The Theory of Evolution

The Theory of Evolution and Its Impact
.
Aldo Fasolo
Editor
The Theory of Evolution

and Its Impact
Editor
Aldo Fasolo
Dipartimento di Biologia Animale e dellUomo
Universita di Torino
Via Accademia Albertina 13
I-10124 Torino
Italy
aldo.fasolo@unito.it
The publication of this book has been made possible by the financial support of the Acca-
demia delle Scienze di Torino.
ISBN 978-88-470-1973-7 e-ISBN 978-88-470-1974-4

DOI 10.1007/978-88-470-1974-4
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Contents
Introduction: The Sand Walk (on the Darwins Steps) . . . . . . . . . . . . . . . . . . . . . 1

Aldo Fasolo
Idola Tribus: Lamarck, Politics and Religion in the Early

Nineteenth Century . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
Pietro Corsi
Darwinism Past and Present: Is It Past Its Sell-by Date? . . . . . . . . . . . . . . . 41

Michael Ruse
Evolutionary Theory and Philosophical Darwinism . . . . . . . . . . . . . . . . . . . . . . . . 53

Paolo Casini
Struggle for Existence: Selection, Retention and Extinction

of a Metaphor . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
Peter Weingart
The Theory of Evolution and Cultural Anthropology . . . . . . . . . . . . . . . . . . . . . 83

Henrika Kuklick
The Concept of Evolution in Linguistics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103

Manfred Bierwisch
Theory of Evolution and Genetics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119

Alberto Piazza
Genes, Evolution and the Development of the Embryo . . . . . . . . . . . . . . . . . . . 131

Giuseppina Barsacchi
v
vi Contents
Evolutionary Mechanisms and Neural Adaptation: Selective Versus

Constructive Strategies in the Development and Plasticity
of the Nervous System . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 159
Ferdinando Rossi
Is the Human Brain Unique? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 175

Gerhard Roth
Aristotle and the Chicken: Animacy and the Origins of Beliefs . . . . . . . . . . 189
Giorgio Vallortigara
Evolution: Remarks on the History of a Concept

Adopted by Darwin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 201
Volker Gerhardt
An Evolving Research Programme: The Structure of Evolutionary

Theory from a Lakatosian Perspective . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
Telmo Pievani
.
Introduction: The Sand
Walk (on the Darwins Steps)
Aldo Fasolo
Abstract To understand the status of Theory of Evolution, highly multidisciplinary

approaches are needed. Thus, the book moves from the historical and philosophical
roots, to follow a long and winding road, passing from anthropology, to linguistics,
genetics, developmental biology, neuroscience, cognitive studies, to find a final lap
on today theories. The inescapable conclusion, quoting the contribution of the
philosopher Michael Ruse, is that in fifty years or a hundred years we will still
have the theory of the Origin around. Great, precisely because it does not stand still,
but remakes itself and grows and changes by virtue of the fact that it gives such a
terrific foundation. Is Darwinism past its sell-by date? Not by a long chalk yet!
Year 2009 celebrated the triumph of Darwin as global superstar, spinning from the
pop icon to the actual understanding to what make him a great innovator, able to
give a turn to the whole modern culture. After such a deluge of books, conferences,
reviews, gadgets, what is today our vision on theory of Evolution and its Impact?
This was exactly the goal of an inter-academy meeting held in Torino (May 2729,
2010) involving the Accademia delle Scienze di Torino, the Accademia Nazionale
dei Lincei and the Berlin-Brandenburgische Akademie der Wissenschaften.
The preliminary question was obviously if we needed another meeting on such a
topic. In the commentary about a book recently published on the first 150 years
since Darwin [1], reporting the dramatic expansion of the applications of evolu-
tionary science in recent years and the wages in terms of confirmations and
extensions, David P. Mindell closes saying: Does all this activity mean evolution
has lost its ability to excite fear and opposition? Not yet. As the root for natural
explanations of human origins . . . and ultimate impetus for human moral behaviour
and values, evolution remains the disturbing discovery.
A. Fasolo (*)
Department of Animal and Human Biology, University of Turin, Turin, Italy
e-mail: aldo.fasolo@unito.it
A. Fasolo (ed.), The Theory of Evolution and Its Impact, 1

DOI 10.1007/978-88-470-1974-4_1, # Springer-Verlag Italia 2012
2 A. Fasolo
This is even more relevant, if we step from biological sciences to humanities.

Accordingly, the goal of the meeting was to achieve a broad analysis of the impact,
pinpointing on a few specific, but paradigmatic topics. Even the place was well
tempered, since Torino was in Italy one the main diffusion spot for Darwinian
thought and work, both from the academic and editorial point of view.
The present book collects essentially contributions (except for Rossi and
Pievanis ones) from the meeting, mixing styles, arguments, subjects. This kind
of inter-disciplinary approach may appear erratic, but it conveys flashes of light on
the changing scenarios where the theory of evolution is moving. It is on line with
the idea to reopen the file of the Two Cultures, looking at shared problems, which
are not really the Third Culture invoked by Charles Percy Snow half a century ago,
but they can foster it, at least in such a pivotal domain as evolution.
1 Roots and Buds of Evolutionary Theory
In history of science, for instance, notwithstanding a few crucial contributions, the

intellectual credits of pre-darwinian authors remain rather bad known. The
almost total lack of interest for the state of affairs in the publishing industry of
the period under consideration, and the total lack of interest for what books,
dictionaries, encyclopaedias actually said, made us blind to major debates of
great significance for the history of the life sciences at European level during
the early decades of the nineteenth century. Thus the reconstruction of the ways in
which Lamarck was read, admired, criticized or denounced might be considered a
mile stone of the modern reappraisal of history of evolutionary thought. Thus
Pietro Corsi is crunching the cultural background before and around Darwin,
focussing on the set of easy assumptions concerning the place and reputation of
Lamarck within the French natural history community of the early decades of the
nineteenth century. Such visions acted as true Idola tribus, preventing research
and limiting in considerable ways our understanding of the complex intellectual,
social and political dynamics of contemporary natural history practices and
publishing. In Pietro Corsis views, such absence or paucity of interest for made
us blind to major debates of great significance for the history of the life sciences at
European level during the early decades of the nineteenth century. Accordingly
the ways in which Lamarck was read, admired, criticized or denounced might be
considered a mile stone of the modern reappraisal of history of evolutionary
thought.
Even for philosophy it is not true that les jeux sont faits. Wittgenstein famously
remarked in [16], Darwins theory has no more relevance for philosophy than any
other hypothesis in natural science. Yet today we are witnessing a major revival of
interest in applying evolutionary approaches to philosophical problems, as Michael
Ruse accomplished recently with the Philosophy after Darwin [13], an anthology of
essential writings covering the most influential ideas about the philosophical
implications of Darwinism, from the publication of On the Origin of Species to
Introduction: The Sand Walk (on the Darwins Steps) 3
todays cutting-edge research. Along this same red line, Ruse argues that work
being done today on evolution and philosophy as part of a broader cultural
movement. In some very deep sense, it is part of a movement to see human beings
in a naturalistic fashion, this being set against more traditional attempts to locate
humans in a religious, a spiritual, a non-naturalistic world. One aim, as you might
already have guessed, will be to show that the story is not quite as straightforward as
one might have expected.
Always on philosophy side, Paolo Casini notes that when John Dewey, in his
essay The Influence of Darwinism on Philosophy (1909), remarked The exact
bearings upon philosophy of the new logical outlook are, of course, as yet, uncertain
and inchoate. We live in the twilight of intellectual transition. Nowadays four
decades of controversy concerning evolution had elapsed, and Darwins Darwinism
was eventually accepted, The transition towards evolutionary logic, according to
Deweys subtle analysis, expelled from biology, and from philosophy as well, all
ideal archetypes, the concepts of design and finality, and destroyed the philosophic
idol of eido (o species).
If we challenge the historical roots of evolutionary theory (as the) with its
present day bearings, what remains of emotional ideas like, The Nature, red in
tooth and claw? Peter Weingart notes that the metaphor struggle for existence
takes its origins in everyday language but it was given a specific meaning in the
context of evolutionary theory. Subsequently, the metaphor was transferred back
into everyday use but had also a tremendous impact on the historical and social
sciences. Darwins metaphor is one of the most famous cases of this type of
metaphor transfer into the sciences and back. Accordingly the usages of the
metaphor appear really wide and loose, but nonetheless they had their time.
A search for occurrence of such a phrase in titles and/or abstracts of documents
in both the SSCI and SCI databases revealed only 21 entries for the period
19731999. Evidently it is justified to say that the struggle for existence as a
metaphor has not survived the struggle for use and attention.
2 The Mankind Affair
Mankind evolutionary history can be tackled in several ways, employing tools from
disparate disciplinary fields as cultural anthropology, linguistics, to-date molecular
genetics.
In a fascinating approach, Henrika Kuklick explores the dialectics and the
somewhat contradictory exchanges between Darwinian theory and the new born
social anthropology:
Anthropological fieldwork framed by a Darwinian biographical approach proved extremely
important in changing the discipline, (perhaps paradoxically) leading to a thorough separa-
tion of cultural from biological anthropology. . . . Not until the 1980s would evolutionary
approaches to the analysis of culture that were advertised as authentically Darwinian seem
respectable to more than a distinct minority of socio-cultural anthropologists, but that is a
4 A. Fasolo
development that may be most significant as evidence of the rise of conceptual pluralism in
anthropology.
A key-corner between biology and society is the language. Manfred Bierwisch

draws an elegant and challenging reflection on such a conceptual and experimental
labyrinth, where we ignore how many (or if any) are the exits. At a first glance, we
are relying on some necessary analogies. Human language history can indeed be
logically explained by an evolutionary theory, but its principles are essentially
different from those that govern the development of biological species. Then the
question is as to whether we have identified a principle of evolution that is
universally applicable to the historical development of language and more broadly
to sociocultural structures. Favoring the supposition of a fundamental role assigned
to language as a basis and ingredient of veritably every sociocultural institution
enabling the capacity for unlimited expression, bounding language symbols to
agreed convention, Bierwisch notes that there are nonetheless grounds for reserva-
tion stemming from two considerations, the domain-specificity of the language
faculty and the intentionality of social behavior, including the creativity of lan-
guage use. The compelling close is that The faculty of language is the prerequisite
of human history, but it does not determine its course.
The theme of the evolution of the language immediately calls us to the extraor-
dinary researches on the genetics of ancient human populations, where pioneering,
monumental studies were performed by Luca Luigi Cavalli Sforza, Paolo Menozzi
and Alberto Piazza. In the present book, Alberto Piazza is arguing on the role of
natural selection, a major factor in Darwinian evolution which is elusive and
difficult to dissect, especially when the case of human evolution is dealt with. In
August 1858, Charles Robert Darwin and Alfred Russel Wallace presenting to the
Linnean Society of London their independent discovery of the theory of natural
selection, suddenly and altered our understanding of life on Earth. He is focusing
his attention on five major advances of genetics on the analysis of human evolution,
and especially on the comparisons between human and chimpanzee genomes and
on the very recently published DNA draft sequence of the Neanderthal genome.
The very questions for modern humans are:
To which extent has natural selection influenced, at the scale of the entire
genome, the degree of population differentiation?
Which type of genetic variants have been preferentially targeted by selection?
Genes and gene variants under strong selective pressures can highlight regions
of the genome explaining the current population phenotypic variation?
The final challenge is methodological: how can we evaluate the relevance of the
sexual selection in humans, starting from the many conjectures and working
hypotheses put forward by Darwin, which are very plausible for animals, but very
difficult to test for humans, especially in modern times when cultural factors on
sexual selection may completely shadow biological pressures.
3 Development, a Persistent Problem of Evolutionary Theory
If the powerful genetics is sitting in the core of modern evolutionary theory, a blow
of new ideas comes from its theoretical belt, as Evolutionary Developmental
Biology (Evo-Devo). Thus Giuseppina Barsacchi analyzes the relationships
between the processes of individual development and the phenotypic changes of
the organism during evolution. Methodological advances such as gene cloning, gene
expression screening and visualization of gene activity in embryonic tissues
facilitated the emergence of a major theme of the current Evo-Devo research, the
evolutionary developmental genetics program. Its foundational achievement was the
discovery of extensive similarities in developmental regulatory genes and gene
networks among distantly related species. The program concentrates on the evolu-
tion of genetic tool-kits and signaling pathways and on the regulatory logic that
underlies organism development. Mapping the expression pattern of gene networks
and signaling pathways and analyzing their correlation with the constructional
features of body architecture, provides information on their possible role in pheno-
typic evolution. Major morphological transitions in evolution are presently
recognized to be accommodated by a few key developmental genetic changes
(part of a developmental reprogramming) and case studies in snakes, ducks,
bats, dolphins, insects, and finches, providing valuable insights into principles of
evolutionary change, are presented. On the other hand, the molecular changes are
rooted in an otherwise conserved developmental genetics tool-kit (e.g., the Hox
genes for anterior-posterior patterning, the network for eye formation etc.) that
substantiates the deep homology underlying diversity of forms. On this ground,
the relationship of the deep homology of genes working through development with
classic morphological homology is in the Evo-Devo field of exploration. How
environmental agents can instruct changes in development, for example altering
gene expression in broad sense searching for a link between proximate causes of
development and natural selection-, falls also in the perspectives of newly growing
and exciting knowledge, where Evo-Devo integrates with Ecology. The problems
are many and very interwoven, as Alessandro Minelli remarks: a real Evo-Devo
biology is now growing in extent and importance, but integration between the two
disciplinary components is still basically fought on the battlefield, case by case ([9],
p.118). The case is for instance the principle of developmental inertia, raised by
Minelli himself, like the arguments about regeneration, developmental pathways,
epigenetics, multiplicity of centers of local development dynamics as opposed to
global control, and so on. . . Summing up, future work may further give reason for
the Charles Darwins appraisal of the importance of Embryology for Evolution.
4 Brain Evolution and Plasticity
Overcoming the traditional dichotomy opposing neural selectionism to construc-

tivism is the goal of Ferdinando Rossi. In an extreme synthesis, Rossi is arguing in
a syncretistic fashion, along the following lines of reasoning. The correct function
6 A. Fasolo
of the nervous system requires complex neural networks bearing precise

connections. In principle, the high structural specificity of neural circuits could
be achieved by genetically-determined processes, selected and refined during
evolution. Highly conserved gene networks regulate some crucial steps of neural
development, such as the regionalization of the neural tube and the initial phases
of neurogenesis and synaptogenesis. A totally hardwired nervous system may
meet the requirements of adaptation and natural selection at the population level,
whereas it would be fully inadequate to allow individual organisms to cope with
rapid changes of environmental conditions. Neural adaptation to external
constraints can be partly achieved by introducing selective mechanisms in neural
development. Accordingly, neurons are generated in excess and then partially
eliminated to match the actual extension of innervation territories. Such
mechanisms, however, are restricted to a set of potentialities, which must be
predetermined in the ontogenetic program. On the other hand, constructive
mechanisms, in which external stimuli directly influence structural modifications
of neural circuits to produce adaptive responses, may allow individual organisms
to cope with a wide variety of unprecedented situations. Thus, in the last ontoge-
netic period as well as in the adult, when the organism actively interacts with the
external milieu, experience exerts a strong growth-promoting effect on neural
circuits and connections inducing the emergence of specific functional properties.
By this mechanism, which requires strict inhibitory control to prevent aberrant
growth and dysfunction, the nervous system exploits external stimuli to create
adaptive responses to unexpected situations.
Such syncretism represents a good way to handle the enormous wealth of data on
brain development recently acquired. Nevertheless this approach raises some
reflections on tricky concepts such as evolvability [7,11] and exploratory properties
in complex systems, namely in neural tissues.
Evolvability is an organisms capacity to generate heritable phenotypic varia-
tion. Metazoan evolution is marked by great morphological and physiological
diversification, although the core genetic, cell biological, and developmental pro-
cesses are largely conserved. Metazoan diversification has entailed the evolution of
various regulatory processes controlling the time, place, and conditions of use of the
conserved core processes. These regulatory processes, and certain of the core
processes, have special properties relevant to evolutionary change, reducing the
interdependence of components and conferring robustness and flexibility on pro-
cesses during embryonic development and in adult physiology.
Even more ambitiously, we can ask: how our brain evolved? In a masterly way,
Gerhard Roth shows that the human brain is not unique in terms of general
structure, since it exhibits the basic pattern typical of mammals and more specific
of primates. In addition, humans do not have the largest brain either in absolute or in
relative terms, although they possess a brain that is seven to eight times larger than
expected from general mammalian brain allometry (defined as the study of the
change in proportion of various parts of an organism as a consequence of growth).
Through an elegant analysis of many other quantitative data, Roth concludes that
the greatest differences between humans and all other mammals/consist in (1) a
strongly increased growth period of the human brain exposing it to a much higher
degree to education, and (2) the presence of the Broca speech area which is a
necessary prerequisite of syntactical language. While these two traits appear to be
minor steps in human biological evolution, they had enormous consequences for
human culture.
5 Old/New Concepts
One major methodological and pragmatic problem is an old and persistent one:
what is the meaning of similarities, in genetics as in anatomy or in developmental
processes? Three old/new friends may help to understand the nature of similarities
and their bias.
Among new or renewed conceptual tools, one emerging clue is homoplasy [14].
Homoplasy is the independent acquisition of the same trait in unrelated lineages.
Parallelism/convergence homoplasy occurs when the same trait is present in two
lineages that lack a recent common ancestor. Reversal homoplasy occurs when a
trait is present in an ancestor but not its immediate descendants; but appears later in
a subsequent descendant. Understanding the diversification of phenotypes through
time has been the focus of evolutionary biology for 150 years. If, contrary to
expectations, similarity evolves in unrelated taxa, researchers are guided to uncover
the genetic and developmental mechanisms responsible. Similar phenotypes may
be retained from common ancestry (homology), but a phylogenetic context may
instead reveal that they are independently derived, due to convergence or parallel
evolution, or less likely, that they experienced reversal. Such examples of homo-
plasy present opportunities to discover the foundations of morphological traits.
A common underlying mechanism may exist, and components may have been
redeployed in a way that produces the same phenotype. New, robust phylogenetic
hypotheses and molecular, genomic, and developmental techniques enable
integrated exploration of the mechanisms by which similarity arises.
On the other hand, the trendy interest in development can effectively enrich our
definitions of homology and our methods to individuate it. The study of develop-
mental processes calls for a comparison at different developmental stages,
overcoming the restriction to adults, which has been the focus in classical compar-
ative studies [4]. Too often, comparative neurobiologists have considered brain
evolution as the transformations of adult brains over time A more extensive interest
in dynamic processes can help unveiling the plastic changes of the brain throughout
life. To give a simple example, the developing human brain seems to be different at
the functional neuroanatomy level from the adult brain, even in processing single
words. Another puzzling problem is the genesis of novelty and its adaptive value.
Interestingly enough, very recent molecular investigation on primates suggest that
the human brain has probably experienced pronounced evolutionary changes in
gene expression during its most recent history [3] and that the evolution of human
cognitive abilities was accompanied by adaptive changes in brain metabolism [6].
8 A. Fasolo
These results are open to different theoretical hypotheses and should not over-
interpreted, but they suggest that processes of fast genetic reorganization might
sometimes occur.
In the light on these considerations, a main question presents itself: what are the
adaptive pressures behind brain and behavior novelties in evolution? We have no
answer yet, but we can agree with the original statement by Williams, in his [15]
Adaptation and Natural Selection, frequently quoted in evolutionary psychology,
but not so frequently exploited: Evolutionary adaptation is a special and onerous
concept that should not be used unnecessarily, and an effect should not be called a
function unless it is clearly produced by design and not by chance. When
recognized, adaptation should be attributed to no higher a level of organization
than is demanded by the evidence. A new emphasis on homology in evolutionary
biology (the persistence of theoretical problems notwithstanding), may offer new
powerful tools for an effective comparative analysis, and may thus help
distinguishing between strict biological correspondence and loose metaphoric
representations of behavior, which are the mere result of an uncritical assumption
of an evolutionary stance. Especially in cases of highly complex behavior, ethics
being a paradigmatic example, biology and culture are certainly tightly entrenched:
the claim that these kinds of behavior have evolutionary bases is simply a truism.
The interesting point would be the possibility to identify the characters, if any,
which show continuity and can be challenged by a homological analysis. The
evolution of the brain involved a complex set of relationships among individual
structures, both at the quantitative and the qualitative level. As aforementioned,
there is some controversy concerning this idea, but the core problem (e.g., whether
changes are directly selected or not) remains unsolved. It seems plausible, however,
that some processes are related to environmental pressures, while others emerge in
response to the need for more flexible answers, and still others are part of a less
specific and foreseeable ecological niche. Likewise, brain structures have devel-
oped along several lines, and one usually finds a mosaic-like pattern even within a
particular line.
In such a mosaic of integrated parts, whatever the evolutional process might
have been, at least a part of the variation has not been selected per se, but it
represents a collection of exaptations [5]. For instance, the molecular evolution of
ASPM gene in hominoids may indeed be an example of a molecular exaptation, in
that the originally selected function of ASPM was for something other than large
brain size, since the ASPM gene sequence shows accelerated evolution in the
African hominoid clade, and this precedes hominid brain expansion by several
million years [8]. The idea that novelty may arise from and exaptation (functional
cooptation in Darwin, then pre-adaptation in Ernst Mayr) can have strong
impact on our views. Three typologies of processes, i.e. classical Darwinian
adaptations by natural selection; the functional shift, by natural selection, from a
previous function to a secondary one; spandrels and other side effects with no
adaptive reasons in their beginning, possibly co-opted by natural selection in new
external conditions can extend the taxonomy of fitness [5,10].
6 Cognition and Reasoning
Moving from comparative neuroanatomy to modern cognitive neuroscience,

Giorgio Vallortigara explores recent research updating Darwins implicit sugges-
tion that there maybe primitive neural pathways that ensure a bias toward sensory
cues about other living things, in particular members of the same species. There has
been of course a long road from the primitive animacy detectors that we can see
operating even in simple brains to the intricacies of agency attribution and theory of
mind of human beings. Nonetheless, the origins of beliefs in supernatural things and
of our intuitive dualism seem to be deeply rooted in natural history.
Thus reason does not have to keep repeating why it holds itself to be so
important if it can see how it became necessary and under what conditions it is in
fact indispensable. Volker Gerhardt believes that evolutionary theory can liberate
reason from the burden of its thousands of years of self-confirmation and lead it
back to the conditions that preceded it that are themselves not yet rational. It might
not exist any other problem that the natural sciences and the humanities should take
a greater interest in. For it is in the natural elucidation of the origin and the potential
achievements of reason and consciousness that both fields have the chance to shed
light on themselves as well and to clarify why they not only emerged from the same
impulses of curiosity, knowledge, and rational guidance, but continue to depend on
one another.
Finally, Pievani reflects on the current status of theory of evolution: how it
changes and grows, remakes itself keeping alive and reinforcing its Darwinian
explanatory core. The starting point is the awareness that the capacity of assimi-
lation of scientific novelties by Modern Synthesis (MS) seems to be progressively
declining. The problem is seemingly no longer its incompleteness, but the
adequacy of the whole conceptual structure of the theory [5]. Using Imre Lakatos
methodology, Pievani argues that the transition in progress from the MS to the so
called Evolutionary Extended Synthesis (ES) [12] could be represented as a
shift from a previous evolutionary research, and a new evolutionary research
program, with an extended Neo-Darwinian core and a protective belt of new
assumptions and auxiliary hypotheses with a pluralistic and integrative explana-
tory approach. Promising and advanced researches like those concerning evolu-
tionary developmental biology (Evo-Devo), epigenetics, multiple ways of
speciation and the role of structural internal constraints find in this perspective
a realistic interpretation as theoretical and empirical novelties with huge
implications, nevertheless not incoherent with an extended Neo-Darwinian
explanatory core. Such approach seems also useful discussing the extension of
evolutionary models in non biological fields, in order to avoid just metaphorical
forms of ultra-Darwinism.
Summing up, the debate on evolution is still open and strives us to exchange
and change ideas. What may be our philosophical and scientific endeavour, we
can agree with Michael Ruse, expecting that in fifty years or a hundred years we
will still have the theory of the Origin around. Great, precisely because it does not
10 A. Fasolo
stand still, but remakes itself and grows and changes by virtue of the fact that it
gives such a terrific foundation. Is Darwinism past its sell-by date? Not by a long
chalk yet!
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Idola Tribus: Lamarck, Politics and Religion
in the Early Nineteenth Century
Pietro Corsi
Abstract There is no doubt that traditionally the history of evolutionary ideas has
been and is Darwin-centred. I have no dispute with this, being a convinced
Darwinian, in spite of years of work I have devoted to study Lamarck and the
many non-Darwinian theories of evolution current in Europe and the United
States before and after 1859. Whereas historians have paid some attention to
post-Darwinian, non Darwinian theories, pre-Darwinian theories have been much
neglected. Attention is usually paid to so-called Lamarckian attitudes present in
European natural history debates from the early 1800s to the 1850s, only to
conclude that Lamarck played no role, was almost unanimously neglected and in
any case unanimously vituperated. This was hardly the case. However, the aim of
my paper is not to vindicate Lamarck, but to argue that even concentration on
Lamarck would amount to gross anachronism. After analysing reasons essentially
political and religious that have been given to explain the alleged oblivion into
which Lamarcks works had fallen (if they ever rose to attention) I will examine
evidence concerning the wider debate on Lamarcks ideas within the medical
literature of the 1810s and the 1820s. This will open up a new research area,
focussed on the translation into French of major German authors (Meckel,
Tiedemann, Carus, Treviranus, Burdach, Oken) and on the attempts to re-formulate
key Lamarckian tenets in the terms of German natural philosophy, comparative
anatomy and embryology, and medicine. The debate on the development of life
historical and embryological was wider and much more interesting than the
debate on Lamarcks own theories, which in any case well deserves to be rescued
from oblivion.
P. Corsi (*)
The Old Boys High School, University of Oxford, Oxford, UK
e-mail: p_corsi@yahoo.com

12 P. Corsi
All differences taken into account, Lamarck and Darwin shared the common
destiny of being often identified with doctrines they never upheld, or not exactly
in the form history has attributed to them. Over the last century and one half, wave
after wave of the recurrent debate on Lamarckism vs. Darwinism, and the
repeated rituals of centenary and other anniversary celebrations have done much to
obscure the real contribution of the two naturalists to the debates on evolution. In
saying so, I am of course taking for granted several assumptions, some of which
will be spelled out and discussed in the following pages. A major assumption which
will not be critically scrutinized, and is presented here as a comment at the end of
one year of world-wide celebrations, is that in-depth and easily available historical
studies on Lamarck and Darwin have been rarely read or consulted by a good
number of commentators who during 2009 have been very active explaining who
Darwin really was. Nor have they been consulted by the much lower number of
those who remembered that 2009 marked not only the bicentenary of the birth of
Charles Darwin, but also of the publication of the Philosophie zoologique, one
of Lamarcks key evolutionary texts. The impression one gets, after reading,
viewing, or listening to a statistically relevant portion of what has been said on
Darwin during 2009 through several continents and languages, is that his works, as
those of Lamarck, are not that well known, and that the work of professional
historians who have engaged the primary sources is scarcely taken into account.
This is not a novelty, after all: since the early 1800s much of the debate on what we
call today evolutionary doctrines was carried on without much attention to the
actual articulations of the scientific arguments under discussion.
Before stating and developing the key themes of my paper, let me provide only
one example of what I peremptorily stated above. There is no doubt that the
doctrine of the inheritance of acquired characteristics is universally regarded as
the cornerstone of Lamarcks theory and the major point of difference with Darwin
and Darwinism. Yet, as Jean-Gayon has persuasively argued, and a rapid search by
word of the Lamarckian corpus available on line will confirm, Lamarck never spoke
of the theory of the inheritance of acquired characteristics.1 He most surely believed
that new needs originate new behaviours, and new behaviours increase or decrease
the size and functions of the solicited organs, to the point that new species and
genera are formed. Life is thus constantly transformed, since the process is cumu-
lative through inheritance. This was a conviction he shared with many authors
active at the end of the eighteenth and at the beginning of the nineteenth Centuries,
to the point that early critics of Lamarck rarely complimented or reproached him for
this. The key issue when discussing Lamarck was always whether the process of
change he had described was sufficient to overcome the species or the genus barrier
a point some were ready to concede as well as higher divides (family, order)
which very few granted. Fifty years later, the same reaction characterized the early
1
Jean Gayon, Lamarck Philosophe, in P. Corsi et al., Lamarck philosophe de la nature, Paris,
PUF, 2006, pp. 935. See P. Corsi, http://www.lamarck.cnrs.fr/ for the complete edition of
Lamarcks theoretical works, his manuscripts and herbarium.
Idola Tribus: Lamarck, Politics and Religion in the Early Nineteenth Century 13
(and later) debates on natural selection: many saw it as a plausible mechanism to

explain the fixation of varieties, which could in no way put in doubt the constancy
of species, or of genera.
The interesting question that emerges from carefully comparing the relevant
Lamarckian and Darwinian texts, is that the capability of organisms to change and
to pass on to the next generation whatever was gained or lost during their lifetime
was severely limited in the case of Lamarck, and less so with Darwin. For Lamarck,
only very young organisms, in which the tissues were still very soft, and the
circulation of blood, nymph, and the nervous and other fluids was particularly
brisk, showed a potential for adaptive change: never the adults. This was not
Darwins opinion. When presenting the ill-fated and little studied theory of pan-
genesis, among other phenomena of heredity Darwin sought to explain how a
change that occurred at a given point in the life of one organism tended to appear
again at the same stage of individual development in his progeny. Furthermore,
whereas Lamarck simply took up a widely shared, almost commonsensical belief
that the characteristics of the parents were passed on to the next generation, Darwin
spent time and ink to understand how this was possible, even discussing similar
theories put forward by authors such as Georges-Louis Leclerc, Comte de Buffon
(17071788) and Charles Bonnet (17201793), on whom Huxley had called his
attention.2 Lamarck spent much less time on the matter: he simply argued that since
the male seminal fluid (akin to electricity and magnetism) acquired specific
peculiarities within each type of organism, it was legitimate to infer that the same
fluid would take up slightly different anatomical and functional properties by
circulating through an organism that had undergone a very slight change during
the early phases of its life. Indeed, for Lamarck, fully blown characteristics were the
end result of a cumulative process of very minor changes within the fluid dynamics
internal to all and every organism. Thus, if a new need was requiring a more
pronounced use of a given organ, thereby increasing the flow of blood, nutritional
and nervous fluids to that part, what was passed on to the next generation (provided
the young individuals that had gone through the same process reproduced together
when adults), was not a character that as yet did not exist, but the slight change in
the pattern of the fluid dynamics and the slightly modified features of the seminal
fluid. On the contrary, Darwin admitted the inheritability of changes occurring in a
single parent, and asked himself how these could be maintained through successive
generations. This is not to conclude that Darwin was more Lamarckian than
Lamarck, but to insist on the fact that the mere reading of the works of Darwin
and Lamarck would prevent all hasty and easy generalizations.
Even though rarely read by those who should, the scholarship on Darwin of the
last 40 years has been on the whole excellent and has powerfully contributed to a
2
See C. Darwin, The Variation of Animals and Plants under Domestication, 2 vols., London, J.
Murray, 1868, vol. 2, Ch. XXVII, Provisional hypothesis of pangenesis, pp. 357404.
14 P. Corsi
less anachronistic appreciation of the man, his career and doctrines.3 The case of
Lamarck, on which I will devote the main bulk of my paper, is to some extent quite
different. There are of course excellent studies of his work and career, though
the writings of the French naturalist have not been translated or edited with the
same alacrity and systematic dedication.4 There is no correspondence left worth
mentioning, no notebooks, no autobiographies or diaries. Much of the Lamarckian
manuscripts are in fact drafts or final versions of printed works. Lamarck was very
parsimonious with information about himself, his life and thoughts, to the point that
much of the scarce biographical hints we have are due to members of his family, his
enemy Cuvier or to a young medical practitioner who interviewed him in the early
1820s.5 Moreover, whereas over the last 20 years or so important scholarship has
appeared offering insights into the wider natural history scene (institutional, intel-
lectual and social) of the United Kingdom, the same cannot be said of France during
the times in which Lamarck was active. In other words, the scholarship on Lamarck
has not incited new studies on the wider scientific and institutional context
characterizing the life sciences during the early decades of the nineteenth century
in France.6
The rare albeit excellent exceptions to the rule have not helped us to gain a less
anachronistic view of contemporary priorities, actors and debates. The set of
traditional assumptions concerning the context of Lamarcks work remain
stubbornly unchanged, in spite of growing evidence that should advice historians
to enlarge the scope of their research. It is a few of these implicit, often untold
assumptions I wish to tackle in the following pages. Basically, they turn around a
major conviction, the total or almost total isolation Lamarck lived in. This assump-
tion generates in its turn a host of further assumptions if not prejudices asked to
perform a causal role in the narrative. They can be ranged, historically and
thematically, from the (usually French) patriotic and whiggish explanation that
Lamarck was born too early, or that he was seeing too far, to the less charitable
(usually Anglo-American, pro-Darwinian) view that he was simply wrong,
overwhelmed by top brass of science such as Georges Cuvier (17691832), Pierre-
Simon Laplace (17491827), or Antoine Lavoisier (17431794) and his pupils.
3
I will only refer here to the biographies by A. Desmond and J. Moore, Darwin, London, Michael
Joseph, 1991 and Darwin, Oxford, Oxford University Press, 2007; and J. Browne, Charles
Darwin, 2 vols., London, Jonathan Cape, 19952002.
4
See for instance R. Burkhardt, The spirit of system: Lamarck and evolutionary biology,
Cambridge, MA, Harvard University Press, 1977, 1995.
5
Georges Cuvier,. Eloge de M. de Lamarck, lu a lAcademie royale des sciences le 26 Novembre
1832, in Memoires de lAcademie royale des sciences de lInstitut de France, 13 (18311833),
pp. ixxx; Isidore Bourdon, Lamarck, Dictionnaire de la conversation et de la lecture, 34
(1837), pp. 265269.
6
See for instance J. Secord, Victorian sensation: the extraordinary publication, reception, and
secret authorship of Vestiges of the Natural History of Creation, Chicago and London, University
of Chicago Press, 2000, and J. Endersby, Imperial nature: Joseph Hooker and the practices of
Victorian science, Chicago and London, University of Chicago Press, 2008.
He had no chance to be listened to in a world moving towards disciplinary

specialization and epistemological rigour. A second assumption concerns the inev-
itability of Lamarcks isolation in the increasingly conservative political climate of
the Consulate and the Empire, and in the ultra-conservative intellectual atmosphere
of the restored monarchy. His materialistic biology and transformist doctrines (it is
claimed) were unacceptable to authorities determined to curb any form of political
and intellectual subversion. Finally, the third assumption we are going to examine
below is the one concerning the audience of Lamarcks works. Followers of various
versions of assumptions one and two will find this third point completely superflu-
ous. To them, Lamarck had no audience worth talking about, at least until the
1820s, and even then the few who paid any attention to him did not, in fact, support
his views as the old naturalist would have wished. In France as well as in Europe,
Lamarcks materialism found sympathetic hearing only within the radical fringes,
thereby adding to the already long list of reasons people had to dismiss him
outright.7
The way in which the assumptions we have sketched above have been argued by
historians does not lack plausibility and evidential support. Yet, consensus has been
gained at the price of restricting the research horizon to the point of neglecting
major features of natural history practices and debates of the early nineteenth
century, in France as well as elsewhere in Europe.
1 Lamarck Versus Institutional Science
Very few historians of early nineteenth century life sciences appear to doubt that the
major educational and institutional reforms introduced by successive revolutionary
governments, the Directory, the Consulate and the Empire deeply changed the
social and intellectual practices of research within the complex articulation of
disciplines still constituting the histoire naturelle. To a significant extent, they
are absolutely right. In 1792, 1793 and 1799, two naturalists occupying the opposite
sides of the epistemological spectrum in the debate over natural history agreed that
France was not doing much, after all. Jean-Claude de la Metherie (since 1793
simply Delametherie) and the then still little known Cuvier insisted that Germany
was better equipped than France in several sub-domains of natural history. Cuvier
pointed out that almost every German university town was publishing its own
scientific or medical journal and hosted important private and public collections.
In France, almost everything was concentrated in Paris, and Parisian naturalists
were too happy to sit on the top of their monopolistic privileges to care about
sharing their knowledge with colleagues in the provinces and abroad. As a
7
The best known and best argued representative of the view that Lamarck was acceptable only to
extreme radicals is Adrian Desmond, The Politics of Evolution. Morphology, Medicine and
Reform in Radical London, Chicago and London, University of Chicago Press, 1989.
16 P. Corsi
consequence, France was rather poor in periodical publications, since only a

