Elwick 2007 Styles of Reasoning in Early To Mid-Victorian Life Research

Journal of the History of Biology (2007) 40:35–69 Ó Springer 2006
DOI 10.1007/s10739-006-9106-4
Styles of Reasoning in Early to Mid-Victorian Life Research:

Analysis:Synthesis and Palaetiology
JAMES ELWICK
Science and Technology Studies
Faculties of Arts and Science and Engineering
York University
4700 Keele St.
M3J 1P3 Toronto, ON
Canada
E-mail: jelwick@yorku.ca
Abstract. To better understand the work of pre-Darwinian British life researchers in

their own right, this paper discusses two different styles of reasoning. On the one hand
there was analysis:synthesis, where an organism was disintegrated into its constituent
parts and then reintegrated into a whole; on the other hand there was palaetiology, the
historicist depiction of the progressive specialization of an organism. This paper shows
how each style allowed for development, but showed it as moving in opposite
directions. In analysis:synthesis, development proceeded centripetally, through the
fusion of parts. In palaetiology, development moved centrifugally, through the ram-
ifying specialization of an initially simple substance. I first examine a community of
analytically oriented British life researchers, exemplified by Richard Owen, and certain
technical questions they considered important. These involved the neurosciences,
embryology, and reproduction and regeneration. The paper then looks at a new
generation of British palaetiologists, exemplified by W.B. Carpenter and T.H. Huxley,
who succeeded at portraying analysts’ questions as irrelevant. The link between styles
of reasoning and physical sites is also explored. Analysts favored museums, which
facilitated the examination and display of unchanging marine organisms while pro-
viding a power base for analysts. I suggest that palaetiologists were helped by vivaria,
which included marine aquaria and Wardian cases. As they became popular in the
early 1850s, vivaria provided palaetiologists with a different kind of living and
changing evidence. Forms of evidence, how they were preserved and examined, and
career options all reinforced each other: social and epistemic factors thus merged.
Keywords: analysis, Charles Darwin, George Newport, museums, neurosciences,

palaetiology, regeneration, reproduction, Richard Owen, Robert E. Grant, serial
homology, styles of reasoning, synthesis, T.H. Huxley, W.B. Carpenter
36 JAMES ELWICK
Introduction
This paper introduces a new dichotomy for historians of nineteenth

century biology: analysis:synthesis and palaetiology. These two catego-
ries reveal differences unexplored by other dualisms such as form versus
function, philosophical anatomy versus natural theological functional-
ism, or epigenesis versus preformationism. Nor do they seem linked
with contemporary concerns about materialism. Most importantly, this
dualism allows us to better appreciate the aims of British pre-Darwinian
life researchers in their own right. I focus on the life sciences in Victo-
rian Britain – specifically London – between the late 1820s and the early
1850s, emphasizing marine invertebrate research. Obviously the usual
warnings about subtle differences and fine nuances ignored by this
theoretical scheme apply – all maps must omit certain points if they are
to highlight new and interesting features of the terrain.
Analysis:synthesis and palaetiology were styles of reasoning, self-
reinforcing norms for what counted as good research. I use ‘styles of
reasoning’ to group various historians’ terms for these categories. They
include Alastair Crombie’s ‘‘styles of reasoning,’’,further developed by
Ian Hacking; Ludwik Fleck’s ‘‘thought-styles;’’ Gerald Holton’s ‘‘the-
mata;’’ Nicholas Jardine’s ‘‘scenes of inquiry;’’ and John Pickstone’s
‘‘ways of knowing.’’ For Hacking, a style of reasoning sets out what it is
to reason rightly. It is epistemological. Historically, a style made certain
kinds of inquiries possible, and yet by helping a researcher commit to
solving certain problems, it also restricted and excluded alternative
inquiries.1 This partly stemmed from how evidence was used in different
styles – someone using one style of reasoning presupposed certain kinds
of evidence to be more relevant than other kinds. In turn how evidence
was generated, stored and examined helped to shape the favored
workplaces of each style. This helped set out possible career alternatives
for researchers. Exploring styles of reasoning helps us fuse researchers’
social and epistemic commitments.
This paper first compares analysis:synthesis and palaetiology. It then
closely examines the style of analysis:synthesis in the life sciences, and
some of the areas considered important by analysts:2 the neurosciences,
classification by nervous structure, development, and regeneration/
reproduction. Richard Owen was one researcher committed to analy-
1
The best overviews of this are in Crombie, 1988; Iliffe, 1998. Another discussion is
in Kusch, 1991, p. 94. See also Collingwood, 1939, pp. 29–30; Holton, 1988, pp. 41–42,
83–84; Harwood, 1993; Jardine, 2000, pp. 3–4, 77; Hacking, 2002, pp. 181–182.
2
For brevity I use this term to refer to those practicing analysis:synthesis.
STYLES OF REASONING IN EARLY TO MID-VICTORIAN LIFE RESEARCH 37
sis:synthesis. The paper then moves on to consider palaetiology in more

detail, and its emphasis on von Baerian embryology. T.H. Huxley and
W.B. Carpenter exemplify palaetiology. The paper ends with Owen’s
1858 attempt to reclassify mammals in an analytic:synthetic way, an
attempt which Huxley reinterpreted (or misinterpreted) palaetiologi-
cally.
A Comparison of Analysis:Synthesis and Palaetiology
A quick way to contrast the two styles is to appropriate two different

images used by Georges Canguilhem to explain the history of mor-
phology. Canguilhem saw morphologists as guided by one of two
irreconcilable pictures. Some saw living things as discontinuous: as
composites of discrete parts. Others saw organisms as continuous: all
growing out of an initial simple and plastic material.3 Canguilhem’s first
image nicely captures how analysis:synthesis portrays living things as
composites. His second can be used to depict palaetiology’s emphasis on
continuity and unity.
Analysis:synthesis has been closely examined by the philosophers of
biology William Bechtel and Robert C. Richardson. And it has been
extensively discussed as a ‘‘way of knowing’’ by historian of medicine
John Pickstone.4 Analysts thought that the best kind of knowledge was
obtained by disintegrating systems or organizations into their simpler
constituent parts, called elements, and then seeing those systems as
nothing but aggregations of those elements. Organisms were one such
kind of system. Analysis:synthesis was thus a doublet. It gained strength
in Britain in the late 1820s – imported largely from France, by British
researchers with an inferiority complex. They used it to strengthen their
hold on museums, where analysis:synthesis mattered the most.
Museums and analysis:synthesis went well together because museums
could store vast amounts of elementary structures, which could then be
compared. This favored the sedentary researcher over the field natu-
ralist, a point noted by Georges Cuvier himself.5 This kind of evidence is
unchanging. In marine invertebrate zoology of the time, for instance,
museum evidence consisted of bleached specimens kept in wine-spirit
filled glass jars. The individuals, their tissues, and their cells could not
interact or transform.
3
Canguilhem, 1994, p. 163.
4
Bechtel and Richardson, 1993, pp. 18–21; Pickstone, 1993; Pickstone, 1994.
5
Outram, 1996, pp. 259–262.
38 JAMES ELWICK
Meanwhile the term ‘palaetiology’ was coined by philosopher Wil-

liam Whewell in his 1837 History of the Inductive Sciences.6 Palaetiol-
ogists espoused that the best kind of knowledge was understanding how
organizations and systems changed over time, becoming more complex
out of simple origins. Whewell saw palaetiologists studying how ‘‘phe-
nomena at each step become more and more complicated, by involving
the results of all that has preceded, modified by supervening agencies.’’7
He was not so much a pontificator - setting out rules that ought to be
followed by researchers – as someone who recognized similar practices.
Whewell can thus be seen as listing ‘‘exemplar disciplines’’ to be prof-
itably copied by other fields.8
Palaetiology in turn was largely brought into Britain from
Germany – for example, Martin Barry imported von Baerian embry-
ology in 1837.9 It was thence used by some British life researchers to
challenge an older generation of British analysts. They cited evidence
that changed over time, which meant that palaetiological life research
usually occurred in different sites than analytical research. Against the
analysts’ museum, one relevant locus of palaetiologists’ activities was
the ‘vivarium.’ In the case of marine invertebrate zoology, this in-
cluded such places as marine aquaria and the Wardian case. Vivaria
facilitated the close control and observation of specimen-evidence over
longer periods. An observer, comfortably indoors, could over several
months watch a hydroid eventually change into a medusoid. Female
water fleas could be isolated from male ones to ensure that one was
indeed observing asexual reproduction. Vivaria only became wide-
spread by the late 1840s.
Many historians of biology are likely familiar with these styles,
particularly ‘palaetiology.’ But to call palaetiology ‘developmentalism’
is imprecise, for both styles of reasoning allowed for development. I
have chosen new terms because each style depicted different directions
of development. Consider again Canguilhem’s two images: the first,
discontinuous, style of analysis:synthesis showed development occur-
ring centripetally. An analyst saw an embryo appearing at a number of
separate points and then fusing together in a new center. It developed
6
Whewell, 1837, pp. 3:481, 486; Whewell, 1967b. I am deeply indebted to John
Beatty for suggesting this name and this approach. More recently there has been dis-
cussion of palaetiology on mid-1990s listservs, such as Robert J. O’Hara’s at http://
rjohara.net/darwin/palaetiology.html, retrieved March 3 2006.
7
Whewell, 1967a, p. 3:399. On the importance of palaetiology, see Hodge, 1991.
8
Thoughts on ‘‘exemplar disciplines’’ are found in Jardine, 2000, pp. 103–104.
9
Barry, 1837a; Barry, 1837b.
by moving inward – making ontogeny the fusion, or synthesis, of

