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M A T T E R, L I F E,. · A N D
GENERATION
Eighteenth-century embryology ·and
- . the Haller~Wolff debate
S H I R L E Y A. .RO E
Departmem of the History of Scienc,
Harvard tfnivmity
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lMSTITUTO DE B!OC!ENClAS
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Contents
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1 _Introduction: mechanism and embryology 1
INSTITUTO DE BIOCIENCIAS"
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Illustrations
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Although clashes occurred between preformationis~s and was first formulated in the 1670s with the work oIMalebranche,
epigenesists throughout the Enlightenment, perhaps the most , Swammerdam, Perrault, and others. These theories of preexist-
important of these was the debate that took place between ence, based for the m.ost part on the concept of emboitement
_Albrecht von Haller ( 1708-77) and Caspar Friedrich ,Wolff (encasement), did not grow directly out of the earlier preforma-
(1734-94). Haller, a renowned scientific figure, announced his tionist positions (see note. 1). Rather, they arose in response to a
support for preformation in 1758, just one year prior to the set of difficulties and concerns that were prompted by -the
publication of Wolff's doctoral dissertation, whic.h strnngly appearance in the mid-seventeenth century of several epigenetic
endorsed-epigenesis. The ensuing controversy, lasting for over theories of development, pi:opounded by ·H~rvey, Descartes,
a .decade, crystallized many of the ,key issues of eighteenth- Highmore, Borelli, and others. '
century embryology. The role of mechanism in biological.ex- Both Aristotle and Galen had proposed theories of epigenesis,
planation, the r~lationship o_f God to his Creation, the question albeit with important differences between their two systerns 2 ;
of spontaneous generation, an.d the problems of regenerarion, and it is largely within the context\of Renaissance scholasticism,
, hybrids, and monstrous births-aHthese were poii-1ts ofissue in the heir to these two great thinkers, that William. Harvey's
the Haller-Wolffdeba.te. Exercitationes de generatione animalium ( 1651) must be understood.
More importantly, the controversy betweeil Haller and Wolff Written largely as a commentary on the thepries ·of Aristotle
iHustrates the fun.damental tie between biological and philo- and Fabricius ab Aquapendente, Harvey's work was based on
sophical questions that existed in the Enlightenment period. observations that· he had made on deer and on incubated
Philosophical concerns were in fact largely responsible for the chicken eggs. Harvey combated the Galenic and Hippocratic
rise and popularity of preformationist theoriesover epigenesis. two-semen theory then in vogue by dai,ming that he could find
The d~sh between Haller and Wol'ffepitomizes this philosophi:- no female semen and by showing that in female deer dissected
cal nature of eighteenth-century embryology, for Haller and shortly af_ter copulation there was no evidence of male semen
Wolff came from widely divergent philosophkal 'backgrounds. entering the uterus. Consequently, the embryo could no~ be the
Haller, a Newtonian mechanist and a deeply religious man, result of,the mixing together of male and female seminal m~te-
held beliefs about the nature of the world and about scientific rial. Furtl:lermore, through his famous dictum "Omne vivum ex .
explanation that differed -fundamentally from' those of Wolff, ovo," Harvey proposed that all organisms, viviparous as well as '
.whose own viewpoint- derived largely fram the tradition of . oviparous, develop from a. primordial egg. Not a preexisting
German rationalism. Their debate over erribryol<,>gical develop- germ, the egg was thought by Harvey to be a product of con- ·
ment can be fully understood only when viewed as a controversy ception. Finally, Harvey's detailed observations of day-by-day
over these. underlying philosophical differences .. Furthermore, development of chick embryos led h~m- to conclude "that the
as representatives of two majQr Enlightenment schools of g~neration pf the chick from ·the egg is. the result of epigenesis
thought, Haller and Wolff illustrate two important cases of the ... and· that all it~· parts are not fashioned simultaneously, but
w:ays in which philosophical issues guided much of eighteenth- emerge in their due succession and order" (1651, 1847 trans.:
century embryology\ As s1;1ch, an analysis of their work· and 336).l . .. . .
..
their_ controversy .sheds light on a m.im~r of aspe~ts of biolog- Although Harvey's work provided significant improvements·
ical, thought during .this period. , - over the theories of his predecessors,. his views had limited
success, partly because, by 1651, his Aristotelian mode of argu-
me_ntation had begun tQ seem out of date. Contemporaneous
THE RISE OF PREFORMATION THEORIES with Harvey, however, was another.promulgator of.epigenetic
Although the.re were those who, before the late seventeenth deve!opment, Rene Descartes~ who ,was the first to offer an
century,_ believed that the· embryo was in some fashion pre-. explanation for generation based solely on matter and motiou.
formed in the body of the parent before conception, the' notion Descartes dealt with animal physiology in his Traite de l'homme,
that all eµibryos had existed .frotn the beginning of.the world the second part ofhisM~. which was wr~tten in the 163os but
\ '
4 Matter, life,.andgeneration :nuroauc,wn !)
· s'1ppressed by Descartes because of. its Copernican viewpoint. · This insufficiency of mechanical explartations of gradual de-
(Galileo. had recently been con~emned for like views by the velopment was an important element in the rise of preexistence
Catholic church.) Published after his death, Descartes.'s'Traiti de the9ries. As Nicolas Malebranche noted in 16S8 concerning
l'hom~ proposed mechanical explanations for the phenomena Descartes's theory of epigenesis, "The rough sketch. given by
of sensation, muscular movement, digestion, the circulation of . this philosopher may heip us understand how the laws of
the blood, and other vital functions.• Yet generation was left motion are sufficient to bring about the gradual growth of ,the
untreated, for Descartes rema,_rked in a let.ter to Mersenne that parts of the animal. But that these laws could form them and ..
he had given up trying to deal.with this subject.in his treatise link them together is something that no one will ever prove"
(Descartes 1964-74, 1:254). In the late 1640s, when Descartes ( 1688:264}; It does not seem possible that' mechanicallaws
returned again to generation, he described this earlier stage in could both fashion and organize the parts of the organism. "I tis
his thinking. about the animal organism, noting "I had almost easy. to see," Malebranche declared, "that the general laws of the
. lost"hope of finding the cau~s of its· formation." Yet, he pro- communication of motion are too simple for the cons~ruction of
claimed in this same letter, "in meditating thereupon, I have organic bodies" (p. 252). Mechanical causes may be part of the·
discovered so much new land, that I hardly doubt that I can process 9f development, but they cannot account for; reproduc..:
complete the whole of the Physics according to my. desire." tion itself.
(5:261). Descartes saw the generation of animals as the last Malebranche waslhe first,to fully art:iculate, in ·167.4, a theory
segment to come under the wing of his mechanical philosophy, 9f preformation by embo~. In discl,lssing the ljmits of our
forming there-the completion of Cartesian.physiology. . senses, especially vision, Malebranche turned to bioiogical ex-
The result of Desc~rtes's delibera:.tions·in the late 1640s w.as a amples. If one looks closely at_ a .tulip bulb" using a magnifying
small treatise De la formation de l'animal, published posthumously lens, one can see aH the parts of the future tulip folded up in
along with the Traiti de l'hotnme in 1664. Here Descartes pro- miniature inside the bulb. And one can assume that tl)e same
posed an explanation f~r development bc1sed entirely upon the may be the case in the -seeds of all trees and plants. ''It does not ·
movement of particles. According ta Descartes's Sfstem, repro- even seem unreasonable,t' Malebranche declared, "to think that
duct.ion begins with'the mixing of semen from the. mfile and there are, infinite trees in a singte g~rm, since it does not contain
fem ate, resulting in a Jerme:ntation of particles. The move01ent only the tree which is the seed, ,but also,a very great number of
of these particles leads to the formation of the· heart, followed other seeds, which are all enclosed in those of the new tree .. : .
by the other embryonic parts. "If one knew what all the parts of one can say'that in a single apple pit, there would be apple trees,
the ,semen of a certain species of animal are, ht particular, for apples, and the seeds. of apples .for infinite or almost infinite
example, of man," Descartes declared, "one could deduce from . centuries" (1674:82) .. Each seed, then, would contain the seeds
this alone, by reasons entirely mathematical and certair:i, th~ of all future individuals, encased .within one another. This no-
whole figure arid conformation of each of its members" ( '1664: tion can be extended to cover animals as well: "One sees in the
146). Through matter and motion alone, one can explain not germ of the bulb of.a tulip the entire tulip. One sees also in the
only the inanimate but the ariimate world as. well. "It is no less · germ of a fresh egg, and which has not been covered, a chicken
natural for· a. clock, made of a certain number of wheels, to which is perhaps entirely formed". One sees frogs in the eggs of
ind1c.ate the hours,"he proclaimed; "than fora tree born from a frogs, arid one will see other animals in their germs, when one
certain seed, to produce a particular fruit" ( i644:326). has appealed to and experimented enough to discover them"
Descartes's explanation I for embry<>logical development by (pp. 82-83). From all of this evidence we can conclude, accord-
mechanical causation was not a successful one; ln particular, it ing to Malebranche, "tbat all the bodies of men c1nd of animals,
failed, as did those of other mechanistic epigenesists, to explain which have been born up to the consummalion-of the century,
why development proceeds as it does, that is, why the proper have perhaps bee-n produced as long ago as the creation of the
organism, with its parts perfectly arranged, is formed. How world" (p; 83; see ~lso &:,hrecker 1938).
does a process based on matter and motion alone result in a In his discussion of animal embo~t~ Malebranche referred
co~plex living organism? - · to the work of Marcello Malpighi on chick eggs and of Jan
·.
6 Matter, lij.e, ana generation • , •• , vuu.1,,••v1•
ment), made explicit his concern over the limits of mechanical anism and blind chance. Matter was viewed by most as entirely
laws; As he remarked in 1680, "I do not know if oric can passive, put into motion only through mechanical laws. But
comprehend how a work ofthis quality would be the 'effett of , since these laws of motion are blind, that they could know ho;w
the ordinary forces of nature.... for I find finally that it .is to form a living organism seemed out of the question. Both
scarcely more inconceivable ... that the world has be~n able to self-active matter and a Goc:t actively involved in each. instance of
form. itself from matter our of chaos, than ail- .ant can form generation were ruled out in the mechanistic universe of late-
. .
seventeenth-century thinkers. The . theory of preformation
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role• was left ·for the Divine Creator? Preformation through Outing the first half of the eighteenth tentury,'regeneratjon
emboitement provided a solution to this difficulty while still pre- became a subject of incr~asing study. Both Thevenot" and
serving a mechanical universe, All organisms were 'formed by Perrault ~ad described regeneration in lizard tails in the 1680s,
God at•t1'eCreation arid e_ncased within one another, so that at but the first major work in this area was Reaumur'.s memoir ori
the appointed time each tiny preformed embryo could expand the regeneration of crayfish claws that appeared in 1712 (see
and develop, through mtchanicalmeans, into a full~fledged · Moeschlin .. Krieg 1953; Roger 1963:39o-g6). Reaumur drew
· organisni. Pteexisterice avoided ·the atheistic and materialistic the parallel bet~een regeneration and normal generation and
implications ,of development by epigenes~s; while ·also account- was critical of the notion that hidden. "germs" might be respon-
ing, for the source of animal organization. Embryos develop sible for the development c,f the ne~ appendages. Yet, Reaumur
· into the proper organisms because all of their parts were noted, aU other hypotheses seemed inconceivable as well (Roger
created at one dme and arranged in the proper fashion by God. 1963:392-94). . .. .
The impossibi~ity of mechanistic epigenesis was further en- The. most astonishing development in this area came with
hanced by the identification, commonly made, between mech- Abraham Trembley~s discovery in 1741 of a new animal, the
10 Matter, life, and gtneration
....... !#,:.:...;.....~~. .\~~:........~/:::.: ""
freshwater polyp. Trembley, who observed the common green
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hydra, 'initially assumed the organisrn was a plant. Vet the crea-
ture's ability to 'Contract when, stimulated and to "walk" by
attaching successively its, posterior and anterior ends to the
surface of an object raised doubts in,his mind~ Trembley'then
proceeded to cut a polyp in two, assuming that the two halves
might live if it was,a plant.' "Ho~ever," he admitted, " ... 1
eKpected to see tqe cut pdlyps die" (1744:26). '.To nis surprise,
not only did both halves live, but each grew into a new complete
animal. As Reaumur, to 'Yhom Trem,bley COJ!lmunicated h~
findings, ~xpressed this sentiment, "whenlsaw for the first time
two polyps form little by little from tqe one thit I had cu~intwo,
I could hardly believe niy eyes; and it is a fact that I am not '' )r'
accustomed to seeing after having seen it again' and agai~
hundreds and hundreds of times" (1734-42, 6:liv-lvr). Trembley
1-.
also observed the polyp's normal reproductive method, thatis, •I) )'(
by budding, a further indication o( the plantlike ptopetties of 5"' r·' .( .r.,.J'«-\,,. '
)
this unusual animal (see Figure 2; see also Baker 1952). ,,
Trembley's discovery. of the polyp's ability to multiply par
boutures, by artificial division, created a tremendous stir among
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eighteenth-century intellectuals. Reaumur, who·· anrioimced
Trembley's findings and demonstrated his experiments before /
~
.th~ Paris Academy of Sciences in 1741, remarked; "I haye, seen "
no one Who has believed this on the-first account he has heard of
it" (1734-42, 6:li). The official report of these sessions captures
some of the excitement and sense of wonder that ntusl have
been present:
l'he story of the Phoenix that is rebor11 from its ashes; wholly fabµfo~s
as it is, ?~fers nothing more marvelous than the discovery Qf which we
are about to speak. The chimerical ideas of the palingenesis or
regeneration of plants and animals, ·which some akhetnists . have•
thought possible by the assembly and rt:union of t~eir essenti~I parts, i
only te,nded to restore a plant or an ammal after 1t1 qestrucuon; the
serp~i;it cut,,in two, .and said to join toged1er again, only g::\ve one and,
the same sei;pent; but l:tere i~ nature going f;uther tl:tan our fancies.
, [Histoire de'tAcadimie Royalt des Sciences, 1744:33-34)
' ' . l \• ' • ' ' .·
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,1
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12 Mauer, life, and generation lntr()(iuction 1~
develop if an accident occurs. Others' saw in the polyp ,unmis- Maupertuis proposed that the semirial fluid in both the male
takable evidence for. epigenesis. · Finally, there were those who_ and female parents contains particles sent from each part of the
simply refused to acknowledge the facts. As Tren:ibley note~. in body. When these two fluids mix, the proper embryo results by
describing the reaction of the Royal Society to· his discoveries, a union of the particles from each part of the parents' bodies.
"The singular fact_s that are contained. in the history of these "If there are, in each of the seminal seeds," ~laupertuis ex.:.
small animals are· the admiration of a great many people: but plain~d, .. partides predetermined to form theiieart, the h~ad~
. several people have been .he$itant t<>,. admit tllem. T!iere are the entrails, the arms and the..Iegs, if these particular particles
those who ha~e even said that they ~ill.{lot believe_ it when they had a special attraction for those which are to be their im-
see them. Apparently these men ha.ve•some cherished system mediate neighbors in the animal body, this would lead ·to the
that they are afraid ofupsetting" (M. Trembley·1943:165-66). formation of the fetus" (p: 56). As an analogy for this formation
The episo<!e of the polyp illustrates well the close dependence of the embryo by attractive partides, Maupert!,liS suggested the
of eighteenth-century theories of generation Ori-philosophical, arbre ,de Diane, a. treelike figure that forms on the surface of
. metaphysical, and religious issues. The, polyp."1as instrumental water from the. mixture of silver, nitric acid, and mercury.
in the. conversion of $ome to.epig~rtesis (Bodemer 19'?4), and Other similar examples .of "organited" nonliving formati_oqs
became a central motjf in the materialist theories Of epigenesis were known to be abundant in chemical phenomena. Why may
proPounded by Diderot and La Mettrie (s~e Vartanian 1950). these not serve as examples of the way embryos form? .. Al-
Yet after creating such an initi~l stir, regeneration ih the polyp though these seem less highly organized than the bodies of most
and in other organisms was quickly subsumed by preformation- animals, might they not depend on the same mechanisms ~nd
ists like Charles Borinet under ad hoc explanations based on on similar laws?" Maupertuis asked (p. 55). ,
preexistent part germs. As Roger has not~d, it W<>uld seem "as if Maupertuis opposed preformation on the grounds thatit is
this system ... was more valuable than the facts the°!selves''. _ . no easier to explain how all organisms were formed at one tim.~
(1963:385). . , · in. the past than at each new instance of generation. "What has
14 Matter:, life, and.generation Introduction 15
natural science lost by the idea: that animals are formed succes- many different organs be placed and assembled in their proper
sively? For God, is there any reat difference between one order? We see with the ivostglaringevidence, that in order t<1arrive
1
moment in time and ,the ne;xt?" (p. 42). There were two classes at the formation of so,complicated a piece of work, it is not enough to
~have m4kiplied and varied the lawsofattraction at pleasure, and that
of evidence that Maupertuis felt made preexistence impossible. one must besides attribute the most compleat stock of knowledge to·.
The first was Harvey's bbservations on the gradual formation of · that attraction. [ 1749, 1750 trans.:463) ;,
em~ryos. Upon opening the .ut~ruses of does shortly after
mating, H~rvey· found .no tiny <;mbryo developing. Rather, .he Without guidance, how could the seminal particles become
found, no. change at all for some time arid t.heri only ~ gradual organized·into a devel~ping embryo? SQ.ch questions were·im ..
development. Second, Maupertuis felt that neither. ovist nor possible to answer on strictly mechanistic grounds, even when
animalculist preformatlon could be reconciled with the fact that ' attractive force~ were invoked. · ( · · '
offspring show resemblance to both parents. Hybrid organisms· Maupertuis's response was to move away from a rigidly
are an even rriore striking example of thisc. Thus, the embryo mechanical view of matter. Hinted at in a concluding .. Doubts
must be a composite' of ge~erative material'from both the and Questions" section of the Vinus physique and developed
mother and the father. Conseq~ently, Maupertuis concluded, more fully in his Systime de la nature (1751), Maupertuis's solu-
we should return _to the ancient notion, promoted also by tion was .to attribute to tpe smallest particles that make up living ·
D~scartes, that theembryo forms out of seminal fluids from organisms a capacity of intelligence and r:n,emory. By "remem-
both parents. bering" their former locations and by possessing an instinct to
Maup~rtuis,did not explain his theory of generation in.any unite, the seminal particles would be able to arrange themselves
, further detail, ·either with regard, to the seminal particles or the into ·an embryo in the proper fashion. "A uniform and blind
' attractive force. y ~t because of the phenomena of resemblance attraction,.'~ Mauper_tuis admitted in the Systime de la nature,
"3nd heredity! Maupertui~ felt thathis was the only viable alter- • "diffused in all t4e particles of matter, wou·ld not be.able to
native to preformation .. Maupertuis.'s · theory was not w_idely serve to explain how these particles arrange themselves to fonn
accepted, and it met with. much initial criticistn; Reaumur, in a, the· body whose o'l'ganization is lhe most simple... ;_ it is neces-
work devoted to techniques of incubation for raising domestic sary to have recourse to SQ.me principle of intelligence, to some-
chickens, attacked in particular Maupertuis's use·of the concept thing resembling w!iat we call desire, aversion, memory" ( 1751,
of attraction in· his explanation of generation. Rejecting first . 2: 146-47). . •, )
De.scattes's theory on the grounds that blind fermentation of In ·pi:oposing this solution to the sour:ce-of-organization di-
seminal material coul.d never be capable of forming a liv.ing lemma, Maupertuis rejected the traditional mechanistic view of
organism, Reaurriur brought similar argum·ents to bear against passive mattet and adopted instead a Leibnizian notion of·
Maupertuis (without mentioning the anonyrt:tous author by matter as fundamentally active (see Roger 1963:479). Adding a
· name).. It had become ,fashionable, Reaurriur charged, for psychical quality to dead particles of matter, Maupertuis was
occult qualities like attraction, which had:been banished_from able to answer the objections of those who, like Reaumul', saw
philos9phy, to be used once again in phy~i.cs and t9 lx(extcmded epigenesis by blind attraction as. an untenable explanation.
into other areas as well, even the formation of the fetus. Yet, · ~aupertuis's solutiori was to offer an alternative not only to
R.e;mmur countered, . preformation but also to the rigidly mechanistic philosophy on
which it 'fas based. · ·
how will attractiO~S be\able to give to SU.ch and SUfh a m~SS the form
and.structure of a heart, to a~other that ofll stomach; to a third one
that 'of an eye, and to another tha:t of ~near? H<>w will they frame Buffon
will
other masses into v~ssel~. valves, etc. [A]lltheir tendency amount
,. .
During these same years, another antipreformationist dieory of/
barely t9the reunion;ofthe similar parts intosolid)nasses:What law
of attra<;tion shall one imagine for the making of that small bone of generation appeared in the second volume of Buffon's Histoire
the ear, whose figure makes it to be .called the stirup? How shall so nature/le .(1749). Buffon's t_heory was based on a distinction
16 Mafter, life, o:nd generation . Introduction 17
admitting that we do·not knowiiow it i.s accomplished but also maintained, the internal mold is passed on to the ·offspring,
renouncing all desire to conceive of i_t'' (174g-89, 2:28). In- there becoming the basis for growth. When the organism ,.
, stead, Buffon r>rqposed that the.formation 0f tpe embrro takes matures, excess organic. particles are collected and through
place aftet the seminill fluids from. bQth parents mix. This them the internal mold is passed on again to new ·offspring,
process is accomplished by two .agents; the organi~m's moult completing the cycle of growth and reproduction.
intmeur ("internal mold") and the existence of spedal "pene- , In order to explain how-the inte.rnal mold accompHshes these
trating forces." The embryo forms immediately upon the mixing tasks, Buffon invoked·c~rtain ''penetrating forces,";analogous
of the fluids, then proceeding. to grow by the assimil"'tion of / to the forces that govern gravity, magnetic attraction, and
organic particles.through nutrition. 9 chemical affinity. "It is apparent ... ," Buffon claii:ned, "that
· Buftpn's concept of the internal mold was intended to be his forces exist in Nature, such as gravity, which relate to the
answerto both preexistence and 1nechanical epigen.esis. "lnthe · exterior..qualities of bodies, but which act .on the most intimate
same manner as we can make mo.ds by which we. give- to the parts and penetrate to all points:... for, in the same manner as
exterior bf bodies whatever figure we please," ~e explained, the·force of gravity penetrates the interior ofall matter, the
"suppose that Nature can make molds by which it gives [to force that pushes or attracts the organic parts of nourishment
' organic bodies] not only exterior figure but also internal form; penetrates into the interior of the organized body" (pp. 45-46).
· would this· not be a means by which reproduction could· be The penetrating forces cause the organic· particles absorbed
brougl:it about?" ( 174g,-:-89; 2: 34). Could we not suppose that . from food to be properly assimilated to the internal mold in
· ·there is some kind of organizational matrix that is responsible each p~rt of the organism's body.Just so, with regard to repro-
for the proper assembly of particles during both growth and· duction, "May not a force si!llilar to that which is necessary
reproduction? When an animal or plant reproduces, Buffon to •.• produce,growth, be sufficient tobring about the reunion
18 Matter, life, and gt_!&eration Introduction 19
~
of these organic particles, and indeed to assemble them into an t,hat are consta~tly balanced against o~e another ( 1750:22 1 n.)~-
organized form similar lo that of the body from which they are Through a continual tendency to expand, checked by a co-
'derived?" (p. 62). The penetrating force.~nsures that the repre- hesi~e force of re~istance, living matter was thought by Needham
sentative organic particles, and the internal rriold they car,ry, . to produce the observed phenomena ofvitality.
become properly assembled in the n~w: embryo. Needham's system had its greatest impact on his contempo-
Buffon's theory of generation shared several of the difficulties · raries in its support for spontaneous generation. Needham
encountered _by Maupertuis.• Both saw the seminal fluids as claimed that he had found animalcules spontaneously forming
as
being made up of particles acting -representatives. from -the. in boiled meat infusoria. Decried as impious and materialist by
-parts of the parents' bodies. Yet neither fully explained in what some, Needham's views led, others, notably Lazzaro Spallan~ni,
this. similarity actually consists. Buffon's internal mold was to attempt experimental ref~tation. Spallanzani, himself a pre-
intended to acc:ount ·for this, yet he never disclosed exactly what formationist, believed his work proved that Needham was in
the internal mold is. Both Maupettuis and, Buffon invoked error, and that ·the appearance of animalcules in infusoria was
attractive forces to guide the formation of the embryo ~u-t of the the. result of Needham's having used 'insufficient sterilization
seminal fluid, yet both left themselves open to Reaumur's techniques (Spallanzani 1_765). 11 Needf1.am responded in 1769
. charge that blind attractiqn could never accomplish this task by publishing an annotated French version of Spallanzani's
. (although, as we saw, Maupertuis attempt~d to solve .this di{fi-· book in which Needham claimed that the excessive heat used by
culty through "intelligent" particles). It is significant, with re- Spal.lanzani had destroyed( the vegetative force· and had im-
gard to the!r utilizatio~ ofattractive forces, that ?°th Maupertuis paired the elasticity of the air within the flasks, thus rendering
~nd Buffon were early supporters of Newton..,on ,the Conti- them incapable of producing animalcules. "What confusion,
nent.10 The freedom that-Newtonian mechamsm offered to· what obscurity prevail~ in his notes to my _microscopical ob-
biology - the addition of force· to rnatter. ana motion as the . servations! What monstrosities in ,his thoughts!" Spallanzani
fundamental categories of explanation -. played an important compla1ned to Bonnet (cited in Dolman 1975:556); Resolved to
role in. embryology as well as in other areas of physiology. · silence Needham once and for all, Spallanzani devoted himself
to further experiments, producing. his masterful Opuscoli di,
fisica animale e vegetabile ( 1776). His work was warmly received
·Needham by Bonnet and othetpreformationists, who uniformly opposed
A third theory of generation.contemporaneous.with those of spontaneous generation. ·
Maupertuis and Buffon was put forward byJohn Turberville The three systems of· Maupertuis, Buffon, and Needham
Needham. Following his laboratory collaboration with Buffon, represented a <::ombined challenge to preexistence theories at
Needham·· proposed his own explanation of generation in a midcentury. The"similarities among their vie\ts·may be due in
· paper published in the Philosophical/Transaction$ of the Royal part to the fact that they were in close comrpunication during
Society for the year" 1748 and in·an expanded French version,. the late )74os. Needham and Buffon collaborated in some of
which appeared in 1750' (Nouvelle$ Qbsetvations microscopiques). their experiments; and,. accor,cling to Needham (1748:633),
.Needharn postulated that th~re exists a univ~rsal semen from ·Buffon and· Maupertuis often discussed the subject of genera- -
whichall getieration·occurs. At the lowest leve.l are formed the tion following the publication of the Vinw phys'ique (1745). Yet
spermatic animalcules and the animalcules ·~ound in infusoria the similarity in these theQries also represents a common dis,;,
· and in pond water. (Contrary to Btiffon, Needham classified all satisfaction with the notion of preexistence, with its inability to
of these as living organisms.) The constant combination, sep- deal with the phenomena of resemblance and hybrids, and,
aration, and recombination of organic matter are all produced, more especially, with the seeming absurdity of the idea of erriboite-
Needham claimed, by a universal vegetative force, the source of ment. Furthermore, all three turned to the concept of attractive
iUHfe activity. This veietative force may further be broken forces to. res?scitate efi~en<:sis following its overly mechanistic
dowOinto two forces, one of expansion and one of resistance, formulation m the fermentatmn models of Pescartes and otheri;.
20 Matter, life, and generation
· . Ha~le_r) third ~~jor objection .to B1uffort's the?ry ~as the structions [salts, snowflakes, etc.] to the simplest animals, which
most important with regard to Hailer's ?Wn changmg views on in all their parts are a similar and uniform jelly? ... Where does
generation, for· it raised the .whole issue of how forces. are the power of universal laws stop? Where is the boundary, within
involved in embryological development. Haller was unable to . which they form, and outside of which.they can build no more?"
see how Buffon's moule interieu,r could possibly organize all the .( 1752a:vi). Hailer's answer is that it is God who directs and
org:inic particles int<> a coll!plete organism. Even if we assume limits the operation of univer~l laws and forces. Matter alone is
that all the particles sent from all the parts of the body are in the powerless, yet when given force~ by God it is capable of pro-
seminal fluid, how are they put together? ·.. M. Buffon has shown ducing all of the natural phenomena we observe . ."If matter has
no cause," Haller claims, "thatbrings them in order, that unites forces that allow it to build thing~, it does not possess them
the eye particles of. the father _with the eye particles of the blindly. They are surrounded by eternal limits, and build always
mother; ... There is missing a building master, who lays down. perfectly not mechanical equals but something si~ilar, some-
30 Mauer,· life, and generation Haller's views on em.bryology 31
' ~hi{tg that is prescri~d in an inviolable pla~: but with·~ diversity always such as ,was the first carnation ..(175-ib:87). God is the.
that excludes the constraint of blindly working matter'' (p. xv). mastermind of the developmental process, not matter alone.
There are two key ai:gumentsinvolv:ed in this last passage that Hailer's conclusion on this)issue of the relative roles of forces
Haller used .to prove that forces alone could not govern the and God in,development is simply that ''The invariable produc~
world. First, Haller. believed that matter possesses no abilities tion of always similar, always qivin.ely constructed animals,.
whats~ver on its. own, without these having been designated.by appeai:5 to be too great for the simple forces that produce ... a_~
God. As he explains in bis review of Buffon's theory, "Matter salt crystal" ( 1752a:xvi)~ Th·e boundary between the prQduction
does not possess these abilities itself. It could be without gravity, of simple structures like crystals and the formation oniving
without elasticity,. without irritability; added qualities, but es- .· ·organism.s is drawn by God; Yet is this not the exact opposite of
sential to the structure of plants and animals. These qualities do what Hall~r had argued only a few years earlier in support of his
not take part in its essence; they are foreign to it; they are not theory of epigenesis based on attractive fqrces? There he had
common to aU parts of matter.... A first callse has thus allotted likened the laws. governing development to those mrough
to different classes of matter abilities and forces· calculated which· crystals and metals are formed. Now he maintains that
according to a general plan,.and it is there that we recognize .the development is too much for these simple forces of attraction to
hand of the Creator" (1751b:86). Matter can do absolutely be able to accomplish by themselves. ·
noth~g without the forces that Cod giv~s to it. Thus, that forces The central issue · in all of these discussions of forces is
of matter produce natural phenomena does ·not threaten the guidance. Haller never que~tioned the idea that simple forces of 1
existence of a Creator. .matter are active in development. His concern was with how they
Hailer.'s second argument rests !)n the- fact that both variety act to produce their effe_cts, sin<:e he did not believe thai: they
and ,constancy of form ex:ist among living organisms. If matter · could atcom plish the task of development unaideci. God, Haller
formed living creatures blindly, why would species be preserved? believed, direc~ and limits the operation of fon:es in producing-
"If a mere attractive and repulsive force in the semen built a the developing embryo. Yet how this guidance is manifested,
man or a deer; if this occurred by chance, why does th~re never that is, what the mechanism .is through which God controls
develop from this .. '. matter ... instead -of a man, a mon"k.ey, development, is not made clear by Haller in _his discussions of
which has so many similarities with man?" ( 1752a:xv).'13 Hailer's Buffon's theory~ J
answer again is that it is God, ·not matter, that is ultimately Hailers ideas on generation had clearly altered since his
responsible for the formation of organisms. A great deal of earlier statements in support of epigenesis. No longer coµld he
variety. exists in specific structures within living creatures. Re- postulate that the attractive forces of matter by themselves
production doe~ not produce an exact duplicate but rather an produce fibers, membranes, and organs from an originally
organism that is similar in its essential details. "'this invariable unorganized liquid. He was n<?t ye)t a preformationist, y~t·he
constancy of species, which permits a slight variety but, which was clearly at this time in a transitional stage, moving away from
never deviates ih these essential characters, appears to me to be I ~is earlier epigenetic views, beginning to question the relation-_
one of the most evident. proofs of the hand of' the Creator" ship between forces of matter and development. On the one
( 1751b:87.). Conside·r, Haller suggests, the case of a carnation. hand, he still believed that development occurs by means of ,,I.
The matter out of which it is formed may produce a carnation forces of matter; on the other hand, he was reluctant to allow
of a different color or even of a different ~tructure, but it never · that these forces alone could· be responsible for generation.
produces a tulip. "It is God who permits a double carnation tQ God!s wisdom must somehow direct development. But how
be ·made, different from the crimson carnation native to the forces of matter are guided by God and exactly how they operate
lakes of the Alps, but always .a carnation, which in a thousand on the material of the embryo was still not clear to Haller.
generations will never produce anything but a seed of a carna- Haller dispensed with the views of Needham even more
tion. All is not chance, otherwise the carnation would become a cursorily in his discussions of Buffon's work on generation. ·
tulip: all is not necessity~ otherwise the carnation would remain Needham's vegetative force was just tqe kind of force Haller
I•
32 Matter, life, and generation· Hall;er's vuws on embryology 33
could not allow. "And yet," he declared, "I am hot troubled by ;f·. .recently died, the·heart must beat from some "unknown cause"
the very remote proof of these building forces. Truth is. like .a ~ that "lies hidden· in the fabric of the heart itself" (1739-44,
corr~ct calculation, e!5tablished on all sides; everything must · ',J,
._,,_i_'
. 'b
2[ 1739]: 129, n.i). Although Haller made simil;1,r remarks in the
help support its structure; only etror falls apart as soon as one ',~ fou'rth volume of the Praelectiones (4( 1743]:586, n. a) and in his
takes away its only support, because all else conflicts against it" Prima,: lineae pkysiologiae (1747:51; 1751c:252), his first major
1
(1752a:xiv). Similarly, he-r~marked at the conclusion of his exposition of the theory of i,rritability and sensibility ~as his
preface, "We can thus wait. tranquilly to see if the experiments famous paper, "De partibus corporis humani sensilibµs e~ ir-
· of the learned will confirm or contradict the vegetative and .:ritabilibus," which was presented in 1752 to the Royal Society of
animating forces of M. Needham. They will lead us always Sciences of Gottingen and published in its journal the followjng
nearer to the truth, and this to God" (p. _xvi). By the time· • year. Here he argued that irritability, the capacity of muscles to
Hailer's preface was reprinted in his Samrnlung ltleiner Haller~ . contract,, is completely separate from sensibility, that is, sensa-
ischer Schriften (2nd edition, r772b), Haller was able to add tion. Haller ascribed sensibility to the nef\l"es and irritabilitY. to
footnotes referring to Spallanzani's experimental refutation in the muscles, which possess an inherent force that causes them to
1765 of Needham's views on spontaneous generation (see Chap- _contract when stimulated. Irritability, called in later publica-
ter 1 ). And in i 767, when Haller included a Latin version of this tions the vis insita, 1s "originates.in the very fabric of the irritable I
. preface in his Opera minora, aH references to Needham's views part" _(1752b:i34; see also Haller_ 1936). Both the existence of ,I
'!
we~ simply deleted.'" · involuntary motions in live animals that are entirely indepen-
Haller rejected both Needh~m·s vegetative force and his dent of the will, and the contractions that occur in muscle tissue ·
promotion of the idea that living organism~ could spontaneously in a dead animal or_in muscles separated from the body, point to
'generate from unorganized material. As we· shall see in later· . the independent property of irritability in muscles. "What
chapters, the issue of spontaneous generation was intimately therefore should hinder us," Haller proclaimed, "from grant-
tied for Haller to the larger question of the nature of generation ing irritability fo be that property of the animal gluten in the
in all organisms, and it played a role in his rejection of Wolff'.s muscular fiber, such that upon being touched and provoked it
theory of epigenesis. It may be that the parallel, drawn explicitly contracts, to whjcb moreover it is unnecessary to assign any
· by Needham, between epigenetic development in higher organ- - cause_, just as no probable cause of. attraction or gravity is
isms_ and the spontaneous genera_tion of lower forms of life assign,ed to matter [in general]. It i$ a physical cause, hidden in
contributed to Hailer's growing unease with the theory of epi- the intimate fabric, and d~vered through experirµents, which
genesis. Spontaneous generation,~ if true, would certainly call are evidence enough for. demonstrating its existence, [but]
into question the::. role of God in the creation of new organisms. which are too coarse to investigate further its cause in the
And the formation of higher org~nisms, as described -in· both• fabric" ( 1752b: 154). Irritability is a force inherent in. a particu-
Needham's and Buffon;s theories, bore.a dangetou~ resem- lar type of matter (a~imal muscle tissue) that operates auto-
.
blance.
to the 'formation of animalcules in Needham's.. infusoria.
'
matically under proper conditions of stimulation. Lixe gravity,
irritability can be deduced. from phenomena, yet its ultimate
cause lies, for the time being at least, beyond the reach of
Force and the concept of irritability
experiment and observation.
During these same years (late 1740s to early 1750s), 11aller was During the 1750s. Haller further developed his ideas on the
developing his ideas on another force concept, that of irritabil- nature of irritability, particularly in relation to the theories of
ity. Hailer's first reference to the notion of irritability may be Robert Whytt, who -believed that muscular contraction is based
found in a footnote in his edition of Boerhaave's lectures, on the soul, and in relation to the ,materialist position of La
Pra,:lectiones academicae. Here, commenting·on Boerhaave's de- Mettrie. From these various ~ontroversies, there emerge, .a . - !
scription of the systolic motion of the heart, Haller remarked picture of Haller attempting t!) steer a middle course between
that since movement persists in hearts in c1niinals that have animism on the one hand and materialism on the other. 16 ,
Maller, life, and generation Haller's views on embryowgy 35
34
Haller believed that irritability is a force of m;tter not based on
the soul or any other immaterial force, yet he was al~ con-
cerned to show that matter could not on its own· possess the
power of irritability. This ability is given to matter by God: "I
piously acknowledge that God is the mover of all nature. For
this rea~on, ·neither the elasticity of expanding air, nor the
', weight of a stone, nor the effervescence of acids mixed with
alkalis; nor the contraction of a dissected muscle ough"t to be
attributed to incorporeal forces. God gave to bodies an attrac-
tive force and other forces, which once received are· exercised''
( 1757-66, 7:xii). God is the source of irritability, gravity, arid
the other forces that matter possesses.
Haller's ideas on the nature of irritability are important in
two ways for our understanding of the development of his views
on. embryology. First, they offer us an example of Hall~r's
philosophical beliefs about biology and about how forces operate .
in biological ph~nomena.·Forces are not inhe11ent in .matter but
are given to matter by God. They operate on a mechanical basis,
however, once bestowed upon matter, automatically acting
under specified conditions. ·
This view of forces is not inconsistent with the kind of epi-
genesis Haller had suggested in 1747, when he claimed that
development proceeds via attractive forc~s and "divine laws."