handful had survived the revolutionary years, and there were not that many even
before 1789, for that matter.8
In the space of a few years the situation changed dramatically and unpredictably.
The Revolutionary armies engaged in the systematic plundering (which they called
confiscation) of conquered lands, to finance the huge state deficit and the costs of
the war. Cash, precious minerals, paintings and sculptures, natural history and
scientific instrument collections took the road to Paris in hundreds of over-charged
wagons. The Museum national dhistoire naturelle, established in June 1793, was
the ideal place were natural history collections could be hosted, catalogued and
studied on behalf of the Republic of knowledge, which did not know of frontiers or
wars. Confiscations were undertaken with a higher view in mind, the benefit of
mankind, French authorities insisted.9 By 1802, Paris hosted the largest and richest
natural history collections ever assembled in Europe. Naturalists from all over the
Continent had to pay frequent visits to the French capital: some, undoubtedly, to
pay due homage to the new rulers; others, because they had to keep up with their
own work. The local German collections Cuvier had extolled in 1799 were no more
sufficient to guarantee cutting edge research.
French scientific publishing also benefited from the new impulse successive
governments accorded to the practice of science.10 The at times purely symbolic
8
Georges Cuvier, Extrait dune Notice biographique sur Bruguiere, lue a la societe
philomathique, dans sa seance generale du 30 nivose an VII, in Magasin encyclopedique, 5th
year, vol. 3 (1799), pp. 4257; Louis Marchant, Lettres inedites de Georges Cuvier a C. H. Pfaff
sur lhistoire naturelle, la politique et la litterature, Paris, Victor Masson, 1858, p. 78: Les
sciences ont aujourdhui peu de dignes pretres en France, et cette pauvrete est dautant plus
penible, que lon se souvient encore de lancien eclat dont elles ont brille; Jean-Claude
Delametherie, Discours preliminaire, in Journal de physique, 42 (1793), p. 7. See also A.-L.
Millin, Journal dhistoire naturelle, in Magazin encyclopedique, 1, n. 8 (8 decembre 1792), pp.
5760, LAllemagne voit parotre un grand nombre de collections, et de recueils dhistoire
naturelle, p. 57.
9
For two recent systematic studies of the accumulation of collections in Paris see B. Daugeron,
Apparition-Disparition des Nouveaux mondes en Histoire naturelle, Enregistrement-Epuisement
des collections scientifiques (17631830), Paris, EHESS, These de doctorat, 2007, 2 vols.and P.-Y.
Lacour, La Republique naturaliste. Les collections francaises dhistoire naturelle sous la Revolu-
tion, 17891804, Florence, European University Institute, Ph. D. Dissertation, 2010, 2 vols. It is
interesting to point out that in his biographical notice of Bruguiere (see n. 8) Cuvier complained
that the collections amassed at great public expense were now collecting dust at the Museum, since
no one appeared to work on them. This too was soon to change, but systematic exploitation of the
conquered natural history riches only started after 1802, that is, after Cuvier became full professor
there. The political innuendos of Cuviers astonishing biography of Bruguiere, his equally
astonishing veiled attacks against colleagues working at major State institutions have never been
analyzed in detail.
10
Amongst many, the testimony of Louis Marchant, Lettres inedites de Georges Cuvier, is telling,
p. 30: Cetait une epoque tres-favorable pour les sciences et ceux qui les cultivaient; le premier
consul se trouvait tres-honore du titre de membre de lInstitut, il le mettait en tete de tous les autres.
Les premiers hommes de la science, comme Laplace, Chaptal, Monge, etaient en meme temps les
encouragement lasted until the end of the 1790s, followed by real investments
especially as far as the Museum dhistoire naturelle was concerned, during the early
1800s. Yet, already in the second half of the 1790s the rhetorical and ideological
support for science-oriented activities was sufficient to massively increase the market
for natural-history publications: collective works of Buffon, dictionaries and
encyclopaedias, textbooks and innovative surveys of new or renewed domains of
research (among which Cuviers Lecons danatomie comparee), manuals for the new
school systems found eager buyers and were exported throughout Europe and the
Americas.11 Unfortunately, this feature of the French publishing market has been
little studied and even less appreciated. Furthermore, no attention has been paid to the
dense population of naturalists surviving thanks to their pen. Historians have tended
to ignore a highly articulated editorial, epistemological and research scene and have
instead concentrated their attention on a handful of individuals and institutions.
As it is often the case within the history of science, our contemporary concept of
proper scientific practice deeply influences our reading of the past. In spite of the
pioneer work of Dorinda Outram, published in 1984, historians insist (at times very
ably indeed) that the natural history scene was dominated by Georges Cuvier, who
exercised an undisputed leadership, set the standards, and marginalized whoever
did not conform to his anti-speculative, matter of fact approach to natural knowl-
edge.12 The Museum national dhistoire naturelle and the Institut de France were
the strongholds from where Cuvier, the so-called Napoleon of intelligence,
manoeuvred his troops and kept epistemological order. Outram richly documented
the difficulties Cuvier experienced, the set-backs he had constantly to suffer, and
pointed out that many of the institutional gains the naturalist scored were often the
consequence of his political shrewdness and power rather than the indication of his
undisputed scientific authority. When in 1818 a distant relative and admirer of
Cuvier, Jean-Jacques Coulmann (17961870), visited London, he was amazed to
notice that the anatomist was better known in the English capital than in Paris.13 As
I have myself documented, Cuviers leadership was constantly challenged, in some
premiers aux affaires. Linstitution grandiose du Jardin des plantes, a laquelle des savants speciaux
dune grande celebrite etaient attaches pour chaque branche de lhistoire naturelle, pour la
geognosie, la geologie, pour la chimie theorique et pratique, a laquelle se reliaient les grands
musees nationaux, avait surtout une grande part dans cette sollicitude et ces encouragements.
11
For further contemporary testimonials, see, among others, F. W. Blagdon, Paris as it was and as
it is, London, C. and R. Baldwin, 1803, vol. 2, p. 582, and passim. See also P. Corsi, After the
Revolution: Scientific Language and French Politics, 17951802, in M. Pelling and
S. Mandelbrote, eds., The Practice of Reform in Health, Medicine, and Science, 15002000,
Aldershot, Ashgate, 2005, pp. 223245
12
D. Outram, Georges Cuvier: Vocation, Science and Authority in Post-Revolutionary France,
Manchester, Manchester University Press, 1984.
13
J -J Coulmann, Reminiscences, Paris, Michel Levy freres, 18621869, vol. 1, p. 233: Coulmann
was the younger brother of the wife of General Frederic-Louis-Henri Walther (17611813),
Cuviers first cousin. Outram, 1984, has called attention to the role Walther played in introducing
Cuvier to powerful figures in Paris in early 1795, when Walther was already a significant figure
within the army.
18 P. Corsi
cases openly derided, and the publications directed to the general public he spon-
sored failed miserably (his famous and very successful specialized works were too
expensive to reach the ordinary reader), to the joy of his opponents. Cuvier did try
his best to convince contemporaries that he was the depositary of true scientific
values and achievements. His eloges of deceased academicians, his reports to the
Emperor on the progress of science in France and Europe spared no effort to turn
celebration and highly selective information into prescription. Yet, year after year
the Journal de physique devoted the entire January issue to review scientific
publications and achievements throughout Europe. Delametherie, its editor, made
no secret of the fact that he was challenging Cuviers rival accounts. For instance,
Cuvier never mentioned current debates on spontaneous generation, to him a non-
subject, whereas Delametherie informed readers and the historians of how many
variants of spontaneous generation doctrines were available in Europe during the
1800s and the 1810s.
I mentioned above the highly competitive market of dictionaries and collective
works. During the 1790s, Charles-Nicolas-Sigisbert Sonnini de Manoncourt
(17511812), a former collaborator of Buffon, launched an edition of his masters
work, to which scores of volumes were added, to turn it into a Complete course of
natural history. 127 volumes were published and sold to over 1,400 subscribers
throughout France and Europe.14 In 1803 the same editorial team launched the
Nouveau dictionnaire dhistoire naturelle, completed in the space of 2 years.15 The
24 volumes, relatively cheap set was sold to some 2,300 readers at Continental
level.16 Cuvier reacted with anger, as I have already documented elsewhere. In
1804 he issued an announcement inviting readers to subscribe to the forthcoming
Dictionnaire des sciences naturelles, placed under the editorship of his brother
Frederic (17731838). Adopting a tone of ease and superiority, Cuvier warned
readers not to buy the rival publication, which was not issued by the Professors of
the Museum as his own dictionary was, but by amateurs without much experience.
14
Charles-Nicolas-Sigisbert Sonnini de Manoncourt, ed., Histoire naturelle, generale et
particulie`re, par Leclerc de Buffon. Nouvelle edition, accompagnee de notes, dans lesquelles les
supplemens sont inseres dans le premier texte, a la place qui leur convient. [. . .], Paris, F. Dufart,
Year VII (1798)-1808. The list of more than 1400 subscribers does not take into account direct
sales from the printers and publishers.
15
Julien-Joseph Virey, Nouveau dictionnaire dhistoire naturelle appliquee aux arts, a lagricul-
ture, a leconomie rurale et domestique, a la medecine, etc., par une Societe de naturalistes et
dagriculteurs, 18031804. 1st ed., 24 vols., Paris, Deterville. 18161819; 2d ed., 36 vols., Paris,
Deterville. Already in 1807 the publishers were considering a second edition.
16
For an interesting contemporary comment on the Nouveau dictionnaire see H. Redhead Yorke,
Letters from France in 1802, 2 vols., London, H. D. Symonds, 1804, vol. 2, p. 333: This precious
work is published at so reasonable a price, that the sale will scarcely defray the expenses of paper
and printing. It is essentially a patriotic undertaking by Sonnini, Virey, Parmentier, Huzard, Bosc,
Olivier, Latreille, Chaptal, Cels, Thouin, Du Tour, and Patrin, men possessed of great knowledge
of the subjects of which they treat.
Cuvier badly lost the war of dictionaries. Only volume 5 of the Dictionnaire des
sciences naturelles appeared and only a handful of subscribers received copies of
volume 6. In 1816, the publishers of the Nouveau dictionnaire started a new
expanded edition. This time, 36 volumes were distributed in the space of 3 years
to almost 3,000 subscribers. Cuvier immediately retorted by changing the cover of
the first 6 volumes left unsold in 18041805, and launched the Dictionnaire des
sciences naturelles a second time over. Volume 60 was issued only in 1830, and a
careful perusal of the entire set (a task historians have surprisingly failed to
undertake) reveals that Cuvier and his brother did not, on the whole, spend much
time on it. Cuviers dictionary is a rather disappointing publication, lacking a firm
editorial line and, at time, even basic supervision. Embarrassingly, determined
enemies of Cuvier were allowed to write entries that contradicted what the famous
naturalist was preaching since his coming to Paris, as it was the case with the entry
Matiere verte, confided to Jean-Baptiste Bory de Saint-Vincent (17781846),
who did not miss the ironic chance of promoting spontaneous generation on the
pages of a publication nominally under the tutelage of Cuvier.17 One final point
needs to be emphasized. In the printed announcement we mentioned above, Cuvier
insisted that his dictionary represented institutional science, and his rivals were only
members of the unruly crowd of unreliable if not desperate amateurs. If one
compares the list of contributors to both dictionaries, one will immediately realize
that Cuvier did not tell the truth. The rival dictionary could boast contributions by
Jean-Antoine Claude Chaptal (17561832), member of the Institut and until 1804
Minister of the Interior; of Andre Thouin (17461824), the famous head gardener of
the Jardin des Plantes and himself member of the Institut; and of Antoine-Augustin
Parmentier (17371813), the ultra-popular pharmacist, lover of potatoes, and expert
in food preservation, among many other accomplishments. Parmentier, it is appro-
priate to mention, was the main sponsor of the Nouveau dictionnaire dhistoire
naturelle: in no way could he be considered an amateur. Thus, even Cuvier was at
pain to draw a clear line between the institutional science he was supposed to lead
and the actual articulations of natural history practices and publications over which
he clearly exercised little control.
During the 1810s and the 1820s, a series of popular generalist encyclopaedias
and of more specialized medical and natural history dictionaries did not hesitate to
openly challenge Cuviers authority. By 1816, Cuvier almost completely withdrew
from all teaching engagements, paid assistants and secretaries to do his job, and
became a full time member of the political establishment. Authors ranging from
Louis Antoine Desmoulins (17941828), Etienne Geoffroy Saint-Hilaire
(17721844) and Francois-Vincent Raspail (17941878) did not refrain even
from alluding to the fact that Cuvier was limiting himself to the writing of prefaces
to the various, very expensive volumes of the Re`gne animal or the Histoire
naturelle des poissons compiled by a host of ghost writers and researchers: he
17
J.-B. Bory de Saint Vincent, Matiere verte, in Dictionnaire des sciences naturelles, 29 (1823),
pp. 314336.
20 P. Corsi
had become an author of coffee table books.18 The question is not whether the
allegations (undoubtedly exaggerated) were true or not. The point to be emphasized
is that during the 1820s Cuviers authority was challenged at various institutional
and editorial levels, and the only power he could exercise was on his own salaried
staff: within certain limits, as we shall see below.
Thus, the view that Lamarck was silenced by Cuviers dominant and domineering
position within the hierarchy of contemporary natural sciences cannot be
substantiated. Cuvier hardly managed to silence anyone, even colleagues he
hated, such as the geologist and monarchist Barthelemy Faujas de Saint-Fond
(17411819, whom he nick-named Sans fond in reference to his colleagues
constant need for money), Geoffroy Saint-Hilaire (who was writing with the style
of a cook, Cuvier mused), Lamarck (a lonely man dominated by his imagination),
or wizards of the natural history publishing industry such as Delametherie (editor of
the then famous Journal de Physique) or Sonnini de Manoncourt. The latter
repeatedly and publicly made fun of his junior colleague, who pretended to disci-
pline all the rest of the community of naturalists by waving at them his pathetic
school teachers stick: people knew better, and bought the works he, Sonnini, was
producing.19
If one pays close attention to Cuviers career after the very early 1800s, one is
struck by the decreasing time he spent teaching and undertaking first hand research.
He was often absent from Paris, and when in town he devoted much of his
seemingly boundless energy to perform administrative duties and to keep his
many State jobs. Staying in power is never a given, and Cuvier, as contemporary
witnesses such as Stendhal, de Chateaubriand, or Cuviers distant relative we
mentioned above testify, worked very hard to keep surfing the treacherous, inces-
santly breaking waves of contemporary power.20
The assumption that Cuvier exercised a decided and unquestioned authority over
the natural history disciplines of his time can be maintained only at the price of
ignoring four fifths of the natural history scene of the time. As we will argue below,
Cuvier did not exercise full authority even on his direct collaborators, who were
wholly dependent on his good will to continue to earn a salary. They obeyed, did
what they were told, but in turn actively supported naturalists who did share their
admiration for the master, but felt free to pursue lines of research Cuvier was very
critical of, or openly despised.
As a client of power, Cuvier was hardly in a position to challenge power, even
when decisions concerning his own career and appointments were taken that forced
18
Antoine Desmoulins, Histoire naturelle des races humaines, Paris, Mequignon-Marvis, 1826,
p. viii; Francois-Vincent Raspail, Coteries scientifiques, Annales des sciences dobservation,
3 (1830), pp. 151159, p. 157; on Geoffroy Saint-Hilaires repeated attacks against Cuvier, see
Corsi, The Age of Lamarck, ch. VIII.
19
I have discussed Sonninis defiant attitude against Cuvier in The Age of Lamarck, pp. 3638.
20
Stendhal, The Life of Henry Brulard, translated and with an introduction by Jean Stewart and
Bert C. J. G. Knight, London, 1958, p. 180
him to swallow very unsavoury compromises. One single example will suffice.
When the highly respected former collaborator of Buffon, Louis-Jean-Marie
Daubenton, died at the end of December 1799, Cuvier aspired to succeed him to
the Chair of mineralogy at the Colle`ge de France. This was to him an important step
forward, and a prestigious, highly symbolic promotion. Delametherie, who had
spared no ink in publicly deriding his junior colleague, whom he depicted as an able
yet arrogant social climber, felt the chair belonged to himself: he was after all (in his
own eyes at least) one of Europes most prestigious mineralogists. Politics played a
key role in the appointment. Delametherie was supported by General Bonapartes
independently minded brother Lucien (17751840), and by his own brother
Antoine (17511804), an opaque yet faithful yes-man of the First Consul
Bonaparte. Antoine had helped Lucien to orchestrate and execute the coup-detat
of Brumaire 1799 that imposed the Consular regime and established Bonapartes
leadership. Almost on the day of the appointment Lucien and Napoleon quarrelled,
and Luciens authority was greatly diminished. The chemist and successor to
Lucien to the Ministry of Interior, ad interim in the fall of 1800, and fully in
1801, Chaptal, and his friend Bernard-Etienne de Lacepede (17561825), a natu-
ralist also very close to General Bonaparte, favoured Cuvier, who was duly
appointed. Yet, political debts to Antoine Delametherie had to be paid as well,
and the letter of appointment specified that Cuvier was granted the chair, but Jean-
Claude Delametherie was accorded the position of assistant. His salary was going to
be one-third of the salary allotted to the full chair, which Cuvier had to pay directly
to the hated rival.21 It was only in November 1802 that Cuvier was finally given a
full chair at the Museum, the same year in which he was appointed perpetual
secretary to the First Class of the Institute. As Dorinda Outram pointed out several
years ago, Cuvier would have been very pleased to know that historians would
make him the undisputed emperor of natural sciences. This would have consoled
him of the enormous efforts he kept producing to maintain his many jobs and his
many salaries in the real world of politics. He had little patience with academic
rituals which he did nevertheless perform as an accomplished master and did his
best to revenge himself against his intellectual and political enemies. Whether he
succeeded is of course another matter, and to assume that he managed to condemn
Lamarck to a bitter isolation is precisely a groundless assumption.
2 The Politics and Religion of Science
As we mentioned above, Lamarck has left precious little archival material; the
thousands of pages of manuscripts still preserved are in fact, for the most part, drafts
of his published works. Thus, we have only his printed works to try to understand
21
H. D. de Blainville, Observations sur la chaire dhistoire naturelle du Colle`ge de France, Paris,
1832.
22 P. Corsi
his standpoints on dramatic and burning issues such as politics and religion before
and after the French revolution and during the Empire and the Restoration. As far as
politics is concerned, Lamarck kept a low profile, though in 1794 he dedicated his
Recherches sur les principaux faits physiques to Jean-Paul Marat, refusing, he
added, earlier suggestions to inscribe the work to Louis Capet, the deceased
King. As far as religion is concerned, some commentators have insisted on his
mentioning the Deity here and there as evidence that Lamarck was at least a deist,
though they failed to notice that reference to a Superior Being increased with the
increasingly conservative intellectual and political climate of the Consulate, the
Empire and the Restoration. As an eternalist who claimed that nothing can
be created or destroyed in nature, since all elements exist since eternity; as someone
who wrote that religious ideas were created and spread by self-interested elites
aiming at dominating the majority of the population; and as someone who at the end
of his life affirmed that there was no metaphysical principle sustaining human life
and consciousness no soul, in other words, there is little doubt that he was not a
fervent nor even a lukewarm Christian, and I personally doubt he was even a deist.
What we know for sure is that he waited a relatively long time before baptizing his
children, and did so only in 1808.22 This was well into the Empire, when it was
abundantly clear to everyone that the regime would not tolerate open profession of
atheism.
Lamarcks post-1800 views, his transformist doctrines in particular, were indeed
accused of leading to, or even of advocating atheism. Julien-Joseph Virey (17751846)
explicitly linked Lamarck to atheism in the successful Nouveau dictionnaire dhistoire
naturelle, and repeated his charges several times until the 1830s. A precious testimony
has recently been unearthed by Herve Ferriere in his ground-breaking doctoral disser-
tation on Bory de Saint-Vincent.23 In 1804 Borys best friend, the entomologist Jean-
Marie Leon Dufour (17801865) was in charge of seeing through the press Borys
Voyage dans les quatre principales les des mers dAfrique since the author was away
from Paris on military duties. Dufour unsuccessfully tried to convince Bory to
suppress a chapter, in which the young naturalist sketched a non-Lamarckian
evolutionary model for the history of the Earth and of life, inspired by current
debates on the Theorie de la Terre: was he not aware, Dufour insisted, of what
priests and bigots (pretraille and bigotaille) were saying against Lamarck?
Prudence was called for, to avoid unnecessary danger and polemic. Though further
research is required to establish the actual foundation and origin of this rumour, there
22
Raphael Bange, Les ressources de letat civil parisien pour lhistoire des sciences. Lexemple
de Lamarck, Bulletin de la Societe dhistoire et depistemologie des sciences de la vie, 1 (1994),
pp. 3041.
23
H. Ferriere, Bory de Saint-Vincent (17781846): naturaliste, voyageur et militaire, entre
Revolution et Monarchie de Juillet; essai biographique, These de doctorat, 2 vols., Universite
Paris 1, Pantheon-Sorbonne, 2001 and Bory de Saint-Vincent: levolution dun voyageur
naturaliste, Paris, Syllepse, 2009.
is no doubt that Dufour is referring to something people talked about within natural
history circles.
Historians have often insisted on the atheistic tendency of Lamarcks tenets to
account for his alleged isolation, and I have myself pointed out instances which
show Lamarcks awareness of the risks he was facing. Though Lamarck has been
credited to be one of the earliest proponents of the word biologie (yet by no
means the first, as some historians love to repeat), the systematic analysis of the
Lamarckian corpus reveal that the proud announcement of his new project, the
establishment of biology, in January 1802, was followed only 6 months later by a
stern disclaimer: his age, commitments and bad health would prevent him from
carrying the project forward. In the Preface to the Philosophie Zoologique, the
famous two-volume work he published in 1809 as textbook for the new Imperial
University, Lamarck reassured readers that his biology was completely abandoned.
Surprisingly, in the preface to the first volume of the Histoire naturelle des animaux
sans verte`bres (1815) Lamarck informed his readers that he was working to
establish a new discipline, for which not even a name existed, which he proposed
to call biology. In a study I published a few years ago, I reconstructed the political
reasons that made Lamarck aware, in July 1802, that his project, and the word he
chose, biologie, could be associated to another word and project, ideologie,
then at the centre of intense political debate. As the ideologues wished to reform
philosophy and dispense with metaphysics and religious tutelage, Lamarck wished
to establish a science of life equally free from religious and philosophical
preconceptions, and solidly grounded on a set of explicitly materialistic assumptions.
In 1803 the ideologues were severely punished by General Bonaparte because of their
republicanism, but also for their opposition to the partial re-establishment of the
Catholic religion in the country, thanks to the Concordat of 1801. The second section
of the Institut, devoted to the moral and political sciences, was closed down, and its
members dispersed throughout the other sections. Many ideologues retained a certain
measure of influence, mainly through their work within the field of education and
legislation, but were kept at a safe distance from any form of political influence and
power.24
A man too proud to court powerful patrons, or simply socially inept, as Cuvier
suggested, Lamarck well knew that he was in a much weaker position than
representatives of ideologie. A campaign against his tenets and philosophical
propensities could have led to his dismissal, which would have meant total destitu-
tion for himself and his family. He therefore spared no words to reassure his readers
and potential critics that he had learned the lesson, and would not persist in pursuing
a line of research people saw as contrary to the sound principles of religion. In 1815,
Lamarck believed, as many Frenchmen did, that the Chart the restored King Louis
XVIII granted to the French people guaranteed full freedom of opinion and of
expression. He thus took up again his biology project, though the language was now
24
Pietro Corsi, Biologie, in P. Corsi et al., Lamarck, Philosophe de la nature, Paris, PUF, 2006,
pp. 3764.
24 P. Corsi
more guarded and prudent. The last works he composed from 1816 through 1820
expanded upon his materialistic interpretation of psychological and intellectual
phenomena, though here and there he paid lip service to the superior truths of
Revelation, which were guiding and correcting research when the highest moral and
metaphysical concerns were involved. During the 1820s, Lamarcks health
deteriorated, his blindness became total after 1818, and he withdrew from public
life and teaching. The Institut granted him the privilege of getting the token extra-
compensation accorded to members who attended sessions: the old naturalist could
not afford missing one.
The question to be asked, one I did not ask myself in my article of 2006 on the
politics of biologie, is the following: can we generalize the situation Lamarck
experienced, and conclude that the practice of natural history was subjected to
close scrutiny if not censorship during the Consulate, the Empire and the Restora-
tion? The answer is complex, and all the evidence at our disposal suggests great
prudence before embarking upon hasty generalizations. That Lamarck was afraid
for his job and livelihood does not mean that others were as well. Then, as now, the
danger of defending a minority or a fringe position was inversely proportional to the
social and political weight of the single individual going public. A few examples
will suffice to clarify this point.
We have already referred to Delametherie in connection with his appointment as
assistant to Cuvier at the Colle`ge de France. Delametherie was known as a die-hard
materialist. Though he often used a language compatible with main stream tradi-
tional eighteenth century deism, Delametherie liked to entertain foreign visitors on
the different shadows of contemporary French atheism.25 In one particular instance,
when accompanying a party of Englishmen on a private visit to the Museum
national dhistoire naturelle, he made fun of one of his guests, who, astounded
by the beauty of the display of the richness of living nature, had made a comment on
the clear indication the splendid collections offered of the existence of God: He
smiled and returned for answer, that I ought to recollect I was in an ecstasy, his
guest Henry Redhead Yorke (17721812) recalled.26 During the 1800s and the
early 1810s, Delametherie closely mirrored Lamarcks publications, taking up
almost the same topics and issues, to the point that a visitor like the Italian geologist
count Giuseppe Marzari Pencati (17791836) could mistake works by Lamarck for
works published by Delametherie, as Cuvier himself did (for different reasons,
needless to say). The differences between the two authors were significant, espe-
cially as far as the doctrine of the transformation of life throughout the history of the
Earth was concerned. Delametherie believed in a primeval Ocean in which rocks
25
On Delametheries atheism, see H.. D. de Blainville, Notice historique sur la vie et les ecrits de
J.-C. Delametherie, in Journal de Physique, 85 (817), pp. 78107, p. 89.
26
H. Redhead, Letters from France in 1802, vol. 1, p. 225. See also J. A. C. Sykes, ed., France in
Eighteen Hundred and Two Described in a Series of Contemporary Letters by Henry Redhead
Yorke, London, William Heinemann, 1906, p. 93. Sykes appeared to ignore the original work. His
edition is marred by hilarious spellings of the names of the main actors of Parisian life in 1802.
and life were formed through countless ages thanks to processes of crystallization,
and maintained that all life forms known to man developed from a restricted
number of prototypes equally generated by specific forms of crystallization all
of which Lamarck firmly denied. Yet, there is no doubt that Delametherie thought
that Lamarck had copied some of his ideas, and wrote extensively on the action of
habits in giving new shapes to organs within the well defined limits of the original
crystallization process that had established the prototypic form, one should hasten
to add. On this subject his discussion was more detailed and sustained than
Lamarcks. Delametherie too was a civil servant, not very well off, having decided
to repay the heavy debts his brother incurred at gambling. Yet, his political
connections and his high European reputation as editor of the Journal de physique
a periodical historians are very unwise to ignore bought him a degree of free
expression, higher than the one Lamarck felt was allotted to him.
We have already mentioned the dark prophecy Leon Dufour addressed to his
friend Bory de Saint-Vincent. The Catholic party would not forgive him for his
materialist account of the history of the Earth and of life on it. It is interesting to
note that Dufour too, as Marzari Pencati and Cuvier, appeared to believe that
Lamarck endorsed the hypothesis of a primeval Ocean. What is more interesting
from the point of view of our discussion is that no Catholic reviewer took up the
challenge. As Herve Ferriere has brilliantly shown, Catholic reviewers were not
nice to Bory, and even accused him of having deserted the famous expedition to the
South Seas led by Captain Nicolas-Thomas Baudin (17541803) which was
indeed true, since Bory (together with other naturalists of the expedition) jumped
ship at the Ile de France, todays Mauritius, and refused to continue the journey.
Yet, no one mentioned the materialist and possibly atheistic tendency of the book.
Favourable reviewers also avoided any comment on the incriminated chapter,
though only 3 years earlier the debate on the Theory of the Earth had aroused
generous comment, and Delametherie was never tired of providing new evidence
for his own geological and biological views in the pages of the Journal de physique.
So, why, contrary to the prediction made by Dufour, and in spite of religious attacks
against Lamarck and his doctrines, friends and foes alike avoided commenting on
Borys equally objectionable doctrines? We cannot know for sure and here again
further research is required. Yet, the fact that Borys uncle and surrogate father (the
young naturalist was an orphan), Bernard Journu-Auber (17451815), was one of
the richest men in the country, and a close collaborator of General Bonaparte as
Regent of the Central Bank of France, may explain why no one dared to attack a
naturalist placed under such a tutelage. Even the serious accusation of desertion
came to nothing: indeed, Journu-Auber even proposed that his protege should be
promoted to the rank of Captain for the services rendered to the State during the
expedition he deserted.
A final example, concerning an amateur naturalist well known in his time
throughout Europe, and completely ignored by historians, will reinforce the point
that during the Consulate and the Empire the danger of maintaining unsavoury
philosophical or scientific tenets was inversely proportional to the social status of
the proponent. Jean-Baptiste Fray-Fournier (17641835) was a surgeon and
26 P. Corsi
amateur naturalist who lost the use of his right hand and opted for a lucrative career
as a purveyor for the army (Commissaire Ordonnateur des Guerres) and organizer
of military hospitals for the Grande Armee.27 He travelled and worked through
several German States, and resided for sometime in Berlin, Magdeburg and Ulm,
from where he engaged in correspondence with leading German naturalists. Frays
passion, and in his mind his title to consideration, were his experiments on the
spontaneous generation of organic molecules. He got Pierre Jean Georges Cabanis
(17571808) interested in his work, and even performed at Arcueil, under the eyes
of Claude Louis Berthollet (17481822) and his assistants. Fray, a believer and a
Christian, was convinced that organic molecules were formed thanks to the action
of solar rays, and they could combine to give birth to elementary forms of life. Once
life started developing, everywhere in the world it would climb the ladder of
complexity from monad to man. Though the process was the same everywhere on
Earth, the end result (as well as the intermediate steps) was marked by the physico-
chemical peculiarities of the locality were the process started. Thus, he explained,
on the top of the Pyrenees one can find trees that are specific to the locality, since
they developed from spontaneous generations made up of basic molecules typical
of the physico-chemical constitution of those mountains. Though Fray did not
doubt that the process had been providentially designed by an all-benevolent
creator, his account of the origin and development of life on Earth bore here and
there close resemblance to points Delametherie, Bory and other materialists
advocated. Frays work attracted favourable attention in Germany. Top naturalists
such as Friedrich Tiedemann (17811861), Johann Friedrich Meckel (17811833)
and Wilhelm August Eberhard Lampadius (17721842) praised his work, and
adapted it to their own views of the origin and development of life forms. In France,
Antoine Desmoulins quoted with approval without mentioning the name of Fray
the explanation for the peculiar flora and fauna showed by isolated geographical
areas such as the picks of the Pyrenees, whereas during the 1820s Bory de Saint-
Vincent and others denied that Fray had preceded them in elaborating the theory of
the spontaneous generation of organic molecules. In the British Isles, Frays work
was well known to John Barclay (17581826), a famous teacher of anatomy at
Edinburgh, who in 1822 devoted a long chapter of his An Inquiry Into The
Opinions, Ancient And Modern, Concerning Life And Organization to a refutation
of the Frenchmans work (whom he considered a better thinker than Erasmus
Darwin), filled with quotations in French. Barclays even bothered to reproduce
the paragraphs Cabanis devoted to the first experiments performed by Fray in the
early 1800s.28 I do not need to expand upon the fact that in France no one appears to
27
Joelle Jezierski, De fleur et de sang. Parcours dun herbier napoleonien, in Machine a feu.
Revue du livre et de la lecture en Limousin, 25 (2007), pp.4041
28
Jean-Baptiste Fray, Essai sur lorigine des corps organises et inorganises, et sur quelques
phenome`nes de physiologie animale et vegetale, Paris, Mme Ve Courcier, 1817. An earlier and
shorter version of the work, Nouvelles experiences extraites dun manuscrit qui a pour titre: essai
sur lorigine des substances organisees et inorganisees had been published in Berlin, L. Quien,
have challenged Frays tenets or asked him to distance himself from current
materialist and atheist explanations for the history of life on earth. His trusted
role as a pillar of the medical services of the Grande Armee acted as a very effective
shield.
What about the Restoration? After an initial tolerant attitude towards the free-
dom of the press and of opinion, successive ultra-monarchic administrations
introduced increasingly restrictive measures. Censorship was exercised with the
utmost severity on theatre productions, or on the teaching of history in schools
preparing students for the Agregation.29 In 1825 an anti-blasphemy legislation was
passed, so extreme that even a famous writer and right wing politician such as
Francois-Rene de Chateaubriand (17681848) felt that Chares X had done what no
King of France had ever dreamt of. One might expect scientific doctrines openly or
implicitly favouring atheism and materialism to be repressed, denounced or at least
censored, Lamarcks in primis. Systematic perusal of periodicals, encyclopaedias,
dictionaries and single works published during the Restoration reveal that this was
not the case. Conservative writers, including Cuvier, denounced as leaning towards
pantheism, if not atheism, Lamarckian transformism, the doctrine of the unity of
composition prevailing according to Geoffroy Saint-Hilaire and his allies
throughout the animal and vegetable kingdoms, or the theory of embryonic recapit-
ulation, according to which the phases of development of the embryo summed up
the major steps in the development of life throughout the history of the Earth.
Cuviers or Vireys strictures only added to the success of dangerous doctrines
with the reading public. From the pages of the internationally successful
Dictionnaire classique dhistoire naturelle (which travelled with Darwin on the
Beagle) he edited from 1822 to 1832, Bory de Saint-Vincent blasted against the
Jesuits and the theory of the immortality of the soul, as well as against Cuviers
conservatism, and extolled the virtues of Lamarcks dedication to scientific truth
and his colleagues transformist doctrines. Bory vigorously campaigned in favour
of polygenism, and his many works provided readers in Europe and the United
States with accurate summaries of the various standpoints debated by French
naturalists. He avoided mentioning, needless to say, that Lamarck believed in the
unity of the human species.
During the 1820s, the golden decade for Lamarcks reputation in France and
Europe, his doctrines were subjected to a variety of criticisms and only a few
commentators insisted on the dangerous leaning of his teaching and theorizing.
From Edinburgh to G ottingen, from Turin to Paris, it was common to pay homage
to the old naturalist, who had left a monument of taxonomic achievement such as
the Histoire naturelle des animaux sans verte`bres (7 vols., 18151822). The fact
and Paris, chez Nicolle, 1807. J. Barclay, An Inquiry Into The Opinions, Ancient And Modern,
Concerning Life And Organization, Edinburgh, Bell and Bradfute, 1822, pp. 126142; for the
quotations from Cabanis, see pp. 127128.
29
V. Granata, Politica del teatro e teatro della politica: censura, partiti e opinione pubblica a
Parigi nel primo Ottocento, Milano, Unicopli, 2008.
28 P. Corsi
that Lamarck had become blind in 1818, and was now too sick to take part in public
and scientific life, only added to the respect surrounding him during his last years. It
is therefore clear that the assumption that Lamarck was isolated because of the
religious and philosophical implications and consequences of his transformist
doctrines has no foundation, and can be maintained only by ignoring the actual
state of affairs in contemporary French and European scientific, political and
cultural life.
3 Authors and Audiences
Historians of science have traditionally shown a remarkable reluctance to accept the

simple fact that at every given moment, the production of knowledge in a given
society is as varied and diversified as the social, political or the religious scene. Since
some individuals or groups of individuals appear to share our concept of science, or
to approach it the most, they are taken as the only ones worth spending ones time on.
In doing so, a host of very interesting phenomena and events are completely ignored,
even the ones that should appeal to the historians of the development of scientific
truth. The case of Cuvier and his collaborators we mentioned above, or the
composite nature of the natural history scene during the decades in which Lamarck
acted his own life, deserve comment. I will deal in particular with selected features
of debates within the medical profession; the latters changing political and intellec-
tual allegiances; and the editorial ventures that represented the viewpoint of promi-
nent factions within it. This will open up new and fascinating perspectives on the
relationship between France and Germany during the first three decades of the
nineteenth century, and on how Lamarck was read by different audiences.
I hinted above that Cuvier was hardly in a position to dominate the natural
history scene of his time, and experienced some difficulty even with his own
collaborators, who depended on his good will more than anyone else. I will limit
my comments to the two young authors who put Cuviers lectures on comparative
anatomy in good form and published it. As is well known, Andre Marie Constant
Dumeril (17741860) took care of volumes 1 and 2 of the Lecons danatomie
comparee (5 vols, 18001805), whereas his colleague Georges Louis Duvernoy
(17771855) edited vols. 35. Dumeril was undoubtedly the more independent of
the two. A brilliant and precocious anatomist and medical lecturer, he arrived in
Paris in 1800, and was immediately recruited by Cuvier to help him in all sorts of
tasks, including the editing of his lectures. Dumeril also engaged in teaching in the
school of medicine of the capital that had taken the place of the abolished Medical
Faculty. Duvernoy was coming from Montbeliard, Cuviers home town, and was a
distant relative of the naturalist. He was not as brilliant as Dumeril, and his career
was less conspicuous. Yet, even Duvernoy played a role we still know little about,
and an important one, for that matter. As I pointed out years ago, his views of life
before entering Cuviers service were not as Cuvierian as one might assume.
He saw an important role the medical philosopher could play when reflecting on
the properties of living beings. He appeared to see life in terms of a fluid dynamics
not dissimilar to the one Lamarck endorsed, though he rejected Delametheries and
Lamarcks materialism. He also expanded upon the role of use and lack of use in
shaping organs.30
We pointed out at the beginning of this article that the gathering of important
collections in Paris forced naturalists from all over Europe to spend time in the
French capital in order to take advantage of the unprecedented wealth of
specimens. A young ambitious German anatomist, Johann Friedrich Meckel
(17811833), spent the years 18041805 working in Cuviers laboratory. Back
to Germany, Meckel translated the Lecons danatomie comparee before
embarking on his own ambitious editorial projects. The role of Meckel in early
nineteenth century German comparative anatomy and embryology is now well
known, thanks to important studies published during the last 20 years.31 Yet, it is
always assumed that during his stay in Paris the young anatomist worked with
Cuvier. Now, this is again an easy and reasonable assumption, albeit one that it is
difficult to prove. During the very early 1800s Cuvier was dividing his time
between extensive travel through the French provinces, as Inspector General of
higher education, and his growing political commitments. What we know for sure
is that Meckel embarked upon the careful dissection of a limited number of human
foetuses under the direct supervision, and collaboration, of Duvernoy.32 If
we follow Duvernoys highly interesting narrative of events, published in 1849,
we learn that Meckel worked on nine foetuses (a rare specimen to obtain even in
Germany, though less so in Paris), and called upon Duvernoy to attest the
reliability of his observations. Back home, in 1806 (in fact, already in 1805)
30
G.-L. Duvernoy, Reflexions sur les corps organises et les sciences dont ils sont lobjet, in
Magasin encyclopedique, 5th year, vol. 3 (1799), pp. 459474. See Corsi, The Age of Lamarck,
pp. 7576.
31
S. Gliboff, H.G. Bronn, Ernst Haeckel, and the Origins of German Darwinism: A Study in
Translation and Transformation, Cambridge, MA, MIT Press, 2008. P. Hunemann, ed., Kant and
the Philosophy of Biology, Rochester N.Y., University of Rochester Press, 2007. R. Richards, The
Romantic Conception of Life: Science and Philosophy in the Age of Goethe, Chicago, University
of Chicago Press, 2002. T. Lenoir, Kant, Blumenbach, and Vital Materialism in German
Biology, in Isis, 71 (1980), pp. 77108, and The Strategy of Life: Teleology and Mechanics in
Nineteenth Century German Biology, Dordrecht, Reidel, 1982. T. Bach, Biologie und Philosophie
bei C. F. Kielmeyer und F. W. J. Schelling, Stuttgart, Bad-Cannstatt, 2001. S. Schmitt, Les forces
vitales et leur distribution dans la nature: un essai de systematique physiologique. Textes de
Kielmeyer, Link et Oken traduits et commentes, Paris, Brepols, 2007.
32
G.-L. Duvernoy, Ovologie, in Dictionnaire universel dhistoire naturelle, 9 (1849),
pp. 281353.
30 P. Corsi
Meckel published in German an essay, Fragments sur lhistoire du

developpement du foetus humain (we are of course following Duvernoy, who
gave titles and texts in French), which ended with the conclusion:
Je suis loin de regarder comme une idee simplement ingenieuse, celle de Kielmeyer, qui
pense que le foetus humain passe par les divers degres de developpement auxquels
sarretent les animaux inferieurs. Un trop grand nombre de faits viennent le confirmer.33
One can therefore surmise that Meckel and Duvernoy discussed several issues in
comparative anatomy and embryology, and it is possible that Meckel told Duvernoy
of the many new ideas Carl Friedrich Kielmeyer (17651844), whom he had met
and listened to, was expanding upon in his lectures and manuscripts. Of course,
Cuvier too knew rather well what Kielmeyer was speculating upon. Yet, Duvernoy,
who in his contributions to the Dictionnaire universel took pains even to show that
Geoffroy Saint-Hilaires transcendental anatomy was in the last analysis due to
Cuvier, is completely silent on the role his master played in directing Meckels
work. Indeed, Cuvier does not appear to have played any role in Meckels research.
As hinted above, the young German anatomist asked Duvernoy to witness his
anatomies: he would not have missed the chance to say that the mentors role had
been fulfilled by the already famous Cuvier, who by the way never paid much
attention to embryology and deeply opposed the doctrine of embryologic recapitu-
lation. It is possible that Antoine Etienne Renaud Augustin Serres (17861868), the
French main proponent of embryologic recapitulation, discussed recapitulation
with Meckel, whom he met in 1805. Serres was a consumer of German works
and became a keen reader and admirer of Lorenz Oken (17791851). The point
which is important to stress is that Meckel became proficient in the French language
and forged close personal links with several French colleagues, especially among
the young aspiring naturalists and medical researchers courting notoriety through
their anatomical research. Further research is required to reach a better understand-
ing of the workings of Cuviers laboratory and the laboratories of other Professors
of the Museum and the modalities of exchange with foreign visitors and
researchers.
Dumeril was apparently more successful than Duvernoy in attracting young
talent, though what follows only constitute the result of a first survey in need of
more sustained research. A precocious anatomist and medical teacher at 19,
Dumeril, as we have already pointed out, reached Paris at the beginning of 1800.
He was quickly noted by Cuvier, who associated him to his Lecons danatomie
comparee, asked him to teach his class at the Ecole Centrale du Pantheon one of
the many jobs Cuvier was starting to hand out to faithful pupils and in 1804 even
33
Duvernoy, Ovologie, p. 348. J. F. Meckel, Fragmente aus der Entwicklungsgeschichte des
menschlichen ftus, in Abhandlungen aus der menschlichen und vergleichenden Anatomie und
Physiologie, Halle, Hemmerde und Schwetschke, 1805, pp. 277381, now available at Edition
Classic VDM Verlag Dr. Muller, 2007. See L. G obbel and R. Schultka, Meckel the Younger and
his Epistemology of Organic Form: Morphology in the pre-Gegenbaurian Age, in Theory in
Biosciences, 122 (2003), pp. 127141.
passed on to him the task of writing a textbook on natural history he had no time to
engage in, commissioned by the Government for the Lycees. Dumeril showed
himself a reliable follower. In the textbook, he never mentioned Lamarck as an
expert on invertebrates, but only Cuvier. All innovation in the classification of
invertebrates was due to Cuviers anatomical research which was to a great extent
true- and Lamarck had played no role in reforming this important branch of
zoology. Thus, no mention was made of the Syste`me des animaux sans verte`bres
Lamarck had published in 1801, and his name appeared briefly in the text in the
sections devoted to botany.34
Yet, if one lists the pupils Dumeril himself took under his wings, and one
considers what he himself published, one can see that he kept a good measure of
independence from his master. In the very early 1800s he taught a young doctor,
Jean Burdin, author of a Cours detudes medicales (1803) which contained the first
published hint that the cranium could be seen as formed of expanded vertebrae.35
The three volumes work was translated into English and German within the year,
even though we have no idea of who paid for this possibly Henri de Saint-Simon
(17601825), then still quite wealthy, and apparently close to Burdin. Early in
1808, Dumeril himself proposed a vertebral theory of the cranium to the Institut, but
was laughed down when someone in the audience uttered the ironic comment here
is the thinking vertebra.36 He also endorsed the theory of the unity of plan
prevailing throughout the animal kingdom, about which Cuvier was expressing
growing reservations. Another pupil of Dumeril was the irascible, highly original
Antoine Desmoulins, who during the 1820s became a fierce and relentless opponent
of Cuvier. It must be said that Desmoulins managed to quarrel with everybody in
Paris, and even succeeded in making Geoffroy Saint-Hilaire defend Cuvier in print,
so outrageous were his attacks against the famous naturalist.
The last pupil Dumeril coached and protected of whom I am aware is Antoine
Jacques Louis Jourdan (17881848), an author deserving close attention. After
studying medicine and opting at first for the lower qualification of surgeon, Jourdan
left France to join the French army in Germany. He was mainly based in
Konigsberg and Berlin, and remained in the German states from 1808 until 1814
(with the exception of short stays in Paris, when training at the military hospital of
Val de Grace). We have no idea of whether he had met Meckel in 18041805, when
he was already a pupil of Dumeril. What is certain is that he forged personal links
34
C. Dumeril, Traite elementaire dhistoire naturelle, Paris, Crapelet, 1804, 2nd ed., 2 vols. Paris,
Deterville, 1807. It is to be pointed out that in 1803 Dumeril was appointed assistant to Lacepede,
who had taken up heavy political and administrative duties.
35
C. G. Carus, Traite elementaire danatomie comparee, suivi de recherches danatomie
philosophique ou transcendante [. . .] traduit de lAllemand par A.-J.-L. Jourdan, 3 vols., Paris
et Londres, J.-B. Bailliere, 1835. See pp. 45 for comments on Burdin, and on the contemporary
development in France and Germany of ideas about the vertebrae composing the cranium. Carus
also pointed out that Italy and England had contributed precious little to the philosophical
developments in anatomy.
36
See Corsi, The Age of Lamarck, pp. 237238.
32 P. Corsi
with many German anatomists and medical researchers, Meckel included, and
started a systematic and gigantic project of translating German works into French.
In his long career, he signed more than 70 major translations, including the works of
Christian Friedrich Samuel Hahnemann (17551843), the founder of Homeopathy,
a medical doctrine Jourdan endorsed and introduced to France.
From the point of view of Cuvier, Jourdan showed a remarkable propensity to
translate all the authors the great anatomist deeply disliked, Karl Friedrich Burdach
(17761847) Gottfried Reinhold Treviranus (17761837) or Carl Gustav Carus
(17891869) among others. Jourdan became a very active collaborator to the
Dictionnaire des sciences medicales (60 vols. 18121822) a remarkable achieve-
ment of the French publishing industry. In 1818 he also became one of the chief
authors and probably one of the editors of the Journal complementaire du
Dictionnaire des sciences medicales, a periodical that, as the title suggests, was
designed to update the entries already published in the dictionary, as well as to
provide welcome summaries of medical progress throughout Europe. It is signifi-
cant that the editorial statement printed in issue n. 1. (signed by the publishers, but
bearing clear marks of Jourdans style) listed 30 medical periodicals published in
German, 6 in Italian, 5 in English and 1 in Dutch. Last but not least, Jourdan and the
Journal complementaire paid particular attention to Meckel and his innovative,
systematic and truly impressive attempt to unite the study of comparative embryol-
ogy and the study of comparative anatomy.
No polemical hint was made at least until the mid-1820s but medical people
well understood the meaning of the translation campaign from German and the
policy pursued by the very successful Journal complementaire and other medical
publications The fact is that several representatives of the medical profession did
not like Cuviers attempt to establish his own superiority in comparative anatomy
and human anatomy. Many openly extolled the work of Marie Francois Xavier
Bichat (17711802), the true reformer of French anatomical studies: no need to add
that, in their view, Cuvier was not. During the late 1810s, Geoffroy Saint-Hilaire
early incursion in and statements on transcendental anatomy, supported by eminent
representatives of the medical profession, such as Serres and members of the
Societe danatomie, as well as Geoffroys work on teratology, were perceived as
constituting a satisfactory vindication of the autonomy of medical research with
respect to research conducted within natural history disciplines. An alliance could
be forged with Geoffroy, because of his strong links with the medical profession, and
his growing confrontation with Cuvier. Moreover, starting with the early-1820s,
severe repressive measure against leading members of the medical community,
including the then old and extremely respected Philippe Pinel (17451826), threw
leaders of the profession and ranks and file doctors into determined opposition to the
Government. Medical journals and encyclopaedias, pamphlets and reports of
meetings of medical organizations joined naturalists such as Geoffroy and his son
Isidore or Bory de Saint Vincent in deprecating the poor state of natural sciences in
France, mirroring the poor state of political freedoms.
What about Lamarck in all this? And what about the larger picture of Franco-
German scientific relationships during the early decades of the nineteenth century?
Jourdan expressed his admiration for Lamarck in a review of the Histoire naturelle
des animaux sans verte`bres and in a long entry, Germe, devoted to the question of
epigenesis and preformation, published in volume 18 of the Dictionnaire des
sciences medicales.37 Contrary to his colleague Virey, who also wrote extensively
for the dictionary and kept insisting that Lamarcks doctrines lead to atheism,
Jourdan summed up Lamarcks main tenets with fairness and admiration, though
he felt free to disagree on several key points. Jourdan, who had attended Lamarcks
lectures in 1806, was reading his teachers work with German eyes, so to speak.
Thus, for instance, he argued that Lamarcks insistence on the role of organic fluids
and on fluid dynamics in explaining biological phenomena was very similar to the
doctrines put forward by Johann Christian Reil (17591813), the founder of the
famous Archiv f ur die Physiologie (17961815).38
Jourdans account of the doctrine of spontaneous generation proposed by his
former teacher Lamarck correctly included reference to the different origins of the
plant and animal kingdoms, and possibly of several branches of main types of
animal organization (p. 161). His interpretation of the progress of organization
made no reference to the tendency of life to develop more complex anatomical
and functional structures. Jourdan rightly stressed that, according to Lamarck, from
time to time the dynamic interaction between the organism and the environment
required a change in the distribution pattern of fluids and nutrients, thereby
favouring those parts that were solicited by a change in the habits of animals. In
its turn, the change of habits was the consequence of a change in the physical or
biological environment. More diversified and complex distribution channels for the
fluids also implied a more specialized, and in the end more rapid circulation. The
nervous system in humans constituted the best example of extremely fast
movements of fluids inside a very sophisticated network of nervous fibres
(pp. 167180). Thus, the only progressive change in living bodies was constituted
by the inevitable increase of the speed at which fluids moved within organisms, due
to the specialization and refinement of conveying networks and ultimately of the
fluids themselves. This did not, however, determine the way in which the organism
was going to develop, since the creation of new organs depended on the actual
circumstances in which animals found themselves and the actual challenges they
had to face (pp. 163164). Jourdan was not a convert, however. He expressed his
conviction that the species barrier could not be overcome. Advanced experiments in
domestication showed that even though much could be achieved by breeders, never
a zebra had become a horse, he ironically concluded.
37
A.-J.-L. Jourdan, Histoire naturelle des animaux sans vertebres, in Journal universel des
sciences medicales, 2 (1816), pp. 145181, and Germe, Dictionnaire des sciences medicales,
18 (1817), pp. 226277.
38
Jourdan devoted a long article to the career and doctrines of Reil, again pointing out the
similarities with Lamarck, Litterature medicale allemande. Sur la connaissances et le traitement
des fievres, par Jean-Chretien Reil, in Journal universel des sciences medicales, 2 (1816),
pp. 217239.
34 P. Corsi
The entry in the medical dictionary, published a few months after the review we
have alluded to, adds interesting dimensions to Jourdans attitude towards Lamarck.
As in his review of the Histoire naturelle, the tone was respectful and sympathetic.
Lamarck was to Jourdan one of the chief supporters of epigenesis, one who had
convincingly shown the weakness of the preformist doctrine. Jourdan summarized
with favour the theories that saw embryos and individual adults develop thanks to
the addition of parts made possible by nutrition or produced thanks to environmental
chemical and physical agents. Jourdan stressed that both preformism and epigenesis
impinged upon our conception of the origin and the development of animal and
plant life on Earth. Whereas preformists tended to embrace a strong or a weak
version of creationism (all germs of all animals were created at the beginning of
time, or successive creations of new germs occurred to fill gaps and losses), the
followers of epigenesis were open to the idea that organisms were formed in
succession through endless ages and endlessly changing environments. Lamarck,
according to Jourdan, was the most coherent representative of the latter view.39
Once again, Jourdan was not a convert. His grasp of the niceties and
complexities of the Lamarckian theoretical corpus is often well above what
historians and other commentators have customarily said of Lamarck. Yet, he
was not convinced that use and lack of use, and more generally the dynamic
interaction between organisms and their environment was sufficient to explain
macro-evolution. The Lamarckian mechanism accounted for the fixation of
varieties into good species, perhaps for the production of new genera, but could
not explain the emergence of new and more complex anatomical and functional
structures. For this, one had to call upon the still unknown laws of development, at
the level of the embryo, as well as at the level of the history of life on earth. In later
years, Jourdan agreed with Meckel, Tiedemann, and his colleague Serres, that the
development of the embryo showed the steps life had to climb in order to produce
increasingly perfect beings. Once life was created, indeed, as Lamarck had pointed
out, once several forms of spontaneous generation had appeared, each endowed
with its own specific structural properties, each form could climb only according to
the potential for growth its structure allowed. The laws of development were the
key factor, both in ontogeny and in phylogeny. Lamarckian mechanisms only
explained how birds adapted to all the environments they were found in, not how
the anatomical and functional type bird had originated, or, better, had developed
from less complex vertebrates.
Jourdan commented on Lamarcks ideas on several occasions, and always with
the utmost respect. To him, as well as to Bory or Geoffroy, Lamarck deserved the
full respect of the scientific community. He had dared to be wrong, whereas Cuvier
simply censored other peoples ideas. And he had left a scientific legacy which
Cuviers compilations could never match. There is of course no space to analyse the
attitudes of other members of the medical community towards Lamarck authors
39
A.-J.-L. Jourdan, Germe, Dictionnaire des sciences medicales, 18 (1817), pp. 226277.
such as Nicolas Philibert Adelon (17821862), for instance, who wrote one of the
rare reviews of Lamarcks last work, the Syste`me analytique des connaissances
positives de lhomme (1820), and kept referring to Lamarck in every successive
edition of his successful medical textbook.40
It is by now abundantly clear that the restriction of the analysis of the reception
of Lamarcks doctrines to what Cuvier or a handful of naturalists had to say
seriously distorts our understanding of debates on the life sciences during the
early decades of the nineteenth century. Not only the vast majority of practitioners
of natural history and authors of natural history publications have been denied any
hearing, but the vociferous and politically very active population of medical writers
and practitioners has been completely ignored. As far as France is concerned, it is
not uninteresting to mention that among the most prolific medical authors of the late
1810s and the 1820s a good number were former medical officers of the Napoleonic
armies, left without pay, as all former army senior staff had been, by a vindictive
decree of the Restoration Government. Many wrote as much as possible simply to
implement their income. Their economic needs, their need for recognition and their
wounded pride contributed to create an explosive mixture that added to the mount-
ing tension leading to the July 1830 revolution. It is not by chance that after July
1830 many medical radicals of the 1820s returned to their professional occupations,
lowered their tone and relented their assaults against official science. Many
journals, such as the Journal complementaire, even abandoned their campaign in
favour of German medicine and anatomy. It was time to stop imitating foreigners.
True, other journals did take up the fight, and dictionaries of the 1830s, 1840s and
1850s kept discussing transcendental anatomy or Lamarckian doctrines. Simply,
those discussions had lost the political pregnancy and urgency they had taken on
during the 1820s.
One final comment is called for. The question of the relationship between French
medical authors and their German colleagues cannot be looked at as a parochial
anecdote or a minor episode within the larger picture of European debates on life of
the early nineteenth century. Several historians have recently argued for the impor-
tance of German anatomical, embryological and medical doctrines for the debates
on the life sciences that marked the period 18201850 in Scotland and England, for
instance, and for the formation of Charles Darwins view of nature and of life.41 It is
perhaps useful to recollect that Jourdans translations of Meckel, Tiedemann,
Carus, Burdach, Treviranus were in fact destined to the British book trade as well
40
N. P. Adelon, Systeme analytique, in Revue encyclopedique, 9 (1821), pp. 257267, and
Physiologie de lhomme, 4 vols., Paris, Compere jeune, 18231824, see vol. 4, pp. 34, 103104,
114115, 232, for favorable summaries of Lamarcks ideas. Adelon was one of the editors of the
Dictionnaire des sciences medicales.
41
See R. Richards, The Romantic Conception of Life: Science and Philosophy in the Age of
Goethe, Chicago, University of Chicago Press, 2002, for an authoritative presentation of this
argument and the relevant bibliography. See also Philip F. Rehbock, The philosophical naturalists:
themes in early nineteenth-century British biology, Madison, University of Wisconsin Press, 1983.
36 P. Corsi
as to the French one. It was the entrepreneurial genius of Jean-Baptiste Marie