parts. A good example of synthetic development was insect meta-
morphosis, when the serially repetitive segments of a larva coalesced
and concentrated during the pupal and imago stages. Analysis and
synthesis were related, as analysis disintegrated a system into its
components, and synthesis put those components back together again.
Perhaps in using this style, early 19th century life researchers sought to
copy the most successful science of the day, Newtonian natural phi-
losophy.10
On the other hand palaetiologists saw development occurring cen-
trifugally, with the embryo emanating outward from a single, and
simple, starting point. Embryos spread outward from this single spot.
Well-known examples of this centrifugal developmental direction in-
clude von Baerian embryology – the emergence of the heterogeneous
from a homogeneous mass – and the Darwinian ramifying tree of des-
cent with modification .
Historians as early as 1912 (Theodore Merz) pointed out a kind of
grand style that I have subsumed under ‘palaetiology.’ Those following
Merz have used different names – hence Dov Ospovat noted the
emergence of a ‘‘theory of diverging development’’ and saw von Baer’s
embryology as simply one manifestation of this theory.11 Meanwhile,
historians long before Pickstone have noted a prevalent style of analy-
sis:synthesis, including Merz and Elie Halévy, the famous economic and
political historian.12
Most importantly, seeing Richard Owen as an analyst:synthesist
makes his researches more coherent. In this light I suggest that
Thomas Henry Huxley especially succeeded in marginalizing his
bitter foe, Owen, by rewriting many parts of the history of biology
to make it seem as if the only kind of good life research was
palaetiological.13
10
Newton’s prism experiments are discussed below. Gravitation is another example.
Consider Scottish anatomist John Goodsir’s modification of Newton’s law of gravita-
tion for anatomy, imagining a ‘‘law of production’’ based on the inverse cube of the
distance of two bodies rather than Newton’s square. Goodsir et al., 1868, pp. 2:212–213.
11
Merz, 1965, pp. 2:279–280; Ospovat, 1976, p. 27; Coleman, 1977, p. 10; Farley,
1982, pp. 75–81; Alter, 1999.
12
Merz notes how the ‘‘morphological period,’’ ending in 1860, was replaced by the
‘‘genetic period’’ – Merz, 1965, pp. 2:214fn, 2:270–275. Halévy, 1928, p. 502.
13
Thus after Owen’s death Huxley wrote a review essay (much like Cuvier’s back-
handed elegies) on Owen’s anatomical work, in Owen, 1894.
Table 1. Styles of reasoning compared 40
Analysis:synthesis Palaetiology
What constituted valid knowledge? Disintegration of system (organism) into simple Understanding system’s growing complexity and
components; reintegration of those components into specialization from its simple origins
system
Contemporaries who noticed a James Mill; John Stuart Mill, Jeremy Bentham James Cowles Prichard; William Whewell
style
Historians who have noticed a style Elie Halévy; Theodore Merz; Georges Canguilhem; Theodore Merz; Alastair Crombie; William Cole-
Randall Albury; John Pickstone man; Dov Ospovat; John Farley; Stephen Alter
Institutions/sites Museums ‘Vivaria’ (aquaria, Wardian cases)
Major Continental researchers and E.B. de Condillac; Antoine Lavoisier; Georges Cu- K.E. von Baer; Matthias Schleiden; T. von Siebold
their styles vier; E. G. St-Hilaire; F-J Gall; H. Milne Edwards;
E. R. A. Serres. (J.F.Meckel)? (C.G.Carus)?
JAMES ELWICK
Major British researchers using/ Robert Grant; Thomas Laycock; Richard Owen; Martin Barry; W.B. Carpenter (convert); T.H.
importing styles George Newport; E.S. Forbes; Marshall Hall; Huxley; George Allman (convert); George Busk;
T. Rymer Jones Charles Darwin (convert)
Exemplary scientific fields Lavoisier’s chemistry; Geoffroy’s comparative anat- Comparative philology; Ethnology; Geological uni-
omy; Gall’s phrenology formitarianism; Paleontology
Focused life research questions Reflex theory; Homologizing; Cephalisation; Reca- Von Baerian embryology; sexual reproduction;
pitulationist embryology; Parthenogenesis/Metagen- Darwinian descent with modification
esis
Analysis:Synthesis Part 1: Analysis
Let us begin by dividing the style of analysis:synthesis into its own

respective components of analysis and synthesis. Analysis disintegrated
systems into their simplest elements. Following the Cartesian method,
Etienne Bonnot de Condillac stated that a machine could be best known
by decomposing it and studying each part separately; when the parts
were put back, the entire machine would be known perfectly. Analysis
was important in Lavoisier’s chemistry and Cuvier’s functional anat-
omy, with Lavoisier crediting Condillac’s method for his success.14 Other
early 19th century life researches using analysis:synthesis included bot-
any (Augustin-Pyramus de Candolle), morphology (Geoffroy Saint-Hi-
laire), and organology/phrenology (Franz-Josef Gall).15
Pickstone notes how analysis largely occurred in the physical site of
museums. Places such as the Muséum d’Histoire Naturelle were foun-
ded or reformed upon analytical lines, gathering researchers, specialized
tools, money and collections in one location.16 We can build on his
point: museums helped define research problems by entrenching anal-
ysis:synthesis. This term ‘entrenchment’ implies that museums were
shaped by myriad research choices and practical considerations; these
sites were the physical manifestations of small, local commitments to
learn about organisms by disintegrating them. This crystallization
reinforced the utility of analysis:synthesis, making it more reliable and
channeling people’s expectations that good life research would be car-
ried out analytically:synthetically.17
Many Britons saw analysis:synthesis as originating in France. Yet
many French authors saw it as having British roots – their favorite
example of analysis:synthesis being Isaac Newton’s use of prisms to
14
Condillac, 1980, pp. 79–81; Pickstone, 2001, pp. 84–85; Bechtel and Richardson,
1993, pp. 18–21; Simon, 2002, pp. 3–4.
15
Elements were not reducible into different domains – thus mental faculties would
not be reduced into chemical compounds. Pickstone, 1994, p. 117; Albury, 1977, pp. 90–
91.
16
Pickstone, 1994, pp. 117–118, 123–124; Appel, 1987, pp. 11, 18. Pickstone includes
teaching hospitals as ‘‘museological institutions,’’ ‘‘collecting’’ patients and classifying
them by universal diseases instead of by individual humoral imbalances.
17
On the entrenchment and ‘crystallization’ of research strategies I have been in-
formed by Gerson, 1998, pp. 26–28, Bourdieu, 1999, p. 36, and Schank and Wimsatt,
1986.
42 JAMES ELWICK
disintegrate white light into its constituent colors, then reconstitute that
spectrum into white light again.18 In the late 1820s analysis:synthesis
appeared or returned to London in great strength, because many British
researchers who had studied in Parisian medical schools and museums
came back to London. Others came down to London from Francophile
Edinburgh medical schools, as noted by Adrian Desmond. Indeed,
‘‘philosophical’’ or ‘‘transcendental anatomy’’ might be seen as ana-
lytic:synthetic, where bodies were depicted as collections of – and
anatomized into – simple, universal, and comparable parts. Figure 1
shows how social networks and styles of reasoning were intertwined: all
were commitments of one form or another.
Those using analysis:synthesis in the late 1820s and early 1830s
ranged from Radical to Conservative.19 They were unified by a sense of
inferiority to French science; amidst fears about a British ‘decline of
science,’ they imported analysis:synthesis from France to reform British
science. It is significant that museums sprouted up in Britain from the
late 1820s onward, in what Nicolaas Rupke has aptly called ‘‘the age of
museums’’ – during Richard Owen’s lifetime (1804–1892), some 200
British museums were built or renovated.20 I suggest that one reason for
this was to facilitate analysis:synthesis.
From workplaces we can move to technical issues. Analysis:
synthesis shaped various researches, including the neurosciences,
classification, comparative anatomy, development, and reproduction.
Findings in one field shaped others – neuroanatomical discoveries
were deployed to explain why complicated animals did not easily
regenerate. It is thus important to draw connections between these
contemporaneous topics.
18
Thus d’Alembert’s article ‘‘Analytic’’ in the Encyclope´die cited Newton’s decom-
position of white light into the rainbow as an example, quoting from the Opticks. A
common use of analysis:synthesis might also explain why certain researchers were de-
picted as the ‘‘Newton’’ of their fields. Bichat sought to become the ‘‘Newton’’ of
physiology; meanwhile Bentham was described as the Newton of the moral world by
Thomas Southwood Smith (recall Bentham’s attempt to explain all ethics as the
interplay simply of pain and pleasure). Albury, 1972, pp. 60–64, 230. Webb, 2004,
p. 312.
19
Conservative Francophobic analysts, however, tended to emphasize the British
roots of the style.
20
Rupke, 1994, pp. 13–14. Relevant museums include the British Museum (renovated
1837); the Museum of Practical Geology (founded 1835, moved in 1848); the Hunterian
Museum (renovated 1836); and the Cambridge University Museum of Anatomy (moved
to an enlarged location in 1832).
R.E.Grant J. Quain W.S.Macleay 1819 J.Dalyell 1814