Yet, as both Roger ( 1g63:709:-10) and Gasking ( 1g67: 114) have
pointed out; the kind of epigenesis Haller could accept, based
. only on mechanical forces, was simply .not viable., How could
such forces account for the organization of the embryo? What
was to serve as Haller;s "building master"? In his statements
su pporti.ng epigenesis and in his discussions of Buffon's theory,'
Haller turned t9 God as the source of this ·guidance. Yet no-
wqere does he ouuine hou, God directs the developmental
process~ how the automatically acting mechanical forces in mat.;.
ter are marshaled to produce the proper offspring. ·
Haller's work on irritability and his reflections on the implica-
tions of the Buffonian system of generation forced him to come
face to face ·with the .problem of .reconciling his belief in the
world as a Divine Creation with mechanical explanation. Jn
criticizing Buffon's theory, Haller began to realize that simple
forces of attraction were not sufficient to account for the de..
velopmental process. But what forces would be sufficient? How
Figure 5. Re~arching irritability and sensibility. (From the frontispiece to Haller, can one: explain development on a mechanical basis and still
MefflOires sur la nature sensible et irritable des parties du corps animal, 1756-6o; courtesy of
the Francis A. Countway Library of Medicine, Harvard Medical School) retain a place for the guidance of God?
36 Matter, life, and generation
Haller first mentioned the problem of generation in a letter ·. Haller began his first series of observations on ipcubated
. ~ritten shortly after the instigation of their correspondence in chicken eggs in August 1755. Two:major subjects occupied his
1754, where he remarked that he int~nded to begin wotking on · att~ntion: the.formation of bone, which he was to investigate all
the subject in the near future.· To ·this Bonnet re~ponded, along in his chick obseiyations and t~ which he devote9 ·a
"When you occupy. yourself, Monsieur, with the great mystery separate publication ( 1758b); and second, the formation of the
of generation, the eyes of all physicians will be upon you" heart and its relationship to the lungs. Haller had repeatedly ·
(Haller MSS, 28 September 1754). Asking in the same letter used Malpighi's observation that the_ he.art is originally only a
what Haller~s views on the question were, Bonnet, received a simple tµbe that later deyelops chambers as an argument in
reply that indicates Haller was still thinking along epigenetic · favor.of epigenesis. Yet in Septembe.r,_Haller·wrote to Bonnet,
lines: "I have no system of ge~eration. Without prejudice .... have made many efforts to pe~etrate the mechanism of the
against evolution [preformation], I see more and more a sim- . heart.... I believe that. I have caught a glimpse of that which
ple, sticky material forming and shapfog itselflittle by little. The does not change in its structure and that it is always equally·
egg of a sheep is for a long time all only a visc~us fluid; it formed of two auricl~. and two ventricles.... I spend two or .
remains in this state 17 or 18 days. But it is necessary to repeat three hours everyday reexamining this marvelous heart to give
these experiments ... " (Bonnet MSS, 14 October 1754). 19 me an exact idea, which the vivacity of its movements renders
Hailer's own experiments were nott6 begin until the:follow- difficult" (Bonnet MSS, 6 September 1755).
ing summer, and th~n they were on chickens rather thari sheep. Haller!s second series of observations, made during the late
During the ir:itervening winter months, Haller and Bonnet sum~er and early fall of 1756, cleared up the question of the
considered the question of the original fluid state of the em- development of the heart and lungs.. As Haller. reported to
bryo. As Haller remarked, still in relation to his experiments Bonnet, "For the heart, all the mystery that I investigateq is,
with.;Kuhlemann, "For the matter of gen~ration, I believe that· redm;ed to a very simple thing. The lung, with its al'.tery a~d its
one part is clear.... A viscosity ve·ry surely successively congeals vein, is very small and invisible in ,the first _times. Little by lit~le it
and becomes th~ membranes uf the fetus .... [Yet] All· is trans- grows, and its artery which through its fineness and_its'trans-
parent and invisible" (Boµnet MSS, 26 November 1754). In his parent liquid was hidden from our eyes, becomes asecond very
next letter, Haller hinted at how the transparency dilemma considerable branch of the aorta from which two small branches
might be resolved: "I am strongly for your opinion, Monsieur, lead to the'lring, etc." (Bonnet MSS; 7 December 1756). Haller
about the organic glue. 20 A salt dissolved in water retains, in- had been unable to reconcile the·formation of the pulmonary
visible qabes~ And there would. be a jump if the small animal, circulation with !dalpighi's notion of an originally tubular heart
becoming visible through the help of spirit of wine at the 18th that curls up on itself to form the four-chambered heart, for the
day, already all formed, had.been _fluid the dcly before. It ex- vessels of the lungs would have to break in at some point in this
isted, without ~ ~oubt, approximately the· same; but its trans- . linear arrangement. Yet Haller solved this difficulty by propos~
. parency hid it from us" (Bonnet MSS, 4January,1755). ing that, in early stages of development, the lungs and their
As an epigenesist, H;aller had used the original fluid state of vessels exist in an invisible state, connected to the heart as they
the embryo as an argument against preformation and in favor will ·be in their later·appearance. The heart itself develops from
of gradual development. Yet now he suggested that transpar- four originally existing but not clearly visible chambers. All
ency may hide structures that actually exist in the viscous liquor. simply become visible durin-g development, with no fundamental
Although l,ie did not develop the. idea fur.ther at' this point, it changes taking place. ' . . .
should be noted that Haller later made wholesale use of the · Hailer's final series of observations were made during the
argument that transparency hides existing structures in his sprin~ and summer of J.757· It was during this period that
campaign for preformation. It was also later to become an issue _ Haller first announced his conversion to the theory of prefor-
of controversy in his debate with Wolff. ination. To Ignazio Somis, an Italian physician with whom
. - '
40 Matter, life, and gentration Hailer's views on embryology 41
Haller frequently corresponded, he declared in July, ··Hind that 175 7. These were publishe9 the following year in a two-volume
all the changes of the egg relate m~ch more to the system of work, Sur la f orrnati!m du coeur dans le poulet. The basic tene~ of
. evolution.than to that of epigei:tesis" (Hirttzsche 1965:40). 21 The Hailer's new theory was that all. essential structures of ~e em-
final piece of evidence emerged wh~n, having finished his obser- bryo-exist first in. the female egg. When conception occurs, the
vations on the heart, Haller began to turn his attention to the yolk heart. is stimulated and, because ·of its inherent irritability,
and its· relationship to the embryo. And in doing so, Hallet · begins to beat. 22 The beating heart sends out fluids through the
discovered what he thought was a definitive proof of preexist- · fold~d parts of the transparent embryo; the embryo's struc-
ence. This he announced to Bonnet on 1 September 1757: tures begin to solidify and to becotne more opaque; and the '
. '
visible fetus emerges. "All of the parts of the animal body,"
The ~embranes of the yolk are in a larger form the-me~branes of the Haller explains, "are born from fluid, apparently organized,
intestines, but more developed. The yolk itself is only an expansion of
· the small intestine of the chick. It is from this that one can draw a very which becomes mucous, and which acquires little by little more
plausible reason to give the female the true beginnings of the fetus; definite boundaries and a consistency that resists pressure"'
for, in short, the yolk exists before the approach of the male; its (1758a, 2:175). When fluids of increasing opacity, containing
membranes are continuous with those of the. intestines of the chick; more and more visc<;ms and solid particles, are pumped by the
the chick thus appears to have existed before this approach. [Bonnet heart through the vessels, the· parts of the ·embryo become
MSS] , increasingly visible as development proceeds. "The man~er by
Hailer's "membrane-continuity" proof rested on the observa- which these same parts become visible from being invisible is of
tion that, in later stages of development, the embryo's intestines the most grand simplicity. It is the effect of growth, ·but even
are connected to the yolk sac in:sµch a maner that the yqlk sac mor~ 1 the simple effect of opacity" .(p. 176). As Haller sum-
seems to be simply a contimiation of the intestines (see Figure marizes: , ,
12, Chapter 3). Haller thought he had found the membranes of It appears very probable to me that the essential parts of the fetus
the yolk sac in the unfertilized egg; and he inferred from this, exist formed at all ~imes; not ,it is true in the way that they appear in
· on logical grounds, that the embryo must exist at this time also, · the adult animal: they are arranged in such a Cash.ion ttl~t certain
even though it c~nnot yet be seen. Consequently, the embryo prepared causes, hastening the growth of some of these parts, im:-
exists in the fem ale before fertilization by the m:,de. peding that of.others, changing positions, rendering visible organs
To I-faller's announcement of his conversion and his new that were formerly.diaphanous, giving consistency to the fluidity and
proof of preexistence, ~onnet~ himself a staunch preforma-· to the mucosity, form in the end art animal that is very different from
the embryo, and yet in which there is no part that did not exist
tionist;jubilantly re.sponded: · . · · )
essentially in the embryo. It is thus that I explain development. [p.
It has been disputed for a century whether the embryo comes origi- ' 186] ' '
C
nally from the male or.the female .... Monsieur de Haller opens an
· egg of the chicken, he examines it with eyes that have come to see, and All organs and structu~es of the adult organism exist in the tiny,
he discovers that the yolk is an expansion of the smc1.ll intestine of the transparent embryo and are made apparent by mechanical
chicken. The embryo thus existed.in the female before the approach causes. 23 · '
of die male. This discovery is assuredly one of the most importantthat Hailer's theory of preformation provided solutio~s to the
anyone has made in the obsetire matter of generation. [Haller MSS, 7 difficulties ·he had previously, raised when he had supported
September 1757] epigenesis. With regard to the initial fluid state of the embryo, ·
Later, after Haller had published these observations, Bonnet Haller now saw no .problem for p'reformation. Structures. do
remarked simply, "Your chickens enchant me. I had not hoped exist in this initial viscous liquor (recall the dissolved sa,lt anal-
that the secret of generation would be discovered so soon" ogy), eveh though they cannot be seen. Furthermore, HalJer
(Haller MSS, 300ctober 1758). · had found that, by using alcohol or vinegar, he could cause _
· Haller read a report of his experiments before the Royal structures to become visible before their normal time of appear-
~ociety of Scienc~s of Gottingen in September and December ance (although he also found that there was a point prior to
j
which this technique did not work). "After these observations," existentgerms ( 1757-66, 8( 1766]: 170-71). When the organism
Haller remarks, "one should be on guard against the tempta- is injured, these germs can be called into play. Never expanding
tion to pronounce. that such and such a part of an animal is on the nature of these preexistent "head germs" and ,"tail
newly produced and that it has not existed befo~. It could have .germs," Haller argued in direct contradiction to his earlier
been too small; it could have only been transparent" ( 1758a, views that regeneration is not a case of gradual, epigenetic
2: 177). Appearances should not deceive us into thinking that, development. Some organisms possess. preexistent ger!11s and
gradual development is actually occurring. · , · are thereby capable of formhig a lost partthroug1l the develop-
Haller was also now able to sotve the problem of changing ment of these germs. Neither Halter's explanati~n for regen-
structu.res in the embryo. As an epigenesist, he had pointed to eration nor that for hyb,rids was very satisfactory, yet he .no
the alterations one observes· in" major organs, especially the . longer saw these two areas of objection as fatal ~o the theory of
heart, as supporting gradual development. Yet through.his own preformation. ·
observations, Haller was able to discount Malpighi's description . Haller's ~ew explanation of embryological development
of the formation of the heart, for Haller believed he could show_ , solved entirely the dilemma concerning the operation of forces
that all of the changes one observes are only apparent. This that h.ad arisen in his critiques of Buffon's theory. As we can
conc4-tsiop he broadened to apply to the whole embryo, and.he recall, i.n dealing w.ith Buffon's ideas on generation, Haller
postulated that differential rates of growth and increasing came face to face with the problem of how development is
opacity could account for seemingly drastic alterations in struc- organized and directed. Because of his own beliefs, Haller
ture. As Haller later remarked, "If I myself, more than twenty could_11ot accept either self-directed material forces or immate,.
years ~go, before my repeated observations on eggs and on rial, vital forces. He was thus left with forces of a material but
pregnant quadrupeds, once used the argu.ment that the fetus is simple nature, operating mechanically, in the sameway that the
· very different from the fully grown animal and therefore that force responsible for irritability operates'. Yet these forces could
ther,e are not found in the rudiments of the new animal tho~ not, on'their own, produce development without being directed
parts that are in"the same, now fully grown ani~al, I recognize in their operations by Go<J. It was this key issue of guidance,
manifestly now that all the very same arguments that I brought which Haller had left unexplained ii! .his epigenetic theory, that
forih against. evolution·[preformation] were in support of it" reopened the problem of generation fo~ him and prompted
(1757-66, 8[1766]:148-49). The changes one observes in the first the experiments on sheep with Kuhlemann and then.the.
embryo's strtictur~ are not evidence for gradual foi:mation but observations ·on incupated chicken eggs. ·
rather for development from preexisting parts. ·· . Hailer's theory of ovist prefonnati~n provided a clear mech-
Hailer's other two previous objections to preformation - the anism for the operation ·of forces in development and for God's
problems of ·hybrid organisms and regeneration - were now . guidance. All forces that are part of the develppniental process
accounted for as well. ·Hybrids resemble the male p~rent, Haller . are ~imple and mechanical in nature; they possess no self- -1I
proposed, bec.ause the male semen can cause greater growth by directing abilities whatsoever. Their utilization and guidance is i
speeding up the. movement of fluids· in. certain parts of the based ultimately on God who, at the Creation, constructed all of ·
embryo (1758a, 2: 189-"91). Hence, for instance, the long ears of the preformed organs of each future embryo and, ~n addition,
the mule .~nd other unusual structures found in hybrid organ~ organized in these structures all of the forces that would be
isms. Yet exactly how the male semen causes this increased active in the embryo's later' development. As Haller explab1ed
movement of fl.u1ds in specific embryonic parts is never clearly ,iri his Elementa physiologiae:
articulated by Haller. · · If the first rudiment of the f et.us is in the mother, if it has been built in
Haller explained regeneration in a similar ad hoc manner. In the egg, and has been completed to such a point that it needs only to
his Elementa physiologiae, he adopted the position, a~gued by receive ·nouris,hmen~ to grow from this, the greatest difficulty in
Bohnet among others, that organisms such as the polyp and the building this most· artistic structure from brute matter is solved. tn
crayfish _can regenerate lost parts because they contain pre- this hypothesis, the Creator himself, for whom nothing is difficult,
44 Matter, life, and generation
has built this structure: He has arranged at one time, or at least before
the male force [of fecundation] approaches, the brute matter accord-
ing to foreseen ends and according to a model preformed by his
Wisdom. [1757-66,8(1766): 143]
When the tiny embryo is stimulated by the male semen, the
3
heart, because of its inherent irritability, begins to beat, initiat-
The embryological debate
ing the entire preprogrammed sequence of development. Every-
thing takes place by natural, material forces, yet all is ultimately
under the guidance of God.
Hailer's observations on incubated chicken eggs provided
two major sources of evidence for his preformation theory. Within a year of the publication of Hailer's Sur la formation du
First, he thought he could now show that the heart does not coeur dans le poulet there appeared a work by a young Berlin
develop as Malpighi believed, but rather in a manner entirely physician,.Caspar Friedrich Wolff, that strongly supported epi-
consistent with preformation. Second, Haller felt that he could genesis. Wolff sent a copy of his Theoria generationis, which had
actually prove that the embryo exists in the unfertilized egg been his doctoral· dissertation at _the University of Halle, to
• through his membrane-continuity proof. 24 Hailer's observations Haller, in the hope that Haller would give Lip his new defense of
on chicken eggs combined to provide the catalyst for his conver- preformation. Little did Wolff suspect that his actions were to
sion, for they enabled him to work out a viable theory of em- provoke a ten-year controversy with Haller, who, rather than
bryological development based on preformation. Yet it is also being convinced by Wolff's work, became preformation's
clear that the need to develop such a theory, one that would be staunchest and most rigid defender. In this and the next chap-
consistent with his fundamental religious and mechanistic be- ter, I shall analyze the debate that ensued between Haller and
liefs, prompted Haller to begin these observations and in this Wolff. This chapter will concentrate on the embryological
sense prepared him for the discoveries that he then made. The aspects of the debate, while Chapter 4 will focus on the under-
period between the late 1740s and the observations on chicken lying philosophical themes of controversy. By way of introduc-
eggs in 1755 to 1757 witnessed Hailer's realization of what the tion, let me present here an overview of the major events of the
· implications and problems of epigenesis were, and his growing debate. (The reader is referred also to Appendix A, "Chronol-
uneasiness with how these difficulties could be resolved in a ogy of the Haller-Wolff Debate.")
manner consistent with his own philosophical beliefs. 25 These The controversy between Haller and Wolff was manifested in
factors combined to set the stage for Hailer's observations and, a series of publications and in several letters that passed be-
through them, to produce his con.version to preformation. - tween the two. 1 Additionally, Haller published reviews of each
of Wolff's works in the Gottingische Ameigen von gelehrten Sachen.
It was in response to the first of these reviews, in which Haller
criticized Wolff's dissertation, that Wolff published his second
book, Theorie von der Generation, in 1764. Not a translation of the
Theoria generationis, as some historians have surmised, the Theorie
von der Generation contains rather a restatement of Wolff's
theory and a polemical attack on the preformationist views of
both Haller and Bonnet.
Haller published a brief review of this work, but his major
response appeared in the eighth volume ( 1766) ofhisE/ementa
physiologiae corporis humani, which also contains a more detailed
account of his own theory of preformation.. During the next
two years, there appeared Wolff's "De formatione intestinorum"
Matter, life, and generation The embryological debate 47
46
in the journal of the St. Petersburg Academy of Sciences ( 1766---
67, 1768). This work was devoted· to demonstrating the epi-
genetic development of the intestinal tract and was perhaps the
. most influential of all of Wolff's publications, at least with regard
to his nineteenth-century successors in the field of embryology.
Wolff moved from Berlin to St. Petersburg in 1767. There-
after, little communication took place between Haller and Wolff,
and their debate effectively came to an end. Haller died in 1 777,
before the publication of Wolff's last major embryological work,
Von der eigenthiimlichen und wesentlichen Kraft ( 1789), which con-
tains both a further explanation of Wolff's theory and some
reflections on Wolff's debate with Haller. Wolff's own death
came in 1794, before he was able to complete his last major work,
on the subject of monstrous births.
Following an analysis of Wolff's theory of embryological devel-
opment, this chapter will be organized around the principal
themes of controversy that emerged in the Haller-Wolff debate.
These embryological points of contention fall into three major
areas: the formation of blood vessels in .the area vasculosa, the
· development of the heart; and Hailer's membrane-continuity
proo'f of preformation. I shall attempt both to describe the -
exchanges between Haller and Wolff on these issues and to
discuss their controversies in more modem embryological terms.
Finally, Wolff's work on.the formation of the intestines will be
presented, followed by some concluding remarks on embryo-
logical observation and on the debate as a whole.
plants and in animals" (p. 115, § 242). There is no need to The development of the vitelline vessels in the area vasculosa
assume the existence of preformed parts in the embryo; every- ex\Iibits striking changes as incubation continues. In early
thing is formed gradually, through the secretion and solidifica- stages of development, the area vasculosa presents a mottled
tion of fluids under the guidance of the essential force. appearance, due to the formation of blood islands by the end of
Wolff's theory was markedly different from Hailer's. Well- the first day. These begin to form at the periphery of the area
argued and based on observational evidence, Wolff's epigenetic and later extend into regions closer to the embryo. After about
explanation of development understandably presented a clear 33 hours of incubation, the area vasculosa begins to look more
challenge to Haller, who had not long before rejected his own netlike because '?f the .extension and joining together of blood
epigenetic beliefs in favor of preformation. Let me turn now to islands to form blood vessel channels. Within the newly forming
the major issues of controversy that emerged in their ensuing vessels, primitive blood corpuscles begin to appear, and the
debate. rudiments of the sinus terminalis become visible. This network
continues to develop, progressing inward toward the embryo to
join with the major vein and artery (the omphalomesenteric
THE EMBRYOLOGICAL DEBATE vessels) that connect the embryo's heart to the vitelline circula-
tory arc through the yolk stalk. By abouqo hours of incubation,
The area vasculosa the vitelline blood channels are completed, and the heart devel-
Haller and Wolff devoted more time to debating the formation ops a sufficiently strong heartbeat for circulation to commence.
of blood vessels in the area vasculosa than to any other single As mentioned earlier, the formation of blood vessels in the
issue. The gradual appearance of this vascular network is in- area vasculosa is one of the most visible aspects of development
deed a striking event in chick development, one noticed by the in the early days of incubation. Noted by Aristotle, Fabricius,
earliest investigators. These blood vessels, known today as the Harvey, and others, the stages of development of the area
vitelline circulatory arc, serve to bring nourishment from the vasculosa were first most completely described by Malpighi.5
yolk to the embryo. They form in the area vasculosa, a region of Beginning with an embryo of 12 hours' incubation, Malpighi
the blastoderm surrounding the developing embryo, and eventu- described the appearance of what he called the umbilical ves-
ally cover a sizable portion of the yolk's surface, bounded on the sels, offering a remarkably accurate account in gross detail (see
outer edge by a circular blood vessel called the sinus terminalis. Figures 1 and 9). Malpighi never clearly separated the vitelline
In the developing chick embryo, three separate circulatory from the allantoic circulation, nor did he fully understand the
arcs are formed, serving different functions. The embryonic analogy of the avian umbilical vessels with the mammalian;
circulation, which develops within the body of the embryo, and these confusions were clarified to a great degree by Hailer's
the vitelline circulatory arc, which develops in the area vasculosa, observations. Between Malpighi and Haller, little new became
form approximately contemporaneously. The third circulatory known about the area vasculo~a, and it was really the work of
arc, the allantoic, is formed later and carries blood to the al- both Haller and Wolff that represented the first major advance
lantois, a saclike structure where waste products are stored and over Malpighi's observations.
where the blood is oxygenated through a region of the allantoic Before the controversy between Haller and Wolff, the area
blood vessel network (the chorioallantois) that lies just under vasculosa did not figure as a major issue among either prefor-
the shell. In mammals, which have only a small vitelline circula- mationists or epigenesists. Harvey believed that the umbilical
tory arc (and no appreciable amount of yolk), the allantoic vessels are form.ed gradually, yet the area vasculosa never be-
circulatory arc is commonly called the umbilical circulation. came a principal example of epigenetic development for him
Through its connection with the placenta, the umbilical circula- (1651, 1847 trans.: 235,238, 392-97). Malpighi expressed the
tion carries on the functions of nourishment, waste disposal, belief that the umbilical vessels preexist, only gradually coming
and oxygenation, functions that are separated in the chick into view as they are filled with fluids and finally with red blood.
between the vitelline and the allantoic circulatory arcs (see Yet one must be careful to note, as Adelmann has pointed out
Adelmann 1966, 3: 1104-5; Patten 1971). (1966, 2:885-86), that when Malpighi maintained that a struc-
52 Matter, life, and generation The embryological debaJe 53
strating the nuances by which a soft and semifluid material can Yet Wolff's model, similar though it may be on gross observa-
pass to a state entirely different from its.primordial state, by the tional terms to Malpighi's, was unique to Wolff's epigenetic
simplest of evolutions" (p. 174). system. The movement of dissolved yolk substance into the
Thus, for Haller, the area vasculosa became a prime example embryo provided evidence, Wolff believed, for the existence of
of his mechanism for development from preformed parts. the vis essentialis. Because a heart cannot be observed in the early
Originally a semifluid, transparent material, the area vasculosa hours of incubation, it could not be responsible for the observed
becomes transformed into a complex web of blood vessels sim- movement of fluids. Therefore, "it follows that the nourishing
ply by the pumping through transparent vessels of fluids that particles pass from the egg into the embryo, and that there
become more and more opaque and gradually colored. Through exists a force, through which this is accomplished" (1759:73, §
this same sequence of events, solidification, expansion of parts, 168). It is the essential force that is responsible for the move-
and increasing opacity are produced in all of the embryo's ment of these fluids and thereby for the formation of the blood
preexisting primordial structures. The area vasculosa was for vessels of the area vasculosa.
Haller the most striking case of this developmental process. After the initial formation of rings in the area vasculosa, one
In Wolff's Theoria generationis, published a year later, the area observes, according to Wolff, the breaking up of these rings
vasculosa was again used as a principal example, but this time into "islands" surrounded by dissolved material. Gradually
for epigenesis. In early stages of development, the embryo is these islands become smaller and smaller, until a network of
surrounded, Wolff maintains, by a series of "rings" that form in connected channels is formed. 6 These channels, Wolff claims,·
the area vasculosa (called by Wolff the area umbilicalis). These are the initial rudiments of the blood vessels, obtaining vessel
rings, illustrated in Figure 10 (top), are formed, according to walls.only when the material immediately surrounding them
Wolff, by the movement of fluids· from the yolk to the embryo. begins to thicken. (This occurs after the blood is propelled
During the first moments of development, the yolk substance through the vessels by the newly formed heart.) As the network
begins to dissolve under the warmth of incubation and to move of channels increases in complexity, major branches appear, .
toward the embryo, guided by the essential force. The rings are and the completed vitelline circulation emerges (see Figure 10,
formed, Wolff says, from nonnutritive deposits that are left bottom).
behind as the dissolved yolk moves to the embryo. Wolff's Wolff sent Haller a copy of his dissertation on 2 3 December
description of the movement of yolk substance inwards toward 1759. 7 In his accompanying letter, Wolff indicates that he
the site of the developing embryo is reminiscent of Malpighi's hopes that his work might sway Haller's recent conversion to
discussions of the movement of colliquament (embryonic fluid) preformation and convince him again of epigenesis. "What
through the umbilical .area toward the fetus. Malpighi had then prevents me," Wolff writes, "having completed my little
stated that, in the earliest stages of development, the area work, from submitting it now to your most penetrating judg-
vasculosa in a chick egg seems to undergo a melting process. It ment as reasons fighting for the other side?" (Haller 1773-75,
"was not everywhere a solid structure," Malpighi noted, "but, 4:269). Little could Wolff have guessed the reaction his treatise
like a hill washed into and flooded by gushing springs, was was to provoke from Haller.
being gradually dissolved by colliquament invading it from Less than a year later, Haller published an anonymous review
outer rivulets" (1673, trans. Adelmann 1966, 2:947). He even of Wolff's Theoria generationis in the Gottingische Anzeigen von
speculated that these rivulets might be the primordial umbilical gelehrten Sachen. 8 Opening with the words, "We have not read in
vessels, which later become visible as colored fluid moves a long time a work as important as that of Carl Friedrich Wolff"
through them, yet he returned again to the idea that these blood ( 1760: :i 2 26), Hailer's review presents a fair summary of Wolff's
vessels are preformed (Adelmann 1966, 2:949). Even the term theory of embryological development. Yet Haller charges that
"colliquament" gives rise to comparisons with Wolff's later Wolff's entire model rests upon an unwarranted assumption:
observations, as it was coined by Harvey to denote the fluid out that if one does not observe a structure, then one can conclude
of which the embryo is produced, formed from the collique- that it does not exist. "But,"· Haller responds, "whoever has
scence, or melting, of the albumen and yolk. practiced much himself with the magnifying glass will have
~ .1,'.i,:~~,
.
The embryological debate 57
informed himself particularly in the mesenteric veins of frogs
Q5,'•
~-·~a.
~
that indeed strong colors make parts visible and transparency
(.
ft.r··•(t~~~~
-
,!.. ···-,~
-... ~ ~·-~,.-,.~-
makes them invisible; and in the grown animal, whose veins
certainly have in other cases visible membranes, very often the
~/ "--o'f,f,i::: '• blood corpuscles are visible, without one being able to recognize
'" .•<" - ·i, ~- ~
' . '-·. (;/!!,·.' ., '"'· ,.. _; ..,. ,· --,
"',. ....
.. , ·:-:, -; .. ·=··' the membranes of the veins" (p. 1229). There are clear cases,
Figure 10 (cont.).
forming rings and islands; t, the embryo seen through the amnion. Bottom, after 3 Y,
days of incubation: a, the embryo; c, d, t, f, g, h, i, major blood vessels of the area
vasculosa; I, vena terminalis (sinus terminalis). (Top, from Thtoria generationis, 1759;
Figure Io. Wolff's illustrations of chick embryos and the area vasculosa. Top, after 118 courtesy of the Francis A. Countway Library of Medicine, Harvard Medical School.
hours of incubation. The coarser material (a) is broken up by fluid material (b), Bottom, from "De formatione intestinorum," 1766-67)
58 Matter, life, and generation The embryological debate 59
Repeating his description of the formation of islands and location. This could not possibly be explained by Hailer's ac-
channels in the area vasculosa, Wolff asks, "Can I therefore say, count, for "that they are vessels or even only passages that ...
that in the circumstances, in which the area is ... [in early stages were expanded and thereby made visible through the force of
of incubation J, vessels are already contained in it? Rather, must the heart by means of the liquids pressed into them is impos-
I not say; that they are not only engaged in building but that sible, because these cavities do not communicate with one
only the very first stage in the building of vessels has begun? and another, and no passages toward the heart are present through
that consequently true vessels are not yet _there?" ( 1764:87). which the liquids would have had to come" (p. 262). The pump-
There is no basis, Wolff asserts, for maintaining that blood ing action of the heart could not possibly have been responsible
vessels preexist in the area vasculosa when one can actually for the appearance of the area vasculosa at this early stage in
observe their gradual formation. Consequently, Wolff continues, development.
"do I assert ... that there are no vessels because I do not see Wolff sent Haller a copy of his new book and wrote that for
them, and I make this conclusion on the principle what I do not the book he had "undertaken new experiments on incubated
see, is not there? Rather do I not conclude these truths from eggs, so that I might render the theory of generation more
that which I really see, 'because I see that the vessels for a long firm" (Haller 1773-75, 5:210; letter of 20 December 1764).
time are not finished, that they are scarcely first begun and Wolff calls special attention to his observations on the area
actually not yet vessels?" (p. 88). True vessels do not exist in the vasculosa, enclosing a series of illustrations (see Figure 20,
~arly stages of development. This one can conclude not by Appendix B; see also Belloni 1971). Two of Wolff's drawings
inference but because one can actually see them form. depict the secretion and solidification of the limbs from the
Wolff charges that Haller's account of the unfolding of pre- spinal column. The remaining two illustrate the formation of
existing vessels is contrary to observation in two ways. First, blood vessels in the area vasculosa and the islands and channels
Wolff claims that as the material surrounding· the channels produced by the movement of fluids during early stages of
increases in density one can observe the formation of vessel development. Wolff reiterates his argument that, because these
walls. Consequently, these vessel walls could not have preexisted islands and cavities begin to form first in the outer portions of
in a transparent state, as Haller claimed. Second, Wolff asserts, the area vasculosa rather than in the region nearest to the
if preexistent blood vessels become visible only when the heart embryo, they could not be produced by the pumping of the
pumps colored fluid through them, then one should observe heart. As he concludes to Haller, "The integrity of the area in
the islands of material in the area vasculosa separate from one the places closest to the amnion [and the embryo], which cannot
another in a regular, orderly manner, "as if they were cut with a be divided even by a needle without evident violent laceration,
knife from one another" (p. 94). But what one actually observes prevents you, illustrious one, from being able to regard these
is large islands separating irregularly from one another into fissures as perfected vessels that have only been dilated by the
smaller and smaller pieces. "You do not see the slightest trace, force of the heart and by blood propelled from the embryo"
but rather the opposite," Wolff asserts, "of an actual expansion (Haller 1773-75, 5:211-12).
and extension by the penetration of blood pumped from the Haller received Wolff's letter and the accompanying book
heart through such vessels, which are already there, and were sometime in February 1765, as he reports in a letter to Bonnet.
only folded together. And that is thus the second contradiction By March, Haller decided to resume his observations on in-
with nature the ideas of Herr von Haller make" (p. 95). cubated eggs. 9 Writing to Bonnet he explains, " ... this M.
Furthermore, Wolff argues in an appendix to his book based Wolff forces me to begin again experiments on eggs. It is a
on new observations on incubated chick eggs, if one looks at the matter of recognizing if the traces [in the area vasculosa] are
area vasculosa at an even earlier point in time, all you see is vessels or if these are globules without partitions. Is he not
cavities in the granular material, with channels not yet formed. amazed that the globules are formed from a circular vein and
These do not communicate with one another and are more can come back to the heart? The globules diffused in the
numerous at the periphery of the area than near the embryo's mesentery
. of a frog have certainly not undertaken a similar
62 Matter, life, and generation The embryological debate 63
perfected vessels, it darkens the blood little by little in the veins, formation of the blood vessels by claiming once again that,
but it does not do it all at once as it does when it touches the before red blood penetrates them, the vessels, and the pale fluid
blood immediately." If the blood vessels were .channels with no they initially contain, are too transparent to be visible. "Thus,"
walls, then the vinegar should reach the blood all at once, and an Haller maintains, " ... the vessels of the yolk that form the.figura
immediate change in color should take place. But, Haller venosa have been present for a long time even before the blood
argues, a gradual color change indicates that the vessels have with its globules came to it. It is only the pale fluid contained in
membranes because the vinegar reaches the blood more slowly. them that has hidden them" (1757-66, 8[1766]:276). Wolff's
This is what one observes when one places vinegar on mature observations relate to appearances only.
vessels. Consequently, true vessels must exist in the area vas- Finally, Haller asks, how is circulation established on Wolff's
culosa. Haller concludes triumphantly to Bonnet, "Thus, my system? Why do the small veins, for1:11ed in the outer regions of
illustrious colleague, this will make the foundation of a new the area vasculosa, come together in larger and larger branches
supplement on eggs; I believe that M. Wolff will be quieted and finally reach the heart? And how are the arteries also
down." To which Bonnet responds, "Your new observations on formed in a system parallel and complementary to the veins?
the chicken, my illustrious friend, have given me indescribable ''The more I reflect on the developing embryo," Haller re-
pleasure ... " (Haller MSS, 17 September 1765). marks, "the more completely I am persuaded that the related
With the results of these experiments in hand, Haller com- parts have been present at the same time; the arteries together
municated them to Wolff. As he reported to Bonnet, "I have with the veins, the organs, the accompanying nerves, and the
written to M. Wolff; I do not know ifhe will capitulate" (Bonnet bones. Chance would never have brought together an artery
MSS, 28 September 1765). This idea certainly appealed to originating from the heart, a vein continuous with the artery,
Bonnet, as he remarked in reply, "I am very impatient to know and a nerve from a completely different origin" (p. 278). The
the part that M. Wolff takes after having read and meditated on vessels are not formed gradually by an essential force, nor are
your critique. It pleases me to think that he will capitulate in any of the other organs. "That an animal is built from uh-
true philosophy, and this victory that you obtain over him will formed matter by a single propelling force," Haller concludes,
be as glorious as you yourself" (Haller MSS, 8 November 1765). "appears to me to be the same as expecting a river to arise from
Hailer's next published work on embryology was the eighth the Lake of Geneva whose branches have the figure of an eagle"
volume of his Elementa physiologiae, which appeared in 1766. (p.279i -
Responding to Wolff was clearly on Hailer's mind, as he de- In an Addenda section to the Elementa physiologiae, Haller
voted more pages to discussing Wolff than to any other single describes the appearance of the area vasculosa at different
figure (with Buffon a close runner-up). The text of this volume hours, remarking "I see all of this [in a manner] similar to what
must have been written before Haller's summer egg observa- the illustrious Wolff has .seen; we differ in our conclusions"
tions, because these are discussed only in a section of Addenda ( 1757-66, 8[1766], pt. 2:218). Reiterating the two experiments
at the end of the volume. In the text, Haller presents several he had reported to Bonnet, Haller concludes simply, "There-
other arguments against Wolff'.s views on the formation of the fore that at all times the vessels have had membranes I believe I
blood vessels in the area vasculosa. am able to take for granted" (p. 219). Haller also added these
First, Haller argues that most mature blood vessels have two experiments to a revised Latin version of his Sur la formation
muscular fibers in their vessel walls and nerves wrapped around du coeur dans le poulet, published in volume 2 ( 1767) of his Opera
them. But no one, he maintains, has ever found that muscle or minora. This version was considerably increased in size, as
nerve fiber can be newly produced in the body. Thus, how can a Haller included observations on chick eggs made during the
vessel be formed initially by the passage of fluids through summers of 1763, 1764, and 1765.
material? The vessels must exist, along with the associated Wolff turned his attention to the area-vasculosa again in his
muscles and nerves, before development. Haller then counters work on the development of the intestines ( 1766-67, 1768),
Wolff's argument that one can actually observe the gradual where he presents a significantly more detailed description of
64 Malter, life, and generation The embryological debate 65
the develop.ment of the vitelline· vessels than he had in his embryo can be observed to begin to develop before a heart can
earlier works (see Figure 10, bottom). Wolff here introduced the be seen to exist, there must be a force, Wolff concludes, that is
term area vasculosa, as he did several others, including vena responsible for this early movement of fluids. It is not surpris-
terminalis for the circular boundary blood vessel, altered after ing, then, that the issue of the heart's formation became a focal
Wolff to sinus terminalis, the term used today. Wolff continued point of the Haller-Wolff debate.
in this work to argue that the observed development of the area From the punctum sanguineum ("point of blood") of Aristotle
vasculosa is inconsistent with Hailer's preformationist descrip-
tion and that it supports instead an epigenetic interpretation.
Years later, in a publication written after Hailer's death,
Wolff, recalled his controversy with Haller over the area vas-
,
f
!
i
to Harvey's punctum saliens ("leaping point"), the appearance of
the heart in early stages of development was auniversally noted
event among observers of the embryo. Indeed as Adelmann has
remarked, "The beating heart, making its appearance while the
culosa. "I did not want at that time," he writes, "to dispute l other parts are still largely so inconspicuous as to escape the
against this great and admired man any longer. But it is clear " untrained observer, is perhaps the most striking, indeed, the
that he, not I, has concluded incorrectly from the appearances most dramatic feature of the early embryo" (1966, 3:1300).
that are the same overall for him and for me" ( 1789: 14 n.). Most early embryologists consequently believed that the heart is
Wolff describes the two experiments that Haller had performed the first organ formed, and many concluded that it was from the
to disprove his theory on vessel formation. Yet, Wolff counters, heart that other structures then develop (see Adelmann 1966,
these experiments prove nothing. If Haller was experiment- 3: 1300-1389). As Harvey proclaimed, "In a word I say, -from
ing on the area vasculosa at a time when it possessed colored the cicatricula [blastoderm] ... proceeds the entire process of
blood vessels, then the experiments would be expected to tum generation; from the heart the whole chick, and from the
out as they did. By this stage in development, Wolff claims, umbilical vessels the whole of the membranes ... that surround
vessels do indeed have thickened matter around them, because it. We therefore conclude that the parts of the embryo are
the heart h<;1s begun beating and the impulsion of blood causes severally subordinate, and that life is first derived from the
an increase in density of the material around the vessel channels. heart" (1651, 1847 trans.:397). Harvey believed thauhe heart.
