that towards the end it gives ofi a slight smell of burning. It shows absolutely no sign of the presence of any alkali, when I test it in every way, as you can see for yourselves; nor does it reveal any trace of acid, when tried how you will. Here you see pounds of this water, but in the bottom of the now open retort see, I beg of you, how little substance remains. Behold, there are fragments contracted into a very small space in comparison with the former quantity. They are endowed with a golden oolour especially where they have touched the glass, but yet they are transparent after the manner of coloured glass. When I take them out I find them very light, very hard, quite fragile, and breaking apart with a crack, smelling slightly of empyreuma, with a taste rather bitter from the fire, and without any flavour of alkali or acid. This is the first part of the analysis. Now I take these remaining fragments in a glass retort (retorlam) in such a way that two-thirds remain over. I put the retort into a stove of sand, first arranging a large receiver. Then thoroughly luting all the joints I distil by successive grades of fire and finally by the highst which I all supprrstianis. There ascends a spirit, running in streaks [striatim] fat and oily, and at the same time, volatile salts of solid form everywhere on the walls of the vessel, rather plentiful in proportion to the dried fragments but small in proportion to the whole albumen before the water had been removed from it. Finally an oil appears besides the light golden material mixed with the first, black, think, and pitchy. When by the extreme force of the fire this oil is finally driven forth, then the earth in the bottom, closely united with its most tenacious oil, swells up and is rarefied and rises right up to the neck of the retort so that had the retort been overfull it would have entered into the neck and clogged it up even causing it to burst with danger to the bystanders. The operation is to be continued till no more comes out. That first spirit, oily and fatty, is clearly alkaline by every test, as you may tell from the way it effervescm when acid is poured on it. If we rectify it we resolve it into an alkaline volatile salt, an oil, and inert foetid water. The salt fixed to the walls is completely alkaline, sharp, fiery, oily, and volatile; and the final oil is specially sharp, caustic, and foetid. The black earth which remains in the retort is shiny, light, thin, and fragile, foetid from the final empyreurnatic oil, and soft because of it. If then it is burnt on an open fire, it ‘mrres a ‘i':'t‘t're ‘i-med ta-nh 1v‘rnt.’n is ‘wloote, , tasteltn, and odumitss, from which scarcely any salt can be extracted, but only a very heavy dusty powder (poIIx'nem).‘

(Processus H3.) Tlrefrerh albumen :1] an Egg zcillputrefy. Sound eggs kept at 7o“ for some days will become foetid and stink. . . . \Ve have learnt then that this is the nature of the material which will shortly be changed into the struc- ture, form, and all the parts of the animal body. Repose and a certain degree of heat produce that effect in that material. We observe therefore the spon- taneous eorruption and change of the material, ahd what is extremely re- markable, if an impregnated egg is warmed in an oven (in hypocaurrir) to a

‘ Cl. the dry distillation of egg-white by Pietet 5: Crime: in 1919.

heat of g: degrees it employs these attenuated parts changed by such a heat

0 fl¢_"1HSl1. Incrensc._nnd complete the chick for 2x days. But in this chick nothing alkaline, foettd, orputnd L! found, henoe observe, 0 doctors (rnedt'¢1') the remarkable manifestations of Nature—by repose and a certain degree of heat a thick substance becomes thin, a viscous substance becomes liquid, an odourlex substance becomes toetid, an insipid substance becomes sour and extremely acrid and bitter to the taste, a soothing substance beoomea caustic, a non-alkali become: alkaline, a latent oil becomes sweet and putrid. Let these results be compared with the observations of Llarcellus Malpighius on the incubated egg, and we shall observe things which shall surprise us. I took are to investigate only the albumen of the egg Erst of all, separating the other parts off where possible, for the albumen alone forms the whole of the material which proceeds to feed (in pabulum) the embryo. The other constit- uents of the egg only assist in changing the albumen, so that when it is changed, it may be applied to farming the structure of the chitin

Boetl1aave's treatment of thae subjects has only to be compared with that of Joachim Beecher, who wrote in 1703, to show how thoroughly modern in outlook it is. Beccher's Pllyrita Sublerranea contains a whole section devoted to the growth of the embryo, but it is extremely con- fused and very alchemiml in its details.‘ The advance made in the thirty years between Beecher and Boerhaave was immense, but, if the bio- chemistry of development advanced so fast, its biophysics was not far behind, as is shown by the work of G. E. Hamberger and I. B. Mazin.

H:trnl>erger’s most important contributions, contained in his Plryn'o- lagia Media: of x751, were his quantitative observations on the water- content of the embryo and its growth-rate, in which he had no fore- runners. Hamberger showed that there are much less solid pans in the foetus than in the adult. The cortiml substance of the brain of an embryo loses 8694 parts in ro,ooo on drying but in the adult it only loses 8096 and that of the cerebellum from 8t parts is reduced to 12. The maxillary glands of the embryo lose out of ro,ooo parts 3459. the liver 8047, the pancreas 7863, the arteries 8278 and even the cartilage: lose four-fifths of their weight decreasing from xo,ooo to Buy}.

The corresponding figures for the adult were: liver 7r92, and heart 7836. These figures do not widely difler from those obtained In recent

Mazin published his Conjecture: physiro-medico-12;,-drottoiicae d: Respiratione Foetu: in 1737 and his Tradatur Zlleniico-meelxonxta in 174-3- In the first of these works Mazin supports “hat is rsserrtially Mayow s theory of embryonic respiration, without, however, mentioning hlayow more than once. It had not been popular since 1700, though Prtcarm 1 BL :, mt. iv, ch. 4. p. 1111, “De mixfione

had defended it. Mazin put the liquids of eggs under an air-pump, and observing that air could be extracted from them aflirmed that the air was hidden in them and that the embryo could therefore respire. He spoke of “aerial particles" in the amniotic liquid, and discussed the respiration of fishes in connection with this. The specific gravity of the embryo also interested him, and he did a great deal of calculation and experiment on it. His most interesting passage, perhaps, is that in which he mentions the “eolipilc" of the Alexandrians, the primitive fonn of the steam- engine, and says thatjust as the heat of the fire makes the water boil, so the heat of the viscera makes the amniotic liquid boil, giving ofl’ respirable vapours. The time-relations of this analogy are interesting, for by I712 Thomas Newcomen had succeeded in making a stearn~ engine which worked with considerable precision, and the question of steam-power was widely dismissed. Possibly Mazin was acquainted with the Marquis of VVorcester’s Century of the Name: and Standing: of Inventions, which had been published in 1663, and which had contained an aeolipile or “water-commanding machine.” England was the centre of this movement and other countries employed Englishmen as engi- neers; Humphrey Potter, for instance, erected astearn—engine for pump- ing«at a Hungarian mine in 1720.

As for the discovery of oxygen, it was near at hand, and Scheele in 1773 and Priestley in 1774 were soon to supply the knowledge without which Mazin could not proceed further.

In his second book, Mazin reported many quantitative observations on the specific gravity of the embryo. He found that it diminished as development proceeded, being to the amniotic liquid as 282 to 274. in the fourth month and as 504 to 494 in the fifth month. His work on this subject was continued by Joseph Onymos, whose De Natura Foetur appeared in r745. .

R. I. Raisin also contributed to this wave of precise measurement in embryology. His dissertation of 1753 took account of the difference between the pulse rates of infants and adults, and contained an arith- metical argument about the prenatal secretion of the foetal kidneys. But it also gave a list of the relative weights of organs, showing that some decreased and others increased relatively to the weight of the body as a whole. Thus the brain was one-tenth part of the body in the foetus and onc~twenty-fifth part in the adult (see Table II, overleaf).

It was the first mention of heterauxctic growth,‘ save for the isolated observations of Leonardo (see p. 98).

About this time. we get occasional references to the obscure mechan- isms oontrolling animal growth. Although the brilliant speculations of Marci (see p. 81) had long been forgotten, some writers, such as


F tut I Cenbnm . . . . . r]'ro 11/12?” Pulmo . . . . . r/66 r/r7 77‘J>mu: . . . . r/37.4 1/4560 90' - . . . I/189 r/1 r4 HfP'"' - . . . 1/23 1/21 I-"" - < - - - 1/324 1/175 Pancreas . . . 1/907 1/445 Vmtntulur vacmu . . . 1/767 1/212 Rene: . . . . . r/154 I/136 Surrenaler glarzdulae . . I/324 1/3040

Iames Parsons, were groping about for the rnorphogenetic controls. In his Philoroplrital Observation: of 1752 Parsons had a good deal to say about “primary" and "subordinate organisations,” notions which have a certain resemblanoe to the field theories of modern embryology, for a short account of which the article of Waddington may be consulted.

Parsons said of his organisations:

There can be no more natural Way of answering a Question proposed bya Gentleman of Penetration in Philosophical Knowledge, which is Why do not Animals and Vegetables grow on without End? Why do not Seeds, when they are perfectly form'd, grow on in their Poet, Husks, or other Receptacles? Because, says he, when a Body has once begun to grow, the same Propensity for growing on ought still to continue, and, the Particles of Matter increasing too, it ought not to cease.

The answer of Parsons was that the organisation comes to "its full Power of Distention, so far as is consistent with its natural Form," after which further nourishment becomes useless and the structures all

Parsons was on the verge of a field theory, for he gave much consideration to the regeneration arperiments carried out by Trembley and others on fresh-water coelenterates (cf. Baker). But he did not develop his idea far enough to escape the objection that the "organisa~ tions" were mere abstract sirnulacra of the visible forms of the animals and plans themselves. For the test, he was a convinced ovist, accepting Nuck's experiment (see p. 163) in the wrong sense (contrast with Mamuet, see comment on p. 209), and desirous of explaining all genera- tion as budding or "propagation." Like Galen long before him. ‘he conducted a lively polemic against all formulations of the C13 plartztd, but in favour, unfortunately, of the direct action of Cyod. In the succes- sive action of his primary and secondary

I The classical modem treatment of the relative growth-nit! of mm cf °rz=fl"=m' is of umne that of Huxley.

approach rather closely the modern conception of a succession of organisers or inductors in development (see Aristotle's ideas on this, p. 48 passim, and the references to modern embryology there given).

These writers, together with Haller himself and J. C. Hefiter, who handled the problem of embryonic growth rate, contribute to one of the best, because most quantitative, aspects of eighteenth-century embryology.

4. Albrecht van Ballet and the Rise of Techniques

Boerhnave‘s greatest pupil was Albrecht Von I-laller} Like Oliver Wendell Holmes at Harvard, Haller occupied a “settee" rather than a "chai.:” at Gfittingcn, and taught not only physiology but also medicine and surgery, botany, anatomy and phamtacology as well. Nor did he merely deal with so many subjects superficially; in each case he published what amounted to the best and most complete text-book up to then written. Haller was made professor in 1736, and for many years worked at Gottingen, devoting much of his time to embzyological researches, which, with those of his opponent Wolfi, stand out as the greatest between Malpighi and von Beer. In 1750 he published a series of disser- tations and short papers on all kinds of physiological subjects, which would have been the direct ancestors of the modem compilations by groups of experts, had they been more systematically arranged. The volume on generation repays some study. The contributions relevant to the present d'scussion. had been written at various times during the previous seventy years, and may be summarised selectively as follows:

IV. Christopher Sturmius, Deplrmtarum animalimnque Generation: (first published 1687). In this paper Sturrnius argues on behalf of the pre- forrnation theory, "which in our times does not lack supporters,” quoting Perrault, Harvey and Desmrtes. He contents himself with countering arguments which had been urged against it, as, (a) spon- taneous generation, (6) annual recurrence of plants, (c) insect meta- morphosis, (-1) generation without copulation.

V. Rudolf Jacob Carnerai-ius,' Specimen Eajverimentomm pIr_y.rt'o- lagim-thrrapeuiicomm circa Gmerationevn hominir el animalium. The most interesting thing about this is that Camerarius mentions the observations of D. Seiller, a sculptor, who had ascertained that the body is five times the size of the head in the embryo but seven and a half times the size afit in the adult. This is in the direct line between Leonardo and Scammon.

XV. Philip Gravel, D: Superfzlalione (first published 1738).

‘ See d'I1sx3'. ' Afterward: furious as the discoverer of Izxualiry in planu.

n.L—~r3 I93 A rrrsrorw or-' rmaxronoay

XVIII. Adam Brettdel, De Embrymrz in 0911/0 mile amteplum pm:-e.u':tan1: (first published x703). Breudel "stands for :1-.¢ Gmfim hyPo_ thesis. Unfortunately, he was also_a preforrnstionist and believed that every limb, organ and function qisted not potentially hut alaséueally in the urtfemlised eggbefore its passage down the Fallopian

XXII. Camillus Faloonnet, Non at friui Szmguir Itzgzmm az;,,,,,,;g (am

V published 1711). This is the first of the French u)mfibufi¢m_q M m,

book; they are all very markedly shorter than the Gemnn ants and much less heavily omnmented with irrelevant quotations. 1-‘alommgg

is concerned to prove that the maternal and foetal circulations are separate, and he describes in an admimbly eoncise manner ztn etperiment in whid: he bled a female dog to death, after which, opening the uterus, he discovered that the embryonic blood-vessels were full of blood although those of the mother had none in at all. Ar-antius was therefore justified. Falconner was soon confirmed by Nunn.

XXIII. Jean de Dir.-st’s Sui Sargguinix rolur gag’/es: Fetus at [first rzublished 173 5) was written to prove a similar point. He refers to the experi- ment of Falcounet and the injections of F. Hofimann, and criticises Cawper’a experiment in which mercury had been injected into the umbilical vessels and found in the matemal circulation. on the grounds that mercury is so “tenuous and voluble" that it might pass where blood could not pass normally. He also objects to the view that the foetus is nourished by the amniotic liquid.

XXIV. Francis David Herissant, Szmrrdinaeferui Pulmonir pnmfnnl aflicid,

1 sanguine maltnw Fetus non alum (firs: published in X741). An excellent paper in which the respiratory function of the placenta is proved by the observation that the foetal blood-Vessel leading to the placenta is always full of dark venous blood, while that leading away from the placenta is light and arterial (flan‘dx'an‘ mrdnmque mlore, ul iprzrnel obsemzd). Herissant adduces also the cases of acephalic

monsters, such as that of Brady, which could not poxibly have drunk up any amniotic fluid, and yet were fully formed in all other respects. He concludes that the urnbiliml cord serves for respiration and nutrition.

XXV. After these three French workers, there is a great drop to Johnrtnes Zeller, whose Injanlicidm mm abralt-it nee a tarlrna liberal Pubnmmn Infantir in aqua nab:-idrnlia (first published 1691) is a long-winded discussion of the floating lung test in forensic medicine. I-{is memory deserves a word of ohloquy for his vigorous insrst.ence‘upon death and torture for infanticide even during puerpenl insanity. Perhaps it was Zeller who called forth the noble answer of de la Mettrie to this inhumanity in his Man a fllaclxint.’