1830 J.R. H.Mayo 1830
Bennett M.
d. 1831
Hall R.Owen
G.
P.M. Newport
d.1854
Roget
?
S.Solly
1835 T.Wharton Jones 1835
W.
Sharpey
R.D.
Grainger
C.
Darwin
M. Barry
W.B.
T.R. Carpenter
1840 Jones 1840
E.S.Forbes
d.1854 W.Baly
J.
Goodsir C.Darwin
H. Goodsir
d.~1845
T.
1845 Laycock G.Busk 1845
Researchers placed at
approximate rise to prominence
formally taught G.Allman
patron-client (arrow size
indicates importance) W.B.Carpenter
A.Henfrey
phrenological sympathies
1850 trained in: J.D. Hooker 1850
Paris T.H. Huxley
Edinburgh
London Univ./UCL G.Allman
ANALYSIS:SYNTHESIS PALAETIOLOGY
Figure 1. Some Life Researcher Commitments, 1830–1850.

44 JAMES ELWICK
The Analysis of the Nervous System
In the 1830s and 1840s, British researchers sought to reveal the simplest
possible elements of the nervous system. They followed Cuvier by using
nervous structure to classify. And this principle underlay any ideological
disagreements. On this Adrian Desmond’s exemplar Radical compara-
tive anatomist, Robert Grant, was similar to Desmond’s exemplar
Conservative anatomist, Richard Owen. Grant renamed the four dif-
ferent Cuvierian embranchements by using their nervous structure as an
index.21 Desmond explains this by pointing to Grant’s Lamarckian
transmutationism, claiming that Grant sought to unify Cuvier’s four
separate groups into a single series.22 But the anti-Lamarckian Owen
also used the nervous system in the same way – he coined new terms,
such as ‘‘homogangliata’’ and ‘‘heterogangliata’’, which denoted two
natural animal groupings.23 This use of nervous structure by ideological
opposites – both in 1836 – reveals problems other than the support or
dismissal of transmutation.
In that same year, Owen explicitly noted that he was following
Cuvier’s method: Cuvier deemed the animal’s primary characteristic to
be ‘‘sensibility,’’ a property best revealed by its nervous system.24
Owen’s contemporaries also thought that using the nervous system as a
taxonomic index was useful: one of Grant’s reviewers thought that this
method was ‘‘generally admitted by comparative anatomists’’ to be
best.25 The brain researcher Samuel Solly used Grant’s system, also in
1836. Later, Owen’s protégé, Thomas Rymer Jones of King’s College
London, noted how Owen, Grant and Cuvier all used the nervous
system for a ‘‘more natural method of classification’’.26
The ‘ganglia’ to which Cuvier, Owen, Jones and Grant referred was a
useful ‘‘element’’ because it was the simplest possible unit that com-
pounded into increasingly complex nervous structures. In the 1830s and
1840s the ganglion was seen as a nervous ‘‘knot’’ made of gray
21
Thus Grant renamed Vertebrata the ‘‘Spini-Cerebrata’’; Mollusca the ‘‘Cyclogan-
gliata’’; Articulata the ‘‘Diplo-Neura’’; and Radiata the ‘‘Cyclo-Neura.’’ Grant, 1836,
pp. 107–108.
22
Desmond, 1989, pp. 86–87.
23
‘‘Homogangliata’’ (Cuvier’s Articulata) denoted the repeating ganglion in each
segment; ‘‘Heterogangliata’’ (Cuvier’s Mollusca), their irregular dispersion. Owen,
1836b, p. 547.
24
Owen, 1836a, p. 244. On Cuvier’s use of the nervous system for hierarchical clas-
sification, see Coleman, 1964, pp. 85–91.
25
Anonymous, 1835, p. 381.
26
Solly, 1836, p. 4; Jones, 1841, p. 3. Jones also included John Hunter in this group.
‘‘neurine’’ – a tissue providing nervous power. Though there were dis-

agreements about how far the term Ôganglion’ ought to be extended,
researchers increasingly saw all of the nervous system – including the
brain and spinal column – as compounded ganglia.27 Simpler animals
had more decentralized nervous systems – the lower the organism, the
more dispersed its ganglia. Ganglia were often depicted as subordinate
brains, because their dispersion explained why the separated body parts
of simpler invertebrates could move independently. The distributed
ganglia of centipedes were described in Jones’s standard textbook as ‘‘so
many brains presiding’’ over nerve functions, while Cambridge Pro-
fessor of Anatomy William Clark described ganglia as ‘‘subsidiary
brains.’’28 Owen also noted that ganglia performed ‘‘the function of so
many brains and for a certain period [were] sufficient for nervous sen-
sibility after the animal has been cut in pieces.’’29
This applied to vertebrates too. Herbert Mayo – protégé-turned-
enemy of Sir Charles Bell, author of several well-regarded textbooks,
and Professor of Anatomy at King’s College London by 1830 – con-
cluded that each segment of the vertebrate spinal cord was comparable
to the ganglion of an invertebrate segment. In 1842 he even noted that
surgically bisected vertebrates could become two ‘‘sentient beings’’ – if
only for an instant. (Invoking the judgment of Solomon, an incredulous
reviewer invited Mayo to try).30 Indeed, one way to determine if a
researcher was an analyst is to see if he openly mused about the pos-
sibility of compound individuality – whether certain lower organisms
were actually aggregates of subordinate agents, such as ganglia-brains.
27
Solly, 1836, pp. 15–16; Grainger, 1839, p. 371; Owen, 1836a, pp. 244–245; Jacyna,
1984, pp. 70–72.
28
Clark, 1835, pp. 102–103; Jones, 1841, p. 692. For Jones’s popularity see Desmond,
1989, p. 274.
29
Owen saw Cuvier as partly founding the group Articulata upon the ‘‘divisibility of
the body, and the power which the fragments possess of retaining a kind of independent
vitality corresponds to the distribution of the nervous system into as many centres as
there are corporeal fragments.’’ Owen, 1836a pp. 244–245. He also noted Virey’s
method, in Owen, Definitions from Museum Lectures on the Animal Kingdom, ND,
OPAP, RCS. On compound individuality see Elwick, 2003, pp. 50–53; Elwick, 2005.
30
Mayo, 1833, pp. 230–231, 220–222; Mayo, 1842, pp. 28–29; Anonymous, 1842, 20,
22.
46 JAMES ELWICK
For the analyst of the 1830s and 1840s, compound individuality was a
valid problem.31
Meanwhile, neurophysiology reinforced compound individuality. In
addition to being materialist, Hall’s 1837 reflex doctrine can also be seen
as analytic:synthetic. Following L.S. Jacyna, I suggest that the reflex arc
was the simplest neurophysiological element, mediated by the simplest
neuroanatomical element of the ganglion: all behavior was made of
compounded reflexes. Ganglia were redefined as centers that mediated
reflex arcs. Demonstrations on invertebrates played an important role in
gaining acceptance for the reflex arc.32 Just as simpler invertebrates were
shown to have more decentralized nervous systems, so too was their
behavior deemed more ‘reflex’ and involuntary.
Conversely, the reflex arc was deployed to explain complex human
mental activities. In 1844, Thomas Laycock proposed that human
consciousness was a series of increasingly compounded reflexes. An
analyst, Laycock was a phrenological sympathizer who denied the
existence of a single perceptual center, a sensorium commune or ‘seat of
the soul.’ Instead mental activity emanated from the interaction of
separate mental faculties. Laycock later replaced phrenological mental
faculties with ganglia, citing the dispersed nervous systems of inverte-
brates as evidence of human nervous structure and function.33
Figure 1 shows Laycock’s place in a network of analytic:synthetic life
researchers. In the early 1830s Laycock took Robert Grant’s compar-
ative anatomy lectures at London University, telling the phrenologist
George Combe that he was Grant’s favorite pupil. His classmates
31
Briefly, I suspect that one reason why beliefs in compound individuality accom-
panied the style of analysis:synthesis was because that style delegated agency to lower
levels of organization. For a researcher to gain clarity and certainty, relationships
between components of a system were treated as properties of those components. That
is, when one analytically disintegrated a system, one ignored that the system’s elements
were members of a larger system with its own peculiar properties arising out of those
relationships. Instead those relationships were depicted either as belonging to each
individual unit, or as irrelevant. When one then synthetically compounded those units
back into a system, that system was reinterpreted as a population of unrelated elements
instead of related members of a whole. For instance, consciousness was seen as the
byproduct of each ganglion; life was seen as the aggregate result of tissue or cellular
activity. For this clarification I am indebted to Eli Gerson. See Albury, 1977, pp. 90–91,
who notes Bichat’s depiction of the entire organism’s life as nothing more than the
contribution of each tissue; on this, Bichat was assessed by Huxley, 1894, pp. 3:366–367.
32
Jacyna, 1982, p. 235. Thus see Carpenter, 1838; Newport, 1843, p. 264; Anony-
mous, 1845, p. 496. Elwick, 2005 notes Newport’s millipede vivisections in more detail.
33
Laycock, 1845; Laycock, 1847; Seccombe, 1892; Leys, 1990, p. 316; See Laycock’s
musings on ‘‘composite animals’’ in Laycock, 1976, pp. 2:243–246, 259–260.
included George Newport and William B. Carpenter. Carpenter also