Bailliere (17971885) that saw the opportunity for profit to be gained in England,
and from his shop in Regent Street the French translations of German works were
sold to private and public medical libraries. More than that: the English language
edition of Tiedemanns seminal work Anatomie du cerveau: contenant lhistoire de
son developpement dans le foetus was undertaken from the French translation, not
from the German original.42 The introduction by Jourdan, as well as his annotations
to the text, was probably considered a kind of added value, though one cannot
exclude that during the 1820s and the 1830s it was easier to find in London a
translator from the French language rather than from the German one. Meckels
works circulated in England in the French editions, rarely in the original German.
Even famous anatomists who knew German, such as the polyglot Robert Edmund
Grant (17931874), or Robert Knox (17911861), owned Meckel in the French
edition, not the German one. An English language edition of the handbook of
anatomy by Meckel was nevertheless published in the United States from the
French edition edited by Jourdan, though the translator informed his readers that
he had consulted a German speaking medical man, in order to correct a few
mistakes and inaccuracies present in the French edition.43
The set of easy assumptions concerning the place and reputation of Lamarck
within the French natural history community of the early decades of the nineteenth
century has traditionally acted as true Idola tribus, preventing research and limiting
in considerable ways our understanding of the complex intellectual, social and
political dynamics of contemporary natural history practices and publishing. The
almost total lack of interest for the state of affairs in the publishing industry of the
period under consideration, and the total lack of interest for what books,
dictionaries, encyclopaedias actually said, has made us blind to major debates of
great significance for the history of the life sciences at European level during the
early decades of the nineteenth century. The reconstruction of the ways in which
Lamarck was read, admired, criticized or denounced cannot be undertaken without
reconstructing the actual articulations of the contemporary natural history and
medical scene, in all its institutional, social and political dimensions.
42
The French translation of Tiedemann appeared in 1823; the English language edition, The
anatomy of the ftal brain: with a comparative exposition of its structure in animals [. . .]
Translated from the French of A. J. L. Jourdan, by William Bennett, M. D., appeared in Edinburgh
in 1826, J. Carfrae and Son.
43
Manual of general, descriptive, and pathological anatomy, by J. F. Meckel [. . .] Translated
from the German into French, with additions and notes, by A. J.L. Jourdan and G. Breschet.
Translated from the French, with notes, by A. Sidney Doane, 3 vols., Philadelphia, Carey & Lea,
1832. Doane, a graduate from Harvard University, had studied in Paris during 18301832. He
translated several French medical textbooks and specialized monographs into English.
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foetus: avec une exposition comparative de sa structure dans les animaux; traduite de
lallemand . . . par AJL Jourdan. JB Bailliere, Paris
62. Tiedemann F (1826) The anatomy of the foetal brain: with a comparative exposition of its
structure in animals [. . .] Translated from the French of AJL Jourdan by William Bennett MD.
To which are added some late observations on the influence of the sanguineous system over the
development of the nervous system in general. Illustrated by fourteen engravings. J. Carfrae
and Son, Edinburgh
63. Virey JJ (181619) Nouveau dictionnaire dhistoire naturelle appliquee aux arts a lagriculture
a leconomie rurale et domestique a la medecine etc. par une Societe de naturalistes et
dagriculteurs 18031804. 1st edn, 24 vols, Paris Deterville ; 2nd edn, 36 vols
64. Virey JJ (181619) Nouveau dictionnaire dhistoire naturelle appliquee aux arts a lagriculture
a leconomie rurale et domestique a la medecine etc. par une Societe de naturalistes et
dagriculteurs 18031804. 1st edn, 24 vols, Paris Deterville; 2nd edn, 36 vols, Paris Deterville
65. Yorke H (1804) Redhead letters from France in 1802, 2 vols. HD Symonds, London
Darwinism Past and Present: Is It Past
Its Sell-by Date?
Michael Ruse
Abstract The year 2009 was the 200th anniversary of the birth of the English
naturalist Charles Darwin, and also the 150th anniversary of his great book, On the
Origin of Species by Means of Natural Selection, or the Preservation of Favoured
Races in the Struggle for Life. No one who takes science seriously would begrudge
Darwin his fame, but there is a major question that is worth asking. Do we honor
Darwin as an important figure in the history of science, but not necessarily as one
whose thinking still speaks to us today? Or are there aspects of Darwinian thinking
that are still important today? Is it possible, desirable indeed, to be a Darwinian in
the sense found in the Origin?
1 Evolution as Fact
I want to go right to the heart of the matter, asking whether in the Origin Darwin
really made a case for evolution for evolution through natural causes and
whether this was successful and lasting. Did Darwin in the Origin give us an
argument to merit taking evolution to be factual, in any reasonable sense of the
term? Did Darwin give an argument that is still central to discussions today? Did
Darwin give an answer that should be compelling to reasonable people? My focus is
more on the fact of evolution rather than its causes, in the Darwinian case rather
than on natural selection, or what today is usually cashed out as differential
reproduction. It is obvious, however, that although ideally one might like to
separate questions about the fact of evolution from questions about the cause or
mechanism of evolution, in practice this is not really possible. This is especially
true in the case of the Origin, for Darwin so often runs the two together. And in
truth, really one would not want to separate the questions entirely. If evolution did
M. Ruse (*)
Department of Philosophy, Florida State University, Tallahassee, FL, USA
e-mail: mruse@fsu.edu

42 M. Ruse
occur (fact) then there must have been some reason why this happened (cause).
Hence, although the focus here is more on fact than on cause, I do not take the
discussion to be irrelevant to the significance of natural selection.
In his Autobiography, written towards the end of his life, Darwin wrote of the
Origin as containing one long argument ([6], p. 140). But what was this argument?
Actually it came in several (at least three) parts. In a letter written a year or two after
the Origin was first published, Darwin made explicit mention of his strategy:
In fact the belief in natural selection must at present be grounded entirely on general
considerations. (1) on its being a vera causa, from the struggle for existence; & the certain
geological fact that species do somehow change (2) from the analogy of change under
domestication by mans selection. (3) & chiefly from this view connecting under
an intelligible point of view a host of facts. (Letter to George Bentham, 22 May 1863
[7]- Vol. 11, p. 433)
Mainly because I and others have elsewhere discussed the first two parts at
length, but also because Darwin (as in this letter) thinks the third part the most
important, I am going to focus on this view connecting under an intelligible point
of view a host of facts. But before I turn to the Origin, I want to set some
background, asking about influences and the methodological authority to whom
Darwin would have turned in making his case.
2 The Consilience of Inductions
One authority above all stands out: William Whewell, formerly professor of
mineralogy at the University of Cambridge, later professor of moral philosophy
and Master of Trinity, writer of textbooks, authority on the tides, and above all for
Darwins purposes a good friend and mentor and an expert on scientific methodol-
ogy [30]. Darwin had known and respected Whewell when an undergraduate, he
was much in Whewells company when he returned from the Beagle voyage (first
directly in Cambridge and then in London through shared involvement in the
Geological Society), and he read and knew of Whewells beliefs about methodol-
ogy. He read twice in the year of publication [33] the three-volume History of the
Inductive Sciences and he knew well the contents of the later (1840) two-volume
Philosophy of the Inductive Sciences from having read a detailed review by the
astronomer-philosopher John F. W. Herschel [18], author of influential discussions
of scientific methodology [17], later noted by Darwin [5].
Whewell came through loud and clear on the issues worrying Darwin. Why
should one accept a causal theory when one doesnt see it in action at least, one
does not see it in action enough to do everything that is claimed? Whewell framed
his inquiry in Newtonian terms. What would the great Sir Isaac Newton have done
and said? Well, we know what he did. He explained everything in terms of the force
of gravity. But precisely what Newton said about this was frankly ambiguous.
Apparently we are to invoke true causes, verae causae. What is a vera causa?
Here Newton was less than helpful, but Whewell was happy to help out.
Darwinism Past and Present: Is It Past Its Sell-by Date? 43
Clearly influenced by the rationalist strain in philosophy (Whewell was much

enthused with the Kantian system), he argued that a true cause is something that
comes at the center of what he called a consilience of inductions ([34], p. 2, p.
230). And what is this precisely? It is a situation where we have a hypothesis about
a cause, which may well not be observed or encountered directly, but which
explains a wide range of empirical facts. In other words, as in a detective story,
the causal culprit is identified not through the act itself (which is generally unseen
and unknown), but through the clues the bloodstains, the footprints, the tobacco
ash (a favorite of Sherlock Holmes), the broken alibi, the motive. The culprit did it,
explains the clues, and conversely (in a feedback argument) the clues convict the
culprit.
Those of us who are professional philosophers of science know that this kind of
argumentation varieties of which have also gone under the name of abduction,
the term of Charles Sanders Peirce [2], and inference to the best explanation,
popularized by the late Peter Lipton [22] has been discussed ad nauseam. Here, I
am going to assume that even though there are debates about why it is a good form
of explanation, it certainly can be a good form of explanation. It can properly
convince you of the truth of the binding premise. Note that we are not now asking
for logical necessity, and note that often supporters of this kind of explanation
demand that (having made the argument) one look for fresh, hitherto-unknown
evidence of its truth. Whewell was strong on this. A real consilience shows it worth
by pointing to unknown facts, perhaps even totally unexpected facts, thus convinc-
ing that it is really about objective reality and not just an ad hoc construal to build a
pretty picture. All of the clues point to one potential culprit and then you find that
(as in The Hound of the Baskervilles) he is the unknown next-in-line to the family
fortune. Now thats convincing!
The young Darwin heard all of this, took note, and put it into practice. Let us now
run quickly through the Origin and show precisely how Darwin put his Whewellian
methodology to work.
3 Darwins Argument
The first part of the Origin sets the scene, introducing natural selection, in part (as
Darwin said in his letter) through the analogy with artificial selection and in part
(also in the letter) by deducing natural selection in its own right. There is also
discussion of subsidiary issues like sexual selection and the ways in which evolu-
tion branches and lines go their different ways (what Darwin called the principle of
divergence). Next come discussion of problems like heredity, all of which clears
the way for the positive treatment that takes us through the main part of the work. In
turn, we take up behavior or instinct, hybridism and the links and gaps between
groups, paleontology and the fossil record, biogeography and the distribution of
organisms, systematics, anatomy, embryology, and ending with a gasp at rudimen-
tary organs.
44 M. Ruse
In the case of instinct we get the explanation through selection of a beautiful

example of what Richard Dawkins [9] has called the extended phenotype. Why is
it that honey bees build hexagonal spaces for their young? Why not squares or
circles or whatever? Through a number of rather ingenious experiments (involving
the use of colored wax to see exactly how and when the bees use their building
materials) Darwin was able to show that this is the most efficient use of the wax and
as strong as you are ever going to get. He also showed that this seems not to be
something that arrived in one instant, but that less efficient insects suggest that this
ability of the honey bees is something that developed gradually. Moving on to the
fossil record, much of the time Darwin was on the defensive, trying to show why it
is that there are so many gaps in the record. But then he started to make the positive
points, for instance about the roughly progressive nature of the record, a clear
indication of descent with modification (to use Darwins phrase for evolution).
Darwin made much of the extent to which one finds earlier fossils in the record,
fossils that look like the combination of very different extant organisms. Lying
behind a discussion such as this was the kind of Germanic thinking that led the
anatomist Richard Owen [27] to his archetypal theory, where organisms within a
group (like vertebrates) are seen as modifications of a basic ground plan or
archetype, what Stephen Jay Gould was to call a Bauplan [14]. For an idealist
like Owen, there was considerable doubt as to whether the archetype was a real
organism, but for Darwin it was always an ancestor.
Biogeography was a winner for Darwin, since it was the denizens of the
Galapagos that set him on the road to evolutionism. Of course, there had to be a
lot of discussion about how organisms could cross large oceans Darwin always
favored rafts and like phenomena rather than now-vanished land bridges but
basically it was gravy all of the way, as Darwin showed how the oddities of
geographical distribution fall away under the gaze of natural selection. The
Galapagos naturally had a starring role, as did the fact that the organisms of islands
tend to resemble their neighbors on local land masses rather than animals and plants
on lands far away.
Darwin moved on through the fields. Systematics makes sense because of
common descent. Anatomy comes to life because of evolution through selection.
Phenomena noted and mysterious since the time of Aristotle the homologies
(as Richard Owen labeled them) between animals of very different lifestyle are
truly inexplicable if one is thinking purely in terms of function. What can be more
curious than that the hand of a man, formed for grasping, that of a mole for digging,
the leg of the horse, the paddle of the porpoise, and the wing of the bat, should all
be constructed on the same pattern, and should include the same bones, in the same
relative positions? ([4], p. 434) But of course they make perfectly good sense
within the Darwinian picture. The explanation is manifest on the theory of
the natural selection of successive slight modifications,each modification being
profitable in some way to the modified form, but often affecting by correlation of
growth other parts of the organisation. In changes of this nature, there will be little
or no tendency to modify the original pattern, or to transpose parts (p. 435).
Embryology, as we draw to a close, was a particular point of triumph for Darwin,

and I suspect may have been a case where Darwin thought he was going beyond his
original consilience and turning up new confirmatory information of a kind that was
unexpected. Everyone knew that the embryos of organisms very different human
and chick frequently look very similar. How can this be? Descent with modifica-
tion, obviously. But then Darwin went into detail, showing how selection and his
analogy from the world of breeding can provide real insight. The Darwinian
argument is that adults have been torn apart by selection in the struggle for
existence. But the young, by and large, are protected from the struggle, so there is
no reason to expect them made different by selection. Turning to the domestic
world, Darwin hypothesized that since breeders are generally interested only in the
adults, we should find the young are far less different than their grown parents.
Checking on horses carthorses versus race horses and dogs greyhounds versus
bulldogs Darwin found his prediction to be correct, even though breeders had
assured him that there was nothing to his supposition.
And so (after some remarks about vestigial organs) we come to the end of the
Origin, with that final flowery passage about entangled banks, singing birds, flitting
insects, and crawling worms. There is grandeur in this view of life, with its several
powers, having been originally breathed into a few forms or into one; and that,
whilst this planet has gone cycling on according to the fixed law of gravity, from so
simple a beginning endless forms most beautiful and most wonderful have been,
and are being, evolved (p. 490).
4 From Then to Now
With the Origin now published and before us, I am going to jump 150 years. It is
well known that, in its day, the Origin had mixed success [28]. Almost overnight,
the world converted to evolution. Natural selection had to wait for acceptance. No
one denied the mechanism outright, but few thought it as powerful as Darwin
argued. It was in fact not until the 1930s, with the mathematical basis of Mendelian
genetics well established, that people generally recognized the importance of
selection and it started to take the place that it still has today. Without further
ado, let us turn to the next question: Is the Darwinian consilience still active today
and does it play the same role as it did in the Origin?
Let us go once again through the range of biological phenomena seeing if and
how Darwinian ideas explain and are in turn given support. Let me say straight out,
however, that if we find ourselves saying no more than was said 150 years ago,
I shall be surprised and disappointed. As Thomas Kuhn [21], above all others,
pointed out, good science is not static. It moves forward solving problems (Kuhn
called them puzzles) while staying true to the basic ideas. The structure and the
spirit may stay the same but if the content is unchanged then truly we have a
problem. Even if Darwin was right, why should anyone care? At least, why should
anyone care other than as a matter of historical interest?
46 M. Ruse
5 The Consilience Today
As with the discussion of the Origin, I am brushing right past such topics as
artificial selection and the ways in which todays biologists attempt to reproduce
natural situations in the laboratory. Also, ways in which one might hope to find
direct evidence of evolution in nature. I will focus exclusively on the argument
across the whole of the life sciences. Start as before with instinct, by which Darwin
very much meant social behavior. In the past half century, this whole topic has
exploded outwards, in theory and in experiment and in study in nature. It has even
been given its own name of sociobiology. Such evolutionary biologists as
William D. Hamilton [15] and John Maynard Smith [23] in Britain and George
C. Williams [35] and Robert Trivers [32] in America offered selection-based
models explaining social behavior and following this theoretical work there was
massive empirical attention. By the middle of the 1970s, the worlds leading ant
biologist, Edward O. Wilson of Harvard, was able to write his magnificent overview
Sociobiology: the New Synthesis [36] and Richard Dawkins of Oxford penned his
popular account The Selfish Gene [8].
Take a first-class exemplar of this kind of work, Wilsons own study of the leaf-
cutting ants of the Amazon, the Atti. They have many castes the large soldiers, the
smaller foragers and then the leaf-cutters, the gardeners who tend the growing of
fungi on the chewed-up leaves, the nursery workers who tend the young, and of
course the massive queen. All told, the ant genus Atta has seven classes of workers.
A key feature of Atta social life. . . is the close association of both polymorphism and
polyethism with the utilization of fresh vegetation in fungus gardening. . . An additional but
closely related major feature is the assembly-line processing of the vegetation, in which
the medias cut the vegetation and then one group of ever smaller workers after another takes
the material through a complete processing until, in the form of 2-mm-wide fragments of
thoroughly chewed particles, it is inserted into the garden and sown with hyphae. ([37],
p. 150)
Why the different castes and why the proportions? Wilson ran experiments
(usually involving removing whole castes) to determine what is the most efficient
way of using resources. If the ants want to get the most bang for the buck, less
metaphorically get the greatest results in the sense of producing new, fertile, gene
bearers for the minimum amount of effort, what role do the various castes play and
is the proportion of one to another the most efficient? This kind of optimality
thinking [26] what would natural selection do to achieve the greatest adaptive
efficiency paid big dividends. What A. sexdens has done is to commit the size
classes that are energetically the most efficient, by both the criterion of the cost of
construction of new workers. . . and the criterion of the cost of maintenance of
workers. I hardly need say that this is way beyond any kind of thinking that Darwin
had, and yet in another sense is completely Darwinian. Most especially in making
the division of labor absolutely crucial. Listen to Darwin in the Origin, speaking of
sterile workers: we can see how useful their production may have been to a social
community of insects, on the same principle that the division of labour is useful to
civilised man ([4], p. 241).
The fossil record always fascinates. Again, in respects, we are far ahead of
Darwin. For a start, we have absolute dates now. The beginning of the Cambrian
540 Mya, for instance. More than this, we have evidence of pre-Cambrian life [20].
To his embarrassment, Darwin had none and gave text-book examples of scientific
adhockery the fossils would be where the oceans are now and, even if we could
drill, the weight above would have squashed them into non-being. Now we have a
pretty good record of life from its beginnings and moreover this is life which starts
simple and gets more complex, precisely as expected. We have lots more transi-
tional fossils, most recently the fish-amphibian Tiktaalik from the upper reaches of
snowy Canada. Or if you want a sequence, then the line leading up to humans is
excellent [25]. Think about Lucy, Australopithecus afarensis, about 3.5 million
years old, less than 4 ft tall, on her hind legs although not as good a walker as we
(but probably a better climber), with a chimpanzee-size brain 400 cc as opposed to
our 1,200 cc [19].
Much contested is the question of whether Darwin was right in seeing (as he did)
the history of life as a smooth process, leading from one form to another. For
Darwin, this was part and parcel of his commitment to adaptation. Rapid changes
would take organisms out of adaptive focus. He had no place for hopeful
monsters. Famously, the late Stephen Jay Gould together with fellow paleontolo-
gist Niles Eldredge argued that the true course of history is much more one of stop
and go periods of relative evolutionary inaction (stasis) broken by times of rapid
change [12]. Much ink has been spilt over this theory of punctuated equilibrium,
including the question of how smooth a Darwinian gradual change must necessarily
be. As with so many controversies, there is some truth in the Gould-Eldredge
hypothesis but nothing like as much as was claimed at times by enthusiasts [31].
Rates of change do vary, but then what would you expect? At the micro-level,
change is probably going to be pretty gradual, although precisely what one might
mean by pretty gradual can be contested. Dimensions might be distorted drasti-
cally, very quickly, with major implications. Richard Dawkinss [10] example is of
a 747 Jumbo Jet being stretched, virtually overnight from the original form, to make
for more passengers. We might get a change in (say) reptilian form pretty quickly
this way. It is however unlikely that we would get something like the overnight
change from reptile to bird. And of course the bird-reptile Archaeopteryx shows
that this did not happen. (The Archaeopteryx fossils started to emerge in the early
1860s, and soon found their way into the Origin.)
Biogeography was one of Darwins strongest supports for his theory and it
continues to be so today. But here the changes have been truly staggering, because
we are now in the era of plate tectonics, leading to continental drift. Although no
one denies the powers of bird-dispersal and the like, there is no need of fabulous
hypotheses and especially not fabulous hypotheses about now-missing land links
because we now know that the continents move around the globe on massive
plates, things which arise out of the earth, move in a stately fashion across the
surface, and finally return into the bowels in an almost Wagnerian fashion. You can
48 M. Ruse
explain for instance why it is that the Permian reptile Lystrosaurus, a slug of a brute
if ever there was one, can be found in Africa, India, and Antarctica. It did not go
traveling under its own volition. It stayed where it was and let the moving plates do
the hard work [29].
Systematics, morphology, embryology they start to bring the consilience to a
close. One could write full-length essays on each of these topics. Change and yet
adaptation due to natural selection is the story. Systematics was transformed in the
1970s, thanks to the coming of cladistics, the system of classification based on the
somewhat idealistic system of the German biologist Willi Hennig [16]. Although
there were some (rather extreme devotees of the philosophy of Karl Popper) who
questioned whether classifications need at all reflect history, generally it was (and
still is) agreed that cladistic classification is firmly evolutionary. It is about paths or
phylogenies. Less obvious, particularly at first, was whether it was always very
Darwinian. For instance, change in a line without branching would not be (could
not be) recorded. As methods have become more sophisticated and refined, how-
ever, particularly thanks to the coming of molecular techniques, it does seem that
the gap between cladistic practices and results and Darwinian processes have come
closer together.
As it happens, like paleontology, morphology has been very controversial and
for much the same or at least related reasons. Stephen Jay Gould argued that form,
as in homology, is basic, and function, as in adaptation, comes afterwards. Certainly
(according to him and co-writer Harvard geneticist Richard Lewontin) it does in
many cases. Most particularly, because form puts its mark on organisms, much that
we find has little or no relation to utility. Often, seemingly important adaptations are
spandrels, that is to say by-products of other characteristics, which may or may
not themselves be adaptive. Spandrels are the triangular spaces at the tops of
columns in medieval buildings. Frequently, as in the church of San Marco in Venice
with beautiful decorations all over them. The design is so elaborate, harmonious,
and purposeful that we are tempted to view it as the starting point of any analysis, as
the cause in some sense of the surrounding architecture. But this is to reverse cause
and effect. The system begins with an architectural constraint: the necessary four
spandrels and their tapering triangular form. They provide a space in which the
mosaicist worked; they set the quadripartite symmetry of the dome above ([14],
p. 148). Warming to his theme, arguing that much of the living world is non-
adaptive, Gould sneered that those who seek adaptation are too often like
Dr Pangloss in Voltaires Candide, forever seeing value or use when there is
none the best of all things in the best of all possible worlds. According to
Gould, Darwinians spin just so stories, at one with the fairy tales of Rudyard
Kipling, who tells us that the elephant has a long trunk because a crocodile pulled it!
Yet that the organic world is marked by adaptive complexity is, despite Goulds
rhetoric, simply a commonplace. Since Darwin, again and again the most outland-
ish feature has been shown tightly tied to reproductive utility. Take the triangular
plates running along the back of the Stegosaurus, a monstrous dinosaur discovered
in the American West in the decades after the Origin. What can they be for? Some
have suggested sexual attraction; others have proposed that their utility comes as
weapons fighting or defence. The popular view today is that they are for tempera-
ture control, for heating and cooling [13]. The brute needed to warm up its blood
first thing in the morning and then, at mid-day, particularly with the heat produced
by its herbivorous diet, it needed to get rid of the heat. The plates are just like the
fins one finding in electrical cooling towers, where they likewise are used for heat
transference. Supporting the hypothesis, the fossil evidence is that the blood was
moved from the main body (and back again) in just the efficient way one would
suppose were heat transference the goal in mind.
As sociobiology has transformed the study of instinct, so the new field of
evolutionary-development has transformed embryology. The molecular biologists
have moved in, giving new insights and empirical discoveries, and offering new
theories. This is no bad thing, if only in the sense that the neo-Darwinian synthesis
of the 1930s followed the population geneticists as regarding development as
something of a black box you have genes (genotypes) on the one side, and you
have physical features (phenotypes) on the other side, and no one cared much about
what went on in between. Pigs in, sausages out, and no questions please. Now this
connection is brought into the sunlight, as biologists trace in detail the paths from
the nucleic acids to the performing physical organisms. And some of the
discoveries are simply staggering, again and again underlining the fact that the
evolutionary consilience is so powerful in the ways that it points to discoveries
altogether unimagined when the scientists started working and theorizing. A beau-
tiful example centers on molecular homology. The late ornithologist and systema-
tist, Ernst Mayr, writing in Mayr [24] about homology, having described it as the
best of all kinds of evidence for evolution, warned non-biologists not to look for
impossible isomorphisms between organisms as different as humans and fruitflies.
You are not going to find them. One hopes that wherever he is, looking up or
looking down, Mayr now celebrates the fact that at the molecular level there are the
most incredible homologies between humans and fruitflies. It turns out that the
genes that control development, the Hox genes, are nigh identical between the two
organisms and with many others too [3].
Does this threaten Darwinism? Is natural selection downgraded and the hunt on
for a new, post-Darwinian theory? Why one should feel this need, why any of this
should impact negatively on natural selection, is hard to imagine. We now know
that organisms are built on the Lego principle the same building blocks can make
the White House or King Kong, a fruitfly or a human. But natural selection has no
less of a role. Going back to earlier discussion, it is true that evo-devo shows that
often you can get fantastic changes by altering what you have rather than by starting
anew the stretch 747 but selection is no less important if the plane does not fly,
then it is of no use. From Thomas Henry Huxley on, there has been a downplaying
of adaptation, especially by bench biologists who never see organisms alive and
well, in their native habitats. But organisms do flourish, in their native habitats, and
if biologists forget this fact, if they forget it is one thing to make an organism but
then it must succeed in lifes struggles, they will never get the full story of
evolution.
50 M. Ruse
6 Conclusions
Darwinian evolution is a theory and a fact! We still have so identifiably the theory
of the Origin. The consilience was crucial and effective back then; it is crucial and
effective right now [11]. And yet, as in the best kind of science, the story is far
richer than one of simple identity. The pre-Socratic philosopher Heraclitus said:
You cannot step into the same river twice. Everything changes. The pre-Socratic
philosopher Parmenides said: How could what is perish? How could it have come
to be? For if it came into being, it is not; nor is it if ever it is going to be. Nothing
changes. It was the genius of Plato to combine these two into one synthetic system
and how right he was to do so. Everything has changed since the Origin. Nothing
has stood still. Thank goodness! And yet nothing has changed. Darwins argument
rules triumphant. Thank goodness! In the history of science as in the world of
organisms, all is evolutionary change and yet continuity and harking back to
earlier successful forms. The analogy I like is with the peoples car, the
Volkswagen, of late 1930s Germany. Today we have the Beetle, the Bug. Not one
item of Dr Porsches brilliant design can be found in todays car, and yet the Beetle
is so obviously the Volkswagen of 70 years ago. I expect that in 50 years time we
will still have the Beetle around. I expect that in 50 years or a 100 years we will still
have the theory of the Origin around. Great, precisely because it does not stand still,
but remakes itself and grows and changes by virtue of the fact that it gives such a
terrific foundation. Is Darwinism past its sell-by date? Not by a long chalk yet!
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Evolutionary Theory and Philosophical
Darwinism
Paolo Casini
Abstract After the early reactions of the scientific community to Origin of the
species, Darwins hypothesis was widely discussed by a growing number of
professional historians and philosophers as well. This paper provides a short survey
of historical research into the pre-Darwinian scenery of biological transformist
intuitions of the so-called forerunners. This is an essential link to the following
outline devoted to the divergent views of five relevant philosophical interpreters of
the evolutionary theory: Spencer, Huxley, Haeckel, Nietzsche and Bergson. The
intricacies of the Darwin-Spencer relationship are a necessary prelude to Huxleys
well-known Darwinian orthodoxy. In a way, Spencers monistic metaphysics was a
no mans land for friends and foes of Darwins Darwinism. Haeckel, naturalist and
philosopher, translated Evolution into a systematic speculative Weltanschauung,
while Nietzsche was first influenced by Darwinism and Spencerism and later
rejected both. His Uebermensch myth was accompanied by an attempt to develop
a biological-speculative basis for psychology. Bergsons general critique of the
experimental method and of the system of Spencer introduced his reinterpretation
of evolution as a creative elan vital exclusively known through the inner
perceptions of time, self-consciousness and intuition.
1 Philosophical Darwinism: A Short History
Four decades of controversy concerning evolution had elapsed, and Darwins

Darwinism was eventually accepted, when John Dewey, in his essay The Influence
of Darwinism on Philosophy (1909), remarked: The exact bearings upon philoso-
phy of the new logical outlook are, of course, as yet, uncertain and inchoate.
We live in the twilight of intellectual transition [1]. The transition towards
P. Casini (*)
Department of Philosophy, University La Sapienza, Roma, Italy
e-mail: caspal@tin.it

54 P. Casini
evolutionary logic, according to Deweys subtle analysis, expelled from biology,

and from philosophy as well, all ideal archetypes, the concepts of design and
finality, and destroyed the philosophic idol of eido (or species). But, if we consider
the transition crisis from a historical viewpoint, its deeper roots should be men-
tioned. In fact, a variety of earlier, pre-Darwinian evolutionary intuitions had
already undermined the Biblical myth of created species and the Aristotelian-
Scholastic taxonomy of class-genus-species became dubious. Geology and the
study of fossil remains suggested alternative solutions of the mystery of
mysteries. The great uniformitarian geologist Charles Lyell, a historian in his
own right and one of Darwins authorities, was well aware of this historical
background when he refuted Jean-Baptiste de Lamarcks Philosophie Zoologique
in volume II of his Principles of Geology (1832). Other contemporary historians
tried to put the reverse question by asking: to what extent was Darwin influenced in
his turn by the philosophers who had foreseen the problem of transformism with all
their implications? Darwin wrote: I was as ignorant as a pig about his subjects of
history, politics and moral philosophy [2]; but, as everybody knows, he was
familiar with the works of the Reverend William Paley, William Herschel, William
Whewell, Robert Malthus, James Mackintosh, Herbert Spencer, and a number of
quotations in his youthful Notebooks show that he was impressed by certain themes
of the empiricist philosophies of David Hume, John Stuart Mill, and Monsieur Le
Comte [3]. In the Historical Sketch devoted to his precursors, he only mentioned a
second-hand Aristotelian topic and then proceeded by quoting naturalists like
Buffon, Lamarck, Geoffroy Saint-Hilaire, his grand-father Erasmus Darwin and
30 more pioneers of evolutionism, including Herbert Spencer [4]. Just after the
publication of On the Origin of Species and of The Descent of Man, some scholars
rediscovered the scattered intuitions of the Presocratics, the anti-creationist cos-
mogony contained in Book V of Lucretius De rerum natura, some writings of
Eighteenth-century French materialists like Demaillet, Lamettrie, Maupertuis,
Diderot and others [5]. The uninterrupted tradition of these old and recent
authorities was considered as alternative to the myth of biblical creation, and they
were read as anticipators of certain features of both Lamarckism and Darwinism.
When two pioneering historians of biology, Emil Radl and Adolf E. Nordenskjold,
outlined the genealogy of the sciences of life, the research concerning the
forerunners of Darwin was a growing scholarly industry.
By 1910 the Darwinian revolution was already considered as a significant
chapter of the history of culture. The new archives of the history of ideas were
opened by the comparative researches of Arthur Lovejoy, who brought into focus a
full genealogy of pre-Darwinian intuitions in Maupertuis and Kant, Herder and
Schopenhauer, Buffon and Lamarck. In the years 19091911 he studied old and
new evolutionist cosmogonies ranging from the anonymous Vestiges of the Crea-
tion (1844) to Levolution creatrice (1907) of Emile Bergson [6]. Moreover, in his
classic The Great Chain of Being, Lovejoy outlined the course of the idea of the
scala naturae in the history of western thought from Plato to Schelling [7]. His
pioneering work was an outstanding contribution to the philosophical background
and to the pre-history or rather to the anti-history of evolutionism. A rather
Evolutionary Theory and Philosophical Darwinism 55
different background was outlined by Ernst Cassirer who, a few years before his
death, wrote the concluding chapters of his Erkenntnisproblem (1940), devoted to
the Darwinian revolution [8]. In this book a short account of Darwins theory is
preceded by an analysis of the concepts of Form, Metamorphosis,
Enwicklungsgeschichte, which were commonplaces in the German philosophical
culture of the eighteenth and nineteenth centuries. As a result, according to
Cassirer, Darwins evolutionary theory appeared to stem from a composite back-
ground including Leibniz metaphysics of living organisms, the biological ideas of
Caspar Wolff, Kants Kritik der Urteilskraft, the Romantic Naturphilosophie, the
morphology of Goethe and von Baer. The role of the idealistische Morphologie in
the growth of an historical view of Nature and Life seemed to be paramount for
Cassirer (maybe he echoed Spencer), but he rejected correctly any comparison
between Darwinism and Hegelian dialectics, a topic which had been discussed
seriously, among others, by Friedrich Engels, Kuno Fischer, and Friedrich
Nietzsche. Essentially, in Cassirers view, Darwins theory of descent was perfectly
in line with some earlier intuitions of the Naturphilosophen. In a wider sense,
Origins of Species deserved a place of high distinction for the epistemologists of
the Marburg, neo-Kantian school. Paradoxically enough, Cassirers historical pedi-
gree of Darwinism involves a curious blunder: in spite of Darwins general critique
of special creations and of teleology, his laws of natural selection and of the
survival of the fittest were interpreted by this historian, in the fullest sense, as
teleological principles. Thus Darwin, as a theorist of the inherited successful
variations favourable to the species, seemed to be a follower of critical finalism,
in particular of Kants theory of regulative teleological judgement in biology
as discussed in the Kritik der Urteilskraft. Finally, the complex history of pre-
Darwinian, speculative evolutionism was fully treated by a Swiss biologist, Emile
Guyenot, in his book devoted to the origins of Lidee devolution (1957), a
landmark of Levolution de lhumanite, the series directed by the historian Henri
Berr [9].
2 Spencer and Darwin on Evolution
These early surveys of the historical-philosophical aspects of the evolution theory

seem to be obsolete today, in comparison to the work-in-progress of recent
historians, whose researches proceed much more analytically into an open field,
ignoring any disciplinary restriction and studying in depth the multifarious facets of
the Darwinian revolution. One of the most interesting topics is the misleading
relationship between Herbert Spencer, the philosopher of Evolution, and Charles
Darwin, the naturalist and rigorous practitioner of the experimental method in
biology. This topic was ambiguous enough to give way to most confusing philo-
sophical, ethical, sociological variants of the evolution theory, obviously including
the so-called social Darwinism. For example, Marwin Harris and Ernst Mayr
formulated opposite views concerning the Darwin-Spencer relationship: Harris,
56 P. Casini
historian of anthropology, emphasized Spencers priority and originality, at least as

a pioneer of the social sciences; Mayr, biologist and historian of biology, radically
minimized his merits [10]. In the meantime, the controversial figure of Herbert
Spencer, who was extremely popular until the end of the nineteenth century,
re-emerged slowly from an enduring eclipse.
In his youth, Herbert Spencer was deeply influenced by the peculiar scientific
and naturalistic culture of the Derby Philosophical Society, where his father was a
leading figure. He was educated as a mechanical engineer, but he was also an
amateurish geologist, palaeontologist, and phrenologist. He was familiar with the
eighteenth-century idea of progress in Nature pervading the proto-evolutionary
views of Erasmus Darwin, naturalist and poet, author of Zoonomia, The Temple
of Nature and The Botanic Garden, who enjoyed a high reputation among the
founding father of the Derby provincial academy [11]. However, Spencer devel-
oped further his philosophical opinions as an autodidact in London, in the intellec-
tual circle of the publisher John Chapman, the director of the radical Westminster
Review. The growth of his reflections about society and politics was variously
influenced by the opinions and theories of his closest friends: George Lewes,
George Eliot, Harriet Martineau, and George Combes, who were strongly
influenced in their turn by the Cours de philosophie positive of Auguste Comte
and by the System of Logic of John Stuart Mill [12]. Spencers first books, Social
Statics (1850) and the Principles of Psychology (first ed. 1852), beared the typical
imprinting of this eclectic background, where also the roots of his much more
ambitious synthetic philosophy are to be found.
Lamarcks evolutionary hypothesis was a most favourite theme of speculation
for the young Spencer since the 1830s [11]. In his Autobiography he postponed in
time his early initiation to the Philosophie Zoologique, affirming that he was first
converted to the idea of the transformation of species in 1840, as a reaction against
Charles Lyells critical exposition of it in the second volume of his Principles of
Geology [12]. He generalized Lamarcks transformation theory into a scheme of
cosmic progress, including the nebular theory of the origins of the universe and
some suggestions of the German Naturphilosophie, that he collected indirectly via
Samuel Coleridge, Thomas Carlyle, and even an obscure Hegelian, Benjamin
Heldenmayer. He was not among the enthusiast admirers of the Vestiges of Crea-
tion, but was initiated by the zoologist William Carpenter to Goethes theory of the
metamorphosis of plants and to von Baers embryology. Spencer declared that
he was deeply indebted to these naturalists:
In respect to that progress which individual organisms display in the course of their
evolution, this question has been answered by the Germans. The investigations of Wolff,
Goethe, and von Baer, have established the truth that the series of changes gone through
during the development of a seed into a tree, or an ovum into an animal, constitute an
advance from homogeneity of structure to heterogeneity of structure. [13]
These last words, in all their vagueness, are one of Spencers earliest statements
of the development hypothesis, the basic catchword of his system: in his views
the homogeneity-heterogeneity scheme had the broad epistemological status of a
law of Nature (in Auguste Comtes sense). In fact, it was the uncritical dogma he
worked out for a lifetime in his System of synthetic philosophy, increasingly
extending and applying it to all the sciences of nature, man and society.
In his article The Development Hypothesis first published in 1852 and in
several other essays written before 1857, Spencer not only outlined his theory of
evolution, but discussed its physico-theological implications [14]. He rejected the
current idea of special creations of the living beings and adopted Lamarcks
explanation of their variations under the direct pressure of the environment, in
terms of adaptation and of an elementary process of survival of the fittest. The
Principles of Psychology contained a most coherent application of the Lamarckian
law to the evolution of human and animal mind.
In receiving the first collection of Spencers papers [15], Darwin praised their
general argument, but distinguished carefully his own professional approach to
the problem of species in the forthcoming Origin:
I have already read several of them [the Essays] with much interest. Your remarks on the
general argument of the so-called Development Theory seem to me admirable. I am at
present preparing an abstract of a larger work on the changes of species; but I treat the
subject simply as a naturalist & not from a general point of view; otherwise, in my opinion,
your argument could not have been improved on & might have been quoted by me with
great advantage. [16]
This letter is usually quoted in order to confirm that Darwin was not indebted to
Spencer. Reciprocally, Spencer was not indebted to Darwin: this is plainly proved
by the chronology of his youthful Essays, and was openly declared for prioritys
sake in his subsequent works and letters [17]. However, when Spencer read On the
Origin of the Species, wrote to Darwin that the book obliged him to change his mind
on a capital point:
You have wrought a considerable modification in the views I held - While having the same
general conception of the relation of species, genera, orders as gradually arising by
differentiation & divergence like the branches of a tree & while regarding these cumulative
modifications as wholly due to the influence of surrounding circumstances. I was under
the erroneous impression that the sole cause was adaptation to changing conditions of
existence brought about by habit, using the phrase conditions of existence in its widest
sense as including climate, food, & contact with other organisms (for general statement of
this view see Essay p. 41 45) But you have convinced me that throughout a great
proportion of cases, direct adaptation does not explain the facts, but that they are explained
only by adaptation through Natural Selection. [18]
In spite of this afterthought, Spencer remained a staunch follower of Lamarck.