Consequently, the vessels would behave as if they had true does not beat before it is filled with red blood - indeed, he
membranes. Thus, Hailer's experiments really proved nothing argued that the blood is made even before the heart. Others,
against his own theory, Wolff claims, because they were per- Malpighi among them, thought that they could observe a heart-
formed too late in the developmental process. beat before the heart is red.
It was again Malpighi who produced the first detailed obser-
·vatipns on the structure of the early heart. Although they were
The formation of the heart
not free from error, Malpighi's descriptions of the morpho-
Intimately tied to the debate between Haller and Wolff over the logical changes in the embryonic heart, as it develops from a
area vasculosa was their controversy over the formation or U-shaped tube to a four chambered structure, parallel essentially
preexistence of the heart. For Haller, the heart was the prime the gross anatomical descriptions given today. Malpighi believed
mover of development. At conception, according to Haller, the that the heart, like other structures, preexists before incuba-
tiny invisible heart, with its inherent irritability, is stimulated; its tion. Yet he also showed that it undergoes complex structural
subsequent beating produces the gradual unfolding and devel- changes before it reaches its final form. · ·
opment of the preexisting embryo. Thus, it was absolutely We know now that the heart does not really begin to form
necessary for Haller to maintain that the heart exists at all stages · until the end of the first day of incubation. By 29 hours, its
of development. For Wolff, on the other hand, it was equally structure is that of a tube, which, because it is growing faster
necessary. to demonstrate that there is no heart during the early than the rest of the embryo, soon bends into a U-shape. Then at
hours of development, so that. conclusive evidence could be about 40 hou.rs of incubation, the heart begins to twist into a
exhibited for the operation of the vis essentialis. Because the loop, at the same time that the embryo is twisting onto its side (as
J
The embryological debate 67
66 Matter, life, and generation
several new parts: but experience itself has shown me that the irregular contractions, which, however, pumped no fluids.
changes in this principal organ are only superficial and that they are Movement, Wolff concludes, "does not arise all at once, as I
born from its primordial structure by successive degrees, which is a once imagined, but by degrees; it gradually becomes stronger
proof for evolution [preformation] instead of being opposed to it. and at last finally passes over from a slow spastic movement, as I
[1758a, 2:172-73]
have described it, into an instantaneous, convulsive movement,
Haller now argued that the heart only appears to be devoid of its as we still see in adults" (p. 268). Thus, we cannot only observe
four chambers in initial stages of development. They are there the heart forming, we can also view the gradual development of
from the very beginning, just as the rigtit ventricle exists while the heartbeat. "Should all this still not be sufficient," Wolff
only the left is visible. They attain their visible state· as the concludes, "to prove my theory of the essential force, of the
gradual unfolding process of development takes place. gradual formation of the parts and the gradual attainment of
Haller believed that the heart appears to begin beating at the characteristics that ... they must have in adults? I shall leave
about 48 hours of incubation. "Why did movement begin at this the whole matter to the judgment of Herr von Haller" (p. 270).
time?" Haller asks, since one observes development in the chick As we have seen, Haller reported to Bonnet that, because of
before this time. The answer lies, once again, in the heart's Wolff's book, he felt that it was necessary to repeat his observa-
transparency; for "because it was transparent and did not strike tions on incubated eggs to satisfy himself "on the preexistence
the eye in one place, nor in another, nor in its passage from the of the heart, which this physician claims to have seen born"
first to the second," its earlier beating was not visible. "One does (Bonnet MSS, 1 1 April 1765). Yet Haller added little to his
not see the wind," Haller declares (1758a, 2:178). The heart discussions of the heart in the Elementa physiologiae. He repeats _
exists from tlie first moments of development, tiny and trans- again his belief in the heart's preexistence, and even reports
parent, but nevertheless capable of functioning to produce the without comment Wolff's observations on the weak beats of the
embryo's development. 29-hour heart. Haller simply concludes, ''[Wolff] repeats in
Wolff's discussions of the heart in the Theoria generationis several places that the heart has definitely not existed before 24
revolve primarily around when the heart becomes visible. In an hours .... I doubt strongly the truth of this phenomenon. This
embryo of 28 hours of incubation, he reports, no heart can be illustrious man has seen the heart at 29 hours, and I [have seen
seen ( 1759:72, § 166). By 36 hours, the heart is visible in the it] already formed and perfect at 48 hours. If it was not visible
form of a white ring, not connected to vessels and not beating earlier, it is because its.smallness, its transparency, its whiteness,
(pp. 72-73, § 167; see Figure 8). Yet "that the embryo is nour- and its resemblance to a mucous cellular substance concealed it"
ished at this time by the substance of the egg is demonstrated by ( 1757-66, 8[ 1766]: 116-17).
the increase in its volume .... it follows that the nourishing An issue closely related to this controversy between Haller
particles pass from the egg into the embryo, and that there and Wolff was Wolff's explanation of why animals have hearts
exists a force, through which this is accomplished, which is not and plants do not. Briefly, Wolff argued that the substance out
the contraction of the heart and the arteries" (p. 73, § 168). of which plants are formed solidifies much more quickly and to
Upon the existence of this force, and the absence of the heart, a more rigid degree than does animal substance. Therefore,
'rests Wolff's whole explanation of development. · . during development, only parallel vessels have time to form as
In the Theorie von der Generation, Wolff repeats that he has fluids move through plant substance. But in animals, Wolff
seen the heart at 36 hours and that it was not beating. "I am claims, branching vessels can form because solidification of
· certain," he writes, "that the heart has never moved in these structures proceeds much more slowly. Branching vessels,
embryos,just as I am certain that there is a time, namely before though, necessarily entail one common branch, and this, says
24 hours, when there is as yet no heart there at all" ( 1764: 265). Wolff, is the heart (1759:92-95, §§ 215-16; 1764:8-10).
Wolff also reports new observations that he made on an egg Haller comments, in his reviews of both the Theoria genera-
incubated 29 hours. To his surprise, he could see a small heart. tionis and the Theorie von der Generation, that he simply does not
Not only that, but he could observe that it made weak and · understand Wolff's argument. Yet Wolff thought a great deal
70 Matter, life, and generation The embryological debate 71
.External membrane
of his explanation for the difference between plants and ani- ,/ of Intestine.
mals, as will be made evident in the discussion of Wolff's ~
philosophy of science in Chapter 4. That Wolff's argument was
lost on Haller is symptomatic of a wide gulf separating the two Inner membrane of
in their attitudes toward scientific explanation.
..
Intcitine.••
tinuous with the skin and with the intestine of the fetus, it must
have existed with it: it is truly part of the fetus. The yolk existed unfertilized egg still in the mot~er. Therefore, he concludes,
in the abdomen of its mother independently of the approaches the embryo must be there also, connected to the yolk sac
of the male; the fetus must have likewise existed there, although through its intestines, but too small and transparent to be visible.
invisible and enclosed in an amnion, always apparently upon It is thus demonstrated that the embryo preexists in the mother.
the yolk, but invisible because of its smallness and its trans- Haller's proof was actually based on faulty observations. Be-
parency" (p. 188). fore fertilization, the egg of the chicken is indeed covered by a
Hailer's argument has been illustrated by F. J. Cole (1930), membrane, the vitelline membrane, which surrounds the yolk
reproduced here in Figure 12. Haller claims that one can ob- substance, the cytoplasm, and the nucleus. _After fertilization,
serve that the internal and external membranes of the embryo's the ovum acquires the albumen (egg white) and the shell. And
intestines are continuous with the internal and external mem- as the embryo develops, the yolk-sac membranes and the em-
branes of the yolk sac. One can also observe, according to bryo's intestinal membranes can indeed be seen to be continuous.
Haller, these same membranes covering the yolk sac in the But what Haller missed was that the yolk-sac membranes of the
72 Matter, life, and generation, The embryological debate 73
developing embryo are different from the vitelline membrane and the exterior membranes with the peritoneum (the abdom-
that covers the unfertilized egg, and that the vitelline mem- inal lining). On the basis of these later observations, one can
brane actually shrinks away as the yolk-sac membranes spread conclude, Maitre-Jan proposed, that the little white body (the
further and further over the yolk (see Figure 13). Thus, al- rudimentary embryo) and these yolk membranes were all
though the embryo is connected through its intestinal mem- formed at the same time in the ovary of the female. "And in
branes with the yolk-sac membranes, it is not continuous with fact," he maintained, "when the smallest eggs in an ovary are
the original vitelline membrane. examined, they are found to be white and covered by very
Haller's error is certainly understandable when one considers delicate membranes" (p. 2167).
what was known among eighteenth-century embryologists about Although Maitre-Jan's argument is certainly similar to Hai-
the membranes covering the fetus and the yolk sac. There was ler's, Maitre-Jan did not carry it to the extreme that Haller did
considerable confusion over the relationship between the blas- with- regard to proving preformation. In his Observations sur la
toderm, the yolk-sac membranes, and the vitelline membrane. formation du poulet, Maitre-Jan was most concerned to refute the
No one before Haller had described the situation any better animalculist view of preformation and to offer instead argu-
than he. It was really not until Wolff's observations on the ments in favor of ovism. He believed that the little white body
formation of the intestines, published in the late 1760s, that could be shown to exist in the unincubated egg and, by in-
some clarification of the relationship between the embryo and ference, in the ovary as well. Yet Maitre-Jan did not use his
the yolk sac was established. argument to support the preexistence of germs through em-
The origins of Hailer's "proof" of preformation are not en- boitement, claiming only that the little white body and the yolk
tirely clear. We know that he devised the membrane-continuity membranes are formed at the same time in the ovary. Nor did
proof during the observations on incubated chicken eggs he he ever clearly state that the little white body is a complete
made in 175 7, although there are no hints of it before his letter to miniature of the embryo.
Bonnet on 1 September. Additionally, Haller refers in Sur la There is no doubt that Haller was fully aware of Maitre-Jan's
formation du coeur dans le poulet and in the Elementa physiologiae to work. Referring to Maitre-Jan's book more than once in his Sur
the work of Antoine Maitre-Jan when discussing the membranes la formation du coeur dans le poulet, Haller notes with regard to the
of the yolk sac. Maitre-Jan had, in his Observations sur la formation structure of the yolk that "Maitre-Jan is almost the only author
du poulet ( 172 2), reported extensive observations on the yolk sac who has made any progress in discovering this" ( 17 58a, 2: 138).
and its membranes, and on the yolk stalk (the short canal that It is quite possible that, in the summer of 1757, when Haller
connects the embryo's intestinal region with the yolk sac). More began to investigate the yolk, he turned to Maitre-Jan's book for
significantly, Maitre-Jan had suggested an argument similar to guidance and that, in consequence, he repeated Maitre-Jan's
Hailer's. As Maitre-Jan explained, "Since the little white body is observations on unincubated and unfertilized eggs. Although
the chick in abridged form [en abrege1, the parts of which unfold Haller made significant advances in his own observations on the
gradually and are fashioned as the egg is incubated, ... and since yolk and its relationship to the embryo over those ofMaitre-Jan,
the exterior membrane of the yolk is an extension of the perito- he was sufficiently impressed with Maitre-Jan's continuity argu-
neum, and the interior one, covering the humor of the yolk, is an ment to see its use as a major proof of preformation. Never does
extension of the common membrane of the intestines, there is Haller hide his debt to Maitre-Jan, who is always duly footnoted
every reason to believe that this little white body and these · when Haller discusses his membrane-continuity proof; yet
membranes are formed at the same time in the ovary" ( 1722, neither does Haller ever contradict those who, like Bonnet,
trans. Adelmann 1966, 5:2167). Maitre-Jan believed that the were to ascribe this proof to Haller alone.
chick develops from a "little white body" 10 that can be observed Whether Haller was either entirely correct or entirely origi-
in unincubated eggs floating on top of the yolk. After the embryo nal with his membrane-continuity proof is less significant than
begins to develop, Maitre-Jan claimed, one can see that the yolk the importance he and others attached to his proof that the
membranes are really just extensions of membranes of the chick, embryo is preformed in the mother. As Bonnet declared in his
for the inner yolk membranes are continuous with the intestines Considerations sur Les corps organises, "I have said ... that one day
74 Matter, life, and generation The embryological debate 75
we would extract from Nature its secret. One oT its most cher- certain that the membrane of the yolk, with which the embryo is
ished favorites, M. le Baron de Haller, has recently interrogated continuous, is already present before incubation? It is precisely
it as it demanded to be, and he has obtained from it responses in this certainty that the entire proof drawn from the observa-
that extend the boundaries of our knowledge. It is the interior tion should consist" (1764:27 4). Yet Wolff claims that his obser-
of an egg of the chicken that has rendered to him its oracles" vations show that this is not the case - that the two membranes
( 1762, 1: 124). Bonnet enthusiastically endorsed Hailer's mem- are indeed not the same. "I am ashamed," he admits regarding
brane-continuity proof as a truly great discovery: "I had not his earlier discussion, "of the ridiculous dispute that I carried
hoped," he confessed to Haller, "that the secret of generation on in such detail, so clearly and so plainly, against the conclusion
would be discovered so soon" (Haller MSS, 30 October 1758). based on continuation; for such a continuation has never in the
Yet Hailer's proof also had its critics, notably Wolff. in his world existed in eggs; and this time I am like the scholars of old
Theorie von der Generation Wolff severely criticized Haller's argu- who disputed so ardently about the way the golden tooth arose"
ment both on logical and on observational grounds. To begin (p. 275).
with, Wolff challenged Haller on the relationship between con- Before presenting· Wolff's new observations, let me review
tinuity and simultaneous creation. As he queries, "if ... to put it briefly the sequence of formation of the various embryonic
briefly, the yolk of an egg continues into the embryo- can one, I membranes in modern terms, so that the full extent of the
say, conclude from this that the two elements, the egg and the Haller-Wolff exchange on this subject can be appreciated. Be-
embryo, therefore necessarily must have begun to exist simul- fore fertilization, the yolk, cytoplasm, and nucleus are enclosed
taneously at any given time, either before or after copulation or in a simple membrane, the vitelline membrane. Following fertil-
at the creation?" ( 1764: 105). Can one infer the constant co- ization, the albumtn (egg white), the shell membrane, and the
existence of two things from their observed continuity? Wolff's shell are added, and the egg is laid by the hen. As the embryo
answer is no. In his own account of development he had shown begins to develop (one can picture it as sitting on top of the yolk,
that parts of the embryo are secreted from other parts, for as it were), a membrane called the serosa (or, alternatively, the
example, the wings of the chick from the ~pinal column. Futher- chorion) begins to spread out from the embryo over th<: whole
more, the two structures thereafter remain connected together. embryo-yolk complex, while at the same time the yolk-sac mem-
Yet the wings and spinal column, Wolff asserts, were clearly not brane (together with the area: vasculosa) spreads out from the
formed at the same time but rather one after, and out of, the bottom of the embryo over the yolk (see Figure 13). As this
other. As Wolff challenges, "I do not see at all the connection happens, the original vitelliile membrane shrinks away before
between these propositions: A part of a thing is directly con- the advancing serosa and yolk-sac membranes. A third major
tinuous with the other part of the same thing; therefore the one structure then begins to develop, the allantois, a sac that serves
part of it could never have existed without the other part, but to collect wastes from the embryo. The allantois continues to ·
the whole thing must have been produced instantaneously" (pp. expand until it surrounds the entire embryo and yolk, and its
105-6). Consider, Wolff says, a wall built from bricks and outer membrane fuses with the serosa (allowing oxygenation of
covered with a common coating. After the wall is completed all the embryo's blood). The two joined membranes are now called
its parts are continuous with one another. But were they neces- the chorioallantois. While all of this has been happening, the
sarily produced all together at the same time? Clearly not, Wolff albumen has decreased in bulk so that it now occupies a very
claims, and thus Hailer's entire proof rests on a logical flaw. small portion of the egg. Consequently, the chorioallantois (the
In an appendix to his Theorie von derGeneration, Wolff returns fused serosa and allantoic membrane) is now right' next to the
to Hailer's membrane-continuity proof, criticizing it now on the shell membrane, and the embryo, yolk sac, and allantois, which it
basis of new observations he had made on chicken eggs. Hailer's contains, occupy almost the entire egg. All of this takes place by
proof, he correctly states, rest ultimately on the assumption that about the 14th day of incubation (Hamilton 1952; Patten 1971).
the membrane that covers the yolk sac and is continuous with Wolff reports from his observations that when one opens an
the embryo's intestines is the same membrane as the one that egg incubated for 12 to 16 days, carefully removing the shell,
covered the yolk before fertilization. "But," Wolff asks, "is it one first encounters an opaque membrane surrounding the
76 Matter, life, and generation The embryological debate 77
come to view" (1766-67:46CHi1; trans. Adelmann 1966, 4: formation. Wolff refutes each of these in turn, particularly one
1667). If the stomach, for instance, were preexistent, only on the membrane-continuity proof, "to which," he writes,
gradually becoming visible as it unfolded, we would have to be "twenty years ago I responded two things: 1) that it [the con-
able to see it, Wolff argt:ies, as a whole stomach from its first tinuity of membranes] does not exist as it is said, and as it would
moments of visibility. If it were preformed, as soon as it could be have to appear in order to prove evolution; [and] 2) that if it
seen, "however small it might be," it "would have to have the existed, as the late M. de Haller would have it, it would prove
true form and appearance of the adult stomach; but it would nothing." Wolff continues, "The solid refutation of my objec-
never be half a stomach, would never be open; and would never tions by the late M. de Haller consists of an observation by which
be joined with parts not belonging to it" (p. 455; trans. Adel- he believed he had seen that at a certain time of incubation (it
mann, p. 1664). Yet one can observe the stomach in these could only have been at the beginning of the third day), the
half-formed stages; therefore, organs do not exist in a tiny, vessels of the umbilical area exist already all formed. M. de
complete form, expanding during development to their adult Haller (in a letter to M. Bonnet) added a single word of scoffing.
size. "This, I say," Wolff asserts, "does not seem to be Nature's He believed, he said, that after this I would be quieted down."
design, but she has intended rather that the formation of This is indeed what Haller had written, in his letter to Bonnet
organic bodies should in general be-left solely to the natural (25 August 1765) announcing his two experiments on blood
forces inherent in animal and vegetable matter" (p. 455; trans. vessel formation, later published by Bonnet in his Oeuvres
Adelmann, pp. 1663-64). Nature dictates epigenesis, not pre- ( 1779-83, 5[ 1779):278-79). "But," Wolff continues to Euler, "I
formation. wrote after this my two dissertations on the formation of the
Haller reviewed Wolff's "De formatione intestinorum" in the stomach and the intestines. M. de Haller has called them very im-
Gottingische Anzeigen von gelehrten Sachen for the years 1770 and portant, and he has never responded to them. It seems, then, that
1771. 12 Both reviews contain Hailer's customary summary, and M. de Haller was quieted down himself" (Castellani 1971 :513).
it is only at the conclusion of the second review that we find any Wolff's "De formatione intestinorum" caused little stir among
reaction from Haller. Here he challenges, "But shoul1 it not his contemporaries. In fact, it was not until]. F. Meckel trans-
remain possible that a highly transparent intestine has appeared lated and published it in 1812 as Uber die Biulung des Darmkanals
to be simple and flat under the magnifying glass when it was im bebruteten H unchen that Wolff's work on the intestines became
already actually closed? At least the begip.ning of the thorax is so widely known. Its influence was particularly marked on Pander
extremely transparent, that it would be invisible without the and von Baer, both of whom recognized the germ layers, a·
acid that brings it out, and_ one day earlier this beginning can be concept toward which Wolff's work was a definite step.13 Von
entirely present indeed, and yet cannot be brought to visibility Baer, in fact, hailed Wolff's work as "the greatest masterpiece
by any acid" ( 1771 :415). It is not surprising that Wolff's account we know of in the field of the observational sciences" (1828-37,
would be dismissed by Haller in this way; the defense of trans- pt. 2:121 n.).
parency was used by Haller throughout the controversy to
undercut true development of parts. Yet one cannot help feeling
OBSERVABILITY AND EXISTENCE
a sense of regret that Wolff's remarkable observations should
have been relegated to the class of optical illusion by Haller, for A constant subtheme in the exchanges between Haller and
they were indeed a milestone in descriptive embryology. Wolff on the area vasculosa, the heart, the membrane-continuity
Wolff noted Haller's less than enthusiastic response to his· proof, and the formation of the intestines was the issue of the
work in a letter to Leonard Euler, written in 1783, six years after relationship between observability and existence. Can one pos-
Hailer's death. The subject was the appearance of a new edition tulate that an embryonic structure does or does not exist if it
of Bonnet's Contemplation de la nature in 1781 (in Bonnet's cannot be seen? Clearly this is a central issue for embryology,
Oeuvres), which contained some additional footnotes on pre- particularly in the seventeenth and eighteenth centuries when
Matter, life, and generation The embryological debate 85
84
good compound microscopes were not yet available (see Brad- aid to observing existing parts that are hidden only because of
bury 1967). Simple microscopes, and some compound, were temporary transparency.
indeed in use, but the earliest stages of embryological devC';lop- Epigenesists had to maintain, of course, that one can ,prove
ment remained hidden from view. It is pertinent in this-regard that embryonic parts are newly born during development and
to note Gasking's argument ( 1967:45-46) that the invention that they do not exist before this time. Wolff addressed himself
· .and use of the microscope in the seventeenth and eighteenth to this issue in his dissertation:
centuries indirectly aided the preformationists' position by In general, it cannot be affirmed absolutely that whatever is not
demonstrating that minute structures do indeed exist that are accessible to our senses therefore does not exist. However, applied to
invisible to the unaided eye. As a result, the possibility that even these experiments, this principle has in fact more elegance than
smaller structures might exist, not yet made visible by micro- truth. The constitutive parts, of which all parts of the animal body are
composed in their first state, are globules that always yield to [i.e., are
scopes, became much more plausible (see also Bodemer 1964: 2 3). discernible by] a mediocre microscope. But who would say that he
Indeed, Malebranche's first enunciation of the emboitement was unable to see a body because of its smallness, when nevertheless
theory in his Recherche de la virite was made within the context of the parts of which it is composed are precisely because of their
a discussion of the limits of our senses. There Malebranche .smallness unable to esc:ape notice? No one has ever uncovered with
noted that microscopes had revealed animals much smaller the help of a stronger lens parts not also detected immediately with a
than had ever be.en imagined; perhaps "there could be smaller microscope of cheaper quality. Either they are detected through no
and smaller animals to infinity, although our imagination is means, or they appear sufficiently large. Therefore, that parts are
alarmed at this thought" (1674:81). With lenses one sees the hidden because of their infinite smallness, thereafter gradually emerg-
ing, is a fable. [1759:72, scholium to§ 166]
germ of the tulip in the bulb, and one can conclude "the same
thing ... in general of all sorts of trees and plants, although this Wolff believed that all parts in plants and animals, as well as the
cannot be seen with the eyes or even with the microscope" (p. substance out of which these parts form, are composed of
82). The existence of germs encased within one another may be globules (not cells in the modern sense). 15 Because one can
beyond the realm of our senses, Malebranche admitted, "But it always observe these globules, he argued, which are certainly
is not necessary that the spirit stop with the eyes: for the view of smaller than the organs they make up, one can conclude that if a
the spirit is much further than the view of the body" (p. 83). larger organ is not visible, it does not yet exist.
The concept of emboitement could be accepted only on this Haller noted this argument in a letter to Bonnet, written
basis, that dimensions of size far smaller than we can even when he was reading Wolff's dissertation for review. Here he
imagine must be possible. The use of the microscope aided this remarks, "This Wolff, defender of Epigenesis, maintains that
notion by revealing a world below our senses. Microscopes all the parts of the embryo are comJ>Qsed of globules. Is this
furthered the preformationist cause also by making visible really true? I have indeed seen globules in the blood, grains in
smaller and smaller parts within embryos. That these parts the umbilical area, types of balls that are the germs of the
could all have existed in an invisible state since the Creation vertebrae, but have you ever seen the cellular tissue, [or] the
became a plausible idea. membranes in the form of globules?" (Bonnet MSS, 3 October
A common technique used to aid visibility was to apply 1760). In his review, Haller elevates Wolff's observability argu-
vinegar or alcohol to embryonic fluids, causing structures to ment to the status of the foundation of his system, claiming that
appear through the coagulation of the viscous fluids. t 4 In many "In the pr~duction of animals, one must pay attention to a
cases, artifacts were undoubtedly produced by this method, but principle, which stands directly at the beginning [of the section
organs were indeed made visible slightly earlier than normal on animals], and according to which that which is not there, one
through this technique. Haller often spoke of seeing the lung, does not see. The basis for this is, according to Herr Wolff, that
for instance, a day earlier than usual by applying vinegar or everything in animals arises from little globules; but these are
wine. Wolff also used this technique but claimed that it was an visible, consequently no part can be assumed [to exist] which is
86 Malter, life, and generation . The embryological debate 87
invisible and still ,present" (1760: 1228-29). "But," Haller re- was fibers, vessels, and viscera" (1757-66, 8[1766]: 116). Struc-
sponds, "whoever has practiced much himself with the magni- tures do exist in transparent states; one can prove that this is so.
fying glass will have informed himself ... that indeed strong Consequently, Haller declares: ·
colors make parts visible and transparency makes them invisi- No part therefore in the animal body has been made before any.
ble" (p. 12 29). Even in the adult animal, parts are sometimes other, and all created parts exist at the same time. If some have said
invisible because they are transparent. But this does not allow that the first origin of the new animal is the spine, the brain, or the
one to conclude that they are not there, back, or the heart; if Galen taught that the liver is the first to be
Wolff's response to Hailer's challenge was to argue, first, that produced; if others regard the abdomen with the head, [or] the spinal
his discussion of the relationship between observability and cord and the brain, as first, with the condition that the remaining
parts are produced from these, I believe that these illustrious men
existence was put forward in a scholium. It could not be a
mean nothing more than that the heart and the brain are visible to the
fundamental principle or axiom, because a scholium is by eye at a time when the remaining parts lie hidden: and that some parts
definition not part of the main argument. Wolff proceeds to of the body are developed to such a degree in the first days of the fetus
reiterate his globule argument, claiming that "the heart, the to be perceived. [p. 148]
vessels ... , the wings and feet, these parts, which all consist of
globules, [and] which are actually only white and indeed some- All is apparent development only; no true epigenesis takes
what as all bodies when they are small and thin, but in no way are place.
completely transparent as water or crystal, that these parts The issue of the relationship between observability and ex-
istence was certainly not one that could be settled, with any ·
because of their smallness or transparency could perhaps re-
amount of argument, between Haller and Wolff. Their differ-
main invisible, appears to me, if I should say as I think, to be an
ing viewpoints on this subject, as on other key points of conten-
egotistic excuse" ( 1764:91).
tion, were too fundamental to their separate positions to be
Second, Wolff maintains that such a principle as "what one
does not see, is not there" is not even valid. If something is not alterable.
visible, then one cannot conclude anything about it at all, be-
cause one has no grounds on the basis of which to draw a
THE EMBRYOLOGICAL DEBATE IN RETROSPECT
conclusion. But one can prove existence or nonexistence from
corroborative evidence. And this Wolff believed he had suc- In considering the Haller-Wolff debate in general, one is struck
ceeded in doing with regard to the blood vessels in the area by how inconclusive it was. None of the three principal issues of
vasculosa. "Transparency," Wolff asserts, "is only something controversy was really settled one way. or another, and the
one is forced to take refuge in. I adduce only observation" debate itself seems to have ended principally because Wolff
( 1764: 1I8). The preformationists, not he, had erred on the moved to Russia and Haller ceased doing embryological re-
observability-existence issue. "In short," Wolff remarks, "the search. Of course, one could argue that the issues that con-
whole story of invisibility is only a chimera. It can cause con- cerned Haller and Wolff could not be conclusively cleared up
fusion, but if we build anything upon it, it always deceives us" (p. during their day because of a lack of full observational knowl-
127). edge of embryological development, due in part at least to poor
Hailer's response to Wolff's remarks was simply to repeat his optical equipment. Yet this does not seem to be the complete
transparency argument in the Elementa physiologiae. It is cer- answer, for there is no doubt that if, for instance, the formation
tainly true, he admits, that the embryo appears to form out of an of blood vessels in the area vasculosa had been fully described in
unorganized primitive glue, but it does not follow that this glue their day, Haller and Wolff would still have disputed about how
does not contain the parts of the embryo. "I have often given this description was to be explained. It is on the level of explana-
consistency to this jelly," Haller claims, "with spirit of wine tion that one must seek the roots of the inconclusive nature of
alone, in order [to show] that_ what appeared to be pure jelly, the debate, and even the source of the controversy itself.
88 Matter, life, and generation
terms "that had become barbaric" into science, "that domain finally, from these liquids both our bodies are preserved and the
that Bacon and Galileo had torn away from the schools" ( 1746b: loss of human generations is replaced by new offspring" (p. i).
356-57). Scholastic rationalism has little to say about the real Sensation, motion, digestion, assimilation, growth, reproduc.,
world, in Haller's opinion, for "God has created individuals, tion - these are the functions of the living organism the physiol-
bodies,· movements, and one amuses oneself by contriving ogist must explain. And his explanation must provide, through
classes, modes, and qualities" (p. 356). force mechanics, "a description of those movements by which
Haller seems to have rejected rationalism from his earliest the animated machine is activated" (p. v). Physiology, Haller
student days. In an amusing story, Haller relates an encounter proclaims, i~ "animated anatomy" (1747:5).
with Cartesianism when he was studying as a young teenager Like his teacher Boerhaave before him, Haller endorsed the
with his stepuncle,Johann Rudolf Neuhaus. "The old man was application of mechanical laws to living processes. Yet this must
a determined Cartesian," Haller reports; "he began by making be done cautiously, for, Haller remarks, "in the animal machine
me study the principles of Descartes, and every page revolted there are many things that are very different from the common
me. 'From where do you know that the particles of the.second mechanical laws" ( 1757-66, 1:v-vi). Water flowing through a
element are round and that those of magnetic material are like a pipe, for example, is not totally analogous to liquids flowing
screw?' These questions came again at all moments and drew through living vessels, which act in various ways to speed up or
from me poor response" (Bodemann 1885:89). Haller's nega- slow down ,the movement of fluids. Simple hydraulics will not
tive attitude toward rationalism was undoubtedly reinforced by explain the motion of the blood through the organism. Haller
Boerhaave, Hailer's teacher at Leiden, who frequently expressed concludes, however, "I would not for this reason believe in
his own anti-Cartesian attitudes (see Lindeboom 1968:266-68, discarding the laws by which moving forces outside of the
270). Thus it was that Haller, a professor at a major German animal body are governed; I propose that they never~. trans-
university, was so opposed to the then dominant German Jerred to our animated body machines unless experiments
philosophical viewpoint. Trained in Switzerland and in Holland, agree" (p. vi).
imbued ~ith the principles of empiricism and Newtonianism, Haller was a mechanist in his physiological outlook, yet he was
Hallet rejected both Cartesian and German rationalism in favor not a total reductionist. Rather than reducing vital phenomena
of an empirical, Baconian approach (see Toellner 1973). The to the known laws that govern inorganic bodies, Haller pro-
philosophical gulf between him and Caspar Friedrich Wolff posed to create a distinct "animal mechanics." Here the basis for
could not have been wider on this point. explanation was to be laws that operate in the same manner as
physical laws, but which are not necessarily the same laws. Living
organisms thus may possess forces that are not found in non-
Mechanism and forces
living matter; yet these forces operate mechanically in exactly
"Whoever writes a physiology," Haller declared in the preface the same way as physical forces do.
to the first volume of hisElementa physiologiae, "must explain the As we can recall from Chapter 2, Haller believed that matter
inner movements of the animal body, the functions of .the is essentially passive and that forces are added to it by God.
organs, the changes of the fluids, and the forces through which "Indeed," Haller comments, "the great world bodies move
life is sustained" ( 1757-66, 1:i). Haller believed that the science themselves, and thus all parts of the earth and sun with them.
of physiology is the science of movement in living bodies, move- But this movement is foreign to them; it is imparted to them.
ment based on mechanical forces. The physiologist, he con- ... Consequently, movement of matter is not grounded in its
tinues, must explain the forces "through which the forms of being" (1775-77, i:223). And although motion, and the forces
things received by. the senses are presented to the soul; through responsible for motion, can be identified through observation,
which the muscles, which are governed by the commands of the we can never fully understand motion itself. "The measure of
mind, in turn have strength; the forces through which food is forces," he explains, "consists in their effects, for the nature of
changed into such different kinds ofjuices; and through which, motion itself, which is a most familiar thing, no one in philosophy
98 Matter, life, and generation The philosophical debate 99
has yet comprehended'.' ( 1775-77, 1:426). Speaking specifically ena. The science of animal mechanics can exist, Haller main-
about irritability, Haller remarks, "One will never know the tained, as yet another proof of the wisdom and design so
mechanical source, from which the movements that follow ir- evident in God's creation. ·
ritation arise, but one will approach [this], one will perhaps
succeed in measuring exactly the effect" ( 1777c: 105). All forces H aller's N ewtonianism
operate on a mechanical basis; all can be known through observ-
ing and measuring their effects-but their ultimate source is not Hailer's philosophical beliefs about science bear fundamental
matter but God. With regard to muscular movement, Haller similarities to those of Newton, both as expressed by Newton
writes, "We create no movement: our soul wills that the arm lift and as seen by his contemporaries. Evidence exists to show that
itself. But it gives to it neither the force nor the movement; God Haller was familiar with the works of Newton and of Newtonian
has put the force in the muscle" ( 1775-77, 3: 148). proponents and tQat Haller held Newton in high esteem as the
Haller defended his theory of irritability against the animist paragon of proper science. 7 During Hailer's years of study at
Robert Whytt and the materialist La Mettrie. 6 Haller was con- Leiden; the faculty included not only Boerhaave, one of the first
cerned to show that irritability is a force possessed by animal Newtonians on the Continent (Lindeboom 1968:268-70), but
muscle tissue that is completely independent of the soul. Like also 'sGravesande, an early popularizer of Newton (see 'sGrave-
gravity and other forces of matter, irritability is a property of a sande 1720-21). After Haller received his medical <;{egree, he
particular kind of matter that is activated by specific stimuli. traveled to England, where he was enormously impressed with
The source of all of these forces is the same, for, as Haller English intellectual life. "In the sciences," he wrote in his diary,
remarks, "God gave to bodies an attractive force and other "it appears that no land is superior to England" (Hintzsche and
forces, which once received are exercised" ( 1757-66, 7:xii). Balmer 1971 :93; see also pp. 94 and 98). Haller learned to read
Furthermore, with God the source of all material forces; Haller English after he returned from his journey, and the influence
argued, they cannot be used to support materialism, as La of English styles of poetry is evident in Hailer's own poems. 8
Mettrie had attempted to do. In his L'Homme machine ( 1748), La Newton received high praise in several of Hailer's poems. In
Mettrie had used irritability phenomena to bolster his argu- "Gedanken iiber Vernunft, Aberglauben, und Unglauben"
ment against the existence of the soul or any other immaterial ( 1729), for example, Haller writes, "A Newton exceeds the
substances. Yet, Haller countered, he had shown that irritability limits of created minds,/Finds nature at work and appears as
is completely separate from the soul; therefore, both must exist master of the universe;/He weighs the inner force, that is active
(1752b:158). in bodies,/That makes one fall and moves another in a circle,/
Along with his eighteenth-century contemporaries, Haller And he breaks open the tables of the eternal laws,/Once made
identified materialism with atheism. If there were no spiritual by God and never broken" (Hirzel 1882:46, lines 51-56). 9 Prais-
soul and if matter could, on its own, possess active powers, then ing Newton's achievements in another poem - the infinitesimal
the need for God as creator and ruler of the universe would be calculus, gravitation, color theory, and Newton's theory of the ·
seriously threatened. As we saw earlier, the dangers of material- tides - Haller declares, "He fills the world with clarity/He is a
ism and atheism figured significantly in Hailer's eventual rejec- continual source of unrecognized truth" (p. 73, lines 267-68). 10
tion of epigenesis. No material forces could by themselves That Newton should appear in such a praiseworthy manner
produce a living creature out of unorganized matter; otherwise in eighteenth-century poetry does not seem all that unusual.
what role would be left for God to play in generation? "Be- Yet, as Richter ( 1972 :66) has pointed out, this is quite unique
ware," Haller wrote to Bonnet, "that it is very.dangerous to with regard to the German-speaking world. In fact, Richter
admit the formation of a finger by chance. If a finger can form claims, Haller was one of the first, if not the first, to refer to
itself, a hand will form itself, and an arm, and a man" (Bonnet Newton in his poetry. References to Newton also abound in
MSS, 27 May 1766). Yet atheism and materialism need not be Hailer's scientific writings, where Newton is often represented
the result of a mechanistic approach to physiological phenom- as the champion of proper scientific rnethod. 11
100 Matter, life, and generation The phiwsophical debate 101
It is not surprising, given the scope and volume of Haller's for "we meet with very little Motion in the World, besides what is
reading, that he should have been acquainted with the major owing to these.active Principles" (1730:399). Hailer's views on
Newtonian publications of his day. Yet their effect on him was the relationship of matter to forces were, as we have seen with
far deeper than mere familiarity. In each of the principal as- regard to both embryological development and irritability,
pects of Hailer's philosophy of science discussed previously one nearly identical. 14
can trace a clear and fundamental Newtonian influence. In my Yet if forces must be added to matter, where do they come
opinion, Haller consciously sought to emulate the Newtonian from? For both Newton and Haller, the answer lay in the reli-
program in his scientific work and to construct, in particular, a gious context of their scientific views. Haller saw God as govern-
new physiology based on the canons of the new philosophy. ing the operations of the world through the forces that He
Concerning the proper method of science, Haller was in close imparted to matter at the Creation. This w~s also a salient
agreement with Newton. Observation and experiment should feature of Newtonianism in the eighteenth century. As Heimann
replace unbridled hypothesizing. Newton's famous declaration ( 1973: 1) has noted: "Newton's ideas were originally presented
that he will "feign no hypotheses" concerning the cause of and disseminated (by the Boyle lectures) in a form which stressed
gravity, because hypotheses not based upon phenomena "have the theological dimension to Newton's philosophy of nature.
no place in experimental science" (1713, 1934 trans.: 547), 12 For early eighteenth century thinkers, Newtonian doctrines of
finds its echo in Haller's silence concerning the causes of irrita- the passivity of matter, of the primacy of forces in nature, and of
bility and ~ensibility. "But the theory," Haller cautions, "why gravity as a power not essential to but imposed upon matter
one or the other of these properties is not in these parts, or is in expressed a theologyofnature." 15 For Newton and manyofhis
other parts 13 of the human body, such a theory, I say, I certainly contemporaries, nature an~ the laws of nature were· seen as
do not hope to give. For I am persuaded that the origin of both dependent upon Divine Providence. An intelligent agent created
abilities lies hidden in the intimate fabric, and is placed far the world, not blind fate. "Such a wonderful Uniformity in tqe
beyond the power 9f the scalpel or the microscope: beyond the Planetary System," Newton declares in the Opticks, "must be
scalpel or microscope I do not make many conjectures" ( 1752b: allowed the Effect of Choice. And so must _the Uniformity in the.