extended the reflex arc to higher mental operations, coining such terms
as ‘‘afferent’’ and ‘‘efferent’’; in the early 1840s he convinced his then-
patron Owen to use the reflex arc.34 Grant’s other pupils included Peter
Mark Roget (of Thesaurus fame, but who also wrote a Bridgewater
Treatise on physiology), and William Baly (translator of Johannes
Müller’s Elements of Physiology).35
Analysis:Synthesis Part 2: ‘Development’ as Synthesis
Synthesis is the reverse of analysis, integrating separate pieces into a

whole. In an 1842 paper Etienne Augustin Serres declared the com-
parative anatomist to be an analyst, using dissection and maceration to
disintegrate his specimen: ‘‘Association has united and as it were
confounded the elements entering into their composition; disassociation
isolates and separates them anew; art acts in an inverse sense to nat-
ure.’’36 Pre-von Baerian embryologists, then, saw development as syn-
thesis. It was akin to metamorphosis. Analysts might decompose parts
like the nervous system into its elementary units, but nature itself syn-
thesized ganglia together, forming compounded nervous structures.
Like Cuvier, Serres also used the nervous system to classify animals,
in 1824 associating simpler systems with younger animals. Thus mature
molluscs and immature insects (larvae) both had a nervous system with
two separate strands of ganglia and nerve fibers. Parallels could be
drawn between those two animals. As the insect larva continued to
metamorphose, however, its two strands fusing and concentrating
around its esophagus, it could be portrayed as moving ‘past’ the dis-
persed mollusc nervous system. This recapitulationist process – of
nervous systems fusing and concentrating in the head, in both the
individual and in the group – was later called ‘‘cephalisation.’’37
Serres was particularly insistent on the direction in which the embryo
developed. The embryo did not start from a central point and then
ramify outward – instead, the parts of the embryo appeared at a number
34
Carpenter and Carpenter, 1889, pp. 10, 22; Leys, 1990, pp. 310–311, 228; James,
1996, pp. 23–24; W.B. Carpenter to R. Owen, 26 Jun 1842, OCORR, NHM, 6/302–303.
35
Roget, 1834; Müller, 1838–1843.
36
Serres, 1842b, p. 116.
37
Serres, 1824, pp. 379–380. The OED notes ‘‘cephalisation’’ was coined in 1861 by
James D. Dana to describe animals’ tendency to concentrate their nervous systems into
heads as they developed.
48 JAMES ELWICK
of independent points and then fused inward. Organs placed along what
was to become the embryo’s central axis started out double or multiple,
then fused together into a single unit. Single and dispersed nerves ap-
peared first, then fused into a nervous cord. Serres contrasted this
direction of ‘‘centripetal’’ embryological development against what he
saw as the obsolete Hallerian ‘‘centrifugal’’ direction. He proclaimed
that the law of centrifugal development – which in his eyes supported
pre-existence – was refuted, replaced by the law of epigenesis.38 Yet by
epigenesis, Serres did not mean differentiation and specialization. The
dichotomy of preformationism versus epigenesis overlooks this crucial
directional difference.
In Britain, Serres was enthusiastically taken up; his work supported
Cuvier’s belief that the nervous system provided a taxonomic key. Thus
one 1828 reviewer noted that ‘‘The more the volume of the brain exceeds
that of the spinal cord, the higher the animal is placed in the scale of
being.’’ Other reviewers celebrated Serres’s ‘‘‘Centripetal or Eccentric
Theory of Development,’’’ and scolded other British researchers for
assuming that the brains and nervous cords appeared before nerves.
Didn’t they know that development occurred in the opposite order and
direction?39 In the 1834 edition of his well-known Elements of Anatomy
(celebrating Condillac’s method, Elements was explicitly a work of
‘‘Analytical Anatomy’’), the London University Professor of General
Anatomy, Jones Quain, noted that ‘‘the fruit of modern research’’
showed how embryos started out as separate parts, which then fused
together.40
Robert Grant’s 1833 London University comparative anatomy lec-
tures also emphasized the nervous system becoming more complex and
concentrated as it developed. Appearing in the Lancet, his lessons noted
how organs showed the comparative anatomist that the animal king-
dom developed ‘‘from simple to compound.’’41 And he drummed this
message into his students in other ways. Questions on his July 1831
London University exam included ‘‘Where do you find the Nervous
System begin to manifest itself in ascending through the animal king-
dom; and what are the principal forms it assumes in the different classes,
before you arrive at animals possessing a Brain?’’ and ‘‘State the
changes which are observed to take place in the Nervous System of
38
Serres, 1842a, p. 19; Russell, 1982, p. 81; Appel, 1987, pp. 122–123.
39
Smith, 1828a, p. 186; Smith, 1828b, p. 459; Anonymous, 1837, pp. 87–88; Anon-
ymous, 1840a, pp. 234–235.
40
Quain, 1834, pp. v–vi, 16–19.
41
Grant, 1833, pp. 44–45; Grant, 1833–1834, pp. 1:399–400.
Insects during their metamorphosis to the Pupa and the insect state’’.42
Those trained by Grant, including Carpenter, Laycock, Newport, Baly
and Roget, would likely have had to answer these, or similar, questions.
The former question, about the manifestation of the nervous system,
was worth the most marks on the exam. The latter question, about the
appearance of the nervous system in insects, was followed by some
similar 1834 Royal Medal winning researches by former student George
Newport. In fact they were so similar that Grant later accused Newport
of plagiarizing from him and from continental entomologists like Pierre
Lyonet.43 Meanwhile an anonymous review of Joseph Swan’s 1836
textbook on the nervous system charged Swan with neglecting New-
port’s 1834 Royal Medal work, as well as research by Swammerdam,
Lyonet again, Herold, Meckel, Steviranus, Dufour, Straus-Durckheim,
Weber, Audoin and Milne Edwards.44 The specifics of the charges
matter less than the larger point that cephalisation was very widely used.
Regeneration as Reproduction, Reproduction as Regeneration
Focused on the simplest elements of the body, analytical life researchers do

not seem to have strongly distinguished between sexual and asexual
reproduction. Indeed the OED shows that the London botanist John
Lindley was the first to use the term ‘‘asexual’’, and only in 1830. ‘‘Asex-
ual’’ was imported into zoology in 1858, by G.H. Lewes, to refer to how the
simplest marine invertebrates reproduced. Before ‘‘asexual’’ was used in
British zoology, then, researchers often used the word ‘‘vegetative’’
reproduction, denoting not so much an activity as an inherent power.
This ‘‘vegetative’’ power was strongest in lower animals like ‘‘zoo-
phytes’’ (animal–plants), for like plants, they could make copies of
themselves by shoots, cuttings or graftings. Owen was especially inter-
ested in this ‘‘vegetative’’ power. In his annotated copy of his 1843
Lectures on Invertebrates, he bracketed his own passage: ‘‘Wishing to
ascertain if the vegetative power was inexhaustible, Bonnet cut off the
head of one of these worms, and, as soon as the new head was com-
pleted, he repeated the act; after the eighth decapitation the unhappy
subject was released by death, – the execution took effect – the repro-
ductive virtue had been worn out...’’. Owen placed two exclamation
42
Questions from Grant’s exam, Sat, 8 July 1831, from Freeman, 1964, pp. 9–11.
43
The charges in the 1837–1838 Lancet include Newport, 1837–38a; Newport, 1837–
38b; Newport, 1837–38c; Newport, 1837–38d; Grant, 1837–38a; Grant, 1837–38b; Hall,
1837; Hall, 1837–38.
44
Anonymous, 1836, p. 437.
50 JAMES ELWICK
marks next to this section, penciling in, ‘‘Dalyell succeeded in artificially

