History of Developmental Introductory article

Biology . Introduction
Article Contents

Tim Horder, University of Oxford, Oxford, UK . Glimpses of Embryology in Pre-scientific Times

. Early Scientific Period

. The Era of Molecular Biology and Genetic Engineering:

1985–

. Final Observations

Online posting date: 15th November 2010

Developmental biology – or ‘embryology’ – is a subject subjects. This is even true of evolution theory; despite its
with a long and distinguished, but uneven, history. It is complexity and temporary reversals, a central core of ideas
remarkable for the list of leading scientists and phil- runs through and unifies the history of the subject. But the
osophers who have contributed to it from antiquity up to ‘history of embryology’ probably does not conjure up any
the present era. Because embryos are typically minute and
overall ‘story of progress’; despite accumulation of much
data, it is notable for lacking any obvious sustained,
inaccessible direct study and understanding has been
coherent structure over time. The interesting question is
slow and relatively recent: before the mid-nineteenth whether this is a reflection of a subject that has simply not
century theories of reproduction and the origin of living yet reached a sufficient state of scientific maturity, or
forms were largely speculative. Since then embryology whether there is something distinctive about this particular
has varied greatly in its importance within mainline bio- discipline which makes any ideal of coherence an illusion.
logical thinking. At one time it provided key evidence for Nonetheless, the history of embryology is also remark-
evolution; later for early theories of genetics. Recently, able for the long list of distinguished scientists who have
and even now it can be argued, biologists are divided and contributed to it in one way or another. Indeed, most of the
unclear about how best to demarcate the subject because greatest philosophical and scientific thinkers up to recent
it overlaps with many other biological themes as diverse times came to consider issues that we would now describe
as ‘developmental’. And yet the subject lacks obvious
as reproduction, malformation, aging and cancer. Today
heroes, comparable to Darwin or Watson and Crick. The
it tends to be approached through the techniques
subject is notable in one other respect: its relative neglect by
of molecular biology and molecular genetics, and has modern historians of science. Interest in the whole subject
acquired renewed interest in the context of fertility area has undergone major ups and downs. Recently
enhancement, stem cells and genetic engineering. (through the arrival of molecular techniques) there has
been yet another resurgence in its popularity as a research
field, but around 1960 it was hard to feel any enthusiasm; it
was widely regarded as an old-fashioned subject without
any discernible future. In the current era, some people
claim that molecular genetics has at last provided the long
Introduction hoped-for definitive account of embryogenesis. But if
during its long history hitherto the subject has failed to
When compared to the history of most biological discip- achieve a stable consensus in terms of a core ‘problem’ or
lines, the history of embryology – or developmental biol- ‘theory’ structure, then one certainly cannot easily assume
ogy as it is now called – seems episodic if not frankly that matters have now suddenly changed.
nebulous. By comparison, the histories of heredity (gen- By the end of this survey I hope to offer suggestions that
etics), biochemistry or molecular biology, for example, go some way to explaining these features. I take the view
trace out a pattern of more or less steady, linear forward that the nebulous or episodic nature of the development
movement over many past decades, with an eventual of the subject of embryology is not due to any intrinsic
stabilising of widely agreed, core principles defining the marginality as a discipline within biology or to its lack
of definable problem issues. Indeed embryology is self-
evidently a distinct discipline that everybody can nowadays
ELS subject area: Science and Society recognise; it is essentially and easily definable as what
happens between egg and adult. Paradoxically, it can be
How to cite: argued, its apparent historical failure to mature may stem
Horder, Tim (November 2010) History of Developmental Biology. In: from the fact that it lies at the very heart of biology.
Encyclopedia of Life Sciences (ELS). John Wiley & Sons, Ltd: Chichester.
Logically speaking it is central within biology because it is
DOI: 10.1002/9780470015902.a0003080.pub2
where evolution, genetics, ecology and morphology (i.e.

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 History of Developmental Biology

the structure of complete, adapted adult organisms) all known about reproduction. What is remarkable is how
meet. Its very complexity and centrality has perhaps made much insight was achieved in the circumstances. See also:
it hard to attack head on; yet, being central, it has repeat- Needham, Joseph; Philosophy of the Life Sciences
edly provided starting off points for other more coherent, Babylonian and Egyptian concerns with congenital
focused and linearly developing subjects. It has perhaps abnormalities and ‘freaks’ are documented. We know that
been the victim of fashion, technique and shifting views of the Greeks took a serious interest in the subject. Hippocrates
what is important in biology generally. Whatever the rea- studied chicken embryos. Clearly informed by much direct
son, the end result throughout the past century was the observation, Aristotle classified animals according to whe-
general impression of marginalisation and especially of ther they reproduced by means of eggs, spontaneous gen-
fragmentation (or even of being ‘taken over’ by other eration (i.e. in smaller species and insects) or viviparity. He
more clearly defined subject areas). All these features can described placental fish (a finding that had to be redis-
immediately be appreciated if one looks at any of today’s covered by Johannes Müller 2100 years later). Yet Galen,
texts on ‘developmental biology’; they present a rich but despite his wide observations, could still disagree with
incoherent, even random, mix of diverse themes covering Aristotle over whether the embryo came from the white or
much of modern biology. So the territory included in this the yolk of the hen’s egg! To both, it was clear that the semen
historical survey is arbitrarily defined, but throughout I played a role; they discussed, for example, whether there was
shall try to show the relevance of people’s thinking to the an equivalent contribution from the female (and its relation
ultimate aim of achieving an integrated view of what to menstrual blood) and also speculated on their respective
actually characterises embryogenesis. material involvements in forming the embryo and its mem-
Each historical period in science tends naturally to be branes. See also: Aristotle of Stagira; Galen of Pergamum;
dominated by a single set of leading interests, themes and Hippocrates of Cos; Müller, Johannes Peter (1801–1858)
methods. How the discipline we now call ‘developmental
biology’ was demarcated and what areas of concern were
included have changed considerably at different times. Early Scientific Period
Thus my division of the history of embryology into rough
chronological sections is not entirely artificial. Discussion Similar patterns of interest continued in much the same
will often focus for convenience on individual named sci- vein until the ‘scientific revolution’ of the seventeenth
entists. The choice is highly selective and arbitrary, but this century. From his notebooks and drawings, we know how
device allows brief summaries of viewpoints usually shared fascinated Leonardo da Vinci was by fetal anatomy. Des-
widely by many others. Advances in thinking can often be pite their accuracy in many respects his drawings are
marked out by key publications; embryology has certainly thought to be composites of human and animal obser-
been richly endowed with landmark documents. My vations. The placenta and umbilical cord were recognised
account will rarely address national differences, though as supplying nutrition to the fetus; maternal and fetal cir-
these are often interesting and important. culations were possibly already seen as separate. Vesalius
(1543) addressed the subject of development yet remained
firmly wedded to the preconceptions still inherited directly
from Galen; he was much pre-occupied by the fetal mem-
Glimpses of Embryology in branes and placenta. He depicted the human fetus with a
Pre-scientific Times placenta which he must have seen in the dog. Beliefs such as
the notion that round eggs were male, elongate female, still
In a sense embryology has always been of interest to indi- held sway. Ambroise Paré included in his surgical textbook
viduals contemplating their own nature; the mystery of life of 1579 a chapter on abnormal fetuses. But already, in
and death, sex, pregnancy, birth, life cycles in plants and the active criticisms and rebuttals that ensued between
animals all relate to it and, even in ancient civilisations, Vesalius and a number of contemporary scientists, one can
required concepts about development and the origin of see that appeal to evidence based on direct observations
living organisms. Ideas about development must have been (rather than classical authority) was increasingly recog-
closely linked with religious beliefs, concerning the entry of nised as essential. See also: History of Classical Anatomy;
the soul into the body for example. Our ability to recon- Vesalius, Andreas
struct earlier ways of thinking is no doubt severely limited, Vesalius’ heir (and Harvey’s teacher), Fabricius, is
but the evidence of myth, early art, and occasional early sometimes thought of as the author of the first illustrated,
scientific texts (some including illustrations) gives us at descriptive text on embryology (1604). William Harvey’s
least worthwhile pointers to the interest in such matters 1651 work on embryology (the result of his own long,
(see, e.g. J Needham’s A History of Embryology). Until intensive observations of hens’ eggs) is regarded as
the seventeenth century the facts of natural life must have important because, perhaps for the first time and as the
presented a prospect largely defying homogeneous beginning of a breaking away from Aristotle, the prospect
explanations and, given the limited possibilities of direct of a unified principle of development and reproduction –
knowledge of embryos themselves, concepts of develop- applicable across all species – was envisaged (Figure 1).
ment were largely filled in by inference from what little was Harvey interpreted the ovary in the deer and the pupa of