115). And even though we do not know the cause ofirritability, Bodies of Animals" ( 1730:402). Or, as Newton phrases it in the
we can postulate its existence from its observed effects "to which General Scholium to the Principia, "Blind metaphysical neces-
moreover it is unnecessary to assign any cause,just as no prob- sity, which is certainly the same always and everywhere, could
able cause of attraction or gravity is assigned to matter [in produce no variety of things. All that diversity of natural things
general]. It is a physical cause, ... discovered through experi- which we find suited to different times an"d places could arise
ments, which are evidence enough for demonstrating its exist- from nothing but the ideas and will of a Being necessarily
ence" ( 1752b: 154). Irritability, itself an attractive force according existing" (1713, 1934 trans.: 546). All of the uniformity, diversity,
to Haller, operates in exactly the same manner as gravity. and.design in the universe gives testimony to God's authorship.
That gravity, and especially the fact that its origins are un- Hailer's beliefs concerning the relationship of God to His
known, would have been used by Haller as an analogy for his creation present a similar viewpoint. Matter possesses-no forces
own unexplained force is not particularly unusual. In the or powers except through God's agency. "A first cause," Haller
eighteenth century, gravity was frequently called upon by declares, "has thus allotted to different classes of matter abilities
physiologists to sanction a myriad of unknown principles and and forces calculated according to a general plan, and it is there
powers that could not be explained but must, it was argued, be that we recognize 'the hand of the Creator" (1751b:86). Blind
postulated to exist (see Hall 1968). Yet in Hailer's case, we find a forces, on their own, could never produce the ordered, yet
utilization of Newtonian views on matter and forces that ex- diverse, world that we know. The constancy we observe testifies
ceeds the simple analogy of irritability and gravity. As Newton to God's will~ not chance, as the cause of phenomena. And the
argued in the Opticks (Query 31), matter alone possesses only diversity is evidence for God's iiltelligence, not necessity, as the
the passive principle of inertia. To matter forces must be added, guiding factor. Speaking .of the similarities and varieties one
102 Matter, life, and generation The philosophical debate 103
fiQ.ds among flowers, Haller observes, "All is not chance, other- Marburg for several years, Christian Wolff, now quite well
wise the carnation would become a tulip; all is not necessity, known, was recalled to HaHe in 1740 by Frederick the Great,
otherwise the carnation would remain always such as was the and he remained there as chancellor of the university until his
first carnation" ( 1751 b:87). An intelligent God, through His own death in 1754. The two professors of medicine, Hoffmann and
free choice, created the universe and the laws that govern it. Stahl, taught at Halle from the first years of its inteption,
Haller saw science as leading toward a deeper appreciation of Hoffmann remaining there almost continuously until his death
and reverence for God and away from the dangers of atheism in 1742, Stahl until 1715. None of the three was still at the
and materialism. Newtonian philosophy was also seen as a bul- university when Caspar Friedrich Wolff arrived in 1755, yet the
wark against irreligion both by Newton and by his contempo- context of philosophical and biological ideas fostered by them
raries. Richard Bentle.r's Boyle lectures '( 1693), the final three played an important role in Wolff's intellectq.al development.
of which were titled A Confutation of Atheism from the Origi.n and Christian Wolff, and the "Wolffian philosophy," was of enor-
Frame ofthe World, were designed to make just this point, as were mous influence on Caspar Friedrich Wolff. Imbued with the
those delivered by other Boyle lecturers. As Roger Cotes de- principles of rationalism, C. F. Wolff set out to create the first
clared in his preface to the second edition of the Principia, rational embryology. But this embryology also had its roots in
"Newton's distinguished work will be the safest protection the mechanism-vitalism controversy that had arisen between
against the attacks of atheists, and nowhere more surely than Hoffmann and Stahl during the early years of the eighteenth
from this quiver can one draw forth missiles against the band of century. Remnants of this debate were still existent at Halle
godless men"·(Newton 1713, 1934 trans.: xxxiii). Haller saw his when Wolff began his studies (Gaissinovitch 1956-57; 1961:
own work as adding another arrow to the quiver and as further- 214), and his own orientation toward biological explanation in
ing the Newtonian cause·of science in support of religion. his embryology represents a synthesis of the two views through
In the three major areas of his philosophy of sdence, Haller the principles of rationalism. Never a total mechanist or an
was fundamentally inspired by Newton and the Newtonian absolute vitalist, Wolff attempted instead to steer a middle
philosophy. Much of this influence ~ame not from Newton course between the reductionism of mechanism and the in-
directly but through other Newtonian proponents, especially explicability of vitalism.
Herman Boerhaave, whose own example as a scientist and
religious man Haller sought to emulate. In Hailer's emphasis on
'
experiment, in his views on the relationship of matter to forces, Rationalism
and in the religious context of his mechanistic outlook, Haller It was Caspar Friedrich Wolff's intention to be the first person
expressed a characteristic eighteenth-century philosophy, one to apply the principles of rationalism to embryology and thus to
derived fundamentally from the Newtonian world view. be the first to offer a bona fide explanation for generation.
"Since the reasons of the organic body are to be given in th~
WOLFF'S PHILOSOPHY OF SCIENCE theory of generation," Wolff claims, "this will give philosophical
knowledge of it, and therefore it will be defined as the science of
The most formative influence on Wolff's philosophy of science natural organic bodies. Furthermore, since anatomy teaches
was his education at the University of Halle, which he attended purely historical knowledge of the same subject, the theory of
from 1755 to 1759. Founded in 1693, Halle was one of the generation will be to anatomy·... what philosophical knowledge
major intellectual centers of the eighteenth century, its initial of a thing is to historical knowledge of the same. And therefore
faculty including Christian Wolff, Friedrich Hoffmann, and one could properlycallitrationalanatomy" (1759:5-6, §§ 1<>-11).
Georg Ernst Stahl. Christian Wolff joined the faculty as profes- Caspar Friedrich Wolff's distinction between philosophical
sor of mathematics in 1706, only to be banished from Prussi.a in and historical knowledge was taken directly from Christian
172 3 at the urging of a group of Pietist theologians whom he Wolff. who, in his Philosophia rationalis (1728), distinguished
had aroused with his philosophical writings. After teaching at among three types of knowledge: historical, philosophical, and
The philosophical debate 105
development and his vis essentialis. "The essential force," he more, determines the differences between plants and animals;
declares near the end of his dissertation, "along With the ability from this central determinant, "all the rest of the properties,
of nutrient fluid to solidify, constitutes the sufficient principle through which the animal organic body differs from the organic
of all vegetation [development] .both in plants and in animals" body of a plant with regard to its composition, necessarily must
(p. 115, § 242). . . follow." Consequently, Wolff concludes, "if thus I say in this
Wolff believed that before him only descriptions of genera- way someone comprehends the building of this organic body
tion had been given, that no bona fide explanations had ever from the nature of the forces that form it and thus compre-
been offered. "I believe that I do not err very much," he hends it as a result of its causes, then he has a philosophical
remarks in his Theorie von der Generation, "when i say that · knowledge o( it which is very different from merely historical
despite the many works that in ancient as well as recent times knowledge" ( 1764: 10).
have been published on generation, nevertheless up to now no Wolff believed that scientific method should be based both on
one has given a true system [Lehre] of generation ... or has logic and on empirical findings. The scientist shoul~ aspire to
really explained generation" (1764:2). Wolff, however, makes philosophical knowledge, but philosophical knowledge must be
one concession, for his fellow rationalist'Descartes, whose work based on historical knowledge. Wolff's own repeated anat6mi-
on generation Wolff claims not to have read when he wrote his cal investigations and observations on plants, chicken eggs, and
dissertation. Although his system was "as false as possible," other organisms testify to the importance of empirical data in
Wolff asserts, · Descartes "showed what an explanation must his research. Yet one must not stop here, for it is through logical
look like, and he taught how one must philosophize if one wants reasoning that one can understand why observed phenomena
to do it really and not just to have the appearance of having are as they are. Wolff's vision, based as it was on rationalism, was
done it. And in this lies the most distinguished merit of Des- thus of an interconnected, logical universe, accessible to those
cartes" (p. 6). What Wolff seems to have admired the most who follow the proper method of inquiry. And through logic
about Descartes's fermentation model of development (see and empirical investigation, one can uncover the rationalstruc-
Chapter 1) was Descartes's attempt to show in a deductive . ture of the world, and one can explain the logical necessity of
manner how the parts of the embryo must necessarily develop the phenomena that it exhibits. This is what Wolff believed he
on the basis of certain principles about matter and motion. had done for the science of embryology.
Descartes failed in his effort, yet, Wolff maintains, his is the only
t.fieory of generation before Wolff's to offer a real explanation. Mechanism, vitalism, and reductionism
Wolff presents an example in the Theorie von der Generation,
which had also appeared in his dissertation, of a proper philo- The crux of the conflict that had occupied Hoffmann and Stahl
sophical explanation. His topic is the difference between plants in the early part of the eighteenth century was the extent mech-
and animals, and why animals possess a heart while plants do anism could be used in explaining physiological phenomena. 17
not (see also Chapter 3). Wolff argues that one can deduce the Hoffmann argued, in liis Fundamenta medicinae (1695), that
reasons for this from the observable fact that animal substance "Medicine is the art of properly utilizing physico-mechanical
solidifies much more slowly than plant material and in fact principles, in order to conserve the health of man or to restore it
never reaches the same degree of rigidity as plant substance if lost.... Like all of nature, medicine must be mechanical"
does. Because of this difference in solidification capabilities, the ( 197 1 trans.: 5-6 ). Stahl believed, on the contrary, that life could
movement of fluids propelled by the vis essentialis through plant not be reduced to mechanical causes; rather the soul (anima)
substance will be able to form only parallel vessels, while in controls the vital functions and resists the decomposition the
animal substance, because it is much less quickly rigid, branch- material body would undergo ifleft to physical forces alone (see
ing vessels will be formed. Branching vessels, however, neces- Stahl 1 708).
sarily entail one common vessel, namely, the heart, from which C. F. Wolff has often been cited in the secondary literature as
all others ultimately arise. The possession of a heart, further- a vitalist, principally because of his concept of a vis essentialis.
108 Matter, life, and generation The philosophical debate 109
Yet, although not a reductionist, Wolff was wary of vitalism also. it; but consider such an explanation as a proposition lacking a
He turns to just these issues in his dissertation when he asks, -~ demonstration, and add one to it" (p. 127, § 255, scholium 2).
"how are life and the machine mutually connected together in The. connection between cause and effect, the sufficient reason
natural organic bodies? Do they accordingly both depend on a f1i · for the existence of the effect, must be. dem.onstrated. If the
common cause, or indeed one upon the other_? And if the latter .'ff;
propqnents of mechanical medicine had followed this proce-
is true, what does life contribute to the machine or the machine } dure, Wolff testifies, they would ha:ve realized the folly of their
l'
to life?" ( 1759:9, § 36). Wolff's answer is presented in the undertaking. _
closing section of the Theoria generationis, where he discusses the
l But iflife-is not to be reduced to mechanical principles, is it to
"I-ii
nature of "mechanical medicine." be explained vitalistically? Wolff answers no to this also, al-
Wolff decried the total reductionism he saw in the application though somewhat ambiguously, in his dissertation. First of all, it
of mechanism to biology being made in his day. Mechanical is clear that Wolff separates the soul from the vegetative process.
causes, that is, those resulting from the way in which the body is As he argues,
made up of its parts, are accessory, not essential, to vital proc- f
l
what do we mean properly by vital actions? Life is attributed to the
esses. Consequently, Wolff alleges, "mechanical medicine, animal either insofai: as it thinks and moves voluntarily, and thus on
whether it exists already developed or may hope to be further account of the acting soul; or for the reason that through various
refined, is thus far an imaginary system ... to which there cor- motions occurring in animals, whatever these may be, the maintenance
responds nothing that exists in the nature of things" (1759: 124, and augmentation of the body are continually accomplished. Both
§ 255). In the process of digestion, for example, chewing and are observed in animals; only the latter is observed in plants. Thus, on
the one hand, sensations, voluntary motion, ratiocination,· and the
swallowing are mechanical accessory causes. But the assimila-
remaining [aspects of] thinking, [and] on the other hand, chylifica-
tion of nourishing fluids is a vegetative process, not based on tion, sanguification, and whatever actions contribute to conservation
mechanical causes. "If you now compare," Wolff argues, "the can be called vital actions, seeing that in both cases life is composed of
importance of these [mechanical] actions ... with the ability [of these. But the former are called animal, the latter are called natural.
the animal] to maintain itself continually, constantly building [1759: 120, § 250, scholium 2]
and growing ... will you be able to doubt either, as. I have Those vital activities that pertain to the soul in animals (volun-
pointed out already, that the machine and whatsoever actions tary _motion, _sensations, thinking) are added to those of the
that depend on it are now to be distinguished from the animal vegetative body. Consequently, the soul is in no way part of the
itself? or indeed, that all these enumerated mechanical actions vegetative process itself.
are to be· considered only as an unimportant appendage of Wolff also distinguishes his views from those of Stahl, who
animals?" ~pp. 125-26, § 255, scholium 1). Because vital activi- attributed all vital actions to the activities of the soul. In a
ties are not mechanically based, mechanical medicine is not passage near the end of his dissertation, Wolff declares,
capable of accounting fully for the activities of the organism.
Wolff also. rejects mechanical medicine on philosophical All those functions of the body that I have denied to be mechanical, I
have not explained in any way, inquiring in fact into the connection
grounds, arguing that its proponents have not followed proper that exists between the machine and life, but by no means searching
explanatory procedures. They have merely pointed to things further for the causes of this where it has no dealings with the
that occur together and called one the cause and the other the machine. If therefore you should wish to int~rpr«:t my mind on this,
effect. But they have neglected, Wolff charges, to demonstrate benevolent reader, you could easily err in this. And certainly indeed
the sufficient reason connecting cause and effect in their argu- and especially I would suffer from it [paterer], 18 if you should impute
ments. Mechanical causes, Wolff admits, are part of vital proc- to me the opinion of Stahl, or that received from him and slightly
esses, but they are not their true cause. i.Do not think," Wolff altered that Whytt and other more recent scholars have proposed, in
which, namely, the functions that occur in our body are attributed to
cautions, "that the whole matter has been brought to light
the power of an immaterial soul, whether.acting directly and freely,
through a demonstration of the existence of this thing which or coerced by the inconvenience inflicted upon it. E1759: 130-31,
you take for a cause, and through a deduction of the effect from § 255, scholium 4)
110
Matter, life, and generation The philosophical debate I I I
Although Wolff agrees with Stahl that vital processes cannot be formation. Using the idea of pre~xistence of germs, Haller was
explained on the basis of mechanical reductionis~, he does not able to explain development in mechanical terms while retain-
concur on the attribution of these functions to the soul. Rather, ing a place for God as the ultimate cause of the production of
on Wolff's system these vegetative processes are to be explained new life. "If the first rudiment of the fetus is in the mother, if it
through the essential force and the secretion and solidification has been built in the egg, and has been completed to such a
of fluids.
point that it needs only to receive nourishment to grow from this,
Wolff's philosophy of biology thus rests on his separation of the greatest difficulty in building this most artistic structure
"vegetative" processes in living organisms from both mechani- from brute matter is solved. In this hypothesis, the Creator
cal processes and those, like sensation and thought, that depend himself, for whom nothing is difficult, has built this structure"
on the soul. Wolff consistently used the term "vegetative" to ( 1757-66, 8[1766]: 143).
refer to the activities of nourishment, maintenance, growth, Haller must have challenged Wolff on just this point in a
and development in the living organism, that is, the vital func- letter, for Wolff turns to the relationship between religion and
tions. Clearly, it is the essential force that is the key to these embryological development in a reply to Haller, written on 17
vegetative processes and thereby the key to Wolff's philosophi- April 1767. Wolff admits that preformation, if true, would be
cal views on the nature of life. Denying total reductionism, yet an outstanding proof for the existence of God. Yet, "it is true,"
unwilling to ascribe to vitalism either, Wolff sought to create an he continues, "that nothing is demonstrated against the ex-
explanation for life processes that was mechanical in ~ts own istence of Divine Power, e·ven if bodies are produced by natural
right yet also unique to living creatures. II
i~
forces and natural causes; for these very forces and causes and
nature itself claim an author for themselves, just as much as
THE PHILOSOPHICAL DEBATE organic bodies do" (Haller 177 3-7 5, 5: 318): Explaining devel-
;~,-
opment on the basis of natural causes does not threaten the
Although the debate between Haller and Wolff centered around
such issues as the development of blood vessels in the area
~ existence of God, for these natural laws must have been created
by God. Besides, Wolff asserts, "by far the clearer and better the
;~
vasculosa, the formation of the heart, and Hailer's membrane- proof would be, if, in contemplating the state of nature, we were
i'
continuity proof, the positions each took oh these points of to find that a single product of it or organic bodies had had need
contention were intimately tied to their differing philosophical of the Creator, and that nothing organic could have been
views. Their attitudes toward empiricism and rationalism, produced through natural causes" (p. 318).
mechanism and vitalism, religion and science all played signifi- This is an argument that Wolff expounded in his Theorie von
cant roles in their debate. Occasionally, some of these under- der Generation ( 1764 :40-46). Here he asks, why should we admit
lying issues surfaced in the debate, yet their presence can be felt preexistence of germs for living organisms if we can find no
throughout the controversy. The two points that received ex- other example in nature of development from causes arranged
plicit discussion concern the questions of religion and of bio- at the beginning of the world and hidden until their moment of
logical forces. Both of these were closely connected to each operation?
person's embryological theory, on the one hand, and to their Wolff objected to the preformationists' reliance on God
views on the nature of scientific explanation more generally. rather than nature as a cause of generation. "It is of particular
importance to me," he remark~ at one point in his dissertation,·
Religion "to discover the principles and universal laws of generation a
posteriori, and especially to show in any event that the finished
The issue of God's relationship to his'Creation played a major plant is not something to whose production natural forces are
role, as we have seen, in Haller's views on embryological devel- completely insufficient and which requires the omnipotence of
opment. His inability to reconcile epigenesis with his religious the Creator: inasmuch as this has been observed [in plants],
views was one of the_ factors that led to his conversion to pre- nothing will stand in the way of. our allowing the same for the
I 12 Matter, life, and generation The philosophical debate 113.
rest of natural organic bodies" (1759:38, § 71, scholium 2). changeable and capable of taking all sorts of forms?" (1757-66,
Preformation offers no real explanation for generation; for 8[ 1766]: 117). How could a force be capable of such organizing
"those who teach systems of predelineation," Wolff asserts, "do activities? Haller continues:
not explain generation but deny that it occurs" (p. 5, § 3). To Why does this force always build without any error a chicken out of
offer a scientific explanation of embryological development the unorganized material of a hen, a peacock out of [the material
one must show how natural forces and causes are themselves irom] a peacock? Nothing is assumed [by Wolff] other than an ex-
responsible for generation. panding and progressive force. I would expect nothing more from
In the "fheorie von derGeneration, Wolff laments what our view this than that the net of vessels would necessarily become larger, as
of nature would be if preformation were indeed the case: long as the expansive force can overcome the resistance. Why, at the
site of this net, are formed a heart, a head, a brain, a kidney? Why in
All organic bodies [would] thus be ... miracles. Yet how very changed each animal is there its proper arrangement of parts? To these ques-
would our conception be of present nature, and how much would it tions no response is given. [p. 117]
lose of its beauty! Hitherto it was a living nature, which through its
own forces produced endless changes. Now it is a work that only Wolff's essential force, Haller argues, cannot be the source of
appears to produce changes, but that in fact and in essence remains as organization, because by Wolff's own admission, it is a simple
unchanged as it was built, except that it gradually is more and more force that operates only through the movement of fluids.
used up. Before it was a nature that·destroyed itself and that created "We do not believe," Haller declares forcefully with regard to
itself again anew, in order to produce endless changes, and to appear ,2: all theories similar to Wolff's, "that there is any force that,
again and again from a new side. Now it is a lifeless mass casting off ·t without being guided by wisdom, can act on matter, foilowing ·
one piece after another, until the affair comes to an end. [ 1764:73] l
directions that are constantly different and that ·are properly
But nature is not this way, Wolff contends, and science must not
make it so.
i~i
combined together in such a manner that this brute matter is
formed into bones, muscles, organs, and vessels, and so that cJ.ll
The contrastbetween Wolff's view of the relationship be- of these are joined together in a certain order. All that is
tween God and the world and Haller's-conception is indeed produced spontaneously, even those full forms of snowflakes
striking. Where Haller saw a threat of materialism and atheism [produced] by artistry, following a single direction, is always
if material forces were used to explain development through formed in hexagons, always in points" (p. 118). Simple fo011a-
their actions alone, Wolff saw instead a case of proper scientific tion of snowflakes, crystals, and the like always takes place in the
explanation based on natural causes. The sterile world of the same manner and always produces the same'regular structure.
preformationists was what Wolff rejected. Both believed that But, Haller claims, in the formation of the eye, for example,
embryological development is carried out by means of material simple forces could never produce the intricately related mem-
(orc~s and natural laws, yet Wolff was able to allow more flexibil- branes in just the right arrangement so that light produces
ity, as it were, for their operation in nature-. Yet how did his epi- vision. "But these [objections] are infinite ... ," Haller concludes.
genetic theory solve the problems that Hailer's had encountered? "I do not think that this illustrious man has seen anything other
than the growth of the chicken, which is guided by heat and the
heart" (p. 1 18) . .
Forces and embryological development
Wolff argued in his dissertation that the movement of fluids
Haller objected td Wolff's vis essentialis on much the same in plants and from the yolk to the embryo in chicken eggs
grounds that he had earlier brought to bear against Buffon's provided observational evidence for the existence of the vis
"penetrating force." How, Haller asked, could Wolff's force essentialis. Furthermore, he distinguished the essential force
account for the source of embryonic organization? "Why," he and the process of solidification from the expansive and resis-
challenges, "does this essential force, which is unique, form tive forces of Needham, noting that Needham's forces operate
parts in the animal that are so different, always in the same place on a completely different physical basis than the essential force
and always according to the same model, if inorganic matter is ( 1759: 106-7, §§ 233-34). Yet beyond this, Wolff does not ex-
l
114 Matter, life, and generation The philosophical debate 115
plain the nature of his essential force any further, concluding body, repair and growth are produced, and the new parts of the
simply that "The essential force, along with the ability of nu- embryo are formed. Because this occurs in plants and in ani-
trient fluid to solidify, constitutes the sufficient principle of all mals dudng embryonic development, the question states, it
vegetation both in plants and in animals" (p. 115, § 242). The cannot be due to the heart and must t_herefore be produced by a
movement of fluids by the essential force, and the solidification i force. "Therefore it is asked," the prize question concludes,
of thes~ fluids, produces all of the phenomena of development. What is the nature of this force? In the first place whether this is the
Wolff was not unaware that he had left the nature of the same as the universal attractive force of bodies, or rather, in what way
essential force partly unexplained. Yet this was not a major is it seen to be different from this and proper only to living animal
concern for him, as he tells us in the Theorie von der Generation: substance and to the vegetable substance of plants? If this latter is
true, it is asked further, what are its particular effects, and by which
It is enough for us to know that it is there, and to recognize it from its properties is it distinguished from the universal attractive force and
effects, as it is demanded purely and simply in order to explain the its singular and special nature made evident? [Blumenbach and Born
development of parts. In the name we call it by lies still less; only this 1789:ii-iii]
must I remind you of, that it is that force through which in the
vegetative body all those thin·gs are accomplished on the basis of Wolff's treatise, in part a commentary on the essays of Blumen-
.which we attribute life to it; and on this basis I have named it the bach and ~orn, is primarily a presentation of his own answer to
essential force of those bodies, because, namely, a plant would cease this question.
to be a plant if this force were taken from it. In animals it occurs as in Wolff proposes a very simple model for how the essential
. plants, and everything that animals have in common with plants force operates. In living organisms, he contends, like substances
depends solely on this force. [ 1764: 160] ·
attract one another, whereas unlike substances repel one an-
The essential force, derived from observational evidence, is that other. On the basis of this phenomenon; Wolff claims, one can
which constitutes life, that which distinguishes the living crea- explain all vegetative activities. In nourishment, for example,
ture from the nonliving. liquids are brought to the different parts of the plant or animal,
Over twenty years later, Wolff published a work devoted to each part attracting out material that is similar to it, which can
the essential force, Von der eigenthiimlichen und wesentlichen Kraft therefore be used for growth or repair. The attraction is pro-
der vegetabilischen sowohl als auch de,: animalischen Substanz ( 1789). d uced by the fact that the two substances are similar in nature
"I flatter myself," Wolff proclaims, "that this essential force, and is caused by the presence of the essential force in both the
which I laid down indeed at that time [in the early works] as the nourishing liquid and the parts of the organism.
foundation and also proved, but in no way explained wherein it In embryological development, attraction and repulsion are
exists, now ... through this present treatise will be explained" responsible for the process of secretion and solidification pro-
( 1789: 50 n.). Although Wolff's work was written several years posed earlier by Wolff to explain the gradual formation of
after Hailer's death, I shall discuss it here in the context of the · structures. A solidified part secretes, through repulsion, mate-
debate because it sheds light on how Wolff conceived of the rial that is dissimilar to it. This· then solidifies to become a new
essential force both in his early and later works. structure and can grow by attracting material similar to itself
Wolff's treatise was written as a response to and published from the nourishing liquids. In animals such as the chick, Wolff
along with two essays on similar topics by Johann Friedrich explains, development commences when the yolk is warmed
Blumenbach and Carl Friedrich Born. The Blumenbach and and begins to dissolve ~nto fluids, which are then repelled from
Born papers had been selected for first-place honors in a com- it and are attracted to the site of the new embryo. The result of
petition held by the St. Petersburg Academy of Sciences for the this mov.ement of fluids is the blood vessel network of the area
best essay on the nature of the "nutritive force." The prize vasculosa.
question, originally suggested by Wolff and made publicby the Wolff believed that through the attraction and repulsion of
Academy in 1782 (see Lukina 1975:414), defined nutrition as the essential force he could explain all vegetative activities,
the process whereby nourishment is brought to all parts of the including nourishment, sanguification, embryonic development,
116 Matter, life, and generation The philosophical debate 117
and even irritability and sensation in animals. He argues again, one such part, at another time, another part" ( 1789: 66 n.). If a
as he did in his early works, that mechanical causes (for exam- building force produces, for example, a root at one·end of a
ple, the pumping of the heart after it is formed) influence seed, should it not then have to produce a root at the other end?
vegetative activities but do not cause them. One could hypo- If building forces were responsible for development, Wolff
thetically construct, for instance, a model of a plant that was admits, there would re no reason why development should
exactly like a real plant in inner and outer structure. But would proceed the way it does. "One must either deny," he concludes,
this model grow and reproduce? "I think this model would "that nothing in the world can occur without sufficient reason,
remain the same," Wolff answers, "and I think that even the or one must admit the impossibility of a buildingTorce" (p. 67
most eager defenders of mechanical medicine would not at- n.). Either we must give up our- belief that everything has a
tribute such activities to the model" ( 1789:39). The real plant is sufficient reason or we must reject the idea of a building force.
alive because of the essential force, which "must be peculiar to But how is Wolff's force different from a building force? The
this plant and animal substance, because no material other than essential force, Wolff explains, is a force that always acts th:e
plant and animal substance is nourished, vegetates, or repro- same in every situation- it attracts when like substances are near
duces its kind. And because, moreover," Wolff concludes, "the one another and repels when unlike substances are near, nothing
whole life of plants [and animals], their nutrition, growth, vege- more. It is the particular situations that arise with regard to like
tation, and reproduction, rests upon it, one can call it a char- or unlike substances being near one another that determine the
acteristic and essential force. For where this force is absent, all eventual outcome of the essential force's action. The essential
vegetable processes cease" (p. 39). The structure of the organ- force itself "produces the different parts of the organic body no
ism cannot be alive without the essential force. longer merely through itself and according to its nature, but
Wolff again distinguishes his force from the vitalism of Stahl, rather with the help of countless other concurring causes; and
even more explicitly than he had earlier in his dissertation. what it does through itself alone, becomes a completely simple
"This characteristic and essential force," he asserts, "appears to ·effect, as attraction or repulsion, and is worlds apart from the
be that, if I do not err, .... .whose existence Stahl very certainly building of organic bodies" ( 1789:67 n.). All of the processes of
recognized, but which he, incorrectly I di.ink, attributed to the vegetation are produced by the essential force plus different
soul. It consists in nothing further than a particularly defined circumstances. Among these circumstances the chief or;ie is
kind of attractive and rep~lsive force" ( 1789:42): The essence certainly similarity or dissimilarity of substances, which Wolff
of life,.Wolff contends, need not be attributed to a soul but can, attributes at one point to "chemical properties," although he
rather, depend upon an attractive and repulsive force. · does not specify what these may be (p. 53). In order to explain
Wolff further articulates the nature of the essential force in a how any particular structure arises in development, one must
discussion of Blumenbach's Bildungstrieb ("building power"; show how the force of attraction or repulsion has interacted
also called the nisus formativus). Here he discusses the kind of with the existing circumstances to produce the observed phe-
<>bjection Haller had raised about Wolff's force, when Haller nomena. Thus, the essential force itself is simple and auto-
had questioned how the vis essentialis could be responsible for matically acting, b~t it produces different ultimate effects in·
development. Wolff explains here that his own force is not a different situations.
"building force" and agrees_ with Haller's position that forces Wolff likens the essential force to the universal attractive
can riot be capable of fashioning, on their own, new organisms. force, and notes that most phenomena seem to depend on
Forces, Wolff argues, are simple in nature and must always attractive and repulsive forces. Perhaps, he at one point specu-
produce one invariable effect. Consequently, "the generation ,.
"-'
lates ( 1789:70), all phenomena are ultimately produced by one
or building of the different parts of the body could not depend l single force that is determined differently in different substances
immediately merely upon one force. For no reason would exist ~',' (for example, magnetic, electrical, living). Yet, even if this were
i
for why this force should work at one time in such a way, at the case, the essential force would still be a separate force,
another time in another, or why it should produce at one time
instance, gravity is in its mode of operation, yet it produces the sophical beliefs shaped his view of biological phenomena and
phenomena of life only in plant and animal substance. dictated support for either preformation or epigenesis. Let me
Wolff complains in this treatise on the essential force that now summarize the major elements of the philosophical split
Haller and others had misunderstood his initial presentations between Haller and Wolff in terms of their differing concep-
of his theory and had overemphasized the vis essentialis to the tions of the nature of scientific explanation.
exclusion of the rest· of his ideas. "One could therefore have The key parameter in Hailer's philosophy of science was
omitted it and could have attributed the movement ofliquids to certainly his belief in God as creator and ruler of the world. All
another cause, as one wanted; or one might have supposed no of his scientific work has reference to this guiding notion, and
cause for it, and left the movement unexplained; so this move- religious themes are clearly evident in Hailer's approval or
ment of liquids would itself not be denied; and the manner of disapproval of his predecessors and contemporaries. Hailer's
production and building of parts, as the main point in a theory admiration for Boerhaave and Newton, his comradeship with
of generation, would have then still remained the same" ( 1789: Bonnet, his criticisms of Buffon and Voltaire, his acrid contro-
50 n.). Haller made too much of the force as a name and versy with La Mettrie, his rejection of Wolff's embryological
neglected the theory of development based on the movement of work, all turn on the theme of reverence for God and condem-
fluids as a total process. Wolff makes a similar point in an nation for atheists and materialists.
unpublished note where he objects to Blumenbach's having When we turn to Wolff, we find a far different situation.
compar~d the vis essentialis to his own formative force (nisus Wolff was not an atheist; he clearly believed in a divinely created
formativus). "Does this most illustrious gentleman not see then," universe. Yet Wolff saw no threat of atheism in science. As he
Wolff queries, "that the motion .of humors through a plant is argued in response to Haller, preformation.would certainly be
one thing, whereas the formation of a plant is something else? an· excellent proof of the existence of God, but its opposite,
And that therefore the force that moves humors is different epigenesis, did not lead to a denial of God's existence at all. For
from the formative force? Does he not see that by supposing the where do the laws and forces used to explain generation have
motion of humors I do not suppose formation, and that by their origin if not in God's wisdom? Wolff's belief in a lawlike,
supposing a moving force I do not suppose a formative force?" logical universe, that could be understood through deductive
(Wolff 1973:255). The term "essential force" could even have reasoning, the principle of sufficient reason, and the other
been eliminated from his theory, Wolff maintains, and his ex- tenets of rationalism, allowed him a certain freedom, as it were,
planation of development on the basis of movement and solidi- from the more confining, more personal God of Hailer's be-
fication of fluids would have remained unchanged. "The handle liefs. One is reminded of the Leibniz-Clarke controversy, where
to this calumny, which, however, I endure with good spirit, was Leibniz's more· metaphysical God, as creator of a universe based
given once by the distinguished Haller,. who wrote that I derived on necessity, is contrasted with the intervening, more closely
formation from some force that I called 'essential,' as if the ruling God of the Newtonian Samuel Clarke (see Clarke 1715).
whole matter hinged on this force and the whole expfanation of The differing attitudes of Haller and Wolff on the role of
formation consisted of this giving of names. After this, a horde God in the world and on the danger of atheism resulting from
went on to follow this great man" (1973:255). scientific theories are reflected in their contrasting views on
spontaneous generation. One of Haller's initial reactions to
THE NATURE OF BIOLOGICAL EXPLANATION
Wolff's theory of epigenesis, expressed in his revi_ew of Wolff's
dissertation, was that Wolff showed "almost a Needham-like
My purpose in this chapter has been to contrast the philosophi- opinion" in ascribing de~elopment to an essential force ( 1760:
cal views of Haller and Wolff and to illustrate the role these 1227). In a letter to Haller, Wolff responded to this, claiming "I
differing perspectives played in their debate over embryologi- would hardly think that the opinions of Needham could gain
cal development. I have tried to show how each person's philo- strength from my work, illustrious one, since he dealt with
120 Matter, life, and generation The philosophical debate 12 l
another matter entirely different from mine" (Haller 1773-75, anything blind and devoid of intelligence could be capable of
5:85; letterof29December 1761). Needham, Wolff points out, forming animals according to foreseen purposes, suitably ar-
dealt with the generation of animalcules from decaying matter, ranged for filling their place in the chain of beings" ( 1757-66,
whereas his own work endeavors to explain normal generation 8[ 1766]: 112). 19 At no level of simplicity or complexity could
in plants and animals. "But the remaining obscure metaphysical Haller allow for the operation of material forces as sole creator
speculations, which Needham adds without any experiments, ofliving organisms. O'Epigenesis," like its counterpart spontane-
merit hardly any attention, in my opinion at least" (p. 86). ous generation, "is totally impossible" (p. 14 7).
Haller must have challenged Wolff further on this issue in a The different attitudes of Haller and Wolff toward religion
letter in response, arguing that even if Wolff's work is different and spontaneous generation find expression also 'in their con-
from Needham's, Needham's ideas are consistent with Wolff's ceptions of biological forces. In all of Hailer's discussions of
and one can, Wolff r~ports from Haller's letter, "prove them embryological development- in his critique of Buffon's theory,
through my [Wolfffs] principles as through new grounds" in his rejection of Needham's and of Wolff's-he expressed the'
(Wolff 1764:31). Wolff answers, "Against this I can in a fair same attitude toward material forces. They are the mechanism
manner object.to nothing. For it is true. And in fact the words of development, yet they are not responsible, on their own, for
cited [from Haller's review and letter] hold nothing further in the formation of the new organism.
them than that through my theory, if it is correct, Needham's As presented in 1789, Wolff's views on "building forces"
opinions also receive at the same time a great probability" (p: 31 ). expressed much the same sentiment as Hailer's: building forces
Wolff seems almost nonchalant in his admission of a consis- simply do not exist. Forces must be simple in nature and must
tency between his and Needham~s views on generation. Needr operate in the same invariable manner. A building force, Wolff
ham was indeed an epigenesist, and for this Wolff would claimed, would have to vary its mode of operation according to
certainly have approved of Needham's approach. Distinguish- variable intended outcomes. Yet this kind of activity would
ing his own specific theories from those of Needham, however, violate the principle of sufficient reason, because there would
Wolff makes no further effort to condemn Needham's work. be no reason why the building force would act in one manner at
This is a far cry from Haller, for whom the mere hint of one time, in another manner at another time. Thus, Wolff
spontaneous generation was an anathema. That matter could in agreed with Haller that forces do not, by their actions alone,
any sense fashion itself spontaneously into a living creature, produce the formation of a new organism.
however small or primitive, was unthinkable for Haller, for it Although their positions on forces were similar in that they
contradicted all of his fundamental beliefs about the creation of both rejected building forces in favor of simple mechanical
life. Even when he was himself an epigenesist, Haller had ex- forces, Haller and Wolff exhibit. differing motivations for their
pressed grave doubts, in his <l:iscussions of Buffon's work on views. The context of each person's conception of forces varied
generation, about Needham's idea_s. Later, as a preformation- in significant aspects, involving contrasting views on the nature
ist, Haller repeatedly identified Needham's views on spontane- of scientific explanation. For Haller, the overriding concerns in
ous generation with epigenesis, for both represented the same explaining natural phenomena were empiricism, mechanism,
thing to him; both allowed matter t9 form living organisms out and religion. An explanation must be derived a posteriori from
of unorganized material, completely on its own. In the Elementa observational and experimental evidence; if it involves forces,
physiologiae, Haller rejected Needham's views on much the same these must be mechanical in operation; and finally, one's ex-
lines as he did Wolff's. Referring to Needham, Haller charged, planation must exhibit rather than challenge the role of God as
"there is in hi,s experiments something that conflicts with my designer and ruler of the world. Hailer's Newtonian mechan-
reflections: There is a corporeal force that, alone and without a ism and his deeply religious beliefs, also Newtonian in signifi-
parent, produces filaments and even spontaneous animals from cant respects, joined together to. produce a clear conception of
an inorganic paste.... It appears ·to us extremely difficult that how s~ience should explain the phenomena of the world. Haller
122 Matter, life, and generation The philosophical debate 123
rejected the materialism of La Mettrie and the animism of Stahl solution implicit in Wolff's theory, based on his concept of
and Robert Whytt on just these grounds. And he rejected epi- "vegetation" and his proposal, in 1789, that circumstances are
genesis, in all its guises, from the fermentation model of Des- responsible for the outcome of the essential force's actions.
cartes, to the spontaneous generation theory of Needham, to These ideas are not discusse<;l in any detail by Wolff in his
the vis essentialis of Wolff, as violating the criteria of a proper published works, but they are more clearly spelled out in un-
scientific explanation. published materials that concern the nature of heredity. In
For Wolff, on the other hand; the key aspect of explanation dealing with the question of how traits are passed on to descend-
was rational'ism. Opposed to mechanical reductionism, to vital- ants, and of the existence of varieties in the structures of organ-
ism, and to explanations resorting to Divine Omnipotence, isms, Wolff reveals his solution to the problem of embryonic
Wolff sought to explain life processes in a manner consistent organization, a solution that was once again, as we shall see in
with his rationalist beliefs. Wolff saw a logical universe, de- the following chapter, based on the principles of rationalism.
signed by a rational God, that man could come to understand
using the proper philosophical method of investigation. Wolff
certainly used both observational evidence and mechanical
fo,rces in his explanations, yet these were placed in a framework
of deductive reasoning, grounded on the principle of sufficient
reason. Wolff's was a more a priori approach than Hailer's, at
least ·more explicitly so, as was the case with his precursors
Descartes, Leibniz, and Christian Wolff. He rejected Rrefor~a-
tion, as he did "mechanical medicine" and Blumenbach's Bil-
dungstrieb, all on the basis of their not fulfilling his criteria for
proper explanation. And he admired Descartes as having been
the only other person to have offered a bona fide, though false,
explanation ror embryological development. Wolff's quest to
create the first "rational anatomy" thus found expression in his
epigenetic theory, for through his model of development and
the vis essentialis Wolff believed he had found the sufficient
reason for generation.