propagating a Sabella [a marine worm] in the same way’’.45
Plants and simpler animals such as the exemplary Hydra polyp of
Abraham Trembley were thus seen not merely as structurally similar –
both sessile, with branching shapes - but also as reproductively sim-
ilar. They were compoundly individual. In 1833, Lindley stated that a
tree – a ‘‘congeries of vital systems, acting indeed in concert, but to a
great degree independent of each other... [with] myriads of seats of
life’’ – resembled a polyp. In that same year, Grant likened polyps to
trees, and in the following year Roget’s Bridgewater Treatise in
physiology noted how most researchers saw plants and polyps as
aggregations of individuals. Roget cited Erasmus Darwin on this
point, although like Newport he also allegedly plagiarized from
Grant. Scottish researchers Allen Thomson and John Dalyell invoked
the polyp-plant likeness. So too did Owen, who in this claimed to
follow the legendary father of British surgery and natural history,
John Hunter.46 Hence Charles Darwin was in no way unusual when
in 1838 he mused about trees as ‘‘great compound animals united by
wonderful and mysterious manner’’, or when he compared the ‘‘tree
of life’’ to the ‘‘coral of life’’.47
The term ÔreproductionÕ was often used in place of the word
ÔregenerationÕ. The ability of, say, lobsters to regenerate lost limbs was
an instance of vegetative reproduction. George Newport thus entitled
one paper ‘‘On the Reproduction of Lost Parts in Myriapoda and In-
secta.’’ And as President of the Entomological Society in 1844, he
identified this problem as one of high importance.48 The subject’s most
promising young researcher was John Goodsir’s brother Harry, who
45
R. Owen, 1843, Lectures on the Comparative Anatomy and Physiology of the
Invertebrate Animals (Annotated copy), OCORR, NHM, 7, p. 144.
46
Lindley, 1833, p. 32; Roget, 1834, p. 1:89fn; Thomson, 1839, pp. 424–425; Dalyell,
1847, pp. 1:5–7; Owen, 1837, p. 4:xxviii.
47
Darwin, 1987, M41 p. 589. This entry is dated about autumn 1838. See also
footnote 41-1, on Erasmus Darwin’s note on the resemblance between the tree – a
‘‘congeries of living buds,’’ and the coralline (a marine invertebrate) – a ‘‘congeries of a
multitude of animals.’’ For ‘‘coral of life’’, idem, B25–26:177, written some time in
1837–1838. M.J.S. Hodge and Phillip Sloan both show how Darwin the ‘‘generation
theorist’’ linked zoophytes with trees, seeing both as bud-colonies; this allowed him to
reinterpret entities across levels of organization. Species were deemed similar to indi-
viduals; buds and cells were likened to individuals or species. Hodge, 1985, pp. 209–213;
Sloan, 2002.
48
Newport, 1844a, p. 5; Newport, 1844b; Newport, 1845.
investigated precisely where new limbs budded on crustaceans.49

Unfortunately in 1845, Harry Goodsir went to the Arctic on the ill-fated
Franklin Expedition. A naval assistant-surgeon like Huxley, Goodsir
was never heard from again.
Others were also interested in the reproduction of limbs. Richard
Owen (in addition to his interest in Bonnet’s and Dalyell’s work) cor-
responded with Welsh researcher John Blackwall, who cut off spider
limbs to study their regeneration.50 Owen’s proposal of dueling mor-
phological forces can be situated in these local research problems, in
addition to any naturphilosophie. From above Owen proposed the
workings of the teleological ‘‘adaptive force’’; from below came ‘‘vege-
tative repetition,’’ which caused phenomena like serial homology.
Nicolaas Rupke thinks that Owen chose the term ‘‘vegetative repetition’’
because repeating body structures (segments, vertebrae) hinted at the
same structure as a plant, such as plant leaves.51 I suggest we go further:
Owen’s ‘‘vegetative repetition’’ in animals was caused by the same kind
of force at work in plants.52 More serial homology meant that lower
animals were more plant-like.
Owen held that lower animals and plants exhibited greater vegetative
repetition. This repetition meant that lower animals were also far more
compoundly individual. In On Parthenogenesis, he noted that polyps
were the individual digestive organs of a ‘‘compound organism’’, like
plant leaves. In 1848, writing to his Oxford friend, the Reverend Daniel
Conybeare, Owen used a tapeworm to illustrate vegetative repetition: it
could have hundreds of segments, of which no individual segment was
so important to the entire worm that its removal would harm the entire
animal. A segment might even survive on its own. But if the adaptive
force grew stronger, each segment lost its independence, becoming part
of a larger group-individual; its removal would harm or kill the rest of
the tapeworm’s segments because of their new mutual dependence.53
49
Goodsir, 1844a; Goodsir, 1844b; also Harry Goodsir, ‘‘The mode of reproduction
of lost parts in the crustacea,’’ pp. 74–78, in Goodsir and Goodsir, 1845. Goodsir was
also Conservator of the Museum of the Royal College of Surgeons in Edinburgh.
50
Blackwall, 1848. On their correspondence see Gruber and Thackray, 1992, pp. 36–
37.
51
Rupke, 1994, p. 197.
52
Whewell saw science emerging from the interplay of ideas and things, and Phillip
Sloan thinks Owen was inspired by Whewell’s ‘‘antithetical’’ methodology, particularly
in his proposal of two duelling morphological forces. This is if we see Whewell as
‘‘pontificating’’ on how science ought to be carried out. Fisch, 1991; Sloan, 2003.
53
Owen, 1849, p. 56; R. Owen to W.D. Conybeare (Draft), 13 Mar 1848, OPAP,
RCS.
52 JAMES ELWICK
Thus Owen’s adaptive force was not simply teleological. It was also
integrating, which for him solved the problem of compound individu-
ality. This also explains his proposal of parthenogenesis, where less
complex organisms had more vegetative reproductive power, and thus a
greater regenerative ability. Frederick Churchill has explained Owen’s
proposal of parthenogenesis as emphasizing sexual reproduction. But
sex should not be overemphasized – for Owen replaced the word
‘‘parthenogenesis’’ with ‘‘metagenesis’’ by 1851. Owen favored this new
term, for it better conveyed a process of metamorphosis spanning dif-
ferent individuals, a kind of ‘metamorphosis-plus.’54
W.B. Carpenter and the Growth of Palaetiology
Famously, Owen’s work on parthenogenesis was later savaged by T.H.

Huxley as overly mystical and mired in confusion.55 Yet at the time
analysts welcomed Owen’s proposal. Edward Forbes especially enjoyed
Owen’s lectures on parthenogenesis, for he too had speculated about
compound individuality and the polyp – plant resemblance.56
One key dissenter from Owenian doctrine was William B. Carpenter.
In 1846 he was an analyst, discussing cephalization in a landmark
paper.57 But by 1848 he had switched to palaetiology. This mid-career
switch was partly because of a social acquaintance with fellow Bristolian
James Cowles Prichard, to whom Carpenter looked up ‘‘with an almost
filial reverence and gratitude.’’ Prichard was an important palaetiolo-
gist: Whewell himself noted that Charles Lyell cited Prichard’s 1813
ethnological Researches into the Physical History of Man more fre-
quently than any geology book. In 1848, Carpenter penned an anony-
mous and glowing Edinburgh Review piece about Prichard’s Researches.
Carpenter described how Prichard’s ethnology was similar to philology:
both sciences were genealogical, showing that both languages and races
fanned out from common origins and a geographic center. Carpenter
even cited tree-like images of descent, quoting one philologist who
thought that we would learn more about languages by learning about
54
Churchill, 1979, p. 150; Owen, 1851. In October 1851 Owen was congratulated by
Julius Victor Carus for this name change, because metagenesis ‘‘wonderfully’’ expressed
the analogous German word, ‘‘Generationweschel.’’ J.V. Carus to R. Owen, 1 Oct 1851,
OCORR, NHM, 6/365–366. See also Steenstrup, 1845.
55
Huxley, 1858, pp. 537–538.
56
E. Forbes to R. Owen, [1848], OCORR, NHM, 12/320–323; Forbes, 1844.
57
[Carpenter], 1846, pp. 503–504.
‘‘...the roots, and by the application of the principle of secondary for-