He was always convinced that the vera causa of the modification of species was not
to be found in inner or indirect factors, but in the direct impact of the environ-
ment. Darwins answer was magnanimous: Of my numerous (private) critics, you
are almost the only one who has put the philosophy of the argument, as it seems to
me, in a fair way, as a hypothesis which explains several groups of facts [19]. But
2 days later he revealed his wholly different feeling in a perhaps more sincere vein,
rejecting Spencers verbiage: I have just read his Essay on Population - he told
Charles Lyell - in which he discusses life and publishes such dreadful hypothetical
58 P. Casini
rubbish on the nature of reproduction [20]. When Spencers First Principles

(1862) were published, Darwin was far from being fascinated by the grandiose
project. He wrote to his close friend Hooker that the great book left him greatly
disappointed: all words and generalities . . . and I could grasp nothing clearly [21].
Since 1858 Darwin had a changing opinion of Spencer and his philosophy; over
the years his attitude swung between the extremes of praise and sarcasm. On the
other side, Spencer defended the priority, chronological as well as theoretical, of his
general law of development. He wrote to his father in 1864 that Darwins natural
selection is seen to be absorbed in the general theory of evolution as I am
interpreting it [22]. In his bulky Principles of Biology (1864) Spencer quoted
throughout Dr [Erasmus] Darwin about living organisms, but made only a few
references to the grandson Mr. [Charles] Darwin. As for the Darwinian law of
natural selection, Spencer mentioned it only en passant. He reprinted fully in the
Principles of Biology his Theory of Population, and in a footnote appended to this
chapter he pointed up clearly the cleavage he posited between his own principle of
the survival of the fittest and Darwins natural selection:
This paragraph [. . .] shows how near one may be to a great generalization without seeing
it. Though the process of natural selection is recognised, and though to it is ascribed a
share in the evolution of a higher type; yet the conception must not be confounded with
that article Mr Darwin has worked out with such wonderful skill. In the first place natural
selection is here ascribed only as furthering direct adaptation only as aiding progress by
the preservation of the individuals in whom functionally-produced modifications have
gone on most favourably. In the second place, there is no trace of the idea that natural
selection, by co-operation with the cause assigned, or with other causes, produces
divergences of structure; and of course, in the absence of this idea, there is no implication,
even, that natural selection has anything to do with the origin of species. And in the third
place, the all-important factor of variation spontaneous or incidental as we may
otherwise call it, is wholly ignored. [23]
Spencer s semantics is misleading: what he meant in speaking of natural

selection was not exactly Darwins metaphor but, more or less, a new label for
the same deterministic process he pretended to have described in terms of the
evolution of all things from homogeneity of structure to heterogeneity of struc-
ture. Living organisms are subject to the laws of mechanics, physics, chemistry,
thermodynamics; therefore they dont possess any inner or spontaneous tendency
to change. They progress only under the direct stimulus of the environment;
consequently they inherit adaptive acquired characters. In the System, as far as the
transmutation of species is concerned, there is no room for chance or random
variations. Obviously, this means that Spencer did not accept the Darwinian
mechanisms of sexual selection, gradual chance variation, and modification
with descent, and that he had no doubt about the rigid speculative character of his
own neo-Lamarckian biology. At least, he had no real concern for empirical
evidence, and apparently underrated the radical divergence of his a priori method
from Darwins experimental epistemology. But he did not hesitate to pretend that
he shared with Darwin and Wallace the role of co-discoverer of organic evolution.
Darwins reaction to Spencers speculative variant of evolutionary biology is
recorded in another witty and biting letter to Hooker:
I have now read the last No. of H. Spencer [i.e. the Principles of Biology]. I do not know
whether to think it better than the previous number, but it is wonderfully clever, and I dare
say mostly true. I feel rather mean when I read him: I could bear, and rather enjoy feeling
that he was twice as ingenious and clever as myself, but when I feel that he is about a dozen
times my superior, even in the master art of wriggling, I feel aggrieved. If he had trained
himself to observe more, even if at the expense, by the law of balancement, of some loss of
thinking power, he would have been a wonderful man. [24]
We may conclude that the long-debated problem of reciprocal influence is, in a

sense, an idle question. But this is not the end of the story. Rather surprisingly, in
the fifth edition of Origin (1869), Spencers paramount formula of the survival of
the fittest was accepted by Darwin pro domo sua:
We have seen that man by selection can certainly produce great results [. . .] But Natural
Selection, as we shall hereafter see, is a power incessantly ready for action, and is as
immeasurably superior to mans feeble efforts, as the works of Nature are to those of Art.
I have called this principle, by which each slight variation, if useful, is preserved, by the
term Natural Selection, in order to mark its relation to mans power of selection. But
the expression often used by Mr. Herbert Spencer of the Survival of the Fittest is more
accurate, and is sometimes equally convenient. [25]
Furthermore, Darwin first introduced Spencers favourite term evolution in the

1872 edition of Origin and in Descent of Man. As it has been often remarked, these
purely terminological or metaphorical changes did not alter substantially Darwins
long argument in support of the variations of species by natural selection.
3 Chance and Determinism: Spencer, Darwin and Huxley
Spencers theory of evolution was particularly at odds with Darwins theory of

descent as far as the dilemma of chance and necessity was concerned. This highly
significant topic deserves a further reflection, within the philosophical context of
Evolutionism. Spencers faith in determinism was absolute. His Principles of
Psychology a book that Darwin praised at the end of the 6th edition of Origin and
quoted in The Descent of Man - were founded on a rigid deterministic basis. The
analysis of the development of the psyche recapitulates the evolutionary process: in
animals and humans alike, reflex action, instincts, memory, reason, are but conse-
quential stages of a passive adaptation to the environment. This means, according to
Spencer, that the mechanistic pattern, extended to human will and mind, destroys
the illusion of free-will. The negation of chance and fortuitous events is, by
definition, an obvious corollary of the general law of evolution from the homoge-
neous to the heterogeneous. The causal chain of psychical events is continuous and
without breaks, and the Laws of Nature are no empirical or contingent rules, but
normative, absolute, teleological principles. This means that in the System of
Synthetic Philosophy the Evolution of Life is not an open-ended process. Notwith-
standing his positivistic refusal of metaphysics and of final causes Spencer
reaffirmed paradoxically that cosmic evolution aims to a supreme goal, and
natural evolution culminates in social evolution: it is a work-in-progress pointing to
60 P. Casini
perfection [26]. Closing the circle, Spencer returned back to a semi-providential

worldview. In his systematic style, he reproduced some features of the pre-Darwinian
theologizing ideology of cosmic progress which had been shared by both Erasmus
Darwin and Robert Chambers, the author the Victorian bestseller Vestiges of the
Creation. Probably Charles Darwin had in mind all these aspects of the Synthetic
Philosophy when, in an often-quoted passage of An Autobiography, he
summarized in retrospect his mixed feelings concerning Spencer.1 Instead,
Darwins methodical strategy in face of the chance/determinism dilemma was
flexible and undogmatic. He remarked in Origin that when we speak of chance
variations we should keep in mind that this incorrect expression serves to
acknowledge plainly our ignorance of the cause of each particular variation [28].
Such a bracketing of the causal problem did not mean, however, a definitive
suspension of judgement concerning the verae causae of the variation phenomena.
Upon second thoughts Darwin worked out his pregnant metaphor of the skillful
architect who realizes his project following a rational, teleological design by
means of scattered chance-produced materials [29]. But in metaphysical sense the
interaction of chance and necessity is a sort of quibble: I feel most deeply as he
wrote to Asa Gray - that the whole subject is too profound for the human intellect.
For any attempt to solve the dilemma rises a difficulty as insoluble as it is that of
free will and predestination [30]. In the Autobiography, recapitulating the
fluctuations of his beliefs in creation, Darwin hints briefly to the skeptical-
materialistic doctrine of blind chance or necessity as opposed to his youthful
belief in Paleys physical theology:
When thus reflecting I feel compelled to look to a First Cause having an intelligent mind in
some degree analogous to that of man; and I deserve to be called a Theist. This conclusion
was strong in my mind about the time, as far as I can remember, when I wrote the Origin of
Species; and it is since that time that it has very gradually with many fluctuations become
weaker. But then arises the doubt. Can the mind of man, which has, as I fully believe, been
developed from a mind as low as that possessed by the lowest animal, be trusted when it
draws such grand conclusions? May not these be the result of the connection between cause
and effect which strikes us as a necessary one, but probably depends merely on inherited
experience? [31]
1
Herbert Spencers conversation seemed to me very interesting, but I did not like him particu-
larly, and did not feel that I could easily have become intimate with him. I think that he was
extremely egotistical. After reading any of his books, I generally feel enthusiastic admiration for
his transcendent talents, and have often wondered whether in the distant future he would rank with
such great men as Descartes, Leibnitz, etc., about whom, however, I know very little. Nevertheless
I am not conscious of having profited in my own work by Spencers writings. His deductive
manner of treating every subject is wholly opposed to my frame of mind. His conclusions never
convince me: and over and over again I have said to myself, after reading one of his discussions,
Here would be a fine subject for half-a-dozen years work. His fundamental generalisations
(which have been compared in importance by some persons with Newtons laws!)which
I daresay may be very valuable under a philosophical point of view, are of such a nature that
they do not seem to me to be of any strictly scientific use. They partake more of the nature of
definitions than of laws of nature. They do not aid one in predicting what will happen in any
particular case. Anyhow they have not been of any use to me [27].
The earliest philosophical vulgate of Evolutionism pieced together uncritically

Darwin and Spencer. The initiates were troubled by the impossible task to concili-
ate their divergent views about the chance/determinism dilemma. During the last
two decades of the nineteenth century the problem was discussed in the context of
the dispute about the laws of nature by philosophers of science of various schools:
critical positivists, materialists, contingentists, neo-idealists. This wider scene
should be kept in mind in a brief survey of the solutions proposed by a few
preeminent theorists of philosophical evolutionism who were at work between the
end of the nineteenth and the beginning of the twentieth centuries, like Huxley and
Haeckel, Nietzsche and Bergson.
Thomas Huxleys attitude was extremely significant. His own research as a
marine experimental physiologist, his friendship with Spencer, his fierce rebuttal
of Vestiges of the Creation, his pioneering and enthusiastic reception of Origins,
followed by his stubborn militancy of Darwins bulldog, finally his straightfor-
ward rejection of Spencers synthetic philosophy and of the politics of administra-
tive nihilism, seem to be as many uncertain steps into a contradictory path. And yet
Huxley followed a coherent line of thought in his reflections concerning the
freedom of the will, liberty and necessity, chance and determinism, particularly in
his commentary of David Humes critical account of these problems [32].
According to Huxleys mature epistemology, Darwins theory of descent was a
successful working hypothesis, that should wait for proof by future experimental
tests; but he was also convinced that no final evidence will ever be able to convert
natural selection into a teleological, absolute, necessary and universal law of nature,
taken in Spencers (and Kants) normative sense.
In his book on Hume, Huxley fully agreed with his authors conclusions about
freedom and necessity, and the empirical connection of the cause-effect
relationships. He extended Humes analysis to the phenomena and laws of living
creatures. He thought the evolutionary biologist, like the astronomer and the physi-
cist is, by definition, unable to transcend the contingent horizon of the phenomena,
and to formulate laws of absolute certainty. Natural selection and the survival of the
fittest are no exceptions to the probabilistic standards of experimental philosophy.
A further inquiry into the substance of matter, living or non living would also
mean to absolutize, like Spencer did, the laws of nature, and in particular to relapse
into the abstractions of Scholasticism and the traditional dichotomies of dualistic
metaphysics: chance and finality, spirit and matter, creation and predestination. In
short Huxley, a skeptical, methodical materialist, did reject systematic material-
ism, and even teleological evolutionism, as a dangerous basis of metaphysical
dogmatizing.
Darwin firmly excluded the final causes Bacons barren virgins from the
transmutation of species. Huxley fully agreed, yet recognized that the descent of
man assures to the human mind a dominant place in nature: but even mind, this
culminating epiphenomenon of the transmutation process is subject, like any other
phenomenon of nature, to the standard causal sequences. The theological illusion of
freedom of the will stays and falls with the ideas of design and finality. But for
Huxley this did not mean a rejection of the ethics of responsibility. On the contrary,
62 P. Casini
in his essay On Evolution and Ethics, without mentioning Spencer, Huxley

energically reacted against the formers postulate of a deterministic continuity
via the adaptation-in heritage scheme of nature, ethics, and society. The ethics of
sympathy which Darwin adopted in his analysis of the animal and human
behaviour, enabled Huxley to counter the brutal struggle-for-life rule by the
observable spontaneous growth of parental and social instincts, in man as well as
in animal species. From the viewpoint of an anti-Spencerian and pro-Darwinian
analysis of the descent of man the ethical progress of society depends not on
imitating the cosmic process. . . but in combating it [33].
4 The Case of Ernst Haeckel
The problem of chance was one of the main philosophical issues of the discussions
concerning the evolutionary hypothesis in the United States and on the Continent.
The German zoological, morphological, embryological, and anthropological cul-
ture was particularly receptive to darwinism thanks to the timely translation of
Origins by Heinrich Bronn, and the immediate pro-Darwin support which was
assured by the young naturalist Ernst Haeckel [34]. The speculations of Goethe,
Schelling, Schopenhauer and the Romantic Naturphilosophie on living nature
animals and vegetables undermined the purely taxonomic approach to the species
by advancing new speculative hypotheses like Urph anomenon, Metamorphosis,
Entwicklung. An obvious semantic shift occurred in the new usage of the last term:
Bronns translation of Evolution by Entwickelung was, in a sense, the retranslation
of a well-known German term that had been formulated by Ernst von Baer in 1825.
Ever since his very first writings Haeckel was busy in founding the new evolution-
ary biology on the biogenetisches Grundgesetz, the supreme Law of recapitulation
that he considered as the alpha and omega not only of organic morphology, but of
natural history and of the physical world as a whole. If a single embryo recapitulates
the transmutations of several preceding species and forms, the sequences of all the
variations cannot be but intrinsically deterministic. The morphological changes of
all living beings may be reduced to the universal scientific and philosophic postu-
late of the biogenetic law. Even human conscience, the culminating stage of the
evolutionary process, is included in it. Haeckel agreed with the deterministic
suggestion of Emile du Bois Reymond, who defined the freedom of the will
one of Die Sieben Weltr athsel, the seven enigmas of the world. Haeckel too denied
freewill, but believed that his own biogenetic Law was the only clue which could
resolve this one as well as the other six Weltrathsel.
In his several popular expositions of science von Haeckel outlined a growing
encyclopedic repertory of the natural enigmas that had been already laid open.
However, proceeding in the evolutionary chain from physical to chemical phenom-
ena, from inert to living matter, from spontaneous generation to the tree of life, from
mammalians to anthropogenesis, in order to explain all variation, heredity and adapta-
tion phenomena von Haeckel introduced an ever-growing range of ad hoc laws,
never thought of by Darwin. Finally a lot of laws, underlaws, counter-laws,

reduced the biogenetisches Grundgesetz into a fragmentary network of contingent
rules, leaving considerable room for the random, unpredictable events of natural
selection, much akin to Darwinss chance variations. In this sense Haeckel, like
Darwin, saw evolution as an open-ended process, and his researches helped in
re-opening the problem still unknown causes of natural selection. Von Haeckel
tried to extend even to the history of scientific ideas his biologisches Grundgesetz:
he recapitulated, so to say, the history of the Abstammungslehre as a cumulative
progression from Empedocles of Agrigentus to Giordano Bruno, from Goethe to
Kant, from Oken and Lamarck up to Darwin, including the great philosopher
Herbert Spencer. According to him, the whole story culminates in a new monistic
and deterministic faith in Nature, based on the revival of a neo-Spinozistic and
pantheistic phoenix through the Romantic Naturphilosophie. The Nat urliche
Schopfungsgeschichte (1868) was the propagandistic tool of a post-religious, eclectic
and secular ideology: a kind of Wagnerian symphonic poem stuffed with striking
dissonance and fascinating experimental suggestion. Haeckel had many admirers
and many detractors, and helped in transforming Darwinism into a Weltanschauung
which enjoyed an ephemeral world-reputation.
5 Romantic Evolutionists: Nietzsche and Bergson
The evolutionary theories of both Darwin and Spencer, blended into one and the
same ideology of Evolution cosmological, biological, and social influenced in
depth the rhapsodic philosophizing of Friedrich Nietzsche by introducing into it a
sign of contradiction. Ever since 1870 he assimilated the ideas of natural selection
and survival of the fittest through the writings of some German commentators:
David Strauss, Paul Ree, Karl Wilhelm Nageli. Soon afterwards Nietzsche read
extensively Spencers works and reacted against the philosophers sociological and
ethical doctrines. He began by pointing out Spencers complicity with the average
gregarious morals of the Christian tradition, or with certain features of Spinozas
Ethics. In his devastating satire of the German philistine David Strauss, Nietzsche
inveighs against the whole descent of man from the earlier animal stage up to that
of the cultivated philistine [35]. Elsewhere he scoffs at the myth of the creation of
Adam, and at the descent of human beings from such a sublime origin, now
made futile by the forbidding image of the most self-conscious ape grinding her
teeth [36].
Nietzsche accepted the descent of man from lower creatures as a true but
murderous theory, and stressed in an ultra-darwinist mood the ancestral
instincts, the aggressive pulsions, the war and competition among all living species.
His personal conception of the struggle for life and of the survival of the fittest
echoed the violent, lawless, hypothetical state of nature of Hobbes and
Schopenhauer, and wholly denied the tribal and parental instincts shared by both
man and animals, including the moral springs of sympathy and solidarity which
64 P. Casini
Darwin commented upon in The Descent of Man. Nietzsche eventually reproached

the evolutionary naturalists with their inconsistency; in fact, they showed their
incapability to foresee the moral and social consequences of the hypothesis.
In several aphorisms of Daybreak (1880) e The Gay Science (1886)
Nietzsches devastating analysis of false moral values is fully deterministic, and
free will is denied, not only as a vain theological postulate, but as a self-deceptive
mask, whereupon the traditional maxims of virtue and merit, individual responsi-
bility and conscience are founded. On the other side, echoing in his own fashion
the epistemological critique of the contemporary contingentist philosophy,
Nietzsche justifies his skeptical attitude by radically denying any firm order in
Nature, rejects the scientists faith in a naive principle of causality, and the
postulate of necessary causation as well. Thus the experimental method, paradox-
ically including even the evolutionary hypothesis, was no more than a deceitful
source of pseudo-truths.
The Nietzsche scholars have traced in the minutest details his readings and the
various stages of a tortuous itinerary culminating in the Anti-Darwin invectives
of the years 18871889 [37]. The Zarathustra prophet not only blent together
Darwins and Spencers ideas, but translated into commonsense psychological
and moral terms always within a neo-Lamarckian context the pros and cons
of the German evolutionary philosophers concerning topics like the instincts of
survival, the adaptation to environment, the inheritage of useful variations, intra-
specific competition and so on. As a result, Nietzsche sketched in his notebooks an
idiosyncratic, non-scientific and deeply emotional version of individual and social

evolution, which eventually became a typical feature of the Ubermensch myth.
In his Genealogy of Morals Nietzsche identified Spencers instinct of self-
preservation with the Spinozistic maxim of the perseveration of all things in their
own being. He also criticized Spencers principles of adaptation and inheritance of
the acquired characters, misunderstanding it as it clearly appears from the context
as a mere moral or psychological tendency, a second-class activity, a mere
capacity of reacting:
This definition, however, fails to realise the real essence of life, its will to power. It fails to
appreciate the paramount superiority enjoyed by those plastic forces of spontaneity,
aggression, and encroachment, consequently the sovereign office of the highest functions
in the organism itself [. . .] One remember Huxleys reproach to Spencer of his adminis-
trative nihilism but it is a case of something much more than the administration. [38]
Clearly, Niezsches skepticism concerning the experimental method was the

clue behind his bizarre translation of the biological laws, and of scientific research
as well, into a misleading philosophical language, a rather Spencer-like by-product
of Darwins working hypothesis. In so doing, Nietzsche tried to reaffirm the priority
of pure speculation against the positive knowledge founded on the natural sciences.
He even indulged in a kind of sociobiological explanation of the wrong face of
English Darwinism:
That todays natural sciences have become so entangled with the Spinozistic dogma (most
recently and crudely in Darwinism with its incredibly one-sided doctrine of struggle for
existence) is probably due to the descent of most natural scientists: in this regard they
belong to the people, their ancestors were poor and lowly folks who knew all to
intimately the difficulty of scraping it. English Darwinism exudes something like the stutty
air of English overpopulation, like the small peoples smell of indigence and overcrowding.
As a natural scientists, however, one should get out of ones human corner; and in nature, is
not distress that rules, but rather abundance, squandering even to the point of absurdity.
The struggle for existence is only an exception, a temporary restriction of the will to life;
the great and small struggle revolves everywhere around preponderance, around growth
and expansion and in accordance with the will to power which is simply the will to life. [39]
Nietzsches philosophical reflection fell into a kind of epistemological vacuum:

his vain effort consisted in trying to refute the naturalistic theory of evolutionary
change of living organisms through the process of adaptation, competition, natural
selection etc., and in opposing to it the enthusiastic notion of the Wille zur Macht,
a revised version of Schopenhauers metaphysical myth of the all-transcending
Will. For Nietzsche, in other words, the Will to Power was the new fundamental
Law of Nature superseding the pseudo-laws of positive science. Within the single
organisms as well as in human society, the Will to Power is supposed to subdue to
her peculiar ends the defects and the recessive involutive and degenerative
characters. This is the anti-Darwinian kernel of Nietzsches aristocratic ideology
condemning the human mass, obedient to the biological-spiritual stimulus of the
Wille zur Macht, to be subservient to the prosperity of a higher and stronger human
race [38]. The Anti-Darwin posthumous fragments and some aphorisms of the
Twilight of the Idols summarize Nietzsches supreme grievances against the
Darwin-Spencer doctrine. At the end of his parabola, Nietzsche exploited also
some suggestions, found in F. A. Langes Geschichte des Materialismus and in
the biological theories of Wihelm Roux, in order to support his voluntaristic
metaphysics of the Will to Power, an Ersatz of natural selection. He pretended
that this law represented the triumph of a non-Spencerian psychology or, more
exactly, of the fr
oliche Wissenschaft exalting an elementary force of nature ignored
by positive science. The new myth, with its corollaries the advent of the
Ubermensch, the eternal return were the extreme, pathological manifestations,
of a variant that Lovejoy defined romantic evolution. The visionary exaltation of the
last Nietzsche announced, on the border of madness, the irrationalist and nihilist
drive that was to degenerate from its psychological origins to the dead end of racism
and political fanatism.
A few years later Henri Bergson accepted the challenge of evolutionary biology
on its own ground. He delineated a critique of positive science, attacking the
experimental method from the viewpoint of his direct intuition. He developed a
metaphysical approach quite different from what he called the traditional pseudo-
science of cause, substance or essence. In his beginnings Bergson was imbued by
Spencers synthetic philosophy, but he realized soon that levolutionnisme
spencerien etait a peu pres a refaire. In his Essai sur les donnees immediates de
la conscience he turned upside down both the principles of the positivist theory of
knowledge, and the fundamentals of Kants Critique of Pure Reason, by
introducing a radical dichotomy betwixt our external and internal perceptions
66 P. Casini
of time and space. He tried to demonstrate that the notions and measures of space
and time, as currently used in the hard sciences, are mere abstractions. For
physicists and astronomers deal with standard events which are regular and pre-
cisely predictable, apparently following uniform and deterministic laws. But this
kind of quantitative knowledge ignores altogether the true essence of time,
la duree profonde. Real time may be discovered by way of direct intuition
only, and this is the approach which unveils beyond the fixed abstractions of
the intellect the qualitative flux of conscience. This uninterrupted sequence of
individual, uncomparable, unique instants is the source of human psychology, of
our free will, of the artists fancy, as well as of open morality, open society,
mysticism and faith.
If Spencers endeavour consisted in reducing biology, psychology and the social
sciences to save the deterministic laws of mechanics and physics, on the contrary
the job of Bergsons Introduction a la metaphysique was to clear the way for a
complete redefinition of the evolutionary laws of nature, beyond and outside the
space-time framework of classical mechanics. The aim of pure philosophical
intuition was to dig into the sources of the spontaneous flux of creative energy,
lelan vital. The immediate perception of the most shifty instants, past and present,
of self-consciousness was instrumental even in penetrating from the inside the
growing sequences of the phenomena of organic life and the evolutionary process
of living species.
In Bergsons view, all the gaps of the evolutionary doctrine of Spencer, Darwin
and their followers might be filled, all the problems of the ongoing experimental
research might be resolved from the viewpoint of the creative flux of duration.
Bergsons main endeavour in his Levolution creatrice (1907) consisted in a
painstaking, systematic analysis of the various theories discussed at that time
among neo-Darwinian and neo-Lamarckian biologists concerning the true causes
of variation, natural selection, inheritance of characters, and in re-examining
thoroughly the old dilemma of chance and deterministic laws.
Bergson put on one and the same level both traditional kinds of absolute determin-
ism: the theological doctrines of providential design, and the mechanical-materialistic
axioms of natures necessary laws. Both represented in his view a closed project, and
the arbitrary projection into nature of our own abstract, anthropomorphic so-called
categories. But finality and necessity are not forms given a priori to the
trascendental ego, in Kants sense; they are rather by-products of our practical
action, artificial schemes of our own skills. Bergson, aiming to a third solution,
insisted on the creative power of nature, depending on une cause dordre
psychologique. . . lelan originel de la vie [40]. Thus Bergson returned, by begging
the question, to his starting point, the metaphysical distinctions between time and
duration, abstract and real, intellect and mind. He claimed he had based on such a
foundation une idee plus comprehensive, quoique par la meme plus vague,
du processus evolutif. The belief in the firm existence of the fixed, static forms
of the zoological taxonomies, created or uncreated, is a mere illusion, equally
shared by materialists and creationists. If we are able to uproot this prejudice, our
conscience is able to oppose to the phantom of passive, outside matter his own active
energy, that is a whole cosmogony in motion car lunivers nest pas fait, mais se fait
sans cesse. . . La conscience est essentiellement libre, elle est la liberte meme [40].
Duration, spontaneity, free will, conscience, evolution the catchwords of
Bergsons mystic vitalism were opposed by a whole generation of philosophers
to the positivistic faith in science. Thus a philosophical variant of the evolu-
tionary hypothesis achieved a paradoxical task: Bergsons speculation seemed to
have digested and de-materialized the Darwin-Spencer vulgate of evolution. As
Dewey wisely commented, the exact bearings upon philosophy of the new logical
outlook are, of course, as yet, uncertain and inchoate [1]. He was perfectly right:
the twilight of intellectual transition engendered not only a new science of nature
and a new anthropology, but a most aggressive idealistic reaction against science
and reason.
References
1. Dewey J (1910) The influence of Darwin on philosophy, Popular Science Monthly, July
1909, reprinted in The influence of Darwin on philosophy and other essays in contemporary
thought. Indiana university Press, Boomington, p 9
2. Darwin CR (1958) The autobiography. In: Barlow N (ed). Collins, London, p 55
3. Manier E (1978) The young Darwin and his cultural circle. Reidel, Dodrecht/Boston
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of favoured races in the struggle for life, 3rd edn. John Murray, London, pp XIIXIX
5. Various essays are devoted to these authors in the symposium edited by Glass B, Temkin O,
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(1979) Die Stellung der Biologie in den neukantischen Systemen von E. Cassirer and
N. Hartmann. Acta Biotheoretica XXVIII:218226; Ferrari M (1996) Ernst Cassirer. Dalla
scuola di Marburgo alla filosofia della cultura. L.S. Olshki, Firenze, pp 105106
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10. Harris M (1968) The rise of anthropological theory. Crowell, New York, p 290 ff, 299302;
Mayr E (1982) The growth of biological thought. Diversity, evolution and inheritance. Harvard
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Spencer, Darwin and the question of reciprocal influence. J History Biol 24:3. A general
discussion by various author is contained in a sourcebook: Herbert Spencer. critical assess-
ment. In: Offer J (ed). Routledge, London/New York, 2000
11. Elliott P (2003) Erasmus Darwin, Herbert Spencer, and the origins on evolutionary worldview
in British Scientific Culture, 1770-1785. Isis 94:129; see also Duncan D (1904) The life and
letters of Herbert Spencer. Appleton, New York; Rumney J (1937) Herbert Spencers sociol-
ogy. Williams & Norgate, London; Burrow JW (1966) Evolution and society: a study in
Victorian social theory. Cambridge University Press, Cambridge; Peel JDY (1971) Herbert
68 P. Casini
Spencer. The evolution of a sociologist. Heinemann, London; La Vergata A (1995) Herbert

Spencer: biology, sociology and cosmic evolution, in biology as society, society as biology:
metaphors. In: Maasen S, Mendelssohn E, Weigart P (eds). Kluwer, Dordecht; Taylor MW
(2007) The philosophy of Herbert Spencer. Continuum Books, London
12. Spencer H (1904) Autobiography, 2 vols. I. Williams and Norgate, London, p 176
13. Spencer H (April 1857) Progress: its law and cause. The Westminster Review, vol. 67, Apri1
1857, p 445465
14. Spencer H. The development hypothesis (first published with no authors name in the periodi-
cal "The Leader", 20 March 1852, p 280281), was reprinted in Essays, Scientific, Political &
Speculative, London, 1891, 3 vols, I, p 1-7. Autobiography, cit., and p 7;http://victorianweb.
org/science/science_text/spencerdevelopment_hypothesis
15. Spencer H (1858) Essays: scientific, political and naturalistic. Williams & Norgate, London
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darwinproject.ac.uk
17. For instance, Autobiography, cit., II, p 27
18. Spencer to Darwin, 22 February 1860, ibid., n. 2706b; also Spencers Autobiography, cit., II, p 50
19. Darwin to Spencer, 23 February 1860, The correspondence project, cit., n. 2714
20. Darwin to Charles Lyell, 25 February 1860, ibid., n. 3126
21. Darwin to Hooker JD, 23 June 1863, ibid. n. 4212
22. Quoted in Spencer H, Autobiography, cit., II, pp 99100
23. Spencer H (1852) The development hypothesis. Westminster Rev 57:490, cit.
24. Darwin to Hooker, 10 December 1866, The correspondence project, cit., n. 5300.
25. Darwin C (1869) On the origin of species. John Murray, London, pp 7273
26. Taylor MW (2007) The philosophy of Herbert Spencer, cit., p. 74
27. The autobiography of Charles Darwin, 18091822, with original omissions restored and edited
with appendix and Notes by his grand-daughter Nora Barlow, Collins, London 1958,
pp 108109
28. Darwin C (1860) On the origin of species, p 131
29. Darwin C (1868) The variation of animal and plants under domestication, vol II. Murray,
London, pp 43031
30. Darwin to Asa Gray, 22 may 1860, The correspondence project, cit., n. 2814; and Darwin, The
variation of animal and plants under domestication, cited, p 432
31. Darwin C Autobiography, cit., p 91
32. Huxley TH (1898) Hume, with helps to the study of Berkeley. Appleton, London, pp 214229
33. Huxley TH (1895) Evolution and ethics. Pilot Press, London
34. Richards RJ The Tragic Sense of Life. Ernst Haeckel and the Struggle over Evolutionary
Thought. The University of Chicago Press, Chicago and London, 2008; S. Gliboff, H.G.
Bronn, Ernst Haeckel and the Origins of German Darwinism. A Study in Translation and
Transformation. The MIT Press, Cambridge Mass. and London, 2008
35. Strauss D der Bekenner und der Schriftsteller, 1873 (my translation from Unzeitgemasse
Betrachtungen I }7: www.nietszchesource.org.text/eKGWB DS). See: Stegmaier W Darwin,
Darwinismus, Nietzsche. Zum Problem der Evolution, in Nietzsche-Studien, 16 (1987),
p. 26487; W. M uller-Lauter, Lorganismo come lotta interna. Linflusso di Wihelm Roux su
Friedrich Nietzsche, in G. Campioni A. Venturelli (eds) La biblioteca ideale di Nietzsche,
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Geschichte bei Nietzsche, De Gruyter, Berlin-New York, 2003, pp 238253
36. Nietzsche F Daybreak (1881) } 49 (my translation from the German Morgenr ote, www.
nietszchesource.org.text/eKGWB M)
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NF-1888.14, 123 and 133).
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40. Bergson H (1969) Levolution creatrice. Presses Universitaires de France, Paris 1969, p 248 ff
Struggle for Existence: Selection, Retention
and Extinction of a Metaphor*
Peter Weingart
Abstract Darwins famous phrase of the struggle for existence is taken as a

metaphor. Semantically it has its origins in everyday language but was given a
specific meaning in the context of his theory. Subsequently, the metaphor was
transferred back into everyday use but also had an impact on the historical and
social sciences. Darwins metaphor is one of the most famous cases of this type of
metaphor transfer into the sciences and back. The analysis focuses on the German
context where the phrase was translated into Kampf um Dasein. The Kampf
metaphors role in various different contexts (discourses in the popular press)
reveals the multifariousness of its usage. It reaches from the navy policy to
colonialism and commerce and ultimately played a normative role in Nazi race
hygiene policy.
Darwins famous phrase struggle for existence is taken as a metaphor. Semanti-

cally it has its origins in everyday language but was given a specific meaning in the
context of his theory. Subsequently, the metaphor was transferred back into every-
day use but also had an impact on the historical and social sciences. Darwins
metaphor is one of the most famous cases of this type of metaphor transfer into the
sciences and back.
Friedrich Engels wrote: The entire Darwinian theory of the struggle for exis-
tence is simply the transfer of the Hobbesian theory of bellum omnium contra
omnes and the bourgeois-economic one of competition as well as Malthuss
demographic theory from society into organic nature. After having accomplished
*This contribution is an excerpt from S. Maasen, P. Weingart, Metaphors and the Dynamics of
Knowledge, Routledge 2000, chapter 3. The material for this study was collected in a research
project carried out together with Kurt Bayertz and J
urgen Kroll.
P. Weingart (*)
Institute for Science and Technology Studies (IWT), University of Bielefeld, Bielefeld, Germany
e-mail: weingart@uni-bielefeld.de

70 P. Weingart
this trick . . . it is easy to transfer these theories back from natural history into the
history of society and to claim one had proved this thesis as an eternal natural law of
society. This famous interpretation of Darwins theory has been repeated by
Nietzsche, Spengler, and countless other less prominent scholars to this day. It
also underlies the general understanding of Social Darwinism and contains a
description of the origin and function of metaphors in science. It provides an
example of three different ways in which metaphors are used as media of exchange
of meaning: (1) from everyday language to scientific language, (2) from the
language of one scientific discipline to that of another; and (3) from scientific to
everyday language. The history of science is full of examples of each of these cases,
much has been written on them, and it can no longer come as a surprise that
metaphors reflect the links between scientific, social, and political discourses.
In the first case, concepts used in science have their semantic origins in everyday
language, but are given a specific meaning linked to a theoretical (scientific) context.
This is often the case prior to the invention of artificial terms designed to distance
scientific from everyday language. In the (third) case of the transfer back from
science into non-scientific contexts, or in Engels words, from natural history into
the history of society, the authority of science is carried by or attributed to the
metaphor. It may be (and often is) taken as representing a proven theory, an eternal
natural law, and not just as an illustrative analogy that could easily be replaced by
another one. This recourse to the authority of science, be it implicit or explicit, by no
means requires a precise or justified (from the viewpoint of the respective disci-
pline) application of the metaphor. In fact, the metaphor may be misused and/or
abused, that is, a different meaning in the new context or a normative meaning may
be given to it. In these cases, ideologies, entire world views, Weltanschauungen, are
elevated to the level of scientific truths. This can be seen as an aspect of the process of
scientification. While the large majority of transfers involves single concepts with
very limited connotations, some cases stand out where a few concepts represent a
self-contained theory and develop into a coherent world view.
The account to follow will show (thereby giving further empirical support to
some studies) that Social Darwinism was less prevalent in those contexts where it
was claimed by historians to be strongest. More importantly, it will give evidence to
the phenomenon that the transfer of the metaphor back from science into public and
political discourse entails an immense expansion and diversification of meanings.
It will show further, how the meaning of the metaphor changes as its conceptual
environment changes.
1 Kampf ums Dasein: Popularization and Contexts

of Usage: Darwins Usage and German Translation
In one form the metaphor already appears in the subtitle of the Origin of Species as
the preservation of favoured races in the struggle for life. The third chapter of the
book is entitled Struggle for Existence, and in it Darwin spells out the bearing of
Struggle for Existence: Selection, Retention and Extinction of a Metaphor 71
the struggle for life on the process of natural selection and species differentiation.
Here he also remarks on the status of the term struggle for existence: namely, that
he uses it in a large and metaphorical sense, including dependence of one being on
another, and including (which is more important) not only the life of the individual,
but success in leaving progeny ([8], p. 116). It should suffice here to refer to some
of those studies that have looked very carefully at the origins of the metaphor.
Young traces it primarily to Malthus Essay on the Principle of Population and to
Charles Lyells Principles of Geology stressing the common context of biological
and social theory ([37], pp. 10945; [36], p. 43 et passim). Bowler has shown the
discontinuities in detail in the use of the concept of struggle between Darwin and
Malthus which centres around the difference between intraspecies competition and
interspecies struggle ([7], pp. 63I50). Leaving the interpretation of the finer grain
to the historians of Darwin, one obvious conclusion is that a common term like
struggle was used widely by scientists, was taken from everyday language, and
employed consciously as a metaphor. An important aspect is pointed out by Bowler.
Regardless of the differences between them, Malthus and Darwin both took part
in a period of intellectual change during which the view of nature and society
as a harmonious system gave way to one characterized by the law of struggle ([7],
p. 644).
Since the concern here is with the German career of Darwins metaphor, the
first issue is its translation. Albeit a story all by itself, a few remarks are in place.
In the first translation of the Origin by H. G. Born, favoured races was translated
to vervollkommnete Rassen which actually means perfected races. That did
part of the damage (notwithstanding the correction in the second translation by
Victor Carus in 1867). Darwins interchangeable use of struggle for life and
struggle for existence was translated to Kampf ums Dasein [2]. That implied
even more damage since struggle for life could be translated more adequately into
Kampf ums Leben which would easily encompass both meanings Darwin had in
mind, namely the struggle for survival of a species in a certain environment of
other species under particular ecological conditions, as well as the individualistic
struggle between members of the same species. Kampf ums Leben or perhaps
even more adequately Kampf ums Uberleben would suggest the unconscious,
general struggle for survival in the natural environment while Kampf ums
Dasein assumed the connotation of an individual, conscious, and ultimately
lethal conflict. In this connotation the metaphor is suggestive of the Hobbesian
vision of the bellum omnium contra omnes
The damage was complete with the combination of perfection and the
struggle metaphor to Vervollkommnung durch den Kampf ums Dasein (that
is, perfection through the struggle for existence), because this elevated the
metaphor to the level of the normative and instrumental as was later exemplified
by the eugenicists strategy to perfect the human race by eliminating the
unfit [27].
When speaking about damage we do not intend to leave our neutral position of
observation. Rather, a dividing line between neutral science and value-laden
72 P. Weingart
political use of the Darwinian metaphor is implied. The very fact that the seemingly
simple translation from one language into another could produce the change in
meaning of which it is also known to have occurred in the transfer from one context
(scientific) to another (political) within the same language demonstrates the elusive
nature of metaphor: floating freely between contexts of use.
2 The Kampf Metaphor and the Reception of Darwinism

in the Popular Press
The popularization of the metaphor is illustrated by its career in Germanys

foremost encyclopedia. By 1872 it was already listed as a commonly used term in
connection with Darwinism, but by 1898 it had achieved the elevated status as an
independent entry in the famous Brockhaus Lexikon. Beyond this the picture
becomes more complicated. Studies of the public impact of scientific theories suffer
from the well-known difficulty that usually little is known apart from the circulation
figures of books and journals and more or less well-informed guesses about their
readers. From there it is still a long way to empirical evidence of their impact, that
is, by shaping opinions and attitudes.
With respect to Darwinism, numerous studies with diverging results have been
undertaken [10, 19, 20]. It cannot be claimed here that the methodological diffi-
culties have been overcome, but only that the evidence has been substantiated. We
will first look at a selection of popular science journals, the assumption being that
they had their readership primarily among the Bildungsburgertum and, thus,
among the opinion-leading elite of the German empire during the last quarter of
the nineteenth century. Then, that analysis will be supplemented by a look at a
number of political arenas with which Darwinism in its analogical use has most
often been associated. Here, periodicals, newspapers, and newsletters of pertinent
associations are the basis of the analysis. Finally, we look at publications submitted
to the famous Krupp Prize Competition. While this procedure cannot claim to be a
systematically drawn sample it does cover a broad range of publications supporting
the assumption that we should get a fairly representative picture of the different
discourses in which the Kampf metaphor played a role.
One of the first popular science journals to take up Darwins ideas in Germany was
the weekly Ausland (Abroad). Although sceptical at first, by 1865 it had become a
fervent supporter of the new biological theory. Its self-posed question what makes
Darwin so popular? the journal answered by stating that although the theory was
not proved, and with respect to its claims about the past probably never could be
proved, it represented a unified principle or Weltanschauung. Because of the
theorys analogical character for many neighbouring fields (see [1]), it had brought
about a new age. At the beginning of the 1870s, the journals editor, Friedrich von
Hellwald, like many other authors at the time of the Franco-Prussian War, resorted
to Darwinism as proof that war was a natural law. The theory of the struggle for
existence that applied to societies must also be applicable to the relationship

between peoples. The war between the civilized people (Kulturvolker) of Europe
against the primitive people (Naturv olker) had to be understood as Kampf ums
Dasein.
During the decade of 18701880, Ausland became the chief popularizing journal
for Darwins theory but, as Zmarzlik points out, its glorification of the Kampf ums
Dasein ran counter to the ethical values held by the bourgeois public. Therefore, in
the 1880s, it radically changed its editorial policy [38]. Von Hellwald, who had
been the chief propagator of Darwinism in the journal, gave up his editorship in
1884, complaining to his friend Ernst Haeckel that his efforts had not been very
successful now that the times had turned away from the theory of evolution. Seven
years prior to that he had welcomed the appearance of another new journal that had
taken over the lead in representing the Monist Weltanschauung: Kosmos.
Kosmos was a Monist journal based on the theory of evolution in connection
with Charles Darwin and Ernst Haeckel which first appeared in 1877. As an
explicitly Darwinian journal, it self-confidently anticipated resistance from the
public and, thus, a Kampf ums Dasein (Kosmos 1877, pp. 13). Although at
first embracing the humanities as well as the sciences, the journal shifted its focus to
evolution after 5 years. Most prevalent were discussions on theories of heredity
(notably Weismanns) and selection. In the latter context a wide range of
applications of the Kampf metaphor can be found. From the interpretation of
human history to the theory of all living organisms, from the history of science
and competition between ideas to the struggle between cells, the concept was used
as a modelling device.
One theme which attracted special attention in the journal was also the promi-
nent example of the Social Darwinist debate. This concerned the question whether
Darwins theory was socialist or aristocratic, an issue that arose out of the famous
dispute between Virchow and Haeckel in which Virchow had attacked Darwinism
as a theory that could be adopted by Social Democrats and was, thus, dangerous.
Haeckel had responded by pointing out that Darwinism was aristocratic rather
than democratic and even less socialist because of its principle of the survival of the
fittest which had been translated into victory of the best. During the first 5 years of
its existence, Kosmos was a Social Darwinist journal propagating the aristocratic
orientation of Darwins theory of selection. In transporting political messages with
Darwins theory it was an exception at the time, but did introduce a new dimension
that was to gain ground later on.
In the early 1880s the increasing number of critical voices indicated that the
enthusiasm for the Darwinist fad had subsided. The Kampf metaphor came into
disrepute as a vague and ambiguous term with which so many Darwinists in lack of
a definite category were befogging themselves ([34], p. 355). What to some
appeared as loss of meaning, to others was evidence of success. After 10 years of
publication, the editor of Kosmos announced its demise, turning defeat into victory
claiming that the theory of evolution had not only gained unquestioned authority in
biology but had also become a self-evident and irrevocable prerequisite in other
74 P. Weingart
disciplines (Vetter 1886, p. 92). Contrary to this evaluation, the journal had
obviously lost the Kampf ums Dasein which it had anticipated at its birth.
In other periodicals Darwinism played a much smaller role, but its diffusion and
popularization followed very similar patterns. One of the most widely read journals
at the time, the Deutsche Rundschau, until the turn of the century carried relatively
few articles on Darwins theory and almost none with a Social Darwinist flavour,
with the notable exception of Oscar Schmidts Darwinismus und Socialdemokratie
[25]. More prevalent were philosophical essays on the implications and limitations
of the theory of evolution for ethics, the history of ideas, etc.
Another widely circulated review for politics, literature, and art, Die Grenzboten
(Frontier Messengers), which until the formation of the empire had been the most
influential organ of the liberal bourgeoisie, had by 1871 allied itself with the
National Liberal and anti-Catholic camp of Bismarck allies. At the same time it
was decidedly anti-Darwinian, and by 1900 the journal declared that the twentieth
century did not want to have anything to do with Darwinism. The axiom of the
Kampf ums Dasein was attacked as the same sophistry which had led to imperi-
alism and anarchy ([12], p. 51).
By the early 1890s, authors even commented on the loss of attention to Darwinism.
It had become common knowledge rather than being limited to science, and in this
process it had lost its attraction. Even the theologians had recovered from their
shock. At the same time science, that is, biology, had calmed down, the revolution
had gone into its second generation, specialization set in and for the lay public it
became ever more boring [5]. Throughout this time the debate over the aristocratic
versus the democratic or socialist character of Darwins theory continued,
demonstrating not only the suitability of the theory as a Weltanschauung, but
also its ambiguity with respect to conflicting political positions. The ultimate issue
was whether society was based on a selective struggle for existence with the survival
of the best or whether that struggle led to their degeneration and eventual demise.
Struggle as the agent was undisputed.
The relationship of Marxist and Social Democratic authors to Darwinism must
be seen in the context of social, political, and economic events occurring in the last
quarter of the century. In 1873 the great depression set in and initiated a change in
the political climate, notably a discrediting of political Liberalism and enthusiastic
Manchester capitalism. The dramatic structural changes, industrialization, and
urbanization, brought the working class into politics and created widespread fear
of the socialist movement among the middle class and the aristocracy. While the
defence and legitimation of power interests on the part of the ruling classes is
usually taken as the basis of Social Darwinism, the reception of Darwinism by the
political Left is overlooked.
During the election campaign for the Reichstag in 1877, August Bebel attacked
Prussian militarism in a brochure, pointing out that war and the military system had
degenerating effects on the population. In support of his thesis, he cited a passage
from Haeckels Nat urliche Schopfungsgeschichte which Haeckel himself omitted
from later editions of the book. This episode is typical for the ambiguity of the
Kampf metaphor with respect to its evolutionary or degenerating effects, which
recurs in the shift of arguments from eugenicists before and after World War I.
Typically, Marxists and Socialists denied the interpretation referring to Darwin
that the struggle for existence had to be an inevitable conflict between individuals;
they focused on the negative results of individual competition and pointed to the
superiority of mutual help. Since the difference between the individualistic stance
of Darwinism and the collectivistic philosophy of Socialism seemed to represent a
contradiction between them, a flood of publications was devoted to the defence of
the commensurability of Socialism and Darwinism [3, 16, 18, 26]. Socialist authors
used Darwins theory as proof for the materialist alternative to religious
explanations of mans creation and the role of such explanations in legitimating
the existing social order, notably the monarchy. The scientific authority of evolu-
tionary theory was seen to coincide with the claims to the scientific nature of Marxs
theory of social evolution. Evolution in nature counted as scientific proof that social
progress had to take place with the same law-like necessity. This association of
Socialism with Darwins scientific theory was carried on through the 1890s and
explains the impact on and fascination of Socialist authors like Kautsky and Bebel
with eugenic schemes, in spite of the fact that they were much more explicitly
selectionist and Social Darwinist than the various applications of the Kampf
metaphor.
The same association had already motivated Virchow in 1877 to criticize
Haeckel and the attempts of his followers to include Darwins theory in school
curricula. In his famous talk before the 50th convention of the Gesellschaft der
Naturforscher und Arzte (then the foremost German scholarly association) he
insinuated in calculated vague terms a relation between Darwinism, Socialism,
and the Paris Commune, most likely because he feared that if the same were done
by reactionaries, the relative freedom gained by science after the unification of the
empire could be jeopardized [33]. It did not help Haeckel to insist that if any
political tendency were to be attributed to Darwins theory, it could only be an
aristocratic and hardly a socialist one.
In summary, scanning the range of major periodicals attached to various
political positions, two points stand out. The reception of Darwinism, notably of
struggle for existence, follows the pattern of fashion, that is, it declines as it
becomes more widely diffused; and its use as an ideology is not limited to one
group whose interests it supposedly matches, but rather by many groups who
interpret it as they choose. This usage, both ubiquitous and heterogeneous,
renders it problematic, if not outright false, to assume a specific impact of the
theory be it instrumental or legitimating. However, while the usages are always
specific and, hence, diverse, they are united by the fact that they all take recourse
to the authority of science and natural laws in order to give credibility to their
specific brand of ideas, opinions, and arguments. The pressure toward
scientification thus enforces the usages of (that is, Darwinian) terms and concepts,
which, in the course of this happening, assume their (at times bewildering)
connectivity to a multitude of discursive arenas.
76 P. Weingart
3 Darwinism in Political Arenas
3.1 War and the Military
It comes as no surprise that, given its origin in everyday language, the Kampf
metaphor assumed different meanings in different contexts and at different times.
The most conspicuous example for the ambiguous use of the metaphor came to
dominate public and scientific discussions about the impact of Darwinism: the
association of the metaphor with war between peoples and nations. The connection
may have contributed to the idea that war means progress although even that idea
was not new. A look at the popular scientific literature in which the metaphor
appears shows, that from the late 1860s onward, the metaphor is used extensively
and with increasing radicalism to justify war as a natural principle. After the turn of
the century a radical author like Klaus Wagner writes in Darwinian terms of the
selective struggle for new space (Auslesekampf um Neuland), the great cultural
significance of the selective struggle for existence (hohe Kulturbedeutung des
auslesenden Daseinskampfes), the selection of peoples as natural selection
(Volkerauslese als naturliche Auslese), and propagates explicitly the enslavement
of the lower peoples in what was or was to become the colonies [35].
The identification of the Kampf metaphor with the German war ideology became
a major topic of English and French popular writings during World War I and
served to associate Social Darwinism and German militarism. This helped to put the
blame for both on the specific German use of the term and to overlook two
important aspects: War Darwinism as La Vergata calls it, had adherents else-
where, and it added little or nothing to the militarist defence of war other than a new
metaphorical repertoire and its scientific prestige, as well as giving evidence of the
almoszt unlimited variety of interpretations of Darwins theory [21].
The usurpation of the Kampf metaphor by saber-rattling militarists makes it
seem as if there were no other voices. However, the Social Democrats gave a
different reading of Darwin. For Alfred Dodel-Port, Darwin had not taught that the
evolution of the human race was to be sought in bloody fight between peoples, but
in the care and strengthening of intellectual powers and social instincts [9]. And
Karl Kautsky attacked vulgar jurists, economists and historians who did not have
any clue about the sciences as well as scientists and medical doctors who did not
know anything about the social conditions and developments for giving any struggle
in history the same name, no matter how it originated: to them they were all
Kampfe ums Dasein [17]. Analysts within the peace movement deplored the
popularity that the metaphor and the theory about the struggle for existence had
assumed because of the Franco-German war and attacked the mindless analogy
between nature and human society. The same critical stance can be identified after
the First World War when M uller-Lyer branded Social Darwinism as a cultural
zoology that had become an intellectual plague after the war of 18701871 ([22],
p. 95). The meaning of struggle had become so broad that it no longer differed
from other concepts such as competition.
Thus, one reads about peaceful Eskimos fighting the struggle for existence in the
Arctic, and the millions of Chinese doing the same while living from their garden-
ing, even though it could only be a struggle with their vegetables [11].
These examples shall suffice to illustrate the association of the Kampf metaphor
with war and, more specifically, the ways in which it was applied. There, the
popularity of the metaphor is best explained by its versatility, instigating incessant
debate. A related arena may be examined in which the language of struggle and
Social Darwinism has played a role: the build-up of the German Navy.
3.2 The So-Called Flottenfrage
The build-up of the German Navy became a prominent political issue in the last
decade and a half before the turn of the century. At the time of rising nationalism
and intensifying competition with the leading imperialist power Great Britain,
support of the navy assumed an integrative function for the bourgeoisie, which
was directed against the rising influence of Social Democracy. The Imperial Naval
Office (Reichsmarineamt RMA) set up a special news bureau commissioned to
mobilize mass support with popular and scientific propaganda. At the same time a
number of organizations took part in these efforts, among them the All German
Association (Alldeutscher Verband), the German Naval Association (Deutscher
Flottenverein), and the Free Association for Naval Treaties (Freie Vereinigung
fiir Flottenvertr
age). In all of these, academics, professors, teachers, artists, and
writers played a leading role, and the obvious focus of the RMAs propaganda was
the Bildungsb urgertum.
The speeches, articles, and communications give a fairly accurate picture of the
position of German scholars on the topic, and if Social Darwinist thinking was
present it might be expected to be seen there.
However, a review of the RMAs annual publication Nauticus, of the collection
of speeches and opinion statements from German university teachers on the signifi-
cance of the Flottenfrage, and of a survey conducted by the M unchner Allgemeine
Zeitung, comes to a disappointing result. The Kampf metaphor had been
generalized and segregated from its Darwinian context. The competitive struggle
(Konkurrenzkampf) between classes, nations, peoples, shipping lines, for food and
world markets, was the dominating terminology. An exception was Max Sering
arguing explicitly against the theory of the Kampf ums Dasein, which he branded as
non-scientific and only a pragmatic political argument used by the British Prime
Minister. Repeatedly, and in spite of an abundance of Kampf and Dasein, in this
discourse one can find explicit rejection of Darwins theory as British and not in line
with German interests. Among the 50 odd scholars answering to the survey of the
M unchner Allgemeine Zeitung in 1897 were such renowned names as Karl Binding,
Felix Dahn, Hans Delbr uck, Max Weber, and the Darwinians Ernst Haeckel and
78 P. Weingart
August Weismann. Haeckel was the only one to speak of a German Kampf ums
Dasein next to other major powers in Europe ([2], p. 3).
Thus, Steinberg may be right in a very general sense in saying: The permeation
of German academic thought by Social Darwinian conceptions led to widely-held
views about the nature of politics as a struggle for survival. The state was a living
organism engaged in a life and death battle ([29], p. 108). But here, when analysing
the rhetoric of struggle, it is advisable to heed Kellys point to distinguish between
those who occasionally appropriated a Darwinian phrase or two and those who
undertook a sustained and detailed application of Darwinism to human society. The
first group those vast ranks of saber-rattlers, socialist-baiters, and self-righteous
rich who happened to live in a Darwin conscious age can be called Social
Darwinian only in the loosest sense of the word ([19], p. 102).
It is precisely the loose usage, however which makes the Kampf metaphor such
a general currency and keeps it alive.
3.3 Colonialism, Commerce, and Trade
In another related policy arena, that is, colonialism, commerce and trade, Social
Darwinism is expected to be strongest if one is to rely on its almost proverbial
association with capitalism. Surprisingly, especially in the area of trade, the almost
complete absence of any allusions to Darwinism alongside the self-confident
assertion of the right of the entrepreneur and the denial of equality to the workers
is conspicuous. The only notable exception is Alexander Tille, who called himself
in Haeckelian fashion a social aristocrat and sought to turn Darwinism into a
social ethic along the lines of early eugenic arguments. In his treatise Struggle for
the Planet, he also followed the common racist topos that the lower races were
less efficient and would thus be pushed off the face of the earth [31]. Tille, when
asked by Haeckel to collaborate in the Krupp Prize turned down the offer saying
that to apply evolutionary theory to social theory the most elementary building
blocks had not yet been assembled ([28], p. 10). The discourse on colonialism was
primarily concerned with the imagined dangers of interracial marriages, and well
into the first decade of the new century a biological concept of race in the strict
sense can hardly be detected in colonialist writings. Purity of race was a distant
derivative of the topos of the struggle between nations insofar as it was considered
an important prerequisite for success in that struggle, but race was mostly used in
a humanist connotation prevalent in anthropology at that time.
Friedrich Ratzel, the founder of bio- or anthropogeography, who was later
claimed by the Nazis as one of their own, in his more than thirty monographs and
some 1,240 articles was far more sophisticated on issues such as the origin of the
Aryan race and the demarcation between and the unity of races than was palatable
for them. Thus, he had to be heavily censored in order to serve ideological purposes
(see [13]). Ratzel started out as a Darwinian influenced by Haeckel, but by 1875
had turned against Darwins selectionism, characterizing it as the crude hypothesis