XXVI. Zeller’s De Vila humamz 2: F rm: pendent: (first published X692) is no

' For a detailed hisrorial amount of this tut and its unreliability, lee Knmmer.

better, though at the time, perhaps because of its striking title, it was famous. It deals with the ligation of the umbilical cord at birth.

Such were some of the typical papers printed by Haller in his 1750 collection. He retired from the Gottingen chair three years later, and in 1757 the first volume of his Elementa Phyrialagine was published, probably the greatest text-book of physiology ever written. It appeared only by slow degrees, so that it was not until 1766 that the embryo- logical section was available. This volume contains a discussion of a mass of literature, most of which had arisen during the preceding twenty-five years, for although many of the names mentioned by Haller occur also in Schurig, many are quite new.

Holler himself published in 1767 a volume of his collected papers on embryology, most of which were concerned with the developing heart of the chick, which he worked out very thoroughly, in collaboration with Kuhlemann. Kuhlemann had already shown in the sheep what Harvey had proved for the doe. But Haller was a convinced prefomia- tionist, a fact which was largely due to his researches on the beds egg, where he observed that the yolk had a much more intimate connection with the embryo than had previously been supposed. Since the whole yolk was part of the embryo, as it were, the preformation theory seemed to him to fit the facts better than epigenesis.‘

Haller went further than Schurig in that he usually gave an opinion of his own after summarising those of other people, but his views were by no means always enlightened, and the atmosphere of Buflon is, on the whole, more congenial to us than that of Haller. Haller, for example, believed that the amniotic liquid had nutritious properties, and that the nutrition of the embryo in mammalia was accomplished first of all per or and afterwards per umbilimm. He denied that the placenta had any respiratory function, and indeed his whole teaching on respiration was retrograde. He mentions, however, an experiment of Nicolas Lemery's, in which it had been found that indigo would penetrate the shell of a developing hen's egg from the outside. Consequently. air might do so too, and Vallisneri had shown that, if an egg was placed in boiled water

undcr an air-pump, the air inside would nrsh out through the shell and appear in the form of bubbles.

I-laller was much more progressive in holding the origin of the amniotic liquid (according to him a subject of extraordinary difficulty—- rolurianem non promitlmn) to be a transudation from the maternal blood-vessels. He followed Noorrwyek in asserting the separateness of the maternal and foetal circulations in mammalia. He opposed the

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pr; 2:. Fatrinfile 0/a ms of his obnnwttfau on

I: in A, can Hallzf: “Elexnmtx Phy-u‘u]ogine" 1111766, mm: flu gvowlh In llnglh and weight of berm III the (In: .

existence of eggs in Vivipara—“\Ve may conclude from all this,” he said, "that the ovarian vesicles are not eggs and that they do not contain the rudiments of the animal." But he accepted it in the restricted sense that the embryonic membranes resembled an egg, thus:

If we call an egg a hollow membranous pocket full of a humour in which the embryo swims, we may admit the opinion of the older authors who derive all animals from eggs with the exception of the tiny simple animals of which we have already spoken. It was in this sense that Aristotle and Empedoclcs before him, said that even trees were oviparous. This has also been confirmed by the experiments of Harvey on insects, fishes, birds, and quadrupeds.

Haller’s most original work was in connection with the growth-rate of the embryo; here he struck out, for once, into entirely new country. He made a beginning with the quantitative description of embryonic growth, and one of his tables showing the changing lengths of the bones is reproduced herewith (Fig. 21). He wrote:

The growth of the embryo in the uterus of the mother is almost unbeliev- ably rapidr \Ve do not know what its size is at the moment of its formation, but it is cenainly so small that it cannot be seen even with the aid of the best microscopes, and it reaches i.n nine months the weight of ten or twelve pounds. In order to clear up this speculation, let us examine the growth of the chick in the egg. In this case again we are unable to measure its size at the moment when the egg is put to incubate but it cannot be more than 1%.: in. long, for if it were, it would be visible, and yet 25 days later it is 4 ins. long. Its relation is therefore as 64 to 64 millions or r to r million. This growth takes place in a singular manner; it is very rapid in the beginning and continually diminishes in speed. The growth on the first day is from 1 to gr}, and what Swammerdam calls a worm grows in one day from one twentieth or one thirtieth of a grain to seven grains, i.e. it increases its weight by t4o to 24o times. On the second day the growth of the chick is from r to 5, on the third day, from x to not quite 4, on the fifth day from r to something less than 3. Then from the sixth to the twelfth day, the growth each day is hardly from 4 to 5, and on the twentyiirst day it is about from 5 to 6. After the chick has hatched, it grows each day for the first 40 days at an approximately constant rate, from 29 to 21 on each day. The increase of the first twentyfuur hours is therefore in relation to that of the lastns 5462 to 5 or r45 to r. Nowas the total increase in weight in the egg is to that of the whole growth period (up to the adult) as 2 to 24 ozs. all the post-embryonic growth is as 1 to xz, i.e. it is to the growth of on: day alone early in incubation as 1 to 7§. . . . The growth of man, like that of the chid-z, decreases in rapidity as it advances. Let us suppose that a man, at the instant of conception. weighs a hundred-thousandth of a grain and that a one-month-old embryo weighs 3:: grains; then the man will have acquired in that time more than 3oo,ooo tima the weight that he had to begin with. But it‘ a foetus of the second month weighs 3 ozs. as it approximately does, he will only now have a uired 8 times the mi ht e beginning of the period. This is a ;l’qDdlgi0llS decrease in sgeedl alriildantl end of the ninth month he will not weigh more than about ro5 o-zs. which is not more than an average increase of 15 per montln A child three years old is about half the size of an adult. It‘ then the adult weighs 225:: ozs. the three- year old child only weighs 28i ozs. which is an eighth of the adult weight. Now front birth to 3 years he will grow {mm ms to 281 or as 5 to 14, but in the following 22 years he will only accumulate 2150 ozs. or eight times vthat he had at 3 years. The growth of 3 man will therefore be in the first month of intra-uterine lifeas r to 300,000, in the second as t to 48, in each ofthe others as r to 15. In the first; years ofextra-uterine lifehis growth will hefrom :64. to 281 and in the succeeding 22 years from 281 to 384., and the growth of the first month to the last will he as 300,000 to -5%. or i36,8oo,ooo to 28, or 4,885,717 to 1. The whole growth of rnan will consequently be as io8,ooo,ooo

to I.

In spite of the rather unfamiliar language in which these facts are described, and the theory of the growth of the heart which Haller subsequently put forth to explain them, they remain fundamental to embryology. Their quantitative tone is indeed remarkably modern. In my opinion, when all the voluminous writings of Haller are carefully searched through, nothing more progressive and valuable than these figures can be found. Heller and Hamberger stand thus between Leonardo on the one hand and Minot and Brady on the other. That they stood so much alone is only another indication of the extraordinary reluctance with which the men of past generations assented to the truth contained in Robert M:iyer’s immortal words, “Eine eirizige Zahl hat mehr wahren und bleibenden Wert als eine kostbarc Bibliothelr van Hypothusenl”

Of development as a whole, Haller spoke thus,

In the body of the animal therefore. no part it made before any other part, but all are formed at the same time. If certain authors have said that the animal begins to be formed by the backbone, by the brain, or-by the heart, if Galen taught that it was the liver which wastirst formed, others have said that it was the belly and the head, or the spinal marrow with the bum, adding that these parts malte others_in turn; I think that all these authors only meant that the heart and the bfllfl or whatever organ it wrm, were nsibie when none of the other pam yet were, and that certain parts of the embryoruc

body are well enough developed in the first few days to be seen While 015"! are not so until the latter part of development; and others again not till after birth such as the beard in man, the antlers in the stag, the breasts and the second setof teeth. If Harvey thought he dscried an epigenetie developruent. it was because he sawfirst I little cloud, then the rudunents of the head, with

I Iflrinne Sdtrijlm mi Bride, p. :26.

the eyes bigger than thewhole body, and little by little viscera being formed. If one oompares his description with mine, one will see that his description of the development of the deer corresponds exactly with mine uf the develop- ment of the chick. If more than twenty years ago, before I had made many observations upon eggs and the females of quadrupeds, I employed this reasoning to prove that there is a great difference between the foetus and the perfect animal, and if I said that in the animal at the moment of oonception one does not find the same parts as in the perfect animal, I have realised abundantly since then that all I said against preforrnntion really went to support it.

The reasons for this change of opinion are not clearly given in Haller's

Writings, and Dareste concluded that it would always remain a mystery. This mystery has, however, been almost completely elucidated by F. J. Cole. In 1744 Haller was certainly an epigenesist,‘ in X758 undoubtedly a prefonnationist; in the intervening period he had made his own embryological researches. How was it that they had the unfortunate efiect of carrying him further from the truth rather than towards it? In Cole's words:‘

The yolk, Haller asserts, is the continuation of the intestine of the embryo chick. The inner membrane of the yolk is continuous with the inner mem- brane of the intestine, and is thus identical with the inner membrane of the gut generally and the skin and the ectoderm. The external membrane of the yolk is an extension of the external membrane of the intestine, and is hence continuous with the mesentery and peritoneum. The envelope which covers the yolk during the last ten days of development is the skin of the foetus. Therefore it is no absurdity to say that from the beginning, and before fertilisation, the intestine of the foetus is no more than a small hernia of the membrane of the yolk. Now if the yolk is continuous with the skin and in- testine of the foetus it must be contemporaneous with it, and is truly a part of the foetus. But the yolk was present in the abdomen of the hen, and was a part of the hen, independently of any congress with the cock. Hence the foetus, enclosed in the amnion, must have existed at the same time, though invisible on account of its smallness and transparency.

It is not difficult, with the aid of Fig. 22, to follow Haller‘s argument. The “inner membrane of the yolk" is the endoderm, which it is true does become continuous with the skin and epidermis 11]!!! the gut cavity has been com- pleted. The “cxtemal membrane of the yolk" is the splanchnic mcsoderm, and the "envelope which covers the yolk during the last day: of incubation" is the allanto—chorion, 's\ hich, however, is not the skin of the foetus. I-lallefs procedure is typical of the time, in which observation and inference had only the remotest relations with one another. For example, the statements "Now if the yolk is continuous with the skin and the intestine of the foetus, it must

not:g“on.l:.t'o;t'l!:zasa.vea Pnulcmonzx, vol. 5, pt. :1, pp. 497 fi’. (1744 ed.).

amu1 main...

Fir. 22. Dmgmn of the mmbmm o/ the rhiclt embryo, to tflurtmle I-IalIn'r argument: (from F. 7. Cole).

be contemporaneous with it" and "the yolk must have arteries and veins as without them it could not have been brought into existence" are pure assumptions, and beg the very question he has set out to prove. If these assumption: can be shown to be baseless, as they are, the whole argument collapses.

In a word then, Haller confused the vitelline membrane with the yolk-sac, and assumed a pn'z7n' that foldings, outgrnwths, etc., of cell- layers could not take place, i.e. that epigcnetic processes did not exist. Substantially the same argument as Haller’s had been used some twenty years before (in 1722) by Maitre-Jan, according to Cole.

The zmbollzmznt aspect of prefomution presented no difficulties to I-Ialler. Speaking of the generation of Volvax, he said,

It follows that the otary of an ancestress will contain not only her daughter but also her granddaughter, her greatgr-anddaughter and her gre:Itgfeat- granddaughter, and if it is once proved that an ovary can contain many generations, there is no absurdity in saying that it contains them alL

The following passage is interesting.

We must proceed to say what is the efficient cause of the beautiful machine which we call an animal. First of all let us not attribute it to chance, as Ofrai‘ would have us do, for although he pretends that all animals come from earth, he is not attached to the ancient opinion, and nobody now believes what Aelian says, namely that frogs are born from mud. . . . Vallisneri has found the fathers and mothers of the little worms in galls, a quest of which Redi despaired, and Redi‘ in his turn has made with exactitude and precision those experiments which Bonannus, Triumphet, and Honoratus Faber had only sketched out imperfectly. Moreover, no seed, no clover. . . . This was the received opinion but in our century a proscribed notion has been revivified and some great men have pretended that there are little animals which are engendered by an equivocal generation without father and mother, and that all the viscera and all the parts of these animals do not exist together, but that the nobler parts are formed first by epigenesis and that then the others are formed little by little afterwards.

This is an admirable illustration of how the ideas of spontaneous

generation and of epigenesis were bound up together. Haller goes on to say:

M. Needham does not admit an equivocal generation but he does admit epigenesis, and a corporeal non-intelligent force, which oonstruets a body from a tiny little germ furnishing the necessary matter for it. He says that there are only the primitive germs which were made at the original creation and that germs organised like animals do by no means pre—exist, for if they did, mole: ulninae, encysted tumours, and the like, could not come into being.

Haller then goes on to describe Needham’s experiments with meat broths, etc., and objects to his “system," largely on the ground that “blind forces without any intelligence, could hardly be able to form animals for ends foreseen and ready to take their places in the scheme of beings." He considers that Needham's theories are completely dis- proved by experiments such as those of Spallanzani, though, curiously

enough, he does not quote the latter author in this connection. I shall return to this later 21 t). He continues,

Nobody has upheld epigenesis more than M. Wolff, who has undertaken an examination to demonstrate that plants and animals are formed without a mould out of matter by a certain constant force which he calls “essential" (in his Tlieorfa Gmnalionil). . . . I have indeed seen many of the phenomena which he describes, and it is certain that the heart seems to be formed out of

' is this not Iulien Offrxy de la Mtttrie? Hallcr had A habit of using Christian nazncv.