mation, overgrowing, sometimes luxuriantly, the ancient stock of
roots.’’58 From this review onward, Carpenter applied the image of life
forms and other phenomena spreading out from a central origin-point.
Meanwhile Carpenter grew bolder at opposing Owen because of a
simmering priority dispute. Who had first used von Baerian embryolog-
ical principles in Britain, after its introduction by Martin Barry in 1837? In
1840, Carpenter was credited by a reviewer for doing this. But in the late
1840s and early 1850s Owen pressured Carpenter to give way; Owen
claimed to have first used it in 1843.59 This alienated Carpenter, desperate
to be seen as an ‘‘independent discoverer’’, not just a textbook-compiler.60
Carpenter used palaetiology to emphasize a new distinction between
what we now call sexual and asexual reproduction. In 1848 and 184961
he anonymously portrayed Owen’s parthenogenetic musings as con-
fused. Owen had claimed that parthenogenesis explained the mystery of
the ‘‘alternation of generations’’: why successive ‘‘generations’’ of dif-
ferent morphological types were produced in the same species. For in-
stance an aphid of type A produced a series of aphids of type B, which
eventually gave rise to type A again. But Carpenter claimed that the
word ‘‘generation’’ shouldn’t refer to a morphological form but to an
activity. Here, I build on John Farley’s interpretation.62
Carpenter used Martin Barry’s 1837 statement that British research-
ers were looking at the tree of life from the wrong direction – that since
58
Whewell, 1967a, pp. 3:397–399; [Carpenter], 1848a,b, pp. 432–433, 473–474, 477. In
the point on American languages, Carpenter quotes Chevalier Bunsen’s 1848 BAAS
report. On ‘‘Filial reverence’’ see Carpenter and Carpenter, 1889, p. 26.
59
This raises the question: since Owen used von Baerian principles, why can’t he be
considered a palaetiologist? Answer: first, partly because he used von Baerian principles
for different purposes – to sort out the resemblance or identity of static body parts.
Owen used von Baer to distinguish between the homology (identity) and mere analogy
(functional resemblance) of two body parts – homologies could exist only between
adults of the same embranchement. Richards, 1977, p. 218; Panchen, 1994, p. 50; Rupke,
1994, pp. 153, 170. Second: partly because he used von Baerian principles incorrectly.
Evelleen Richards also maintains that Owen misunderstood von Baer until he had read
Huxley’s 1853 translation of Entwicklungsgeschichte, a point raised by Huxley himself.
Richards, 1987, pp. 142–143. For Carpenter being credited with first use of von Baer,
see Anonymous, 1840b, pp. 112–113; on the priority dispute see W.B. Carpenter to R.
Owen, 20 Oct 1851 OCORR, NHM 6/333–334, and 2 Aug 1853 OCORR, NHM 6/335–
336.
60
For Carpenter’s need to be seen as an ‘‘independent discoverer’’ see W.B. Carpenter
to R. Owen, 2 Aug 1853 OCORR, NHM 6/335–336.
61
[Carpenter], 1848b; [Carpenter], 1849.
62
Farley, 1982, pp. 80–81.
54 JAMES ELWICK
development was now known to occur in a centrifugal direction from a

single point, what now counted was the origins and beginnings of ani-
mals – in the case of zoophytes, precisely how and where they repro-
duced.63 In the analytic:synthetic view, plants and lower invertebrates
were disintegrated into nominally independent components, with agency
delegated to each part. This made the question of compound individu-
ality viable. But Carpenter used palaetiology to redefine the biological
individual as the entire product of a sexually fertilized ovum – a new
historicist definition of individuality. Though admitting that cells and
body parts often possessed ‘‘independent vitality’’, he stated that this
independence should no longer be the criterion for biological individu-
ality. The agency of parts was no longer important.
More subtly, Carpenter rejected any ‘‘vegetative’’ reproductive
power, deeming this simplistic. He instead distinguished between two
modes of plant reproduction – ‘‘gemmiparous’’ reproduction (the
budding of new cells) and ‘‘oviparous’’ reproduction (occurring only in
germs produced by special organs, such as flowers). Oviparous repro-
duction happened only after two sexual organs interacted. In turn, two
kinds of reproduction meant two kinds of reproductive parts. Gem-
miparous generation took place in buds; oviparous generation happened
in ova. Frederick Churchill sees this as a fundamental distinction, yet
while this is now obvious to us, at least two of Carpenter’s contempo-
raries had trouble with his division. Carpenter claimed that the only way
to distinguish between the two parts was how they reproduced. While
the bud had its own spontaneous power of producing new tissues, an
ovum was matter ‘‘not yet organized’’ and could develop only after
impregnation.64 But if sex could not be observed, then how could any
distinction be made between the two parts?
A more charitable way to look at Carpenter’s point is to see it as
shaped by how evidence was used. Carpenter’s distinction between the
two kinds of germs and their modes of reproduction could only be
appreciated by other researchers who watched those germs change over
time. His point could only be appreciated by those who were able to see
the same kinds of processes at work. In turn this emphasizes the
importance of how evidence was kept, and in turn the places where that
evidence was kept. This brings us to palaetiological sites – ‘‘vivaria,’’
controlled environments that housed living specimens.
63
[Carpenter], 1848b, pp. 188–189. He paraphrased from Barry, 1837b, pp. 362–364.
64
[Carpenter], 1848b, pp. 188–189, 192–194, 204–205; Churchill, 1979, pp. 150–
151.
To bolster his novel claims, Carpenter extensively cited the work

carried out by Scottish baronet Sir John Dalyell.65 Dalyell was one of
the first British researchers to extensively use glass vessels for the long-
term observation of the marine creatures he collected from the seashore,
for he believed that hasty scrutiny caused the most mistakes in natural
history. Field work at tidal pools and on dredging expeditions was time
consuming, and offered only snapshot glimpses of these creatures. But
used properly, Dalyell’s glass vessels allowed observations to be made
over months or even years. To use Matthew Goodrum’s evocative
image, these glass vessels allowed tidal pools to be uprooted and
brought indoors. The entire marine organism life cycle could now be
observed. Specimens could be separated from one another too. Hence
John Lubbock later used ‘‘vivaria’’ to keep male Daphnia (water fleas)
separate from female ones to determine whether sexual reproduction
had in fact occurred. Without such isolation one could not distinguish
between sexual and asexual reproduction in these animals.66
To be sure, places such as the Muséum d’Histoire Naturelle had
attached zoos and gardens. How, then, did museums differ from
vivaria? Museums were places where unchanging, not alive, organisms
or organs were disintegrated into simpler and smaller pieces. In this
light, places like the Muséum’s zoological gardens provided spectacles
for the public and specimens for analysts, such as comparative anato-
mists. One key distinction can be found in the uses to which those
gardens were put. In one British case – the Zoological Society – Sir
Stamford Raffles provided money both for a park and a museum. But
the park was to be used to restock aristocrats’ hunting and fishing
grounds just as much as it was to help British natural history. In the late
1820s and early 1830s, it was the Society’s museum that was the center
for London scientific reformers.67 Another key distinction between a
museum’s gardens and a vivarium is the degree of control exerted over a
specimen’s environment: my case of hard-to-study marine invertebrates
better highlights the differences. Vivaria not only facilitated the close
observation of living specimens by keeping them alive and isolated; they
allowed researchers to manage those specimens’ surroundings (tem-
perature; salinity; nutrients; brightness; living space).
65
Thus [Carpenter], 1848b, pp. 196–198.
66
Lubbock, 1857, pp. 79–81; Goodrum, 1997, pp. 278–283.
67
Desmond, 1985, pp. 232–233. See Booth, 1839, pp. 57–58 for the initial purpose of
the Zoological Society – the domestication (and consumption) of non-native animals.
56 JAMES ELWICK
In the case of marine animals, vivaria were usually built out of

expensive glass. David Allen notes how Dalyell could afford to have
‘‘capacious glass vessels’’ made despite high taxes on glass (making it
four times as expensive as its production cost). Dalyell also had to have
seawater carted to his Edinburgh residence each morning, another costly
undertaking. Yet because of his vivaria, he could observe the Prome-
thean recoveries of his hydroid specimens: in 550 days he cut 22 hydroids
that regenerated from the single stem of a branched marine invertebrate.
In a single evening Dalyell watched a sea anemone produce 230 young.68
Glass vessels also meant that specimens could survive in the heart
of a city. In addition to aquaria (then called an ‘‘aqua-vivarium’’) the
best-known vivaria were Wardian cases, first built in 1829 by
Nathaniel Ward; these were sealed environments allowing plants to be
grown even in polluted city centers. After 1829, Ward found that
animals such as goldfish and even a robin could live inside them for
long periods. Wardian cases were made famous at the 1851 Great
Exhibition. On this list one can include other kinds of vivaria, such as
artificial incubation machines: one 1839 London guidebook advertises
the ‘‘Eicallobion’’ located at 121 Pall-Mall, where observers could
break sequences of eggs, examining birds in their ‘‘‘nascent or partly
formed state’’’ for a shilling a break.69
The more that creatures’ entire life cycles could be observed, the more
that palaetiology was strengthened. But even in the late 1840s, when
Carpenter made his novel points, there were fewer vivaria than muse-
ums. This implies that researchers were less familiar with palaetiological
evidence. Instead it would have been far easier to work with pickled
marine invertebrates fished out of spirit-filled glass jars. Perhaps this is
why Carpenter was quickly rejected by contemporaries. To someone
looking at the reproductive parts of dead specimens preserved in alcohol,
it would be difficult to grasp Carpenter’s notion of ‘‘generation’’ as an
activity; his distinction between ‘‘buds’’ and ‘‘ova’’; or his view of bio-
logical individuality as the sum total of the contents of a sexually fer-
tilized ovum. He was criticized by the Scottish physiologist Allen
Thomson and by Edward Forbes, both of whom were unable to
68
This particular sea anemone, called ‘‘Grannie’’ for its fecundity, lived to be 66 years
old, surviving Dalyell by 36 years. It was obituarized in The Times and The Scotsman.
Allen, 1976, p. 132; Dalyell, 1847, pp. 1:36–37; Dalyell, 2004, p. 8. John Reid, also of
Edinburgh, kept marine invertebrates in vivaria too, maintaining one jellyfish colony for
at least 17 months in the 1840s. Reid, 1848, pp. 25–26, 33–34.
69
Booth, 1839; Anonymous, 1851, pp. 1: 466–467.
acknowledge Carpenter’s evidence and his new definition of an indi-