of the survival of the fittest in the struggle for existence [30]. Ratzel is often
identified as a Social Darwinist because of his coining of the term Lebensraum
which became the catch metaphor of colonialist and later Greater German expan-
sionism. Actually, he reinterpreted the Darwinian Kampf metaphor in Malthusian
fashion. To a large extent Darwins often misunderstood and abused struggle for
existence, in his opinion, had to be a struggle between living organisms for space
([24], p. 51).
Thus, wherever one looks before 1890 and the list could be easily extended
various public discourses, which are usually associated with Social Darwinism,
actually show very little, if any impact of it. They do reveal a very loose usage of
Darwinian thought and terminology, that is, of Darwinian metaphors, the most
widespread of which is the Kampf metaphor.
Kellys conclusion is corroborated: Social Darwinism, in whatever form, never
achieved a mass popularity. And the few popular writers who talked a lot about
struggle and race had such tenuous and ambiguous relationships to Darwinism that
it would be absurd to call them Social Darwinists ([19], p. 109).
This is not to be confused with the fact that the Kampf ums Dasein metaphor had
become a highly inflated currency, together with elements of evolutionism and
selectionism, that was being traded in all kinds of contexts with absolutely no
relation to Darwins theory. Popular Darwinism, in contrast to Social Darwinism,
had reached millions of readers by World War I not least through the writings of
Wilhelm B olsche, whose reading of Darwin was that nature not only showed brutal
struggles for survival but also cooperation and love, especially on the higher levels
of evolution [6].
I will skip the story of the Krupp Prize competition of 1900 which asked what
can we learn from the principles of the theory of descent with respect to the internal
development and legislation of states? except to say that the majority of entries
leaned toward a very differentiated and careful adaptation of Darwins theory to
society. Some advocated versions of state Socialism, mechanisms to protect the
weak, and institutions of social welfare. Schallmayers eugenics was explicitly
antiracist and, compared to authors like Tille and Woltmann, was moderately Social
Darwinian. Kelly is correct in his judgement that most of the authors, despite their
biologistic thinking, remained committed to humanitarian values. This is an
important point to keep in mind when drawing parallels between Social Darwinism
and Nazism ([19], p. 108).
3.4 The Normative Turn of the Kampf Metaphor
True as that may be when reading the essays of the Krupp competition, Kelly
underestimates the fact that, after 1900, Social Darwinism appeared in a different
disguise. The relatively vague Kampf metaphor had been translated into a fairly
precise normative scheme of selectionist demographic and eugenic policy.
80 P. Weingart
To Alfred Ploetz, eugenics, or racehygiene as he called it, was essentially a

program to determine an optional organization of the internal Kampf ums Dasein,
that is, the struggle for reproduction and living conditions between individuals of
one race, and of the external struggle, that is, between races ([23], p. 22). In the
racehygienic discourse, coupled with Weismannism and shortly after the turn of the
century with a crude Mendelism, the notion of the fit and the unfit in the Kampf
ums Dasein was translated by 1913 into a normative economic calculation of the
costs of the inferior to the state [15] and by 1920 into the supposedly humane
suggestion to destroy life not worth living (see [4]).
In the context of Nazi ideology, inferior assumed two meanings, both of which
were apparent in the eugenic/racehygienic movement from its outset. To the
anthropologically oriented wing of the movement (then to be called racebiology),
inferior pertained to the non-Aryan races including Jews; to the medically and
social hygiene-oriented wing, its meaning combined supposed hereditary diseases
like alcoholism and forms of behaviour judged to be antisocial.
Although neither Social Darwinism in the narrow sense of the word as it was
represented by the authors of the Krupp competition nor popular Darwinism like
Haeckels or B olsches were officially accepted by Nazi propaganda, this does not
warrant the conclusion that Darwins ideas had lost their effect or were ostracized.
The National Socialist Weltanschauung was shaped from a garbled mixture of
biological holism equating notions of organism, race, and Volk, of evolutionism
and hereditary theory. This mixture was declared to be the truly German Biology
and was peddled by the biological profession as the core of public education. At the
same time, the Nazis themselves made repeatedly clear that National Socialism was
a political and not a scientific movement, and that it drew a sharp distinction
between what science had determined as reality and the various research areas
and theories of individual scientists. Thus, neither Lamarck, Darwin, or Haeckel,
regardless of their importance for the progress of science, nor any of their followers
or opponents, could be equated with the movement.
The leadership strategy of calling on the authority of science without becoming
involved in ongoing debates and academic quarrels, and thereby retaining the
charismatic authority of the leader, had consistently been employed by Hitler
himself. In his notorious book Mein Kampf, a whole chapter was devoted to Volk
and Rasse. Hitler never cited any of the sources he used, but it is obvious that he
must have read much of the literature on racehygiene and race biology at the time
(that is, 1924), as Fritz Lenz later proudly claimed. The chapter opens with an
account of Darwins principle of the struggle of existence and selection, without a
mention of Darwin, without even using the Kampf metaphor in full, and
analogizing the struggle between species and the struggle between races.
Kampf, he concluded, is always a means to promote the health and vigor of the
species and thus the cause for its advancement. What follows is the murky
language of racehygiene and race biology in which the state is to take an active
role in the selection of the fit and the preservation of racial purity ([14], pp. 313,
47580).
4 Conclusions
The most striking observation about the metaphor is the multifariousness of its
usage. But, nonetheless, the metaphor has had its time. A search for struggle for
existence occurring in titles and/or abstracts of documents in both the SSCI and
SCI databases revealed only 21 entries for the period 19731999 (SSCI: 15; SCI: 6
entries). Evidently it is justified to say that the struggle for existence as a metaphor
has not survived the struggle for use and attention.
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37. Young RM (1969) Malthus and the evolutionists: the common context of biological and social
theory. Past Present 43:109145
38. Zmarzlik HG (1974) Osterreichische Sozialdarwinisten. Ein Beitrag zur Brutalisierung des
politischen Denkens im spaten 19. Jahrhundert. Der Donauraum 19:147163
The Theory of Evolution and Cultural
Anthropology
Henrika Kuklick
Abstract The relationship between cultural anthropology and Darwinism is com-

plex. Contrary to what was once received wisdom, cultural anthropology was not
inspired by Darwins ideas. Many nineteenth-century anthropological arguments
predated Darwinism by a century or more. Yet, Darwinians figured prominently in
organized anthropology in late-nineteenth century Britain, and when Darwin wrote
The Descent of Man he drew upon the writings of an international population of
anthropologists although his most important sources were British. But cultural
anthropology changed dramatically at the end of the nineteenth century, when its
practitioners left their armchairs and took to the field and conceptualized cultural
variation in terms of Darwinian biogeography. Arguably, these practitioners, such
as Baldwin Spencer, were influenced by the Darwin who wrote On the Origin of
Species, not the Darwin who wrote The Descent of Man. And the disciplinary result
of their labors was paradoxical: cultural anthropology informed by notions derived
from Darwinian biology factored biological elements out of explanations of cultural
variation.
I must begin with some definitions. To speak of cultural anthropology in the mid-
nineteenth century, which is when my story necessarily begins, is to speak anachro-
nistically. There was no such enterprise at that time. The family of inquiries that are
collectively termed anthropology had three branches: one, prehistoric archaeology;
two, anthropology which meant investigations of humans physical characteristics;
and three, ethnology which then meant investigations of behaviors that humans
learned (rather than performed instinctively) and which was a fundamentally histori-
cal pursuit. What was then often called ethnology was the member of the anthropo-
logical family closest to modern-day cultural anthropology, also known as social
H. Kuklick (*)
Department of History and Sociology of Science, University of Pennsylvania, Philadelphia, PA, USA
e-mail: hkuklick@sas.upenn.edu

84 H. Kuklick
anthropology (The names social anthropology and cultural anthropology have some-
times denoted different approaches, which have been linked to different national
styles, but today socio-cultural anthropology is a relatively international enterprise).
Because the bible provided an axiomatic background to early-nineteenth century
ethnology, degeneration figured prominently in its narratives (as, of course, the
biblical story of humankind was of descent from the perfection of Eden). In the second
half of the nineteenth century, progress became the statutory story, and degeneration
was understood as unnatural in the absence of some sort of conflict between peoples.
In the early decades of the twentieth century, socio-cultural anthropology became
sharply distinguished from ethnology; at that time, the objective of constructing
historical backgrounds for their subjects then-current behavior became optional (if
not irrelevant) for many practitioners. The rejection of historical analysis was an
unanticipated consequence of research informed by evolutionary theory.
We must also note that, with the possible exception of anthropologists in France,
the division of labor among nineteenth-century anthropologists was hardly as
clear-cut as it became in the twentieth century.1 Consider, for example, the British
baronet and long-serving Liberal Party Member of Parliament John Lubbock
(18341913), the child of a friend and neighbor of Charles Darwin who served
as an informal tutor to the young Lubbock. In 1871, Lubbock became the first
president of the Anthropological Institute of Great Britain and Ireland (in and after
1907, the Royal Anthropological Institute), which incorporated all branches of
anthropology. The scientific work for which Lubbock was best known was on
insects, but his anthropological specialty was ethnology. Lubbocks work was
frequently favorably cited by Darwin in The Descent of Man, first published in
1871; indeed, although my generalization is based on impressions rather than
calculations, I estimate that Darwin cited Lubbock more frequently than any
other anthropologist in this book and he cited an international congeries of
anthropologists in it. Lubbock was also well known for his work in archaeology,
and chose to become Lord Avebury when he was elevated to the peerage in 1900,
naming himself after prehistoric megaliths he had studied and saved from
destruction.2
Consider Lubbocks contemporaries outside Britain. The best-known American
anthropologist was the lawyer Lewis Henry Morgan (18181891) probably most
often remembered today for his influence on Karl Marx and Friedrich Engels
whose social theory Lubbock treated somewhat critically, as he did in his presiden-
tial address to the Anthropological Institute.3 Darwin was also critical of Morgan
(although Morgans anthropology seems to have been of less interest to Darwin
1
See Emmanuelle Sibeud, The Metamorphosis of Ethnology in France, in Henrika Kuklick, ed.,
A New History of Anthropology (Oxford/Malden, MA: Blackwell, 2008), 96110.
2
Henrika Kuklick, The Savage Within. The Social History of British Anthropology, 1885-1845
(New York: Cambridge University Press, 1991), 456.
3
John Lubbock, On the Development of Relationships, Journal of the Anthropological Institute
1 (1872): 129.
The Theory of Evolution and Cultural Anthropology 85
than what one might call anachronistically his ethology of the beaver). Given
that Morgan had an identity as a naturalist, it is not surprising that his model of
the course of human history had a biological component, rather than being strictly
social: for example, he asserted that as humans brains grew in size as they
negotiated the social developmental stages from savagery to civilization (a judg-
ment with which Darwin agreed).4 And consider Germanys pioneering physical
anthropologist and ethnologist, respectively, the noted pathologist Rudolph
Virchow (18211902) and Adolf Bastian (18261905) who studied medicine
with Virchow. Virchow, who was, like Lubbock, a liberal parliamentarian, pro-
moted all types of anthropological inquiries; but, as Bastian also did, he insisted that
these inquires were entirely discrete. Their example was followed by Johannes
Ranke (18361916) who taught both cultural and physical anthropology when he
was appointed in 1886 to the first university chair in anthropology in Germany, in
Munich the first professorship in the subject anywhere, to the best of my
knowledge; but Ranke did not present his two subjects in tandem.5 In sum: although
the intellectual projects of ethnology and physical anthropology may have been
represented as distinct by nineteenth-century scientists, these projects coexisted
within the persons of individual scientists, many of whom (inevitably) made
intellectual connections between them. Note also that personal as well as intellec-
tual ties joined many of the figures I am describing.
How did Darwin describe his account of human history? In The Descent of Man
he paid tribute to the eminent German biologist Ernst Haeckel, who was Virchows
student and protege, but became his former mentors scientific opponent as a major
proponent and popularizer of Darwins ideas in his country.6 Darwin said of
Haeckel, Almost all the conclusions at which I have arrived I find confirmed by
this naturalist; and Darwin claimed (however accurately) that he would not have
completed The Descent of Man if he had not done so before Haeckels functionally
4
See, e.g., John Lubbock, On the Customs of Marriage and Systems of Relationship Among the
Australians, Journal of the Anthropological Institute 14 (1885): 292300; Lubbocks attack is
indirect here, on Morgans Australian disciples, Lorimer Fison and A. W. Howitt. For Darwin on
Morgan, see Descent, 75, 84. On Morgan, see, e.g., Bernhard J. Stern, Lewis Henry Morgan:
American Ethnologist, Social Forces 6 (1928): 344357.
5
Andrew D. Evans, Anthropology at War. World War I and the Science of Race in Germany
(Chicago: University of Chicago Press, 2010), 30. I do not consider comparable the appointment of
Daniel Garrison Brinton as professor of ethnology and archaeology in the Academy of Natural
Sciences in Philadelphia in 1884; technically, his appointment as professor of ethnology at the
University of Pennsylvania in the same year as Rankes appointment, 1886, was the first in North
America, but, as Regna Darnell observes, he received no salary and had no identifiable students;
see her North American Traditions in Anthropology, in Kuklick, ed., 38. E. B. Tylor was
promoted from a readership at Oxford to a personal professorship in 1896.
6
See, e.g., Robert J. Richards, The Tragic Sense of Life (Chicago: University of Chicago Press,
2008), 28.
86 H. Kuklick
equivalent work appeared.7 So Haeckels assessment of the development of anthro-

pology as an enterprise is of some interest. In 1913, he delivered a lecture entitled
Fifty Years of Anthropology to an International Medical Conference in London.
In it, he identified three disciplinary foundational moments that occurred in 1863: a
lecture he gave on Darwins theory of evolution at the Congress of Natural
Historians at Stettin; the publication by the German Zoologist Karl Vogt of his
Man, his Place in the Universe and in the History of Earth; and the publication by
Thomas Huxley of his Mans Place in Nature. All of Haeckels intellectual markers
represented identification of the family relationships joining humans to other life
forms as well as corroboration of his Biogenetic Law: The human embryo repeats
in its growth the history of the human race. Darwin had certainly found clues to
evolutionary relationships in embryonic forms, and Haeckel argued (somewhat
self-servingly) that the study of comparative embryology was what distinguished
the Darwinian understanding of human evolution; thus, in Haeckels account, there
had been only a brief interval between the publication of On the Origin of Species
and the emergence of the distinct scientific field of anthropology.8
Haeckel failed to mention perhaps out of ignorance that 1863 was also the
year in which the Anthropological Society of London (ASL) was founded, seceding
from and claiming a more uncompromisingly scientific approach than the Ethno-
logical Society of London (ESL).9 The foundation of the Anthropological Society
was inspired by that of the Societe dAnthropologie de Paris, organized in 1859 by
Paul Broca. In both groups, the predominant view was polygenism, the characteri-
zation of types of humankind as separate species, rather than varieties of a single
species. Darwin explicitly rejected polygenism as inconsistent with the principle of
evolution. The Darwinians among mid-century anthropologists largely remained in
the ESL; indeed, the only Darwinian active in the ASL was Alfred Russel Wallace,
usually remembered as the co-discoverer with Darwin of the process of natural
selection and he tried to reconcile the two anthropological factions.10
Did others agree with Haeckels judgment that On the Origin of Species inspired
the development of anthropology? Certainly, there were others who agreed with his
estimation of the scientific importance of the biogenetic law. For example, in 1890,
the Oxford zoologist E. B. Poulton observed that the great impetus given to
biological inquiry by the teachings of Darwin ha[d] chiefly manifested itself in
7
Charles Darwin, The Descent of Man, Second Edition (New York: A. L. Burt, 1874), 3.
8
Ernest Haeckel, Fifty Years of Anthropology, The North American Review Vol. 198, No. 696
(November 1913): 609616; quote 610f. For Darwins embryology, see, e.g., Descent, 10. For
Darwins rejection of polygenism, see, e.g., Descent, 200.
9
The definitive study of the split between the ESL and ASL as well as their reunion in 1871 as
the Anthropological Institute is George W. Stocking, Jr., Whats In a Name? Whats in a name?
The origins of the Royal Anthropological Institute, 18371871. Man n.s. 6 (1971): 36990.
10
For Wallaces efforts to reconcile the ASL and ESL viewpoints, see his The Origin of Human
Races and the Antiquity of Man Deduced from the Theory of Natural Selection, Journal of the
Anthropological Society of London 2 (1864): clviii-clxxxvii.
the domain of Comparative Anatomy, and especially in that of Embryology.11

A reading of The Descent of Man might suggest that the formulation of the
biogenetic law had been equally inspirational for anthropologists, since it was
translated into generalizations about the course of human development by the
many authorities cited by Darwin who included not just Lubbock and Morgan
but also such figures as Herbert Spencer, John McLennan, John Beddoe, and
E. B. Tylor, the most prominent anthropologist in nineteenth-century Britain, the
first to gain an academic position. In 1884, Tylor was appointed Reader at Oxford,
and he is conventionally credited with formulating the definition of culture as
learned behavior that remains foundational today.12 Darwin cast his net widely in
searching for anthropological justification for the arguments he made in Descent.
Indeed, the theoretical positions of Darwins anthropologist-sources ranged from
the biblically-inspired notions of the early-nineteenth century British physician
James Cowles Prichard (a pillar of the Ethnological Society of London) to the
polygenism of the Americans Josiah Nott and George Gliddon. But Darwin
seconded the judgments of prominent anthropologists of his day. If we focus on
the persons whose theoretical views mattered to him, we are not surprised to see
that they were overwhelmingly British, although British schemes bore considerable
resemblance to those of the American Morgan and the German Gustav Klemm
(whose voice in his country was a minority one).13
What was conventional wisdom among British anthropologists around the time
that The Descent of Man was published? Humans in their primitive condition were
engaged in a Hobbesian war of each against all, according to Thomas Huxley,
who has been conventionally represented as Darwins Bulldog because he
became Darwins public defender after the publication of On the Origin of Species
in 1859 and who played a major role in the anthropological community during the
last third of the nineteenth century, not least as broker of the union of the ESL and
the ASL as the Anthropological Institute. Nevertheless, The savage is possessed of
human reason and speech, Tylor wrote, indicating that his brain power . . .
enables him to receive more or less of the education which transforms him into a
civilized man. This statement enunciated the principle of the psychic unity of all
varieties of the human species, which implied that humans everywhere were
capable of independent invention of the rudiments of civilization, if not its
11
Quoted in Richard Burkhardt, Jr., Patterns of Behavior (Chicago: University of Chicago Press,
2005), 2f.
12
Tylors first appointment at Oxford was in 1883 as Keeper of the university museum, and he
became Reader in anthropology in the following year.
13
A major point of considerable contention between British anthropologists and Morgan was
whether the original type of relationship between the sexes as what Morgan termed primitive
promiscuity which also seemed unlikely to Darwin; on this point see my Humanity in the
chrysalis stage: Indigenous Australians in the anthropological imagination, 18991926, British
Journal for the History of Science 39 (2006): 535568. On Klemm, whose ideas were possibly an
inspiration for Tylor, see Harry Francis Malgrave, Gustav Klemm and Gottfried Semper: The
Meeting of Ethnological and Architectural Theory, Anthropology and Aesthetics 9 (1985): 6879.
88 H. Kuklick
highest refinements. Everywhere, the course of peoples evolutionary advance

through an ordered series of culture complexes was the same, although peoples
negotiation of this course varied in pace and consistency; some developmental
anomalies so called survivals of past practices invariably endured, even
though they no longer contributed to the orderly operation of social life. As John
Lubbock observed in 1870, progress was not inevitable: if a population had
achieved perfect adaptation to its environment, it would not modify its way of
life unless prompted to do so by some sort of novel stimuli. Nevertheless, when
there was change, it had a virtually inevitable direction.
To anthropologists, all of these evolutionary trends were consistent: from migra-
tory lifestyles to settled ones; from practice of mating customs that violated
Victorian standards of morality and represented defective understanding of
biological kinship to the development of the monogamous and patriarchal family;
from an undifferentiated political-social structure to a defined class system
associated with an established government order that incorporated routinized
patterns of succession to offices, which ruled by law rather than force; from simple
to sophisticated technology; and so on. In general, evolutionary trends represented
the triumph of rationality, science, and compassionate morality all of these some
functions of personal discipline. In Tylors summary formulation, the wildman of
the forest [was] forgetful of yesterday and careless of tomorrow . . . so wanting in
foresight to resist passion and temptation.
Social evolutionists believed that contemporary primitives were much like the
earliest humans, and all of these relationships were parallel: primitives to advanced
peoples; the lower orders to the upper classes (mental defectives, the unredeemable
poor known as the residuum, and the criminal classes were the most primitive of
all); women to men; and children to adults. The development of the child
recapitulated the development of the human species in both biological and social
terms. Growing children in advanced societies recapitulated the moral stages of past
ages while playing with toys that were the practical tools of their ancestors the
bow and arrow of the hunter and the rattle of the witchdoctor.14 Tylor often called
anthropology the reformers science, the findings of which would enable the
great modern nations to understand themselves, to weigh in a just balance their
own measures and defects (prominent among these defects were survivals of the
primitive past, which would presumably be eradicated once they had been identified
and recognized for what they were). Thus, in the nineteenth century (unlike in the
twentieth) the disciplines purview was all types of human societies at all times.
Nevertheless, anthropologys most urgent task was recording what could be learnt
about primitive peoples, whose culture (if not very physical existence) was destined
to disappear with exposure to Western civilization. Compelling evidence was
14
The general discussion is in Kuklick, 1991, esp. 8088. See 86 for the quotation from Huxleys
Social Diseases and Worse Remedies (1891), 80 and 86f for the quotations from E. B. Tylor,
Anthropology (1881) and 82-3 for the discussion of John Lubbocks The Origin of Civilization and
the Primitive Condition of Man (1870).
provided by the extinction of the Tasmanians, who were imagined as the most
primitive population who had survived until the modern era, the last full-blooded
member of whom had died in 1876.15
Darwins account of human development was much the same. The human
embryos developmental stages represented negotiation of the process by which
the human species developed from lower forms. Useless biological features
persisted in adults survivals that preserved attributes of our remote semi-
human progenitors and were especially notable among mental defectives.
Classifying human varieties as different species was unjustified, not least because
interbreeding was evidently viable; there were not clear demarcations among types,
since they graduate[d] into each other. Certainly, among contemporary
primitives, relics of human biological development were more conspicuous than
they were among advanced peoples. Primitives brains seemed smaller than those
of advanced peoples (Australian Aborigines the smallest of all), and there was
somewhat less physical variability among primitive than advanced populations.
Moreover, different human types differed in mental qualities, chiefly . . . in their
emotional, but partly in their intellectual faculties. Nevertheless, the differences
among human types were relatively inconsequential, not least because considerable
mental powers were required to survive in unimproved nature. And even the most
savage of peoples, such as the Fuegians, displayed many little traits of character
showing how similar their minds were to ours; the Fuegians behavior was but one
expression of a general phenomenon evident from Mr. Tylors and Sir
J. Lubbocks interesting works, Darwin said. Independent invention followed
from the consistency of humans mental endowments: ethnographic evidence
proved that even the most primitive peoples were capable of making evolutionary
progress. As primitives advanced, they developed aesthetic and moral sensibilities,
notions of private property, fixed abodes, and formal government. Nevertheless, in
the relatively near future the civilized races of man [would] almost certainly
exterminate and replace the savage races throughout the world. Some climates
were distinctly inhospitable, but the civilized would prevail everywhere that they
could acclimatize (and civilized peoples capacities to adapt to varied climates were
greater than primitives were). The limiting case of the Tasmanians (at the time of
Darwins writing hovering on the brink of extinction) exemplified the process by
which primitive peoples disappeared, being a product of both colonists deliberate
efforts to eliminate them and their constitutional vulnerability.16 Most important, as
we all know, when Darwin wrote about the human species, he was at his most
Lamarckian which is to say that he was at his least Darwinian, as we have come to
understand this approach. In Darwins account, just as in the accounts of his
15
These characteristic remarks of Tylor happen to come from his introduction to Friedrich Ratzel,
The History of Mankind (1896), quoted in Kuklick, Ibid, 7. For Tylor in 1893 on the Tasmanians,
see Ibid, 251.
16
Descent, quotations, in order, 14, 199, 203, 191, 178; see also 10f, 17, 22, 32, 44, 60ff, 164, 105f,
151, 214, 209.
90 H. Kuklick
anthropologist-sources, habits became instincts and human history had an inherent

direction.
What does the historian of anthropology make of the relationship between
Darwins ideas and those of his anthropologist contemporaries? Did On the Origin
of Species inspire a congeries of thinkers to formulate arguments that Darwin
naturally found congenial when he came to write The Descent of Man? Recall
that this was Haeckels judgment, and it became received wisdom.17 But when the
history of anthropology began to develop as a specialized scholarly subject several
decades ago, it seemed especially important to establish that there had been no
Darwinian impetus behind the emergence of the field itself. In such seminal works
such John Burrows Evolution and Society, first published in 1966, much was made
of the pre-Darwinian background to socio-cultural anthropological analysis. As
Burrow insisted, it was essential to dispel the widespread belief that the growth of
social anthropology in the third quarter of the [nineteenth] century [w]as a kind of
joyful and conscious imitation of the natural sciences.18 Rather, the discipline had
a number of antecedents in earlier thought of a strictly social character. In particular,
conspicuous features of the model anthropologists shared with Darwin the
recapitulation hypothesis, the comparative method, the notion that human develop-
ment proceeded through an ordered sequence of stages in which survivals from
antecedent forms made progress somewhat disorderly derived from the thinkers
of the Scottish Enlightenment. Indeed, Enlightenment thinkers formulated the very
definition of the minimal characteristics of civilization as settled habitation, private
property, and stable formal government.19 Also central in Enlightenment thought
was the notion of sympathy, the essential social instinct for Darwin, the
existence of which was axiomatic for many late-nineteenth century social thinkers
in Europe and America.20
It is evident that Burrow was right: the social theories that informed late-
nineteenth century anthropological thought had a considerable lineage, and did
not derive from biological thought. It is also the case that Burrow was writing at a
time when socio-cultural anthropologists considered their enterprise altogether
distinct from biology, even when their colleagues included physical
anthropologists, as has been traditional in anthropology departments in North
America: anthropologists then wanted a past consistent with their present. Burrow
did not trouble to consider the possibility that biological thought might have shaped
cultural anthropology at the very end of the nineteenth century. But it was then, as
I will argue in the remainder of this paper, that Darwinian ideas of the sort that we
17
For one effort to establish anthropologys Darwinian pedigree, see A. C. Haddon, History of
Anthropology (London: G. P. Putnams Sons, 1910).
18
J. W. Burrow, Evolution and Anthropology in the 1860s. The Anthropological Society of
London, 186371, Victorian Studies 7 (1963): 137154, p. 141.
19
J. W. Burrow, Evolution and Society (Cambridge: Cambridge University Press, 1966), 1016.
20
For one discussion, see Susan Lanzoni, Sympathy in Mind (18761900), Journal of the
History of Ideas 70 (2009): 265287.
now consider truly Darwinian ideas typical of the author of the On the Origin of
Species as opposed to the author of The Descent of Man in fact become very
important in analysis of social behavior. The unanticipated consequence of this
development was the emergence of a cultural anthropology thoroughly
differentiated from biological anthropology.
Intellectual change had followed methodological change. That is,
anthropologists perspective was altered as they turned away from what is conven-
tionally termed armchair anthropology the production of synthetic judgments on
the basis of data that theorists did not collect themselves but culled from all manner
of sources, a type of analysis widespread among diverse types of naturalists, of
course. In the late-nineteenth and early-twentieth centuries, anthropologists from
Britain, Germany, France, and North America took to the field to collect their own
data. These fieldworkers had precursors most notably the Berlin-based Adolf
Bastian, whose career as a museum anthropologist included considerable fieldwork
but it was not until the end of the century that there was a substantial movement of
academically-trained persons out of the shelter of their studies. In the received
history of anthropology, the value of fieldwork was appreciated only as the result
of a historical accident: in 1914, at the time that World War I broke out, Bronislaw
Malinowski happened to be visiting Australia (to attend a meeting of the British
Association for the Advancement of Science); as a citizen of the Austro-Hungarian
Empire, he was identified as an enemy alien and trapped in the area for the duration
of the war; he found himself with no option other than to pursue funding that allowed
him to live in the field for an unprecedented period, and retrospectively recognized
the merits of his protracted field research.21 But Malinowski himself acknowledged
that he was inspired by a number of figures.22
Conspicuous among Malinowskis anthropological forebears was Baldwin
Spencer, foundation professor of biology at the University of Melbourne. And
after reading Malinowskis earliest ethnographic efforts, Spencer became an
21
For one reiteration of this story, see David M. Hoffman and Andrew M. Gardner, Fieldwork and
Writing From the Field, in Gardner and Hoffman, Dispatches from the Field (Long Grove, IL:
Waveland Press, 2006), 1 f. In fact, Malinowski had intended to spend two years in the field, and,
thanks largely to the efforts of C. G. Seligman, his supervisor at the London School of Economics,
had the wherewithal to do so; he was able to secure additional funds from the Australian govern-
ment by promising to identify information useful to Australias colonial administrators. Michael W.
Young, Malinowski. Odyssey of an Anthropologist, 1884-1920 (New Haven: Yale University Press,
2004). 245f, 437; see also Malinowski to Atlee Hunt, Secretary, Department of External Affairs,
April 28, 1915, and Hunt to Malinowski, May 4, 1915, Yale University Archives, MS 19, Box 4.
22
Anthropologists fieldwork methods were in no small part borrowed from the methods of other
practitioners of the sciences that derived from natural history, not least because a number of early
field-going anthropologists were scientifically trained. For one account of the development of field
method, which includes an analysis of the origins of Malinowskis approach, see my Personal
Equations: Reflections on the History of Fieldwork, With Special Reference to Sociocultural
Anthropology, Isis 102 (2011): 1-33.
92 H. Kuklick
informal tutor to Malinowski for a time.23 Oxford-trained, Spencer was introduced

to anthropology when he audited E. B. Tylors undergraduate lectures; and in 1885,
together with his then-mentor, H. M. Moseley, Oxfords Professor of Human and
Comparative Anatomy, he assisted Tylor in setting up the exhibits in Oxfords Pitt-
Rivers Museum.24 Taking up his Melbourne post in 1887, Spencer did not enjoy his
work, complaining in a letter to an Oxford classmate that his obligatory teaching of
medical students was the most monotonous and soul killing work imaginable.25 It
is not surprising that he found solace in anthropology, which increasingly occupied
his intellectual attention after he served as the zoologist on the Horn Expedition to
central Australia in 1894. While on the expedition, he met a long-serving civil
servant who presided over Aboriginal affairs, F. J. Gillen (who acted as chief
informant for the expeditions anthropologist, E. C. Stirling), and the two
established a durable research partnership.26 Spencer wrote all of the texts that
were published under his and Gillens names and surely contributed much of their
conceptualization, but he also invariably consulted Gillen. By 1913, Malinowski
said of Spencer and Gillen that half the total production in anthropological theory
ha[d] been based upon their work, and nine-tenths affected or modified by it.27 It is
because their research on central Australia had an international impact that I will
devote most of the remainder of my paper to it.
In the Australian summer of 18961897, Spencer and Gillen did fieldwork for
their first work, The Native Tribes of Central Australia, published in 1899. Focused
on the Arrernte (whom they called the Arunta), it was what was then called an
intensive study, a type of anthropological inquiry that had only just been defined
(in opposition to survey research, such as that conducted by the 1898 Cambridge
Anthropological Expedition to Torres Straits). The intensive study was justified
with a plan for the development of a new sort of anthropology that was, in fact,
never realized. That is, the intensive study would shortly become an end in itself,
but it was initially rationalized as the fundamental building block of comparative
23
One testimonial to Spencers encouragement is Malinowskis letter to his supervisor at the
London School of Economics, C. G. Seligman, 4 May 1915, Malinowski Papers, file 27/3, London
School of Economics.
24
D. J. Mulvaney and J. H. Calaby, So Much that is New. Baldwin Spencer, 1860-1920 (Carlton,
Victoria, Australia: 1985), 60.
25
Baldwin Spencer to W. E. Roth, 30 January 1903, S P, Box 1A. As the Protector of Aborigines
for Queensland, Roth was himself able to devote considerable time to anthropological inquiry.
Spencer also served as the editor of the Horn Expeditions Reports.
26
Frank Gillen was Post and Telegraphs Stationmaster, stipendiary magistrate, and Sub-Protector
of Aborigines for South Australia; his formal schooling ended when he was twelve.
27
Bronislaw Malinowski, Review of Across Australia by Baldwin Spencer and F. J. Gillen, Folk-
Lore 24 (1913), 278.
analyses: as detailed studies of delimited areas accumulated, their results would be