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 History of Developmental Biology

Figure 1 The frontispiece of Harvey’s book on the Generation of Animals (1651). It depicts Zeus liberating living beings from an egg (shown in detail on the
right). The picture symbolises his view of the uniformity of the principles of development across species.

insects as equivalent and as comparable initial steps in embryos was unnecessary. (It mattered little whether the
development. Others see Harvey as being the first to go preformed animal was seen in the egg or the sperm.) This
beyond a purely descriptive view of embryos (based at best was the age of ‘enlightenment’ or rationalism and of the
on a few static stages) to one that treated embryonic encyclopaedists and it can be argued that approaches to
development as a dynamic process requiring causal embryology reflected this general intellectual spirit.
explanations of the sequences of events. Descartes also Charles Bonnet emerged as the strongest advocate of
wrote an embryological treatise (printed 1664) in which he preformation (bolstered by his own rediscovery of par-
tried to apply mechanical explanations. See also: Des- thenogenesis, first noted in Harvey’s time). For Bonnet
cartes, René; Harvey, William; Reproduction and Life preformation represented a crucial part of a theological
Cycles in Invertebrates; Reproduction in Vertebrates: position. Others were equally strongly opposed; like Har-
Overview vey they favoured ‘epigenesis’, the gradual emergence of
Harvey used only weak magnifying lenses. Malpighi embryonic structure as a function of the developmental
(1673, 1689) made remarkably accurate drawings of chick process. The continuing debates, especially those involving
embryos, based on his pioneering use of microscopic Albrecht von Haller and Caspar Wolff, attracted wide
observations. By 1677 Leeuwenhoek had first described interest. Needham lists the many names of scientists
spermatozoa, thought by some actually to be parasites. involved in these controversial issues. But, despite the
Possibly triggered by Malpighi’s claim to have seen a fully philosophical nature of these debates, it is interesting that
formed chick embryo in an unincubated egg, put together they increasingly drew in wider and wider sources of data
with speculations on the age of the Earth and Descartes’ gleaned from the expanding knowledge of the range of
mechanical philosophy, there now emerged the concept of natural phenomena also characteristic of the time. Phe-
‘emboı̂tement’ (elaborated especially by the philosopher nomena like spontaneous generation, parthenogenesis,
Malebranche), the notion that all organisms are preformed vegetative reproduction, sexuality in plants, regeneration
in the germ of the egg and that the succession of generations (Trembly and Spallanzani), grafting, teratology, larval
had also been preformed at the time of the Creation. A stages, metamorphosis and the first ideas on genetics
number of early microscopists claimed to see fully formed (parental blending of characters) were all drawn in, often to
‘homunculi’ in eggs or sperm (Figure 2). See also: provide ingenious supporting arguments for one or other
Leeuwenhoek, Antoni van; Light Microscopy; Malpighi, position. See also: Bonnet, Charles; Haller, Albrecht von;
Marcello Spallanzani, Lazzaro; Wolff, Caspar Friedrich
Of progress in the eighteenth century, it is easy to con- However, there was really only one idea available;
clude simply that theory ran ahead of data. It has been whether development was a matter of preformation (also
suggested that, given the potency and clarity of the idea of known as ‘evolutio’, meaning overt manifestation of the
emboı̂tement, further direct observation and description of preformed germ) or its opposite, variously and vaguely

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 History of Developmental Biology

Figure 2 Illustration of the concept of preformation. On the left Hartsoeker’s drawing (1695) of a spermatozoon and on the right Malpighi (1673) of a
fertilised but unincubated hen’s egg, showing what was taken to be an already formed embryo (but was probably a misinterpretation of the deceptive
inhomogeneities that can occur in the early blastodisc). Both illustrate the idea that future structure is already present at the first stage of development.