Yet I must raise one further question with regard to Wolff's
theory. Did his explanation of development, even as presented
in 1789, solve the problems that Haller had raised with regard
to both Buffon's theory and Wolff's? Did Wolff's final model
really account for the source, of embryonic organization? We
saw that Wolff agreed with Hailer's position on forces - that
building forces could not exist- and that Wolff defined his own
essential force so as to avoid its being thought of as a building
force. But if the vis essentialis is not responsible for the process of
development, what is? What guides the gradual formation of
the embryo? Wolff is not explicit in his answer to these ques-
tions, for nowhere does he address himself directly to the
source-of-organization dilemma. Yet there is an avenue of
Wolff's later work 125
il
ft
5
Woljf 'slater work on variation
and heredity
The subject of monstrous births interested Wolff throughout interfere with God's preordained program for development.
his career. He included a brief section on monsters at the end of i
~
And on the other hand, if monsters are preordained, this chal-
his dissertation ( 1759: 134-35, § 262) and published three sepa- lenges the wisdom of God, for why would he create malformed
rate papers on the subject in the St. Petersburg Academy journal
126 Matter, life, and generation Wolffs later work 127
academician Peter Simon Pallas, who participated in a number quently, "species is something in the plant that cannot be ob-
of natural history expeditions to different parts of Russia in the served and that is altogether different from form and structure,
late 1760s and early 1770s (see Stresemann 1962; Uschmann because, together with the climate of St. Petersburg, it produces
1962 ). When o~e transports plants from Siberia to St. Petersburg in St. Petersburg the form of St. Petersburg, whereas together
and vice versa, Wolff reports, one often observes no changes at ·With the climate of Siberia it produces in Siberia the form of
all for- several generations. But then gradually the plants begin Siberia, etc." (p. 171, § 35). Wolff thus defined species in such a
to change until they become almost unrecognizable as their way that _it was totally separate from the inward or outward
Siberian forebears, Yet, Wolff maintains, the species of plants structure of the organism. This structure is produced by the
do not change, only their outward structure. Furthermore, he species (in conjunction with environmental conditions), but
argues, neither the Siberian. nor the St. Petersburg form is more there is no aspect of the organism's form that is essential to the
"natural" than the other. Both are the product of the plant's ~ species. Consequently, any part of the organism's structure is
species as it is affected by differing environmental conditions. variable, and no traits are more closely related to, or more
Wolff bases his claim that the plant's species does not change expressive of, the organism's species.
even though its structure does on the following evidence: Having distinguished species from varieties, and having
Whenever you transfer a plant from one climate to another, he shown that an organism's species cannot be equated with its
relat~s. it always changes its structure in such a manner that it structure, Wolff turns to his model of vegetation to explain
will remain distinguishable as a separate species. "Therefore," what the nature of species really is. In the vegetative body, he
Wolff maintains, "the species, even if the former figure of St. maintains, one may discern three processes: vegetation, f!IOde
Petersburg does not remain, nevertheless produces another of vegetation, and degree of vegetation. As he explains, "Vege-
figure in Siberia that is peculiar to it as its effect, and in this way tation produces the 'vegetable,' or the plant as such; the mode
it distinguishes itself both from the other congeneric species of vegetation produces orders, genera, species, and classes; the
that remain· in St. Petersburg and from the effects of these degree of vegetation produces varieties" ( 1973: 168, § 34).
species ... that have been transferred to Siberia as well- no less Vegetation is the actual process of production of structure in
than this species distinguishes itself in St. Petersburg from its the organism, and it depends primarily on the distribution,
congeneric species" (Wolff 1973: 166, § 32). . secretion, and solidification of humors in the developing organ-
Conversely, if a Siberian plant is brought to St. Petersburg ism. The mode of vegetation relates to the qualities vegetable
and, after a few generations, it changes into a form that is matter possesses and thereby to the organism's species. The
identical to one found normally in St. Petersburg, we should not degree of vegetation pertains to the size and quantity of struc-
conclude that one species changed into another. Rather, Wolff tures produced and is affected by changes in living conditions,
argues, we must conclude that we had formerly mistakenly particularly climate. It is through degree of vegetation that
believed that a Siberian and a St. Petersburg plant were separate varieties are produced. ·
species, whereas in truth they are simply varieties (1973: 167, The mode of vegetation, which is responsible for the exist-
scholium to§ 32). "It is clear therefore," Wolff concludes, "that ence of species, depends primarily on what Wolff terms materia
species, genus, etc., are one thing, whereas external form or quali.ficata vegetabilis ("qualified vegetable matter"). Tile quali-
structure are another; that species is quite different from this ties that vegetable matter possesses, he explains, are "attributes
structure or form; that each one of this pair of things has been proper to the vegetable substance by which the forces of vegeta-
badly confused in all natural history and also in the natural tion are variously determined and which therefore have an ,
sciences; that species, whatever it is, is the cause, whereas struc- essential influence on vegetation" (1973: 158, § 18). It is through
ture and form are the effect; and that species itself escapes us these qualities of vegetable matter that the organism produces
and our sensations, while only the forms - its effects - are the structures peculiar to its species. "There exist qualified
evident and observable" (p. 167, § 33). The species of a plant vegetable matters," Wolff claims, "because all matters that v_ege-
does not change, even in the production of varieties. Conse- tate; vegetate in a peculiar and determinate mode so long as
130 Matter, life, and generation Wolff's later work 131
they produce a peculiar and determinate organic body through die change in structure, for they have no influence on the
vegetating, and no plant exists that continues to vegetate, fir.st in qualified vegetable matter. "It was not therefore the climate or
one way and then in another" (p. 169, § 34, scholium 1). All soil, if you want to hit the nail on the head," Wolff claims, "but
organisms, even when they change structures after a change in the plant in i~ species, its qualified vegetable matter, that,
environment, continue to produce that structure proper to disturbed by the soil and climate, produced a new form and
those conditions. Never do changes occur arbitrarily. "There structure.by vegetating" (1973:181, § 7). The qualified vege-
exist qualified vegetable matters," Wolff maintains, "which can table 'matter of the plant remains unchanged when changes in
vegetate' but which can each of them vegetate only in its own. outside conditions occur. It responds by producing new struc-
peculiar mode" (pp. 168-69, §. 34). tures that are better able to cope with the new soil or climate.
Wolff points to solidifiability as the most important quality For example, if a plant is moved from rich soil to poor soil or
vegetable matter possesses. Differing rates and capabilities of vice versa, changes in the size of its vessels will result, as the plant
solidification had been used, we can recall, by Wolff in his adjusts to receiving more or less nutriment. Yet this adjustment
earlier works to explain such things as why animals have hearts is not a direct response of the vessels to the soil; rather the mode
and plants do not. He does not elaborate further in the "Objecta of vegetation responds to the changed conditions by producing
meditationum" on what the different qualities of vegetable altered vessels. .
matter may be, maintaining only that through the qualifications Those who hav~ identified the structure of organisms with
of vegetable matter, the process of vegetation (that is, the dis- species, Wolff argues, have been in error. "Now therefore it is
tribution, secretion, and solidification of humors via the essen- certainly true," he explains, "that genera and species exist if you
tial force) produces different structures in different organisms. understand by species and genera nothing else but what is
"Vegetability is so c-onnected with qualification," Wolff asserts, constant and immutable. This will be false if you place that thing
"that neither unqualified vegetable matter nor qualified non- which is constant and immutable in the figure and structure."
{
vegetable matter could exist, ... perhaps qualification exists in All aspects of external and internal structure are changeable.
vegetability the way that an attribute exists in a thing" ( 1973: 178, "Similarly," Wolff continues, "it is true that varieties alone are
§ 36, scholia to scholium 3). · ' mutable if you attribute the whole external figure (and the
• Mode of vegetation, or the qualities of vegetable matter, is whole internal structure) to the varieties. But this in turn will
responsible for the production of those structures that are likewise be false if after the fashion of the Botanists you under-
proper to the organism's species. But, Wolff reiterates, those
structures themselves are not the species; rather, the qualifica-
t stand by varieties certain marks a~d characteristics in the ex-
ternal form taken apart from other marks in the same form,
'I
tions of vegetable matter are. As he expresses this: since whatever is observed in the form ... is equally mutable
Now since we have seen ... that the mode of vegetation and the and able to be propagated" (1973:202:...3, § 37). Constancies
qualities or forces of qualified vegetable matters are those causes by appear in structures, Wolff asserts, only because environmental
which those struc.tures and forms are produced that are usually conditions are common or remain the same, not because these
wrongly taken to be species or genera and orders, it is also clear that structures constitute signs of the organism's species. "There-
those very qualities or forces by which vegetable matters are qualified fore," Wolff declares, "the genera and species of the Botanists
(or, if you prefer, modes of vegetation) are, properly speaking, those are reduced to nothing" (p. 203, corollary to§ 37).
things that we obscurely sense under the name of species, genera,
orders and that we are so very well conscious of having observed to be Even though all structures are potentially alterable on Wolff's
constant. r1973: 171, § 36] theory, we must remember that these changes occur only in a
determinate manner, tha(Js, all organisms of the same species
It is the materia quali.ftcata, or mode of vegetation, that is to be change in the same way. New species therefore can never arise
. identified with species, not the external form of the organism. from varieties:
When varieties are produced, as in the case of plants that Although now the whole organism is mutable and able to be propa-
change their structures in response to a change in climate, the gated (conditionally able to be propagated, so that it can only be
changes in environmental conditions themselves do not cause propagated so long as the mutating causes continuously remain the
132 Matter, life, and generation Wolff's later .work
I
are constant can in no way be established for this reason, and that the structure. (In fact, in the original manuscript text of this sen-
new structures that are therefore formed by the monster-making tence, the word "botanical" is written in, but crossed out, as a
potency, however much they depart from the usual structure, are not modifier of "species.") Thus, when he says that change.s in
constantly but only conditionally able to be propagated. r1973:206, structure "continuously produce new species, as it were, from
§44]
When a variety is produced by the qualified vegetable matter in
the kingdom of possible species," he is arguing simply that there I
response to changes· in living conditions, that variation will
continue in the variety's offspring only as long as the mutating
are an unlimited number of varieties in structure that can be
produced from the realm of possible structures. Therefore, I
varieties of one species do not approach the form of those of
causes remain the same. As soon as these causes cease, the struc- another species. Yet the species of each organism remains con-
ture of the organism will revert in its offspring to its original stant, because on Wolff's own definition, species is not exhibited
structure. Never is a new species created, because the qualified by structure.
matter itself is not altered by changes in external conditions. Wolff's model of development through vegetation thus pro- ·
The distinction between species and varieties is thus that be- vided him with a basis for defining species and distinguishing
tween the qualified vegetable matter and external living condi- them from varieties, and for formulating a distinction betw~en
tions. "Without doubt," Wolff asserts, "the usual structure is a internal cause and external form that parallels the modern
product of the qualified vegetable matter joined with the usual genotype-phenotype dichotomy. Through the way an organ-
kind of life. Monsters, in contrast, and all other varieties, are ism develops and forms its structures, it becomes and remains.
products of the same qualified vegetable matter joined with an an identifiable member of a species. Built into this process,
unusual or monster-making kind oflife" (p. 207, § 44). however, is ari adaptability that allows the organism to produce
Thus, no matter what chang~s are produced in the structures variations in its offspring in response to changed conditions.
of an organism's offspring, be they varietal or monstrous, none Yet the new offspring retains the same qualified vegetable mat-
occur equivocally, that is, indeterminately. Furthermore, Wolff ter and the same characteristic mode of developing that its
reiterates, changed organisms do not alter themselves in such a parent possessed, and thus it remains a, member of the same
way that they begin to look like another species. In discussing species. A
this point, Wolff makes a statement that has misled some com-
mentators to allege that he believed in the mutability of species,
or at least in some kind oflimited transformism. 4 Wolff remarks: Monsters and hybrids
Although the mutations that different climates and kinds of nutri- Wolff distinguishes between monsters and varieties on the basis
ments or different degrees of vegetation and quantities introduced
into any vegetable body are themselves determinate and peculiar to of the quantity versus the quality of the nutriments. Monsters,
that body, nevertheless determinate mutations do not for this reason he claims, possess structures that are eit_her too large or too
arise so that a new structure of a plant introduced by mutation may small, or they have too many or too few of the same part. "A
return to or at least approach hence some one of the settled species. man may have two heads, but they will be altogether human; he
Rather, as the new structure goes away from and departs from the may have ,three arms, four .feet, six fingers, two skulls, three
former structure in varieties and monstrosities defined up to now, so eyes, two noses, but always these parts will be human in a man,
similarly it would go away from and depart from all other species and bovine in a calf, and ofa chicken in a chicken; and there will
therefore continuously produce new species, as it were, from the
kingdom of possible species, since mutation has gone so far that the never be any part in monsters, for example in human monsters
former character has been extinguished by it and a new one intro-. . .. that is not a conflation of two portions of two parts, in which
duced. And species is in the usual fashion placed in the structure of portions you recognize the same old ordinary human struc-
the plant or indicated by its structure. [ 1973:234, § 90) ture" (1973:229, § 80). This occurs in plants as well as in ani-
In the final sentence of this discussion, Wolff makes it clear that mals, for in all organisms monstrous structures are produced
he is ·using the word "species" in what he calls the traditional sirri ply by an unusual quantity of nutriments.
t •
1 34 Matter, life and generation Wolff's later work 1 35
the correct number, but they are formed in a different wa}, be it
with a different structure, color, or the like. "An Ethiopian has a
black color distributed through all his skin; his bones are yellow,
his blood is black. In the usual way there is only one nose, but it is
formed differeritly. 5 •• : [In plants] The Siberian variety has its
own peculiar properties distributed through all its parts. In this
r way there are the varieties of the forests, of the field, and of the
gardens" ( 1973:230, § 82). Furthermore, one and the same
organism may possess both monstrous and varietal structures,
and they are not always easy to tell apart. Yet their origins are
different, for "As all monstrous structure should be ascribed to
the degree of vegetation and to the quantity of nutriments
proportionate with the digestive force, so in contrast every
variety depends upon the mode of vegetation (upon the mode
without doubt of production) and upon the quality of the nutri-
ments, or upon the foreign substances that are present in
nutriments" (p. 231, § 85). _
Wolff's terminology here is somewhat inconsistent with his
earlier distinction. between mode and degree of vegetation as
being that between the production of species and the produc-
tion of vari<;ties. Now he is ascribiug only monsters to the
degree of vegetation, and moves varieties to the mode of vegtta-
tion. Although this is not consistent usage, we should not be led
to any confusion about the distinction between species and
varieties. Had Wolff actually polished his treatise for publica-
tion, the inconsistency would most likely have been cleared up.
Suffice it to say at this point that what he means by the mode of
vegetation's being responsible for the production of varieties is
simply that it is through qualitative rather than quantitative
influences that varieties arise. These qualitative changes in
living conditions have no effect on the species of the organism,
only "upon the mode without doubt of production." There is
certainly room in Wolff's concept of the mode of vegetation for
both the unchanging qualified vegetable matter and the mode
Figure 19. Wolff's drawing of Siamese twins from his manuscript materials. (From of production, that is, vegetating that actually gives rise to an
Wolff, Ttorija zarozlidtnija, 1950)
organism's structures. By distinguishing this from the degree of
vegetation, Wolff sought to underscore his differentiation be-
tween monsters and varieties on the basis of the quantity versus
the quality of their structures.
Varieties arise, on the other hand, from different qualities of Almost all variations and monstrou!! traits are inheritable
nutriments. In varieties, the organisms's individual parts are only in the sense that if the offspring experiences the same
136 Matter_, life, and generation Wolff's later work 1 37
conditions that produced the vanat1on in its parent, it will proper sort of organism results from propagation. Offspring
produce a structure similar to its parent's. If the conditions resemble parents because they share similar qualified matter
and because they experience similar living conditions; offspring
causing the ·original mutation change, for example, if the j differ from parents because their qualified matter produces
organism is returned to its original climate, it will produce
structures that no longer resemble its parent's. These kinds of I altered structures in conjunction with differing environmental
variations Wolff calls "unprescribed" or "free," and they make
up the largest class of variations. Yet there is another group of
mutations that are inherited as such, irrespective of external
conditions, and these Wolff terms "prescribed." The principal
I
l
l
f
influences.
example of this kind of variation is hybridization. When two f. Wolff's views on the nature of species, variation, and heredity
organisms of different species ·produce an offspring, that off- are of sufficient novelty for us to inquire what relationship they
spring receives a heterogeneous qualified vegetable matter and may have had to those of Wolff's contemporaries. First of all, it
exhibits structures that are produced by qualified vegetable is clear that Wolff's notions place him squarely in the "essential-
matter from both parents. Other examples of prescribed varia- ist" or typological camp of species concepts (Mayr 1957, 1968,
tion are, according to Wolff, grafting, where a part of a plant' 1969), according to which species are defined by certain essen-
from one spe~ies is made to grow from the stalk of another, and tial characters, which may or may not be morphologically ex-
the case of an organism being nursed by another, for example, pressed, but which remain constant amid accidental variations.
an Ethiopian woman nursing a European child. The plant that Within this group one c3:n also place John Ray, Joseph de
results from grafting will be prescribed to produce structures Tournefort, Carl Linnaeus, and other eighteenth-century tax-
resembling both species, and the child, Wolff speculates, might onomists, who ·saw species as constant and varieties as imper-
in some way change its color. Finally, there are prescribed manent, and who based their systems of classification upon
monstrosities, such as sexdigitism, where a monstrous structure morphological distinctions between characteristic and acciden-
is inherited from one of the parents (or even from a grand- tal structures of plants. For Tournefort, the flower and the fruit
parent). In all cases of prescribed variation, the new organism's were most utilized in classifying plants; whereas for Ray, it was
materia qualificata is altered. This is the only way the qualified the seed, as well as the petals, calyx, and leaf arrangement (see
vegetable matter can ever be changed, for in unprescribed Tournefort 1694; Ray 1686-1704). For Linnaeus, several dif-
variations it remains unaffected. ferent structures were viewed as exhibiting specific characters,
In the case of hybrid organisms, Wolff admits, because the the sexual parts of the plant taking precedence as the basis for
offspring receives mixed qualified vegetable matter, its species Linnaeus's binomial nomenclature, the foundation of modern
is in that sense changed also. "Since only qualified vegetable taxonomy (see Linnaeus 1735, 1753; Larson 1971).
matter of vegetable bodies is constant, and since true species .Wolff's views on the nature. of species present significant
consist in this matter alone, it is clear that heterogeneous semen differences from those of Linnaeus and the oth~r taxonomists,
changes both the qualified vegetable matters of v.egetable bodies for Wolff was not particularly interested in classification. Al-
and therefore their true species" (1973:239, § 98). But, Wolff though his interest in plant generation is well established by the
cautions, two things must be kept in mind about this. First, amount of discus.sion he devoted to the subject, particularly in
hybrids tend either to be sterile or to revert to one of the his dissertation, where he formulated the concept of metamor-
parent's species when propagated. "Second," Wolff asserts, "it phosis (see Chapter 3), Wolff was not a taxonomist himself. It is
must be noted that hybrid plants or animals are not new species difficult to believe that a field naturalist would have formulated
but old established species that have now been mixed. There- a definitjon of species like Wolff's, which rests on necessarily
fore, nature does not go beyond prescribed limits in producing invisible characteristics. Wolff's separation of species from out-
and propagating vegetable bodies" (p. 239, § 98). The materia ward structure, his belief that all external characteristics are
qualificata is what provides these limits, what insures that the variable, and his identification of species with qualified vege-
""!
'
138 Matter, life, and generation Wolff's later work 139
table matter delineate him clearly from his contemporaries. hers of the same genus to split into separate species. 7 Thus, for
There is no doubt that when Wolff proclaimed "the genera and example, the organisms of the New World could be viewed as
species of the Botanists are reduced to nothing'' he had in mind degenerations from Old World species, the alterations in form
the taxonomists Tournefort, Ray, Linnaeus, and others. Their having been caused by environmental influences. Wolff quite
systems, relying on outward characteristics of plant structure, possibly knew of Buffon's views, even though his only reference
can never reveal the true species of the natural world. to Buffon in the "Objecta medit~tionum" concerns Buffon's
A further point must be noted about Wolff's views. In his theory of generation. Wolff, like Buffon, allowed environment
treatment of hybridization, Wolff allowed for the creation of to play a determining role in the production of plant and animal
new species, although, as we saw, he regarded this as only a form. Yet, unlike Buffon, Wolff saw climate as causing only
mixing of established species. Moreover, he pointed out, hybrid varieties to be produced, not species (although Buffon's dis-
organisms tend either to be sterile or to revert to one_ of the tinction between species and variety was never as clear-cut as
parent types when they reproduce. (This occurs especially in Wolff's). For Wolff, environmental conditions could never af-
plants that are back-crossed with one of the parent species.) fect the qualified vegetable matter and thus could never alter an
Linnaeus had also been aware of plant.hybridization and, in the organism's species.
l 7 40s and 1750s, hybridization phenomena led him to alter his Wolff's discussions of the nature of species and varieties show
original view of species as fixed and constant. He began to his awareness of other contemporary views, at least those of the
believe that God had created only the gen~ra and that all species botanists, even though his own views were not in agreement.
had been formed by crossings between each genus's original Wolff struck an independent course from plant taxonomists in
species and those of other genera. 6 Although Wolff never went propounding a theory that w;as even more "essentialist" than
as far as Linnaeus .in his views on the significance of hybridiza- theirs. His own definition of species in terms of qualified vege-
tion, both clearly recognized that hybrid organisms represent table matter owes more to Wolff's· rationalist heritage than it
altered species. Wolff was very likely aware of Linnaeus's views; ctoes to eighteenth-century taxonomy, for the matcria qualificata
for, even though he does not cite any of the works in which united Wolff's model of vegetation and epigenetic develop-
Linnaeus's hybridization theories are expressed, he does refer ment with his belief in the fixity of species.
to other works by Linnaeus in both the Theoria generationis and
the "Objecta meditationum." Yet Wolff's conclusions concern-
Aptness
ing the impact of hybridization on the fixity of species were far
more conservative than those of Linnaeus, who, in his final Wolff includes a rather lengthy section in the middle of the
theories, saw hybridization as the means whereby the earth had "Objecta meditationum" on the subject of "aptness." Here he
become populated with species. Other botanists adopted posi- explains why organisms are so suitably adapted to their en-
tions more similar to Wolff's; Joseph Koelreuter, for example, _, vironments and what the origins of adaptation are. If organisms
who dealt extensively with hybridization phenomena, saw no ;.t were apt for their living conditions through natural necessity,
threat in them -to the fixity of species. Most hybrids are sterile, l that is, through a causal mechanism based only on internal and
4';
Koelreuter noted, and those few that are fertile are not God's
but ·man's artificial creation (see Koelreuter 1761-66; Glass
1959c).
I!
:!_'
e~ternal conditions, Wolff notes, then this would mean that the
ends we observe in nature are imaginary. But, he warns, "Be-
ware of precipitating yourself into a most pernicious error I It is
Wolff's emphasis on the influence of climate and other en- by this argument alone, and by no other, that the most manifest
vironmental conditions on the structures of plant and animal ends in nature would also be destroyed, ends that alone, in my
organisms is also something that was discussed by his contem- opinion, provide us with a solid and firm argument for the
poraries. Buffon, for example, saw environment as playing a existence of God, and by the destruction of which the way would
fundamental role in species relationships. Although Buffon's therefore be opened to Atheism" (1973:180, § 2). If the suit-
ideas on species changed during his lifetime, he believed, in his abilities we see in the structures of plants and animals were the
final theory, that different living conditions could cause mem- result of simple efficient causes, rather than a final intelligent
140 Matter, life, and generation Wolff's later work 141
cause, Wolff admits, then no longer would the design we wit- Vet they are fully consistent with his earlier expressed beliefs,
ness in nature be evidence of Divine Wisdom. for they further demonstrate his view of a rationally ordered
For something to be "n~cessarily apt," Wolff argues, the thing - universe, operating on the basis of natural causes. As he stated
for which it is apt must be the cause of its suitability. For in one of his letters to Haller, ascribing epigenetic development
example, the formation of teeth, and the arrangement of in- to natural laws does not threaten the existence of a Creator,
cisors and molars, would have to be the product, in some sense, because these natural laws themselves must have their origin in
of chewing. But this is not the case. Rather, Wolff claims, the God. That all organisms have been allotted the ability to adapt
wide structure of molars and the sharpness of the incisors are to changing environmental conditions, through a mecha~ism
produced by the fact that molars are secreted from a thick of natural causes, is further proof of Divine foresight. The
arterial trunk, whereas incisors are secreted 'from the extreme organism's qualified vegetable matter, its species, provides the
end of the artery. The teeth that form thus necessarily possess means through which this. adaptability, based on an intelligent
these structures. They are also apt for their functions. But they cause, is expressed. Variation thus serves a purpose in nature,
are not neccessarily apt; their formation must be due to an one based ultimately on the wisdom o_fGod.
intelligent cause ( 1973: 192, scholium to§ 31).
The most convincing evidence for the role of Divine foresight
THE "DISTRIBUTIO OPERIS"
in the formation of organisms, Wolff claims, is their ability to
vary their structure when environmental conditions alter. "And Among Wolff's unpublished papers, there exists a short manu-
now, as I promised," Wolff remarks, "you see a stupendous script that has bearing on the subjects treated in the "Objecta
miracle of Divine Acuity. Not only has an apt fabric been given meditationum." Titled "Distributio operis," this document pre-
to plants to correspond to the soil, the climate in which each sents a plan for the part and chapter contents of Wolff's major.
lives, and the country to which each belongs ... but also a fac- work on monsters. Part 1 was to be an anatomy of monsters that
ulty has been put in these plants of such a nature that if it ever would presumably have included the several. anatomical de-
were to happen that the plant would be taken away from its scriptions of monsters that Wolff had already completed. Part
country and transferred into another that is very different ... it 2, titled "Physiology, or on the Nature and Generation of Mon-
builds a new form and structure for itself." The plant, however, sters," was slated to include chapters on the natural orders of
does not lose its species, for if it is returned to its original monsters, on the generation of animals, on the origins of
climate, it soon recovers its former struqure. "The same crea- monsters, on the nature of propagation (where, presumably,
ture," Wolff continues, "as it were, makes itself a new creature ·' the "Objecta meditationum" would have been used), on the
for each new country by nevertheless remaining the same as far souls of monsters, and on the purposes of monsters. 8 A monu-
as its nature is concerned. This seems to me at least to be mental work, Wolff's treatise could easily have stretched into
exceedingly much, and I do not think that I can marvel enough several volumes.
at it" ( 1773: 187, § 15). It is because organisms possess qualified '
t·' The description of Wolff's chapter on the natural orders of
vegetable matter, which allows them to alter internal and ex- tl monsters is the most extensive section of the "Distributio operis."
ternal structure in response to changing living ccmditions, that
they show such aptness for their environments. Not only plants,
ii The principal question Wolff proposes to examine is whether
monsters exhibit the same kind of order that is found in normal
but animals and human beings exhibit this adaptability as well. organisms. In nature, Wolff asserts, there is a remarkabie com-
At one point, Wolff likens the apparent foresight that is uni- bination of change with constancy. In monsters, he notes, one
versal among organisms to "souls," and he includes human observes this as well. Although there seem to be as many dif-
judgment as an apt suitability (pp. 189-90, §§ 21-25). ferent abnormalities as there are different structures, similari-
Wolff's v!ews on adaptability as expressed in the "Objecta ties are evident. For example, Wolff observes, one-headed
meditationum'' show us a fuller picture of his attitude toward monsters often resemble each other in having two hearts and in
God and design in nature than is evident in his earlier works. having similarly constructed throats, digestive canals, and brains.
142 Malter, life, and generation Wolff's later work 143
"From [this] one must conclude," Wolff maintains, "in monsters process. God created in organisms the capability of reproduc-
of nature, as in the genera of animals and plants, changes ing in their offspring both their traits and their ability to adapt
combine with preserved similarities in the same remarkable and these traits to their living conditions. This takes place through a
no less miraculous way" (Gaissinovitch 1961:526-27). And al- process of epigenetic development based on the organism's
though monsters do not tend to live to an -age at which they vegetative powers.
could reproduce, if they did, their traits would be transmitted to I have discussed Wolff's "Distributio operis" in soine detail
their descendants, :which would "become species in future cen- because it has been used by commentators in the past to docu-
turies" (p. 527). ment Wolff's alleged belief in evolution (Rajkov 1964:610-12).
Wolff now ·asks, on what does the observed constancy in Others have argued that Wolff believed in a limited version of
nature depend? He answers, as we would expect; that the simi- transformism, based on mutability of species within fixed gen-
larities that species exhibit "have a sufficient reason of their own era (Gaissinovitch 1961:449-50). Yet I do not believe that
constancy in the vegetation or vegetation impulse imprinted as Wolff's views constitute either evolution or limited transform-
if by God in the vegetative substance and which are transmitted ism. Wolff allows for the transmission of mutations to offspring
to th~ir descendants" (p. 527). No other vegetation can be only when the materia quali.ficata, the process of vegetation that
transmitted to offspring. The purpose of this constancy is thus governs the formation of structures, is affected. But this hap-
"the preservation of the species, genera, orders, and the original pens in very rare instances - in hybrids and in some kinds of
forms, as intended by God" (p. 528). m~nsters (most of which do not reproduce). Only these kinds of
Yet Wolff also discusses a second purpose of this constancy: variations are prescribed for the next generation; most varieties
the stabilization of new traits. If monsters could ljve to a repro- and most monsters arise from unprescribed variation, which is
ductive age, they would transmit their traits to their descendants. the result of the influence of external living conditions on the
Two heads would become "stabilized" as a trait just as six fingers organism. In hybrids and in those few abnormal traits, like
has been. Perhaps one can view monsters as nature's "trials," sexdigitism, that are propagated in a prescribed fashion, the
Wolff asserts; for if a new trait finds access to the vegetative altered characteristics of the offspring are indeed the result of
process, then it is transmitted. Sexdigitism is an example of this heredity. Yet hybrids tend to be sterile or to revert to a parent
having occurred, for a six-fingered child may be born even species when propagated. Furthermore, Wolff maintains, hy-
from parents with normal fingers if a grandparent possesses six brid organisms do not represent new species but rather mix-
fingers. The case of national traits among humans, however, tures of old ones. In the case of monsters, most never reproduce.
Wolff maintains, is not an example of different species having If they did; Wolff admits, their line could form a new species of
arisen. Rather, the facial characteristics of, for example, Ethio- . organisms. Yet this does not occur; only some abnormal traits
pians, are a result of the effect of climate and soil on vegetation. are inherited, and these do not constitute new SP.ecies. Thus,
"Therefore," Wolff concludes, "there are no necessities" Wolff's views do not lend themselves to a transformist interpre-
(Gaissinovit~h 1961 :530). Constancy of form is due to the per- tation. He certainly recognized the immense amount of vari-
sistence of vegetation, not to necessary development. "Thus," ability that organisms exhibit, in fact more than most of his
Wolff declares in opposition to the preformationists, "how un- contemporaries did; yet he retained a place for unchanging
likely it is that the endless legions of germs in the organic body, species through his concept of qualified vegetable matter.
already molded and manufactured, could have come from the
very hand of God. Apparently, the omnipotent God created WOLFF'S VIEWS IN RETROSPECT
only substances that were endowed with their own forces, not
apprehensible by our senses and unknowable, becoming ap- Although the material covered in this chapter was written by
parent only in .their activity" (p. 530). Rather than a static Wolff long after his debate with Haller and indeed after Haller's
progression of predetermined individuals, Wolff views the death, several aspects of Wolff's theory of variation and hered-
order of natural things as a product of a dynamic developmental ity shed light on the theory of generation put forward in his
144 Matter, life, and generation Wolff's later work 145
earlier works. Furthermore, these later views are consistent ism that led, as we saw in the case of Haller, to its heated
with Wolff's philosophy of science, as discussed in Chapter 4, rejection. (One has only to recall the outrage that greeted the
and indeed clarify some of its key issues. publication of La Mettrie's L'flomme machine in 1748.) Given this
Before attempting a synthesis of Wolff's views, let me treat contemporary definition, one cannot include Wolff among the
briefly an issQe that has arisen in the secondary literature on materialists. Even tliough he did allow a close tie between the
Wolff. In a section of another manuscript dealing with a mon- soul and the body, he never denied that the soul is immaterial;
ster consisting of two humans sharing a common chest region, and, furthermore, he allowed it a separate eternal existence,
Wolff includes a discussion of the relation of the soul to the destined for it by its Creator.
body that has led some historians to conclude that Wolff was a We should recall in this regard Wolff's discussion of God and
materialist. 9 In this passage, Wolff suggests that the soul arises· design in the "Objecta meditationum." There he argued that
with the body and that it is in some sense the "extract" of the the "aptness" of organisms for their living c<mditions shows that
cerebrum and the medullary substance of the body.just as the they were created by an intelligent cause and that the purposes
egg is an extract of the hen and the spinal column is an extract of we see exhibited in nature are not imaginary. Wolff believed the
the egg. (We should recall here that Wolff saw generation as a universe to be a rationally created product of Divine Wisdom.
process of ordered secretion and solidification: the mother Never does he suggest that matter alone could be responsible
secretes the egg in which the spinal column first solidifies; this in for all life phenomena or that only material substances exist.
turn secretes the limbs and other parts.) The soul is perfected, Wolff did not fear, as the preformationists did, that explaining
Wolff maintains, during the life of the body, through its inter- the generation of living organisms on the basis of natural causes
, action with bodily substances and forces. Yet when the body might lead to atheism. Wolff's logical universe could not pos-
'dies, the soul "throws itself off from the body and continues to sibly be the product of material necessity but could result only
live after the body's death, just as we see offspring from other from a rational God.
parts of the body live and vegetate after its death. And now As I remarked at the close of Chapter 4, even in Wolff's 1789
eternal life is enjoyed, to which it was-destined by the Creator. publication on the essential force, one problem remained un-
For our present life is not the true life of the soul, but only its answered, the source of embryonic organization. Arguing
generation and formation; the true life of the soul is this, against the existence of "building forces," Wolff maintained
whereby, liberated from its body it is enjoyed, and which is t_hat his essential force was a simple force that acts in the saine
eternal" (Wolff 1973: 289, §§ 7, 8, 9·). manner (attraction or repulsion)Jn all circumstances. Conse-
Although Wolff seems to allow for the creation of souls with quently, the essential force can in no way be responsible, on its
the formation of each individual, the soul remains separate own, for the formation of the organism. Rather, the differing
from the body and is indeed freed from it at the body's death. In situations in which it acts result in the development of different
some sense the body contributes to the development and per- structures in the embryo. Yet Wolff did not explain how this
fection of the soul prior to its "true life," which is eternal. I do process is governed or why it results in the proper organism.
not see how one can conclude from these passages that Wolff It is in Wolff's unpublished views on variation and heredity
was a materialist, especially if one is careful to use the term as it that we find his answer to the source-of-organization problem.
was used during the eighteenth century. Principally, the epithet Through the materia qualificata, the determinate qualities that
"materialist" was applied to those who denied that an immate- plant and animal substances possess, the transmission of char-
rial soul coexists in human beings with the material body, and acteristics from parent to offspring is effected. Children re-
who sought instead to explain such thing as voluntary motion,
f semble parents because they possess similar qualified vegetable
,,!
sensation, and thinking on a material basis alone. "Materialist" r[ matter. And this materia qualificata determines the mode of
was often used synonymously with "atheist," because a denial of vegetation, that is, the course of development of the organism.