2.1:. Turbervdle {or J. T. Needham. ' See R. Cole.

n congealed humour and that the whole animal appears to have the same consistency. But it does not follow that because this primitive glue which is to take on the shape of the animal does not appear to possess ix; smmum and all its parts, that it has not eiiectively got them. I haxe often given greater solidity to this jelly by the use merely of spirits of wine and by this means I saw that what had appeared to me to be a homogeneous jelly was carnpmed of fibrm, vessels, and viscera. Now surely nobody will say that the vi: mmlialir of the spirit of wine gave an organic structure to an unfcu-med matter, on the contrary it is rather in the removal of transparency and the accession of greater firmnuu to the extremities, as well as the making of: more obvious boundary to the contour of a viseus that one could see the structure of a cellular tissue. which was ready to he formed but which the transparency had previously hidden and the wetness not allowed to be cir- cumscribed by lines. . . . Finally, to cut a long story short, why does this ti: crrentialir, which is one only, form always and in the same place: the parts of an animal uhich are so ditferent, and always upon the same model, if in- organic matter is susceptible of changes and is capable of taking all sorts of forms? Why should the material coming from a hen always give rise to I chicken, and that from a peacock give rise to a peacock? To these question: no answer is given. This was the case because Wolff was not a theorist, but rather an experi- mentalist; his writings are marked by their abstention from the discus- sion of speculative points. The above passage is very interesting. It reminds us of the great difficulties with which the emhryologists of this epoch had to contend. Serial section cutting was unknown, the staining of thin layers and reconstruction were unheard of; even the hardening of the soft embryonic tissues was only just discovered, as is indicated by Haller above. Hertwig has excellently discussed‘ the advances in embryological technique it hich took place during this and the {allowing century. It is true that dyes were beginning to be used, as some instance already given demonstrate, and as is seen from the use of madder in the staining of bones, which began about this time, and was later much used by the Hunters. Hertodt's Crocolagia is important in this connection. Hertodt, by injecting saffron into the maternal circulation, found it afterwards in the amniotic fluid,‘ and his experiment was cited by Heller in support of that t.heory of the origin of the llq-llld. the most im- portant advance in technique was the progress In amfiaa-' tncuil-‘CM-‘L The art, though lost throughorixt the Middle Ages and the seventeenth century, was now to be revive . _ During this period much nor): was done on it. As far back as 1600, de Serra had mentioned some experiments of this nature, but they were not successful.‘ "The chicks." 11¢ Said

1 And more recently Oppenheimer. ' Quzemo W. P- 273- ' m‘- "- ‘hr ‘-

defective or having too many legs, Wings, or heads, nature being inimitable by art." Birch, in his History of the Royal Society, speaks simi- larly.‘ “Sir Christopher Heydon [a relative of Digby's Sir Iohnij to- gether with Drebell, long sincein the Minories batched several hundred eggs but it had this efiect, that most of the chickens produced that \\ ay \vere lame and defective in some part or other." Antonelli states that similar trials were made at the court of the Grand-Duke Ferdinand II at Florence about 1644, while Poggendorfi and Antinori relate that the Accademia d. Cimento, inspired by Paolo del Buono, made trial of artificial incubation between 1651 and 1667.

But the most famous of all the attempts to make artificial as successful

as natural incubation were those of de Réaumur, whose book De l‘ar1 de faire [clove ler Pouletr of 1749 achieved a wide renown. He devotes many chapters to a detailed description of incubators of very various kinds (see Plate XVIII, facing page 204): but he nowhere gives any indiution of his percentage hatch. It was probably low. He speaks also of the “funestes eflets” of the vapours of the dung on the developing embryos, without, however, furnishing any foundation for an exact teratology. In the second volume he describes those experiments on the preservation of eggs by varnish which caught the imagination of Maupertuis and were held up to an immortal but by no means deserved ridicule by Voltaire in his Akakia. For the details of this amusing but irrelevant issue, see Miall and Lytton Strachey.

After de Réaumur, there were numerous continuations of the work which he had started, in particular by Thévenot, La Boulaye, Nelli, Porta and Cedemhieb-n. Much the most interesting of these was the work of Beguelin, who attempted to incubate Qggs with part of the shell removed so as to form a round window. He was not, however, successful in the carrying out of this very modern idea. Probably the most peculiar investigation made on developing eggs at this time was that of Acbard, who is mentioned in a passage of Bonnet's.

M. de Réxurnur did not suspect in 1749 that some day one would try to substitute the action of the electric fluid for his borrowed heat. This beautiful invention was reserved for M. Achard of the Prussian Academy who excels as an cxperimentalist. He has not so far succeeded in actually hatching a chick by means of so new at process, but he has had one develop up to the eighth day, when an unfortunate accident deranged his electrical apparatus.

Bonnet goes on to say that this substitution of electricity for heat him hope that by electrical means an artificial fertilisation will one day become possible. ‘ Vol. 3. p- 455-

References to these experiments and to those of - ~ ._ gators will be found in Heller. By the heginningmnofythlemxiiillezizecilth Century :1 great mass of literature had developed on the subject, and it

possible to hatch out more or less successfully from , Oughfhe losses were stlll great. Early in the nineteenth century Bonncrnaln and Iouard referred to the large number of mgnsms produced, and in r8o9 Paris wrote,

_ During the period that I was at College, the late Sir Busick Hanrood. the ingenious Professor of Anatomy in the University of Cambridge, frequently attempted to develop; eggs by. the heat of his hotbed, but its only raised monsters, a result which he attributed to the unsteady application of the heat.‘

5. Embryos and Theologians

'I:his is the most convenient place to mention theological embryology again. See pp. 22, 65-6, 75. Its place in the eighteenth century was small, and in the nineteenth, with the general recognition that whatever the soul might be it “as not a phenomenon, it altogether disappeared from serious general discussion. F. E. Cangiamila's Embr;-alagia Sacra, how- ever, ran through several editions between x7oo and I77 5. Cagniamila or Cangi:unila' (Pammn. Ecrl. Can. 172:0]. at in tom Sic-il. Regan tantra hatrttftam pm:-[totem Ingmlitore Pror:irla'aIi) deals very fully with the time of animation, quoting a host of writers such as St Gelasius, St Anselm, Hugh of St Victor and Pier) dclla Mirandola. His mind retains :1 quite mediaeval conformation, as the following curious passage illus- trates: Quo! nonfazlus abortive: ex ignomnlia obrtetritmn rt nmmml exdpit lafrfno, quorum zmima, :1’ Baptirmaie nan fmmiarelur, Damn in nztmuzm ridntt. met demllius lumukmduml His instnletiorts fur the baptism of

I ormmam, p. 366. _ ' Canglamlla, whose strnnge personality has recently been sympztlheualiy reviewed by Hutchinson and hv Boldnm, deserves - lmle biognphienl notice. Born at Palen-no in I702, he became Archprtesl at Glrgenti in 1731 and had the dsunctlon ofbzptinlng the fiat infant delivered by Caesarean section in Sicily. He was lecturer It Palermo In X74: and Vicar-Generil md prm incinl Iuquhitor in 1755. Hu Ernbryologiu Sam was 1 bent-ulltr (see Blblingrlphy) and was even translated into modern Greek for the benefit of those Orthodox who desired to rtudy the lengths to which Reason ma the

Latin mind could go. Cangial-nfla'n main ion-nula was mndlnonal; Sr tn 5: upon‘.

ega te bapuza (even to the amnion). _HE nnxaety to bmpuse the embryo led km: to the

-uél-«vars; 1/2(warmer.C,1As''ez:za‘azrb.oz\thAb.2xo.gnud.tbedud.I\1nmntunes

urlier theology had been more modest, _|.nd the Rarnan Rmmb of us. contented ml: with an edlfyzng De Betltdxrtlanefotttu xn mm malfll’. Nevertheless the eondemnanon at‘ the unbaptlscd infant to em-na_l torment goes back to 1 \er)_ early stage of Lima theology and I! deeply embedded in the teazhlng of Councils. t_unta Ind popes. I! 111! been learrledly shown by Coulton in has Rudy of Infant Perdition. The doetnne does

hllmnrf edi . yméiltiamghuia this mhiect an m gum in Cohen’: boot on the Tdmud (9. m).

B tine‘ th ofstur. , _ _ ygunnmtignfilfdgflflmmdmgme ethic: offoeticrdelmy be hand In Armdr; Glmn: and Hughes. 2°4 PLATE XVIII

Fig. 23. Dalerrpaliux’ dlamngr of Imman !p(nnaI0:oa (from Lemme-nimzk).

monsters are also very odd. But theological embryology probably reached its climax in the report of the doctors of divinity at the Sor- bonne on March 30, 1733, in which intm-uterine baptism by means of a syringe was solemnly recommended. This is included.'in Deventer’s book,‘ and has been referred to by Sterne and Spencer. For other aspects of these tracts of thought, see Nicholle and his anonymous antagonist. But Cangiamila and his t:olleagues—-Gerike, Kaltschmied, ete.—are only of decorative importance to our present theme, and for fuller information regarding them, reference must be made to the treatise of Witkowski.' It is interesting to note that as late as 1913, 182 days was fixed as "perfection-time," whatever that may be, by Moriani.

6. Ovism and Anirnalculism

\Ve must now return to the beginning of the century in order to pick up the thread of the main trend of thought. By 1720 the theory of preformation was thoroughly established, not only on the erroneous grounds put forward by Malpighi and Swamruerdam, but on the experi- ments of Andry, Dalenpatius’ and Gautier, who all asserted that they had seen exceedingly minute forms of men, with arms, heads and legs complete, inside the spermatozoa under the microscope.‘ Gautier went so far as to say that he had seen a microscopic horse in the semen of a

‘ Hlsloirt dc: Aezouehmmlr, pp. 13; E _ ' Cf._ the drscuman an Cleopatra (p. 65). Itis of interest that prenatal ba tism, even including the use of syringes, to this day forms pm at ofiiernl Lucia theo on 1! any nte in the Chunk of France (Ortolan). In literature, we catch tn echo of that am- tmvemcs in the essay of Donne (I531), "That virginity is : Yerruc."

' Qdmy:txm' dnvnngs. r roduecd from Leeuwenhoek in Fig. 23, were nlmost certainly : hoax; sec 1-‘. 1. Co e, PD. 63 t1’.

‘ Hamocknh drawing. illustrated in Fig. :4. represented not what he had um himself but what he supposed xpemulazc: would look like if they could be u.-tn Iu.fiiu‘:nLly clearly.

horse (he gave a plate of it) and a similar animalcule with very large ears in the semen of a donkey; finally, he described minute cocks in the semen of a cock. l-laller remarks gently that he has searched for these phenomena in vain. Vallisneri asserted the same kind of thing about the mammalian ovum, though he admitted that, in spite of long searching, he had never seen one. Besides the main distinction between proforma- tionists and cpigenesists, then, there arose a division among the former group, so that the ovists regarded all embryos as being produced from smaller embryos in the unfertiliscd eggs, while the animalculists re- garded all embryos as being produced from the smaller embryos provided by the male in his spermatozoa. The animalculists thus afforded a singular example of a return to the ancient theory mentioned by Aeschylus int-he 076154 (5¢€}3- 43} Their usual view was that of Hartsoekefl and Andry, who pictured each egg as being arranged like the Cavorite sphere in which H. G. \l/ellsl‘: lorers made their way to the moon, i.e. wit

1: nap-door. The spermatozoa, like sobmany ' e men, all tried to occu an egg, '1‘ 13 tnlilenelggs were far fewer than splzxmtowa. 0166 were, when all was over. only 3 few MPPY animalculcs which had been lucky enough to find empty cggg, dimb in and lock the door behind thegiher followers of!:x,t‘t;ni1l0¢k'f§Sb5€:'llx¢d 11141‘ there were apermatie an‘ cu es 0 o setcf.

4 as one oauld see by a slight difference near their

firlgie-121:; 0!/Inl:";,Iilkr::1: tails, that they copulated, that the females beanie =1m"'"""’°"- rcgnant and gave birth to little ammaleu_l5. that young and feeble ones could be seen,‘ that 31,61)’ j1¢dH':1£“' ‘hi: and finally that some had been Observed With '_W’-if '3 5; _ :‘;;z‘1:°_ made good use, on thkes \vholc1:‘,! of lusc:::x;g,y=m D 5°°P"°‘5m' terised all these remar 35 "0 Y °°“.l ' . -

The whole controversy was intimately bound up the hi: spontaneous generation. 501’. “'1m3V°’ am “*5” ":3 . cdemmm’ mg! of animals, if it were true that the lower ones coulh fflim at least must slime, mud or meat infusionnfor Instance, then f :7 P for it mu“ have been made by epizmws 3”’ *3" "3 “Y °' " ‘:tar)iicture oftlut hardly be held that a homogeneous .lnfl.1Sl0‘l"l had afiguls ‘hm thflhin k'md_ And if gpigenesis could occur in the lower am 1 I See p. :75

end of the wedge had been driven in, and it might occur among the higher ones as well. It was in this “ray that the spontaneous generation controversy came to have a peculiar importance for embryology in the eighteenth century.

Driesch‘ has essayed to make the generalisation that all the supporters of epigenesis were vitalist in their tendencies, while those who adhered to the preformation theory were not. But there are too many exceptions to this rule to make it helpful. In so far as it applies, the association doubtless arose from the fact that the continual production in epigenesis of new organs and new relationships between organs already formed seemed to require an immanent formative force of some kind, such as the vi: esrentialir of Wolff, while on the preformation hypothesis, where ernbryogeny was little more than a swelling up of parts already there, it could be explained simply as nutrition. But the failure of the “short- cut" mechanistic philosophers such as Gassendi and Descartes led to preformationism just as much as to epigencsis. A remark of Cheyne's throws some light on this question, for in x71 5 he wrote, rallfing to Gassendi's line of thought for different reasons, “If animals and vege- tables cannot be produced from matter and motion (and I have clearly proved that they cannot), they must of necessity have existed from all eternity."’ Preformationism thus became the only resource if the uni- versal validity of the mechanical theory of the world was to be retained. Stahl, and later Wolff, saw no point in retaining it, and carefully joined together what Descartes had with equal care put asunder.

Von Haller affords some interesting evidence against the identification of epigenesis with vitalism and preformation with mechanism, for he says, “Various authors have taught that the parts of the human body are formed by a mechanism depending on general laws" (i.e. laws not simply of biological validity), "or by the virtue of some ferment, or by heat and cold making crusts out of the different juices, or in other ways. All these [mechanical] systems have some resemblance to that of M. Wolff.” Haller also always speaks of Wolff's vi: erxznlialir as "blind."

The original discoveries of de Grant’ and Stensen were extended by Tauvry in 1690 to the tortoise, and by Lorenzini' in 1678 to the Tar- pedo: so that the eighteenth century began with an excellent basis for ovistic prefarmationism. The greatest names associated with this school were Suarnmerdam, Malpighi, Bonnet, Von Haller, VVinslow, Vallis-

neri,‘ Ruysch and Spallanzani.4 But there were many others, some of whom did valuable work, such as Bianchi,l Bourguet, Bussiere,

I And Ililfltiewiczz see pp. 2:4 and H9 heruftzr. 'Pfn7oxaplu':al I’h‘m-r'p!zr_ ch. a, sect. _m E. p. 6!. ' See p. 167. ‘ Ste Franchiru. ‘ D: Nat. Gem, pp. 417 H’.

C°5°h“”‘zr Fm‘: P¢“'3\1l‘. 5i€m’-y Teichmeyer, Veroelloni, Wdussi and Weygand: The treatises of Imbert and Plonquct were written from :51; point ofvrew, as was the bright little dialogue ofde Houpeville. J. B du Halftttl asserted that he could see the chick embryo in the egg beioxe gegglisation, and Jacobaeus made a like affirmation in the case of the

On the other side, that of animalculistic prefor-mationism, the con- testants were fewer. Their greatest names were Leeuwenhoek, Han. 5°€k€1’. Leibniz and the cardinal dc Polignac.’ In England the physi- cians Keil and Cheyne supported this position, in France Geofroi and the obstetrician la Motte, in Germany Withof and Ludwig, and in Belgium Lieutaud. De Supenille wrote in favour of it in the Philom- phieal Transaction: of the Royal Society, and an anonymous Swedish work of some fame supported it. To the argument of Vallisneri that the existence of so many aniznaleules must be an illusion, since Nature could hardly be so prodigal, the animalculists retortetl by insraneing such observations as that of Baster, who had taken the trouble to count the eggs of a crab and had found that they amounted to 12,444. James Cooke later elaborated a theory! of :1 “orld of the unborn to which the spermatozoa could retire between each attempt to find a uterus in which they could develop-—this avoided V:rllisneri’s argument.