known as ‘epigenesis’ or ‘metamorphosis’. Arguments Embryological phenomena and issues provided pivotal
about these two simple alternative positions had con- grounds on which grand theoretical speculations could be
siderably wider terms of reference than merely the nature of built. Preformation gave way to epigenesis.
embryogenesis. They were co-extensive with questions of As always, it is difficult for us now to imagine ourselves
the nature of life and its origins. Clear distinctions between back into modes of thinking of the time; even the words we
what we now call genetics, evolution and embryogenesis still share have often changed meanings dramatically. The
were impossible to make at the time, and in practice the perspective that now came to dominate was what we know
themes were all lumped together. Even so, by the end of the as ‘recapitulation’. Now the embryo was seen as a reflection
century some extraordinarily pioneering experiments and of the ladder of nature, scala naturae or, more generally,
observations were being made, for example tooth grafting, the ‘great chain of being’. The conception ‘has been one of
bone growth studies by John Hunter, and new epigenetic the half-dozen most potent and persistent presuppositions
interpretations of embryological observations by Wolff. in Western thought. It was _ (until the early nineteenth
See also: Hunter, John century) _ probably the most widely familiar conception
of the general scheme of things, of the constitutive pattern
1800–1880: The Era of ‘Recapitulation’ of the universe’, wrote AO Lovejoy in 1936. The increasing
complexity seen during the developmental stages of the
Around the turn of the century, the steady build-up of embryo was, according to this perspective, related to, and
empirical revelations was overtaken by the overarching paralleled in, the various scales of perceived complexity of
effects of a powerful philosophical movement (Romanti- the species found in Nature. Such parallels had been hinted
cism, Idealism or Naturphilosophie). It was particularly at by Aristotle; now, spurred on by no less a person that
powerful in Germany, the country that was increasingly Goethe, versions of this idea became commonplace. The
taking over from France as the leading scientific force. The idea of evolution was in the air in the long run up to
emphasis in directly studying Nature led to a wide-ranging Darwin. An early step was the notion of relating and
and integrative approach; in place of the fixed, ordering ranking species. Another was to recognise the stark alter-
principles sought in the earlier era, the new prospect was natives of fixity (creationism, emboı̂tement) and transmu-
of open-endedness, the possibilities of variety and tation of species. This approach to understanding embryos
changeability, a breaking out of pre-ordained systems. chimed with wider biological concerns; in other respects

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 History of Developmental Biology

Figure 3 The concept of recapitulation. Haeckel’s summary diagram of (three) developmental stages in eight different vertebrate groups, a fish on the left
and a human on the right. The remarkable similarities of the earliest stages support his claim that they correspond to a common, ancestral form; each species
seems to recapitulate ancestral morphologies during its own development.

embryogenesis remained mysterious. To quote the broad facts he had documented.) Darwin discussed
Encyclopaedia Britannica of 1842, ‘the immediate agency embryology only briefly, but Ernst Haeckel (his main
by which one living being is rendered capable of giving rise spokesman in Germany) made it the centrepiece of his
to another similar to itself is enveloped in the most pro- impassioned presentations (Figure 3). Moreover, he
found and most hopeless obscurity’. And yet ‘generation is returned to an extreme version of recapitulation which
at once the most obscure and the most wonderful of the others had already questioned; he believed that during the
processes occurring in organised bodies’ (Todd 1839, course of its development each embryo literally ‘recapitu-
quoted by Oppenheimer). See also: Evolution: History; lates’ the adult stages of its ancestors. He interpreted the
Goethe, Johann Wolfgang von ‘gastrula’ – the early stage seen in almost all embryos,
Darwin regarded embryology as ‘second to none’ among consisting of two germ layers surrounding a primitive gut
the various categories of evidence for his evolution theory cavity – as exactly equivalent to the common ancestral
because the similarity of early embryos across species was organism. The appeal of such thinking is obvious. If
direct evidence of descent from a common ancestor, and in ‘ontogeny recapitulates phylogeny’, this meant that one
some sense the progressive elaboration of an embryo sug- had direct access to the phylogenetic stages of each species
gested the sequence of changes that was envisaged in evo- by observing embryological stages. Throughout, Haeckel’s
lution. Epigenesis and evolution were parallel, dynamic main objective was the construction of phylogenetic trees.
concepts. The Estonian Karl von Baer (discoverer of the To recapture the power of these ideas one has to imagine
mammalian ovum and the notochord, and pioneer inter- how remarkable it must have seemed, in the wake of
preter of ‘germ layers’) was Darwin’s leading source of data Darwin, that the changes seen in evolution might be dir-
on embryonic stages in different species (i.e. his great Über ectly displayed in the still largely obscure phenomena of
Entwicklungsgeschichte der Tiere of 1828–1837). Von Baer embryogenesis. See also: Baer, Karl Ernst von; Cleavage
followed an anti-evolutionary (Cuverian) approach and and Gastrulation in Mouse Embryos; Darwin, Charles
had already explicitly argued against recapitulation; he Robert; Haeckel, Ernst Heinrich Philipp August
later rejected Darwin’s theory itself. (As Lovejoy has First writing in 1866 (in his aptly named Generelle
argued, it may be that von Baer so overstated his position – Morphologie, to be followed by books increasingly
wrapping it up in outdated philosophical speculations – addressed to the wider public), Haeckel was the ultimate
that Darwin found no difficulty in discounting von Baer’s schematiser, law maker and coiner of new terminology. He
own theoretical interpretation, while still making use of the summed up his ideas (in 1875) as ‘the biogenetic law’ (the

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 History of Developmental Biology