compared, and generalizations about human evolution would be formulated.28
As the worlds largest island, Australia was understood as a protected preserve
for living fossils, ancient types of plants and animals that were extinct on the
continents of the Northern Hemisphere.29 A general Darwinian principle explained
the natural history of Australian life forms. That is, their vulnerability when
confronted with imported forms which had been demonstrated many times over
as the plants and animals that colonists brought with them overwhelmed indigenous
species confirmed Darwins judgment that the geographical isolation necessary to
species differentiation in the limiting case of island enclosure resulted in idiosyn-
cratic types that were unlikely to survive competition with species bred in more
competitive continental conditions.30 If cultural phenomena were analogous to
biological ones, Australias human behavioral forms were also ancient, and the
geographically isolated area of Central Australia was a preserve within a preserve, a
virtual island where persons were relatively protected from the pressures of natural
selection as these processes operated in the cultural realm. As the armchair anthro-
pologist J. G. Frazer put it, in Central Australia the scientific inquirer might
reasonably expect to find [there] the savage in his very lowest depths, to detect
humanity in the chrysalis stage.31 In Spencer and Gillens words, the conditions of
Central Australia had preserved creatures that have everywhere passed away and
given place to higher forms, including human beings that still remain on the
culture level of men of the Stone Age, persons who were exceptionally backward
28
A. C. Haddon to Baldwin Spencer, 5 May 1902, in the Spencer Papers, Pitt Rivers Museum,
Oxford (subsequently, S P), Box 1. Near-contemporaries, Spencer and Haddon were both trained
as biologists, and were professional rivals as such before they became like-minded
anthropologists: Haddon was the runner-up in the competition for Spencers Melbourne chair.
For Haddons endorsement of detailed study of a delimited area, as well as an account of the
vicissitudes of his career, see Henrika Kuklick, Islands in the Pacific: Darwinian Biogeography
and British Anthropology, American Ethnologist 23 (1996): 61138.
29
As Patrick Brantlinger has documented, arguments that Australian Aborigines were inferior
forms of humankind date to the beginning of British colonization of their land. Long before the
promulgation of Darwins evolutionary theory, Europeans in Australia pronounced that
Aborigines constituted the connecting link between man and the monkey tribe, as Peter
Cunningham did in 1827; quoted in Brantlinger, Dark Vanishings (Ithaca: Cornell University
Press, 2003), 117. On the general theme of Australia as a zoological garden of living fossils, see
R. A. Stafford, Annexing the landscapes of the past: British imperial geology in the nineteenth
century, in J. M. MacKenzie, ed., Imperialism and the Natural World (Manchester: Manchester
University Press, 1990), 6789, esp. 812.
30
For a general discussion of the putative plight of Aboriginal Australians, see Russell McGregor,
Imagined Destinies. Aboriginal Australians and the Doomed Race Theory, 18801939
(Melbourne: University of Melbourne Press, 1997). For example, naturalists remarked upon the
triumph of the English bee over the Australian bee, and plotted the rapid territorial expansion of
introduced rabbits. For some naturalists observations, see Charles Nicolson to John Lubbock,
1857, Avebury Papers, British Library, Add.MS 49638; P. M. Byrne to Baldwin Spencer, 18 April
1895, S P, Box 1A.
31
J. G. Frazer, The Origin of Totemism, Fortnightly Review 71 (January-June 1899), 648.
94 H. Kuklick
because they had been shut off from contact with other peoples, deprived of the
developmental stimulus derived from external sources.32 The Arrernte were so
primitive that they were ignorant of the most basic facts of life that pregnancy
followed sexual intercourse and that deaths occurred naturally (rather than through
the operation of witchcraft).33
Spencer and Gillen had no doubt that the way of life of the Arrernte would not
long endure, even though the people had to a degree proved capable of turning the
changes bought by European colonization to their own purposes; they incorporated
rabbit fur in their personal adornments, for example, and adopted European tools.34
Spencer and Gillen were certain that the once large and powerful Arunta tribe
would soon be reduced to a degraded remnant.35 Their judgment followed a
conventional formula, derived from axioms of the evolutionist anthropology I have
described: contact with Europeans so profoundly affected primitive peoples that
they soon lost the last remnants of their individuality; and the larger was the
cultural gap between the peoples in contact, the more rapidly would the less
advanced society disintegrate.36
Nevertheless, Spencer and Gillen undertook to understand the Arrernte sympa-
thetically. They became fully-initiated members of Arrernte society, who were
able to learn secrets previously hidden from white men.37 They produced an
account of Aborigines life in all of its aspects, ranging from everyday practices,
such as making fire, and preparing Emus for eating; to aesthetic sensibilities,
expressed in designs with which Aborigines decorated their bodies, their sacred
objects, and the rocks and caves of Central Australia; to important ceremonies, such
as circumcision rituals.38 Moreover, Spencer and Gillen urged anthropologists to
put themselves into the mental attitude of the native, explaining that Aborigines
lifeways were appropriate adaptive responses to the conditions of Central Australia,
even when they violated European standards of propriety.39 They argued that
32
Balwin Spencer and F. J. Gillen, The Arunta (London: Macmillan, 1927), vii; Native Tribes, 54.
33
See Native Tribes, 265, 356, 53648.
34
Baldwin Spencer and F. J. Gillen, The Northern Tribes of Central Australia (London:
Macmillan, 1904), 720; Native Tribes, 575.
35
Baldwin Spencer and F. J. Gillen, Across Australia (London: Macmillan, 1912), vol. 2, 300; and
see Native Tribes, 78, 1718.
36
This characteristic statement is from A. C. Haddon, Manners and Customs of the Torres Straits
Islanders, Nature 42 (1890), 637642, quote 638. See also Native Tribes, vii. For the translation
of this generalization into a formula, see, e.g., W. H. R. Rivers, Report on Anthropological
Research Outside America, in W. H. R. Rivers, A. E. Jenks, and S. G. Morley, The Present
Constitution and Future Needs of the Science of Anthropology (Washington, DC: Carnegie
Institution of Washington, 1913), 528. And see Native Tribes, vii.
37
E.g., Native Tribes, v. Spencer and Gillens claim was somewhat hyperbolic; they had been
given local identities, but they did not have fully-initiated status, not having experienced the rites
de passage that would have conferred it.
38
Native Tribes, 5846, 61835, 21851.
39
Native Tribes, 48; see also 256.
Aborigines had developed their mental powers . . . along the lines which are of
service to them in their daily life, acquiring formidable memories and detailed
knowledge of their natural environment. They pointed to a gradual improved
development which is still proceeding, noting that Aborigines self-consciously
undertook to implement sensible suggestions for progressive change made by the
exceptionally talented persons among them.40 Indeed, they saw many admirable
features in Aborigines way of life. Aborigines were, for example, devoted to their
children, no less than the average civilized parent, as well as helpful and generous
to white men which, as Spencer and Gillen observed, they might well not have
been, given the many wrongs that had been done to them.41 Even in the absence of
formal government, they had developed a distinctive moral code that was rigor-
ously enforced, which served to maintain social order, even though it differed from
a white mans standard.42
Most important, the Arrernte way of life was not to be understood as some
function of biological endowments. This argument was predicated on Spencer and
Gillens judgment that all of Australias Aborigines were a single people. They
derived from a band of invaders who entered the country from some single port. But
sub-groups of the invading population subsequently wandered all over Australia.
Therefore, their cultural diversity a prominent feature of which was their remark-
able linguistic diversity was strictly cultural. It was to be explained as a product
of their adaptation to the extraordinarily varied ecology of the vast territory of
Australia; each sub-population had adjusted to its particular niche. Distinctive
differences among Aboriginal groups had developed when geological changes
created barriers that isolated [them] from one another for long periods of time.
Aboriginal populations adapted to their local conditions in fashions that were not so
much determined by these conditions as compatible with them, each population
developing skills in the fabrication of particular tools and other artifacts. Eventu-
ally, the Aborigines joined in wide-ranging trade networks to exchange their
distinctive goods (as well as implements introduced by white settlers).
Why was it critical to establish that the Australias Aborigines were a single
people? The Aborigines ability to survive under many conditions denoted their
enormous inherent potential which made possible further adaptation in the
future. Their diversity represented expression of nascent possibilities of develop-
ment along many varied lines, and their capacity and mental outlook should not
be underestimate[d].43 In sum, Spencers orthodox Darwinism (quite different
40
Native Tribes, 26. And see Baldwin Spencer, President, Inaugural Address, Australasian
Association for the Advancement of Science, Report of the Fifteenth Meeting of the Australasian
Association for the Advancement of Science. Hobart Meeting [actually held in Melbourne],
January 1921 (Melbourne: AAAS, 1921), liii-lxxxix, esp. lxvi.
41
Across Australia, vol. 1, 123, 1889; vol. 2, 3078.
42
Native Tribes, 46; and see 56, 96100, 196, 381.
43
Northern Tribes, 16; Native Tribes, 117, 596; Spencer, Inaugural Address, lxvii, lxxiii. And
see Native Tribes, esp. 19, 575, 5878.
96 H. Kuklick
from the stereotypical social Darwinism that some have imputed to him)
contradicted much conventional white settler wisdom: Spencer and Gillen
repudiated the widespread fears that European migrants to Australia would degen-
erate, even in the parts of the country that were least like Europe, its tropical
regions; and they suggested that Aborigines could learn to live in the colonists
social world and would produce healthy children when they formed unions with
Europeans.44 (It is worth noting that in his official capacity, Gillen was an energetic
defender of Aborigines rights and welfare.)45 However we may feel now about the
political significance of advocacy of assimilation, this position represented a dis-
tinctly liberal political view in Spencer and Gillens day.
Spencer and Gillen and the anthropologists with whom they agreed certainly
did not wish to argue that there were no qualitative differences among cultures.
But their approach conduced to the disaggregation of nature and culture. To
Malinowski, for example, it seemed obvious that populations social behavior
could change rapidly, while their biological characteristics remained stable.46
Moreover, it seemed clear that cultural evolutionary change did not take the orderly
course that nineteenth-century anthropologists had imagined. As Spencer observed,
there [wa]s no such thing as an all-around primitive tribe.47 The appropriate
way to understand the essence of evolution, Malinowski said, was not as a
sequence of different forms changing one into another, but [as] a better adaptation
of an institution to its function.48
His anthropological research earned Spencer international fame as well as a
knighthood, conferred in 1916. But his work with Gillen influenced anthropologists
worldwide in ways the men could not have anticipated (and to some extent
deplored). It figured in an international controversy over a general phenomenon
totemism, or behavior based on some putative special connection between a group
of people and a feature of the natural world, an animal or a plant. Because Spencer and
Gillen judged that Aborigines had had a distinctly Australian form of totemism, they
objected that many of the theories putatively derived from their evidence were quite
44
On fears that Europeans would degenerate in Australia, see Richard White, Inventing Australia.
Images and Identity 16881980 (St. Leonards, N.S.W.: Allen and Unwin, 1981), 7071.
45
For example, in one notable instance, in 1891, Gillen attempted to hold a white policeman liable
for the murder of an Aborigine; see John Mulvaney, F. J. Gillens Life and Times, in John
Mulvaney, Howard Morphy, and Alison Petch, eds., My Dear Spencer. The Letters of F. J. Gillen
to Baldwin Spencer (Melbourne: University of Melbourne Press, 1997), esp. 67.
46
See Nicholas Peterson, Studying man and mans nature: the history of the institutionalization of
Aboriginal anthropology, Australian Aboriginal Studies 7, 2 (1990), 7.
47
Baldwin Spencer to J. G. Frazer, 7 June 1903, in R. R. Marett and T. K. Penniman, eds.,
Spencers Scientific Correspondence with Sir J. G. Frazer and Others (Oxford: Oxford University
Press, 1932), 93.
48
Bronislaw Malinowski, Anthropology, Volume 1 of the three supplementary volumes
published for the 13th edition of the Encyclopaedia Britannica (London and New York: The
Encyclopaedia Britannica Company, 1926), 133.
inapplicable to our tribes.49 I must stress that virtually all of the social
anthropologists in the British sphere of intellectual influence who came to profes-
sional maturity around 1920 and afterwards embraced a conceptual framework that
derived from studies that interpreted cultural variation in bio-geographical terms
and that the British social anthropology developed after World War I arguably had
an international influence greater than any other national variant of the discipline.
Many anthropologists outside Britain did not take an approach to cultural analysis
that derived from field research comparable to Spencer and Gillens, however.50
Indeed, in the United States, where anthropology came under the leadership of the
German-born and trained Franz Boas in the early twentieth century, bio-geographical
reasoning was repudiated just as Boass critique of evolutionism in the style of Morgan
became received wisdom. Although Boas allowed that geographical isolation could
preserve a populations older type of culture, his and his students approach to
anthropology was fundamentally historical with an emphasis on situational
peculiarities and a near-hostility to generalization; Boasians judged that geographical
factors explained behavior only in trivial and shallow ways, and that people often
did things in spite of geographical conditions.51 One might argue that Spencer and
Gillens analytic framework was practically irrelevant to the international debate.
Nevertheless, the anthropological argument provoked by the publication in 1899 of
Native Tribes of Central Australia and just as important the second edition of
J. G. Frazers The Golden Bough, published in 1900 and featuring Spencer and Gillens
material, suggests that the decline of the nineteenth-century, unilinear conceptualiza-
tion of cultural evolution was over-determined. Frazer brokered the publication of
Native Tribes with his publisher, Macmillan, not least because he intended to turn its
evidence to his purposes (this was a recurrent theme in Frazers dealings with his
publisher). And as a person with considerable stature in the world of anthropology in
his day (no matter what his legacy is in ours), Frazer was able to shape the totemism
debate that followed. In essence, Frazer expected that Spencer and Gillens data would
prove of vital importance in answering fundamental questions of evolutionary anthro-
pology: One, was it possible to specify the characteristics of each of the stages through
49
See Gillen to Spencer, 17 November 1902, in Mulvaney, Morphy and Petch, 419, 416.
50
Similar assumptions informed the Cambridge Anthropological Expedition to Torres Straits of
1898, organized by A. C. Haddon, a Cambridge-trained zoologist whose career was much like
Spencers (he came second in the competition to become Melbournes foundation professor of
biology, for example), and who corresponded with Spencer and promoted his work. Virtually all
British anthropologists until World War II were somehow professional progeny of the Torres
Straits Expedition; conspicuous among these were students of two members of the expedition,
C. G. Seligman and W. H. R. Rivers, respectively Bronislaw Malinowski and A. R. Radcliffe-
Brown. I describe the biogeographical conceptualization of the expedition in Kuklick, 1996.
51
Franz Boas, Arctic Exploration and Its Object, Popular Science Monthly 22 (1885): 7881;
Franz Boas, Ethnological Problems in Canada, Journal of the Royal Anthropological Institute 40
(1910): 530, 531. The generalizations are Alexander Lessers; see his Franz Boas, International
Encyclopedia of the Social Sciences, David Sills, ed. (New York: Macmillan and The Free Press, 1968),
II, p. 101.
98 H. Kuklick
which human populations passed during the course of their progress from savagery to
civilization? Two (the most important question for Frazer), what were the origins of
truly spiritual religion and monogamous marriage, the signal accomplishments of high
civilization?52
But the prevailing view of social theorists everywhere came to be that
evolutionists problems defied solution. It seemed impossible to extrapolate
from the simplest social state known at present to an absolute beginning, as
Emile Durkheim observed in The Elementary Forms of Religious Life (subtitled The
Totemic System in Australia), now generally regarded as his most important book
and unquestionably the most enduring product of the totemism controversy, which
relied heavily on Spencer and Gillens data and rejected Frazers interpretation of
them.53 Moreover, virtually any account could be dismissed with the charge that
was made against Native Tribes that it reported the behavior of people whose
authentic culture had been corrupted by contact with Europeans, precluding the
possibility that the people represented genuine specimens of any cultural stage.54
And social evolutionists ideal types were belied by reports from the field. For
example, evidence provided by the German missionary Carl Strehlow, who lived
with and also wrote about the Central Australian Arrernte, convinced a number of
persons, including the British Andrew Lang and the Austrian Wilhelm Schmidt,
that there was no association between superior spirituality and material progress;
humans in the lowest known grades of savagery, said Lang, were capable of
beliefs as monotheistic as some Christians.55 The German sociologist Max
Weber concluded that the phenomenon of totemism was evidently neither universal
nor associated with a specific pattern of development.56
Just as important, as the Boasian John Swanton observed in 1906, those who
debated the nature of totemism not only generated conflicting theories but were also
incapable of adopting shared definitions of the terms totem and totemism, as
well as of practices that might be associated with totemism, such as witchcraft,
magic, and fetishism.57 Such confusion made arguments circular. Did totem
ceremonies intended to increase the supply of the specific animal or plant with
which persons identified represent the very earliest form of religion? Or, did these
52
See, e.g., J. G. Frazer to George Macmillan, 23 August 1897, British Library, Macmillan
Archive, Add.MS 55134.
53
E. Durkheim, The Elementary Forms of Religious Life, translated by Karen E. Fields
(New York: Basic Books, 1995 [orig. 1912]), 7.
54
See, e.g., A. A. Goldenweiser, Reconstructions from Survivals in West Australia, AA n.s. 18
(1916): 4768.
55
Quoted in G. W. Stocking, Jr., After Tylor (Madison: University of Wisconsin Press, 1995), 59.
56
Max Weber, Sociology of Religion, posthumously published in 1925, cited in Brian Morris,
Anthropological Studies of Religion (New York: Cambridge University Press, 1987), 71.
57
J. R. Swanton, Review of Lectures on the Early History of the Kingship by J. G. Frazer, American
Anthropologist (subsequently AA) n.s. 8 (1906): 157160; and Review of The Secret of the Totem
by Andrew Lang, AA n.s. 8 (1906): 160165. Swanton was not alone in this critique. See also, e.g.,
A. R. Brown (later Radcliffe-Brown), The Definition of Totemism, Anthropos 9 (1914): 62230.
ceremonies constitute efforts to perform magic, rather than being acts of submis-
sion to the will of some sort of supernatural being?58 Resolution of these questions
required procedural consensus that anthropologists could not effect. The Boasians
critique of the issues at stake in the totemism debate represented dismissal of the
entire nineteenth-century social evolutionist paradigm: totemism was not associated
with the changes that denoted progress to evolutionists such as the shift from
kinship structured by ties to mothers to patrilineal organization, the specification of
marriage regulations that prohibited unions between close kin (however close kin
were defined in a given society), the differentiation of social roles and the elabora-
tion of class hierarchy, and the development of religion in which worshippers
submitted to an omnipotent god. Nor was there a consistent pattern of change:
because peoples acquired traits by diffusion as well independent invention, societies
often incorporated elements that evolutionists had imagined were irreconcilable.59
In sum: the repudiation of social evolutionary theory of the nineteenth century
variety was over-determined. Not least because of the expansion of colonial power
at the end of the nineteenth century, some exotic places became relatively safe
places for Europeans to spend sustained periods in residence; and these were the
areas from which came the evidence that seemed most significant to anthropologists.
Anthropological fieldwork framed by a Darwinian biographical approach proved
extremely important in changing the discipline, (perhaps paradoxically) leading to a
thorough separation of cultural from biological anthropology. Nevertheless, bio-
geographical analysis was not required to discredit linear models of human
development. Abundant evidence became available that served to suggest that
evolutionists tidy scheme of successive stages of evolutionary progress did not
correspond to observed behavior. Socio-cultural anthropology remained focused on
non-Western peoples, but the rationale for studying them ceased to be that their
behavior revealed patterns of evolutionary development (unless evolution meant
only successful adaptation to circumstances). Instead, simple societies were under-
stood as simple organisms in which it was relatively easy to observe the enduring
structures of social life that were essential to all those societies on earth that
managed to survive. Not until the 1980s would evolutionary approaches to the
analysis of culture that were advertised as authentically Darwinian seem respect-
able to more than a distinct minority of socio-cultural anthropologists but that is a
development that may be most significant as evidence of the rise of conceptual
pluralism in anthropology, and is, as such, another story.60
58
See, e.g., J. G. Frazer, On Some Ceremonies of the Central Australian Tribes, Report of the
Eighth Meeting of the Australasian Association for the Advancement of Science. Melbourne: 1900
(Melbourne: AAAS, 1901), 312321; Native Tribes, 170.
59
See especially, John R. Swanton, The social and the emotional element in totemism,
Anthropos 9 (1914): 288299; Alexander A. Goldenweiser, Totemism, An Analytical Study,
Journal of American Folk-Lore 23 (1910): 178293.
60
For a survey, see William H. Dunham, Advances in Evolutionary Culture Theory, Annual
Review of Anthropology 19 (1990): 187210.
100 H. Kuklick
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55134
The Concept of Evolution in Linguistics
Manfred Bierwisch
Abstract Language has been viewed for a long time as a field of evolution where new
things originate through variation. In the early nineteenth century, important insights
were reached within this still pre-Darwinian tradition by historical-comparative
linguistics. The laws of sound change and vowel shift discovered by Grimm and
others are essentially valid today, although development by then was largely
considered as loss and deprivation. A strictly Darwinian view of language change
was advocated in 1863 by Schleicher, who considered languages as organisms to
which biological principles of adaptation and survival were to be applied directly.
The difficulty to reconcile insights of language development with principles of
biological evolution was the requirement to construe languages as species, the
genetic information of which is subject to random variation and completely inde-
pendent of reproduction of their members. On closer inspection, the range of
possible language variation is provided by the human language faculty, and the
domain of competition for reproduction is the actual use of language. This leads to
two competing views of language change. A Lamarckian view considers variants as
the product of functional needs, subsequently inherited by the language used, while
a strictly Darwinian view considers variants as random products of the language
faculty which do survive or not according to conditions of language use. Although
in many, notably lexical, cases the functional origin of variants is beyond doubt, the
Darwinian nature of much language change is still attested by properties that cannot
be due to functional requirements, but emerge from traits of the language capacity
and are not suppressed by functional needs. Language change must thus be consid-
ered as evolution of mixed nature. A short epilogue raises the question whether this
type of mixed evolution could be the general nature of human history and cultural
development. Skepticism is indicated for two reasons. First, the language faculty is
domain specific, its conditions cannot be generalized to arbitrary fields. Second,
M. Bierwisch (*)
Humboldt-Universitat zu Berlin, Berlin, Germany
e-mail: bierwisch@gmx.net

104 M. Bierwisch
there is no reason to assume a universal cultural capacity from which action patterns
randomly arise.
1 The Background of the Humanities
Change per se does not constitute evolution. In this paper, we will use the term
evolution to describe change that gives rise to something that did not previously
exist. What evolution produces need not always represent a higher stage of devel-
opment, nor will it necessarily create an independent entity either in and of itself or
by taxonomic definition. What evolution should always represent, however, is a
genesis of differentiable states or entities. This definition allows for the existence of
various types of evolution, depending on what sort of processes are involved. There
is, of course, the evolution of biological species, but we can also speak of the
evolution of galaxies, of stars and planets, of geological formations and chemical
elements, or even of institutions, communities and technologies. With respect to
linguistics, the phenomenon of evolution is evident in at least three different areas:
phylogenesis (the development of the language faculty in man), ontogenesis
(the development of the internal language in an individual), and language change
(the emergence of the various languages and how they change over time).
That these phenomena are very different in their natures is obvious. Less
obvious, however, is how they relate to and influence one another.
As a species-specific characteristic in the strict sense, the phylogenesis of
language is one facet of the phylogenesis of homo sapiens and as such a subject
of study for evolutionary biologists. But it is also a matter for linguists, since
explaining the biological prerequisites for and the origination of the language
faculty is not possible without an understanding of the specific structure of
human language. In strict linguistic academia, researchers examine variation and
development primarily with respect to the phenomenon of language change and
hence to the emergence of new languages and dialects. One important question
facing researchers is how the development of languages is linked through language
acquisition with the conditions of the language faculty.
Man has since antiquity been aware of the fact that languages change, that their
elements and rules undergo alteration and development. For many years, however,
it was only possible to make isolated observations and in some cases develop
speculative theories, but these can be left aside here. It was not until the early
nineteenth century with the realization of the cultural and historical importance of
Sanskrit and the emergence of comparative historical linguistics that chance
observations were replaced by systematic study in the works of Rask, Bopp,
Grimm and others. The systematic understanding of the historical relationships
between the Indo-European languages and the fundamental concept of sound shift
that allowed for an explanation of these relationships led to the theory of the Indo-
European language family and concurrently to the far-reaching theory of systematic
language change. This theory allowed scientists to comprehend the existing
The Concept of Evolution in Linguistics 105
phonetic and grammatical forms of the various languages as the result of tangible
processes and then to retrace these processes to identify precursors to existing
forms. Thus was born a fundamental concept for describing the principles of
language development. Jacob Grimm summarized one key discovery as follows:
The entire twofold sound shift, which has momentous consequences for the
history of language and the rigor of etymology, can be so expressed in a table:
Greek P B PH T D TH K G CH
Gothic FPB TH T D ... K G
Old High German BFP DZT G CH K
(Jacob Grimm, Deutsche Grammatik, Part 1, [3], p. 584)

In other words, language development is determined by processes of the type:
P > PH, T > TH, K > KH, etc.
or, more generally: Stop > Spirant
Media > Tenuis
Spirant > Media
This grand scheme was fully confirmed by later supplementary work and the
incorporation of revealing details and was further generalized to cover the entire
Indo-European language family. Processes of this kind that lay claim to
exceptionless validity were later considered to represent phonetic laws, which
were explained and justified based on the workings of the human speech organs.
Initially, though, language change was deemed a phenomenon residing solely in the
realm of intellectual history, fully disconnected from biological evolution or any
other processes in the natural environment. In the full spirit of the romantic era,
Jacob Grimm viewed language development basically as a process of decay, as the
slipping away of a hereditary treasure. This even applied to sound change in itself,
which he interpreted as nothing more than a descent to a lower condition:
For just as Old High German has sunk one step down from the Gothic in all three grades,
Gothic itself had already deviated by one step from the Latin (Greek, Sanskrit). (Ibid.)
Wilhelm von Humboldt likewise viewed the development of language as an

intellectual process that was continuously robbing us of a rich inheritance. About
language, he wrote:
Its nature is to pursue a progressive path of development, under the influence of the mental
power, at any time, of its speakers. In this progress there naturally arises two periods, which
must be sharply distinguished, the one where the sound-making impulse of the language is
still in a state of growth and lively activity; the other where, after completed shaping at least
of the outer speech-form, a seeming halt occurs, and there follows a visible decline in that
creative sensuous impulse. ([5], Collected Works, 7, p. 160)
For both Humboldt and Grimm, one of the most prominent symptoms of such
decay was seen in the gradual disappearance of inflected forms. But of course the
obvious changes in language and even in language types that they explicitly
106 M. Bierwisch
examined in their works was not considered a phenomenon that could in any way be
linked to the evolution of biological species.
2 Evolutionary Linguistics: The Initial Approach
Thirty years later, Jena-based Slavist and linguist August Schleicher did make
precisely this link in an initial approach to linguistic Darwinism set forth in a
highly programmatic open letter that was inspired by Ernst Haeckel and entitled
Die Darwinsche Theorie und die Sprachwissenschaft [8]. In the English translation
published in 1869 under the title Darwinism Tested by the Science of Language,
Schleicher states:
Languages are organisms of nature; they have never been directed by the will of man; they
rose, and developed themselves according to definite laws; they grew old, and died out.
They, too, are subject to that series of phenomena which we embrace under the name of
life. The science of language is consequently a natural science; its method is generally
altogether the same as that of any other natural science.
Although in a note Schleicher excludes philology as a historical discipline, he

does state the following:
The rules now, which Darwin lays down with regard to the species of animals and plants,
are equally applicable to the organisms of language, that is to say, as far as the main features
are concerned. [. . .] What Darwin now maintains with regard to the variation of the species
in the course of time, through which when it does not reveal itself in all individuals in like
manner and to the same extent one form grows into several distinct other forms by a
process of continual repetition, that has been long and generally recognised in its applica-
tion to the organisms of speech. Such languages as we would call, in the terminology of the
botanist or zoologist, the species of a genus, are for us the daughters of one stock-language,
whence they proceeded by gradual variation. Where we are sufficiently familiar with any
particular family of speech we draw up a genealogical table similar to the one which
Darwin attempted for the species of animals and plants.
Building on the discoveries of comparative historical research, Schleicher uses a

rigid and somewhat speculative genealogical tree to depict the relationships within
the Indo-European language family as known to that time. Though he firmly
believed that a proto-language formed the common trunk of the family tree, he
nonetheless skeptically posed the question:
But how stands the fact with the creation of the genera? That is to say, in the glossologists
phraseology, with the self-development of those mother-languages which have given birth
to the different families of speech? . . . do those parent idioms again descend from a
common stock, and all these in the end from one single primitive form of speech? [. . .]
This question might be decided with greater certainty if we had examined the primitive
form of a good many more families of speech . . . than we have done.
Above all, the varieties of those special families of speech, which have been carefully
examined, are so great and of such a nature, as to render it impossible for any unbiased mind
to believe in a common origin.
Based on the approach derived from these considerations, Schleicher pursued

the study of the parallel development of species and languages:
Now we observe during historical periods how species and genera of speech disappear, and
how others extend themselves at the expense of the dead. I only remind you, by way of
illustration, of the spread of the Indo-Germanic family and the decay of the American
languages. In the earlier times, when the languages were still spoken by comparatively
weak populations, this dying out of forms of speech was, no doubt, of much more frequent
occurrence [. . .] It is very possible that many more species of speech perished during the
course of that time than the number of those which have prolonged their existence up to
the present day. This explains the possibility of so great an extension as for instance that of
the Indo-Germanic, the Finnic, the Malay and South-African families, which, over a large
territory, branched off into such a multitude of directions. A similar process is assumed by
Darwin with regard to the animal and vegetable creation; that is what he calls the struggle
for life.
Schleichers highly autonomous parallelization of biological and linguistic evo-

lution was based on the analogy between languages and biological species, an
analogy with elicits interesting insights but to a still greater degree reveals unre-
solved problems. The similarity between languages and species at first appears
plausible and unproblematic. Like a species, a language manifests itself within a
group of individuals (a population), with which it grows, migrates or dies out. But
unlike with a species, the extinction of a language does not necessarily mean the
demise of its population. The speakers of one language can continue to survive
speaking an altered language, whereas the members of one species cannot continue
to survive as the representative of another. Related to this finding are two essential
differences that are concealed by Schleichers equating of languages to natural
organisms.
Firstly, though neither Darwin nor Schleicher could have been aware of the true
nature of the changes that give rise to new species or languages, such knowledge is
not necessary to realize that the main issue lies somewhere else altogether, namely
that the traits of a new species are transmitted to the individuals of the next
generation by inheritance. This was Darwins basic thesis. In contrast to biological
evolution, however, the traits of a new language are not inherited by members of the
species; but rather must be acquired.
The second difference, directly related to the first, is the fact that the core claim
of evolutionary theory, which states that successful traits will survive and less
successful ones will die out, found no apparent counterpart in the process of
language development, regardless of whether such development was seen as a
process of enrichment or decay. For it is nowhere evident how changes in language
characteristics could be of any advantage in human reproduction, nor did Schleicher
ever touch upon this subject.
This problem is exacerbated by the fact that Schleicher not only compares
language with species but also with organisms, for example when he states that the
. . . roots are, as it were, the simple language-cells yet unprovided with separate organs for
the functions of verb, noun, etc., and these functions (the grammatical relationships) are yet
as undifferentiated as breathing and digestion are in single-cell organisms or in the
blastocysts of higher life forms.
108 M. Bierwisch
Even if only meant as a metaphor, this statement does suggest a comparison

between language units and organic elements that is invalid with respect to both
how such units and elements come into being and to the function they perform.
There may indeed exist contexts where organic stem cells could be compared with
unspecialized grammatical roots, but the evolution of linguistic elements cannot be
grasped in this way.
Based upon an impressive, systematic background of insights into the structure
and history of natural languages, Schleichers open letter represents a remarkable
surge forward in the development of linguistics and other disciplines. Nonetheless,
this work did not determine the course for the mainstream of linguistics. This fact
was doubtless due more to the growing alienation between the natural sciences and
the humanities than to the yet lacking genetic understanding of evolutionary
processes. In light of todays knowledge, however, we are now called upon to
revisit and reexamine Schleichers theories. The task at hand is to investigate
whether and in what manner the concept of evolution that applies to biology can
also be applied to languages (and hence to other sociocultural institutions).
To this end, it is helpful to first recapitulate the most important terms and
concepts.
3 Evolution and Language: Essential Definitions
Teleonomy, or purposefulness, deriving from evolution (this was the decisive

insight made by Darwin and Wallace) is in principle the result of those random
variations that subsequently prove to be advantageous for the organism and its self-
propagation. In contrast to the earlier Lamarckian theory of evolution, a teleonomic
trait is thus not the result of adaptation or goal-oriented (teleological) change, but
rather comes about through selection from randomly arising possibilities. This
means that the traits available for transmission to the next generation appear at
random and without any control. Only upon confrontation with the natural environ-
ment will any purposeful advantage of a trait be revealed, and this advantage will
ultimately lead to the trait being inherited by offspring. In a population where
individuals reproduce themselves and their constant traits, the following two factors
are decisive for the concept of evolution:
(a) The existence of a repertoire of random variations of traits within a population.
(b) The selection of those variants that succeed in the environment and in
reproduction.
Geneticists and molecular biologists today have a step-by-step understanding of
the details surrounding these factors and the relationship between them. To a large
extent, they have been able to explain the interaction of variation and selection
(a simple given for Darwin) in accordance with the laws of biophysics and behav-
ioral regulation. Of primary essence here is an understanding of the dichotomy
between the genotype as the sum of an individuals genetic information, the
variation of which is the subject of factor (a), and the phenotype that is expressed in
the organism and which is the object of selection in factor (b). The two factors are
therefore not only subjected to different effect mechanisms but also belong to
completely separate spheres of causality. The causes that drive variation and
which exert their effects in the genome (the entirety of an organisms hereditary
information) are unrelated to the conditions under which an organisms
characteristics are tested and selected. Even more important is the fact that the
genotype and the phenotype are governed by fully dichotomous types of natural
laws. In oversimplified terms, there is essentially no connection between the
microphysical conditions that impact the genome and the macrophysical processes
that shape the phenotype. The link between these two spheres is established by the
organisms hereditary information in accordance with the principles of genetic
control. Generalized across the population as a whole, the process of selection
will in time effect the spread of favorable traits in the population due to the
reproduction of successful variants.
Years of subsequent research have enhanced and enriched this basic scheme of
variation and selection by incorporating a number of different factors. Although
these factors do not threaten the fundamental theory of evolution, they certainly
must be taken into consideration when evaluating analogies between biological and
linguistic evolution. For example, the concept of adaptive selection, whereby only
advantageous variants enter the hereditary line, has been complemented by the idea
of exaptive selection, which also accepts characteristics that are neutral, i.e. not
disadvantageous, but which may very well assume an adaptive function under
changing conditions. This may explain, for example, why various decorative traits
have been incorporated into the hereditary material of many different species.
Further, there has been repeated speculation concerning the existence of internal
or structural selectors that act on the genetic information level to influence the
formation of variants, seemingly narrowing the gap between the phenotype and the
genotype. These structural selectors are, however, probably better understood as
factors that control the creation of random variants, thereby acting as regulators for
purely stochastic mutations and making these mutations random with respect to the
phenotype but not necessarily with respect to the structure of the hereditary
information.
Finally, Wilson and Dawkins have shown with their concept of sociobiology that
the conditions for variance and invariance in a species should be understood as the
tendency towards genome constancy, i.e. that the selection to which the phenotype
is subjected serves the self-assertion of the genotype. This highly successful
concept of genome identity preservation has spotlighted an aspect that is not
relevant in classic evolutionary theory. While selection was initially only seen
with regard to the reproduction conditions surrounding the phenotype, sociobiology
equates reproductive success with the constancy or self-assertion of the genome.
From this perspective, the selection of variants which indeed runs counter to such
constancy is considered to be minimal change under altered environmental
conditions. Of course, the genome has no intentions. It is not goal-oriented. On
the contrary, it works causally, like an actor, with the effect of maximum constancy
110 M. Bierwisch
of self-replication. The heteronomy of the genome and the organism remain

unchanged: the constancy and variation of the genetic information are not products
of the causal relationships associated with the phenotype.
Besides anatomical and functional characteristics, other genetically conditioned
and inherited traits include biologically determined, species-specific behavioral
patterns and their underlying organic conditions. Already given central prominence
by ethnologists, this notion is gaining importance in the field of sociobiology as
well. It also covers what are known as epigenetic developments, i.e. dispositions
and behaviors whose framework is biologically fixed but which only find expres-
sion during ontogeny under the influence of individual experiences. Examples
include walking upright in humans or song learning among birds. Human language,
too, would naturally fit into this category.
So what do these realizations about evolution mean with respect to language?
Here, we must clearly distinguish between three aspects that linguists have
discerned since de Saussure [2] as faculte de langage, langue and parole. The
same distinctions are essentially made by Chomsky [1] using the terms faculty of
language, internal language and external language.
1. Faculty of language is a characteristic specific to homo sapiens and is itself a
decisive product of evolution that emerged through mutation and selection and
which must be anchored in the genome. The only controversy here surrounds the
questions of how specific and differentiable this faculty is compared with other,
more general dispositions, and whether it grew out of a complex series of
developmental stages or emerged in a single evolutionary leap. Whatever the
case, the all-important outcome is the ability to create and iteratively combine
symbols, i.e. to make use of complex, convention-governed signs that link
structured signals to cognitively organized meanings. Of all the abilities that
evolution has endowed upon us, the faculty of language is without a doubt the
most characteristically human.
2. Building on the faculty of language, the internal language develops in a largely
spontaneous process as a complex structure within the brain of every normally-
constituted speaker. The internal language encompasses the entire range of
symbols and combination rules that are needed to construct and understand
utterances. The internal language acquired by the members of a language
community is the shared basis of that community and can vary, within narrow
limits, to allow for different elements and combination rules.
3. By external language we mean the structured behavioral habits that the internal
language makes possible, i.e. the concrete spacetime actions in a population
united by a given language, or, in other words, the speech acts and their
manifestations.
To our knowledge, the faculty of language is a universal, species-specific and
biologically hardwired disposition, the foundation of which is anchored in the
human genome. Among different populations, this universal disposition finds its
expression through varying and changing systems of internal and external language.
Taken together, internal and external language form two completely different yet
equally essential aspects of what August Schleicher envisioned as the linguistic

counterpart to the biological species. Internal and external language are asymmet-
rically linked, representing the structure and the execution of linguistic utterances,
borne by the members of a language community, that is, the individuals without
whom there would be no internal language or adherence to its rules.
Nonetheless we cannot as noted above equate the speaker of a language with
an individual of a species. While speakers of a language have the required ability,
individuals of a species are representatives of that species. A speaker belongs to a
species, or even to an ethnic group, but certainly not to language. (You can acquire
a different language but not a different ethnic group.) The difference drawn here is
not one of definition but rather of category, and its significance is greater than
Schleicher was aware.
But despite the fundamental difference between languages and biological spe-
cies, the relationship between internal language and language behavior does reveal
some parallels with the genotypephenotype relationship. Although the transfer
process through which the DNA structure determines the makeup and behavior of
an organism has nothing to do with the way that the internal language structures
linguistic utterances, the internal language does enable and control language
behavior just as the genotype determines the phenotype. As such, the internal
language would be comparable to the genome, with one crucial difference regard-
ing how such determination works: the structures and variants of the genome are in
no way subject to the principles of the phenotype, while internal and external
language intermesh with one another directly and are characterized in the same
terms. To put it briefly, the genotype and the phenotype are subjected to different
causalities within completely separate phenomenal spaces, while internal and
external language, though also determined by different causalities, belong to the
same domain of experience.
What do these complex correspondences and differences say about the validity
and fruitfulness of applying evolutionary concepts to language development?
4 Language Change and Evolution
One problem associated with evolution but which can only be addressed indirectly
or speculatively concerns the genesis of the proto-language or proto-languages in
connection with the phylogenesis of the language faculty, which is, after all,
difficult to conceive of as a purely abstract disposition. Pinker and Bloom [7]
have examined this problem and we must leave it at that here. The question that
we wish to address and answer here is as follows:
Do the principles of evolution apply to language change? Or, to be more precise:
Is language change, like biological evolution, the result of mutation and selection?
Before beginning a serious discussion of this question, a seemingly plausible and
often-made error is to be rejected, which clearly shows what language change is
112 M. Bierwisch
not. One pertinent example that is found in many different guises is the phenome-
non of phonetically reducing unstressed syllables as illustrated here:
we are not going to stay > we arent goina stay > we aint gonna stay
it is not necessarily so > it isnt necessarily so > it aint necessarily so.
We must not try to explain such reduction and assimilation of segments as a type
of mechanical attrition, as seen on worn out tires or well-trodden stairways, but
rather as a change to the internal sound form, i.e. as license within the internal
language. A phonetic effect has to be grounded in the internal language in order to
become a repeatable phenomenon of the external language and manifest itself as an
effect of language-change. Although the notion of language decay has since the
romantic era suggested otherwise, phonetic reduction (along with its opposite,
phonetic expansion) is not a mechanical process but rather a mental one. This
applies all the more to grammatical changes or changes in meaning, such as the
reduction of case distinctions or the shifting of semantic boundaries. All serious
attempts to analyze and explain language change give consideration to this insight.
The qualifying studies that fulfill this prerequisite draw upon a rich repertoire of
carefully documented and analyzed phenomena found in numerous and diverse
languages and dialects, including the classic facts derived from the Indo-European
languages mentioned in Sect. 1 above. We must distinguish between two opposing
schools of thought which, though occasionally approximating one another in their
descriptive details, pursue strictly different explanatory approaches. We may label
them functional/teleological and causal/teleonomic, but in essence they correspond
to the Lamarckian and Darwinian understandings of evolution. Whereas teleology
views the emergence of formal traits as a consequence of functional purposes,
teleonomy sees a causal explanation, independent of function, behind the emer-
gence of these traits, which then subsequently either demonstrate their fitness and
survive, or else disappear.
One thing that the teleological and teleonomic explanations for language change
share in contrast to the mechanical view is the insight that language characteristics
and their changes always, both in the long-term historical and narrower local
perspective, represent an internal language phenomenon and are thus part of that
sphere that corresponds to the genome: only that which reaches the elements and
rules in the speakers mind can pass to the external language and find expression in
speech. The crucial difference between the two types of explanation lies in the
question of how linguistic elements originate.
The teleological explanation states that internal language elements, distinctions,
and rules develop during the formation of external language in a goal-directed
manner, that is, in principle, intentional, though not necessarily involving conscious
reflection. Words, forms and the rules that govern them develop for functional
reasons. They are created, acquired and passed on in order to satisfy functional
needs. This also applies to previously acquired or generated elements, which are
deployed or modified in accordance with intended goals in full adherence to the
Lamarckian concept of evolution.
Following the Darwinian idea, the teleonomic explanation focuses on the trans-
mission of given structures, during which, for any number of reasons, variations
may arise which are then subject to selection, in this case based on whether a variant
promotes or hinders understanding within the language community. What is left is
to explain the character and origin of the variants, which represent the counterparts
of the genome mutations found in biological evolution. A causal parallel between
biological mutations and language variants, however, can at most be seen in the fact
that copying errors may play a role in both processes. As one of the sources for
variants of the language structure in this sense we again look to language acquisi-
tion, where copying errors can play a systematic role, for example by simplifying,
with the aid of the language faculty, input information during processing. In any
case, the critical point is that the root cause of language variants necessarily lies in
the phenotype sphere, i.e. in the external language, while that of biological
mutations is found in the genome, which is not subject to the causality of the
phenotype. With respect to language, the emergence and selective survival of
variants do not occur in separate spheres characterized by different elements and
principles, but are indeed both based on the same structures.
For this reason we must examine, in unavoidably abridged form, the organiza-
tion of the internal language, which (with the above reservation) is analogous to the
genetic information in living organisms. The two primary aspects of language
faculty underlying every possible internal language enable firstly the creation of
complex symbol inventories and secondly the combination of these symbols based
on recursive rules and principles. Although these two aspects are inseparably
meshed, they are associated with very different types of potential variants, an
adequate elaboration or demonstration of which would stretch beyond the scope
of this paper. A relatively easier task is to examine these two complex phenomena:
1. Lexical units derive from a reservoir of possibilities for differentiating signals,
concepts and combination types.
2. Syntactic principles enable processing strategies, the availability of which is
what essentially constitutes the faculty of language.
In very crude approximation, these phenomena correspond to (1) the conditions
of the lexical system, and (2) the organization of grammar. The resulting
possibilities and options circumscribe a terrain that has been intensively explored
in the cognitive sciences, most particularly by linguists. Notwithstanding the many
unresolved questions, the research does license the conclusion that some structured
leeway is guaranteed in the internal language for variations generated due to
different causes. This applies not only to lexical units (both random and function-
ally motivated variants for naming conceptual complexes), but more importantly to
the morphosyntactic conditions and patterns to which the combinatorics are subject.
But there are still two crucial and related differences compared to variant formation
as seen in biological evolution.
The first difference arises from the fact that the internal language is an expres-
sion derived from and bounded by the language faculty, which for its part is a
product of evolution, but which above all functions as a variation framework for
114 M. Bierwisch
which there is no apparent counterpart in the mechanism of phylogenesis. The

faculty of language thus forms a set of conditions which, from a strictly formal
perspective, represents the counterpart to the totality of biophysical conditions from
which emerge the structures of potential variants in the genome. This means that
variant formation in the realm of the internal language is bounded and determined
within a specific domain and in a particular manner, which itself originated as a
result of evolution.
The second, very closely related difference is decisive here since it violates, as it
were, a central tenet of the Darwinian theory of evolution, namely that variation and
selection belong to different spheres of causality. The variants that can potentially
emerge within the limits of phenomena (1) and (2) such as the differentiation or
reduction of case forms or other morphological categories, changes to lexical units,
the modification of syntactic word order conditions or changes to phonetic
characteristics as witnessed in the sound shift all variants of this type belong to
the same sphere of differentiation and selection in which the external language also
resides. With respect to language change, the genotype and phenotype are thus not
caged within separate spheres of causality, even if the conditions underlying the
selection of relevant traits need not at all coincide with the intentional function of
external language. In other words, the communicative intent is by no means
identical to the chosen means of linguistic communication.
So we are apparently left with a stalemate between the teleological and teleo-
nomic arguments. Not only is it difficult in specific cases to distinguish between
function-based variant formation on the one hand and variant selection based on the
fitness of cause-induced variants on the other, the very basis for making such a
distinction also appears to be nonexistent, since in principle the same conditions
apply to both functionally motivated variation and to selection. But this impression
is incorrect, as is evidenced by the wide variety of structural phenomena in
language that can only be explained teleonomically (based on the conditions of
the language faculty), but by no means teleologically (directed by purpose).
One of many examples backing this conclusion is a typical phenomenon found
in the syntax of various Germanic languages in which the verb must be placed at the
end of the sentence under certain conditions but towards the front under other
conditions. In German, we find cases such as the following sentences (a) and (b):
(a) . . . weil sie den Kurs rechtzeitig beendeten
(. . . because they finished the course in time);
(b) . . . denn sie beendeten den Kurs rechtzeitig
(. . . since they finished the course in time).
Replacing the verb beenden in these examples with the synonym einstellen
reveals a further particularity of German syntax: In words that combine a verb
and a prefix, such as einstellen, aufh oren, anlegen, abtreten and countless others,
only the verb constituent is subject to the described word order variation, despite the
fact that the intended meaning is only produced through the combination of verb
and prefix:
(a) . . . weil sie den Kurs rechtzeitig einstellten

(. . . because they terminated the course in time);
(b) . . . denn sie stellten den Kurs rechtzeitig ein
(. . . since they terminated the course in time).
The difference between (a) and (b) cannot be explained teleologically, and the
structure in (b) would even have to be described as almost malfunctional: The
word einstellen (terminate, close down) is split in the sentence in a way that has
absolutely nothing to do with its meaning. This phenomenon and others like it are
not marginal or secondary. They can be explained teleonomically and systemati-
cally as deriving from the principles of the language faculty, assuming that the
relevant criteria are accordingly defined within the variation limits allowed by the
language faculty. Although the details behind this explanation cannot be presented
here, a number of comprehensive analyses on this topic have been published by
linguistics researchers in recent decades, discussed a. o. in Haiders Deutsche
Syntax [4]. The key finding here is that the organization of the language faculty
provides not only for the principles required for language structuring but also for
leeway or license in permitting variation. This situation thus allows both for
characteristics that better fulfill particular functions and are hence selected as
well as for characteristics that are teleologically unmotivated but which satisfy
the principles of the language faculty. Haider [5] summarized the crux of the matter
in his conclusion that not every why has a wherefore.
This, however, also implies that in context of language change the genotype and
phenotype, in contrast to biological evolution, do not reside in separate domains and
that properties can indeed also emerge and be passed on teleologically. It must be
noted that there is an important difference as to just how this works with respect to
the phenomena indicated in (1) and (2) above (or, more simply, for lexicon and
grammar). The majority of lexical units and changes associated with the lexical
system are motivated more or less directly by function. The whys and wherefores of
their existence merge here, something that is not seen, for example, in the syntax of
verb placement. Certain interferences, however, make it difficult to draw a clear
line between the two phenomena. Germans borrowing of verbs such as updaten or
downloaden can on the one hand be explained as serving specific functional needs
associated with computer technology, but such loans can on the other hand lead to
violations of the German rules of morphology and verb placement.
Unless we are willing to generously and vaguely equate evolution with change,
an interesting and by no means trivial situation arises within the study of language
development. On the one side, the processes of language change and those of
biological evolution are principally different due to the fact that language change
has no analogue to biological evolutions categorical distinction between genotype
and phenotype and hence to the separation of variation and selection associated
with this distinction. On the other, variation and function in the field of language
change are also subject to different conditions that are not reducible to one another,
so that the essential logic behind variation and selection also applies to the features
of the language structure. In the interplay between internal and external language,
116 M. Bierwisch
this logic doubtlessly corresponds to the relationship between genotype and

phenotype.
For certain types of phenomena, however, the teleological explanation is perti-
nent either alongside of or in place of the teleonomic argument. In particular,
terminology can be established by agreement or decision, and such terminology
can subsequently exert an influence on other contexts by virtue of existing
principles. Unlike in biological evolution, such hybrid conditions are possible
here precisely because the two spheres of influence the teleonomic and the
teleological are not mutually exclusive.
Viewed in their entirety, these parallels and differences reveal that language
development is an independent field in which evolutionary and functional factors
work side by side with the capacity for mutual influence and interaction.
5 Epilogue: Social Change and Evolution
If human language history can indeed be logically explained by an evolutionary

theory whose principles are nonetheless essentially different from those that govern
the development of biological species (due to the lack of separate genotype/
phenotype causality), then we are faced with the interesting and pressing question
as to whether we have thus identified a principle of evolution that is universally
applicable to the historical development of sociocultural structures.
Seemingly favoring this supposition is the fundamental role assigned to lan-
guage as a social institution and as a basis and ingredient of veritably every
sociocultural institution, as Searle [9] has recently and distinctly reemphasized.
The symbolic character of natural languages, which first enabled the capacity for
unlimited expression, ties language symbols to agreed convention and hence
inevitably to the population that supports such convention. If, therefore, the manner
in which languages form, divide and transform must be viewed as a special mode of
evolution, it stands to reason that this mode would describe the overall pattern
of shift and change in sociocultural institutions and their underlying knowledge
systems.
That there are nonetheless grounds for reservation here stems from two
considerations. The first concerns what is known as the domain-specificity of the
language faculty. The fact that natural languages are combinatorial systems of
symbols does indeed make them whole and complete in the sense that they can
express any cognitive content, i.e. all that is propositional. And yet they are built
upon combinatorial and representational systems that reveal characteristic
possibilities and limits. The principles of the language faculty are essentially
determined by the conditions required for efficiently processing complex symbol
structures. It is by no means obvious that the type of teleonomic evolution
associated with these phylogenetically generated preconditions can be generalized
to apply to other areas.
The second consideration relates to the intentionality of social behavior, includ-

ing the creativity of language use. For the concept of evolution to apply to language
development there must first exist the conditions of randomness from which
variants arise, conditions that would form the sphere of non-intentionality where
potential selection can take place. But such randomness directly contradicts the
goal directedness usually associated with social behavior. What we absolutely may
not presuppose is the premise that actions generally rest upon a disposition that
corresponds to the specificity of the language faculty, i.e. that they have a biological
foundation in a domain-specific and yet universal faculty of action. The faculty of
language is the prerequisite of human history, but it does not determine its course.
References
1. Chomsky N (1986) Knowledge of language: its nature, origin, and use. Praeger, New York
2. de Saussure F (1916) Cours de Linguistique Generale. Payot, Paris, Lausanne
3. Grimm J (1822) Deutsche Grammatik, Teil 1 Dieterichsche Buchhandlung, G ottingen
4. Haider H (1993) Deutsche Syntax generativ. Narr, T ubingen
5. Haider H (2001) Not every why has a wherefore. In: Bisang W (ed) Aspects of typology and
universals. Akdademie-Verlag, Berlin, pp 3752
6. Humboldt W von (1830) Uber die Verschiedneheit des menschlichen Sprachbaus und ihren
Einfluss auf die geistige Entwicklung des Menschengeschlechtes, Gesammelte Schriften, Band
VII. Akademie-Ausgabe, Preuische Akademie der Wissenschaften, Berlin
7. Pinker S, Bloom P (1990) Natural language and natural selection. Behav Brain Sci 13:707784
8. Schleicher A (1863) Die Darwinsche Theorie und die Sprachwissenschaft. Hermann B ohlau,
Weimar
9. Searle JR (2007) What is language: some preliminary remarks. In: Tsohatzidis SL (ed) John
Searles philosophy of language. Cambridge University Press, Cambridge, pp 1545
Theory of Evolution and Genetics
Alberto Piazza
Abstract The main purpose of this contribution is to document the role of natural
selection, a major factor in Darwinian evolution which is difficult to dissect
specially in the case of human evolution. Recent major steps in human genetics
can be summarized as follows: (a) the complete sequence of the human genome;
(b) the Single Nucleotide Polymorphisms (SNP) discovery and characterization;
(c) the HapMap Project; (d) the Human Genome Diversity Project (HGDP)-CEPH
set-up of 1,000 world-wide cell lines publicly available; (e) the 1,000 genomes
project; (f) The very recent draft DNA sequence of the Neanderthal Genome. An
interesting advance from HGDP-CEPH genome-wide analyses on genetic diversity
and population structure at the world-wide level (high resolution from technologi-
cal advance allowing to type more than 500,000 SNP per individual) has been to
resolve migration from adaptation and natural selection. The tests of selection used
were: (a) iHS detection of partial sweeps in the genome scenario from haplotype
patterns; and, more traditionally, (b) Fst differentiation of gene frequencies,
calculated either globally or between broad geographic regions. Interestingly
enough, the color of the skin problem originally pointed out by Charles Darwin in
The Descent of Man, and Selection in Relation to Sex. p. 381, 1981 facsimile edition
by Princeton University Press, has been elucidated by finding some related traits
(SLC24A5, KITLG, MC1R) whose world geographic distribution shows how
natural selection and geographic barriers are interacting. What makes us humans?
The draft sequence of the Neanderthal genome (Green et al. 2010) surprisingly
shows that the Neanderthal DNA signal can be found not only in the genomes of
Europeans, but also in people from East Asia and Papua New Guinea, where
Neanderthals never lived. Even more of interest, a catalogue of features unique to
the human genome has been pointed out. Five genes affected by two substitutions
A. Piazza (*)
Department of Genetics, Biology and Biochemistry, University of Turin and Human Genetics
Foundation (HuGeF), Turin, Italy
e-mail: alberto.piazza@unito.it