All those other attending Anirnalcula, except that single one that is then conceived, evaporate army, and return back into the Atmosphere again, whence it is very likely they immediately proceeded; into the open Air, I say, the common Receptacle of all such disengaged minute sublunary bodies; and do there circulate about with other Sernina, where, perhaps, they do not absolutely die, but live a latent life, in an insensible or dormant state, like, Swallows in Winter, lying quite still like a stopped Watch when let down, till they are received afresh into some other Male body of the proper kind, to

‘ Ch. 3, p. 8. _ ‘Anti-Luneriux. net. 8.

‘ Support: by flrolluton. Actually : urmlar argument had been E\'Dl\:d some seventy yenn previously by F. M. van Hzln-mot, who was worrrine. not, like Cooke. about the losr spermatozoa. but about the ovulared egg: which (nled to get fernhsrd. Xn his strong: discourse on mttunpsychosin, he toy: (p. 15;): “Quation no. 33. More- over. when we find that Children m the womb be formed out of Eggs. of wh-ch there are so great . number in every woman, am we do not find one am bur: w many Children as Ill! hath Eggs, which oh: brought into the V\'orld with her: l\lu1t we not therefore conclude, that the rest of these Eggs were created in win. in use they should not it some time or other main to their full perfection? Now to retrieve this dt-fficulfv. musrwe not conclude. that the Life of these Egg: doth prvplgate xtsgli,-nothzr wry. to the rnd that what doth not arrive or pexfecuon one tune. rmry xturn it or mother] And um mmtm the remaining Egg: must necessarily be revolved In order to am: perfecuon. II which in the production of than. Nature had directed her xntentrdorzi

In like mumer what an we euppose to be the rmon of that gxprtss commln , That no basnrd should enter into the ccn8f98'|"°“

God. which we read, Dent. 23, 2, by means of ten llemlunons. the

of the Lord to the math generation; but this. that evil might be wrought our?

be again set on Motion, and ejected again in Coition as before, to run a fresh chance for a lucky Conception; for it is very hard_to conceive that Nature is so idly luxurious of Seeds thus only to destroy them, and to make Myriad: of them subservient to but a single one.

But Cooke's attractive hypothesis, published in 1762, came too late, as Punnett says, to save the animalculists.

The idea that human seed, or spermatozoa, floated everywhere in the air (obviously derived by Cooke from the Stoic—Talrnudic-Kabbalistic line of thought described on pp. 66, 79) led to the amusing satire of Sir John Hill in x75o mockingly addressed to the Royal Society, Ludrm sine Concubilu. He affected to have invented a machine for trapping the seminal animalcules home on the ‘Nest wind.

Accordingly after much Exercise of my Invention, I contrived a wonderful cylindrical, caloptrical, rotunda-concavo—convex Machine (whereof a very exact Print will speedily he published for the Satisfaction of the Curious, designed by Mr H-y-n, and engraved by Mr V-rtu), which, being hermeti- cally sealed at one End, and electrified according to the nicest Lavis of Elec- tricity, I erected in a convenient Attitude to the West, as a kind of Trap to intercept the floating Animalculae in that prolific quarter of the Heavens. The Event answered my Expectation; and when I had caught a suflicient number of these small original unexpanded Minims of Existence, I spread them out carefully like Silk~worm's Eggs upon White-paper, and then apply- ing my best Microscope, plainly discerned them to be little Men and Women, exact in all their Lineaments and Limbs, and ready to ofier themselves little Candidates for Life, whenever they should happen to be imbibed with Air or Nutrimcnt, and conveyed down into the Vessels of Generation.‘

On the experimental side, Garden came forward with descriptions of little men inside the animzilcules, thus “confirming" the work of Gautier and Hartsoeker. It is fair to add, however, that Garden held quite enlightened views of the mutual necessity of egg and sperrnatozoon. So did Massuet, whose dissertation appeared in 1729 at Leiden. An animalculist, he yet believed both egg and sperm to be needed in generation, the fomier rather as a nidus. He reversed the Malpighian view. Si ovum gallium, he said, non foenmdatum minorrnpio inspexrfis, mzlla in ca animalirfanna apparebit. He gave the correct explanation of Nuck’s experiment (see p. 163), saying that :emx'm'.r aum seu animaltula had first passed up the Fallopian tube. In his plate he figured spemta- tozoa, chick embryos and tadpoles all together, confusing the former with the primitive streak.

Another adherent of enlightened common sense “as Hugh Cham- berlen, one of the famous obstetrical family. In his English translation (1683) of Maux_iceau‘s midwifery (was) he took exception to th. doubIe—seed (Epicurean) opinions of the French writer, and addcd the; following as a footnote:

Our author lying under a Mistake, in his notions concerning the Testida in this chapter, I shall here give my sentiments. We find that the Tcsticles of a Wornanare no more than, as it were, two Clusters of Eggs, which [ie there to be impregnated by the Spirizuous Particles, or Animating Emu. VN1115. Oonveyed out of the Womb through the two Tubes, called by our Author, Defcrent Vessels. . . . Some days after the impregnation of the Egg or Eggs, as in Twins, they decid through those two Tubes into the Womb: where being placed, the Embrio takes up its quarters.

But la l\Iotte maintained that the egg (which he identified with the Graafian follicle) was too big to go down the Fallopian tube, launxrrll‘ Sbaragli, another writer on the animalculist side, agreed with As for the supporters of cpigenesis, they were few, but they included Desmrtes, de Maupertuis, Antoine Maitre-Jan and John Turberville Needham. 1‘/Iinor Writers on the same side were Tauvry, lVelsch, Dartiguelongue, Béttger, Drelinourtius and Mazin. After r75o C. F, Wolff brought an abiding victory to their opinion.

Among the arguments brought forward against the preformationists ‘VETS 3 (I) That it is impossible to explain the production of monsters on a preformation theory. Brunner first brought this point forward in 1683. but its classical statement was that of Etienne Geolfroy de St Hilaire, in his work of 1826 on experimental teratology. On its history, see Strohl and Dareste.

(2) That preformation is incompatible with the facts of regeneration. An intelligence, argued Hartsoelser in 1722, that am reproduce the lost claw of a crayfish, can reproduce the entire animal. This point much impressed Erasmus Darwin.‘ The whole subject was much to the fore in the eighteenth century owing to the brilliant observations of de Réaumur and Trembley!

(3) That the extraordinary resemblance between small embryos of mammals, birds, reptiles, etc., discredits preformation. This was the View of Prévost 8: Dumas, 1834-1838.

Some maintained a quite independent position, Bufion, who welded together an epigenetic theory of fertilisation with a prtfnrmr tionist theory of embryogeny. Pascal (not the great Jansemst) W‘ ‘W’ 'Zoonoam'n, :, 45:137. ' See Merward dte chemical view that fertilisation consisted in a combination between the acid semen of the male and the “lixivious” semen of the female, perhaps because in chemistry acids had been regarded as male and alkalis female.‘ Claude Perrault‘ and Connor also suggested that the formation of the embryo was a fermentation set up in the egg by the sperxnatic animalcule. In this they were following the example of van Helmont, who had originally suggested such a theory.’ In 1763 Jacobi discovered how to fertilise fish eggs with milt; a practical matter which had a good deal of influence on biological theory. De Launay alone still held to the Aristotelian conception of form and matter.

7. Spontaneous Generation

There is no need here to do more than glance at the spontaneous generation controversy itself, for it has long been well known in the history of biology, especially in connection with the subsequent work of Pasteur. J. T. Needham’s books, New I|rIicra.tco1>1‘:ul Dircoverie: of 1745 and Obrmvnliant upon the generation, composition, and decomposition of Animal and Vegetable Subxtmices of 1749. exercised a considerable influence. They were written concisely after the French fashion (Need- ham had been educated at Douai), and with some brilliance of style, and it is hardly true to say, as Rad] does, that their experimental foundation was meagre. That it was inadequate was proved defi- nitively as events tumed out by Spallanzani. De Kruif's picture of the controversy is false and misleading, especially in its estimate of Needham, “ho is much more truly described in the words of Louis Pasteur.‘

Need.ham’s case rested upon the statement that if meat broth was placed in a sealed vessel and heated to a high temperature so that all life in it was destroyed, it would yet be found to be swarming some days later with microscopical animals. All depended, therefore, upon the sureness with which the vessel had been scaled and the efficacy of the heat employed to kill all the animalcules initially present; and in the

ensuing controversy Needham lost to Spallanzani entirely on the question ‘of technique. It may be remarked here without irrelevance that the problem is still in a sense unsolved, for what the experiments of Sp:illan- zani proved was that animals the size of rotifers and Protozoa do not

"2153. Gregory‘; interuting essay on the minute and mechanical model: of 193'» s

' One:-m diverm de Pluyriwe t1|1!cnru'qu4(i7z ), nl. 2, . 439.

' Omu, l, :1. How intercatrdtthcse men vvouldIha‘\"l: bee: in modern researches on the changes at fertilisation of respiration, glyeolytie activity, and enzyme action in the prmoplum of the egg cell.

For an accurate and detailed account at’ the controversy, see Prescott.

originate s ontaneousl

pasta“. “.3: that orgmyivs :03’ Ptl'ov_cd. by lgose of in that way. The knowledge which has been at uirrtili yngmme v 2 now "filter-passing organisms," such as the niosaii V"ll'|1S"i.)ir'e1ci::"t}§a:s of Pl-'15‘. and phenomena such as the bacteriophage of Twort and d’iilei:ll: has re—opened the whole matter, so that of the region between for example, the semi-living particles of the bacterioph:lge(1°—i: Em)’ and the larger-sized colloidal aggregates (mm gmm) We know ,cma,k3M little. The possibility of the new formation of viruses without s ancestry, in the cells of living hosts, is to-day an open qucsting aid we still have no proof that their origin from non-living Organic ',m,c,;31 can never occur. Recently it has proved possible to "synthesise” infec- tive viruses from separate protein and nucleic acid oonstituents of other viruses, constituents some of which alone are incapable of reproduction.‘ The dictum onm: z-ioum 2: viva, accepted with such assurance by the biologists of the early twentieth century, may thus tum out not go mum quite what they thought.

But to dwell further on this would be a digression. The important point was that Spallnnzani's victory was a victory not only {or those uho dishelievcd in spontaneous generation, but also for those who believed in the preformation theory of embryogenyf By 1786, indeed, that view- point was so orthodox that Scnchier, in his introduction to an edition of Spallanzani’s hook on the generation of animals and plants, could treat the epigenesists as no better than atheists.

Spall-anzani's views on embryology were largely drawn from his study of the development of the frog’s egg (see Fig. 19). Here he went far beyond Bose, but in spite of many careful observations he thought he saw the embryo already present in the unfertilised ovum. This led him to claim that Amphibia ought tobe numbered amongviviparous animals. His principal step forward was his recognition of the semen as the actual agent in fertilisation on precise experimental grounds—the narrative of his artificial insemination of a bitch is too famous to quote: he said it gave him more intellectual satisfaction than any other experiment he had ever done. This demonstration finally dbposed of the aura .mm'naIr'.t wlvich Harvey had. found himself ohliged to adopt on the grounds of his dissections of does. But Spallanznni failed to convince himself that the

spermatozoa themselves were the active agents.

‘ Fl-aenlrcl-Canrat has given : good description of this work. There now seems no

mson why it should not be possible In the {ensemble future to nynqmiie . nucleic hid: when introduced into n host cell

acid from other organic izhemiaal ntlnmnes w _ _ would show infecuvity, i.:. n self-rephaxtion or reproduction within the metabolic

' - This was realised at the nine rspecially by Patrin, who am to Needhanfa defence

in 1778 (see Btlikiewiez, p. I47). See on, p. 218.

8. Preformation and Epigenesis

Of all the preformationists Charles Bonnet was the most theoretical.‘ He was an adherent of that way of thinking mainly on the theoretical ground that the organs of the body were linked together in so intimate a manner that it was not possible to suppose that there could ever be a moment when one or two of them were absent from the ranks.

One needs [he said] no Morgagni, no Hallcr, no Albinus to see that all the constituent parts of the body are so directly, so variously, so manifoldly, intertwined as regards their functions, that their relationship is so tight and so indivisible, that they must have originated all together at one and the same time. The artery implies the vein, their operation implies the nerves, which in their turn imply the brain and that by consequence the heart, and every single condition a whole row of other conditions.

Bonnet compared epigenesis to crystal-growth, in which particles are added to the original mass independently of the plan or scheme of the whole, i.e. in contrast with the growth of an organism, in which particles are added on only at certain places and certain times under the guidance of "forces (It: rapport.” Przibram has recently discussed the question of how far such at comparison is admissible, but in Bonnet’s time at any rate it became very famous. Bonnet referred to von Haller’s dis- covery of the intimate relationship between embryo and yolk as evidence for his theory. The embryo begins, according to him, as an exceedingly fine net on the surface of the yolk; fertilisation makes part of it beat and this becomes the heart, which, sending blood into all the vessels, ex- pands the net. The net or web catches the food particles in its pores, and Bonnet supposed that if it were possible to abstract allthe food particles at one operation from the adult animal, it would shrivel and shrink up into the original invisible web from which it originated. Bonnet was no more afraid of the emboflement principle than was Haller; indeed, he called it “one of the greatest triumphs of rational over

‘ \Vhite'| 77!: Phlagittm Thzovy will interest those who Willi to study the rather striking panllel between chemistry and biology in the eighteenth century. Broadly speaking. rationalism in science had too much got the upper hand of zrnnincism. The trace: of preforrnntionism remaining in modern biology are well reviewed by Huxley 8: de Beer. \Vhitrnan distinguished between "predeterrninntion,” I physiological or potential prcfnmistion not capable of rniuoscapic resolution, and "predelmeation," which I! the old morphological or more preformation. Modern embryology might therefore be called "predzterm' ed epigtnesis." “Inddington has recently (.952) intro- duced the term “e~pigeneiies" t nelnde everything etmctrned \\llh the causal analysis of development, that is to say, with the genes Ind thtir effects in embryonic life as well as the merphogenetie rnedunisms themselves.

' Bonnet‘: nu.-tiul contributions to science were not numerous but rather impor- tani. ll: Congflfled in 1745 (r779, 13. 36) Leeuwenhoek‘: dI5C<7VE1'y of the pnnhrno- gum: or aphids (1702), and he announced the formation or new individuals alter the cutting of worms into regrntrits (nee the paper of Erhard). The value of the former

observation I! 1 support for ovism is evident. For further dcuil: concerning him.Ie: Lemaitre and Whitman.

sensual conviction." Many of his argum - Haller's, and he says in his preface that heehziil i:eri:etehEhisml2::li°snosuif time before Haller's papers on the chick appeared but then finding his . . . .

own views confirmed by the expenmentally better founded ones of Haller, he determined to publish what he had set down. Thus in one place he says,

I shall be told, no doubt, that the observations on the development of the chick in the €88. and the doe in the maternal uterus, make it appear that the parts of an organised body are formed one after another. In the chiclr for instance it has been observed that during the early pan of incubation the heart seems to be outside the animal and has a very different form to nhat it will have. But the {eehleness of this objection is easy to apprehend. Some people wish to judge of the time when the parts of an organised body begin to must by the time when they become visible to us. They do not reflect that minuteness and transparency alone can make these Pang inyi5,'],1¢ gg ‘,3

although they really exist all the time.