word ‘biogenesis’ had been coined by TH Huxley to Given the impact of Darwin’s theory and in the absence
describe how life never emerges by spontaneous generation of any real alternative understanding of embryological
but always from earlier living forms). At a time when phenomena, Haeckel’s viewpoint was all but inevitable.
development was widely interpreted in terms of an inbuilt His bold, all-embracing conceptual framework was
impulse of living organisms to unfold and complete seductively simple and briefly dominated the period. JHF
themselves (‘vitalism’), Haeckel’s theories were part Kohlbrugge identified no less than 72 variants of the theory
of an all-embracing ‘philosophy’ of biology which was of recapitulation prior to 1870.
unreservedly reductionist and materialistic. The facts of
sexual reproduction, fertilisation, spontaneous generation 1880–1925: The Ousting of Haeckel by
and the basic structure of the nucleus and cell were still Entwicklungsmechanik
being worked out. Speculative ideas (frequently inspired)
often anticipated the conclusions soon to be resolved Unsurprisingly, Haeckel’s grandiose, speculative style did
in such areas of uncertainty; Haeckel got near to locating not appeal to everybody and a counter-reaction soon set in.
heredity in the nucleus in 1866. And yet it is hard to It came initially (1874) from Wilhelm His who pointedly
avoid the suspicion with Haeckel that detailed evidence was showed how embryogenesis can be seen as the product
subjugated to the prior aim of inferring underlying unifying of simple (mechanical) forces acting within the embryo as
forces, principles and laws. See also: Huxley, Thomas it forms (i.e. as caused in the here-and-now rather than
Henry as products of earlier evolution). Haeckel and His were
In this period embryology was a key concern for most divided by each other’s combative style, by background
biologists. If there was one word that summed up the (zoology as opposed to medicine), the species they studied
driving aim of all nineteenth century morphologists it was and aims. His was not much interested in evolution or
‘form’; their underlying objective was to explain the origin heredity, but much concerned with methods and accuracy –
of ‘form’ (i.e. morphology) and here little distinction was he anticipated later concepts such as those of ‘fate maps’ –
drawn between adult comparative anatomy and embry- and he was no experimentalist either. But one of Haeckel’s
ology. The two subjects went hand in hand as ‘evolutionary own pupils, Wilhelm Roux, was just as important in defining
morphology’. The classical themes of the pentadactyl (five- an alternative. Starting from the evolutionary perspective –
digit) limb, skull segmentation, germ layers, gill arches and he was initially interested in ‘natural selection’ between
so on, were already being thoroughly documented, but now parts of embryos and adaptation to function within the
took on new force. Embryology provided a new means of developing embryo (1881) – he followed a path close to that
establishing ‘homologies’. The second half of the century of August Weismann. In 1888 in an attempt to test and
was the golden age of both comparative and descriptive confirm Weismann’s theory of ‘germinal selection’ (i.e. the
embryology. Moreover, descriptive histology, as well as notion that separate, inherited (particulate and probably
microscopic and preparative techniques, had reached vir- nuclear) determinants come to control separate parts of
tually modern standards by the 1860s. Although, hitherto, the egg and future embryo) he introduced a classic piece
embryos had been analysed primarily in terms of germ of methodology; killing one of the initial cells of a two-cell
layers, now it was in terms of cells. Rudolph von Kölliker stage frog embryo. The result was the development of a
published the first embryological textbook based on cell half-embryo (an example of ‘mosaic’ development; each
theory (and including much physiology) in 1861. Balfour’s part developing as if its distinctive elementary parts
A Treatise on Comparative Embryology (1880–1881) had already been arranged in their final pattern). The
was a landmark, and similar monumental, encyclopedic experimental approach and search for here-and-now causes
accumulations of documentation followed, for example of development was described and promoted by Roux
Wilhelm His’ (1885) followed by Keibel’s (1897–1938) as ‘Entwicklungsmechanik’. He edited the first specifically
series of ‘normal tables’ of various species including embryological journal (1895, Wilhelm Roux’ Archiv für
humans. But a form of fragmentation was also emerging; in Entwicklungsmechanik der Organismen). The two ap-
Otto Hertwig’s massive, multivolume Handbuch der proaches to embryology came to be seen as opposed to each
vergleichenden und experimentelle Entwicklungslehre der other and incompatible. Descriptive study gave way to
Wirbeltiere (1906) and Keibel and Mall’s authoritative experimental technique. The new mood (based especially on
Manual of Human Embryology (1910–1912) develop- physiology) emphasised methodological rigour; that is
mental data were organised on the basis of specific organ against Haeckelian-type ‘speculation’ and against the
systems rather than species. See also: Cladistics; Hertwig, vitalism that was still prevalent. See also: His, Wilhelm;
Wilhelm August Oscar; His, Wilhelm; History of Com- History of Physiology; Roux, Wilhelm; Weismann, August
parative Anatomy; History of the Optical Microscope in Friedrich Leopold
Cell Biology and Medicine; Homology and Homoplasy: A In the period 1880–1900, the ground was laid for the
Philosophical Perspective; Homology in Character Evo- rediscovery of Mendel’s laws. Weismann is representative
lution; Koelliker, Rudolf Albert von; Morphological of a host of biologists (including notably Theodor Boveri
Evolution: Epigenetic Mechanisms; Morphology and and Roux) who were working towards what we now rec-
Disparity through Time; Owen, Richard; Philosophy of ognise as genetics. Their main evidence was cell theory and
Biological Classification; Systematics: Historical Overview the behaviour of chromosomes, particularly in meiosis,

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fertilisation and cell division. Weismann led the way in Zebrafish Embryos; Conklin, Edwin Grant; Driesch, Hans
attacking the concept of the inheritance of acquired char- Adolf Eduard; Evolutionary Developmental Biology:
acters. EB Wilson’s classic review (1896) covers all these Developmental Bias and Constraint; Harrison, Ross
issues; its title, The Cell in Development and Inheritance, Granville; Medawar, Peter Brian; Morphometrics; Regen-
says it all. Another approach was to document morpho- eration: Principles; Sea Urchin Embryo: Specification of
logical variation in all its manifestations, together with Cell Fate; Thompsonian Transformations; Tunicate
patterns of inheritance of such variants, as in William Embryos and Cell Specification
Bateson’s compendium of teratology, Materials for the A new focus gradually emerged through the work of
Study of Variation (1894). This was a fertile period when Herbst and particularly Hans Spemann. They began the
much of our basic knowledge of biological phenomena was laborious task of unravelling the causal mechanisms
first clarified and laid down. Embryology remained underlying embryogenesis. Starting with the formative
important, if for no other reasons than that the relevant influence on the lens by the eye (1901) Spemann elaborated
generation of leading biologists had, almost without a theory of ‘embryonic induction’ by one part of the
exception, been trained initially as embryologists (in such developing embryo of the next part to be formed.
institutions as Naples, Woods Hole and Plymouth marine If one can pick on one event that stands out in this period
biological stations) and that it provided many of the con- as particularly important, it must be the discovery of the
cepts and starting points which led to a hiving off of ‘organiser’ by Spemann in 1924 (Figure 4). Here a small
definitive and distinct subjects such as genetics and cell region of the early amphibian embryo was shown to have
biology. See also: Bateson, William; Boveri, Theodor; the power to influence and control the entire emergence
History of Classical Genetics; Mendel, Gregor Johann; of patterned differentiation throughout the embryo. A
Wilson, Edmund Beecher remarkably elegant technical feat in itself, this turned out to
But if we try to pick out the most important theoretical
issue that now began to emerge for embryologists, the
contrasting implications of the work of Roux and Hans
Driesch sum matters up well enough. Following up Roux’s
study on the frog, Driesch (1891) discovered an entirely
different result in the sea urchin, namely ‘embryonic
regulation’, the formation of a complete normal embryo
from separated single cells. It was this discovery that in
effect defined the most central problem within the field of
embryology: how can organismal structure emerge in all its
ordered and predictable complexity if it starts from a cell
with apparently no fixed initial, inherent organisation? But
it was to take a long time for the full implications of the
opposed findings of mosaic and regulative development to
be understood. The experimental approach was proving so
powerful and revealing that it was sometimes pursued for
its own sake, it seems, rather than for any search for a
unifying core of embryological theory. Regeneration was a
favourite experimental route for some (e.g. Morgan,
Child). New and powerful techniques such as tissue culture
and cross-species transplantation (both pioneered by Ross
Harrison) were elaborated. Concepts such as allometry,
control of growth, involution and malignancy were per-
ceived as related to embryology but the precise nature of
relationships was rarely a primary concern. The tendency
towards specialisation and fragmentation increased as
individual laboratories developed experimental techniques
suited to particular organs and particular species (sea
urchin, frog and later chicken, mammal). Notable schools
of embryologists concentrated on ‘polarisation’ of the newt
limb (led by Harrison) and studies of the fate maps and
mosaic properties in marine invertebrate embryos (e.g.
Whitman, Lillie, Conklin at Woods Hole). See also: Figure 4 Spemann’s discovery of the ‘organiser’. A small region of tissue
Cleavage and Gastrulation in Avian Embryos; Cleavage (at the dorsal lip of the blastopore, shown blue) has a unique, organising
property; when transplanted to a noncorresponding site in a host embryo it
and Gastrulation in Mouse Embryos; Cleavage and causes the formation of a complete, secondary embryo by nearby host cells
Gastrulation in Sea Urchin; Cleavage and Gastrulation (shown below). Elsewhere the host embryo develops normally (shown
in Xenopus laevis Embryos; Cleavage and Gastrulation in above).