It is the qualities of the generative material, and the way they
a spiritual side to human beings challenged the existence of
God. It· was principally the anti religious overtones of material-
j
j
determine th.e direction of the process of secretion and solidifi-
146 Matter, life, and generation Wolff's later work 147
cation of structures, that govern generation. Thus, the proper . animals irritability and sensibility may also be fundamental
organism results from reproduction, and the species is pre- ·qualifications of animal matter. The qualifications of vegetable
served amid variation. . · matter, in conjunction with the actions of the essential force,
lt was Wolff's rationalist view of substance, which was mark- · thus provide the basis for vegetation, and thereby for life, in
edly different from that of many of his contempora1}es, that plant and animal organisms.
provided him with an avenue of solution to the source-of- Second, it would be a mistake ·to conclude that, in his final
organization dilemma. For many eighteenth-century thinkers, theory, Wolff resorted to preformation by allowing for some
matter was a passive entity activated only under the influence of sort of material substrate to be passed on from parent to off-
simple mechanical forces addeq to it by God. This concept of spring. Wolff saw his concept of "qualified vegetable matter" as
matter underlay Hailer's theory, as it did those of most prefor- providing the completion of his epigenetic system. Through it,
mationists in the eighteenth century (see Chapter 1). Develop- he could explain how the embryo develops from its own forces
ment had to be programmed in preexisting structures because ' withoµt resorting to preexisting structures. ~dmittedly, em-
blind mechanical forces acting on simple, passive matter could bryos on Wolff's theory do not start out in a state. of absolute
not possibly be responsible for producing a living organism. For homogeneity. Yet one must be careful not to define epigenesis
· Wolff, however, whose ideas developed within the tradition of so narrowly that clearly epigenetic systems like Wolff's are
dy"namism popularized by Christian Wolff, matter was not such excluded. Gradual development of complex heterogeneity from
a simple entity. Rather than passive extension, matter was simple heterogeneity can provide a valid epigentic viewpoint
viewed by Wolff as something that possesses form, qualities, (see Churchill 1970a: 169-71). In Wolff's system, the embryo's
modes, and attributes. initial heterogeneity is of a potential nature, based only on
Through his view of organism-as-qualified~substance, Wolff physical factors like solidification and a.ttraction and repulsion,
was able to give matter a role in generation where his mechan- which produce the structures of the organism through a gradual,
istic contemporaries could not, and thereby to explain why the but automatic, sequence of events. This is a far cry from prefor-
embryo develops as it does without resorting either to preexist- mation, especially in its eighteenth-century emboitement form.
ing organization or to self-guiding forces. For Wolff, not all Wolff's theory of epigenesis, and the philosophy of science
matter is alike; through the qualities it possesses its nature is upon which it was based, was uncommon for his era. Opposed
determined. Applying this to biology leads to Wolff's view of to mechanical reductionism., to vitalism, to materialism, and to
the organism: qualified matter determines the specific nature, explanations relying only on Divine Omnipotence, Wolff sought
the species, o,f the organism, just as the specific natures of all to explain life processes in a manner·consistent with his rational-
substances are determined, according to the principles of .ra- ist beliefs. Wolff's theories were based on mechanical princi-
tionalism, by their inherent qualities. ples, with their emphasis on secretion, solidification, attraction,
Still, Wolff's concept of qualified vegetable matter is a prob:. repulsion, and the like. But Wolff's mechanism was not based
lematic one that is not entirely clarified in his writings on the on passive particles in motion, to which life processes were to be
subject. Two things must be kept in mind, however, when reduced. Rather, Wolff's rationalist view of substance allowed
considering Wolff's theory. First, it is important to realize that for a more complex sort of mechanism, as such a view had for
the materia qualificata does not refer to an identifiable entity, Leibniz and for Christian Wolff, and thereby opened up avenues
located at some point in an organism, but rather to a condition of explanation that were closed to so many of Wolff's con-
possessed by living material. Vegetable substance possesses at- temporaries.10 By starting from a different philosophical base,
tributes through which it is capable of carrying on the activities Wolff's "rational anatomy" provided a unique challenge to
of vegetation, that is, through which vital processes are pro- eighteenth-century preformationism.
duced. Differing solidification abilities of plant and animal mat-
ter are the most important qualities of vegetable matter in
Wolff's system. Wolff speculates at one point that perhaps i?
Epilogue 149
i
;
between Haller and Wolff over embryological development can A further aspect of scientific controversy ~hat is exemplified
be fully understood only when one considers the philosophical in the Haller-Wolff debate is the tendency for such debates t9
presuppositions that underlay their controversy. Haller and remain unresolved. The myth of the experimentum crucis mani-
Wolff debated not only the formation of embryonic structures, fests itself here, for most controversies simply are not resolvable
but, more fundamentally, the views of scientific· explanation in terms of experimental evidence. Can one imagine, in retro-
upon which their embryological theories were based. Each per- spect, a single experiment or observation that would have set-
son brought to bear on the embryological points of contention a tled the. Haller-Wolff debate once and for all, that would have
whole host of philosophical asim·mptions toncerning mechani- convinced either Haller or Wolff of the truth of his opponent's
cal explanation, biological forces, logical reasoning, God's rela- position? I think not, for repeatedly in their controversy Haller
tionship to natural phenomena, and the like'- assumptions that and Wolff confronted each other with new, seemingly conclu-
fundamentally colored their respective perceptions of the em- sive evidence, only to find the opponent subsuming it under his
bryologicaf level of debate. At issue was more than simply the own explanatory system.
interpretation of specific observations of chick development, Closely related to the unresolvable nature of the Haller-Wolff
for it was Hailer's whole Newtonian outlook, with its religious debate is the element of incommensurability that is also evi-
and mechanistic parameters, that confronted Wolff's rational- dent. 2 At a number of points in the controversy, one has the
ist, antireductionist beliefs. This.is what led to such a heated clear feeling that Haller and Wolff really are "talking past" one
debate between the two, and what was ·responsjble for their · another, that neither is able to see the same issue, or the same·
controversy's inconclusiveness and its frequent aura of non- evidence, fully from the other's point of view. Although I do not
communication. · wish to argue that HcJ,ller's and Wolff's positions were totally
My thesis is not one that is unique to the Haller-Wolff debate. incommensurable (or that such a thing ever exists in science), in
There is no doubt that major controversies in scien,::e regularly a very real sense Haller and Wolff we-re "living in different
have their extrascientific components, be they philosophical, worlds," as each sought,.to promote his own conception of how
religio1,1s, or political jn origin. Furthermore, it is these under- science ought, and ought not, td explain biological phenomena.
lying, contextual elements that provide the key moti~ational I do not think, for example, that Haller ever showed any under-
forces behind scientific debates. In the history of biology, for standing of Wolff's rationalist approach, of his driving need to
example, there have been numerous such controversies, where subject empirical evidence to logical analysis and to demon-
observational; "scientific" issues provided the forum for clashes strate deductively how an explanation accounts for the facts.
between whole metaphysical systems. The debate between Har- Neither did Wolff ever share Haller's deeply ingrained distrust
vey and Descartes over the nature of the heartbeat is one such of any scientific theory that might threaten the omnipotence of
example, for Descartes's reduction of the heart's muscular con- God. Because of this, Haller and Wolff were never really able to
traction to a combustion model opposed far more than Harvey's see the empirical evidence from exactly the same perspective,
"pulsific faculty." In contention were two syster:ns pf biological
150 Matter, life, and generation ~pilogue 151
for neither ever fully grasped the implications that such evidence In the eighteenth century, most embryologists adopted pre-
held for the other's philosophical position. This is not to say that formation because it was the only mechanistic explanation of
there was no communication or no common ground of under- development that was consistent with the Qominant religious
standing between the two. Rather, their debate exhibits that world view. If one believes that matter is essentially passive,
limited degree of communication and understanding that is activated only through divinely endowed mechanical forces,
necessary for controversy to arise and flourish in the first place. then preformation is really the only viable explanation for
Both the unresolvable and incommensurable aspects of the development one can adopt. Such a viewpoint underlay the
Haller-Wolff debate are closely tied to my general thesis, for it arguments of preformationists from Malebranche through
was because their controversy took place in a much wider con-
text than that of embryological observation alone that it was so
inconclusive. Haller and Wolff were not merely trying to con-
vince one another of the "correct" explanation of blood vessel
Hal.Ier and Bonnet. Those who proposed mechanistic epigenesis
- like Descartes, Maupertuis, or Buffon - either failed to ac-
count satisfactorily for the organizational aspects of develop-
I
~
ment or, like Maupertuis, turned to nonmechanistic solutions
development; they were in' fact out to discredit each other's to obviate this difficulty. As I have argued in prevfous chapters,
philosophical view of scientific explanation. That neither suc- it was the source-of-organization problem upon which mechan-
ceeded in winning their debate is less significant than the fact istic epigenesis continually foundered.
that neither ever really could, because the rules of the game With Blumenbach and the other German embryologists, the
were so different for each. The Haller~Wolff controversy, like situation changed. radically. No longer was the source of em-
so many others in science, thus provides a "laboratory view" of bryonic organization the central problem to be explained.
the relationships between metaphysical commitments, theoret- Rather, organization became the one element of life that was
ical beliefs, and empirical observation as these operate in the taken for granted. Following Kant, these German embryologists
practice of science. uniformly embraced a teleological view of ~mbryological de-
velopment, based on a presupposed original state of organiza-
tion in the generative material. 3_ Agreeing with Haller and the
THE RISE OF TELEOLOGICAL EPIGENESIS
preformationists on one count, Blumenbach and his followers
Although the debate between Haller and Wolff was not resolved rejected the idea that organization could be explained in me-
- by the two of them, history s~ows that after the great preforma- chanical terms. As Kant argued, "Absolutely no human reason
tionist triumvirate of Haller, Bonnet, and Spallanzani, the ... can hope to understand the production of even a blade of
theory of preexistence fell into demise and epigenesis replaced grass by mere mechanical causes" ( 1790, 1966 trans.:258, § 77).
it as the most widely held view of embryological development. Referring specifically to Bluruenbach and his nisus formativus,
Iri the late eighteenth and early nineteen~h centuries, the most Kant remarked similarly, "That crude matter should have
important contributions to embryology came from Germany, '! originally formed itself according to mechanical laws, that life
I
where Blumenbach, Kielmeyer, Dollinger, Oken, Pander, von should have sprung from the nature of whiit is lifeless, that
·Baer, and others pursued developmental researches from an matter should have been able to ~is pose itself into the form of a
epig~netic point of view. Yet none of these investigators ever self-maintaining purposiveness -- this he rightly declares to be
really disproved preformation in any significant experimental ~ contradictory to reason" (p. 274, § 81). Once one accepts organ-
way; most simply rejected it out of hand. As Oken declared in
181 o, "The theory of preformation contradicts the laws of
t ization as a teleological fact, Kant proposed, one can then pro-
ceed to explain on a mechanical basis how this organization
natural development" (1809-11:28). That the German epi-
genesists were able to dismiss preformation in such a manner
tI functions and is maintained. Using the concept of purposive
organization as a regulative idea, as a guiding thread, one
points to a deeper cause than the evidence itself, for the philo- l should then conduct empirical investigations int~ the mechan-
sophical foundations of their biological views were markedly
i
isms that govern vital phenomena.
different from those of their eighteenth-century preformation- This is indeed what the German embryologists proceeded to -
ist predecessors. do, with important descriptive and comparative embryological
152 Matter, life, and generation Epilogue 153
studies as a result. Freed of the need to explain the source of their historical development, arose with the concept of the
organization, they rejected eighteenth-century preexistence archetype - the idea that all animal forms are simply variations of
theories as contradictory to the developmental laws of nature a limited number of ideal types - and with the first enunciations
and turned their attention instead to describing the sequence of of the biogenetic law by Kielmeyer, Oken, Meckel, and others
embryological development in accurate detail. One of the cul- (see Gould 1977). Not only was individual development seen.as
minations of this research was von Baer's enunciation of the the result of an immanent teleological power, but so too was the
germ layer theory in 1828 (see von Baer 1828-37). Yet one must history of life on earth. As Kielmeyer explained, "Since the
realize that these embryologists substantially altered the ground distribution of forces in the series of organized beings [organ-
rules from those on which their eighteenth-century predeces- isms] follows the same order as their distribution in the develop-
sors had based their embryological theories. Indeed, Kant mental stages of given individuals, it fo,llows that the force by
termed this new form of epigenesis a "generic preformation," which the production of the latter occurs, namely the reproduc-
built as it was on "preformed" purposiveness ( 1790, 1966 trans.: tive force, corresponds in its laws with the force by which the
2 7 2, § 8 1). The epigenetic viewpoint proposed by Blumenbach series of different organized beings of the earth were called into
and later German embryologists did not fulfill the criteria of existence" ( 1793:262). That ontogeny recapitulates phylogeny
seventeenth- and eighteenth-century epigenesis, for it relied on, seemed a natural consequence of the belief that the creation of
rather than explained, embryonic organization. As Temkin has both the individual and the species are under the guidance of
remarked in a similar vein, "the theory of epigenesis as advo- immanent, nonmechanical laws of nature. 4
cated in Germany around 1800, contained a strong teleological Such an explicit tie between embryological developmettt and
element and ~as, therefor:e, a compromise between preforma- natural history was absent in the eighteenth century. Although
tion (rejected in the form of eniboitement) and mere mechanistic , Bonnet had drawn a parallel between ontogeny and the perfec-
explanation" ( 1950:230, n. 16). Opposed to preexistence, these tion of species on earth (Gould 1977), and Buffon, in later
embryologists retained the most successful aspect of eighteenth- volumes of the Histoire nature/le, had sought to establish a con-
century preformation -its-assumption of built-in organization- nection between the history of life and environmental influences
while at the same time promoting a far different view of gradual on the moule intirieur (Farber 1972), the identification of onto-
development than that of their mechanistic forebears. genetic and phylogenetic development by German embryolo-
With the German epigenesists, ontogenetic development also gists and naturalists was based on a far different con~eption of
took on wider significance than it had during the preceding nature than had existed in the preceding century. Embryo-
century. Rejecting the search for a causal account of genera- logical research took on a new significance and was used in
tion, these embryologists viewed the formation of the individual answering new varieties of questions. Rather than seeking a
as part of a process of development encompassing the whole of mechanistic, causal account of the formation of the individual,
the organic realm. As Casking has pointed out, "Growth ac- the German epigenesists turned to embryological investigation
companied by change was now regarded as a fundamental to provide examples of nature's developmental tendencies, as
feature of the Universe, and the growth of living things was the well as data for understanding the taxonomic relationships
analogy in terms of which all other processes were to be under- among organisms. Descriptive and comparative embryological
stood. It, therefore, seemed a basic phenomenon requiring no studies were pursued with renewed effort and with remarkably
further explanation. What had been an atypical, almost miracu- fruitful results. Not until the end of the nineteenth century did
lous process from the seventeenth and eighteenth c.entury point descriptive embryology give way once again to causal embryol-
of view, became the paradigm of the natural for the [early ogy, with the experimental work of His, Roux, Driesch, and
nineteenth-century] nature philosophers" ( 1967: 151 ). One con- others, and, as a consequence, to a new round of debates over
sequence of this new "developmental paradigm" was the signifi- preformation and epigenesis. 5 ·
cance that embryological data began to provide for natural In the shift that I have described from eighteenth-century
history. A new view of the relationships among species, and of mechanistic preformation to nineteenth-century teleological
154 ~atter, life, and generation Epilogue 155
epigenesis, what role can be ascribed to Caspar Friedrich Wolff? Even though Wolff was not the sole, or even the primary,
Let me state at the outset that nowhere does Wolff express any cause of the shift toward epigenesis in German embryology, one
of the explicit teleological viewpoint found in Blumenbach and can identify elements of similarity in his and the later epigen-
the other Germans. We should recall that Wolff clearly distin- esists' theories. Although Wolff did not endorse a teleological
guished his vis essentialis from Blumenbach's Bildungstrieb, re- view of the organism, he was much less explicitly concerned
jecting the latter's "building force" on philosophical grounds. about the source-of-organization question than his preforma-
Furthermore, Wolff was none too happy with those who had tionist contemporaries had been. In his early works, he did not
mistaken his essential force for a "building force." Wolff's theory even deal with this pro~lem (much less solve it), leading to
of epigenesis was based neither on the assumed teleological
nature of the organism nor on the presupposition of its in-
herent organization. · ,7
Hailer's justified criticisms in this regard. In his unpublished
manuscripts, I have argued, one can find an account of how.
organization is passed on through generations, via qualified
i•
~
That Wolff's theory of epigenesis was different from those vegetable matter. Yet even here, Wolff did not offer this as a
proposed by his epigenetic successors was recognized by vori direct response to' the organization dilemma, but rather as an
Baer, who was familiar with almost all of Wolff's writings (see explanation for the fixity of species.
von Baer 1866:294-96). Against pre[ormation, von Baer wrote, Through his theory of qualified vegetable matter, Wolff.ex-
Wolff "fought a good deal and triumphantly ... and set up , plained both hereditary resemblance and variation. And even
against it the principle of epigenesis, of the actual new building though his theory represents neither a lapse into preformation
of all parts and of the whole embryo. In this he evidently went too · nor a teleological view of the organism, it does allow organiza-
far. It is true that neither the head, the feet, or any single part tion to be in some .sense passed on rather than created de novo
whatever was there earlier, rather all become; but they become with e.ach new instance of generation. Wolff's unpublished
not through actual new building, rather through the reorgan- work was, of course, completely unknown to his immediate
ization of what already exists" (1866:318-19; see also Oppen- successors, and it therefore had no influence on· the German
heimer 1967:295-307). Reproduction, von Baer explained, is a embryologists' subsequent endorsement of organization as an
continual process of organization being passed on from parent assumed property of living organisms. Yet it is significant to
to offspring. note that neither Wolff nor his epigenesist successors "solved"
Although Wolff's papers on the formation of the intestines the source-of-organization problem in the terms in which it was
were relatively unknown among German embryologists until]. enunciated by seventeenth- and eighteenth-century mechanists.
F. Meckel's translation of 1812, Wolff's support for epigenesis Wolff's rationalis.t program allowed him to redefine the question
in his earlier works was undoubtedly much more widely known. 6 in such a manner that his epigenetic explanation based on quali-
Certainly those who read Blumenbach's works were aware of fied vegetable matter accounted for the hereditary phenomena
Wolff, although they would not necessarily have been led there- of living organisms. Just so, the early nineteenth-century epi-
by to read Wolff in the original. Yet the later German embryol- 1 genesists viewed embryonic organization not as a problem to be
ogists·did not explicitly build their theories directly on Wolff's.
To be sure, Wolff was regularly cited, especially after 1812, as a
i explained, but as a starting point for embryological research.
The rise of teleological epigenesis was not an isolated event in
precursor to the epigenetic theory. Yet Wolff and his work were
out of the mainstream of German biology, because Wolff was at
St. Petersburg for almost all of his active career and trained no
il late-eighteenth-century Germany. Rather, it was part ofa much
broader shift toward developmental and historical thinking in
many areas of German intellectual life. Several scholars have
students there. Ironically, Blumenbach began his studies at f pointed to Naturphilosophie and Romanticism more generally as
1 providing a philosophical climate conducive to teleological
Hailer's former stronghold, the University of Gottingen, and !
he initially accepted Hailer's preexistence theory before con- thinking in biology, and to the reciprocal influence of biological
verting to epigenesis (see Lenoir 1980). Thus, although Wolff's theory on these philosophical movements themselves. Others
views were known among his German successors, his influence stress the contemporaneous rise of progressivist views of human
was less direct than one might initially suspect. history, and the social and political events experienced by
156 Matter, life, and generation
·1
I'i
German intellectuals in the 1780s and 1790s.~ All of these ·~ \-
1{f
analyses point to a fundamental change in philosophical view- APPENDIX A
point that encompassed far more than the biological sciences. fl
if,{
; i
Although it is beyond the· scope of the present study to explore ;: Chronology of the Haller-Wolff debate
these issues further, there is ample evidence to indicate that
German embryologists turned to epigenesis in the late eigh-
teenth and early nineteenth centuries because it filled the same
kind of philosophical need that preformation had during the
previous period. Epigenesis was as compatible with the new 1758" Haller Sur la formation du coeur clans le poulet
progressivist view of human history and natural phenomena as 1759 Wolff Theoria generationis
preformation had been with the religious and mechanistic be- 1759 Wolff Letter I-(23 December)
liefs of the seventeenth and eighteenth centuries. [1760]*. Haller Response to Wolff Letter I
AU of this points once again to my principal message. One 1760 Haller Review of Wolffs Theoriagenerationis
cannot treat the embryological views held during these earlier 176l Wolff Letter II (29 December)
[1762-64] Haller Response to Wolff Letter II
centuries in isolation from their wider philosophical context. Wolff Theorie von der Generation
1764
For Malebranche and Descartes in the seventeenth century, for 1764 Wolff Letter III (20 December)
Haller, Bonnet, Buffon, Wolff, and Blumenbach in the eigh- (1765] Haller Response to Wolff Letter III
teenth, as well as for Kielmeyer, von Baer, and the other German 1765 Wolff Letter IV (5 May)
epigenesists in the early nineteenth century, embryological 1765 Haller Review of Wolffs Theorie von der Generation
issues were not all that was at stake. The nature of matter and of [1765] Haller Response to Wolff Letter IV
forces, the applicability of mechanism to biology, the roles of 1765 Wolff Letter V (16 November)
1766 Haller Elementa physiologiae corporis humani, vol. 8.
the empirical· and the logical in scientific explanation, the re- Wolff Letter VI (6 October)
1766
ducibility of life to nonlife ""." these and similar questions formed [1766-67] Haller Response to WolffLetter VI
the background of embryological research and debate. As a 1767 Haller Opera minora, vol. 2
forum for the confrontation of these more fundamental issues, 1767 Wolff Letter VII ( 1 7 April)
the science of embryology thus provides a particularly revealing 1767 Wolff Moves to St. Petersburg
guide to the intellectual forces that shaped Enlightenment 1768-69 Wolff "De formatione intestinorum"
biology more gener~lly. 1770-71 Haller Reviews ofWolffs "De formatione
intestinorum"
1774 Wolff Theoria generationis (2nd ed.)
1776 Wolff Letter VIII (27 September/8 October)
[1777] Haller Response to Wolff Letter VIH
1777 Wolff Letter IX (7 May)
1777 Haller Death ( 12 December)
1778-83 Wolff Unpublished u::eatise on monsters
1789 Wolff Von der eigenthumlichen und wesentlichen Kraft
1794 Wolff Death (22 February)
*Because Hailer's letters to Wolff no longer exist, the dates given here for
them are estimated.
\
f
i
Wolffs letters to Haller 1 59
l
If, illustrious one, I could somehow please you with this work of
Wolff's letters to Halter mine, of whatever quality it may be, I would consider myself suffi..
ciently happy and would then also be confident of the approval of
i others. For your m!)st solid judgment has been for me up to now and
r
J
will always be for me equiv.aleni: to that of all men. If I have erred or
not satisfied your nature, do not deem me unworthy of your correc-
j tion, however it pleases you to provide it; rather, I would earnestly ask
Wolff sent a total of nine letters to Haller, seven during the years 1759 that you allow me to enjoy the same favor with which you are accus-
to 1767 and the final two in 1776 and 1777. The originals of the first tomed to regard all your disciples.
seven and the ninth letter are in the Burgerbibliothek in Bern ("Briefe I will be most devoted to you all my life.
an Albrecht v. Haller," MSS Hist. Helv. XVIII), while the eighth
letter is in the archives of the Germanisches Nationalmuseum in
Nuremberg. Haller published the first seven of Wolff's letters in LETTER II
volum(is four and five of his Epistolaro.m ab ero.ditis viris ( 1773-75).
These have also been published, with letter nine, in German transla- Berlin
tion by Schuster ( 194 1) and in Russian by Gaissinovitch ( 1961: 5 10- 29 December 1761
25). All nine letters are included below in full, letter eight being
published here for the first time. After I had received your letter, which was full o( the highest
humanity and benevolence toward me, and the works of the most
famous SproegeI; 1 I hesitated in doubt about whether I should re-
spond immediately in order to attest to my most grateful feelings or
LETTER I
rather whether I should not disturb you in matters of great importance
too often without need but should wait for, the publication of your
Berlin opinion and then fulfill all my duties at once. This latter argument
23 December 1759 finally convinced me. Now in the very celebrated Gottingen review,
I am bold enough to send you my inaugural dissertation [Theoria which. reached me very late through the fault of the Berlin distribu-
generationis, 1759], which sets forth my theory of generation, even tor, I have just now read, although they appeared last year, judg-
though I hold you to be the highest and _most acute judge of these _ments and opinions concerning my dissertation [ 1759] that I easily
matters; nevertheless your singular humanity, with which you sup- attribute to you yourself, illustrious one [see Haller 1760]. Where-
port the labors of those who at least make a sincere attempt to fore, I give you my greatest a11d most obliging thanks, both for the
accomplish something, bids me hope that you will regard my efforts benevolence with which you deigned to read my work and to pro-
favorably and fairly as well. nounce your most eagerly awaited judgment on it, as well as especially
Nor, illustrious one, does it frighten me that you have recently for the letter, which I think to have received from you will always be a
honored the opinion opposed to my theory with your illustrious name source of the highest glory to me. I am convinced that the praise that
and authority. For not only have I long been convinced of the sincere you acc~rd me in the very celebrated review, which I had not ex-
love with which you pursue the truth, which I believe I have touched pected, is due more to your benevolence and singular humanity
upon, and which the whole learned world admires in you, but those toward me than to my merits_. I will certainly not treat carelessly your
things that have been advanced in the most welcome experiments advice, e·specially that concerning the vessel walls; but rather I will
undertaken on incubated eggs are not of such a kind that I ought to now see to it with the greatest care that this matter, whatever it may
have abandoned all hope of the tr~th ofmy theory, which, at the time be, will at last be elucidated by new experiments. I admit that I was too
that I first read about the experiments, was almost finished. This is thrifty in reviewing experiments, since for the sake of reducing costs I
what your words surely say, illustrious one, in Part II, p. 172 [of Sur la chose only those experiments from my collection that I thought were
formation du coeur dans le poulet, 1758a]: "I propose the contrary exactly sufficient for demonstrating what I wanted. For the rest, I
opinion, which is beginning to appear to me to be the most probable. would hardly think that the opinions ofNet;dham could gain strength
The chicken has furnished me with reasons in favor of development from my work, illustrious one, since he dealt with another matter
160 Matter, life, and generation Wolff's letters to Haller 161
entirely different from mine. He strove with his infusoria not to or perhaps with other, higher investigations of nature. I have under-
ex'plain but merely to demonstrate the occurrence in nature not of the taken new experiments on incubated eggs so that I might reqder the
normal generation of perfect animals, which results from the union theory of generation more firm; and I have paid particular attention
of parents, but rather of the birth of microscopic animalcules from to what seemed most doubtful to you. The little book that I am
putrid matter without the union of similar animalcules. I, however, sending you [Theorie von der Generation, 1764] .will make clear what the
not• differing concerning birth up to. this point, attempted not to particular results of these labors are. Its first· part contains those
demonstrate that the usual generation of perfect animals and plants things that it seems to me can be elicited by stricter reasoning from
occurs but rather to explain it from its physical causes. But the observations already known before, and also from the nature of the
remaining obscure metaphysical speculations, which Needham adds thing itself. The second. part establishes a briefer theory, which is
without any experiments, merit hardly any attention,·in my opinion at corrected in places. Finally, I have advanced in an appendix those
least. And here too he has nothing in common with me, as, if it were things resulting from repeated experiments that seemed to me to
worth the effort, I could easily make amply clear. I explartied the fact accomplish the most toward my goal. I have added to these things a
that a heart does not arise in plants in § 216 indeed with too much drawing of these observations [see Figure 20], which, although not
conciseness on this basis: in general, the heart arises only if the vessels very ornate, is at least true, in order that you, illustrious one, can .
are so formed that smaller ones are always being gathered into larger judge them yourself. Figure 1 shows that very delicate cellular area
ones, and these finally into one common vessel, which, whatever its surrounding the spinal. column, in which a short time later those
form may be, is itself the true heart; but in plants, for this reason, protuberances that are the beginnings of the feet and wings arise out
collected or branched vessels cannot be formed, and consequently of the new liquid deposited within this area, which is itself the first
also for this reason a heart cannot result in them. Why branched beginning of the abdomen and thorax. Figure 2 shows the same area,
vessels cannot be formed in plants .was explained already in the but now it is much more complete, more determinate at its edges, less
preceding§ 2 15. I gladly admit that up to this point I 1'ave been totally diffuse, more firm and solid in its substance. From this you will easily
' ignorant of the source of irritability; therefore, I would not know how perceive in this matter, illustrious one, the mode of origin from the
to explain the origin of pulsation either, but I think that the remain- successive secretion of liquid, which formerly appeared lighter and
ing attributes of the arteries have been explained in§ 202, 205, etc. more diffuse, but which now is seen to be more solid and contracted. In
Now, illustrious one, may you live safe and sound for a very long truth, in this more solid area, those protuberances are now already
time, for the continuous growth of medicine and of all natural philos- present that reveal the first beginnings of the extremities. You will
ophy, both of which sciences have long owed their better part to your see at the same time, illustrious one, to what extent the heart is
labors. How indeed wholly new, more solid, and more pleasing the covered on both sides by the true beginning of the thorax. I ts anterior
face of the economy and nature of animals has appeared, on account and largest part projects freely, covered by the whole membrane of
o.f your method of investigation! Especially since you discovered the the amnion, which, bent back and continuous .with the skin of the
great principle of irritability, which seems to me certainly to be the embryo, covered the heart altogether. l have discussed all these
first foundation of all animality. How I now congratulate the learned matters in the appendix [of my book] on pp. 257-59. Figure 3 is that
world for so great a treasure as it has received in the Elementa physio- umbilical area [the area vasculosa], with the first beginnings of vessels,
logiae!2 To this the whole of medicine, with whatever form it may ever which l have described on pp. 26o-61. Certainly these places, which
be enabled or embellished, will always owe the firm, unchanging are hardly a little more obscure than the area itself is, and which are
foundation on which it rests. But I cease saying these things to you, full of pale dark fluid, are nothing else but that fluid which clings
for you are more aware of and see better than anyone else how much under the membrane of the yolk, which is prec:;ipitated from the white
natural science owes to you. Now, Iinstead commend myself to your matter that makes up the'area and liquefies in various places, and
further favor, illustrious one, for I will always be your servant. which stagnates; hence the area takes on a white color, opacity, and a
character that is quasi-dry. If the liquefying and dissolving of the area
continue, the fluid flows together from all directions, and the places
LETTER III where it has been stationary up to now become vessels, or rather
streams, as Figure 4 shows. In the interior parts of the area, fissures
Berlin and ruptures appear in various shapes, at the places·where the fluid
20 December 1 764 has not yet flowed together from all sides; in the exterior parts,
however, this fluid has already broken up the area to such an extent
Please do not construe it in bad manner, illustrious one, if I once that it is completely divided into larger and smaller islands, mostly
again dare to disturb you, even if you flre busy with weightier matters oval in shape. The integrity of the area in the places closest to the
"
162 Matter, life and generation Wolff's letters to Haller 163
r: •
ii./ ..· could happen that would please me more. I will be all the more
J t s ) ' ..
J" ,:, .. ~ persuaded of the truth of what you accept, for your most extensive
experience and most solid judgment are for me superior to the
r ;)
.B 7J weightiest arguments. Those things that seem doubtful to you will
certainly require further investigation, and will therefore provide
opportunity for new experiments and new speculations. Therefore,
your advice will always be the most useful of anyone's to me.
As far as the most celebrated Bonnet is concerned, I admit that I
perhaps wrote a little too harshly against him; but indeed, in thinking
the matter over seriously, I do not seem to myself to have sinned
greatly. Certainly it is not the contradictions that affect me. For if he
had contradicted me sincerely and openly, or had attacked me in any
?·~--
£,../,q,r,Ja:::.~~ u,n,.,,,,,, c.,,..£m/,r,p-. lo>,:, ,!Ju.L way, even vehemently or sharply, I would have accepted all this in a
a.tin:,. !a,&. nraa , u r , J , ~ eunr ""'2&Ulu
~ . i i J ~ - ~ · - ~., ~ ir,~-h,,=· rather agreeable spirit, well aware that all vehemence of expression
~
·4& "'~;t,,canJ,;, t_U.« pr,ln'l4 /la,.rurr,c "°/"""""' un'ut1IL -
. ~~~~ !:aawn/""2::- can hardly be avoided in disputes. But Bo_nnet set a trap, and exhibited
',..Jal,,,. ~
Ari!'~ rU"'/a.G;-,·tin4~~
E,.,/,,y~4/;Z;:'f.:tf::: }".~~.~•.--
. 'l" 1'.M... -
/ri·~ signs of an evil mind; I do not think that this matter requires forther
explanation. 3
.~=rr
~!:,.= "'" ,, ,, . -' • i Further, I pray, illustrious one, to the Divine Power that all the days
·~ ~7 < \ " of your life in the future may be happy, and not disturbed by any
,r ho,
.,
Ov,""'-7,.,,.,J.,Jw,,
~i.k.U, vif_,. · \ J
cares or by the envy of adversaries, so that you may be able happily to
finish for your own enjoyment your great work, which the learned
world _eagerly awaits. I will certainly always rejoice with a most grate-
... ful mind that I owe most of my accomplishments to this work and to
your other writings. Farewell .
~-~fi.--,,2,,,,,.,.
.a.. a a,.,.. ~ . 6 Ga,k, ,-,,-,
""""l'°'"'"
'!l"&,&,..,. E. ~ , , ; .
~ ans.z ~ · ~ i . d.
.
errv,yy.,,;:,...
r""¥'.:&w/.&,;.,,r,· ;,nr,,,,u;,,. ..
. e.~&nbry,,,.,,,..~·"l'"rv,u, LETTER IV
,,,.,,.. ~ l,,k.,. ~ , , . , . . , , . , ~ , , ,,,_"
- . . / a,~·.P· ~/."'o/=n!"~.
A. a : , ~ iupwre ,yl.iL.vn ,,.,.,,,.,, - Berlin
l,du~~eg~ r7't..'. I
5 May 1765
l,fH-f-r•.l!'.-4'C1'
Although I am always afraid, illustrious one, that by interrupting
you when you are occupied with very serious matters I may finally
Figure 20 • Drawings sent by Wolff to Haller along with Letter III, 20 December 1764. make myself troublesome and hateful, nevertheless the latest letter
(From Haller, Epistolarum ab truditis viris, 1773-75, vol. 6; courtesy of the Francis A. with which you have seen fit to honor me is of such a sort that, unless I
Countway Library of Medicine, Harvard Medical, School. See also Belloni 197 1.) respond to it, I may rightly be accused of impudence or at least of
gross negligence. Therefore, I think that I may rightly ask you,
illustrious one, to hear my excuses on this occasion at least.
As far as the first circumstance is concerned, about the thoughtless
quotation of the review [Haller 1760; reprinted in the Theorie von der
Generation], I admit that I committed an error that shames me more
amnion, which cannot be divided even by a needle without evident than I can say. Of all those things that you, illustrious one, have
violent laceration, prevents you, illustrious one, from being able to rightly pointed out to me, I was, through some unlucky star, com-
regard these fissures as perfected vessels that have only been dilated pletely unconscious. If any of these things had entered my awareness,
by the force of the heart a~d by ~lood propelled from the embryo. I would not have acted so inconsiderately. I certainly intend to be
If it should please you, illustrious one, to declare once more your careful in the future not to commit any similar error.
opinion concerning these matters and others that seem more note- As far as the second matter is concerned, if I had attacked the most
worthy to you that are contained in the little book, certainly nothing celebrated Bonnet when he was unaware of his fault and innocent, I
i64 Matter, life, and generation Wolff's letters to Hatler 165
would be very ashamed and would never forgive myself for such a seem to me to be nothing other than the lower layer of the area,
great transgression. But, illustrious one, how can it happen that distended by fluid in these places and raised higher, since the other
someone should openly seek to defend one thesis and to refute the places, where the white matter adheres between the two layers, pro-
contrary position, and that he should deliberate for a long time ducing the interstices between the roads, are more constricted and
concerning this matter (as the preface of the book of this celebrated therefore lower.
gentleman shows), but that, before he publishes his own book [1762], I, like all who study nature, await with great longing the expanded
he should neither read nor have read to him another book, in which Latin edition of your most excellent work on the formation of the
the contrary thesis is especially defended, which has only recently chicken and of bones [in Opera minora, vol. 2]. That day will be an
been published, in which new arguments are advanced, and which, especially festive one for me on which I begin to read and understand,
finally, should certainly have been known to him at least from its in the long-desired volume VIII of your Elementa physiologiae, thi::
review? Everyone reads on such occasions even the most worthless universal history of generation, which is a matter so close to my heart.
books that discuss the same arguments, so that he may have at least Farewell, illustrious one, and, if there is one thing I beg of you,
some familiarity with their contents. For these and other reasons I never deprive me of the favor of which you have considered rile
thought that I was obliged to reply to my refuter, who thought me worthy up to now.
worthy of refutation but not of citation. I should wish that whatever
happened in these matters in the meantime had not been done. It is
better to bear than to inflict injury, as I now see and have long known;
LETTER V
but it is difficult in disputes to avoid all harshness of expression. For
when I am eager to explain something, I often neglect the words, and Berlin
words that seem harmless enough to me as I write them, often seem 16 November 1765
harsh to a reader, and rightly so. This is the explanation that I am able
to give for what has been done up to now and for what cannot be On the 13th of November I received the letter that you wrote on the
undone. As far as what I should do in this matter in the future, that 27th of September, illustrious one, and I perceived in.it new indica-
depends upon your will, illustrious one, and upon that of the most tions of your customary humanity and benevolence toward me. If by
celebrated Bonnet. chance I have erred in those things that were concluded by reasoning,
I saw the beginnings of the vessels in the umbilical area [area I do not think that it is a dishonor to me, so long as my mind has always
vasculosa] shortly after the first day of incubation and for the whole been hostile to hypotheses and eager to pursue the truth itself. But I
second day, when the area, examined with the naked eye, appears seek the praise that I know how to venerate justly your most solid .
wholly intact and equal, and the vessels appear to have no color, not doctrine and you, its illustrious a_uthor, and how to form an accurate
even. a dark one. Then with the microscope I discovered spots that judgment of and to acknowledge gratefully all the things - how great
were somewhat darker than the rest of the area, and irregular in they are! - that I owe to you, illustrious one, in the physical sciences.
shape and size and not connected, which subsequent observations I understand that the formation of vessels in the egg, as well as the
revealed were the first delineations of the vessels. When in fact on the whole matter of generation, is a thing of great importance and cer-
third and following days red or dark points and lines begin to be tainly deserving of further investigation. Therefore I shall first await
apparent to the eyes, then the microscope easily reveals connected the new edition of your work [Opera minora] ~nd read of _the new
and branching roads that already have a greater similarity to vessels. experiments, and I shall then, following your footsteps, see what my
Spirit of wine also binds the two layers of the umbilical membrane shoulders can bear in further investigating nature itself.
together in such a way that the places that contain dissolved fluid and Farewell, illustrious one, and continue, as you are accustomed to
are the beginnings of the vessels project out on the lower surface; for do, to honor me with your favor.
this reason, the filled vessels swell and jut out beyond the parenchyma.