Bonnet was therefore what might be mlled an "organic preforma- tionist," for his objection to epigenesis lay in the {act that it did not seem to allow for the integration of the organism as a whole. His mistake was that he assumed the capacities of the adult organism to be present all through foetal life, whereas the truth is that they grow and dilferentiate in exactly the same way as the physical structure itself does. Bonnet's philosophical position, which has been analysed by \Vl1itm:1n, again contradicts the generalisation of Dr-iesch' that all the epigenesists were vitalists and all the preformationists mechanists. For Bonnet an epigcnetic and a mechanical theory wen: one and the same; he hardly distinguished, as Rédl says, between Descartes and Harvey; and it wasjust the neodritalist idea of the organism as A whole that he oould not fit in with epigenesis. Needham and \Voll1' were un- doubtedly epigenesist-vitalists, and Bonnet was undoubtedly a pre- fonmtionist-vitalist, but Maupertuis was equally clearly an epigensisb mechanist.

G. L. Leclerc, Comte de Bufion, the most independent figure in the controversy, stood alone as much bemuse of his erroneous experiments as because of his originality of mind. As has so often been observed. Bufion was not really an expcrimentalist at all; he was a writer, and preferred other people to do his experiments for him. The vol'ume.0n generation in his Hirtoin Natxmlle begins with a very long historical account of the work which had been done in the previous centuries on embryology. At the beginning of the section on reproduction in general he said,

The first and most simple manner of reproduction is to assemble in one body an infinite number of similar organic bodies, and to compose the sub- stance in such a manner that every part shall contain a germ or embryo of the same species, and which might become a whole of the same ltind with that of which it constitutes a part.

Such an idea resembles the ancient atomistic speculations, and was explained by VV. Smellie, the obstetrician, who translated Bufion i.nto

English, as follows:

The intelligent reader will perceive that this sentence, though not very obvious, contains the principle upon which the whole theory of generation adopted by the author is founded. It means no more than that the bodies of animals and of vegetables are composed of an infinite number of organic particles, perfectly similar, both in figure and substance, to the whole animal or plant of which they are the constituent pans.

This conception explains l3ufl'on's curious attitude to the preformation question. An embryo was preformed in its germ because all the parts of the germ were each a model of the animal as a whole, but it was also formed by epigcnesis because, the sexual organs being first formed, all the rest arose entirely by a succession of new origins. Bufi'on’s “organic living particles” bear some resemblance to the “biogen molecules" which later generations were to discuss,‘ and he says that an exactly similar but simpler stnrcture is present in dead matter.

In his discussion of former theories he resolutely rejects the ¢rnI7oi!e- merit aspect of preformationism, giving various calculations to show its impossibility and maintaining that every hypothesis which admits an infinite progression ought to be rejected not only as false but as destitute of every vestige of probability. As both the vermicular and ovular systems suppose such a progression, they should be excluded for ever from philosophy.

He completely destroys the theory which the ovists and animalculists had set up in order to explain resemblance to parents, namely that although the foetus might originate either from egg or spermatic animal- cule originally, it was moulded into the form of its parents by the in- flucnce of the rnatemal organism during pregnancy. This field,’ which was more than once disturbed by the contestants during the course of the century, received systematic attention from time to time by medical writers. There was a memorable dispute on the point between Turner and Blondel, whose polemics, written in an exceedingly witty manner, are still very pleasant and amusing to read. Blondel was the soeptic and

And even more to the Item: of Annsgoru and the Stoic-Knbbslistic "ends." ‘ See p. :9.

Fig. 25. G. L. L. J: Bujon and hi: frialdr xtudying marnnmlian generation. 71m mm an m n 1-ignem in Buflovf: "Him-nre Nanu=l.le" at 11:: head qf

MR7! Lwiv Ddvbwtvl (1715-I799) loglu dawn the miaaxtope. 77:: mrgm in atttnd. one: may be miner Gutnnzu 4. Jormz-emu (1720-1785) or 2‘. F. Dubbard (I703-1779), mo mm eolhbarman af B_uflon. /in um uwlurr mu epzgemm and anmuzl. mhm, and mug in mm mumk: Whit]! ha: nnw bun Md, they mm am-inmx that M luzdfmmd mmmnnzaa in the Gme/Ea: fulluks of xhef.-male ovary. The um: xhomx urnnrining the mmmmluza generative organ: (me pp. 168 am! an: of the "Hisroire Naturelle," Va!‘ 11, r75a). The ingmxhur xdznofizarwn of the pawn is due to Pmfeuar R. C. Pu-rmenr on Guéneau dz Alonlblliard, su Bnmel and Mmquae

Turner the defender of the numerous extraordinary storiu which passed for evidence on this subject. It is interesting to note that Turner believed in the continuity of foetal and maternal blood-vessels. Krause and Ens later supported the opinions of Turner, while Okcs, in 3 Cam- bridge disputalion, argued against them.‘

BuEon's sixth chapter. in which he relates the progress of his on experiments, is unfortunate, in that his main result “as to discover spermatozoa in the liquor fulliculi of ovaries of female animals (see Fig. 25). The explanation of how he came to make such an egregious mistake has never been satisfacton'Iy given, and it was not long before the truth of the observation was questioned by Lederrnuller} It led him naturally to the assertion that the ovaries of mammalia were not cgg~ producing organs but animalcule—producing orgam, and to the Via!’

that the beginning of embryonic development lay in the fusion of the male with the female spermatic anirnalcules——a curious revival of Epi- cureanism.‘ But it is to be observed that he does not mean one male animalcule with one female animalmle, but rather all with all, in a kind of pangenesis.

All the organic particles which were detached from the head of the animal will arrange themselves in a similar order in the head of the foetus. Those which proceeded from the backbone will dispose themselves in an order corresponding to the structure and position of the vertebrae.

And so on for all the organs. The fact that for the organs common to both sexes a double set of nnimalcules will thus be provided does not give Button any difiiculty and is fully admitted by him. Accordingly he could only agree to the aphorism «mm: vivum ex aw in the sense of Harvey namely as referring to the egg-shaped chorion of Vivipara, and definitely not in the sense of von Baer, namely in the modern sense.

Eggs, instead of being common to all females, are only instruments employed by Nature for supplying the place of uteri in those animals which are de- prived of this organ. Instead of being active and essential to the first im- pregnation, eggs are only passive and accidental parts destined for the nourishment of the foetus already formed in a particular part of this matrix by the mixture of the male and female semen.

Biology at this period was still labouring in the dark without the illumination of the cell-theory, and therefore unable to distinguish between an egg and an egg-cell.‘

In spite of his leanings towards epigenesis, Bufion repeated precisely the error of Malpighi.

I formerly detected [he says] the errors of those who maintained that the heart or the blood were firs: formed. The whole is formed at the same time. We learn from actual observation that the chicken exists in the egg before incubation. The head, the backbone, and even the appendages which form the placenta are all distinguishable. I have opened a great number of eggs both before and after incubation and I am convinced from the evidence of my own eyes that the whole chicken exist: in the middle of the cimtrix the moment the egg issues from the body of the hen. The heat communicated

‘ it is interesting that the rejection of the Epicurean theory of female seed by the Latin theologians led, in 2ighteenth—cenl'ury moral theology, to 3 very unequal em- phasis on mumrbuion u I am in main and females. The male (cf. the influence of the nnimnlcuhsts) was to be regarded as little short of I murderer if tfluxio Irwin’! occurred; the female could not thus -in quiz: rennn semenin mxlmfbur nan datunuee CI‘pellmn.nn. pp. 88 R‘.

In 1778 \V. Cruikshank found blasxoeystl in Ll’): Fallopian tubes of the nbbit on lhe third day utter coitus, but his -neunte ohm-vuim-is were not publnhed till 1797.

2:5‘ 5): inculgltian czrpaods the pam only. But we have never been able to °"“"" ‘“ “"-W“? what parts of the foetus are Erst fixed, at the moment of its formation.

The expenmcnt oftaking a look at the cimtrioes of eggs on their way down the parental oviduct seems so obvious that Bulfon may well have thought of it, and it would he really interesting to know what factor in the intellectual climate made him regard such an observation as not worth attempting. His obsenations on the embryo itself were good and in some “my: new; thus he noticed that the blood firs: appears on the "plaoenta” or blastoderm, and for the first few days seems hardly to enter the body of the embryo. He gave an extremely good account of the whole developmental process in the chick and in man, and his opinions on the use of the amniotic liquid and the functions of the umbilical oord were advanced.

J. T. Ncedham, however, spoke very clearly in favour of epigenesis, though he himself did no emhryological experiments.‘ His Idéerommairz of 1776, written against Voltaire. who had called him a Jesuit and who had drawn materialistic inferences from his writings, contain the following passage:

The numerous absurdities which exist in the opinion of pre—existent germs, together with the impossibility of explaining on that ground the birth of monsters and hybrids, made me anbrace the ancient system of epigenesis, which is that of Aristotle, Hippocratns, and all the ancient philosophers, as well as of Bacon and a great number of savants among the neoteriqucs. My

obsen-ations also led me directly to the same result.

Needham’s embryology is mostly contained in his Obrmwtimu nou- celler run In Griréralinn of 1750. He was explicitly a Leibnizian and

postulated a vegetative force in every rnonad.

Needham was not the only thomughgoing epigenesist of this period. Maupertuis, whose Vina: P}1_)'s1'que was published anonymously in 1746, mme out very clearly in {armor of this doctrine.‘ He wrote:

I know too well the faults of all the systems which I have been desc-n'bing to adopt any one of them, and I find too much obscurity in the whole matter to wish to form one of my own. I have but a few vague thought: which I propose rather as thoughts to be examined than as opinions to received, and I shall neither be surprised nor think myself aggrieved if they are

I Hi: valuahle biologial discoveries have beat samewim mnhadawed by th« lpontaneaus geuention controversy. In fact he made ldrnirabée mnueigptrai-slang}: cu-nplzmted physiology, sexual and gains], ofcephalapods -n -.-1].;-11> 5;, mum in pollen grains is maloguu oiepenmtozoa and was the lingo nee mvm

them, and he described the horned eggs of damiobraneh bu.

rejected. It seems to me that the system of eggs and that of spermatic animal- cules are both incompatible with the manner in which Harvey actually saw the embryo to be formed. And one or the other of these systems seems to me still more surely destroyed by the resemblance of the child, now to the father and now to the mother, and by hybrid animals which are born from two different species. . . . In this obscurity in which we find ourselves on the manner in which the {oetus is formed from the mixture of two liquors. We find certain facts which are perhaps a better analogy than what happens in the brain. When one mixes silver and spirits of nitre with mercury and water. the particles of these substances come together themselves to form a vegeta« tion so like a tree that it has been impossible to refuse it the name.

This was the Arbor Dianne, which played a great part in the embryo- logical controversies of the eighteenth century. It has much interest for us, for it was perhaps the first occasion on which a non-living phenomenon had been appealed to as an illustration of what went on in the living body. It is true that Descartes long before had said that the movements of the living body‘ were carried out by mechanisms like clocks or watches, and that they resembled the statues in certain gardens which could be made to perform unexpected functions by the pressure of a rnanipulator’s foot on a pedal, but these instances were all artificially constructed mechanical devices, whereas the Arbor Dianae was a natural phenomenon quite unexplained by the chemists of the time, and the lineal forerunner of Lillie’s artificial nerve, and Rhumbler’s drop of chloroform. VVe know now that its formation is a simpler process than anything which occurs in the developing embryo, but the growth of knowledge has made it undeniably clear‘ that the same forces which operate in the formation of the Arbor Dianne are at work also in the developing embryo. To this extent Mztupertuis is abundantly justified, and Driesch’s comments on him are not in agreement with the facts. Maupertuis continues:

Douhtless many other productions of a like kind will he found if they are looked for, or perhaps if they are looked for less. And although they seem to be less organised than the body of most animals, may they not depend on the same mechanics and on similar laws? Will the ordinary laws of motion sufiice, or must we have recourse to new forces? These forces, incomprehen- sible as they are, appear to have penetrated even into the Academy of Science:

at Paris, that institution where so many opinions are weighed and so few admitted.

Maupertuis goes on to speak of the contemporary deliberations on the subject of attraction.

See (2.; zxlmyle Rinne or Puibnrm

Chymistiy has felt the necessity of adopting this conception and attraaire force is nowadays admitted by the most famous chyrnists who have carried the use of it far beyond the point which the astronomers had reached. lfthis force exists in nature, why should it not take part in the formation ofaniimlsi

Maupertuis was thus an epigenesist and a mechanist at the same time. His opinions have an extremely modern ring, and his only retrograde step was in suggesting that the spermatic animals had nothing else to do except to mix the two seeds by swimming about in them. But that legacy of ovism was common all through the eighteenth century, and thirty years later Alexander Hamilton could still say, “From the discovery of Anim.-ilcula in ::mine man-i:li'rio by Leeunenhoclfls Glasses, a new Theory was adopted which is not yet entirely exploded.”

But the real middle point and fulcrum of the whole period lay in the controversy between von Haller and Caspar Friedrich Wolff, the former at Gottingen and the latter at St Petersburg in the academy of Queen Catherine. Kirchhoff has described this polemic. \Volfi‘s Theorm Generntioni}, uhich was :1 defence ofepigenesis on theoretiml and philo- sophical grounds, uritten in a very forrml. l9g‘°-‘ll and ‘"‘_’“‘d“bl° manner, appeared when he was only tW¢Df}"5'-‘ 3"-'3" Pld» "1 *759- Leilmiz, as Ridl points out, had borrowed from the earher preforrna- fionisrs the conception of a unit increasing in bulk in Order *0 590911“ another kind of unit, but W013. f01_l°W!_"8 N¢‘dh“m3 b°"°‘"d rm"; Leibniz the idea of a inonad destlopmg into an urg=1“‘5‘.“ 5}’ 335"“; its own inherent force, and to this he joined the Stahlialn 31011:; ililra supra-physical generative force in nature. On the Pl’-‘Cm’ V5‘ 5- °_ I 3 work was indeed of the highest lmP°mn°°‘ If the embfio mtvs 5' he argued, if all the organs are actually_prcsent 3! ‘I13 V") ¢3'_1°5‘ 5'38!‘ and only invisible to us even with the highest power; of our m}icr0S0l:P€I3]: then we ought to see them fully formeitl as soon as we see I em a ha“; in other words, at the moment a1 “'l“‘h ‘"7 gm“ om: Ecmcjf the view, it ought to have the form and shape. though “_°‘l; 35”“: um“ same organ when fully completed in the embryo at hirt n ‘hm cm band, if this is not the way In which du'c1cpm=n‘hn8°°‘-‘S; . into ought to be able to see uith the microscope anti 5 Pceh onegggemm another shape. and in fact a series of ap_pe:u'.n_iCcS» :1 words 2 Suit: {mm um which imimdmely pmcfides 1‘, or utitlile “riinitive emhr)“ of advancin§hfi“l;'pt:lllJ::sa:fllil-gtfivl'::lT;lp;|rst: he bloliid-vessels of (ht oriic rn:isS- 0 C - . bkgstodgfin in the chick. for he saw that at one _XflC|:1l'zi1‘:nx:()lth;JSe€clI]:-fl1}l’I?lls ‘"5 “'35 in "~‘55“’"°°’ Wm‘ die mama“ baht’: "that the homogene- mgcmsggpicaheseaxches led turn to the conclusion

ous surface of the blastoderm partially liquefies and transforms itself at these points into a mass of islands of solid matter, separated by empty spaces filled with a colourless liquid but afterwards with a red liquid, the blood. Finally, these spaces are covered with membranes and become vessels. Consequently it was demonstrable that the vessels had not been previously formed, but had arisen by epigenesis.