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be a watershed. This discovery seemed to solve the problem embryology from the main currents of biology. This was
of the origin of embryonic organisation at a stroke. Even in the period leading up to the ‘modern evolutionary syn-
1934, Huxley and De Beer’s well-known textbook, The thesis’ (a definitive linking up between Darwinism and
Elements of Experimental Embryology, was largely devoted genetics) and here again embryology was to prove an
to working through its various implications. The concept irrelevance. In 1934, Thomas Hunt Morgan’s book
of epigenesis was beginning to acquire real meaning Embryology and Genetics unwittingly made it all too clear
because there was now a methodology for defining how deep the divide was between the two subjects, how
sequences of cause and effect during embryogenesis. Spe- little they interrelated, and, by contrast to genetics, how
mann already recognised the equivalence of the nuclei in little embryology was moving forward. See also: De Beer,
each early embryo cell (anticipating later proof by JB Gavin Rylands; Evolutionary Ideas: The Modern Syn-
Gurdon). Thus embryonic differentiation was not the thesis; Heterochrony; Huxley, Julian Sorrell; Morgan,
result of Weismannian ‘germinal selection’ or preform- Thomas Hunt
ation but of epigenetic interactions between cells. See also: There now followed a period that led to a descent into
Beyond the Genome; Cleavage and Gastrulation in confusion for embryology. Related disciplines were mak-
Xenopus laevis Embryos; Ecological Development Bio- ing major progress; biochemistry, nutrition, endocrinology
logy; Epigenetics: Influence on Behavioral Disorders; were all more or less in their golden ages. Following the
Lens Induction; Morphological Evolution: Epigenetic discovery of the organiser, the assumption among embry-
Mechanisms; Secondary Induction: Overview; Spemann, ologists was that gradients and organisers would turn out
Hans to be chemically identifiable and that they would operate
very much like hormones. Needham equated the terms
1920–1950: The Era of Biochemistry ‘organiser’ and ‘morphogenetic hormone’. However, des-
pite much investment of effort and many claims through
On the theoretical side the organiser phenomenon became the 1930s, no such identification emerged. Embryology had
closely associated with a variety of often poorly defined, failed in its bid to follow and even lead advances in the
interpretative concepts such as those of fields and gradi- modern approach to reductionist biology. By the end of the
ents, all ultimately designed to explain the two key prob- 1930s the sense of disappointment and disillusionment was
lems: (a) how does an integrated pattern of structures manifest. See also: History of Biochemistry; History of
emerge and (b) how does the embryo ‘regulate’ its pattern Nutritional Science; Reduction: A Philosophical Analysis
in the face of early lesions and transplants? Embryology,
which has long been closely associated with ‘theoretical 1945–1985: The Era of Cell Biology
biology’, provided key evidence in discussions of funda-
mental issues concerning vitalism and later holism or The combined effects of disillusionment before the Second
organicism (Woodger, Bertalanffy, Weiss). The ‘Theore- World War, the hiatus of the war itself and the emigration
tical Biology Club’, an influential discussion group oper- and career changes of leading figures – during the 1940s
ating in Cambridge and London through much of the 1930s many leading embryologists left the field or deflected into
and involving many of the most prominent researchers, one of the various subspecialties within embryology –
was much concerned with defining methods and concepts brought something of a new start to the discipline. Now, as
appropriate to tackle such problems. Were the various the techniques of cell biology rapidly advanced, the
concepts that were increasingly invoked to explain organ- emphasis was on the cell or subcellular level. The whole
iser effects and embryonic patterning (e.g. twinning, mirror embryo was of less and less concern in laboratory research.
imaging, polarity, competence, double assurance, etc.) It is apparent that, despite its importance, Spemann’s
more than metaphorical (the terms were often borrowed organiser concept was coming to have some unfortunate
from engineering and physics); what was their biological effects on the way in which the discipline of embryology
and cellular basis? was regarded in the context of the rapidly advancing,
In the 1920s Garstang, De Beer and JS Huxley still strongly positivist and reductionist trends of the biological
sought to identify the weaknesses of Haeckel’s position mainstream. The organiser was increasingly perceived
and, on the basis of growing knowledge about ‘larval as a return to ways of thinking more typical of Nat-
forms’, complex life cycles and metamorphosis (and with urphilosophie. Spemann was accused of a ‘vitalistic’
an eye to palaeontology, which was still holding out for the approach. ‘Organicist’ would be fairer and nearer the
concept of recapitulation), introduced the idea of ‘paedo- mark. See also: History of the Optical Microscope in Cell
morphosis’ as a form of ‘heterochrony’. This process Biology and Medicine
involves the exact opposite of the embryonic changes nee- But this was to be an era of further fragmentation, based
ded for recapitulation and was persuasively proposed as to a considerable extent on new techniques in cell biology,
the mechanistic basis for the creation of new evolutionary such as the electron microscope and radioactive tracers.
phyla. This was not only a final nail in Haeckel’s coffin, but Now nucleo-cytoplasmic interactions, cell membranes,
as a perspective on embryological phenomena, it provided cell-to-cell communication channels and extracellular
few connections with embryologists in the Entwicklungs- matrices were of most interest. An example of research that
mechanik tradition and had the effect of further detaching was much admired was that of Gurdon; by transplanting a