In this way a condition that is a little more p~rfect than it is in truth is
produced in the area. For after the first day, when the spots are still
not connected, wrinkles are for this reason formed on the lower
surface that are varied and mostly parallel, which prevJously did not
exist; but on the following days what had earlier been branching
roads appear, under the effect of the binding spirit [of wine), to be
swollen, branching vessels. But even at this time, these swollen vessels
166 Matter, life, and generation Wolffs letters to Haller, 167
LETTER VIII
St. Petersburg
169
I honor the illustrious Bonnet, as I always truly have, with the been closed, it is inserted in the right sinus alone. Indeed, closer to
highest respect. I rejoice to be about to have the occasion to attest this your observations, illustrious one, is the structure that I saw in a
publicly. 8 The very rich storehouse of monsters that has been col- human fetus of three months, where the inferior vena cava passes
lected and preserved over a long series of years in the Imperial directly into the left sinus by a large orifice (which is the same foramen
museum has now been handed over to me, so that I can compose a ovale), which is not opened by a valve but which does have a small
description of them and perform anatomies where I decide to. In this lateral orifice, wider and stopped by the Eustachian valve, by which it
therefore it will be necessary to deal once more both with the origin of opens into the right sinus, which at this stage is.already very large.
monsters as well as with generation in general. 'I know that these things cannot be sufficiently reconciled with the
I have not written to you for many years, illustrious and excellent things that anyone who practices customary dissection has observed
one. This has been because of the long journey that letters would have in nature, or thinks he has observed. But the customary dissection,
to make through many hands and especially because in all this time, which we practice most frequently, and which is most suitable and
since I have often been prevented by sickness, I have written nothing indeed necessary for uncovering, and exhibiting the foramen ovate,
that could have sei:ved as a sufficiently worthy approach to you. does not appear to be sufficient for recognizing its position more
Farewell, and, as you have done up to now, highest one, continue to deeply and for explaining the phenomena. In most illustrations con-
honor me with your favor. cerning the foramen ovale, as in Table 1, Figures 8 and 20 in Trew
[ 1736] and in that very beautiful illustration that is Figure I in the
Table concerning t~e foramen ovale in Fascicle 4 of your immortal
/cones anatomicae [ 1743-54], illustrious one, the right sinus is exposed
LETTER IX from the posterior, or right, side along with the inferior vena cava. In
these illustrations then appears immediately the orifice that is com-
St. Petersburg monly called the foramen ovate. All this region in which this orifice is
7 May 1777 found is believed to belong to the right sinus, and that part of it that is
I received your letter, which is dearer to me than anything else, on visible is believed to be that which produces the septum between the
the 10th of April, illustrious one. Together with the whole learned two sinuses. Therefore, the foramen ovate is thought to be in the
world and with all good men, I grieve, illustrious one, over your bad septum of the sinuses and tQ be observed and seen in the septum of
health! But I hope that without doubt God, who is so great and good, the sinuses.
\\'.ill soon restore your complete health. Now I ask, illustrious one, where in these illustrations or in nature,
On the 11th of April our Academy also received a response from which they represent, is the boundary between the right sinus and the
you. I give you indebted thanks, illustrious one, because you were inferior vena cava that allows us to determine that the foramen ovate
willing to add your most illustrious name to the splendor of our belongs to the sinus and not to the vein? And where in these illustra-
Academy, a signal honor in which I too participate. tions is the orifice by which the inferior vena cava opens into the right
My observations center chiefly around the mature or recently born sinus? by which alone this vein can be distinguished from the sinus,
fetus of humans and of calves; in these I have found that the fora men w~ich cannot appear in these illustrations, since the anatomist de-
ovale, which was thought to be in the septum of the sinuses, belongs to stroyed the orifice by cutting into the vein? So that it can be determined
the inferior vena cava, which ends on top with two orifices or short whether the foramen ovale is in the sinus or in the vein, the sinus must
branches, and that with the right one of these it enters the right sinus, be cut from the anterior side together with the anterior ventricle and
whereas with the left one, which is itself the foramen ovate, it enters the venous orifice of the ventricle (as in my Figure 1 [ see Figure 2 1]),
the left sinus. And yet, in my opinion at least, illustrious one, all this is leaving untouched the inferior vena cava and the orifice by which it opens
most consistent with your most beautiful observations on the incu- into the right sinus. Then in the open right sinus will first appear this
bated egg, which made it clear that only one auricle is present in orifice of the vena cava, by which it opens into the right sinus (my
earliest life together with the left ventricle. Without doubt, this is the Figure 1, numbers 10, 11, 12), above the isthmus of Vieussens (10)
left auricle, since it empties into the left ventricle. It is therefore and below the movable edge of the Eustachian valve (this valve is
necessary that the inferior vena cava should extend into this auricle in separated from the isthmus in the aforementioned illustrations and
this first stage, since the right auricle is not yet present. And it is bent back outside its position; in this way, the orifice of the vena cava
certainly this structure whose vestiges I recognized when I observed disappears). Within this orifice and in the very cavity.of the inferior
a fetus of nine manths. In it, the inferior vena cava still crosses into the vena cava, in its left wall, the foramen ovale, or rather only its larger,
left sinus through the left orifice with half of itself, and with half it higher part, appears (my Figure 1, nos. 13, 14, 15; and Figure 5, x, y,
crosses into the right sinus; whereas later, after the left orifice has z), since the lower part of the ring and foramen, descending deeper in
Wolffs letters to Haller 173
the wall of the vein, is covered by the Eustachian valve, and the whole
ring and foramen ovale only appear when the vena cava is cut, as in
/ your splendid Figure 1 and also in your incomparable eighth Figure
of the same Table, in which you exhibited for view the Eustachian
valve, illustrious one, loose and bent back, and the fossa ovalis which
was filled in in this way, as was proper. Therefore the foramen ovale
belongs to the inferior vena cava, in whose wall it is located, and not to
the septum of the sinuses, which is bounded by the highest part of the
ring, or isthmus (my Figure 1, no. 10). It is nothing else but the orifice
of the inferior vena cava, by which it opens into t_he left sinus.
The following, illustriQus one, are therefore the primary. argu-
ments of my dissertation, which I do not think were ever stated
before: 1) )'he inferior vena cava terminates on top in a fetus of nine
months in not one but two orifices or short branches. 2) By one of
these orifices it enters the right sinus, by the other, the left. 3) The
so-called foramen ovale is nothing other than this very left orifice of
the inferior vena cava, by which the way is opened into the left sinus.
4) The sinuses therefore do not communicate with each other, but
each of them communicates separately with the inferior vena cava. 5)
And the blood therefore does not pass from the right sinus into the
.p, . left, nor from the left into the right, but it proceeds directly from the
"j~-~~·. ····.~-.--~-=
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have abused your patience. .
If you wish to honor me with the second part of your Bibliotheca
anatomicae [ 1774-77], illustrious one, please send it to Basel to Emanuel
ll.....
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~ -· ' -C, .,.JC J .,,, U-+.-· ·····-~ Farewell, illustrio1:1s one! I pray to immortal God that you may be
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-JI strong! and that you may still live safe and sound for a long series of
years and may teach the learned world.
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Figure 2 1. One of Wolff's illustrations from his paper on the foramen ovale, showing Figure 21 (cont.).
the right auricle and ventricle of the heart, opened from the front right side. B, C, D, cava; 1, 2, J, 4, 5, brifice of the coronary vein; 5, 7, 8, 9, Eustachian valve; 8, 10, 11, 12,
right ventricle; E, F, G, H, K, right sinus (auricle);/, apex of the heart; p, superior vena right orifice of the inferior vena cava; 13, 14, 15, cavity of the inferior vena cava, in
which a part of the left orifice (or foramen ovale) can be seen. (From "D~ foramine
ovali," 1775)
Notes to pp. r-24 175
editions (1747, 1751c; trans. 1754) support epigenesis, while the third 17 Kuhlemann's dissertation ( 1753) supported a mild preformationist posi-
( 1 765b; trans. 1803, ·1966) and later editions support preformation. tion, based on a coagulation model; yet because it was not written until
7 See, for example, Jenny 1902; Cole 1930; Rostand 1930; Bilikiewicz after Haller left Gottingen, one cannot conclude from Kuhlemann's views
1932; Needham 1934;rev. ed. 1959; Meyer 1939; Schopfer 1945; Adel- what Haller's own thoughts on generation were at this time.
mann 1966, 3: 1388; and Hintzsche 1972. For a review of the literature on 18 Hailer's letters to Bonnet are preserved in the Bibliotheque Publique et
Hailer's conversion from epigenesis to preformation, see Mazzolini 1977. U niversitaire, Geneva, while Bonnet's letters to Haller are housed in the
8 This footnote does not appear in the first edition of the Scimmlung (1756). Burgerbibliothek, Bern. References to this correspondence will indicate
9 See also Gasking 1967: 107-16; Toellner 1971: 182-88; and Roe 1975. ':Haller MSS" or "Bonnet MSS," followed by the date of the letter. (See
1o Regarding Hailer's authorship of this review, see note 4. ~ all!O Bibliography, "Unpublished Materials," for the titles of these collec-
11 It is interesting to note that Maupertuis, whose theory of generation was tions.) For information on Bonnet, see Whitmann 1895a, 1895b; G.
very similar to Buffon's in this regard, recommended performing a series Bonnet 1929; Savioz 1948a, 1948b; Castellani 1972; Bowler 1973; and
of experiments where a specific part of an organism's body would be Anderson 1976.
destroyed generation after generation to see if deformed offspring would 19 I am indebted to Renato Mazzolini for correcting a translation I pre-
result. There is no evidence that these were ever carried out (see viously suggested for this passage. Mazzolini ( 1977:226), however, argues
Maupertuis 1745, 1966 trans.:78-79). In the late nineteenth century, on the basis of this statement that, at the commencement of his observa-
August Weismann tried a similar experiment to test the inheritance of tions on incubated chicken eggs, Haller had placed himself in a position of
acquired characteristics. He cut off the tails of mice for several genera- "voluntary neutrality" with regard to preformation versus epigenesis.
tions, but without any visible hereditary effects (see Weismann 1891-92; Yet, although Haller clearly recognized that his objections to Buffon's
Churchill 1976). theory challenged his own epigenetic account of development, he still
12 The "tree of Diane" was frequently cited in the eighteenth century as an believed in 1754, as this passage demonstrates, in an epigenetic coagula-
example of how natural laws could produce complex, organized forma- tion model for embryological development. (For a similar interpretation
tions. A treelike figure forming on the surface of water through the of this passage see Bonnet 1762, 1: 139; Schopfer 1945:83-84.) This
chemical interaction of silver, nitric acid, and metcury, the arbre de Diane model was transformed into a 'preformationist one only subsequently,
was used by Maupertuis (see Chapter 1) and others as a model for epi-' culminating in Hailer's conversion to preformationism in 1757.
genetic development. · 20 It is not entirely dear what opinion of Bonnet's Haller is referring to here..
13 This passage is followed by a statement that has led Roger ( 1963:707) and In the correspondence now extant in Geneva there is no letter· from
Toellner ( 1971: 184-88) to erroneously date Hailer's conversion to pre- Bonnet to Haller between Hailer's letters of 26 November 1754, and 4
formation as having taken place by 1751. Here Haller states, "This con- January 1755. Bonnet, who reprinted this quotation in.his Considerations
stancy {of same type of organism resulting from reproduction] has con- sur Les corps organises ( 1762), preceded it with the following: "I had always
vinced me against all the experiments of M. Needham [on the formation thought that a glue that appeared to organize itself was already organized.
of animalcules in infusoria]; there must accordingly be something pre- I had never been able to accept in my mind [the idea] that the parts of a
pared and built in the fertile liquid of man and animals, although it is not a plant or an animal should have formed themselves successively. The more
miniature of a whole body nor a caterpillar of an imagined butterfly" I reflected on such a formation, the more I felt the insufficiency of the
( 1752a:xvi). The "imagined butterfly" is undoubtedly a reference to mechanical means celebrated with such complacency by diverse authors. I
Swammerdam's demonstration in 1669 that a butterfly could be found insisted on the above to M. de Haller, when I received this response"
folded up in both the caterpillar and the chrysalis, which was taken as a (I: 139-40).
proof of pre formation (see Chapter 1). Read in context, this passage from 2 1 For additional brief discussions of Haller's observations on incubated
Hailer's preface indicates only that Haller believed that the material from chicken eggs in his letters to Somis, see Hintzsche 1965: 17, 27, 29. Refer-
which the embryo develops, which is derived from the parent, influences ences to these observations can also be found in Hailer's letters to Gessner
the type of organism that results and ensures constancy of species. That (Sigerist 1923:239, 250, 256, 257, 258-59, 265, 267, 269), Morgagni
Haller had not yet converted to preformation is further borne out by the (Hintzsche 1964:65, 72, 76, 81, 84-85), Caldani (Hintzsche 19-66: 18-19,
fact that Haller reiterated his support for epigenesis in a letter he wrote to 21-22, 27, 28, 29), and Tissot (Hintzsche 1977:42, 43, 59).
Bonnet in 1754 (discuss~d later in this chapter). 22 Haller believed that conception occurs when an '.'odorous"portion of the
14 See also Hailer's discussion of Spallanzani's work in the Addenda to seminal liquor stimulates the preformed heart to begin beating. Although .
volume 8 ( 1766) of his Elementa physiologiae (pt. 2, pp. 216-17). For two he was well aware that Harvey's experiments on deer had shown that no
letters from Needham to Haller, in which Needham defends his epi- . male semen could be found in the uterus after copulation, a conclusion
genetic views against the charge of materialism, see Mazzolini 1976. confirmed at least in part by Haller's own observations on sheep with
Halle r's objections to Needham's theory are also discussed by Duchesneau Kuhlemann, Haller was not convinced that the semen never reaches the
(1979:80-85). uterus. Yet he thought that only a vaporous portion of the seminal liquor
15 See Haller 1765b, §§ 400,402,404,407. In the Elementa physiologiae actually acts to stimulate the heart's irritability (see Haller 1757-66,
( 1757-66, 4:446), Haller referred to irritability as the vis contractilis 8[ 1766J: 18-2 3, 154-55). As Mazzolini ( 1977: 203, n. 54) has noted, Haller's ·
musculis insita ("contractile force innate to muscles"). theory of conception cannot properly be described as one of "fertiliza-
16 See Rudolp~ 1964; Toellner 1967; and Roe 1981. tion" but is rathu one of "stimulation" or "irritation." Although I have
/
'
178 Notes to pp. 4 1-48 Notes to pp. 48-85 1 79
continued to use the terms "fertilized" and "unfertilized" when discussing 3 Wolff's vesicle theory cannot be considered a precursor of the modern cell
Hailer's theory, they should be understood in tbe broader sense of theory. Rather, Wolff's belief that all organic parts acquire vesicles and
"fecundated" and "unfecundated," the terms actually in use in the vessels during development is entirely consistent with the widespread
eighteenth century. belief in the eighteenth century that "cellular tissue" (i.e., connective
23 Both Gasking (1967:112-13) and Duchesneau (1979) point to Hailer's tissue) is the major structural component of organisms (see Wilson 1944).
views on the interdependence of organ systems and on the relationship Wolff stressed that the same process of vesicle and vessel formation occurs
between organic function and structure as having played influential roles in plants and in animals, and that one can therefore consider as analogous
in his adoption and elaboration of a preformationist theory. See especially plant "cells" (observed by Malpighi and Grew) and animal "cellular tissue."
Duchesneau (pp. 90-100) for an analysis of Hailer's explanation of the 4 With his theory of plant development from a "vegetation point," Wolff
developmental process in terms of the structure-function relationship. anticipated Goethe's concept of metamorphosis. Goethe was apparently
24 Gasking (1967:110-11) and Mazzolini (1977:199) assert that'Haller's unaware of Wolff's ideas until after he had formulated his own theory (see
membrane-continuity argument was not a proof of preformation per se Goethe 1817-24, 1:8o-89; see also Kirchoff 1867; Kohlbrugge 1913; and
but only of ovist preformation (as opposed to animalculist preformation). Arber 1946).
Consequently, they argue, Haller's discovery,ofthe membrane-continuity 5 See Adelmann 1966, 3: 1106-27, for these early observers' comments on
proof was not, as Cole ( 1930) and others have argued, the cause of his the area vasculosa. ·
conversion to preformation. Although I agree that Hailer's formulation 6 These islands must not be confused with the blood islands (first correctly
,of this_ proof was only one of many factors contributing to his conversion identified in the nineteenth century) that form in early stages of the area
(as I have shown in this chapter), nevertheless Hailer's proof, by demon- vasculosa. Wolff's islands represent the material broken up by the forma-
strating that the embryo exists before conception in one of the parents tion of blood vessel channels, these channels forming between the islands.
independently of the other, was for that reason an argument for pre- 7 Haller probably received Wolff's dissertation in February, for in March
existence. Its discovery was only one of the observational factors that he remarked in a l~tter to Bonnet, "A young physician of Berlin has sent
played a role in Halter's new explanation of development, yet its im- me a thick thesis in which he ·defends epigenesis.... his name is Wolff"
portance should not be underestimated. For further analysis of Hailer's (Bonnet MSS, 4 March 176o).
membrane-continuity proof, see Chapter 3. 8 Regarding Haller's authorship of this review, see Chapter 2, note 4.
25 Mazzolini ( 1977: 2 28-29) suggests that Haller's reading of Diderot's Pen.sees 9 Haller had continued his observations on.incubated chicken eggs and on
sur l'interpretation de la nature ( 1754) was influential in his conversion to quadrupeds in the summer months of 1763 and 1764, intending to include
preformation. Mazzolini reports that Haller read this work in 1757 and these in a revised Latin version of his Sur la formation du coeur dans le poulet,
that he remarked about it in a letter to Bonnet, "they want to tear from us scheduled to appear in volume 2 of his Opera minora ( 1762-68). The series
the strongest and most popular demonstration that we have of the ex- of observations undertaken in 1765 was designed to refute Wolff's claims
istence of God" (Haller MSS, 6 July 1757). In this work, Diderot had as presented in the Theorie von der Generation. See Hailer's unpublished
briefly presented Maupertuis's theory of generation (as explained in the journal ofexperiments, "Observationes anatomicae Bernenses," vol. 3.
- Systeme de la nature [ 1751 ]), had criticized its giaterialistic implications, and 1o Adelmann ( 1966, 3: 1028-29) identifies Maitre-Jan's "little white body" as
had hinted at his own theory of sensibility, which would be expanded in Le the nucleus of Pander, a concentrated region of yolk that, when_ viewed
Rive de d'Alembert ( 1769) into Diderot's materialist view of vital function- from above, both in unincubated eggs and i!) early stages of development,
ing and of generation. There is no doubt that Haller was disturbed by looks like a small white disc. Numerous observers, Malpighi and Harvey
these views and that, in 1757, they reaffirmed the dissatisfaction wjth among them, frequently mistook this "white point" for either the embryo
epigenetic theories that had stemmed from his critique of Buffon's and or the developing heart.
Needham's views in 1751. Whether Haller read Diderot's book before or 11 For a more detailed explanation of Wolff's theory, see Adelmann 1966,
after he had actually formulated his preformation theory is at this point 4: 1653-57 ..
not possible to conclude for certain. 12 Regarding Halle r's authorship of these reviews, see Chapter 2, note 4.
13 Wolff's work on the intestines was cited by von Tredern, Tiedemann,
Burdach, Oken, Dollinger, Pander, and von Baer, among others. See
Chapter 3. The embryological debate Adelmann 1966, 4: 1679-1739, for the impact of Wolff's work on German
embryology. On the history of the germ layer concept, see Oppenheimer
1 Wolff's letters to Haller (nine in all) are reprinted in English translation in 1967:256-g4.
Appendix B of this book. All references in the text will give page citations 14 See Needham 1934, 1959ed.: 158,185,233; Adelmann 1966, 5:2236-42.
to the original Latin of these letters as published by Haller ( 1773-75). On the microscopes and techniques that Haller used, see Mazzolini 1977:
Unfortunately, Hailer's letters to Wolff are no longer in existence. 220-24.
2 For biographical information on Wolff, see Kirchoff 1868; Wheeler 15 Wolff's "globules" were most likely optical artifacts. As Hall (196g, 2: 186)
1899; Schuster 1936; Schutz 1947; Uschmann 1955; Herrlinger 1959, has remarked, the history of the globular theory is largely the history of an
1966; Gaissinovitch r961, 1978; Rajkov 1964; and Lukina 1975. For a illusion, one due to diffraction haloes caused by improper lighting tech-
contemporary view of Wolff, written by one of his students, see Mursinna's niques and to compound microscopes uncorrected ,for spherical and
letter in Goethe 1817-24, 1:252-56. chromatic aberration.
-~
180 Notes to pp. 89-109 "i
Notes to pp. ro9-4 7 181
(see, for example, the German translation, Wolff 1896, no. 85:81; and the
Chapter 4. The philosophical debate: Newtonianism versus rationalism Russian translation, Wolff 1950:203). In my opinion, the."suffer from"
1 See d'Irsay 1930; Guthke 1962, 1970, 1973, 1975; Sonntag 1971, 1974a, translation is the correct one for two reasons. First, it is consistent with the
1974b, 1975, 1977; and Toellner 1971, 197~. other discussions of the soul in Wolff's dissertation (dted above). Second,
2 See Lundsgaard-Hansen-von Fischer's (1959) catalog of Hailer's pub- in a later work of Wolff's titled Von dn- eigenthumlichen und wesentlichen
lished works, which contains over 300 main entries, a majority of which Kraft (1789), Wolff, in complaining of Hailer's earlier critique of his
are books, and an additional 400 editions and translations. This list does dissertation, remarked that "Haller has thus not been entirely correct
not include Haller's ten- to twelve-thousand book reviews (see Chapter 2, ... when in the judgment of my work he put forward the essential force as
note 4). · the main point, remembering all that properly belongs to this theory
3 See Hochdoerfer 1932; Guthke 1962: 17-19, 1975: 174-92; and Toellner almost with no words and then deliberately mentioning that this thing is
197 1: 1-2 7, who provides a review of the literature on this s4bject. called by me the essential force, but which I completely separatedfrom the soul
4 Original text: "Genug, es ist ein Gott; es ruft es die Natur,/Der ganze Bau of the Stahlians; about which indeed exactly so much as nothing was said"
der Welt zeigt seiner Hande Spur." (p. 50 n.; italics added). Wolff's perception thirty years later was that he
5 Original text: "Der Mensch, vor dessen Wort sich soll die Erde biicken,/lst had dearly distinguished his essential force from the soul and from Stahl's
ein Zusammenhang von eitel Meister-Stiicken;/ln ihm vereinigt sich der ideas. That he _was referring to the passage I am now discussing is borne
Korper Kunst und Pracht,/Kein Glied ist, das ihn nicht zum Herrn der out by the fact that nowhere else in the dissertation are Stahl or the
Schopfung macht." "Stahlians" mentioned. (On the translation of this passage see also Mario
6 On Hailer's debate with Whytt, see Haller 1756-6o; Whytt 1751, 1755; Stenta's comments in Driesch 1911:37g-83, n. 78; and Gaissinovitch
and French 1969. For Haller and La Mettrie, see Haller 1751d; Bergmann 196 1:253-:-55.)
1913; Vartanian 1949; Saussure 1949; Guthke 1962a; and Hintzsche 19 On Haller's critique of Needham's views see Duchesneau 1979:8o-85.
1968. See also Bastholm 1950.
7 See Richter 1972:6<>--08; Roe 1981. In his pu&lications and diaries, Haller Chapter 5. Wolff's later work on variation and heredity
refers to Newton's Optic/rs, the Principia, the Leibniz-Clarke correspon-
dence (Clarke 17 15), 'sGravesande's Physices elementa mathematica ( 1720- 1 For a list of these materials, see Rajkov 1964:625-26 and Wolff 1973:304-
21 ), and Henry Pemberton's View of Sir Isaac Newton's Philosophy (1728). 5. See also von Baer· 1847, for the earliest report of Wolff's manuscripts.
See Haller 1746c and 1774-77, 1:621-22; Sigerist 1923:22-23; Guthke 2 All citations will be to the Latin text in Wolff 1973.
1967: 150; and Hintzsche and Balmer 1971:80, 91, 93. 3 In all, Wolff published thirty-six papers in the Academy of Sciences'
8 See Jones 1925; Price 1926; Teeter 1928; and Richter 1972:57-111. journal during his years in St. Petersburg. Aside from the papers on
9 Original text: "Ein Newton iibersteigt das Ziel erschaffner Geister,/Findt monsters and on the formation of the intestines, these are mostly anatom-
die Natur im Werk und scheint des Weltbaus Meister;/Er wiegt die innre ical investig~tions, including several studies of the muscular fibers of the
Kraft, die sich im Korper regt,/Den einen sinken macht und den im Kreis heart. For a complete list of Wolff's papers, see Rajkov 1964:623-25 and
bewegt,/U nd schlagt die Tafeln auf der ewigen Gesetze,/Die Gott einmal Wolff 1973:302-4.
gemacht, dass er sie nie verletze." . 4 See Gaissinovitch 1961:444-50; Rajkov 1g64:6o5-18; and Lukina
1o Original text:" ... Er fiillt die Welt mit Klarheit,/Er ist ein stater Quell von 1975:420-21. .
unerkannter Wahrheit." 5 Wolff refers to Ethiopians several times in his treatise in discussing varia-
11 See, for example, Haller 175oa:xiii-xiv, xvii, xviii-xix; 1757-66, 4:533, tions in human traits, and at one point he cites a tract called "Ethiopian
557; and 1774-77,'1:621-22. Anatomy," apparently written by himself (1973:221, § 75). Lukina, who
12 I have altered this translation from "frame" to "feign" following Cohen has not been able to locate this paper among Wolff's manuscripts, sug-
1971:241, n. 9. gests that it was written after 1772, when Wolff was sent the body of an
13 I am reading "partibus" for "particulis" in the phrase "Theoriam autem, African man for dissection (see Wolff _1973: 289, n. 21).
cur utravis proprietas aut in his partibus m.illa sit, aut in aliis corporis 6 See Stearn 195 7: 156-61; Glass 195gc: and Larson 1971:94-121.
humani particulis [sic] aliqua .... " 7 See Roger 1963:567-82; Farber 1972; and Bowler 1973. ,
14 See also Hailer's adoption of Newton's ideas on motion in his Elementa 8 See Gaissinovitch 1g61 :444-50 and Rajkov 1964:6oo--005. For a Russian
physiologiae, 1757-66, 4:557-58. translation of the "Distributio operis," see Gaissinovitch 1961 :526-33. All
15 SeealsoOakley 1961;Kubrin 1g67;McGuire 1g68;Cohen 196g;Heimann references will be to Gaissinovitch's translation.
. and McGuire 1971; and Tamney 1979. 9 See Gaissinovitch 1961:426, n. 682; 1978:525; Rajkov 1964:619-20; and
16 On Christian Wolff's philosophy, see Gurr 1959; Blackwell 1961; Burns Lukina 1975:417-18. The Latin text of this section is included in Wolff
1966; Beck 1969; Corr 1972; and Frangsmyr 1975. 1973:288-89.
17 On Hoffmann and Stahl, see Lemoine 1864; Rather I g61; King I g64, 1o I am not arguing that a beliefin rationalism necessarily entails support for
1969; and Duchesneau 1976. epigenesis. Both Leibniz and Christian Wolff, for example, supported
18 Wolff's use of the verb paterer (imperfect subjunctive of patior) has en- theories of spermaticist preformation (see Christian Wolff 1723, § 444;
gendered <,:tmsiderable controversy among Wolff scholars, who have fre- Leibniz 1951:109, 195-97, 276, 526-27, 548-49, 550; and Roger 1968).
quently translated paterer in the sense of"allow" rather than "suffer from" What I do wish to claim is that, in Caspar Friedrich Wolff's case, his belief
182 Notes to pp. 147-63 Notes to pp. 164~73 183
in rationalism, together with his phil9sophy of biology, provided the basis responded by attacking and ridiculing Bonnet's views in his Theorie von der
on which an epigenetic theory of development could be built. Generation ( 1764). Although Haller apparently maintained, in his reply to
this letter, that Bonnet had indeed not read Wolff's dissertation and had
Chapter 6. Epilogue: the old and the new only heard of it through Hailer's letters, Wolff was not satisfied, as can be
seen in his subsequent letter (Letter IV).
1 On the H~rvey-pescartes controversy, see Passmore 1958; Mendelsohn 4 See Wolff's paper, "De foramine ovali, ejusque usu, in dirigendo motu
1964; and Toellner 1972. Regarding spontaneous generation, see Farley sanguinis. Observationes novae," Novi Commentarii Academiae Scientiarum
1977 and, on the Pasteur-Pouchet debate, Farley and Geisorn974. Con- /mperialis Petropolitanae, 20(1775):357-430. After this volume, the Acad-
cerning Stahl and Hoffmann, see Chapter 4, note 17. For the Lawrence- emy's journal was titled Ai:ta Academiae Scientiarum lmperialis Petropolitanae.
Abernethy controversy, see Temkin 1963 and Goodfield-Toulmin 1969. 5 The fora.men ovate is an opening in the septum between the two auricles
On Roux and Driesch, see Churchill 1969, 1973; and Allen 1975. of the mammalian fetal heart. Before birth, the fetal lungs do not operate,
2 On incommensurability and related philosophical and social aspeqs of and the fetus receives oxygenated blood through the umbilical vein. This
scientific controversy, see, for example, Hanson 1965; Scheffler 1967; blood enters the right auricle of the heart through the inferior vena cava.
Kuhn 1970; Lakatos and Musgrave 1970; Suppe 1974; Barnes 1974; and The foramen ovale allows the blood to bypass the fetal lungs by sending it
Mulkay 1979. directly from the right auricle into the left auricle, rather than i_nto the
3 See Temkin 1950; Larson 1979; ano Lenoir 1980. right ventricle and thence to the pulmonary circuit. The foramen ovate
4 Not all of the German embryologists I am referring to accepted the closes shortly after birth when the lungs begin to function and normal
recapitulation theory. Von Baer, for one, was critical of the views of circulation is established. Analogous openings between the two auricles
Meckel and other proponents of recapitulation, substituting his own exist in the fetal chick and in the embryos of other animals with four-
theory of divergent development .... that among the four basic types of chambered hearts. '
organisms embryonic development proceeds from the general to the 6 Wolff uses the word· "sinus" to refer to the entire region of the right or left
specific, that is, for example, from a generalized vertebrate form to a auricle. He reserves the term "auricle" for an ear-shaped portion of the
special kind of vertebrate (see Oppenheimer 1967:221-55; Ospovat auricle that projects out over the surface of the heart. Modern anatomists
1976; and Gould 1977). Meckel and the other recapitulationists believed often distinguish between "atrium" and "auricle" in the same 111anner.
that the embryo proceeds through the adult forms of lower organisms (in 7 See Winslow 1717, 1725a, 1725b; and Rouhault 1728. An earlier con-
an ideal sense) during its developmental sequence. It should also be noted troversy over the function of the fora men ovate also took place at the turn
that these early proponents of the biogenetic law believed that ontogeny of the century between Jean Mery and Joseph-Guichard Duverney. See
recapitulates phylogeny in an ideal, or transcendental, sense rather than tht; volumes of the Histoire de l'Academie Royal des Sciences for the years
in an evolutionary one. The biogenetic law received its first evolutionary 1699, 1701,and 1703.
formulation later in the nineteenth century in the work of Fritz Muller 8 Both Bonnet and Haller, along with several other notable hgures, were
and Ernst Haeckel (see Gould 1977). made ~oreign members of the St. Petersburg Academy of Sciences in
5 See, for example, Oscar Hertwig's tract, The Biological Problem of To-day: honor of its fiftieth anniversary. See Proces-verbaux des seances de l'acadimie
Preformation or Epigenesis1 ( 1896), which opposed· August Weismann's imperiale des sciences ( 1897-1911), 3:272-75, 298.
germ plasm theory of inheritance. Preformation-epigenesis controversies 9 Wolff's account of fetal circulation and of the role played by the foramen
also arose in the early twentieth century in association with debates over ovate was erroneous in several respects·. Although his illustrations of the
Mendelian genetics and over the chromosome theory. See Churchill embryonic heart are quite accurate (see Figure 2 1), Wolff was incorrect in
1968, 1970b; Allen 1975; and Baxter 1976. claiming that the inferior vena cava sends blood separately to the right
6 See Kohlbrugge 1913:218, n: 14; Temkin 195o;and Herrlinger 1966:22- and left auricles. The blood that enters the right auricle from the inferior
28. Herrlinger notes that Wolff's writings were eclipsed by the popularity vena cava passes (for the most part) directly through the foramen ovate
of Blumenbach's Ober den Bildungstrieb und das Zeugungsgeschiifte ( 1781 into the left auricle (and then to the left ventricle). Although some blood
and later editions). does indeed pass from the right auricle to the right ventricle, this is blood
7 See Temkin 1950; Oppenheimer 1967:136-47; Gasking 1967; Gould that has entered the right auricle from the superior vena cava. The two
1977; and Lenoir 1978. streams of blood (from the inferior vena cava and the superior vena cava)
remain quite separate in the right auricle (see Patten 1968). What may
Appendix B. Wolff's letters to Haller have confused Wolff is the fact that, in the fetal heart, the valve of the
1 Wolff had probably received from Haller some of the publications of inferior vena cava (the Eustachian valve) is continuous with the border of
Johann Adrian Theodor Sproegel, one of Hailer's former students, who the foramen ovale. Thus, it would be possible to view the inferior vena
completed his medical degree at Gottingen in 1753. cava as continuing directly into the left auricle.
2 By the end of 1761, three volumes of Hailer's Elementa physiologiae had Haller was apparently unimpressed with Wolff's anatomical investiga~
been published. tion on the fora men ovate. As he wrote Johannes Gessner after receiving
3 Bonnet had defended preformation and attacked epigenesis in his Con- Wolff's paper~ "He says nothing in all.these pages except that the foram~n
siderations SUT Les corps organises ( 1762) without mentioning Wolff or his ovale is a canal that has two openings, but this had been kno.wn already for
theo,ry of epigenesis. Wolff interpreted this as an intentional snub and a long time" (Sigerist 1923:535; letter of 19 April 1777).
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204 Bibliography
Bonnet, Charles (cont.) "De partibus corporis humani sensilibus embryonic organization, problem of Eustachianvalve, 171,173, 183n.9
discovered by, 23; preformation and, et irritabilibus," see Haller, works source of, 8; 151,155; Descartesand,4; evolution (developmental), see
12, 19,20,40,45,57,60,73-74, 150, Descartes, Rene, 156; De la formation de Haller and, 25, 35, 91-92, 112-13; preformation, evolution and
151, 182 n.3; on regeneration, 12, 22- l'animal (1664), 4; epigenesis and, 3-4, Maupertuis and, 15; teleological evolution (of species), see transformism
23, 42-43; St. Petersburg Academy 14, 19, 151; Garden on, 8; Harvey and, epigenesis and, 151-52; von Baer and, explanation, scientific, 103-7, 110, 112,
membership, 183 n.8; spontaneous 148, 182 n.1; Malebranche on, 5; as a 154; Wolff and, 112-13, 122-23, 145- 119, 147, 148
generation opposed by, 19; Traiti mechanist, 3-4; Traiti de l'homme 46, 155
d'insectolot;ie ( 1745), 23; see also Haller, (1664), 3-4; Treatise of Man (1972), 174 embryonic structur,es, observability of, see Fabricius ab Aquapendente,
works reviewed by n.4; see also Haller, Descartes and; observability of emb~yonic structures Hieronymus, 3, 6, 51, 167
Born, Carl Friedrich, 114, 115 Wolff, Descartes and empiricism, go, 95, 96, 110, 121, 122 Fontenelle, Bernard Le Bovier de, 1
Bowler,Peterj.,7, 174n.1, 175n.5, 175 development, divergent, 182 n.4 encasement, see emboitement foramen ovale, structure of: Haller on,
n.7, 175 n.9, 177 n.18, 181 n.7 developmentalism in Germany, 152-53, epigenesis (see also Haller, conversion 171, 183 n.9; Paris Academy of
Buffon, George Louis Leclerc, Comte de 155-56 to epigenesis; Haller, epigenesis, Sciences debates on, 169, 183 n.7;
(su also Haller, on Buffon's generation Diderot, Denis, 12, 178 n.25 theory of; preformation, epi'genesis Wolffon, 169, 170-73, 183n.5, 183n.9
theory; Wolff, Buffon and), 156; on Driesch, Hans, 149,153,181 n.18 vs.): and area vasculosa, formation of forces (see also Bildungstrieb; forces,
animalcules, 16; epigenesis and, 15-18, Duchesneau, Fran~ois, 176 n.14, 178 blood vessds in, 51; 53, 54; Aristotle attractiv~; gravity; irritability, as a
151, 175n.9;onfemalesemen, 16, 17; n.23, 18on.17, 181 n.19 and, 3, 174 n.2; atheism and, 8; force; vis essentialis; vis insita):
Histoirenaturelle(174g-89), 15-18, 26, Duverney,Joseph-Guichard, 125, 183 attractiveforcesand, 13, 14-15, 16, 17- embryological development and, .