Holler replied to this new experimental foundation of morphogenesis without delay, for he was working on the development of the chick at the same time, and held closely to the opposite theory. \Ve have already seen what his one and only argument against \Volff was. He used it time after time in all its possible variations, maintaining stoutly that the chick embryo was so fluid in the early stages that \Volif had no right to deny the presence of a given structure simply because he could not see it. Haller’s explanation of Wolff’s results was that the blood-vessels had been there all the time but that they had not become visible until the moment at which Wolff saw the islands forming.

After I had written the above [said Hallei-], M. Wolff made new objections against the demonstration. Instructed by new researches, he denies absolutely that the yolk-membranes, which he makes two in number, exist before incubation. He pretends that they are new and that they are born at the be- ginning of incubation, and consequently that the continuity of their vessels with the embryo does not in the least prove that in the body of the mother the yolk received vessels from the foetus. I have compared the observations of this great man with my own and I have found that the yolk never has more than one pulpy and soft membrane, part of which is called the umbilical area, and that the fine exterior membrane does not belong to the yolk but to the inner part of the umbilical membrane. . . . I do not believe that any new vessels arise at all, but that the blood which enters them make them more obvious because of the colour which it gives them, and so by the augmenta- tion of their volume they become longer.

Wolff replied by another extensive piece of work, which he called De Formation: Intertinorum, and which appeared in one of the publications of the Russian academy for 1768. It mined preformationism. In it he demonstrated that the intestine is formed in the chick by the folding back of a sheet of tissue which is detached from the ventral surface of the embryo, and that the folds produce a gutter which in course of time transforms itself into a closed tube. The intestine, therefore, could not possibly be said to be preformed. From this as a starting-point Wolff went on to propose an epigenetic theory which applied the same process to all organs. It is interesting to note that the facts brought forward by Wolff have never been contradicted, but have been used as a founda- tion to which numberlcss morphological embryologists have added facts discovered by themselves. It is noteworthy too that although Wolff 5 second general principle, that of increasing solidification during embryonic development, led to no immediate results, it has been abundantly confirmed since then. His observations on the derivation of the parts of the early embryo from "leaf-like" layers were even more important, and acted as a very potent influence in the work of Pander and von Baer.

It happened, however, dint Haller had much the greater influence in the biological world at the time, so that Wold": conceptions did not immediately yield fruit in any general advance. Looking back over the second half of the seventeenth and the first two-thirds of the eighteenth centuries, it is remarkable how little theoretical progress was made in View of the abundance of new facts which were discovered. Punnett, in an interesting paper, has vividly brought this our.

The controversy between the Ovists and Artimalculkts had lasted just I oentury, and it is not uninteresting to reflect that the general attitude or’ science towards the problem of genetation was in 1775 much what it had been in 1675. When the period opened, almost all students ofhiology and medicine were Prefomiationists and Ovisrs; at its close they were for the most part Ovists and Prefotznationists.

Ovisrn sprang in the first instance from de Gra.1l"s discovery of the mammalian “egg,” which gave a new and precise meaning to Harvey's aphoristn. Preformationism, already old as a theory, acquired an apparent factual basis in the work of Malpighi and Swammerdam, and allied itself naturally with twism. With Leeuwenhoek and his sperma- tozoa, animalculism txrne upon the field. The main outlines ofthe battle which went on between the two viewpoints have already been drawn, but it is worth remembering that there were independent minds who were impressed by the obvious facts of heredity and found it difficnlt to call one sex essential rather than the other. Among these Needham and Maupertuis‘ may be counted. Among the lesser writers, James Handley with his Jlletlmnical Ermyr on the Animal Oemrwmy of 1730 ought to receive a mention. Though fond of theological arguments, ‘he upheld the common-sense attitude against ovists and anirmlcullsfs a‘.ike——“\Ve dissent in some things," he said. "bath from Lceuwcnhoeck and Harvey. . . . Both the semen and ova (notwithstanding all “His 1‘-‘in be said) we believe to he a mum sine qua mm in every Generation. But what finally killed anirnalculism was the discovery in so many’ Pl-‘"5 °i small motile living beings. flagellates, Protozoa. l-"E5 V‘l7"°3' 1‘ ‘V13 difiicult to maintain in the face of this new evidence that the spermatozoa were essential elements in generation, though the seminal fluid itself might very well he, as of course was Spallanzanfs opinion. The prefomiatian theory was what was holding up further progress, and when VVolff’s arguments prevailed in the very last years of the eight- eenth century, the way was open for the recognition of the true value of the spermatozoa.

I See the excellent review or Glass which describes. inler olia, zuruv=fl“"' xfivnm investigation of polydanyly in mm.

The physician rl'Aumont, otherwise unknown, who wrote the article on "Generation" in Diderot‘s famous Emyrlopaediu, brought this out in an interesting Way; for himself an ovist he summarised the arguments which in 1747 were destroying the anirnalculist position, and reducing rapidly the number of its adherents.

1. Nature would never be so prolific as to produce such millions of

spermatic animalcules, each one with its soul, unnecessarily.

2. The sperrnatic animalcules of all animals are the same size, no matter how large the animal is; how therefore can they be involved in its generation? 3. They are never found in the uterus after coitus, but only in the sperm.’ 4. How do they reproduce their kind? - 5. \Vhat evidence is there that they are any different from the animalcules (of similar shape, etc.) which are to be found in hay infusion, scrapings from the teeth, and many other places? No one supposes that these have relation to reproduction (Bourg'uet).

9. The Closing Years

The last forty years of the century were not marked by any great movement in a fruitful direction for morphological embryology, an iconographic wave of some merit due to Albinus, W. Hunter, Tarin, Senfi, Rosenrnuller, Danz and Sdn1mering' excepted; and it was not until 1812 that J. F. Meckel the younger translated Wolfl"s papers into German. This was one of the principal influences upon Pander and von Boer. In his introduction, Meckel describes how \Volfl"s work had been disregarded, and points out that Oken, writing in r8oG, had apparently never even heard of it. In the very early years of the nineteenth century morphologiml embryology received a great impetus however. One of the most interesting figures of the new period was de Lézérec, a Breton, whose father had been in the Russian naval service. The son, as a Russian

Loss of motility, and agglutination, doubtless disguised their presence from the investigators of this riod.

' \’nn Baer hirns¢lr(I8z7) believed in the extraneous nature of the xnimaleules, and attempted to zxprtss it by the name “Ape:-rmtoeoa" which he gave them. Not until Kfillsilterg work in r8.u wu theirhistogermu as non-ml |issue—¢l:mmn dernonscmed.

ntaxéal ;adet,b no doubt stunulated by the writings of .\vo1t-2, who had piml ah ‘ 5"“ _‘-"E. usedlo incubate eggson board slup. He eventually left t sea, studied medicine at Jena, and in 1808 wrote an excellent disser- tation on the embryology of the chick, which Stieda has recently brought to light. He then went to Pans, and taking a medical appointment at G“3d¢l°‘-‘PC. was lost to science. Very much more important was the work of Pander in 1817 and von Baer in 1828,‘ but it belongs to the modem penad,' and must be left for the next volume. Here it will sufliee to say that these great investigators established firmly the con- ception of the germ-layers (our now familiar ectoderm. mesoderm and endoderm) and clearly distinguished between their formation (“primary difl'erentiation") and the subsequent processes of histologiml and morphological difierentiation. At the same time the mammalian egg was at last discovered, and before long recognised as a single cell.‘ The nucleus of the egg-cell had been seen in molluscs as long ago as t79x by Poli in Xtaly, but not cleaxly dscribed until the xwtk of the Czech Purkyne on the "germinal vesicle” of the hen's egg in x825. In this my the road lay open for the triumphs of the mid-nineteenth century, when a Kowalevsky could reveal (1867) that such fundamental processes as germ-layer formation and gnstrulation were common to all animal phyla.‘

It is interesting to note, however, that the recapitulation theory, which was first clearly formulated by Von Baer, was already taking shape in ~’ various minds during the closing years of the eighteenth century.‘ Lewes has thus described the thesis of Goethe’s Morplxologie, written in

The more imperfect a being is the more do its individual parts resemble each other and the more do these parts resemble the whole. The more perfect a being is the more dissitnilarare its pam. In the former case the pun are more or less a repetition of the whole, in the latter case they are totally unlflte the whole. The more the parts resemble each other the lm subordination is there of one to the other: and subordination is the mark of high grade of

organisation.‘ “’i|liarn" and John Hunter belong also to the end of the oentury. '-‘For ‘xniumuftion an ‘van But. he Ks:-stv, Atddtfln; A. “I. Maya’. Ind. Stink-

h ‘ ( ' t z. _ H)I,fI“:$>.i‘s°gu:i>e hii:°mg?§Esms:§n.m's work on the development of Amphibxl (J. Pagel).

11,,-, k {S h ' 8 Cf. E. 5. Russell. A

c[_D‘g‘;!d:;rieO‘p'snplenl‘;ein1cet'w;|-glrélr|I?\x1ss3e<ll;md the classical mm at;-‘mt muoux. ‘ The history of the coneepraf rccspmzhuan has recently been gone mm with name thoroughness by A. w. Meyer (I93s).who is ttr-nzely rel!-Icunt '0 511*‘ '“Y6“'““ it earlier than Hunter. On the other lnnd, I am glad to Ice thxt Bxlsl (Iv: ) IS?‘ with me ugnausg Ammo: on this matter (m p. 49 of the vmcm hm‘)-

- L,-/-,, F, 353, ' 5:: Duncan.





ranvomcv mom 1920 a.d.to1p10u.d..



In his book on the anatomy of the gravid uterus, proved _ y an completely the (l'uEh‘0f the new that the maternal and foetal circulations are distinct. HIS injections left no shadow of doubt about ll"? lTlIll_1€l'. and the way was clearly opened up {or the study of am Pl’°P€l'tI¢8 of the Capillary endotheliztl membranes separating the bloods. a study which is still vigorously proceeding both in its histological and physroco-ehemicnl aspects. There was a quarrel between thehrothers over the priority of this dernonstmtion. John Hunter's Essays and Observa- tion: also contain matenal important for embryology.‘ His drawings of the chick in the egg were very beautiful, and are still in the archives of the Royal College of Surgeons. He adopted l\iayow‘s theory of the oflice of the air-space, and anticipated von Baer’s theory of recapitula- tion much as did Goethe.

If we were capable of following the progress of inurease of the number of parts of the most perfect animal as they were first formed in auccesion, from the very first to its state of full perfection, we should probably be able to compare it with some one of the intxztnplete animals themselves. of every order of animals in the creation, being at no stage difierent from some of the inferior orders. Or in other words, ifme were l0 talic 3 "55 05 an-lm3l3. from the more imperfect to the perfect, we should probably find an imperfect animal oorruponding with some stage of the most perfect.

It is interesting to reflect on the curious course which was taken by the essence of the idea of recapitulation in the history of embryology. As first formulated by Aristotle it was as much bodily 3-5 mental» but all his successors until the eighteenth century treated it as a psycho- logical rather than a physiological or m0rphol0glC3l “~ll=°TY. and lit“ themselves in speculations about the vegetativc. Senslllvc and rational souls. Yet the other aspect of the theory was only aslctp. and W35 des- tined to be of great value as soon as investigators began to direct their attention more to the material than to the psyclmlogitzl aspect of tilt developing organism.

Hunter did not absolutely reject preformationlsmi ll‘-ll l’¢Ef1“l¢l'l ll 3-‘ holding good for some species in the animal lilflgdflmi l1= ll1=1’¢f°Y° attached no philosophical importance to it.

Although “'olfl"s work did not lead to the immediate morphological advances which might have been cgpected. it was in mm’ "t"-‘Y5 {"‘“‘f“l- It stimulated J. F. Blumenbaclfs Ubrr Jen Bildifllglhitb Of 1739. fl_“’0fl§ which elaborated the Wolfiian tr‘: trrenlialir into tl1t=_III'-Y1l3[0f7fWl!’|lf. 1* directing morphogenetic force peculiar to living bodies. ll 15 lnlefcfilmg I Detailed and interesting Iurveys of John Hunter

have been published by A. W. Meyer (1915. 1916). In his address at the Hnrnrd Bicentenary Celebi-mom of EM!“-

to note that Blumenbaeh passed through an'exactly opposite succession of opinions to that of Haller, for he was first attracted by preforma- tionisrn, but being convinced by VVolfl"s work,‘ abandoned it in favour of epigenesis. Blumenbach comparu his ninufarmativur with the force of gravity, regarding them as exactly similar conceptions and using them simply as definitions of a force whose constant efiecrs are re- cognised in everyday experience. Blumenbach says that his nimr farmativur differs from \Volfi’s vi: errmlialxlr because it actively does the shaping and does not merely add suitable material from time to time to a heap of material which is already engaged in shaping itself. Woltfwas still alive at this time, but made no comment on Blumenbach’s ideas. He may well have thought the differences unimportant. Both Blumen- bach and Wolf? were mentioned by Kant in the Critique of jiuigment, where he adopted the epigenetic theory in his discussion of embryonic development. A word must be said at this point about the opinions of the eighteenth century on foetal nutrition. At the beginning of it there was, as has been shown, a welter of conflicting theories, and though later on writers on this subject were fewer, the progress made was no more rapid. In 1802 Lobstein was supporting the view (which had been defended by Boerhaave‘) that the amniotic liquid nourished the embryo per us, though Themel had shown forty years before from a study of acephalic monsters that this could be at most the very slightest source of material. These workers had obviously learnt nothing from Herissant and Brady, who had been over precisely the same ground fifty years earlier. On the other hand, Good and Osiander reported the birth of embryos without umbilical cords, so that the solution of this question became, in the first years of the nineteenth century, balanced, as it were, be~ tween the relative credibility of two kinds of prodigy. Nourishment per 0: was defended by Kessel, Hermes and Grarnbs, and was attacked by Vogel, Bernhard, Glaser, Hannhard and Reinhard. The idea lingered on right into the modern period, and as late as 1886 von Otr, who was much puzzled about placental permeability, decided that a great part in foetal nutrition must he played by the amniotic liquid. Weidlich, a student of his, fed a calf on amniotic liquid for some days, and as it seemed to thrive on this diet reported the amniotic liquid to have nutritive properties. The appeal to monsters was still resorted to at the end of the nineteenth century, for Opitz, in order to negative Von Ott's conclusions, drew attention to a specimen in the Chemnitz Polyklinik in which the oesophagus of a well-nourished normal infant was closed

' And by experiments of his own on the regeneration of hydroidr. ‘ fInm‘nm'on¢:. net. 382.