120 A. Piazza
that either change aminoacids or introduce a stop codon, and that have become
fixed among humans since the divergence from Neanderthals have been identified.
The start codon in the gene TRPM1 that is present in Neanderthals and
chimpanzees has been lost in some present-day humans. The TRPM1 encode
melastatin, an ion channel important for maintaining melanocyte pigmentation in
the skin. It is intriguing that skin-expressed genes comprise three out of six genes
that either carry multiple fixed substitutions changing amino acids or in which a
start or stop codon has been lost or gained. This suggests that selection on skin
morphology and physiology may have played an important role on the hominin
lineage. The finding of 20 top candidate selective sweep regions adds further
evidence on how the present technological advances are instrumental for resolving
cases of natural selection in humans too elusive to be found by traditional genetic
tools: the case of skull structure (cleidocranial dysplasia and RUNX2 gene) is
mentioned. Eventually a challenging topic worth of further analyses, the sexual
selection in humans, has been quoted from Charles Darwin in The Descent of Man,
and Selection in Relation to Sex, pp. 605606: in fact, his prophetical thinking has
not been yet fully explored: We cannot positively say that this character, but not
that, has been thus modified; it has, however, been shown that the races of man
differ from each other and from their nearest allies, in certain characters which are
of no service to them in their daily habits of life, and which it is extremely probable
would have been modified through sexual selection.
1 Introduction
The main purpose of this contribution is to document the role of natural selection, a
major factor in Darwinian evolution which is elusive and difficult to dissect,
especially when the case of human evolution is dealt with. In August 1858, Charles
Robert Darwin and Alfred Russel Wallace presented to the Linnaean Society of
London [5] their independent discovery of the theory of natural selection and, in
doing so, they radically altered our understanding of life on Earth. Darwins
meticulously detailed book, On the Origin of Species by Means of Natural Selec-
tion, or the Preservation of Favored Races in the Struggle for Life, published 1 year
later [2], provided a more comprehensive account of his evidence for natural
selection and its role in producing evolutionary change. The immediate and
enduring interest in Darwins and Wallaces work is not only because of the
powerful, unifying, and explanatory theory it provides for biology (and in prospect
for medicine) but also because of its impact on us as humans and on our place in the
natural world. The Origins final paragraph is notable for Darwins insight into
what the volume accomplishes in natural science and his awareness regarding its
impact on its future. Even more remarkable are his predictions on how his theory
would stimulate a search for the causes of variation, presaging the future connection
of genetics and evolutionary theory. The famous passage:
Theory of Evolution and Genetics 121
It is interesting to contemplate a tangled bank, clothed with many plants of many kinds,
with birds singing on the bushes, with various insects flitting about, and with worms
crawling through the damp earth, and to reflect that these elaborately constructed forms,
so different from each other, and dependent upon each other in so complex a manner, have
all been produced by laws acting around us. These laws, taken in the largest sense, being
Growth with Reproduction; Inheritance which is almost implied by reproduction;
Variability from the indirect and direct action of the conditions of life, and from use and
disuse: a Ratio of Increase so high as to lead to a Struggle for Life, and as a consequence to
Natural Selection, entailing Divergence of Character and the Extinction of less-improved
forms. Thus, from the war of nature, from famine and death, the most exalted object which
we are capable of conceiving, namely, the production of the higher animals, directly
follows. There is grandeur in this view of life, with its several powers, having been
originally breathed by the Creator into a few forms or into one; and that, whilst this planet
has gone cycling on according to the fixed law of gravity, from so simple a beginning
endless forms most beautiful and most wonderful have been, and are being evolved.
More than a relevant metaphor for the ecological interrelationship of living

organisms which recalls the lush jungle environment of Darwins early field work
in Brazil and the dense forests that line the banks of the Beagle Channel in Tierra
del Fuego, it is a hymn to the grandeur of the evolutionary perspective, an awe at
the complexity and beauty of the life-forms created in accordance with the opera-
tion of natures laws, freed from metaphysics.
2 Recent Major Steps in Human Genetics
I will focus the attention (in some cases by quoting the relevant web site only) on
the five major advances on the analysis of human evolution which in my opinion
punctuated the field in the last 10 years.
2.1 The Complete Sequence of the Human Genome
Completed in 2003, the Human Genome Project (HGP) was a 13-year project coordi-
nated by the U.S. Department of Energy and the National Institutes of Health. During
the early years of the HGP, the Wellcome Trust (U.K.) became a major partner;
additional contributions came from Japan, France, Germany, China, and others [8].
Project goals were to:
Identify all the approximately 20,00025,000 genes in human DNA;
Determine the sequences of the three billion chemical base (nucleotide) pairs
that make up human DNA;
Store this information in databases;
Improve tools for data analysis;
Transfer related technologies to the private sector;
Address the ethical, legal, and social issues (ELSI) that may arise from the
project.
122 A. Piazza
Though the HGP is finished, analyses of the data will continue for many years. It
is worth noting that the definition of complete sequence referred to the mosaics of
DNA sequences taken from different individuals rather than to the DNA sequence
of one individual. A landmark paper was published in 2008 [10] where the first
high-resolution sequence map of human structural variation in eight individuals has
been showed a prelude to future individual genome sequencing projects.
2.2 The Single Nucleotide Polymorphisms (SNP) Discovery

and Characterization: The HapMap Project
Various scientific endeavors had already started even before the completion of the
first human genome reference sequence to identify unique genetic differences
between individuals. 99.9% of one individual DNA sequences will be identical to
that of another person. Of the 0.1% difference, over 80% will be single nucleotide
polymorphisms (SNPs). A SNP is a single base substitution of one nucleotide with
another, and both versions are observed in the general population at a frequency
greater than 1%. Human DNA is comprised of only four chemical entities, e.g.
A, G, C, T, whose specific chemical order is the alphabet of the genome. An
example of a SNP is individual A has a sequence GAACCT while individual
B has sequence GAGCCT, the polymorphism is an A/G. The most recognized
public effort was spearheaded by The SNP Consortium [9], whose original mission
was to determine and map about 300,000 evenly spaced single nucleotide
polymorphisms within the human genome. Current estimates are that SNPs occur
as frequently as every 100300 bases. This implies in an entire human genome there
are approximately 1030 million potential SNPs. More than 4 million SNPs have
been identified and the information has been made publicly available. Many of
these SNPs have unknown associations. Compilation of public SNPs has produced
a subset of SNPs defined as a non-redundant set of markers that are used for
annotation of reference genome sequence and are thus referred to as reference
SNPs (rsSNPs). Over 2.6 million SNPs have currently been assigned as rsSNPs.
Recent work has suggested that about ten million SNPs that are common in
human population are not inherited independently; rather, sets of adjacent SNPs are
present on alleles in a block pattern, so called haplotype. Many haplotype blocks in
humans have been transmitted through many generations without recombination.
This means although a block may contain many SNPs, it takes a few SNPs to
identify or tag each haplotype in the block. The HapMap Project has the goal to
map such haplotypes along the human genome and provides a very powerful
resource to mark the genome by very stable reference markers to allow comparisons
between different human genomes.
2.3 The Human Genome Diversity Project (HGDP)
A publicly available DNA collection set up and administered by Centre dEtude du

Polymorphisme Humain (CEPH) in Paris, France [1], allowed genome-wide
analyses on genetic diversity and population structure at a world-wide level. Cell
lines of 1,056 individuals from 52 populations have been tested at high resolution
from technological advance: 500,000 SNPs per individual have been typed in an
effort to resolve the effect by migration from that by adaptation and natural
selection in modeling human biological variation [11]. A very simplified summary
of the most recent analyses shows that the genome of each of us differs from the
genome of another individual in about three million nucleotides and that about 95%
of this variation has no effect on genetic characters (phenotypes) but it is influenced
by demographic factors and provides very powerful information to infer the origin
and the migrations of our species. The remaining 5% of human genetic variation
seems to affect phenotypes: how to dissect its possible adaptive role in humans due
to natural selection, it will be shortly explored in the next paragraph.
2.4 The 1,000 Genomes Project
The best way to describe this Project is to quote the abstract of the first publication
by the Consortium The 1,000 Genomes Project [15], an international collaboration
whose goal is to produce an extensive public catalogue of human genetic variation:
The 1000 Genomes Project aims to provide a deep characterization of human genome
sequence variation as a foundation for investigating the relationship between genotype and
phenotype. Here we present results of the pilot phase of the project, designed to develop and
compare different strategies for genome-wide sequencing with high-throughput platforms.
We undertook three projects: low-coverage whole-genome sequencing of 179 individuals
from four populations; high-coverage sequencing of two motherfatherchild trios; and
exon-targeted sequencing of 697 individuals from seven populations. We describe the
location, allele frequency and local haplotype structure of approximately 15 million single
nucleotide polymorphisms, 1 million short insertions and deletions, and 20,000 structural
variants, most of which were previously undescribed. We show that, because we have
catalogued the vast majority of common variation, over 95% of the currently accessible
variants found in any individual are present in this data set. On average, each person is
found to carry approximately 250 to 300 loss-of-function variants in annotated genes and
50 to 100 variants previously implicated in inherited disorders. We demonstrate how these
results can be used to inform association and functional studies. We directly estimate the
rate of de novo germline base substitution mutations to be approximately 10 8 per base pair
per generation. We explore the data with regard to signatures of natural selection, and
identify a marked reduction of genetic variation in the neighborhood of genes, due to
selection at linked sites. These methods and public data will support the next phase of
human genetic research. [16]
124 A. Piazza
2.5 The Draft DNA Sequence of the Neanderthal Genome
A very recent finding (May 2010) which will be explored below.
3 Natural Selection and Population Differentiation
Genetic DNA variants conferring an advantage to better adaptation will be selected

by nature: this process is one ingredient of evolution which distributes genetic
variation in space (population differentiation) and in time as the environment
changes with time. Well known examples are some red-blood-cell disorders (e.g.
sickle cell anemia and thalassemia) which are caused by a pathological DNA
mutation inherited from both parents. The remarkable point is that if this DNA
mutation is inherited from one parent alone not only the disorder is absent, but the
individual who carries it is less susceptible to malaria infection. This better adapta-
tion was strong and detectable still today for malaria, but it is much more elusive for
other traits. Identifying targets of positive selection in humans has, until recently,
been frustratingly slow, relying on the analysis of individual candidate genes.
Genomics, however, has provided the necessary resources to systematically inter-
rogate the entire genome for signatures of natural selection. To date, more than 20
genome-wide scans for recent or ongoing positive selection have been performed in
humans. A key challenge is to begin synthesizing these newly constructed maps of
positive selection into a coherent narrative of human evolutionary history and
derive a deeper mechanistic understanding of how natural populations evolve
[13]. Three questions have been generally addressed:
To which extent has natural selection influenced, at the scale of the entire
genome, the degree of population differentiation in modern humans?
Which type of genetic variants have been preferentially targeted by selection?
Genes and gene variants under strong selective pressures can highlight regions
of the genome explaining the current population phenotypic variation?
One of the parameter to test the effect by natural selection is called Fst, which
measures the differentiation of gene frequencies, calculated either globally at the
world level or between people separated by broad geographic regions.
Empirically: (a) Look more generally at the geographic patterns of SNPs with
extreme Fst; (b) Choose the top ~200 SNPs that differentiate pairs of populations on
the basis of Fst; and (c) Plot the Fst distribution of the genes associated to each SNP
across the Human Genome Diversity Panel populations above. Figure 1 plots the
global Fst distribution of 650.000 SNPs.
It is based on the assumption that selection is sufficiently strong to generate
extreme spatial patterns compared with the rest of the genome. Modified from
Fig. 4b of [12].
Fst Distribution (1% tall)
400
Fst Distribution
50000
300
SLC45A2
Frequency
200
40000
MC1R
KITLG
100
SLC24A5
20000 30000
OCA2
Frequency
0
0.4 0.5 0.6 0.7
SLC24A4 Global Fst
10000
TYR KITLG SLC24A5

SLC45A2 MC1R OCA2
0
0.0 0.2 0.4 0.6 0.8

Global Fst
Fig. 1 Outlier approach for identifying candidate targets of selection
The genes SLC45A2, the most extreme outlier in the figure, and KITLG are
thought to account for ~30% and 20%, respectively, of the difference in dark skin
pigmentation between Bantu Africans and Europeans. At a lesser extent MC1R,
which is found in the 1% tail of the genome-wide distribution, is involved in skin
pigmentation as well. It is a very remarkable finding. Even more remarkable are the
prophetic words by Darwin:
The best kind of evidence that the color of the skin has been modified through sexual
selection is wanting in the case of mankind; for the sexes do not differ in this respect, or
only slightly and doubtfully. On the other hand we know from many facts already given that
the color of the skin is regarded by the men of all races as a highly important element in
their beauty; so that it is a character which would be likely to be modified through selection,
as has occurred in innumerable instances with the lower animals. [4]
4 What Makes Us Humans? The Draft Sequence

of the Neanderthal Genome
Published two weeks before the Meeting, the DNA draft sequence of the Neander-
thal genome signed a major step in the analysis of our species evolution [7].
The first proto-Neanderthal traits appeared in Europe as early as
600,000350,000 years ago. Associated with the Chatelperronian industry, Nean-
derthal man shared their habitat with Homo sapiens and was found in Eastern and
parts of Western Europe, and some areas of Western and Central Asia. In 2008 the
126 A. Piazza
mitochondrial DNA (the mitochondrion is a cellular membrane-enclosed organelle

which generates most of the cells supply of chemical energy) of a Neanderthal was
extracted and completely sequenced [6]. The analysis showed it differs from that of
modern humans but shares a common ancestor with humans around 500,000 years
ago. When compared to modern human it was inferred that Neanderthals donated
little or no mitochondrial DNA to our species. The draft Neanderthal nuclear DNA
sequence has still to be refined, but, despite its limitations, is in any case extremely
useful because human and chimpanzee genomes are available for comparison:
places where humans differ from chimps, while Neanderthals still have the ances-
tral chimp sequence, may represent uniquely human genetic traits. Such
comparisons enabled the researchers to catalog the genetic changes that have
become fixed or have risen to high frequency in modern humans during the past
few hundred thousand years. To help make further informative comparisons, Green
et al. [7] sequenced the genomes of five present-day individuals from different parts
of the world: southern Africa, West Africa, Papua New Guinea, China, and western
Europe. By quoting their words:
Surprisingly the Neanderthal DNA signal shows up not only in the genomes of Europeans,
but also in people from East Asia and Papua New Guinea, where Neanderthals never lived.
The research found the genetic signal of Neanderthals in all the non-African genomes,
meaning that the admixture occurred early on, probably in the Middle East, and is shared
with all descendants of the early humans who migrated out of Africa.
The observation that the Neanderthals genome appears more closely related to the
genome of Asian and European than to that of Africans is particularly striking as there
is, to date, no fossil evidence that Neanderthals existed in East Asia or Papua New
Guinea. Green et al. [7] suggest that between 1% and 4% of the genomes of people in
Eurasia are derived from Neanderthals and therefore gene flow between Neanderthals
and modern humans occurred prior to the divergence of European and Asian
populations (between 70,000 and 50,000 years ago), as sketched in the Fig. 2 below.
5 A Catalogue of Features Unique to the Human Genome
What kind of genes show important changes in recent human evolution? Green
et al. [7] identified a catalog of 72 genetic features unique to modern humans by
comparing the Neanderthals, human, and chimpanzee genomes. Genes involved in
cognitive development, skull structure, energy metabolism, and skin morphology
and physiology are among those highlighted in the study as likely to have
undergone important changes in recent human evolution. The Authors claim to
have identified five genes affected by two substitutions of DNA sequence that either
change aminoacids or introduce a stop codon, and that have become fixed among
humans since the divergence from Neanderthals:
DCHS1 (CCDS7771), which encodes fibroblast cadherin-1, a calcium depen-
dent cell-cell adhesion molecule that may be involved in wound healing.
Fig. 2 Divergence of European and Asina populations between 70,000 and 50,000 years ago
RPTN (CCDS41397), which encodes repetin, an epidermal matrix protein that is

expressed in the epidermis and particularly strongly in eccrine sweat glands, the
inner sheaths of hair roots and the filiform papilli of the tongue.
SPAG17 (CCDS899) sperm-associated antigen-17 that is thought to be impor-
tant for the structural integrity of the central apparatus of the sperm axoneme,
which is important for flagellar movement.
TTF1 (CCDS6948), a terminator of ribosomal gene transcription and regulator
of RNA polymerase I transcription. And
SOLH (CCDS10410), which encodes a protein of unknown function.
To these five genes we could add
The start codon in the gene TRPM1 that is present in Neanderthals and
chimpanzees has been lost in some present-day humans. The TRPM1 encode
melastatin, an ion channel important for maintaining melanocyte pigmentation
in the skin.
It is intriguing that skin-expressed genes comprise three out of six genes that
either carry multiple fixed substitutions changing amino acids or in which a start or
stop codon has been lost or gained. This suggests that selection on skin morphology
and physiology may have played an important role on the hominin lineage.
128 A. Piazza
Table 1 Top 20 candidate selective sweep regions

Region (hg 18) S Width (cM) Gene(s)
chr2:43265008-43601389 6.04 0.5726 ZFP36L2;THADA
chr11:95533088-95867597 4.78 0.5538 JRKL;CCDC82;MAML2
chr10:62343313-62655667 6.1 0.5167 RHOBTB1
chr21:37580123-37789088 4.5 0.4977 DYRK1A
chr10:83336607-83714543 6.13 0.4654 NRG3
chr14:100248177-100417724 4.84 0.4533 MIR337;MIR665;DLK1;RTL1;
MIR493;MEG3;MIR7770
chr3:157244328-157597592 6 0.425 KCNAB1
chr11:3060100-30992792 5.29 0.3951
chr2:176635412-176978762 5.86 0.3481 HOXD11;HOXD8;EVX2;MTX2;HOXD1;
HOXD10;HODX13;HOXD4;HOXD12;
HOXD9;MIR10B;HOXD3
chr11:71572763-71914957 5.28 0.3402 CLPB;FOLR1;PHOX2A;FOLR2;INPPL1
chr7:41537742-41838097 6.62 0.3129 INHBA
chr10:60014775-60262822 4.66 0.3129 BICC1
chr6:45440283-45705503 4.74 0.03112 RUNX2;SUPT3H
chr1:149553200-149878507 5.69 0.3047 SELENBP1;POGZ;MIR554;RFX5;
SNX27;CGN;TUFT1;PI4KB;PSMB4
chr7:121763417-122282663 6.25 0.2855 RNF148;RNF133;CADPS2
chr7:93597127-93823574 5.49 0.2769
chr16:62369107-62675247 5.18 0.2628
chr14:48931401-49095338 4.53 0.2582
chr6:90762790-90903925 4.43 0.2502 BACH2
chr10:9650088-9786954 4.56 0.2475
In the following table (Table 1) a catalogue of the top 20 candidate genome

regions, ordered by decreasing width in centimorgans (cM), the unit of genetic
length, is shown .
They require further, more detailed investigation and correlation with expres-
sion. One gene of interest among them may be RUNX2 (CBFA1) on chromosome 6
which is the only gene in the genome known to cause cleidocranial dysplasia, which
is characterized by delayed closure of cranial sutures, hypoplastic or aplastic
clavicles, a bell-shaped rib cage, and dental abnormalities. Some of these features
affect morphological traits for which modern humans differ from Neanderthals as
well as other earlier hominids. For example, the cranial malformations seen in
cleidocranial dysplasia include frontal bossing, i.e. a protruding frontal bone.
6 The Sexual Selection in Humans: A Challenging Topic

Worth of Further Analyses
The views here advanced, on the part which sexual selection has played in the history of
man, want scientific precision. He who does not admit this agency in the case of the lower
animals, will disregard all that I have written in the later chapters on man. We cannot
positively say that this character, but not that, has been thus modified; it has, however, been
shown that the races of man differ from each other and from their nearest allies, in certain
characters which are of no service to them in their daily habits of life, and which it is
extremely probable would have been modified through sexual selection. We have seen that
with the lowest savages the people of each tribe admire their own characteristic qualities,
the shape of the head and face, the squareness of the cheek-bones, the prominence or
depression of the nose, the color of the skin, the length of the hair on the head, the absence
of hair on the face and body, or the presence of a great beard, and so forth. Hence these and
other such points could hardly fail to be slowly and gradually exaggerated, from the more
powerful and able men in each tribe, who would succeed in rearing the largest number of
offspring, having selected during many generations for their wives the most strongly
characterized and therefore most attractive women. For my own part I conclude that of
all the causes which have led to the differences in external appearance between the races of
man, and to a certain extent between man and the lower animals, sexual selection has been
the most efficient. [3])
The quotation above, as many conjectures by Darwin, includes a series of work

hypotheses very plausible for animals but difficult to test for humans, especially in
modern times when cultural factors on sexual selection may shadow biological
pressures. The technological advances in analyzing thousands of genes at the same
run make it more affordable to pick them out today and appreciate once more
Darwins revelatory vision of nature and of man in it. The full understanding of why
sex is evolutionary advantageous in our species, requires still some time to wait, but
if we are less ambitious and like to understand minor but biologically interesting
aspects of sex appeal (see for instance the popular account by [14]) a rich literature
is available to us: a further evidence, among many others, of the impact the
evolutionary key has and will have for interpreting our past and challenging our
future.
References
1. Cann H et al (2002) A human genome diversity cell panel. Science 296:261262

of favoured races in the struggle for life, 1st edn. John Murray, London
3. Darwin CR (1871/1874) The descent of man and selection in relation to sex, 2nd edn. John
Murray, London, pp 605606
4. Darwin CR (1981) The descent of man and selection in relation to sex. Princeton University
Press, Princeton, p 381 [1871 facsimile edition]
5. Darwin CR, Wallace AR (1858) On the tendency of species to form varieties; and on the
perpetuation of varieties and species by natural means of selection. Proc Linn Soc Lond Zool
3:4650
6. Green RE, Malaspinas AS, Krause J et al (2008) Complete Neanderthal mitochondrial genome
sequenced determined by high-throughput sequencing. Cell 134:416426
7. Green RE, Krause J, Briggs AW et al (2010) The draft sequence of the Neanderthal genome.
Science 328:710722
8. Human Genome Project Information (s.d.) http://www.ornl.gov/sci/techresources/Human_Genome/
home.shtml
9. International HapMap Project (s.d.) http://snp.cshl.org/
130 A. Piazza
10. Kidd JM, Cooper GM, Donahue WF et al (2008) Mapping and sequencing of structural
variation from eight human genomes. Nature 453:5664
11. Li JZ et al (2008) Worldwide human relationships inferred from genome-wide patterns of
variation. Science 319:11001104
12. Novembre J, Di Rienzo A (2009) Spatial pattern of variation due to natural selection in
humans. Nat Rev Genet 10:745755
13. Pickrell JK, Coop G, Novembre J et al (2009) Signals of recent positive selection in a
worldwide sample of human populations. Genome Res 19:826837
14. Roach M (2005) Roots of desire: the myth, meaning and sexual power of red hair. Bloomsbury,
New York
15. The 1000 Genomes Project (s.d.) www.1000genomes.org
16. The 1000 Genomes Project, Consortium (2010) A map of human genome variation from
population-scale sequencing. Nature 467:10611073
Genes, Evolution and the Development
of the Embryo
Giuseppina Barsacchi
Abstract Evolutionary Developmental Biology (Evo-Devo) deals with the

relationships between the individual development and the phenotypic changes of
the organism during evolution. Major morphological transitions in evolution are
presently recognized to be accommodated by a few key developmental genetic
changes (part of a developmental reprogramming) and case studies in snakes,
ducks, bats, dolphins, insects, and finches, providing examples of develop-
mental bases of evolutionary change, are presented. On the other hand, the
molecular changes occur in an otherwise conserved developmental genetics
tool-kit (e.g., the Hox genes for anterior-posterior patterning, the network for
eye formation) representing the deep homology underlying diversity of forms.
Based on a relationship between embryo development and organism evolution,
Evo-Devo represents a synthesis between Developmental and Evolutionary
Biology.
1 Evo-Devo: A New Discipline with Darwinian Roots
Evolutionary Developmental Biology, usually nick-named Evo-Devo, explores

the relationships between the processes of individual development and the pheno-
typic changes of the organism during evolution. Evo-Devo represents a synthesis
between Developmental and Evolutionary Biology, based on a relationship
between embryo development and organism evolution. In fact developmental
reprogramming [1] may generate the phenotypic diversity the Darwins varia-
tion which natural selection can act upon producing evolutionary novelties.
G. Barsacchi (*)
Lab. of Cell and Developmental Biology, Department of Biology, University of Pisa, Pisa, Italy
e-mail: gbarsacchi@biologia.unipi.it

132 G. Barsacchi
Academically, Evo-Devo was born around year 2000, when the first dedicated
Journals and Societies were established, but it is however rooted in an ancient and
noble tradition, including Darwin himself. Indeed, Charles Darwin fully appreciated
the relationship between embryology and evolution. In a letter to the American
botanist Asa Gray (1860) [2] he praises the role of Embryology in providing evidence
for the change of forms: Embryology is to me by far the strongest class of facts in
favor of change of forms. In a complementary manner, in On the origin of species
(1859) [3] Darwin recognizes that Community of embryonic structure reveals the
community of descent. Thus, Darwin fully appreciates that embryo development
can provide evidence for both conserved structures (the community of embryonic
structure), and changes affecting those structures (changes of forms) in due time
and circumstances.
Through his own work Darwin demonstrates that similarities between embryos
may reveal phylogenetic relationships, thus contributing to the newborn nineteenth
century field of Evolutionary Embryology a precursor discipline to Evo-Devo.
For instance, in his huge monograph on Cirripedia Darwin easily adopts Embryol-
ogy as a methodological tool to reveal homologies: by discovering that the larva of
the barnacles looks like that of the shrimps (nauplius), he establishes that the
barnacles are not Mollusks but Crustaceans and is able to revise the erroneous
taxonomy of this group of Invertebrates [4]. In fact Darwin considered the
similarities between embryos, as compared to the dissimilarities between adult
forms, one of the strongest arguments in support of his theory of evolution.
Another example of nineteenth century Evolutionary Embryology prompted an
enthusiastic reaction from the very same Charles Darwin, who, in The Descent of
Man declares [5]: Some observations lately made by M. Kowalevsky will form a
discovery of extraordinary interest. The discovery is that the larvae of Ascidians are
related to the Vertebrata. . . a great discovery indeed, since, based on the adult
morphology, Ascidians were classified either as Mollusks devoid of a shell or as
Worms. Thus, Embryology reveals the proximity of Ascidians (Tunicates) to
Vertebrates . . .in their manner of development, in the relative position of the
nervous system. . . that is dorsal in the larvae of Ascidians as it is in Vertebrates
and in possessing a structure closely like the chorda dorsalis of vertebrate
animals. The following is the Darwins conclusion: It thus appears. . .if we may
rely on embryology, which has always proved the safest guide in classification that
we have at last gained a clue to the source whence the Vertebrata have derived.
Again, here Darwin pays a great tribute to Embryology, but, more importantly, he
recognizes that classifying the Ascidians as Chordates may pave the way to enquire
into the origin of Vertebrates:
We should thus be justified in believing that at an extremely remote period a group of
animals existed, resembling in many respects the larva of our present Ascidians, which
diverged into two great branches, the one retrograding in development and producing the
present class of Ascidians, the other rising to the crown and summit of the animal kingdom
by giving birth to the Vertebrata.
Genes, Evolution and the Development of the Embryo 133
Fig. 1 Thomas H. Huxley

and Julian Huxley (1893)
Recent genomic data have confirmed and extended the Kowalevskys discov-
ery: Tunicates are not only relatives of Vertebrates, but surprisingly they and
not Cephalochordates appear to represent the closest living relatives of
Vertebrates [6].
The great Thomas H. Huxley (18251895; Fig. 1), a strenuous advocate of the
Darwins theory of evolution, also elaborates on the concept of the relevance of
Embryology for Evolution and in a very sharp sentence asserts [7]: So far as
individual plants and animals are concerned, therefore, evolution is not a specula-
tion but a fact; and it takes place by epigenesis. The last part of this statement
interests us most, since epigenesis here means the building of a body structure
through embryo development.
To complement the theory of variation and natural selection, we therefore
would need a theory on how a body is built through development: a theory to
explain how development can generate those changes in anatomy, which selec-
tion can act on. However, a sound understanding of embryo development that
could be formulated into a coherent theory was not available at the Thomas
Huxleys time and was only made possible after the impressive twentieth century
achievements of classic and molecular genetics and of molecular and cell
biology.
In the meantime, in 1942 another Huxley (Julian, Fig. 1) had modernized his
grandfather view by suggesting: A study of the effects of genes during devel-
opment is as essential for an understanding of evolution as are the study of
mutation and that of selection [8]. The present genetic program of Evo-Devo,
on which this paper is focused, is complying with the Julian Huxleys
expectation.
134 G. Barsacchi
2 Genes, Development and Evolution: The Genetic

Program of Evo-Devo
The concepts expressed by Francois Jacob in the 1970s, stemming from his
discoveries on gene regulatiom in Prokaryotes, have oriented our thoughts on
which genes to look for in the Evo-Devo perspective: these are regulatory
genes genes that stand high in the hierarchy of gene activities and control the
function of many other genes. For example, a class of regulatory genes codes for
transcription factors, proteins capable of binding DNA and of generating regu-
latory circuits or gene networks that control the activity of many other genes. In
1977 Jacob wrote [9]:
The genetic program is executed through complex regulatory circuits that switch the
different biochemical activities of the organism on or off. Very little is known as yet
about the regulatory circuits that operate in the development of complex organisms. (. . .)
It seems likely that divergence and specialization of mammals, for instance, resulted from
mutations altering regulatory circuits rather than chemical structures.
The present knowledge of the developmental regulatory circuits is enormously

increased with respect to the 1970s but, as it will be shown later in this chapter (see
paragraph 5) the Jacobs general conclusion holds true. In fact, for Jacob evolution
works by tinkering with regulatory genes:
It is always a matter of using the same elements, of adjusting them, of altering here or
there, of arranging various combinations to produce new objects of increasing complexity.
It is always a matter of tinkering. [9]
Evo-Devo has demonstrated two unexpected phenomena concerning develop-

mental regulatory genes, which represent tools for tinkering.
Firstly, the accumulation of molecular and genetic data uncovered an incredible
similarity of developmental regulatory genes throughout animals. In general, the
conserved regulatory genes have a special function in development: they are
engaged in determining the overall body plan and the number, identity, pattern of
body parts [10]. They therefore constitute the genetic tool-kit that controls
development of an animal from an egg into an adult, driving formation of specific
features. In the developmental tool-kit, genes do not act individually but as
networks of interacting genes, functioning in concert to deploy a given develop-
mental function.
Most tool-kit genes code for two classes of gene products with the most
global effects on development: signaling pathway factors and transcription
factors. The two classes of gene products are functionally connected, since the
signal pathway factors link paracrine factors acting outside the cell to
transcription factors acting inside the cells nucleus (Fig. 2). The signal
transduction pathways the chains of biochemical signaling linking the cell
environment to the cell nucleus are highly conserved in different Phyla.
Conservation of signaling pathway factors and transcription factors is both a
result of, and a substrate for, tinkering.
PARACRINE FACTOR
SIGNAL TRANSDUCTION PATHWAY
TRANSCRIPTION
FACTOR
Gene
Nucleus Enhancer
Fig. 2 The tool-kit genes encoding paracrine factors, signaling pathway factors and transcription
factors are highly conserved in different animal Phyla (Kind gift from Prof. S.F. Gilbert. Modified
from devbio8e-fig-06-13-2.jpg)
As examples, are parts of the developmental genetic tool-kit:

Genes that control formation of the body axes, such as the Hox genes for the
Anterior-Posterior (A-P) axis (see paragraph 4) and the chordin/BMP4 genes for
the Dorsal-Ventral (D-V) patterning;
The Otx and Emx genes, which are involved in anterior patterning and
cephalization;
Pax6 that, in a network with other genes, drives eye development (see
paragraph 3);
Distal-less, which controls appendage formation (see paragraph 5, Heterotypy);
and many more: developmental tool-kit genes are extensively discussed in [10, 11].
Many discoveries have now confirmed the great extent of conservation of the tool-kit
genes across different Phyla, despite the differences between a human and a fly eye
or brain or leg.
136 G. Barsacchi
The flexibility of gene regulation is the second phenomenon allowing and

underlying tinkering. Developmental genes are regulated by multiple DNA
sequences, the enhancers, which can each control the gene activity in a different
body region. Enhancers are like the different keys of a piano that can produce
different melodies according to the pattern with which they are played. For example,
the different enhancers of the Pax6 gene can activate Pax6 expression either in the
pancreas, or in the lens and cornea, or in the neural tube, or in the retina. Thus,
modularity of enhancers allows gene expression to be changed in one organ without
changes affecting other organs. Modularity of enhancers is a perfect example of the
effects and of the potentialities of tinkering.
3 The Concept of Deep Homology and the Case

of Eye Evolution
Based on the discovery of these two phenomena conservation of the develop-

mental genetic tool-kit and flexibility of gene regulation the concept of deep
homology has been proposed to indicate the conserved genetic regulatory
apparatuses that are used to build specific animal features during development
and the features produced by deeply homologous genes can be morphologi-
cally and phylogenetically disparate [12]. On this view, deep homology is
deeply buried inside the cells and not necessarily manifest in the surface of
morphology.
Perhaps the eye represents the most famous case that fits with the deep
homology description: eyes are morphologically disparate and their evolutionary
origin represents a conceptual platform for different views since Darwin. In On
the Origin of species Darwin devotes a whole paragraph of Chap. 6 to the
question of the possible origin of the eye by natural selection [3]. Significantly,
Chap. 6 is entitled Difficulties on Theory and the paragraph dedicated to the eye
is entitled Organs of extreme perfection. To some extent rhetorically, Darwin
asks: Can we believe that natural selection could produce. . . organs of such
wonderful structure, as the eye, of which we hardly as yet understand the inimita-
ble perfection?.
Darwin appears to be fully aware of the difficulties concerning eye evolution.
He approaches this subject carefully and is even sympathetic with skeptics: . . .
though I have felt the difficulty far too keenly to be surprised at any degree of
hesitation but nevertheless he thinks he can recognize a graduated series of
variations in the eyes of some animal groups (i.e., Crustaceans) supporting the
idea that all eyes evolved by natural selection, starting from a simple ancestral
eye.
At variance, based on a wide morphological, structural and embryological
comparison, the influential work of Salvini-Plawen and Mayr [13] proposes that
eyes originated through 4065 independent events during evolution.
3.1 The Genetic Network for Eye Development
Here is where the genetic program of Evo-Devo comes into play. Indeed, over the
past 15 years many insights into the evolution of eyes came from descending beneath
the visible diversity of animal eyes into the genetic machinery that controls their
development, providing us with a textbook example of deep homology [12, 14].
Homologous transcription factor genes are co-expressed in the embryo territories
(eye fields) fated to generate the eye in both Drosophila and vertebrates; furthermore,
the individual genes are connected in similar genetic networks in both invertebrates
and vertebrates, even though these networks specify formation of eyes as different as
the compound eye of insects and the camera eye of vertebrates [15, 16] (Fig. 3). The
networks include transcription factors encoded by members the eyeless, sine oculis
and eyes absent gene families in Drosophila and their homologous in vertebrates.
+ Tll, Optx2 (late)
toy
ey
eya so
dac
Fig. 3 The genetic network for eye specification is conserved between invertebrates and vertebrates.
Top schemes: the expression domains of the transcription factors specifying the eye in Drosophila
(left) and Xenopus (right) are outlined in different colors. Eye specification requires co-expression of
transcription factors in space and time. Middle schemes: proposed gene networks for eye specification
in Drosophila (left) and Xenopus (right). Bottom left: a Drosophila compound eye (From [14],
courtesy of Prof. W. Gehring, with permission from Elsevier). Bottom right: the R. Magritte Le
faux miroir is used to symbolize the vertebrate eye. Top and middle schemes from [15, 16] for
Drosophila and Xenopus, respectively
138 G. Barsacchi
Jellyfish PaxB into fly Squid Pax6 into fly
antenna
antenna wing
wing
Drosophila eyeless/Pax6 into fly Mouse Pax6 into fly
leg leg
leg
Fig. 4 The PaxB/Pax6 gene of either jellyfish, squid or mouse elicit ectopic eye formation in
transgenic flies (Drosophila) (From [17, 18, 20], respectively; courtesy of Prof. J. Piatigorsky and
Prof. W. Gehring; with permission from Elsevier, P.N.A.S. (Copyright 1997 National Academy of
Sciences, U.S.A.) and Oxford University Press). The ectopic eye (arrowhead) generated on a
Drosophila leg by the eyeless/Pax6 gene of Drosophila is also shown (From http://www.dnalc.org/
view/16772-Gallery-37-Drosophila-eyeless-mutant. Courtesy of Prof. W. Gehring)
Conservation of individual genes in the eye networks is so extreme, that the

PaxB/Pax6 genes of a jellyfish [17], or a squid [18], or an ascidian [19], or a mouse
[20], are each able to elicit formation of ectopic eyes in transgenic Drosophila flies
on an antenna, or on a wing or on a leg, of the fly and all these ectopic eyes have
the structure of the compound eye of Drosophila, regardless of the provenance of
the Pax gene (Fig. 4). Thus, formation and evolution of morphologically disparate
eyes depends on homologous genetic regulatory circuits.
3.2 The Origin of Photoreceptors
The conservation of the molecular genetics underlying eye formation, contrasting

with the diversity of eyes, has resuscitated the question of the evolutionary origin of
eyes but at different levels: both at the level of the regulatory genetic mechanisms
underlying eye development as briefly outlined above and at the level of
individual cell types. Special attention is devoted to photoreceptors, the retinal
cells capable of capturing light initiating the process of image formation.
Two main types of photoreceptors, rhabdomeric and ciliary, exist in the different
taxa [21, 22] (Fig. 5a). The two kinds of photoreceptors are differentially
distributed among animals, with a prevalence of rhabdomeric photoreceptors used
for vision in invertebrates and with the ciliary photoreceptors being the only type of
photoreceptors employed for image-forming vision in the vertebrate eye. The two
types of photoreceptors differ as for their structure and morphology; they also use
opsins of different gene families (r-opsins or c-opsins for rhabdomeric or ciliary
photoreceptors, respectively) and distinct phototransduction cascades. Their dis-
tinct cellular type and distribution has represented an argument in favour of the
separate origin of the different types of eyes.
It is now clear that bilaterian animals typically have both types of photoreceptor
cells, but it depends on the phylum as to which type is used for vision. Ciliary
photoreceptors with a vertebrate-type opsin have been discovered in an invertebrate
brain: while the marine ragworm Platynereis dumerilii hosts rhabdomeric
photoreceptors in its eyes, ciliary photoreceptors are present in its brain [23] and
this might be a common trait in many invertebrate phyla (Fig. 5b). Based on such
significant discovery, the Authors propose that early metazoans possessed a single
type of precursor photoreceptor cell that used an ancestral opsin for light detection,
photoperiodicity control and possibly phototaxis. In prebilaterian ancestors, dupli-
cation of the opsin gene into c-opsin and r-opsin allowed the diversification of the
precursor into ciliary and rhabdomeric cell types. Accordingly, urbilaterians the
last common ancestors of bilaterians possessed both ciliary and rhabdomeric
photoreceptors. In the evolving bilaterians and similarly to the setting in
Platynereis, the rhabdomeric photoreceptors associated with pigment cells to form
simple eyes, while the ciliary photoreceptors formed part of the evolving brain,
active in non-directional photoresponse. In the evolutionary line leading to
vertebrates, both photoreceptor types were incorporated into the evolving retina:
the rhabdomeric photoreceptors transformed into ganglion cells (see below), while
the ciliary photoreceptors became the main visual cell types, the rods and cones.
Molecular fingerprinting of the specific retinal cell types [24] supports this view.
Also, the derivation of rods and cones from brain ciliary photoreceptors strengthens
the view that the vertebrate retina evolved as an outfolding from the brain, as
recapitulated during the development of the present day vertebrate eye [24].
140 G. Barsacchi
a b
ciliary photoreceptor
in Platynereis brain
rhabdomeric ph. ciliary ph.
c d melanopsin
Cubomedusae mouse retina
Fig. 5 (a) Schematic representation of the major morphological difference between a

rhabdomeric and a ciliary photoreceptor: the apical cell membrane or the cilium membrane is
expanded in either cell type, respectively (From [21]; courtesy of Dr. D. Arendt and with
permission from The Royal Society). (b) Scheme of a ciliary photoreceptor as found in the
Platynereis brain, based on EM images (From [23]; courtesy of Dr. D. Arendt and with permission
from the A.A.A.S.). (c) The lens eyes of Tripedalia cystophora possess a retina (r) endowed with
ciliary photoreceptors and expressing PaxB (in pink). l: lens. Arrowheads indicate pigment layer
(Modified from [17]; courtesy of Prof. J. Piatigorsky and with permission from Elsevier). (d)
Section of a mouse retina where melanopsin expressing retinal ganglion cells (M1, M2) are
highlighted by immunostaining. Arrows and asterisks indicate two plexuses in the inner plexiform
layer (IPL). GCL ganglion cell layer, IPL inner plexiform layer, INL inner nuclear layer, OPL outer
plexiform layer, ONL outer nuclear layer (From [27]; courtesy of Authors)
Vertebrate eye components have also even been found in the jellyfish eyes [25].
The Cubomedusae (box jellyfishes, e.g. Tripedalia cystophora), tiny jellyfishes
about 1 cm3 large, possess amazing camera-type eyes that host a retina endowed
with ciliary photoreceptors (Fig. 5c). These photoreceptors express c-opsin;
c-opsin, but not r-opsin, has been found in other cnidarians eyes as well [26].
The Tripedalia eyes also use melanin as a shielding pigment, the same as used in
vertebrate eyes. Melanin is enclosed into the cytoplasm of the Tripedalia
photoreceptors, so that the jellyfish photoreceptors appear to combine in just one
cell the characteristics of a photoreceptor and of a pigmented cell, that are instead
split in distinct cells in the vertebrate eyes.
Conversely, and significantly, the vertebrate retina appears to host the

descendents of the invertebrate rhabdomeric photoreceptors disguised as retinal
ganglion cells (RGCs). A subset of RGCs are intrinsically light-sensitive and
mediate perception of day/night cycle: they express melanopsin, an opsin of the
same family as the rhabdomeric r-opsin [27] (Fig. 5d). In addition, precursors of
vertebrate RGCs express transcription factors homologous to those expressed by
precursors of rhabdomeric photoreceptors in the invertebrate eye (e.g., Atonal/
Math, Bar/BarH1) [28, 29]. The Atonal/Math transcription factors are critically
required for both rhabdomeric and ganglion cell formation in the Drosophila and
vertebrate eye, respectively: thus, two evolutionarily diverse eye types require
homologous transcription factors for retinal neuron formation [30]. The whole of
evidence points to a transmutation of rhabdomeric photoreceptors, engulfed into
the vertebrate eyes, to originate retinal ganglion cells that play a major role in image
processing and in entraining the circadian clock to the ambient lighting conditions.
In conclusion, the vertebrate eye represents a composite structure, combining
distinct types of light sensitive cells, the rhabdomeric and ciliary photoreceptors.
3.3 Deep Homology of Eye Development and the Parallel

Evolution of Animal Eyes: A Model
Since both ciliary and rhabdomeric photoreceptors are found in both invertebrates
and vertebrates, it now seems possible to reconstruct the light-sensitive systems that
were present in the ancestors of all animals (Urmetazoa) or of all bilateral animals
(Urbilateria), but we are as yet in an early phase of this reconstruction, facing a few
possible models.
Based on the concept of deep homology and on the information presently
available, a recent model [12] proposes that (Fig. 6):
The Cnidarian-Bilaterian common ancestor had photoreceptors expressing
c-opsin and a Pax gene (PaxB; Fig. 6a);
After the split between the cnidarian and bilaterian lineages, in ancestral-stem
bilaterians rhabdomeric photoreceptors evolved, expressing r-opsin and Pax6
(Fig. 6b);
Urbilateria, the last common ancestors of all Bilaterians, probably had two types
of light-sensing organs: a prototypical eye and a brain photo-clock, with
rhabdomeric and ciliary photoreceptors, respectively, as they are presently
found in the ragworm Platynereis (Fig. 6c);
The photoreceptor types established in the Urbilateria were then incorporated in
different ways in the parallel evolution of diverse eyes in different Phyla
(Fig. 6d). Rhabdomeric photoreceptors were the foundation for the evolution
of compound and camera-type eyes in arthropods and mollusks, respectively.
142 G. Barsacchi
a e
Cnidarian-Bilaterian Animal photoreceptor- Jellyfish
ancestor cell precursor camera eye
(PAXB, c-opsin) (ciliary)
PAX6, r-opsin
Ancestral-stem Rhabdomeric Ciliary

bilaterians photoreceptor photoreceptor
c
Urbilateria Prototype Brain
eye photo-clock
Arthropod Squid Vertebrate

compound eye camera eye camera eye
(rhabdomeric) (rhabdomeric) (ciliary)
Fig. 6 Deep homology of eye development and the parallel evolution of animal eyes: a model
(see text for explanations) (From [12]; courtesy of Authors and with permission from Nature
Publishing Group)
Both types of photoreceptors were incorporated into the vertebrate eye, with
ciliary photoreceptors carrying out photo transduction and rhabdomeric
receptors being transformed into ganglion cells, functioning in image processing
and circadian entrainment;
The ciliary camera-type eyes of box jellyfishes are proposed to have evolved in
parallel in the cnidarian lineage (Fig. 6e).
In conclusion, the developmental molecular genetics of the diverse eyes deep

homology illustrates that similar tools have been used to build a great variety of
structures long thought to have completely independent histories.
4 The Hox Genetic Tool-Kit Specifies AnteriorPosterior