28,153; Maupertuisand, 18, 19,176 n.7 18, 18-19,24-25,31,35;Blumenbach Haller on, 28-32, 35-36, 43, 113; God
n.11; moule intirieur (internal mold) 16- and, 118, 150, 151, 152, 153, 154; as source of, 30, 35, 43, 97-98, 101,
17, 18, 28, 153; Needham, collabora- Elementa physiologiae corporis humani, see Bonnet's opposition to, 6o, 182-83 n.3; 146, 151; Haller-Wolff debate and,
tion with, 16, 18, 19; as a Newtonian, Haller, works Buffonand, 15-18, 151,175 n.9; 101, 121, 148; irritability and, 33, 35,
18; on nutrition, 16; penetrating force emboitement (encasement; see also definition of, 1; Descartes and, 3-4, 14, 43, 98, 100; materialism and, 98, 112;
of, 16, 17-18, 29; preformation preformation), I, 3, 8, 9, 19, 73, 147, 19, 151; Diderot and, 12,178 n.25; matter and, 2g-30, 35, 97-98, 100-
opposed by, 15-16, 19; ,m species, 138- 152, 174 n.1; animalculism and, g; Dollinger and, 150; embryo in fluid 101, 102, 120-u, 151; penetrating,
39; see also Haller, works reviewed by Hailer's belief in, 36; Malebranche's state and, 23, 25, 31, 38; fermentation Buffon's theory of, 16, 17-18, 29; in
Burgerbibliothek (Bern), 36, 158, 177 theory of, 3, 5, 84; regeneration in and,4, 13, 14, 19,24, 106;forcesand, physiology, Haller on, 35, 36, 96-g7;.
n.18 polyp and, 12; Swammerdam and, 3, 6 Hailer's critique of, 28-32; Galen and, vegetative, Needham's theory of, 18-
embryological development (see also area 3; 174 n.2; as generic preformation, 19, 31-32, 113, 120-21; see also Haller,
cell theory, Wolff and, 85, 179 n.3 vasculosa, formation of blood vessels 152; and guidance, problem of, 4, 8, on forces
cellulartissue, 179 n.3 in; emboitement; epigenesis; heart, 14-15, 18,25,4~ 112-13, 122,145; forces, attractive: Buffon on, 16, 17-18;
chicken eggs, observations on (see also formation of; heartbeat; intestines, Haller and Kuhlemann's observations in embryology, 13, 18, 19; Hailer's
Haller, observations on chicken eggs; formation of; observability of on sheep and, 36; Hailer's rejection of, theory of epigenesis and, 24-2 5, 31,
Wolff, observations on chicken eggs): embryonic structures; preformation; 44, 98, 122; Hailer's religious beliefs 35; Maupertuis on, 13, 15; Needham
by Harvey, 3, 167, 179 n. 10; by transparency, development of embryo and,91, 110-11; Harvey and, 3, 14, 51, - on, 18-19; Reaumur's critique of, 14-
Malpighi, 5, 6, 174 n.5, 179 n.10 and): chance and, g, 98, 101, 125; and 65, 174 n.3; and heart, formation of, 15
chorioallantois, 50, 75, 76, 77, 78 guidance, problem of, 14-15, 18, 25, 23, 24, 25, 39, 42; and intestines, frogs' eggs,.Swammerdam's observations
chorion of Mal pig hi, 76, 167; see also 31,35,36,43,44,91-g2, 112-13, 122, formation of, 46, 8o-82; Kant and, on,6
serosa 145 (see also embryonic organization, 151,152; Kielmeyerand, 150,156; La
Churchill, Frederick B., 147, 176 n.11, problem of source of; God, role in Mettrie and, 12; materialism and, 8, 12, Gaissinovitch, A. E., 103, 142, 143, 158,
182 n.1, 182 n.5 development) 98, 112; Maupertuis and, 13-15, 18, 178 n.2, 181 n.4, 181 n.8, 181 n.9, 181
Clarke, Samuel, 119, 180 n.7 embryology: causal, 153; descriptive, 151; mechanistic theories of, 1, 3-4, 13, n.18
Cole, F.J., 70, 71, 175 n.9, 176 n.7, 178 151-52, 153; German, 150-54, 151; mechanistic theories of Galen, 3, 87, 175 n.11
n.5, 178 n.24 155-56; natural history and, 153; inadequate,5,7-8, 14-15, 19,151, Galileo, 4, 95, 96
Collegium Medico-Chirurgicum Newtonian mechanism arid, 18; 152; monsters and, '126; Needham Garden, George, 8
(Berlin),46 philosophical questions in, 2, 8, 12, 88, and, 18-19, 120-21; Oken and, 150; Gasking, Elizabeth, 35, 84, 1511, 176 n.7,
conception, Hailer's theory of, 41, 44, 89,150,156 Pander and, 150; proof of God and, 176 n.9, 178 n.113, 178 n.114, 1811 n.7
17°7 n.22 embryonic circulation, development of, 111, 119, 141, 168; rise of at end of genera, 1118, 1119, 130,138, 1411, 143
Condorcet, Marie jean Antoine Nicolas 50; allantoic circulatory arc and, 50, eighteenth century, 150; spontaneous generation, see emboitement;
Caritat, Marquis de, 175 n.2 77; umbilical circulation and, 50; generation and, 32, 120-21; embryological development;
Cotes, Roger, 102 vitelline circulatory arc and, 50, 51, teleological, 151-52, 153, 154, 155; epigenesis; God, role in development;
66 two-semen theory of, 3, 4, 13-14, 16- preformation
deductive reasoning, Wolff on, 105-7, embryonic differentiation, Roux vs. 18,174 n.2; von Baer and, 150,154, germ layer theory, 811, 152, 179 n.13
119, 122, 149 Driesch on, 149 156; Wolff's religious beliefs and, 111- Gessner.Johann, 177 n.21, 183 n.9
deer, Harvey's observations on, 3, 14, 36, embryonic membranes, formation of (see 12, 141,166, 168;,Wolff'stheoryof, globules, Wolff's theory of, 85, 86, 179
177 n.22 also membrane-continuity proof; 45,4~51, i47 . n.15; Haller on, 85-86
"De formatione intestinorum," see Wolff, vitelline membrane; yolk-sac essential force, see vis essentialis God (see also epigenesis, proof of God
works membranes), 71-72, 75, 76 Euler, Leonard, 46, 82-83 and; preformation, prqof of God and;
208 Index Index 209
God (cont.) 100, 102;onexplanation, 119, 122;on Haller, works: Bibliotheca anatomica 149-50; philosophical issues in, 2, 32,
religion): as Creator, 1, 2, 8, 22, 30, 36, forces, 28--32, 35-36, 43, 96, 97-g8, (1774-77),91, 173, 1800.7, 1800.11; 45,88,89, 110, 118--19, 121,148,150;
43-44,91-g2,96,99, 101-2, 110-12, 12 1; on heart, formation of, 39, 42, 44, "De partibus corporis humani unresolvable nature of, 87, 149
n9, 121,122, 142-43, 145;monsters 64, 66-68, 69-70; on intestines, sensilibus et irritabilibus" ( 1752b), 33, Harvey, William: on area vasculosa,
and, 125-26; role in development, 8, 9, formation of, 82; irritability and 92, 98, 1 oo; Dest:riptio foetus bicipitis formation of blood vessels in, 51; on
14, 29-31, 32, 35, 43-44, 91-g2, 111- sensibility, theory of, 32-36, 98, 100, ( 1739), 22, 126; Dewc Memoires sur la ' colliquament, 54; Pescartes and, ~48,
12, 125-26, 141,142; as source of 160; La Mettrie and, 33, 90, 91, 98, formation des os ( 1758b), 39; Eltmenta 182 n.1; epigenesis and, 3, 14, 51, 65
adaptation, 139-41; as source of 119, 122, 180 n.6; Maitre-Jan and, 72- physiologiat corporis humani ( 17 57-66), 174 n.3; Exercitationes·de generatione
material forces, 30, 35, 43, 97-g8, 101, 73; materialism and, 33, go, 91, 92, 98, 35,42-44,45,62-63,69,72,78,86c...87, animalium (1651), 3, 51, 65, 167; on
146, 151 102, 112,119,122, 145; on matter, 29- 88,94,9o-g7,98,111,112-13,120-21, heart, formation of, 65; on
Goethe.Johann Wolfgang von, 178 n.2, 30, 97-g8, 100-102, 120, 146; 160, 165, 166, 174 n.14, 176 n.15, 177 metamorphosis, 6; observations on
179 n.4 mechanism and, 35, 44, 89, go, go-gg, n.22, 180 n.11, 180 n.14; 182 n.2; chicken eggs, 3, 167, 1_79 n.10;
Gottingen, Royal Society of Sciences of, 102, 110, _121, 148; membrane- Epistolarum ab eruditis viris ( 1773-'75), observations on deer, 3, 14, 36, 177
33, 40 . continuity proof, 40, 44, 70-71, 72-73, 55, 59, 88, 111, 119-20, 158, 178 n.1; n.22
Gottingen, University of, 22, 154 77-79, 178 n.24; Needham and, 31-32, ''Gedanken iiber Vernunft, heart, embryonic, see foramen ovale,
Giittingische A nuigen von gelehrten Sac hen, 119, 120-21, 122, 176 n.13, 176 n.14; · Aberglauben, und Unglauben" ( 1729), structure of
45,55,61,82, 1750.4 on Needham's generation theory, 31- 99; /cones anatomicae (1743-54), 92, heart, formation of (see also Haller, on
Graaf, Regnier de, 9, -175 n.9 32, 120-21, 176n.13, 176n.14, 178 171-73; "Observationes anatomicae heart, formation of; heartbeat; Wolff,
Graafian follicle, 175 n.8 n.25, 181 11.19; as a Newtonian, 2, 89, Bernenses" (1753-65), 36, 179 n.g; on heart, formation of), 65-66;
'sGraxesande, Willem Jacob van, 99, 180 96, 99-102, 119, 121, 148, 180 n.7; Opera minora (1762-68), 32, 63, 77, 94, Descartes on, 4; Hailer's belief in
. n.7 nonscientific writings of, go; on 165, 166, 167, 168, 176 n.14; 179 n.g; epigenesis and, 23, 24, 25, 39; Hailer's
gravity, 99; as analogy in embryology, 13, observation in physiology, 92-g3, 100; Praelectiones academicae ( 1739-44), 21, conversion to preformation and, 39,
17; as analogy in physiology, 100; God observations on chicken eggs, 26, 36, 22, 23-24, 32-33, 174 n.5, 175 n.3; 42, 44, 67-68; Haller-Wolff debate
as source of, 30, 35, 98, 1o 1; irritability 38,3g-40,43,44,5g-60,61-62,63, Preface to Allgemeine Historit der Natur, and, 46, 60, 64-65; Harvey on, 65;
and, 33, 35, 98, 100; nutritive force 66-67, 67-68, 69, 70, 72, 77, 78, 92-g4, vol. 1, by Buffon (175oa),95, 180 n.11; Malpighi on, 39, 42, 44, 65, 66-67
and, 115; vis essentialis and, 117-18 158, 177 n.19, 177 n.21, 179 n.g; Preface to Allgemeine Historie der Natur, heartbeat: development of, 51, 66;
Guthke,Kar!S., 175n.4, 18on.1, 18on.3, observations on shc;ep with vol. 2, by Buffon (1752a), 26-28, 28-- Haller on, 68, 69; Harvey on, 65;
180 n.6, 180 n.7 Kuhlemann, 36, 38, 43, 177 n.22; 30, 31, 32, 36, I 76 n.13; Primae lineae Harvey vs. Descartes on, 148-49, 182
occupations of, go; ovist preformation, physiologiae (1747), 24-25, 33, 35, 97, n.1; as initiator of development,.36, 41,
theory of, 36, 41'-42, 45; philosophy of 175-76 n.6; (1751c), 33, 175-76 n.6, 44, 64, 177 n.22; irritability and, 32-
Haecke-1, Ernst, 182 n.4 science of, go-102, 119, 121-22; on 176 n.15; (1765b), 24-25, 33, 35, 91- 33, 36, 41, 44, 64, 177 n.22; Malpighi
Halle, University of, 45, 46, 48, 102, 103 physiology, 9o-g7; as a poet, go, 92, 99; 92, 175-'76 n.6; Rijlexions sur le systirM on, 65; Wolff on, 68-69
Haller, Albrecht von: on anatomy, as a preformationist, 2, 21, 89, 91, 1 11, de la generation de M. de Buffon (1751a), heredity (see also hybrids; resemblance to
92-g3; animalculist preformation, 120, 158, 176 n.6; Professor of 28; Sammlung lueiner Halltrischer parents): Wolff on, 123, 124, 126, 135-
belief in, 21-22; on animal mechanics, Anatomy, Surgery, and Medicine Schriften ( 1756), 175 n.4, 176 n.8; 36, 143, 155 .
97, 99; animism and, 33, 98, 122; on (Gottingen), 22; rationalism and, 90, ( 1772b), 26, 32, 175 n.4; Sur la Herrlinger, Robert, 178 n.2, 182 n.6
area vasculosa, formation of blood 95-g6, 110; on religion and science, formation du coeur dans le poulet ( 1758a), Hertwig, Oscar, 182 n.5
vessels in, 53-54, 55-57, 5g-63, 83; on 9o-g2, 110; religious beliefs of, 2, 44, 21,26,40-41,42,45,53-54,63,66, Hintzsche, Erich, 40, go-g1, 99, 175 n.2,
ath~sm, 9o-g2, 98, 102, 112, 119; 89,go-g2,97-g9, 101-2, 110-11, 119, 67-68,70,72,73,77,92,158--59,176 176n.7, 177n.21, 18on.6, 18on.7
Bacon and, 95, 96; biographical 121, 148, 149, 178 n.25; returns to n.9 Histoire de rAcadimit Royale des Sciences,
information on, 21, 175 n.1; Bern, 36; as reviewer, go, 175 n.4; Haller, works reviewed by: "De 10-12, 183 n.7
Boerhaave and, 21, 22, 91, 96, 97, 102, Spallanzani and, 32, 176 n.14; studies formatione intestinorum" (Wolff), 82; Hoffmann, Friedrich, 102, 103, 107, 180
119,175 n.3; on bone, formation of, at Leiden, 21, 89, 96, 99; travels to Histoire naturelle, vol. 2 (Buffon), 28, n. 17; Fundamenta JMdicinae ( 1695),
39; Bonnet and, 36, 49, 57, 73-74, 119; England, 22, 99; on vivisection, 92; 30-31, 101-2, 176n.1o;OeuvresdeM. 107; mechanism and, 107; Stahl and,
on Buffon's generation theory, 26-31, Whytt and, 33, 98, 122, 180 n.6; Wolff Thomas, vol. 4, 95; Philosophia rationalis 103,107,149, 18on.17, 182 n.1
35, 36, 43, 62, 90, 1 i 2, 119, 120, 121, (Caspar Friedrich) and, see Haller, (Homann), 95-g6; Principia (Newton), hybrids (see also resemblance to parents):
122,177 n.19, 178 n.25; conversion to correspondence, with Wolff; Haller- 180 n.7; Thtoriagenerationis (Wolff), Hailer's epigenetic theory of, 25;
epigenesis, 21, 22, 23-24; conversion Wolff debate; Wolff (Christian) and, 45,55-56,69,85-86, 119,120,159, Hailer's preformationist theory of, 42,
to ovist preformation, 21, 26, 36, 39- 95-g6 163-64; Theorit .von der Generation 43; Haller-Wolff debate and, 2;
40, 44, 55, 67-68, 98, 110-11, 176 n. 7, Haller, correspondence: with Bonnet, (Wolff), 45, 61, 69; Traiti d'insectologie Koelreuteron, 138; Linnaeus on, 138;
1760.13, 1770.19, 1780.23, 1780.24, 36-40,57,5g-62,66-67,69,72,74,77, (Bonnet), 23, 24, 26 Maupertuis on, 14; preformation and,
178 n.25; Descartes and, 95-g6, 122; 83,85,95,98, 176n.13, 177n.18, 177 Haller-Wolff debate, 2, 45.:..46, 87-88, 14, 19; species and, 136, 138, 143;
Diderot and, 178 n.25; education of, n.20, 178 n.25, 179 n.7, 183 n.3; with 110, 143, 148, 166; embryological Wolff on, 136,138,143
21, 89, 96; empiricism and, 90, 95, 96, Caldani, 177 n.21; with Gessner, 179 issues in, 46, 88, 110 (see also area hypotheses, 95, 100
110, 121; epigenesis, theory of, 24-25, n.21, 183 n.9; with Morgagni, 177 vasculosa, formation of blood vessels
31, 35, 176 n.6; as an epigenesist, 21, n.21; with Somis, 39-40, 177 n.21; with in; heart, formation of; intestines,
38,42,44,91, 110,120, 177n.19; Tissot, 177 n.21; with Wolff, 45, 55, 59, formation of; membrane-continuity insect development: metamorphosis and,
"evolution," use of term, 175 n.5; on 88,111, 119-20, 124,141, 158--73, 178 proof; observability of embryonic 6, 7; as proof of preformation, 6, 176
experimental method, 92-g3, 95, 97, n.1 structures); incommensurability and, n.13
2 10 Index Index 2 I I
intestines, formation of, 78; membrane- definition of, 144-45; La Mettrie and, Lemery on, 22, 125; preformation and, Oppenheimer,JaneM., 154, 175n.7, 179
continuity proof and, 40, 70-71, 72- 12, 33,91, 98, 119, 122, 145; Needham 22, 125-26; Winslow on, 125-26 n.13, 182 n.4, 182 n.7
73, 74, 76, 77, 78, 166; reception of and, 176 n.14 motion: Halleron,97--98, 18on.14; ovism (see also preformation): de Graaf
Wolff's views on, 46, 83; Wolff on, 46, materia qualificata vegetabilis (qualified Newton on, 101, 18on.14 · and, 9, 175 n.8; Harvey and, 3; Maitre-
63, 72, 79--82, 83, 154, 179 n.13 vegetable matter), 129--31, 132, 133, moule interieur (internal mold): Buffon Jan and, 73
irritability: animism and, 33, 98; 135, 136, 137, 137-38, 139, 140, 141, on, 16--17, 18, 28,153; Hailer's critique ovum, discovery of mammalian, 175 n.8
development of preformed organism 143, 145-47, 155 of, 28----29
and, 36, 41, 44, 64, 177 n.22; as a force, matter (see also Haller, on matter): forces Muller, Fritz, 182 n.4 Pallas, Peter Simon, 128
33, 35, 43, 98, 100; God as source of, and,29-30,35,97--98, 100-101, 102, Mursinna,_Christian Ludwig, 178 ri.2 Pander, Heinrich Christian, 83, 150, 179
30, 35, 98; Hailer's conversion to 146, 151, 156; Newton on, 100-101; n.13
preformation and, 35-36; Hailer's passive nature of, 9, 15, 29--30, 97--98, Naturphilosophie, 155 ParisAcademyofSciences, 10,125,169
theory of, 32-35, 98, 1oo, 160; 100-101, 146,147,151; self-active Needham.John Turberville (see also parthenogenesis in aphids, Bonnet's
heartbeat and, 32-33, 36, 64, 177 n.22; nature of, 9, 15; Wolff on, 146 Haller, Needham and; Haller, on discovery'of, 23
materialism and,33, 98; Wolff on, 116, Maupertuis, Pierre-Louis Moreau de Needham's generation theory; Wolff, Pasteur, Louis, 149, 182 n.1
t 147, 160 arbrede Diane and, 13, 176 n.12; on 'Needham and): on animalcules in Patten, Bradley M., 50, 66, 75, 183 n.9
attractive forces, 13, 18; Buffon and, infusoria, 18, 19, 32, 176 n.13; Buffon, Perrault, Claude, 3, 7-8, 9
Kant, Immanuel, 151, 152 18, 19, 176 n. ll; epigenesis and, 13- collaboration with, 16, 18, 19; plants: classification of 137; as different
kidneys, Wolff on formation of, 48----49 15, 18, 151; on Harvey's observa1ions, epigenesisand, 18----19, 176n.14; from animals, Wolff on, 69--70, 106--7,
Kielmeyer, Carl Friedrich, 150, 153, 156 14; on matter, active nature of, 15; materialism and, 176 n.14; Nouvelles 130, 16o; formation of, as evidence for
Koelreuter,Joseph Gottlieb, 138 preformation, opposition to, 13-14; as Observations microscopiques ( 1750), 18, preformation, 5, 48; formation of,
Kuhlemann,Johann Christoph, 36, 38, a Newtonian, 18; Reaumur's criticism 19; Philosophical Transactions ofthe Royal Wolff on, 48; variation and, 127-29,
43,177 n.17, 177 n.22 of, 14-15, 25; Systemede la nature Socidy, paper in (1748), 18; Spallanzani 13~31, 135,139.140
(1751); 15, 178n.25; Vinusphysique and, 19, 32, 149; on spontaneous polyp, freshwater (see also regeneration):
La Mettrie, Julian Offray de: epigenesis (1745), 13-14, 15, 19, 176n.11 generation, 19, 119--20; on vegetat~ve epigenesis and, 12, 24; Haller's
and, 12; L'Hommemachine (1748), 98, Mazzolini, Renato G., 176 n.7, 176 n.14, force, 18----19,31-32, 113, 120-21 conversion to epigenesis and, 22, 23,
145; materialism and, 13, 33, 91, 98, 177 n.19, 177 n.22, 178 n.24, 178 n.25, Neuhaus.Johann Rudolf, 96 24, 25; materialism and, 12;
119,122,145; see also Haller, La 179 n.14 Newton, Isaac (see also Newtonianism): multiplication by artificial division and,
Mettrieand mechanical medicine, Wolff on, 108--g, Hailer's praise for,91, 99, 119; Hailer's 10; preformed germs and, 11, 12, 42-
Lawrence, William, 149, 182 n.1 122 review of Principia, 180 n. 7; on 43; reproduction by budding in, 10,
Leibniz, Gottfried Wilhelm von, 9, 105, mechanism ( see also Haller, mechanism hypotheses, 100; on matter and forces, 11; Trembley's discovery of, g--10
119,122,147,181 n.10 and; Wolff, mechanism and): 100-101; on motion, 101, 180 n.14; Pouchet, Felix-Archimede, 149,182 n.1
Lemery, Louis, 22, 25, 125 Blumenbach and, 151; Descartes and, Opticks(1730), 100-101, 18on.7; preexistence, see. preformation,
limbs, Wolff on formation of, 59, 74, 161 4; embryology and, 1, 2, 3-5, 7-8, 15, Philosophiae naturalis principia preexistence and
Linnaeus, Carl, 13 7, 138 151,156; Haller-Wolff debate and, 2, mathematica (1713), 1,100,101,102, preformation (see also embo,tement:
logic, Wolff on, 105-7, 119,122,149 110, 148; Hoffmann and, 103, 107, 180 n.7; religion and, 101, 102 Haller, animalculist preformation,
Lukina, Ta(jana A., 114, 124, 178 n.2, 149; Kant and, 151; Maupertuisand, Newtonianism (see also Haller, as a belief in; conversion to ovist; ovist
181 n.4, 181 n.5, 181 n.9 15; in the physical sciences, 1; in Newtonian): Boerhaave and, 99, 102; preformation, theory of): animalculist,
lungs, Haller on formation of, 39 physiology, I, 4, 96--g7, 107; Stahl and, Buffon and, 18; 'sGravesande and, 99; 1, 8, 9, 23; and area vasculosa,
103, 107, 149; vitalism and, Stahl vs. Maupertuis and, 18; religion and, 101, formation of blood vessels in, 51, 53,
Maitre-Jan, Antoine, 72-73, 179 n.10 Hoffmann, 103, 107, 149 102 54; Blumenbach and, 154; Boerhaave
Malebranche, Nicolas, 156; Descartes Meckel,Johann Friedrich, 83, 153, 154, nisus formativus, see Bildungstrieb and, 21,211, 23; Bonnet and, 12, 19, 20,
and, 5; on emboitement, 3, 5, 84; on 182 n.4 Novi Commentarii Academiae Scientiarum 40,45,57,6o,73-74, 150,151, 182n.3;
observability in embryology, 84; membrane-continuity proof (see also lmperialis Petropolitanae, 169, 183 n:4 Buffon's opposition to, 15-16, 19;
preformation and, 5, 6, 7,151; Haller, membrane-continuity proof; nucleus of Pander, 179 n.10 definition of, 1; demise of, 150;
Recherche de la viriti ( 1674), 5, 84 Wolff, on membrane-continuity nutrition: Buffon on, 16; St. Petersburg embryo in fluid state ~nd, 41-42;
Malpighi, Marcello, 175 n.6, 179 n.3; on proof): Bonnet on, 40, 73-74; Cole's Academy prize question and, 114-15; epigenesis vs., 11, 53, So, 153, 182 n.5;
area vasculosa, formation of blood illustration of, 71; Haller-Wolff debate Wolff on, 108, 115 evolution and, 175 n.5; Garden and, 8;
vessels in, 51-53, 54; chorion of, 76, and, 46, 78----79; Maitre-Jan and, 72-73; generic, 152; God as Creator and, 8--g,
167; Dissertatio epistolica de formatione ovist preformation and, 178 n.24 Objecta meditationum pro theoria'. 43-44, 91--92, 125-26 (see also God, role
pulli in ovo ( 1673), 6, 5 2, 54; Haller on metamorphosis: Goethe's theory of, 179 monstrorum, see Wolff, works in development); Haller and
heart observations of, 39, 42, 44, 66-- n.4; Wolff's theory of, 48, 137, 179 n.4 observability of embryonic structures (see Kuhlemann's observations on sheep
67; on heart, formation of, 39, 42, 44, microscopes, 6o, 84, 85, 164, 179 n.14, also transparency, development of and, 36, 177 n.17; Hailer's religious
65, 66--67; on heartbeat, 65; 179 n.15 embryo·and), 38, 41-42, 55-57, 57-58, beliefs and, 91--92, 1 10-1 11 ; hybrids
observations on chicken eggs, 5, 6, 174 monsters, formation of (see also 85-87; Haller-Wolff debate and, 83; and, 14, 19; insect development as
n.5, 179 n.10; preformation and, 6, 7, sexdigitism; Wolff, on monsters): by Malebranche on, 84; use of vinegar or evidence for, 6, 176 n.13; and
51-53 accident, 22, 125, 126; Duverney on, alcohol and, 41-411, 811, 84-85, 86-87, intestines, formation of, 8o-81;
materialism (see also Haller, materialism 125; byepigenesis, 126; Haller on, 22, 164-65,167 Kuhlemann and, 177 n.17; Leibniz
and; Wolff, materialism, and): 125, 126; Haller-Wolff debate and, 11; Oken, Lorenz, 150; 153, 179 n.13 and, 181 n.10; Maitre-Jan and, 711-73;
212 Index Index 213
preformation (cont.) religion and science, Haller on, 9o-g2, 31,176 n.13; Linnaeus on, 137, 138; vegetation (see also Wolff, on vegetation):
Malebranche and, 5, 6, 7, 151; 110 Ray on, 137, 138; taxonomy and, 137- degree of, 129, 135; mode of, 129, 130,
Malpighi and, 6, 7, 51-53; reproduction, see embryological develop- 38, 139:Tourneforton, 137,138 131,133,135,145 -
Maupertuis's opposition to, 13-14; ment; epigenesis; preformation spermaticism, see animalculism vegetation point, see metamorphosis
mechanismand,5,8, 15,146,151,156; resemblance to parents (see also hybrids): spinal column: formation of, Wolff on, vena cava, inferior, 169, 17.0, 171, 17 3,
monsters and, 22, 125-26; ovist, 1, 9, Halter's critique of Buffon on, 28; 48, 144; secretion of organs from, 183 n.5, 183 n.9 _
2 3; pans perm ism and, 7; Perrault and, Hailer's theory of epigenesis and, 25; Wolffon,59,74,80, 144,161 vena cava, superior, 183 n.9
7-8; plant development as evidence Maupertuis on, 14; preformation and, spontaneous generation: Hailer's vesicles, Wolff on formation of, 48, 49,
for, 5, 48; popularity of, 9, 174 n.5; 14, 19; Wolf-fon, 127,137,145,155 conversion to preformation and, 32; 80, 179n.3
preexistence and, 174 n.1; proof of Roe, Shirley A., 176 n.9, 176 n.16, 180 Haller'soppositiori to, 32, 119, 120-21, vis essentialis (essential force; see also
Godand, 111,119,168, 178n.25; n.7 122; Haller-Wolff debate and, 2, 32, Wolff, on vis essentialis): as attraction
origins of theory, 2-3; rejection of, in Roger,Jacques, 6, 9, 12, 15, 26, 35, 125, 119, 121;,Needham and, 19, 119-20; , and repulsion, 115-18, 145, 147;
Germany, 150,152; religion and, 151, 174 n.1, 174 n.5, 175 n.7, 175 n.9, 175 Needham vs. Spallanzani, 149; Pasteur Blumenbach's Bildungstrieb and, 1 16,
156,168; resemblance and, 14, 19; n.10, 176n.13, 181 n.7;181 n.10 vs. Pouchet, 149; preformationists' 122, I18, 154; building forces and,
Spallanzani and, 19, 20, 150; Rouhault, Pierre Simon, 183 n. 7 opposition to, 19; Spallanzani's u6-18, 121, 145, 154; gravity and,
Swammerdam and, 6, 176 n.13; Wolff Roux, Wilhelm, 149, 153, 182 n.1 refutation of, 19.32; Swammerdam's 117-18; Hailer's objections to, 112-13,
(Caspar Friedrich), final theory and, Ruysch, Frederik, 92 opposition to, 7; Wolff on, 119-20, 118, 119, 122, 181 n.18; life and, 114,
147; Wolff(Caspar Friedrich), 159-00 116; Needham's forces and, 113;
opposition to, 58--59, 8o--81, 111-12, St. Petersburg, Wolff's move to, 46, 79, Sproegel,Johann Adrian Theodor, 159, Stahl's vitalism and, 116
142; Wolff(Christian) and, 181 n.10 89,168 182 n.1 vis insita (innate force), 33, 176 n.15 -
Primae lineae physiologiae, see Haller, St. Petersburg Academy of Sciences, 45- Stahl, Georg Ernst: animism and, 107, vitalism, Wolff and, 103, 107-8, 109-10,
works 46,47-48, 79,114, 124-25, 168,183 149; Haller and, 33, 98, 122; 116, 122, 147, 18o--81 n.18
n.8 Hoffmanq and, 103, 107, 149, 180 vital principles, Lawrence vs. Abernethy
Rajkov, B. E., 124, 143, 178 n.2, 181 n.1, Schuster.Julius, 158, 178 n.2 n.17, 182 n.1; see also Wolff, Stahl and on, 149
181 n.3, 181 n.4, 181 n.8, 181 n.9 science: debates in, 148--49; stomach, formation of, 80, 81 vitelline membrane, 71, 72, 75, 77, 78, 79
rational anatomy, Wolff and, 103,122, extrascientific influences on, 148; as sufficient reason, principle of (see also Voltaire, Fran,ois Marie Arouetde, go,
147 route to God, 91-92, 102 rationalism): Wolff (Caspar Friedrich) 119
rationalism (see also sufficient reason, secretion and solidification, Wolff's and,49, 105, 108-g, 117,119, 1111, 122, · Von tier eigenthumlichen und wesentlichen
principle of): Boerhaave and, 96; theory of, 48--50, 59, 74, 80, 105-6, 127, 142; Wolff (Christian) and,:. 105 Kraft, see Wolff, works
epigenesisand, 181-82 n.10; Hailer's 110,114, ~15, 118,129,130,144,145, Sur la formation du coeur dans le poulet, see
opposition to, 90, 95-96, 1 10; Haller- 161 Haller, works Weismann, August, 176n.11, 182 n.5
Wolff debate and, 1 10; Wolff (Caspar semen, female: existence of, 16, 17; Swarnmerdam,Jan, 3, 5-6, 6-7, 174 n.5, Whytt, Robert (see also Haller, Whytt
Friedrich) and, 2, 89, 103-7, 110·, 119, Hailer's rejection of, 28 176 n.13 and),33,98, tog, 122
122,123,139,146, 147,-148, 149,155, sensibility: Hailer's theory of, 33, 100; Winslow,Jacob, 125, 126, 183 n.7
181-82 n.10; Wolff (Christian) and, Wolff on, 147 Wolff, Caspar Friedrich: anatomical
1q3-5, 181 n.10 teleology: epigenesis and, 151..:.52, 153,
serosa, 75, 78 papers of; 181 n.3; on area vasculosa,
Ray.John, 137,138 154, 155; German embryology and,
sexdigitism, 126, 136, 142, 143 formation of blood vessels in, 54-55,
Reaumur, Rene Antoine Ferchault de, 9, 151-52, 153; Kantand, 151,152;
shell membrane, 75, 76 57-59,63-64,86,115;161-62,164-65,
10, 14-15, 18,25 phiJosophical basis for, 155-56; Wolff
sinus terminalis, 50, 51, 64 179 n.6; on atheism, 119, 139, 145;
reductionism: Haller and, 97; Wolff's and,154,155 biographical information on, 46-48,
solidification, in plants and animals Theoria generationis, see Wolff, works
opposition to, 89, 103, 108, 110, 122, compared, Wolff on, 69, 106-7, 130, 178 n.2; Blumenbach and; 114-15,
147,148
_Theorie van tier generation, see Wolff, works
146 116,118,154;Bonnetand,45,57,60,
regeneration: Bonnet and, i2, 2·2-23, Toellner, Richard, 90;91, 96, 176 n.9,
solidification and secretion, Wolff's 82-83, 163,164,170, 182-83 n.3; Born
42-43; in freshwater worms, 22-23; 176n.13, 176n.16, 18on.1, 18on.3,
theory of, see secretion and 182 n.1 and, 114-15; Buffon and, 127, 139;
Hailer's conversion to epigenesis and, solidification, Wolff's theory of correspondence: with Euler, 82-83;
22, 23, 24, 25; Hailer's explanation of, Tournefort,Joseph Pitton de, 137, 138
Somis, Ignazio, 39-40, 177 n.21 with Haller, 45, 55, 59, 89, 111, 119-
as preformationist, 42-43; Haller- transformism, Wolfhnd, 132-33, 143
soul (see also animism): Haller on, 24, 98; transparency, development of embryo 20, 124, 141, 158--73, 178 n.1;
Wolff debate and, 2; Perrault on, 9; Stahl on, 107, 109-10, lt6; Whytton, Descartes.and, 1o6, 122; education of,
polyp and, 10, 11; preformed germs and (see also observability of embryonic
33; Wolff on, 109-10, 116,140,141, 46, 89, 102-3; empiricism and, 107,
and,9, 11, 12,42-43;Reaumuron,9; structures), 38, 41, 42, 57, 63, 68, 6g,
144, 145, 181 n.18 110, 122; epigenesis, theory of, 45, 48--
Thevenot on, 9 70,79,82,85,86 / 50, 147; as an epigenesist, 2, 89; on
Spallanzani, Lazzaro: Needham and, 19, Trembley, Abraham, 12, 22, 23; polyp
religion (see also God; religion and 32,149; Opuscolidifisicaanimalee Ethiopians, 135, 136, 181 n.5; Euler
science, Haller on; science, as route to discovered by, 9-10
vtgetabile ( 1776), 19; preCormation and, 46, 82-83; evolution and, see
God): Hailer's belief in, 2, 44, 89, 90- and, 19,20, 150:spontaneous Wolff, transformism and; on
92, 97-99, 101-2, 110-11, 119,121, generation, refutation of, 19, 32 variation (see also plants, variation and; explanation, 103-7, 112, 119,122,147;
148, 149, 178 n.25; Haller-Wolff species (see also variation; Wolff, on species; Wolff, on variation): on foramen ovate, 169, 170-73, 183
debate and, 2, 110, 148; Wolff and, species): archetype and, 152-53; environment and, 126, 127-29, 130- n.5, 183 n.9; Haller and, see Wolff,
llO, lll-12, 119, 122, 139-41, 144- Buffon on, 138--39; essentialist_ 32,133, 135-36, 138--39, 140,141,142, correspondence, with Haller; Haller-
45, 147,166,168 concepts of, 137, 139; Haller on, 30- 143; Haller on, 30-31 Wolff debate; on heart, formation of,
214 Index
Wolff, Caspar Friedrich (cont.) vitalism and, 103, 107-8, 109-10, 116,
55, 64-65, 68-69,·161; illustrations of 122, 147, 18o-81 n.18; von Baer on,
chick development, 49, 56, 81, 161, 83, 154, 179 n.13, 181 n.1; Wolff
162; influence of, 46, 83, 154, 179 (Christian) and, 103-5, 146; works
n.13; on intestines, formation of, 46, reviewed by Haller, 45, 55-56, 61, 69,
63, 72, 79-82, 83, 154, 179 n.13; 82,85-86, 119,120,159,163-64
Linnaeus and, 138; mat<;rialism and, Wolff, works: "De foramine ovali"
144-45; on materia qualijicata vegetabilis (1775), 124, 169, 183 n.4; "De
(qualified vegetable matter), 129-31, formatione intestinorum" ( 1766-67,
132,133,135,136,137, 137-38, 139, 1768),45-46,63-64,72,7g-82,83,
140,141,143, 145-47, 155; 154, 179 n.13; "Descriptio vituli
mechanism and, 89, 103, 108-g, 110, bicipitis" ( 1772), 124, 125, 126;
116,122, 147; on membrane- "Distributiooperis," 141-43, 181 n.8;
continuity proof, 74-79, 83; on Objecta meditationum pro theoria
monsters, 46, 124-25, 126, 132, 133- monstrorum(1973), 124, 126-37, 138,
36, 141-42, 143, 144, 170; moves to St. 139-40, 141,145,181 n.2, 181 n.5;
Petersburg, 46, 89, 168; Needham and, Theoria generationis ( 1759), 45, 48-50,
113, 1 1g-20, 15g-60; observations on 54,55,68,69,80,85, 103, 105-6, 108-
chicken eggs, 58-59, 68-69, 74-75, 75- 9, 111-12, 113-14, 119, 124, 138, 158,
76, 77, 78, 79, 161; on philosophical vs. 159, 160; Theorie von der Generation
historical knowledge, 103, 107; (1764), 45, 57, 58-59, 61, 68-69, 74-
philosophy of science of, 102-10, 119, 75, 79, 80, 86, 100-7, 111, 112, 114,
122,144,147; preformation, 120, 161, 163, 179 n.9, 183 n.3; Uber die
opposition to, 58-59, 8o-81, 111-12, Bi/dung des Darmkanals im bebriiteten
142; preformation and final theory, Hiinchen(1812),83, 154; Vonder
147; Professor of Anatomy and eigenthiimlichen und wesentlichen Kr.aft
Physiology (St. Petersburg), 48, 168; as (1789),46,64, 114, 116-18, 145,181
a rationalist, 2, 89, 103-7, 110, 119, n.18
122, 123, 139, 146, 147, 148, 149, 155, Wolff,Christian, 102, 103-5, 1'22, 146,
181-82 n. 1o; reductionism and, 89, 147, 1800.16, 181 n.10; Haller and,
103, 108, 110, 122, 147, 148; religion 95-96; on historical, philosophical,
and, 110, 111-12, 119, 122; 139-41, and mathematical knowledge, 103-5;
144-45, 147, 166, 168; on species, 126, mathematical method of, 105;
128-33, 135, 136-37, 137-39, 141, Philosophia rationalis ( 1728), 103-5;
142, 143, 146, 155; on spontaneous preformation and, 181 n.10;
generation, 1 1g-20, 15g-60; Stahl and, rationalism and, 103-'5, 181 n.10;
109-10, 116, 181 n.18; studies at sufficient reason, principle of, 105;
University of Halle, 45, 46, 48, -102, Wolff (Caspar Friedrich) influenced
103; teleological epigenesis and, 154, by, 103-5, 146
155; transformism and, 132-33, 143; Wolffian bodies, 49
treatise on monsters, 46, 124, 141;
unpublished materials of, 123, 124,
yolk: Harvey on, 167; Wolff on dissolving
141, 155, 181 n.1; on variation, 123,
of, 48, 54, 55, 115
124,126, 127-29, 130-33, 135-36,
yolk sac, 70-71, 72, 74, 76, 77, 78, 79,
141, 143, 146, 155, 181 ri.5; on
vegetation, 48, 49, 106, 108, 109-10, 166.-67
yolk-sac membranes, 40, 70-72, 75, 76,
114, 115-17, 123,126,127, 129-31,
77,78,80, 166-67
133,135,139,142,143, 146-47;onvis yolk stalk, 51, 72, 76, 80, 166, 167
essentialis (essential force), 48, 49-50,
54,55,64-65,68, 106,110, 113-14,
115-18, 122, 130, 145, 147, 154; Zimmermann.Johann, 92, 175 n.2