2 27 A msronv or cxuanrocoor

at the upper third without the development of the body having been in any way restricted. The fuller possibilities of biochemistry itself have sometimes been exploited in favour of the ancient theory of nourish. mm‘ Per 0!: thus Kfittnitz in 1889 collected some data about the pres- ence of peptones and protein in the human amniotic liquid with this object in view. That the foetus swallows the liquid which surrounds it towards the end of gestation in all anmiota can hardly be disputed, and as there are known to be active proteolytic enzymes in the intestinal tract, no doubt some of the protein which it contains is digcsted—but to maintain that any significant part is played in foetal nutrition by this process, has become steadily more and more impossible since 1600.

But to return to the eighteenth century; all was not repetition, occasionally someone brought forward a few facts. Thus the degluti- tion of the amniotic liquid was discussed by Flemying in 1755 in a paper under the title "Some observations proving that the foetus is in part nourished by the amniotic liquor." “I believe,” he said, "that very few, if any at all, will maintain now-a-days with Claudius de la Courvée and Stalpart van-der-Wiel, that the whole of its nourishment is Con- veyed by the mouth." But he himself had found white hairs in the meconiurn of a mlf embryo with a white hide. Both Aides and Swam- merdam had found the same thing. but Aldes did not think it of any significance, and Swamrnerdam merely remarked that the calf must lick itself in utero.

More interesting was W. Watson’s "Some accounts of the foetus in ultra being diflerently affected by the Small Pox.” This was the earliest investigation of the permeability of the placenta to pathological agents. “'I'hat the foetus," said Watson, "does not always partake of the _Infec- tion from its Mother, or the Mother from the Foetus, is the subject of this paper.” Two of his cases, he said, evince that the Child before its Birth, though closely defended frorn the external Air, and enveloped by Fluids and Membranes of its own, 15 not secure from the variolous Infection, though its Mot.her.has had the D15- temper before. They demonstrate also the very great Subtilrty of the vanolous

Etfluvia. But other cases

are the very reverse of the former, where though from inoculationthe mgst minute portion of Lint mo1sten’d with thc\:anolous Matter and _5PP]_‘°d ‘° slightly wounded Skin, is generally sufficient to pmpngyite this D1Sl_¢'“P¢1’u yet here we see the whole Mass of the Mother’; Blood. urcglaung dunno the Distemper through the Child, was not sulhuent to produce it. . . . Historic: it appears that the Child before IE mnh ought to 5* mm“ “

separate, distinct Organization; and that though wholly nourish'd by the Mother’s Fluids, with regard to the Small Pox, it is liable to be affected i.n a very different Manner and at a very different Time from its Mother.

Doubtless the modern explanation of \Vatson’s discordant results would be that in one case there were placental lesions, destroying the perfect barrier between the circulations, and in others there were not. In the last year of the century (but the seventh of the Republic) Citizens Lévcillé 8: Pnrmentier contributed an interesting paper to the journal de Physique in which they observed the increase in size of the avian yolk an incubation and spoke of a current of water yolkwards.


When the contents of this book was given in the form of lecturu at University College, it bore the title “Speculation, Obsenvagion and Experiment as illustrated by the History of Embryology.” Of the fin: two of these factors we have seen enough, but the third would have necessitated the continuation of the story down to the end of the nine- teenth century, and this must still await our projected second ‘-0113,15 The true science of experimental embryology did not come into being until the time of Wilhelm Roux} The early chemical Obsefijfions an the embryonic liquors (p. r 59) were indeed observations rather than experiments, and there was no systematic study of the changes which the liquors undergo during the development of the foetus; this was not done till the time of John Dzondi (1806). Harvey’s segregation of does at Hampton Court (p. r46) merits perhaps the name of an experi- ment, involving as it did die use of "cantrols;" and an outstanding instance is the ligature of Nuck in x691 (p. 163). As in Nuck‘s case, experiment in the hands of bath Spallanzani and J. T. Needham led to error. Spallanzani confuted his adversary on the question of spontaneous generation and the vegetative force by what amounted to rigid criticism of experimental conditions, but later on denied their proper function to the spermatozoa on exactly the same methodologically faulty grounds. But on the whole, experimentation, active interference with the wurse of Nature and subsequent observation of the resulting system in comparison with systems in which no such interference has taken place. was a characteristically nineteenth-century product as far as biology and embryology are concerned. Only at the prrsent day, indeed, are rte beginning to appreciate the statistical and other difliculties attending upon the full applimtion of the experimental method to living organ- isms, and the manifold obstacles which prevent obedience to the rule that only one variable be modified at one time. But this is no matter of reproach against the older embryologists. Knowledge of form must neoessarilypreoede knowledge of change of fon-n and the factors pro~ ducing it, and so we see during the last seventy years the production of

Info]. Entu-'ckbmgrm:z}:4nilr in 1394. thus naming ta! morphology and arm! embryology.

‘ 185°-tgu. He founded the Art the modern discipline of erpenmen

"Nomialtafeln" or tables of morphological pictures showing normal development; these are the essential basis for experimental studies.

Probably the best way to summarise the influences which have operated in the history of embryology is to concentrate attention on what may be called, borrowing a phrase from general physiologifi-the‘ "Limiting Factors” of advance. We may thus regard the progress of knowledge about generation as governed by a reaction-chain, one link in which may at any given time be slower than all the others, and hence may set the speed for the whole.

Relation of lnvesrigarnn to their environmmt

Co-operation of investigators

Prminl mm“: {mm recrmique<n . Terminologzcll C Wftufli UAIRUSGVE Cancepruzl { Destructive


Balance berween Speculation, Observation and Experiment

Of these limiting factors the first which may be mentioned (though I do not wish to pronounce here upon their relative importance) is the relation of investigators to their environment. The Carlylean tendency

to regard the history of science as a succession of inexplicable geniuses arbitrarily bestowing knowledge upon mankind has now been generally given up as quite mythological. A scientific worker is neoasarily the child of his time and the inheritor of the thought of many generations. But the study of his environment and its conditioning power may be carried on from more than one point of view. We have already seen (p. H 5) what a sharp distinction the culture-historians (Sigerist, Bili- kiewicz, etc.) make between the mental atmosphere of the Renaissance, the Baroque, the Rococo, the "Au£‘k!.amng" period, and so on. There is doubtless much to be learnt from historical investigations carried on in this light, but it may sometimes lead to a hypostatisation of abstrac- tions, and as in the case of ovism and feminism (p. H6) its results may border on the fantastic. The social and political ruling ideas of a dis- tinguishable epoch play, on this view, an overwhelmingly important part in the scientific thought of the time, and may act as limiting factors to further advance. Thus the politiml absolutism of the Baroque period is thought to have mirrored itself in the extreme rationalism of seven- teenth-century biology. There is much more to be done in working out the internal relations of successive intellectual climates and their con- nections with contemporary social situations.

The other principal point of view which may be taken regarding the environment of the scientific worker as a limiting factor is that which emphasises his existence as an eccnoniicunit, andscelts to show horrhis position in a society with such and such a class structure influences the development of his thought. Some reference to this point of View has already been made in the introduction (p. r4). It seems to ofl'er more chance than the preceding method for new discoveries in the history of science, for it directs its attention upon those aspects of human society (trades and techniques, labour conditions, the evcryda ' life of the mine, the factory, the barber-surgeorfs shop) which, precisely because of their assumed inferiority, have not been discussed in the majority of books, written inevitably by members of the goveming classes, or by those who aspired to imitate gentility. Thus the rather sharp cleavage between the philosophic biologist of the Hellenistic age and the contemporary medical man, who might often be a slave._c_on- tributed doubtlss to the sterility of ancient Mediterranean x;nCdlClIlC, including obstetrii: and gynaecology. ln the later Christian \\ est there was not much incentive to embryological study so long as the process of childbirth was left to the charms and incantations of barbarous mid-wives. But for a better insigbl l“‘° am °°°’,‘°m‘° P‘-"5‘"°“ °f °mb’5’°' iogiscs in past ages nearly all the work remains to bedpne. .

Next Domes co-operation of scholars. In the civilisation of the Hellenistic age, it may ‘be said, a considerable measure af_ such 43* operation had been attained;.the iivorks of Aristotle and Hipgsocss as were fairly readily available in iyritten form, and evidence CE: brought {onward (pp. 54: 73>. pm-cuiulywuh r=sardt°J=“r1sb "*g.“§ ~ that this was well used. But we must beware here of sufienng. a is ctr‘; (ion of perspective in the contemplation of antiquity,_lu1l’ It’: ¢35Yu1d exaggerate the co-operation of ancient thought A 5“‘l:' '5 ‘ :‘c::cm consider itself lucky if it passed once in twenty-EVE 1:313 fljcfin Greece and India after Alexander 13- 27)- Am°"E °°‘t'ms B: influences that save fis= *° ‘‘*° '-‘“i“'”?'°.“ "5 "“ ‘-M flinehanci aperation,hamperedbyenoimouslmgltlsfltf d1fli°‘1l“_53c;“ dihcolo _ul and by the diversion of interest £rom scientific to etlhrl 311 C mVc3‘!hc channels on the other, sank to a V60’ 1°" l°‘°l' ah°‘:l°°of‘;is wmem_ |’¢m5“'k“b1° Specude uf 2 Leqnirdo’ many Y?’ Z of fortifiutlonsi pomrics’ and able to am 3 lmngfmly all adimgriries to any living finding it impossible to ooxnmmucate Ksgovks my by 3 mm person. and reduced to 5“-YY“‘§ am“ “‘ “° C M chance available to SC.l‘tOl£il'S of after

Among the most important of limiting factors we must reckon technique, extending the term to cover mental as well as material methodology. The part which the latter has played in the history of embryology can hardly be overrated. Thus until the introduction of hardening agents, especially alcohol, by Boyle 158), the examination of the early stages of embryos was bound to remain crude, and we have seen (p. 185) how embryology attained an entirely different level im- mediately afterwards in the hands of Maitre-Jan. The parallel case of the microscope is too familiar to dwell on, but the work of Malpighi obviously marked a turning-point in the science (p. r63). It may here be noted, however, that even when methods are available, the workers of the time do not necessarily use them, and although Harvey could have employed an early {am of microscope, he restricted himself to the weak lenses, pnrpfdlia, or "perspectives,” which had already been used by Riolanus.‘ A still more striking instance is that of artificial incubation. Carried on in Egypt since the remotest antiquity (p. 22), this process must have been at the disposal of Egyptian physicians, Alexandrian biologists and Arabic scholars for a period of three thousand years, yet so far as we know, no embryological use of it was ever made. In eighteenth-century France and England the technique of the process had to be painfully rediscovered at a time when biologists were only too eager to make use of such assistance. Let us mention, as other instances of the effect of material technique on embryology, the burst of knowledge which followed the invention of the automatic microtorne by Thrclfall and others about 1883, and the great advance which in our own century has followed the successful mastery of graft- ing technique in operations on amphibian embryos by Gustav Born and Hans Spemann.

Just as important, however, as material technique is mental technique. And first with respect to words; on several occasions we have had to notice a standstill on account of the lack of a satisfactory terminology.

‘ As this rtzrterxrent about Harvey may seem Iurprising to lame, it is worth while to rcopitulate briefly the fact: thou! the iruention of the microscope. For the detailed tvidenee the elassrml papers of Singer may be consulted. The introduction of convex lenses Is spectacle: for presbyopin may now be dated very soon after A1). 1286 in Iuly, recording to the ¢X.hl|.I3LlV€ researches ol Roscn. Concave lenses for rnyopin came into use much later, about xsoo, Ind the first bilcnticuhr ryutem was probably due to Leonard Digger (d. r57r), who invented (but never fully described) I rudimen- tzry telescope. The Eu-rt lens combination for a microscope Wu mentioned, again obxcurcly, by Gin:-nblttistx dell: Port: (1540-1615) in his Jllagiae Nannalix (1559), but no practial nppliution was made cm. The rnicroscope mu, begituwith bclurias Jansen (I530-fa. 1630) of Middclburg in Holland, who put together two convex lenses rome time between 1592 and recs, as we know from the detailed Ineount of Pierre Borcl. Comoliu: Drebbel of Alcrnur, nuthcxmticiln to King June! in X619. brought one or these instruments into England. i e. thirty year: before the conclusion of Wrtlrun Haney’: researches on genennon. Perhaps Harvey made :Eom to acquire and me one, perhaps he was too uonren-nL~i-V: and letpticaI—'we do not know.

Thus in the thirteenth century Albertus of Cologne had arrived at a Pom: bcwnd “inch P1987335 was impossible in the absgnee 1‘ words’ when’ {°' ‘“-mPl°o ‘hut Was no other means of descrihingnte}: sero-amniotic ’unction ’ t c h

on the left sidje of the i’-essigl whicherizghsllhébloiig bihn«:hi: htfiic right hand of something else” accuracy was difficult and s eed in‘: C aim“ A P795551)’ Sirnilar position was occupied by Boerlliaave Clflhtcfimh 591""-1'Y. only now in the case of biochemical word; Factd “-kh.s°m° 5"b5“"“°° sud‘ 35 3 "EYCOSY, streaky yellow oil, smelling of 3u“111n¢ 531$." Boerhaave unable to describe it except in thsc corn. m°"‘5_¢D5° "M15: and lacking the means either to submit it to further analysts or to characterise it by accurate physiCo<.hcmiml units, he was forced to admit a large number of ultimates into his schemes which were not ultimate at all.