vidual.70 They saw his proposal was bizarre or unnecessary – the defi-
nition of biological individuality as independence worked well enough
for them.
In 1848 and 1849 Carpenter failed to get much support. But over the
next seven years palaetiology came to challenge analysis:synthesis as a
useful style for life researchers. During this time vivaria began to spread
throughout Britain – made cheaper by the removal of the glass tax in
1845, publicized at the Great Exhibition, and exemplified in the opening
of the 1853 ‘‘Fish House’’ marine aquarium at the Zoological Gardens.
In that same year, Philip Henry Gosse told the readers of his Natural-
ist’s Rambles on the Devonshire Coast that aquatic plants helped keep
animals alive inside the vivarium. This ushered in a fashion for aquaria
in the middle-class British home. Specialized aquarium stores even
began to open in London, revealing a market large enough for com-
mercial viability. By 1855 information about aquaria was codified in
textbooks and other popular works.71 They had become familiar.
Meanwhile palaetiology strengthened in Victorian science and cul-
ture. Other historians have noted this. Theodore Merz’s magisterial
history of the sciences devotes a chapter to the ‘‘genetic view of nature’’,
seeing geological uniformitarianism, Lamarckian species modification,
Vestiges of the Natural History of Creation (1844) and the Origin of
Species (1859) as answers to similar historicist questions. William
Coleman links changes in biology with the search for the common
source of modern languages, and Stephen Alter connects comparative
philology and Darwinian descent with modification with Whewell’s
palaetiology. All proposed ramifying pathways from common ances-
tors.72 Eleven years before Darwin, Carpenter – stimulated by ethnol-
ogy, philology and von Baerian embryology – had already begun to
apply these centrifugal principles to biology.
T.H. Huxley
By the time T.H. Huxley had returned to Britain in December 1850

from his tour on HMS Rattlesnake, the palaetiological style was
70
Forbes, 1848, pp. 87–88; Thomson, 1852–1856, pp. 39, 35–36. Part 2 appeared in
1854.
71
Allen, 1976, pp. 135–137; Goodrum, 1997, pp. 252–256, 278–283.
72
Merz, 1965, pp. 2:363–366, 2:279–280; Coleman, 1977, pp. 10–11; Alter, 1999, pp.
2, 13–14.
58 JAMES ELWICK
strengthening. Palaetiology served Huxley well in his research. On

Rattlesnake, Huxley had ample opportunity to view living and devel-
oping marine invertebrates, and he established himself as an elite
researcher by coming up with a new way to classify some of the mem-
bers of the ‘‘lumber-room’’ of Cuvier’s Radiata: by examining their
embryos, investigating the foundational tissues as they turned into more
specialized and differentiated organs. Before Rattlesnake, while being
trained by the embryologist Thomas Wharton Jones at Charing Cross
medical school, Huxley had already copied out quotes from the German
botanist Matthias Schleiden: zoology could not become a true science
until it was informed by development. And like Carpenter, Huxley also
saw the developmental process of animals resembling the centrifugal
emergence of modern languages from earlier ones.73
On returning to London, Huxley accentuated his palaetiological
leanings. He imported the most up-to-date German research, bringing
such works as Albert Kölliker’s Manual of Human Histology to an
English-reading audience.74 And he revived older work too: Huxley
translated large parts of Karl Ernst von Baer’s writings on the grounds
that von Baer had not only been ignored, he had also been misinter-
preted in Britain. All this occurred while Carpenter was slowly giving
way to Owen in their dispute over who had first used von Baer’s law.
Owen’s pressure became too great for Carpenter – giving up all claim to
priority, Carpenter sadly concluded to Owen that he would mostly be
tied up with his educational duties at the University of London, and so
‘‘...I must be content to see younger men taking the place that I had
hoped to occupy as a discoverer, and satisfy myself with endeavouring to
qualify them for a philosophical appreciation of what they may have the
good fortune to find out.’’75 Huxley was one of these younger men, and
conveniently enough one who had publicly recognized in 1853 that
Carpenter was the only English physiologist to have correctly used von
Baer’s principles. By 1855 Huxley again publicly praised Carpenter as
the only Briton or Francophone (save Martin Barry) who properly
73
Huxley, Account of Researches into the Anatomy of the Hydrostatic Acalephae,
1851, HP, IC 37.12–42, pp. 37.32–33; Huxley, Considerations upon the Meaning of the
Terms Analogy and Affinity, [1846–1847], HP, IC 37.1–21, pp. 37.11, 37.13–15, 37.20;
Winsor, 1976, p. 61; Desmond, 1997, p. 25.
74
Kölliker, 1853.
75
W.B. Carpenter to R. Owen, 11 Feb 1854 OCORR, NHM 6/337–338. Emphasis in
original.
understood and appreciated von Baer’s laws: principles that were ‘‘to
Biology what Kepler’s great generalizations were to Astronomy.’’ By
implication, Owen had not accurately used these principles.76
As von Baer’s law became one of the most important principles in life
research, analytic:synthetic questions – about the quasi-independence of
each body part, or how development proceeded by the fusion of those
parts – in turn gradually lost importance. Development was increasingly
seen as proceeding centrifugally, in a ramifying process from unspe-
cialized ‘homogeneity’ to specialized ‘heterogeneity.’77
Huxley followed Carpenter by also defining an individual histori-
cally, as the entire product of a sexually fertilized ovum, no matter how
many independent parts emerged from this process. He was more suc-
cessful than Carpenter at convincing others about this. From the
botanical term ‘‘phytoid’’ Huxley coined the term ‘‘zoöid’’ in about
1850 to avoid referring to seemingly independent body parts as ‘indi-
viduals’. At Huxley’s very first public presentation in 1852 he used the
word ‘‘zoöid’’ to overturn a seeming paradox: zoöids only simulated
biological individuals. Privately he criticized Owen, in what was to
become a series of escalating attacks. Others who found Huxley’s new
perspective useful – including George Busk and George Allman –
committed to the term zoöid, and in the process their negative opinions
about Owen crystallized too.78 Palaetiology also served Huxley’s social
aims. Huxley’s dire career prospects in the early 1850s – sharply at odds
with his own strong faith in himself – only accentuated his perceived
differences from most of the leading practitioners of British life research.
He seized on the growing split between Carpenter and Owen, justified
the break with Owen by emphasizing Owen’s ‘unpleasant’ personality,
and then engineered a schism.79
76
von Baer, 1853, p. 176fn; [Huxley], 1855, pp. 242.
77
Ospovat, 1976 – the classic account of the British reception of von Baerian
embryology – emphasizes divergence through the influence of Louis Agassiz’s paleon-
tology. Yet by focusing on divergence, Ospovat might overlook cephalization (hence
linear and hierarchical arrangement) in classification, and thus Cuvier’s legacy.
78
Huxley, 5 September 1850, Notebook, 23 August 1850 – 4 August 1851, HP, IC,
63.8; Huxley, On Animal Individuality, 30 April 1852, HP, IC, 38.2–38.52, p. 38.5; W.B.
Carpenter to T.H. Huxley 16 Jul 1855, HP, IC 12.78–79; Allman, 1853, p. 379.
79
On Huxley’s self-estimation see T.H. Huxley to H. Heathorn, 28 Aug 1852, T.H.
Huxley – H. Heathorn Correspondence, Imperial College London, Huxley Archives,
HH 221. It is claimed that later apologists for the scientific naturalists retrospectively
created Owen’s reputation for ‘sneakiness’, a matter needing historical explanation – see
White, 2003, pp. 64–65.
60 JAMES ELWICK
Huxley was an extraordinarily hard worker, a skilled researcher and