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 History of Developmental Biology

nucleus from a gut cell into an enucleated frog’s egg and defined by the technology of molecular biology (and later-
thereby demonstrating that each cell nucleus contained all added signalling molecules such as ‘growth factors’) now
the genetic information needed to mediate full develop- came increasingly to dominate the field. Techniques
ment of a new frog (what we would now call ‘cloning’), originally developed in Drosophila now showed remarkably
Gurdon finally demonstrated what embryology and similar applicability in vertebrate embryos. Christiane
regeneration had long implied; the nucleus contains all the Nüsslein-Volhard’s Nobel Prize, awarded for this work in
information for development, but it is the job of the 1995, was only the second ever to be given to an embry-
embryological process to read it out. Significantly, despite ologist. The first had gone to Spemann. See also: Drosophila
its title (The Control of Gene Expression in Animal Devel- Embryo: Homeotic Genes in Specification of the Anterior–
opment), his book of 1974 was solely concerned with studies Posterior Axis; Evolutionary Developmental Biology:
within the single (egg) cell and had almost nothing to say Homologous Regulatory Genes and Processes; Evolution-
about developmental events beyond that. See also: The Cell ary Developmental Biology: Hox Gene Evolution; History
Nucleus of Molecular Biology
In this period the field acquired a new name (‘develop- In this recent period there has been a swing of interest
mental biology’), echoing and presumably modelled on the towards developmental genetics (a subject that had long
earlier terms ‘molecular biology’ or ‘cell biology’. This struggled to make a broad contribution within develop-
seems to have symbolised the aspiration towards mod- mental biology), and towards evolutionary matters. The
ernity and a holding together of the wide diversity of spe- relevance of embryology to evolution had been recalled as
cialist interests that might shelter under this umbrella. (It an issue, significantly not by an embryologist but by a
certainly implied an aspiration to locate embryology within palaeontologist (SJ Gould, in 1977). But the renewed
biology as a whole.) In the USA ‘The Society for the Study interest in ‘evo/devo’ is largely the result of new sources of
of Development and Growth’ became the ‘The Society for data on molecular sequence homologies within and across
Developmental Biology’ in 1951, and its new journal phylogenetic groups. These data not only directly indicate
Developmental Biology started in 1959. But one gets the shared phylogenies but they are also pointing to remark-
impression that any sense of integration and overall pur- able evolutionary constancies in embryological mech-
pose was strained. One indication of this was the resurgence anisms among different species. See also: Evolution of
of interest in theoretical biology and ‘model building’ Development; Human Genome Project as a Social Enter-
which often drew in developmentalists. The leading UK prise; Human Genome Project, HUGO and Future Health
embryologist, Conrad Waddington, organised a series of Care; Human Genome Project: Importance in Clinical
symposia (later published, as Towards a Theoretical Biol- Genetics; Molecular Evolution: Introduction; Molecular
ogy (1968–1972)). The need to define priorities was simi- Phylogeny Reconstruction; Sequencing the Human Gen-
larly felt in the United States, and the resulting attempt ‘to ome: Novel Insights into its Structure and Function
overcome the trends of overspecialisation by encouraging a Achieved by a distinctly roundabout route, today we are
wider, interdisciplinary perspective and by integrating the seeing a return in developmental biology to many of the
ever-growing volume of accumulated information into a older and classical problems of evolution. The grain of
broad conceptual framework’ was published in the influ- truth concealed in Haeckel’s exaggerated claims may yet
ential volume Analysis of Development (1955). The theo- emerge. History seems almost to be repeating itself.
retical heart-searching did lead to an important outcome in Another trend is the emergence of ‘embryology’ in the
Wolpert’s entirely new (and all-encompassing) concept of form of its applications within medicine. Key steps are
‘positional information’ (1969), which in many respects listed in Table 1. Until very recently embryology as studied
rejected and replaced the traditions built on the study of in university research laboratories or marine stations had
induction and organisers. There followed a new interest little relevance to either medical, veterinary or agricultural
in gradients and their molecular basis (e.g. the putative practice. Attempts to culture embryos and achieve in vitro
morphogen, vitamin A). fertilisation go back to the start of the century, but practical
application dates from the first successful human embryo
transfer and birth of Louise Brown in 1978. Even now the
links between embryological knowledge and medicine are
The Era of Molecular Biology and tenuous. The new medical procedures owe little to theory
Genetic Engineering: 1985– or scientific knowledge, but are often largely pragmatic.
Further merging of medical embryology and molecular
By 1985 molecular biology had not impacted on develop- techniques will presumably come through future ‘gene
mental biology in any significant way, although it was therapies’ and genetic engineering. Looking to the future,
already offering powerful techniques. But it was around this the prospects are of increasing exploitation of stem cells as
time that a bridge was established in the form of the dis- a route to organ replacement. Such applications have given
covery of the role of ‘homeotic genes’ in the control of body new meaning to the word ‘embryology’ in public per-
(particularly segmental) form. The molecular elucidation of ceptions. In their train have come feminist, ethical and
gradients may have failed to reach any definitive resolution, political dimensions never dreamt of by embryologists of
but these new families of specific pattern-forming genes the past. With the prospect of ‘designer babies’ looming,

ENCYCLOPEDIA OF LIFE SCIENCES & 2010, John Wiley & Sons, Ltd. www.els.net 9
 History of Developmental Biology

Table 1 Key events in medical applications involving embryology

Artificial insemination in humans, end of nineteenth century (may have been tried by John Hunter); in horses by Babylonians,
1400 BC; Royal Commissions reject it, 1948, 1960; widely adopted during 1970s
Experimental initiation of egg cleavage (‘artificial parthenogenesis’); Loeb, 1899
First Midwives Act; 1902
First sex hormone identified; (oestrogen) 1923
Pregnancy test; 1930
Mammalian embryo transfer; 1934
First in vitro fertilization (IVF) of human ovum; 1946
Amniocentesis; 1877, becomes ‘routine’ around 1953; first routine prenatal diagnostic techniques; 1956 (for sex-linked inherited
disease)
First contraceptive pill marketed; 1960
First in utero surgery; 1963, fetal blood transfusion (for haemolytic disease of the fetus)
The Abortion Act; 1967; first legal abortion in UK
Ultrasound prenatal monitoring; 1958, becoming routine around 1975
First human egg transfer and birth; Louise Brown, 1978
Embryo cryopreservation in humans; 1984
Human Fertilization and Embryology Act; 1990
First cloning in a mammal; ‘Dolly’ the sheep, 1997
Sequencing of the human genome; 2000