Mental technique as a limiting factor in embryologiml history goes deeper than words, however, for it involves the concepts of the investi- gator. What the Germans mil “Begrifl"sbildung" or the construction of concepts congruent with oertain sorts of natural phenomena. though never conscious in the history of biology, has none the l$s been open. tive. In this field we may remember the doctrine of Galen conoeming the natural faculties (61-rci;:s1;. p. 70), and the immense length of time which was required for biologisw to see that it was nothing more than 8 concise statement of the phenomena themselvms. Not until it was "seen throng " as an explanation was post-Renaissance biology pas. sible. Similarly, the peculiar contribution of Leonardo to embryology was his realisation that embryos could be measured. not merely as to dimensions at one moment but as to dimensions at a suocasion of moments. The applicxtion of the concept of change in weight and size with time, a concept which, as modern biology $hO\\S, admits of much accuracy when properly worked out, was thus first made by Leonardo. In the same way Boyle was the first to see clearly that a problem of mixture is presented by the developing embryo (though Hippoa-ans had stated it dimly some two thousand years before). If the embryo is made up of mixed things, some definite proportion and way of mixture rrnrst exist. And no hope of finding out what this was could be obtained from the Aristotelian elements (heat, cold, moisture and dryness) or from the alchemical principles (salt, sulphur and mercury). Hence Boyle’: emphasis on the corpuscularian or mechanical hypothesis. and all its historical implications (p. 176). Besides this creation of concepts, and the choice of which of them to apply, the mentality of the scientific workers of the past often differed greatly with regard to a fundamental quality which can called audacity. Probably Aristotle's greatest claim to our respect is that alone of his contemporaries and predecessors he had the audacity to suggest that animal farm is not limitlessly manifold or infinite in its manifestations, but that given industry and intelligence, a clzssifizmtion was possible. This alone marks him out above all subsequent biologists. On a smaller scale, we find the same mental audacity in Kenelm Digby, whose discussions of the development of the chick are remarkable for their naturalistic tone (p. 122), for their conviction that the proccssu of development are not beyond the reach of the reason and imagination of man. It is most ironic that Digby, who did little or nothing himself to advance our knowledge, should have spoken thus, while his great contemporary, \Villiam Harvey, to whom we are indebted for so many advances in embryology, was led to despair of understanding develop- ment. Another interesting point that emerges from the same period is that such mental audacity can go, perhaps, too far, as when Descartes and Gusendi built up an embryology more geometiico demomtram, in which the facts were relegated to an inferior position and the theory was all.

But not only must the right concepts be chosen, the wrong ones must be abandoned. One of the principal necessities which has faced investi- gators since the earliest times has been the recognition of silly questions in order to leave time for the examination of serious ones. It was presumably inevitable that the pseudo-problems concerning the entry of the soul into the embryo should be taken seriously until a very late date. But a more typical instance of a meaningless question may be found in the dispute about what parts of the egg farm the chick and which feed it. The tacit assumption here was that since to common- sense food and flesh are different things, there must be in the hen’s egg. bsides sufficient provision of food, some sort of pre-flesh out of which the embryo can be made. Not until 1651 did this pseudo-problem go out of currency in the light of Harvey's demonstration of the unsound- ncss of the assumption.

The expulsion of ethics from biology and embryology forms another excellent example. That good and bad, noble and ignoble, beautiful and ugly, honourable and dishonourable. are not tenns with biological meaning. is a proposition which it has taken many centuries for bio- logists to realise.

Idus of good and bad entered biology partly under the concept of “perfection? In 1260 Albcmrs was maintaining that male chicks alvay: hatched from the more spherical eggs and female chicks from the more oval ones, because the sphere is the most perfect of all figures in ’ solid geometry. and the male the more perfect of the two sexes (p. 87).

We realise today that to ask which is the more perfect of the two sexes 18 1| meaningless question, for we have expelled ethics from science and °=’“}°‘ |’=Lr~1fd any one thing as being more perfect than anything else. A831": describing the course of the arteries in the developing chick

Mb"-""5 “F53 "One of the two pusagm uhich spring from the heart, b"1"°hC3 info ‘W0. One of them going to the spiritual part which contains '-he ha“. and carrying to it the pulse and subtle blood from which the lungs afldotherspiritual parts are formed; and the other passing through the diaphragm to enclose the yolk of the egg, around which it forms the liter and stomach." This distinction between the organs above the dflphmgfn. the lungs, heart. thymus, ete., cxlled "spit-itualia,” and the organs below, the stomach, liver, intestines, spleen, ete., runs through the whole of the early anatomy. It was as if the organs of the thorax were regarded as a respectable family living at the top of an otherwise disreputable block of flats. To us it seems absurd to call one organ more “spiritual" than another, but that is because we realise the irreletance of ethic-:.l issues in biology. Thomas Aquinas, about the same time, dealt in passing in his Summa 77zealagz'm with human generation (p. 93).

The generative power of the female is imperfect compared to that of the male; forjust as in the crafts, the inferior workman prepares the material and the more skilled operator shapes it, so likewise the female generative virtue provides the substance but the active male virtue makes it into the finished product.

This is really the pure Aristotelian doctrine, but St Thomas gives it the characteristically mediaeval twist. Aristotle might make a distinction between form and matter in generation, but the mediaeval mind, with its perpetual hankering after value, would at once enquire which of the two, male or female, was the higher, the nohler, the more honourable.

In the eighteenth century the same frame of mind persisted. It was maintained that in every detail of the visible world some evidence could be found for the central dogma of natural religion, the belief in 3 .l“33 and beneficertt God. Biology was thus not free from the mental [>515 associated with theology.‘ Between 17:20 and 1850 a multitude of books were written which purported to reveal the wisdom and goodness of God in the natural creation. The theologians tool: what suited their purpose and left the rest. It is instructive to see how Goethe. who was deeply committed to the theologiml interpretation of phenomena, re- acted to the ornithological anecdotes of his secretary E'cke1-mano on 18th 0et., x827. He said little while Eckerrnann told him about the habits of the cuckoo and other birds, but when Eckerrnann related how he had liberated a young wren near a robin‘s nest and how he had found it subsequently being fed by the robins, Goethe exclaimed:

1 Far I ‘pg-[king mmple of this. see Edmund Gone‘: I-‘atlnn and San.

That is one of the best ornithological stories I have ever heard. I drink success to you and your investigations. Whoever hears that, and does not believe in God, will not be aided by Moses and the prophets. That is what I call the ornnipresence of the Deity, who has everywhere spread and implanted

1 portion of His endless love.

And so it always was with the theological naturalists; they hailed with enthusiasm the discovery of monogamy in tortoises, or mother-love in goats,‘ but they had nothing to say concerning the habits of the hook- worrn parasite‘ or the appearance of embryonic monsters in man. Not until the beginning of the nineteenth century did it become clear that Nature cannot be divided into the Edifying, which may with pleasure be published, and the Unedifying, which must be kept in obscurity. Experimental embryology then contributed to this clearer vision of the living world by its manifold demonstrations that in spite of the appar- ently deeply teleologiml character of normal embryonic development, once the individual morphogenetic processes have been experimentally "derailed," they laboriously continue their operations so as to imitate (and therefore ultimately to explain) all the possible varieties of naturally occurring monstrosities.' Of course the riddle of nomml integration remains.

In the end we may say that the progress of a branch of natural science such as embryology depends on a delicate balance of three things, speculative thought, accurate observation and controlled experiment. Any modification of the optimum balance will act as a powerful limiting factor on progress. Speculative thought, in particular, has shown a tendency to crystallise too readily into doctrines which, by way of attachment to some philosophicafor theological issue, live a longer life than they deserve. Thus the Aristotelian theory of the {emotion of the embryo by the coagulation of the menstrual blood, built in the first

‘ One find] stfilting parallels for this interest in animal behaviour among the neo- Cornfucian school of philosopher! in mediaevnl China (see Sciznzz and Civtlirntion in China, vol. a). In so far as it contributed to n usnviction of the reality of an evolutionary which for the Chinese thinkers it certainly did, it was useful and commendable.

ut that-uheywere never committed to the idea of a rpeciat creation by uuu-benzfieent personal deity. For them thertforc "gleam: of righteousness” in ants and alien were pres: of that human community which the im enonal Order of Nature (the Tao) wool in due time produce, pieces of evidence 1 out 3 social evolution, not about a personal Creator.

‘The guinea-worrrI(Dra.I.1mnlIu.r irmimemis) had been givzn . drunatic d:_scrjption by Vzlsch in r67.;—nnd indeed by Avicenna long before him. A.n.Lylostorru1ns had been known Ind described in ancient Egypt.

‘ On lethal genes and their action, see the brilliant book of Hxdorn.

instance upon a faulty deduction, became incorporated in the Aristote- lian tradition of form: and materia, and although quite repugnant to ob- servation, remained the official theory throughout the European Middle Ages, and apparently perpetuity in India. So ponerful was the rationalism of a medical education round about 1639 that the physicians to whom Harvey demonstrated the empty uteri of the king's does preferred to believe their books rather than the evidence of their senses. And precisely parallel to this attitude, as we have seen (p. 213), that of the preforrnationists in the following century, who, having dccfl‘-ltd. like Bonnet, that epigenesis was inconceivable, only aooepted such observations as coiilinned their upriari view.

Prefomiationisin as a manifestation of rationality merits’ further examination. The dogmatic manner in which preforinaticnism W-is held during the eighteenth century would not perhaps have bceit 50 fatal if the biologists of that time had been able to take rnatheniatical reasoning more seriously. There um Harvey's yery convincing ai‘B“' inent about the circulation of the blood, and Freind's equally convinc- ing. but unfortunately erroneous, deductions about the quantity of menstrual blood and the weight of the newborn foetus (p. 1'50). If these could have been accepted, it was a pity that I-Iartsoekers argurnglit about preforrriation could not. In 1722 Hartsoeker caleulated t I l0‘°°'°°° rabbits must have existed in the first rabbit, 35-Shummg '11“ ‘he creation took place 6000 5'63” 33° and lb“ }"’bb"s bggm la "Pmduc; their kind at the age of l‘KblOnlll1'|Sd;'nBUt to I135 B‘J}:sn€tY°"c1:“"3)¥' thatitwasalwayspossi 6, Y3 33"“ ‘’““f ' . ' nation under the weight of numbers, and he described !h= P"5‘{"m3“°“ theory as one of the most striking victories oi the understa:_u‘g:Ig 0:: the senses. It would have been better dumbed as one 0 9 In striking victories of the imagination over the understanding. . d ‘E

The fact is that the biologists of the eighteenth century, cxirne Jar: by Prefoririationist theory, tool: emb1'Y°l°EY °“ ‘"3 Pmteagllc ‘he observation became superfluous. They rrould have founm alieepd hm’. sentiment satiriscd by Boyle that “ us. mote E 3:11" P do‘ sophiczil to argue a prion" than 1'1?”-"”“’”' and min ewilidionythnt barred from looking at devclopmg €mbl_'}‘03 by their con‘: ‘her may structure and organisation would Ifertflilili 59 ‘l‘°‘°' W cm {ad I '°°"’d 5°‘ i‘ °‘ “°“ Th‘ "'°{°'"“"°"’“ mwowtify liiiaibhalists {mi “P5550” in bi°l°gY °f ‘he mnufltfly between C rtionalists were the ernpiricists in pliil0S°PhY- The wnmmpomry ta

Pwpk who held that - - - i f’ mpmatien which humwI>=ina= "=r°i*=P‘zss°?“°“°‘°‘"““ ‘3“"”*’ “° “‘ d aids. be were not simply g€fl€1'IlJSal.IOEB from expencntfi, 5“ W“, "C"

used as major premises in arguments concerning Nature. If observations weie not in accordance with expectations founded on such reasoning, they were dismissed as illusions. The empiricists, on the other hand, held that there was no knowledge independent of observation, and that the rationalism’ principles, in so far as they were admissible at all, were generalizations from experience.‘

It is obvious that nearly all the preformationists were rationalists. They thought that Reason was in a position to decide the issue whatever might be the results of observation. "It is remarkable,” as Cole says, in his book on this period, “that the preformationists did not realise that if the point to be established is assumed at the outset all further discussion is superfluous.” In this example, then, we have a disturbance of the balance towards the side of rationalistic speculation.

It would be a mistake, however, to regard this tendency as confined to the eighteenth century. Ample examples of its presence can be collected from nearly every period in biological history. “We plume ourselves,” says Cole, “on that aspect of our work which is vain and argumentative, and condescend to the more modest but enduring labour of observation." There can be no doubt that this state of afiair-s, so unfortunate for science. is one aspect of that contempt for manual labour which has run through the Stratified structures of all societies in the history of civilisation. The manipulator of paper and ink, educated in the classical traditions of his time, has always seemed, by reason of his superficial similarity to the political administrator, a superior being to the empirical mechanic engaged in the manual work of the arts and industries. The tradition is as old as civilisation, yet for the advance of science it must be broken. Not until the manual worker and the audacious theorist are combined in one person will the fullest develop- ment of scientific thought be possible.

On the other hand, there can be no doubt that a plethora of observa- tion and experiment is also had for scientific progress. Modern biology is the crowning instance of this fact. What has been well called a

"medley of ad hoc hypotheses" is all that we have to show as the theoretical background of a vast and constantly increasing mass of observations and experiments. Embryology in particular has been theoretically dxraidbare since the decay of the evolution theory as a mode of explanation. Embryologists of the school of F. M. Balfour thought that their task was accomplished when they had traced a maximum number of evolutionary analogies in the development of an animal. \ViIhelm His, perhaps the first causal embryologist, struggled succtssfully to end this state of affairs.

My own attetrrpts [he wrote in r888 in 2 famous passage] to introduce mm: elementary physiological or mechanical explanatinns inta ernbryolog have not been generally agreed to by mnrphnlogisu. To one it seamed ridiculous to speak of the elasticity of the germinal layers; anatherrbought that by such considerations we put the can befure the horse; and one recent author states that we have something better to do in embryology than to discuss temions of germinal layers, etc., since all ernbryologiml explanation must nccaxtrily be of a phylogenetir: nature.

But this strictly evolutionary dominance in embryology did not last on into the twentieth century. The unfortunate thing is that nothing has so far been devised to put in its place. Etperimental embryology, Morplrologiml embryology, Physialogical cmbryology, and Chemical embryology {am today a vast range of {actual knowledge, without one single unifying hypothesis, for we cannot yet dignify the axial gradient doctrines, the field theorim and the speculations an the genetic control of enzymes, with such a position. We cannot doubt that the most urgent need of modern embryology is a sex-res of advances of a purely thearetitzl, even matlrerrrntiw-logical, nature. Only by samething of this kind can we redre§ the balance which has fallen over to observation and experiment; only by some such cflort an We obtain a theoretrral embryology suited in magnitude and spaciousrtess to the wealth of facts which contemporary investigatnrs are accumulating day by dd)’-


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The following work remain in-accasihla to me in Englfind 1nd 1 MW 110‘ yet seen them: Mzntcllassi, c: Dr'vm1'S:'.rmm':an L: Gtnzraziane. (norcncc, X749-) Rvpfimd in Rama’! dz pike: tie Illldez-in: (Paris, 1763) and in Pféa: inllrmdnfef

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1 would like in express my gratitude to the followifig (07 15° 5°}? “"7 ha" given me in tracing thcs: and other books: Dr 5- Asdfll. M154 Ethtl G- Bmdie, D: R. E. D. cm, Dr E. J. mngwau, Mm Elmer Gregory. Pr Amald C. Klebs, Mr W. B. McDaniel, Dr Want! Pagcl and Mr H. Zurlinger.

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