writer, and a tactical wizard. He even deployed new beliefs about how a
man of science should behave, nicely shown by Paul White.80 But self-
fashioning alone was not quite enough for Huxley to rise, and it does
not quite explain his ability – stunning in retrospect – to depict Owen’s
research as foolish and confused. Huxley not only succeeded in con-
veying to history a picture of Owen as a ‘sneaky’ scientist. By redefining
certain problems as unimportant, Huxley also erased the very real
analytical commitments of the 1830s and 1840s that animated a larger
community of British life researchers. When Darwin’s palaetiological
theory of natural selection arrived, this clouded earlier life research aims
even more, ‘‘like the secretion of a cuttle fish’’.81
For one illustration of Huxley’s successful erasure, consider the
infamous dispute between Huxley and Owen over the structure of the
human brain. It began when in 1858 Owen told the Linnean Society that
mammals should be grouped by brain structure. He divided mammals
into four groups: the lowest was the Lyencephala, ‘‘loose brained’’
because of the ‘‘disconnected state’’ of its cerebral hemispheres. Next
was Lissencephala, ‘‘smooth brained’’ because there were few convolu-
tions (although the hemispheres were connected). Then came Gyren-
cephala – ‘‘winding brained’’ to denote increased convolutions; and
finally Archencephala – ‘‘(over)ruling brain[ed]’’, where the cerebral
hemispheres covered the olfactory lobes and cerebellum. Humans
belonged to this final group, and in two sentences of a 37 page paper,
Owen offhandedly noted that humans had a special posterior lobe that
he called the ‘‘hippocampus minor.’’82 Nicolaas Rupke sees Owen’s
proposal as a novel form of classification.83 Yet Owen’s 1858 scheme
followed his earlier cephalisation-groupings (recall Homogangliata and
80
White grounds Huxley’s rejection of Owen in new views about how researchers
should conduct themselves. In Owen’s perspective – normal in the unequal 1851 world
of gentlemanly science – patrons helped clients in return for deference. But Huxley
rejected this, hoping to speak the truth regardless of social network. White, 2003,
pp. 37–38, 45.
81
This was Samuel Butler’s image. Butler, 1923, p. 292; Lightman, 2006, forthcoming.
82
Owen, 1858, pp. 14, 17–18, 19–20. The exact quote: ‘‘[Archencephala’s] posterior
development is so marked, that anatomists have assigned to that part the character of a
third lobe; it is peculiar to the genus Homo, and equally peculiar is the ‘posterior horn of
the lateral ventricle,’ and the ‘hippocampus minor,’ which characterize the hind lobe of
each hemisphere. The superficial grey matter of the cerebrum, through the number and
depth of the convolutions, attains its maximum of extent in Man.’’
83
Rupke, 1994, p. 266.
Heterogangliata) – it revived his earlier Cuvierian-inspired project to

classify by nervous structure.84
But in a fight that would become almost as famous as his disputes
with Bishop Wilberforce or his later skirmishes with Herbert Spencer,
Huxley seized on Owen’s minor remarks about the hippocampus
minor. He famously proclaimed that no such posterior lobe existed;
hence there was no dividing line between humans and the rest of the
animal kingdom, affirming the new Darwinian view. Historians tend to
follow Huxley’s reasoning, focusing on the presence or absence of the
hippocampus minor, and so we ignore the rationale behind Owen’s
scheme. By default we place Owen in a Darwinian framework, locking
him into a fight over links between humans and animals. We might
avoid this by seeing his work – and that of other pre-Darwinian life
researchers – as informed by analysis and synthesis.
Conclusion – Possible Worlds
Was it inevitable that palaetiology pushed out analysis:synthesis in

key domains of life research? Probably not – analysis:synthesis might
have remained strong in Britain if three key people had not died
early. Glance again at Figure 1 and speculate on a world that might
have been. The promising young Harry Goodsir might not have
disappeared on a failed exploration. E.S. Forbes – who would alone
have considerably restrained the young fanatic Huxley’s attacks on
Owen – might not have succumbed to kidney failure. And the
industrious and well-regarded George Newport might not have died
from a fever caught while capturing frogs in a London marsh.
Though things largely turned out one way – mainly Huxley’s – other
arrangements and research directions were possible. We might instead
imagine a world in which it is Harry Goodsir that delays his departure
by a year, becoming assistant-surgeon on HMS Rattlesnake and trav-
eling to Australasia. In that same world, Huxley advances his itinerary
by a year, taking Goodsir’s post on the Franklin Expedition only to
perish on a desolate Arctic island. Picture the very different kind of life
research that might have appeared as a result – perhaps one in which
palaetiology was not brought so forcefully to Britain. Owen’s program
of parthenogenesis and metagenesis might have been continued by
84
Indeed, Owen cited his own 1842 Hunterian Lectures on the Nervous System –
Owen, 1858, pp. 13–14. For a similar view see [Carpenter], 1846, pp. 503–504.
62 JAMES ELWICK
Harry Goodsir’s regeneration investigations, and Owen might have

been celebrated, not maligned.
It is also important to note that people did indeed change their views.
They were not the creatures of styles of reasoning. This is shown on
figure 1 – by shifting their commitments, Carpenter, Allman, and
Darwin made fruitful new discoveries and raised new questions. But
instead of seeing these changes as the free and rational choices made by
individuals who heroically struggled against the constraints of styles of
reasoning, it is more subtle to see their situations as intercontingent.85
Each researcher was not simply affected by his own choices – he was
also influenced by all of the choices and activities of every other life
researcher. They were susceptible to others’ commitments; they con-
tinuously monitored one another; they defined their work against oth-
ers’ research; they compared their self-presentation with others. Thus
even implacable enemies – Huxley, Owen – were bound up together. As
members of changing networks, they were less autonomous and far
more mutually dependent than they realized.
Acknowledgments
For Polly Winsor. I thank Katherine Anderson, John Beatty, Elihu

Gerson, Michael Ghiselin, Ian Hacking, Bernard Lightman, Gordon
McOuat, Lynn Nyhart, John Pickstone, and the anonymous referees of
the Journal of the History of Biology. I gratefully acknowledge the
permission of the President and Council of the Royal College of Sur-
geons of England to quote from the Richard Owen Papers, and Tina
Craig’s assistance there; the permission of Imperial College London’s
Library Archives and Special Collections to quote from the Thomas
Henry Huxley Papers and Manuscripts, and the assistance of Anne
Barrett and Hilary McEwan there; and the permission of the Trustees of
the Natural History Museum (London), to quote from the Richard
Owen Correspondence and Collection, and Paul Cooper’s assistance
there. This research and writing was supported by a Social Sciences and
Humanities Research Council of Canada Postdoctoral Fellowship, and a
grant from the Joint Initiative in German and European Studies, Univer-
sity of Toronto.
85
On ‘intercontingency’ see the lively Becker, 1998, pp. 34–35, who is following up on
Norbert Elias’s suggestion that we replace the usual antithesis of freedom versus
determinism: Elias, 1978, p. 167.
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Journal of the History of Biology (2007) 40:35–69 Ó Springer 2006

Styles of Reasoning in Early to Mid-Victorian Life Research:

Abstract. To better understand the work of pre-Darwinian British life researchers in

Keywords: analysis, Charles Darwin, George Newport, museums, neurosciences,

This paper introduces a new dichotomy for historians of nineteenth

sis:synthesis. The paper then moves on to consider palaetiology in more

A Comparison of Analysis:Synthesis and Palaetiology

A quick way to contrast the two styles is to appropriate two diﬀerent

Meanwhile the term ‘palaetiology’ was coined by philosopher Wil-

by moving inward – making ontogeny the fusion, or synthesis, of

Institutions/sites Museums ‘Vivaria’ (aquaria, Wardian cases)

Analysis:Synthesis Part 1: Analysis

Let us begin by dividing the style of analysis:synthesis into its own

R.E.Grant J. Quain W.S.Macleay 1819 J.Dalyell 1814

Figure 1. Some Life Researcher Commitments, 1830–1850.

The Analysis of the Nervous System

‘‘neurine’’ – a tissue providing nervous power. Though there were dis-

included George Newport and William B. Carpenter. Carpenter also

Analysis:Synthesis Part 2: ‘Development’ as Synthesis

Synthesis is the reverse of analysis, integrating separate pieces into a

Regeneration as Reproduction, Reproduction as Regeneration

Focused on the simplest elements of the body, analytical life researchers do

marks next to this section, penciling in, ‘‘Dalyell succeeded in artiﬁcially

investigated precisely where new limbs budded on crustaceans.49

W.B. Carpenter and the Growth of Palaetiology

Famously, Owen’s work on parthenogenesis was later savaged by T.H.

‘‘...the roots, and by the application of the principle of secondary for-

development was now known to occur in a centrifugal direction from a

To bolster his novel claims, Carpenter extensively cited the work

In the case of marine animals, vivaria were usually built out of

acknowledge Carpenter’s evidence and his new deﬁnition of an indi-

By the time T.H. Huxley had returned to Britain in December 1850

strengthening. Palaetiology served Huxley well in his research. On

Huxley was an extraordinarily hard worker, a skilled researcher and

Heterogangliata) – it revived his earlier Cuvierian-inspired project to

Conclusion – Possible Worlds

Was it inevitable that palaetiology pushed out analysis:synthesis in

Harry Goodsir’s regeneration investigations, and Owen might have

For Polly Winsor. I thank Katherine Anderson, John Beatty, Elihu

Manuscript sources and abbreviations

The Cyclopaedia of Anatomy and Physiology, 1836–1856. 5 Volumes. Edited by R. B.

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