aspects of embryology are now of wide public interest. with embryology have differed and diverged. In plants
See also: Bioethics in Embryonic Research; Bioethics distinct developmental processes seem to apply and botany
of New Assisted Reproduction; Cloning of Animals in has provided few links with animal embryology. Unlike
Genetic Research: Ethical and Religious Issues; Dolly and most disciplines, embryology could not claim (despite
Polly; Human Cloning: Arguments Against; Human deploying much experimental ingenuity and skill) a single
Cloning: Arguments for; Human Gene Therapy; In Vitro ‘method’ or ‘technique’ that was unique to it or charac-
Fertilization; Stem Cells and Treatment of Neurodegen- teristic of it. But I would highlight two interesting features,
erative Disorders; Steptoe, Patrick which perhaps help explain some of the peculiarities of the
history of embryology. See also: History of Plant Sciences;
Phenotypic and Developmental Plasticity in Plants
Final Observations Firstly, in the long, slowly moving course of the history
of the subject, embryology’s own historical legacy has in
In its long history the discipline of embryology has had a itself played a part in significant ways. The discrediting of
number of highs and lows. It can be argued that throughout the Haeckelian superstructure happened in such a way that
the nineteenth century the subject stood near centre-stage it left a negative attitude to the subject at a critical time,
among the concerns of biological scientists; that it was an which had traumatic and long-term effects. Its replacement
integral part of general thinking and indeed was the source by Entwicklungsmechanik owes something to an overt
of leading ideas. The issue of the origins of form was seen, in opposition to Haeckel. Not only recapitulation theory but
effect, as the very essence of biology. But in the last century, along with it the descriptive and comparative methods so
despite a number of dramatic achievements, the discipline central to embryology became tainted and ‘old fashioned’.
failed to cohere or to prosper in its own right. Generalising, The negative taint has reverberated subtly right up to the
one can say that from around 1880 it has retreated from present. In the repositioning of different biological discip-
being a core subject. It became highly specialised and came lines that occurred in the establishment of the ‘modern
to be ignored, probably with relief, by most biologists. evolutionary synthesis’ in the 1940s, embryology was
Clearly, many considerations contributed to this conspicuously absent. The sceptical and antagonistic
chequered history. Needham (1959) offered some explan- approach to recapitulation taken by embryologists with
ations; he lists a number of ‘limiting factors’, including evolutionary interests in the 1920s (Garstang, Huxley, De
technical, theoretical, terminological, psychological and Beer) was an important determinant in this surprising
interestingly the ‘balance between Speculation, Obser- situation. One consequence has been that, until very
vation and Experiment’. In contrast to many comparable recently, any consideration of the evolutionary impli-
disciplines (e.g. genetics, biochemistry, immunology or cations of embryology remained somehow suspect;
endocrinology), embryology was not ‘useful’; until very Haeckel and the concept of recapitulation are still
recently it derived no focus from any direct practical remembered as a cautionary tale. Even today echoes of
implications or applications in medicine or elsewhere. Haeckelian thought remain embedded and unavoidable in
Throughout history, medical and biological involvements everyday embryological concepts such as germ layers or

10 ENCYCLOPEDIA OF LIFE SCIENCES & 2010, John Wiley & Sons, Ltd. www.els.net
 History of Developmental Biology

gastrulation, but the classical database of descriptive phenomenology. First its descriptive base in ‘evolutionary
comparative embryology remains largely forgotten and morphology’ was challenged and made redundant; then,
work in this area has low status. See also: Haeckel, Ernst with increasing reductionism, the classical phenomena
Heinrich Philipp August revealed in whole, living embryos during the last century
A second factor that influenced embryology in a par- became ever harder to maintain as central explanatory
ticularly confusing way was the effect of a gradual, general targets within the discipline. A broad view of embryology –
trend affecting almost all areas of biology: reductionism a view which remains open to considerations that
(Figure 5). Embryology has played a leading and often were emphasised perhaps much earlier in the history of
honourable role in the countering of notions of vitalism in the subject – suggests that the reductionist path by its
biology. But as biology as a whole moved in the alternative, nature fails to address some crucial problems. In spite of
materialistic/reductionist direction – first through cell its obvious logical, fundamental and comprehensive
theory and physiology, then genetics and biochemistry, explanatory power, the exclusively molecular approach
and eventually molecular biology and molecular genetics – is ill-suited to solving certain embryological issues. As
embryology has increasingly lagged behind. As each all ‘nature/nurture’ interactions demonstrate, ‘environ-
reductionist phase came and went embryology’s search for mental’ factors extrinsic to the genome or the embryo are
its core principles tended to be sought in those terms. The equal partners with genes as determinants of embryo-
boundaries of disciplines became blurred. A succession of genesis. It is hard to see how a molecular account can ever
subspecialties formed on the basis of new techniques and, satisfactorily and fully ‘explain’ something like neural tube
often, a single preferred ‘model organism’. Each passing folding or adaptation of bone structure to gravitational
change of biological focus tended to bring with it its own forces. Mechanical as well as molecular forces are required.
claims to provide the key to a ‘solution’ to the problems of Moreover, many embryological phenomena operate at a
embryology and there has been a long-term failure of the scale and explanatory ‘level’ (i.e. that of tissues or whole,
subject to reach a consensus on defining principles. A intact embryos) above the molecular. Although, in prin-
precondition for any discipline to define itself is the estab- ciple, embryological phenomena (which are typically
lishment of an appropriate, stable, generally accepted, multicausal) must ultimately be reducible to the sum of
integrative set of concepts and methods of its own, which underlying molecular events, achieving this in practice
sufficiently embrace the core database of the subject (i.e. the soon becomes unattainably complex. See also: Nature/
establishing of the terms of reference in which the subject Nurture - A Philosophical Analysis
ultimately needs to be thought about and the sources of Embryology remains ill defined as a discipline pri-
evidence they should cover). Haeckel provided just such a marily because of its ‘complexity’ in terms of potentially
system. Molecular genetics now seems to offer an alluring relevant data and multiple levels of consideration. Single,
new system. See also: Reductionism in Biology direct ‘solutions’ (whether conceptual or through a
All these forces operated against achieving coherent, hypothesised, key controlling molecule) are bound to be
sustained discipline formation. The subject has never inadequate. The future sorting out of priorities in
managed to settle on a conceptual core or on an agreed embryology will require methods for handling complexity
and for the integration of hierarchies of multilevel con-
cepts. It seems likely that, in seeking to achieve a stable
discipline structure, embryology needs to retain some of
its traditional concepts as well as reap the benefits of
molecular genetics. Ironically, although they are not
often spoken of in this way, the dilemmas posed by
molecular reduction recall the most classical and funda-
mental of embryological themes, the preformation/epi-
genesis dichotomy. Molecular reduction poses all the
explanatory problems that we associate with any concept
of preformation. Any such approach in itself explains
nothing because it merely pushes the problem of the
origin of embryological pattern formation back into
earlier stages or more fundamental explanatory terms.
Only through ‘epigenesis’ (Figure 6) can we escape this
circularity, but the notion of epigenesis remains vacuous
until the relevant mechanisms (such as interactions of
parts) have been defined and understood. Only by invoking
Figure 5 A cautionary comment on changing perspectives in cellular interactions occurring during the process of
embryological research methods. Although classically the whole intact embryogenesis can we explain how the passive structure of
embryo was the focus (on the left), now attention is increasingly
biochemical (a homogenate of an embryo, middle frame); or molecular
the genome common to all cells can be deployed differen-
(an isolate of selected molecular components; right). (From Weiss PA tially to make patterned structural elaboration in the
(1968) Dynamics of Development: Academic Press.) embryo possible.

ENCYCLOPEDIA OF LIFE SCIENCES & 2010, John Wiley & Sons, Ltd. www.els.net 11
 History of Developmental Biology

Fertilised ovum
by cleavage divisions to
Anal canal
Thyroid Morula Enamel of teeth
Middle ear
Oral epith.
Digestive tube I Eustachian tube by hollowing out and expansion Hair Proctodaeal
II Tonsillar Pars buccalis
Pharyngeal recesses Sebaceous epithelium
Liver pouches III Primitive thymus Blastodermic vesicle Nails
Stomodaeal of hypophysis
glands
Parathyroids with inner cell mass epithelium
IV Parathyroids Differentiation Sweat glands Nasal and olfactory epith.
Pharynx Post-branchial bodies at primitive streak and olf. nerve
Pancreas Trachea Endoderm Outer epithelium
Bronchi Lungs Mammary glands of body
Allantois Ext. - emb. ent. of
yolk sac Notochord Lens of eye
Ectoderm
Mesoderm Auditory
Urinary Primitive vesicle
Brain
bladder gut Cranial motor
Ext. - emb. ect. of
amnion and serose Inner ear nerves
mechanism Epiphysis
Head mesenchyme Neural tube
Intermediate Pars neuralis
Neural crest of hypophysis
mesoderm
Outer layers Cranial Optic vesicles
Cephalic c.t. Lateral
and muscles
of eye
Dorsal Pronephros Spinal dorsal sensory gang. Retina
mesoderm root ganglion and nerves
Dentine mesoderm and optic nerve
Skull Sensory spinal
Mesonephros
Primary (somites) Sympath. nerve roots
of ductuli efferentes Spinal cord
sex gang.
Sclerotomes Müllerian ducts Metanephros Motor spinal
cells Axial skeleton Kidney tubules Medulla of nerve roots
Oviducts Vagina Metanephric divert. adrenal
Myotomes Ext. - emb.
Appendicular skeleton Uterus Ureters Renal pelvis Cortex of
Collecting tubules mes. of adrenal
Dermatomes Mesonephric ducts Extramb. mes. of yolk sac
Buds of appendages
and allantois
Skeletal muscles amnion and serose
of trunk Connective tissue
Ductus epididymis Splanchnic mesoderm
Ductus deferens
Muscles of layers of skin
appendages Somatic mesoderm Visceral pleura
Pleura Visceral peritoneum
Parietal Pericardium Haemangioblastic Mesenteries
Peritoneum tissue
(?) Gonads Primary sex cells Epimyocardium
Stroma Mesenchyme Blood
Gamete Epicardium
Mother of corpuscles Heart
Myocardium
cells gonads Connective tissue and smooth Endothelium of Endocardium
Gametes muscle of viscera and blood vessels blood vessels

Figure 6 An illustration of the concept of epigenesis. This flow diagram shows the cumulatively elaborate sequence of events in embryogenesis
(as illustrated in a typical recent textbook). Starting from the ovum (top, centre) the branching paths show steps in the development of the range of body
organs. Each involves multiple stages. Most of the arrows indicate experimentally proven (inductive) cause and effect embryonic tissue interactions.

Further Reading Horder TJ, Witkowski JA and Wylie CC (1986) A History of

Embryology. Cambridge: Cambridge University Press.
Allen G (1975) Life Science in the Twentieth Century. New York: Laublichler MD and Maienschein J (eds) (2007) From Embry-
Wiley. ology to Evo-Devo: A History of Developmental Evolution.
Cole FJ (1930) Early Theories of Sexual Generation. Oxford: Cambridge, MA: MIT Press.
Clarendon Press. Lovejoy AO (1936) The Great Chain of Being. Cambridge, MA:
Deech R and Smajdoe A (2007) From IVF to Immortality. Con- Harvard University Press.
troversy in the Era of Reproductive Technology. Oxford: Oxford Meyer AW (1939) The Rise of Embryology. Stanford: Stanford
University Press. University Press.
Gould SJ (1977) Ontogeny and Phylogeny. Cambridge, MA: Needham J (1959) A History of Embryology. Cambridge: Cam-
Harvard University Press. bridge University Press.
Hopwood N (2009) Embryology. In: Bowler PJ and Pickstone JV O’Dowd MJ and Philipp EE (1994) The History of Obstetrics and
(eds) Cambridge History of Science. Vol. 6. Modern Life and Gynaecology. New York: Parthenon Publishing Group.
Earth Sciences, pp. 285–315. Cambridge: Cambridge Uni- Oppenheimer JM (1967) Essays in the History of Embryology and
versity Press. Biology. Cambridge, MA: MIT Press.
Horder TJ (2001) The organizer concept and modern embryology: Russell ES (1917) Form and Function. London: John Murray.
Anglo-American perspectives. International Journal of Devel- Singer C (1931) A Short History of Biology. Oxford: Clarendon
opmental Biology 45: 97–132. Press.
Horder TJ (2008) A history of evo-devo in Britain. Annals of Willier BH, Weiss PA and Hamburger V (eds) (1955) Analysis of
History and Philosophy of Biology 13: 101–174. Development. Philadelphia: Saunders.

12 ENCYCLOPEDIA OF LIFE SCIENCES & 2010, John Wiley & Sons, Ltd. www.els.net