Polarity Through Animals
The Hox genes specify the regional identity of the anterior-posterior (A-P) body
axis in the most diverse animals. The roots of our knowledge on Hox genes lie in the
observations and ideas presented by William Bateson (18611926) in his book on
Materials for the study of variation [31], where he reported on hundreds of
changes from the normal morphology of a species that could help in understanding
how evolution worked. In fact he believed that frank observations of variations
present in nature were the common basis for all views on evolution.
Bateson dedicated special attention to those changes where a structure was
transformed, or formed, as to take the appearance of another existent structure:
for instance, when an insect antenna had become similar to a leg; or a frog first
vertebra, the atlas, had acquired processes like those of other vertebrae; or an
additional vertebra was present in a frog; or a second head had formed in a turtle.
To describe such weird variations he suggested a new word:
I think that a new term is demanded. (. . .) the term Homeosis (. . .) for the essential
phenomenon is not that there has merely been a change, but that something has been
changed into the likeness of something else.
Thus, the concept and the word homeosis were born to stay with us up to the
present. Bateson predicted that:
. . . this particular kind of variation will be found to be especially important and I believe
that in the future its significance and the mode of its occurrence will become an object of
high interest.
He was right: homeosis is due to mutations in homeotic/Hox genes which from

homeosis draw their name and the study of Hox genes has been providing us with
a wealth of information as to how animal form is achieved; a real gold mine for
scientists [32, 33].
Hox genes are evolutionarily conserved from Cnidarians to Vertebrates and are
grouped in a single cluster in the genome of most animals (Fig. 7a). Over the course
of vertebrate evolution, the single ancestral cluster underwent duplications to
produce four gene clusters, which together with gene loss resulted in the 39 Hox
genes found in all extant mammals. Each ideal cluster contains protein-coding
genes all transcribed in the same 50 -to-30 orientation, allowing the clusters to be
considered as having a 50 and a 30 end. This unique chromosomal organization
facilitates coordination of Hox-cluster expression, which is characterized by spa-
tial and temporal colinearity: genes located at the 30 end of the cluster are
expressed more anteriorly and earlier, whereas genes more 50 are expressed pro-
gressively more posteriorly along the A-P axis and at later stages of development
[34]. Colinearity establishes a partially overlapping, staggered arrangement of the
Hox genes domains of expression along the A-P axis (see also [32, 33] for variations
from ideal clustering and colinearity).
144 G. Barsacchi
b
Drosophila
M
A
M
M
A
L
S
Fig. 7 (a) Cladogram representing Hox gene chromosomal organization for representative
animals. At the base is shown a cnidarian (Nematostella vectensis), which has a dispersed genomic
organization of Hox genes. The left branch displays fragmented Hox clusters for both
lophotrochozoan (Schistosoma mansoni) and ecdysozoan (Drosophila melanogaster,
Caenorhabditis elegans) species. The right (deuterostome) branch portrays the Hox cluster of
the sea urchin Strongylocentrotus purpuratus, the prototypical Hox cluster of Branchiostoma
floridae (a cephalochordate), the dispersed genomic organization of the Hox genes of a urochor-
date (Oikopleura dioica), and the quadruplicated Hox clusters of a mammal (Mus musculus). At
the base of the cladogram is the likely Hox cluster organization of the last common ancestor of
bilaterians (Urbilateria) (From [32]; courtesy of Prof. W. McGinnis, with permission from A.A.A.S.).
(b) Depiction of the Hox gene (arrows) complexes of Drosophila and mammals. Loci encoding
microRNA are also indicated (red arrowheads). microRNA genes are conserved between Dro-
sophila and mammals and inhibit translation of more anterior Hox mRNA (red lines) (From [34];
courtesy of Prof. E.M. De Robertis and with permission from Elsevier)
From Cnidarians to Vertebrates the Hox genes shape the A-P axis of the body by
specifying, during embryo development, identity and order of structures in the most
diverse animals, either flies or mice or humans: where to build the head, where the
tail, and where the structures in between. Hox genes accomplish such a sophisti-
cated job by undergoing a complex spatial-temporal pattern of expression during
development.
It is emerging that microRNAs are major players in the complex molecular
machinery of Hox gene regulation. These short cellular RNA sequences about 20
nucleotides long can specifically bind messenger RNA (mRNA) molecules,
inhibiting their translation into a protein. It has been found that some microRNAs
can bind specific Hox mRNA molecules blocking their translation in specific body
regions [32, 34, 35]. In the Evo-Devo context it is of importance that this lately
discovered microRNA-based mechanism appears to be conserved between
invertebrates and vertebrates. Strikingly, in both Drosophila and Vertebrates the
Hox genomic clusters encapsulate conserved sites encoding microRNAs, which are
transcribed by the same promoters of posterior Hox genes and repress the activity
of more anterior Hox genes in the cluster, in posterior body regions (Fig. 7b).
Thus, regulation by microRNA of the Hox genes way of functioning appears to
provide a molecular explanation for the enigmatic phenomenon of the posterior
prevalence discovered by E.B. Lewis on a purely classical genetics ground.
In accordance with the variations discovered by Bateson, when mutations affect
the complex regulation of Hox gene expression, regions along the A-P axis of the
body are transformed to the likeness of other regions [10, 11]. Famous examples
include an improbable dipterous endowed with two pairs of wings, due to the
transformation of the halters into an extra-pair of wings (bithorax); and a fly
carrying a pair of legs on its head, where they replace the antennae (antennapedia).
Mutations in Vertebrates include the appearance of an extra-thoracic vertebra in
mouse, or of an extra-digit in man. Thus, homeotic mutations transform regions
along the A-P axis into one another in the most diverse animals.
4.1 Hox Genes and the Possible Origin of the Snake

Vertebral Column
Homeotic mutations provide us not only with special phenotypes, but also with
hints as to how changes in Hox genes may have generated variations along the A-P
axis during evolution. How, for instance, may the elongated and limbless body of a
snake have diverged from the tetrapod lineage?
By observing the vertebral column in diverse animals (Fig. 8), we notice that
100% of vertebrae have a thoracic identity in the snake and, as such, they carry ribs;
the python has virtually no cervical vertebrae and therefore lacks a neck [36].
Furthermore, the distinct types of vertebrae cervical, thoracic, lumbar etc. are
in different numbers in different species: the cervical vertebrae are 14 in the chick,
146 G. Barsacchi
a cervical thoracic lumbar sacral caudal

7
Hoxc5
14
Hoxc5
17
Hoxc5
Fig. 8 Differences in the vertebral patterning correlate with shifts in Hox gene expression
domains. (a) The anterior boundaries of expression of several Hox genes are shown beneath
somites (circles) and vertebrae (squares). The different identity of vertebrae is matched by the
different colors, as indicated on top of figure. In mammals and birds the anterior boundary of the
Hoxc6 expression lies at the cervical-thoracic transition, where the forelimb buds can form. At
variance, in the python both Hoxc6 and Hoxc8 have a more anterior expression boundary,
reflecting the expanded thoracic vertebral identity and no forelimbs develop (see b) (From [10];
courtesy of Prof. S. Carrol and with permission from Wiley-Blackwell). (b) Lateral view of a
complete skeleton preparation of python embryo at 24 days of incubation. Arrow indicates position
of hindlimb rudiments (removed). The vertebrae anterior to arrow are homogeneous (From [36];
with permission of Nature Publishing Group)
17 in the goose and 7 in virtually all mammals. As a consequence, the transitions

from one vertebral type to another occur at different levels with respect to the head
in the different animals.
The diverse vertebral anatomical patterns correlate with the expression patterns
of Hox genes observed in the presumptive axial skeleton of the embryos. As
examples (Fig. 8a), the Hoxc5 gene is expressed in the presumptive cervical
vertebrae, while expression of Hoxc6 and Hoxc8 is confined to the trunk region;
furthermore, the boundary between Hoxc5 and Hoxc6 expression precisely
coincides with the boundary between the cervical and the thoracic vertebrae
regardless the actual numbers of the two kinds of vertebrae. To complement the
descriptive information, coherent functional data demonstrate that the different
patterns of Hox developmental expression specify the different vertebral
compositions observed in the adult animals.
The inference from the developmental data is that shifting back or forwards the
boundaries of the Hox expression domains (Hoxc5 and Hoxc6/Hoxc8 for the
cervical and thoracic vertebrae, respectively) during evolution, may have deter-
mined the different distribution of the vertebral types we now observe in living or
fossil vertebrates. In turn, developmental molecular data also suggest that changes
in the molecular machinery regulating Hox gene function in the course of evolution,
namely chromatin-remodeling proteins, cis-regulatory sequences, activity of
transposable elements and microRNA sequences, could have caused the observed
shifts in Hox domains of expression along the A-P axis [3238].
The python represents an extreme example of the consequences of shifting the
Hox genes expression domains along the A-P axis (Fig. 8b). In the python embryo,
expansion of Hoxc6 and Hoxc8 domains along the A-P axis correlates with expan-
sion of vertebral thoracic identity [36]. While expression of HOXC6 and HOXC8
proteins is confined to a defined trunk region in the chick embryo, it is expanded
from the head to the tail in the python embryo. Thus, the loss of the snakes neck
and the expansion of its rib-bearing vertebrae correlate with the shift in expression
of HOXC6 and HOXC8. From these and other studies the present model proposes
that snakes progressively lost their axial regionalization, and concomitantly their
limbs during evolution, due to an expansion of the domains of expression of the Hox
genes specifying vertebral thoracic identity [36, 37, 39]. Recent analyses of several
Hox genes expression in snake and caecilian embryos (caecilian amphibians
convergently evolved a deregionalized body plan) suggest that the evolution of a
snake-like body plan involved not only changes in Hox gene cis-regulation, but also
a different downstream interpretation of the Hox code [40]. In conclusion, the snake
case shows that a change in the place of expression of regulatory genes
heterotopy, see below can generate evolutionary novelties.
5 Four Genetic Mechanisms of Tinkering
Wallace Arthur [1] has proposed that changes affecting developmental regulatory
genes in a developmental reprogramming during evolution may be of four types
and has defined their respective evolutionary significance. These changes need not
to be mutually exclusive but more than one change may occur at a time or at
subsequent times during succeeding developmental steps. They appear to accom-
plish the Jacobs idea of tinkering.
Heterotopy, or changes in place: a developmental event is evolutionarily shifted
from one part of the organism to another. Changes in the regional expression of a
gene during development may produce evolutionary novelties. Heterotopy is
thought to have been at work to shape the snakes body.
A novel place of gene expression is also involved in the formation of the webbed
feet in some birds, such as the duck: a significant change, allowing birds to
effectively exploit aquatic niches [41]. In the early embryo, the interdigital mem-
brane is formed in the hind limb bud of either the chick or the duck (Fig. 9a).
148 G. Barsacchi
a Newborn Apoptosis BMP Gremlin

a b c d
Chick
hindlimb
a b c d
Duck
hindlimb
b
untreated gremlin
Fig. 9 A novel expression of gremlin preserves the interdigital membrane in the hindlimb of the
duck. (a) b, b Apoptosis destroys the interdigital membrane of the developing hindlimb in the
chick (b) but not in the duck (b). In (b), areas of interdigital cell death are marked by vital staining
with neutral red (arrows). c, c BMP4 is expressed (blue color) in the interdigital membrane of
the developing feet in both chick and duck. d, d Expression of gremlin in the interdigital
membrane of the duck (d, arrows) but not of the chick (d), contrasts the BMP4 signaling in the
duck hindlimb interdigital membrane. (b, b, d, d: from [41], courtesy of Dr. J.M. Hurle and
adapted with permission from Development; c,c: from [42], adapted with permission from A.A.A.S.;
the setting is a kind gift of Prof. S.F. Gilbert). (b) Scanning micrographs showing the result of an
experimental manipulation of chick developing hindlimb untreated (saline bead) or treated with
gremlin-containing beads in interdigital space. Interdigital cell death is inhibited following
gremlin treatment, producing a duck-like syndactyly in a chick limb (From [41], adapted with
permission from Development)
However, later in development the interdigital membrane is eliminated by apopto-

sis (or programmed cell death) in the chick (b), but not in the duck (b). Apoptosis is
brought about by the action of secreted proteins of the BMP family, that are
expressed in both chicks and ducks interdigital membrane (c,c); however, in
the duck BMP proteins are antagonized by expression of another secreted protein,
called gremlin (d), thus preventing apoptosis to occur. At variance, gremlin is not
expressed in the interdigital membrane of the chicks foot (d), where therefore
apoptosis can take place, separating the digits one another. Thus, the novel devel-
opmental expression of gremlin in the interdigital membrane of the duck hind limb
bud allows formation of the bird webbed feet and this interpretation is supported by
functional experiments (Fig. 9b). In fact, the forced expression of gremlin in the
interdigital space of the developing chick hind limb generates a webbed foot,
mimicking the foot of a duck!
The inference from this case study is that a spatial change in gene expression
during development namely, the novel expression of gremlin in the interdigital
membrane of the hind limb bud may have contributed to the evolution of the
webbed feet in birds.
A similar developmental mechanism operates to produce the chiropatagium that
beautifies the bats forelimbs, rendering bats the only mammals endowed with
powered flight [43]. Modifications of the bat forelimbs include elongation of the
forearm and digits, reduction in bone thickness, and the presence of tissue between
the digits. Maintenance of the interdigital membrane in the forelimb is due to
expression of gremlin in the interdigital membrane of the bats embryo forelimb,
but not in its hind limb nor in the mouse limbs [44] (Fig. 10). By inhibiting the BMP
pathway, gremlin contributes to protect the forelimb interdigital webbing from
apoptosis. Conversely, the BMP signaling plays a role in supporting the dispropor-
tionate elongation of those digits (IIIV) that sustain the chiropatagium (see below,
Heterometry).
Recent studies implicate the Prx1 homeobox transcription factor in supporting
the elongation of the bats forearm bones [45]. Interestingly, a cis-regulatory
sequence of Prx1 a highly conserved enhancer accounts for at least part of the
elongation of the long bones in the wings of bats. The bats enhancer up-regulates
Prx1 transcription more distally in the cartilage and pericondrium of the forelimb,
compared to the activity of the homologous enhancer in mouse. Strikingly, the bats
enhancer is able to impart a bats pattern of Prx1 trancription to transgenic mice,
resulting in elongation of the mouse forearm. This exciting result shows that
differences in Prx1 cis-regulatory sequences in the bat compared with the mouse
affect bone length. While functional tests of changes in regulatory sequences have
largely been restricted to insects, this study has brought this level of Evo-Devo
analysis to vertebrate systems [46].
In the same Chap. 6 of On the origin of species [2] entitled Difficulties on
Theory where Charles Darwin discusses the difficulties posed by the eye (see
paragraph 3), he also faces the case of evolution of bats in a paragraph entitled On
the origin and transitions of organic beings with peculiar habits and structure. He
asks: . . .is it possible that an animal having, for instance, the structure and habits of
150 G. Barsacchi
a b
FGF8
Gremlin
BMP BMP2
Fig. 10 Differential expression of diffusible factors (indicated on the bats wing) is involved in
producing the differential forelimb morphology in mice and bats. (a) An adult mouse,
Mus musculus. (b) An adult bat, Carollia perspicillata. Digits are numbered from anterior (I) to
posterior (V). Bat digits are elongated compared with mouse digits (inset) and maintain webbing
between the posterior digits. Expression of gremlin in the developing interdigital membrane
prevents apoptosis by inhibiting BMP signaling in the bat forelimb. FGF8 also cooperates to
maintain the bat interdigital membrane. At variance, expression of a higher amount of BMP2
correlates with the extraordinary elongation of the bat fingers that sustain the chiropatagium
(see text) (Modified from [44]; courtesy of Dr. S.D. Weatherbee and with permission from
P.N.A.S. Copyright 2006 National Academy of Sciences, U.S.A.)
a bat, could have been formed by the modification of some animal with wholly
different habits?. Indeed, in absence of transitional forms, it is difficult to imagine
how such perfected structures as the vertebrate eye or the bat wing could arise de
novo. Darwin provides indirect evidence of instances of transitional habits and
structures in closely allied species of the same genus, referring to the evolution of
flying squirrels and lemurs. The ancestors of modern bats that first appear in the
fossil record about 50 My ago already have elongated digits, extensive interdigital
membranes, and robust forelimb muscles indicative of powered flight. However,
the common ancestor of modern bats likely originated about 64 My ago, indicating
a gap of about 14 My where possible intermediate forms from quadrupedal to flying
mammals could be searched [43]. In any instance, while it is still unclear whether
modern bats arose gradually or rapidly from their quadruped ancestor, Evo-Devo
began to elucidate some of the molecular changes required to modify the morphol-
ogy of a limb into that of a wing, providing a potential explanation as to how bats
were able to achieve powered flight and colonize the air. The general picture
emerging from these studies is that small changes in the spatial expression of key
developmental genes may have driven evolution of morphological innovations such
as the bat wing.
Heterochrony: change in timing. The rates of development of part(s) of an
organism are evolutionarily shifted relative to each other. A change in the timing of
gene expression during development may also underlie the appearance of novel
structures in evolution.
a b AER
c d
AER
Fig. 11 In the dolphin embryo, the genes for some digit production are active far longer than in
other mammals causing more skeletal nodules to form in the fingers, as it is shown in the cetacean
example in (a). (b) Schematic reconstruction of the distribution of thickened apical ectodermal
ridge (AER, in red) at developmental stages 5, 6, and late 6, respectively, in the dolphin Stenella
attenuata. (c) Digit II phalanges (braces) and distal condensed mesenchyme (arrow) at stage 6. (d)
Boxed area from (c). Asterisk marks thickened apical ectoderm ridge in cross-section; this ridge is
also present in adjacent sections spanning digit II (From [49]; courtesy of Prof. M.K. Richardson
and with permission from John Wiley and Sons)
Perhaps the most dramatic evolutionary reorganization of the autopod has

occurred in cetaceans (whales, dolphins, and porpoises), whose common ancestor
diverged from an extinct group of deer-like artiodactyls approximately 50 My ago
[47]. Changes in limb morphology associated with the return to water include
considerable variation in the number of phalanges, with digits IIIV characterized
by as many as 13 in digit II of some species (Fig. 11a). Hyperphalangy appears to
have evolved three times among extant cetaceans, with two reversals. According to
an extensive analysis among terrestrial and secondarily aquatic extant and fossil
taxa (cetaceans, ichthyosaurs, plesiosaurs and mosasaurs), extreme hyperphalangy,
defined as exceeding a threshold condition of 4/6/6/6/6, is only found among
secondarily aquatic vertebrates with a flipper limb morphology and occurs exclu-
sively in digits II and III among extant cetaceans [48].
Heterochrony appears to support the disproportionate elongation of digits II and
III in cetaceans: the prolonged maintenance of a small section of apical ectodermal
ridge (AER) distal to digits II and III drives elongation of these digits and extra joint
patterning after the basic (terrestrial) digital pattern has been established and the
other areas of the AER regressed (Fig. 11bd) [49]. Developmental prerequisites
for hyperphalangy also include lack of cell death in interdigital mesoderm, produc-
ing a flipper limb. A dynamic molecular pathway has been suggested, in which joint
specification, interdigital cell death and digit elongation are mediated by
interactions between BMPs, BMP antagonists and FGF signaling from the AER.
152 G. Barsacchi
Slight modifications of the pathway may explain how hyperphalangy develops.

Thus, within cetaceans preservation of the interdigital cells and prolonged AER
signaling maintain viable the digit specification pathway to provide the patterning
mechanisms for formation of a limb apt to swimming.
Heterotypy: change in type. Molecular changes in the very same developmental
regulatory genes or gene networks can provide the proteins with new properties and
may generate macroevolutionary changes in the morphology of organisms. One of
such changes may explain why Insects have six legs only [50, 51].
About 400 My ago the Insect clade, provided with six legs only, split from the
crustacean-like Arthropod ancestors, supplied with numerous legs. At present, in
Drosophila melanogaster (and in other insects as well) the homeotic protein
Ultrabithorax (Ubx) is expressed mainly in the abdominal segments, which are
devoid of legs. In Drosophila, the protein Ubx possesses an alanin-rich aminoacid
domain working as a repressor of transcription. This Ubx domain mediates repres-
sion of transcription of the gene distal-less in the Drosophila abdomen; since distal-
less is a master gene for formation of arthropod appendages, no legs develop from
the flys abdomen. The alanin-rich domain is highly conserved among Ubx
orthologues in Insects, but is absent from Ubx in other Arthropods and
Onychophorans. In brachiopod Crustaceans endowed with many legs Ubx is
expressed in both thorax and abdomen, where it does not inhibit the distal-less
gene due to the lack, in its sequence, of the poly-alanin rich repressor domain.
Therefore the unrepressed distal-less gene can activate formation of the many legs
present along the crustacean trunk.
Thus, the mutated Ubx gene, by acquiring an aminoacid domain able to repress
the leg-forming distal-less gene in the Insect clade, may have mediated the evolu-
tionary transition to the six-legs pattern of Exapods. The evolution of this domain
may also have facilitated the great morphological diversification of posterior
thoracic and anterior abdominal segments characteristic of modern Insects. This
exemplifies heterotypy as a tinkering mechanism able to diversify animal morphol-
ogy in evolution.
Heterometry: an evolutionary change in the amount of some developmental
entity. A change in the amount of a developmental gene product can also generate
major morphological transitions.
Heterometry appears to be involved in the extraordinary elongation of the digits
that support the bat chiropatagium [52]. As mentioned above (Heterotopy), the
earliest fossil bats resemble their modern counterparts and the lengths of the III-V
digits (the primary supportive elements of the wing) have remained constant
relative to body size over the last 50 million years. Investigating embryonic
development, it was found that the digits in bats are initially similar in size to
those of mice but that, subsequently, they greatly lengthen due to relatively high
rates of chondrocyte proliferation and differentiation. A dramatic change in the
intensity of Bmp2 expression and BMP signaling in the bat embryonic forelimb,
relative to the bat hind limb and mouse forelimb stimulates cartilage proliferation
and differentiation and increases digit length in the bat forelimb (see Fig. 10b). This
species- and limb-specific change indicates that BMP2 has a major role in the
developmental elongation of bat wing digits. In addition, an up-regulation of the

BMP pathway may potentially represent a key mechanism in the evolutionary
elongation of bat forelimb digits.
Heterometry has been beautifully demonstrated in the most celebrated example
of adaptive evolution, the radiation of Darwins finches. Darwins finches are a
group of about 14 closely related species on the Galapagos Islands and Cocos Island
collected by Charles Darwin and others during the Beagle expedition in 1835.
In 1839 Darwin wrote [53]:
The most curious fact is the perfect gradation in the size of the beaks in the different
species of Geospiza, from one as large as that of a hawfinch to that of a chaffinch, and (. . .)
even to that of a warbler. (. . .) Seeing this gradation and diversity of structure in one small,
intimately related group of birds, one might really fancy that from an original paucity of
birds in this archipelago, one species had been taken and modified for different ends.
The finches have beaks of different sizes and shapes specialized for specific
nutritional tasks: some species (ground finches) have stout beaks apt to cracking
seeds; others (cactus finches) have slender, pointed bills apt for reaching into cactus
flowers (Fig. 12a, b). Even small differences in any of the three major dimensions
(depth, width and length) of the beak have major consequences for the overall
fitness of the birds [54]. The specialized beak shapes are apparent at hatching and
thus are genetically determined. A comparison of beak development in six species
of Darwins finches belonging to the genus Geospiza revealed a striking correlation
between beak morphology and level and timing of Bmp4 expression [55, 56]. In
particular, a higher amount of Bmp4 is expressed in the primordium of the wider
and deeper beak, while a lower expression of Bmp4 corresponds to a longer, thinner
beak (Fig. 12b). In addition, artificially increasing BMP4 levels in the beak mesen-
chyme of a chick beak embryo, is sufficient to alter beak morphology in the same
direction as is seen in the larger beaks of ground finches.
Conversely, the calmodulin (CaM) protein a molecule involved in mediating
Ca2 signaling is expressed at higher levels in the developing long and pointed
beaks of cactus finches than in more robust beak types of other species [57]. When
up-regulation of the CaM-dependent pathway is artificially replicated in the chick
frontonasal prominence, it causes an elongation of the upper beak, imitating the
beak morphology of the cactus finches (Fig. 12c, d). Thus, the CaM-dependent
pathway is likely to have been a component of the evolution of Darwins finch
species with elongated beaks.
To sum up, observations and experimental results may provide an explanation
on how tinkering with BMP4 and calmodulin regulation in the beak primordia
provided the morphological variation acted on by natural selection in the evolution
of the beaks of the Darwins finches. Furthermore, the fact that BMP4 and calmod-
ulin act through two different signal transduction pathways may have contributed to
the flexibility of beak conformation in the different birds during evolution. The
exciting discoveries being made from studies on Darwins finches are a true
celebration of Darwins legacy to modern science.
154 G. Barsacchi
a b Bmp4 expression
c d
Fig. 12 Heterometry (and heterochrony, see text) in development of the beaks of Darwins finches.
(a, b) Differential expression in the timing and amount of key molecular regulators of the beak
formation during development appears to account for the different beak shapes of Geospiza species.
(b) The represented species form two groups: ground (top) and cactus (bottom) finches, with distinct
beak morphologies. At stage 29, Bmp4 (red arrows) is expressed at high levels in the distal beak
mesenchyme of G. magnirostris, the species with the larger and deeper beak. Broad domains of
Bmp4 expression are detectable around prenasal cartilages of G. fuliginosa and G. fortis. A small
domain of strong Bmp4 expression is also found in the most distal mesenchyme of G. conirostris,
and weaker expression is seen in G. scandens and G. fortis. (c, d) Functional analysis of CaM-
dependent pathway in beak development. Whole head views of day 10 chick embryos. (c) Wild-type.
In (d), the CaM-dependent pathway was up-regulated and the beak was lengthened. The length of the
beak is shown with a red line; the depth of the beak at the base and the depth of the beak at the tip are
shown with blue and green lines, respectively. ((a) Kind gift of Prof. A. Abzhanov (b) adapted from
[55]; courtesy of Prof. A. Abzhanov and with permission from A.A.A.S. (c, d) from [57]; courtesy
of Prof. A. Abzhanov and with permission from John Wiley and Sons)
6 Conclusions
Evolutionary Developmental Biology, or Evo-Devo, explores the relationships

between the processes of individual development and the phenotypic changes of the
organism during evolution. Evo-Devo is acknowledged to be revolutionizing our
understanding of how the development of organisms has evolved and of how

development contributes to evolution. Methodological advances such as gene
cloning, gene expression screening and visualization of gene activity in embryonic
tissues facilitated the emergence of a major theme of the current Evo-Devo
research, the evolutionary developmental genetics programme [58]. Its founda-
tional achievement was the discovery of extensive similarities in developmental
regulatory genes and gene networks among distantly related species. The
programme concentrates on the evolution of genetic tool-kits and signaling
pathways and on the regulatory logic that underlies organism development.
Mapping the expression pattern of gene networks and signaling pathways and
analyzing their correlation with the constructional features of body architecture,
provides information on their possible role in phenotypic evolution.
Major morphological transitions in evolution are presently recognized to be
accommodated by a few key developmental genetic changes (part of a develop-
mental reprogramming [1]) and case studies in snakes, ducks, bats, dolphins,
insects, and finches, providing valuable insights into principles of evolutionary
change, are presented. On the other hand, the molecular changes are rooted in an
otherwise conserved developmental genetics tool-kit (e.g., the Hox genes for ante-
rior-posterior patterning, the network for eye formation) that substantiates the deep
homology underlying diversity of forms [12]. On this ground, the relationship of
the deep homology of genes working through development with classic morpholog-
ical homology is in the Evo-Devo field of exploration. Also, how environmental
agents can instruct changes in development, for example altering gene expression
in a broad sense, searching for a link between genetic and environmental influences
on development and the emergence of selectable phenotype variation is in the
perspectives of the newly growing and exciting discipline Ecological Developmen-
tal Biology (Eco-Devo), where Evo-Devo integrates with Ecology by examining the
mechanisms of developmental regulation in an ecological context [59, 60].
We can expect that future work may further give reason for the Charles Darwins
appraisal of the importance of Embryology for Evolution.
Acknowledgements I regret that space constraints make it impossible to cite all relevant work
and I therefore apologize to those whose work could not be cited. Additional references may be
found in the cited papers. I am grateful to all Authors who were the source for this work and in
particular to Prof. S.F. Gilbert, whose seminars and writings raised my interest in Evo-Devo. I wish
to thank Prof. S.F. Gilbert and Prof. A. Abzhanov for their kind and generous gift of some pictures.
I am also grateful to the Academies that organized the meeting on The Theory of evolution and its
impact in Turin, May 2729, 2010, for giving me the opportunity to present some of the present
work in the genetics program of Evo-Devo.
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Evolutionary Mechanisms and Neural
Adaptation: Selective Versus Constructive
Strategies in the Development and Plasticity
of the Nervous System
Ferdinando Rossi
Abstract The correct function of the nervous system requires complex neural
networks bearing precise connections. In principle, the high structural specificity
of neural circuits could be achieved by genetically-determined processes, selected
and refined during evolution. Highly conserved gene networks regulate some
crucial steps of neural development, such as the regionalization of the neural tube
and the initial phases of neurogenesis and synaptogenesis. A totally hardwired
nervous system may meet the requirements of adaptation and natural selection at
the population level. Nevertheless, it would be inadequate to allow individual
organisms to cope with rapid changes of environmental conditions. Neural adapta-
tion to external constraints can be partly achieved by introducing selective
mechanisms in neural development. Accordingly, neurons are generated in excess
and then partially eliminated to match the actual extension of innervation
territories. Such mechanisms, however, are restricted to a set of potentialities,
which must be predetermined in the ontogenetic program. On the other hand,
constructive mechanisms, in which external stimuli directly influence structural
modifications of neural circuits to produce adaptive responses, may allow individ-
ual organisms to cope with a wide variety of unprecedented situations. Thus, in the
last ontogenetic period as well as in the adult, when the organism actively interacts
with the external milieu, experience exerts a strong growth-promoting effect on
neural circuits and connections inducing the emergence of specific functional
properties. By this mechanism, which requires strict inhibitory control to prevent
aberrant growth and dysfunction, the nervous system exploits external stimuli to
create adaptive responses to unexpected situations.
F. Rossi (*)
Department of Neuroscience, Section of Physiology, University of Turin, Orbassano, Torino, Italy
e-mail: ferdinando.rossi@unito.it

160 F. Rossi
1 Introduction
Over the last decades, substantial advancements have been obtained in the elucida-
tion of the cellular and molecular interactions that regulate the development of the
nervous system, govern its function and determine its plastic capabilities in adult-
hood. These discoveries have led to the proposition of concepts and principles that
relate, in a very peculiar manner, developmental neurobiology and neurophysiol-
ogy to evolution. In addition to the obvious influence exerted by evolutionary
processes on neural ontogenesis and on neurobiological mechanisms [57], this
novel relationship stems from the understanding that both the construction of the
nervous system and its operation are continuously scrutinized for their efficacy in
enabling the organism to cope with environmental demands. Hence, the notion that
neural development and plasticity represent the biological substrates of adaptation
has led to propose that these processes are regulated by fundamental mechanisms
that are shared with Darwinian evolution and, notably, the mechanisms of natural
selection [8, 13].
This concept originated from the discovery that some fundamental ontogenetic
phenomena, such as the formation of appropriate numbers of neurons or synapses in
the brain, are subjected to environmental constraints, in a way that is reminiscent of
the regulation of population size in living organisms. For instance, there is now
general agreement that most neuron populations are initially generated in excess
and attain their final numbers by a process of cell elimination, in which death or
survival depend on the extension of innervation territories, the availability of target-
derived trophic substances or the level of neuronal activity [27, 47]. Similar
considerations are usually applied to synaptogenesis, where initially exuberant
contacts are progressively withdrawn according to a set of restrictive parameters,
including levels of activity, spatio-temporal patterns of synaptic activation or
activity-dependent uptake of neurotrophic factors [27, 62].
This large body of evidence highlights the role of selective mechanisms in
aspects of neural development and plasticity that are strictly related to adaptation.
Nevertheless, a purely selective mechanism implies a range of pre-existing
potentialities, which is restricted following confrontation with intervening
demands. In other words, all the available options should be hardwired ex ante in
the ontogenetic program responsible for constructing the organism. Now, is such a
mechanism really compatible with adaptation? How can the variety of pre-existing
potentialities be expanded at an adequate pace to match the speed of environmental
change? Are the discarded options permanently lost or can they be rescued if they
become again advantageous in the future?
A selective strategy is primarily designed to control adaptation at the population
level. Hence, it is most efficient in regulating species evolution or, as we will
discuss later, in defining the number of neurons belonging to a certain category. On
the other hand, the main goal of neural adaptation is to allow individual organisms
to cope with changing environmental conditions. A closer examination of neural
development and plasticity in this perspective actually suggests that the nervous
Evolutionary Mechanisms and Neural Adaptation 161
system must be endowed with an intrinsic capability to construct neural circuits so

to create novel functional properties, beyond the original set of potentialities. As a
consequence, both selective and constructive mechanisms participate to determine
neural ontogenesis and plasticity. Constructive strategies, however, prevail over
selective ones when the individual nervous system has to face contextual environ-
mental demands.
2 Adaptive Mechanisms Can Be Either Predictive or Reactive
Biological modifications set up to cope with environmental changes occur according

to two main modes. On one side, the organism is able to predict the incoming
variation and builds up an anticipated response. On the other, the organism is unable
to foresee the external change and it can only react to novel conditions once
they have been established. Thus, predictive adaptation implies that the organism
is ready to face the novel environmental demand at the time when it materializes,
whereas reactive adaptation will be only unfolded in a subsequent time.
At a first glance, predictive adaptation may appear more efficient in favouring
survival of the organism. Nonetheless, it can be only used in a restricted set of
situations. Actually, predictive mechanisms are only suitable to face extrinsic
changes that happen at a constant pace through a long period of time (essentially
forever). Organisms that spontaneously acquire predictive abilities are favoured
over their counterparts and, hence, these abilities become selected by evolutionary
mechanisms. Accordingly, predictive adaptation is usually sustained by highly
conserved gene networks, whose spatio-temporal patterns of activation correspond
to the time course or space distribution of the related environmental conditions. The
best example of this kind is the regulation of circadian and circannual functions [12,
19]. These functions are operated by molecular cascades endowed with intrinsic
rhythms that match the duration of relevant environmental periods, to which they
become entrained by sensory information. As we will discuss here, predictive
mechanisms operate in some major ontogenetic processes, which are also governed
by highly conserved gene programs. For instance, the gene networks that direct the
building of the body (and neural) plan have clearly evolved to cope with consistent
environmental constraints, such as gravity, the sources of energy or relevant
sensory information (e.g. sunlight) or the mechanics of movement.
Albeit successful, predictive strategies take very long times to become
established and diffused. In addition, it is clear that the vast majority of environ-
mental changes happen according to completely unpredictable frequencies and
locations. Such situations can be adequately faced only by means of reactive
processes, which allow individual organisms or populations to design and set up
novel responses. In these cases, evolutionary processes favour the emergence and
maintenance of certain abilities, but leave ample degrees of freedom in their actual
expression. Most homeostatic mechanisms work in this way. For instance, body
temperature is maintained by a series of evolutionary-selected interdependent
162 F. Rossi
devices, from thyroid hormones to horripilation, whose function is triggered and

modulated by feedback loops that tune every response to the concomitant situation.
The vast majority of external conditions that may influence the function of the
nervous system belong to the latter category. More, I would say that the main
emerging property of the nervous system is to design novel strategies to solve
unprecedented problems. Accordingly, neural cells and circuits must be endowed
with the ability of reshaping connectivity so to generate new functional capabilities
that are not part of the constitutive repertoire of the species. Acquiring new
information or learning new skills are examples of this sort of morpho-functional
modification that underlies neural adaptation. Hereafter, I will argue that these
processes, that are crucial to regulate neural development and plasticity, cannot
be solely explained in terms of selective mechanisms, but require constructive
properties that allow the creative design of new adaptive strategies.
3 Neural Development and Evolutionary Mechanisms
In the perspective of this essay, neural development can be schematically

subdivided in three main phases (Fig. 1): (1) neurulation refers to the formation
of the neural tube and its segmentation into discrete morphogenic regions; (2)
neurogenesis is the process by which neurons (and glia) are generated; (3)
synaptogenesis is the process by which neurons become connected to each other
into functional circuits. These phases comprise both addition (e.g. generation of
new neurons, formation of new synapses) and loss of elements (e.g. physiological
cell death, synaptic pruning). Therefore, the growth of the nervous system actually
results from the balance of concurrent expansive and regressive phenomena.
Neurulation is triggered by inductive signals issued by the notochord,
a mesodermal structure lining the rostro-caudal axis of the embryo, which triggers
profound morphogenic rearrangement of the overlying ectoderm leading to the
formation of the neural tube [3, 27]. The latter is a highly polarised structure, which
soon becomes subdivided in discrete domains that acquire distinctive morpho-
functional specification along the main spatial axes (Fig. 1) [3, 27]. The most
important partition occurs along the rostro-caudal axis, where morphologically
distinct segments appear, corresponding to the major subdivisions of the adult
Central Nervous System (CNS). Within each of such segments, the dorso-ventral
axis defines sensory or motor structures, whereas the medio-lateral axis defines the
relationship linking neural circuits to axial structures (the trunk) and distal
appendages (the limbs).
The regionalization and spatial specification of the neuraxis are determined by
the interplay between diffusible or contact signalling cues and the combinatorial
expression of specific sets of transcription factors [3]. The whole process is
regulated by gene networks, which direct the morphogenesis of the entire body
plan. This gene program has been particularly successful during evolution: it has
been inherited from invertebrates and it is highly conserved through the phyla of
Fig. 1 Regulatory mechanisms of neural ontogenesis. The early phases of nervous system
development are determined by the execution of a gene program that directs neurulation, the
regionalization of the neural tube, the generation of nerve cells and the initial formation of
synapses. While these processes are regulated in a predictive manner, later phases are accom-
plished according to reactive strategies, required to adapt ontogenetic processes to contextual
environmental conditions. Surplus neurons are eliminated before birth by a selective mechanism
depending on the extension of available innervation territories. On the other hand, synaptogenesis
is carried out after birth, when the organism is interacting with the external world. Hence, synapse
formation and reshaping are governed by experience-dependent constructive mechanisms
vertebrates [51]. The program assembles a structural scaffold, in which fundamental

morphogenic interactions are precisely regulated in space and time, securing the
coordinate development of intrinsic neural networks and their appropriate integra-
tion within the nascent organism. On this basic canvas, evolution creates diversity
by introducing domain-specific variations in the rate of growth and in the connec-
tion patterns. In this way, birds have a relatively large mesencephalon, whereas
mammals are characterized by a prominent telencephalon. Thus, neural morpho-
genesis is accomplished, in a predictive manner, by the intrinsic activity of specific
gene networks, whose success is determined a posteriori by natural selection.
164 F. Rossi
Neurogenesis, which is obviously interrelated with morphogenesis, comprises

all the phenomena leading to the generation of neurons and glia from neural stem
and progenitor cells (Fig. 1) [27]. These cells proliferate in germinal structures
located at different levels along the neuraxis, become specified towards different
identities and migrate to specific locations, where they acquire mature phenotypes.
Then, the final size of each neuronal population can be refined through physiologi-
cal cell death. The generation of phenotypic diversity is largely determined by
diffusible molecular cues or cell-to-cell interactions that regulate the expression of
particular combinations of transcription factors [14, 22, 37, 39]. Once cell fate
choices have been taken, however, the differentiation into mature phenotypes is
achieved by the unfolding of type-specific gene programs, in an essentially cell-
autonomous manner. Hence, neuronal differentiation as well as the establishment of
the basic framework of connectivity are also governed by predictive mechanisms
that determine a priori the capability of a given neuron to migrate into a certain
position, orientate the navigation of its axon or recognize appropriate targets.
The situation is different when the regulation of neuron numbers is considered
(Fig. 1). The number of neurons generated for each category is determined by the
interplay between intrinsic properties of neural progenitors and local regulatory
interactions that modulate the rhythm of proliferation, the relative proportion of
cells that initiate differentiation or continue to divide, and the duration of neuro-
genic periods [7, 33]. All these mechanisms operate to regulate neuron numbers by
adjusting their production and, hence, work according to a predictive strategy.
Nevertheless, since the pioneering work of Rita Levi-Montalcini and Giuseppe
Levi [30], it is well known that most neuron populations are actually generated in
excess and the final amount of nerve cells that populate the mature nervous system
is achieved through the elimination of supernumerary elements [42]. Cell death or
survival depend on a set of parameters, including both intrinsic features of the
neurons (e.g. their level of activity) and environmental constraints (e.g. the exten-
sion of the target field or the availability of neurotrophic substances). This process
is suitable to match the size of each neuronal population to the amount of potential
synaptic partners or to the extension of innervation territories in the periphery. It
operates according to a selective mechanism that is most reminiscent of natural
selection: the juvenile neurons compete for limited quantities of available resources
and their fate depends on their intrinsic ability to overcome their rivals [8, 47]. In
this case, however, the mechanism works following a reactive strategy, required to
adjust neural development to individual fluctuations in the dimension of different
parts of the body. Accordingly, the size of most neuron populations can be signifi-
cantly modified by experimental manipulations that increase or reduce the exten-
sion of the available innervation territory [27, 42, 44]. Therefore, the final number
of neurons belonging to each population derives from a dual mechanism, which
combines a predictive component, that determines the initial production of surplus
neurons, and a reactive component, that eliminates supernumerary elements in
response to contextual environmental conditions.
At a first glance, similar mechanisms may apply during synaptogenesis (Fig. 1).
A well-established notion in developmental neurobiology is that synapses are
initially formed in excess and then partially withdrawn to shape the mature connec-
tivity [27, 47]. Since effective function of neural circuits depends both on the
number and on the specificity of synapses, the pruning process would be required
both to reduce the exuberant, supernumerary contacts and to remove aberrant,
wrong connections.
The initial formation of synapses is guided by recognition cues exposed on the
neuronal membrane, whose nature is determined by the intrinsic neurochemical
profile of the partner neurons [5, 61, 66]. Synaptic pruning is driven by activity-
dependent mechanisms that are directly influenced by the functional efficacy of the
developing circuitry [8, 47]. Thus, synaptogenesis also appears to depend on a dual
mechanism. Synapse formation is guided by molecular interactions determined by
the unfolding of neuronal-intrinsic gene programs that work in a predictive manner.
On the other hand, synaptic pruning is driven by an essentially reactive mechanism
that selects good connections on the basis of their functional meaningfulness. Again,
the latter phenomenon appears to follow some fundamental principles of natural
selection.
The analogy is partial at best. It is well established that a number of synapses are
withdrawn to shape appropriate spatial connection patterns on specific target
domains. Nonetheless, it is definitely clear that, when the number of contacts and/
or their functional weight is considered, the final balance of the synaptogenic
process is a positive one: newly-formed synapses greatly outnumber the lost ones
[46, 49, 62]. This has been clearly demonstrated in a variety of experimental
models, including the autonomic nervous system [31, 48], the visual system [60],
or the cerebellar climbing fibres [23], just to cite a few ones. Even in the case of the
neuromuscular junction where mono-innervation of muscle fibres appears to be
solely achieved through the elimination of supernumerary axons, the winner
endplate undergoes a remarkable outgrowth to cover the entire postsynaptic surface
with additional junctional complexes and releasing sites [46, 55]. Therefore, the
reactive component of synaptogenesis is not a selective process, but rather operates
in a constructive manner.
This conclusion has profound implications in terms of structure-to-function
relationship during neural development. Indeed, while the initial formation of
synaptic contacts is essentially aimed at establishing a basic framework of neural
networks capable of initiating the interaction with the external world, the refine-
ment phase is aimed at modifying the structure of such networks to improve their
operational abilities. Thus, a fundamental circuit scaffold, assembled by executing
intrinsic gene programs, is confronted with experience and modified to achieve
adaptive function. The latter process involves the elimination of some unspecific
contacts, but it is primarily characterized by the strengthening of meaningful
connections with the addition of numerous new synapses.
This process of structural remodelling, which involves the simultaneous
outgrowth of both presynaptic axons and postsynaptic dendrites [44, 48], leads to
the emergence of novel functional properties, whose nature is influenced by the
specific features of the contextual environmental conditions. In other words, the
final structure of neural circuits is congruent with the actual experience: a particular
166 F. Rossi
interaction with the external world will always lead to an appropriate pattern of
connectivity [53]. The essentially constructive nature of this process can be best
appreciated in extreme experimental conditions. For instance, severe manipulations
such as monocular deprivation or experimental squid during the critical periods of
visual system development induce extensive changes in the connectivity of the
subcortical and cortical visual pathways [60]. This peculiar structure, albeit
strongly divergent from that of the normal population, is clearly adaptive when
the visual experience of the relevant individuals is considered. Indeed, there is no
reason to leave half of the cortical territory to an eye that is not conveying any
significant sensory information. Similarly, there is no use to form binocular
connections if the two eyes are seeing different scenes. Yet, it is difficult to believe
that such unusual projection patterns result from the selection of pre-existing
connections, rather than being actively constructed by adapting the morpho-func-
tional properties of the circuit to real life experience. Similar considerations apply
to other systems, such as the peculiar tonotopic representation that can be induced
in the auditory cortex by exposure to auditory stimuli of specific frequencies [10].
On the whole, the initial phases of nervous system development, which include
neural morphogenesis, neuronal production and the establishment of basic connec-
tion patterns, are directed by the activity of species-specific gene networks that
operate according to an essentially predictive strategy. These processes lead to
assemble the fundamental framework of the nervous system, which then undergoes
individual-specific morpho-functional adaptation according to reactive strategies.
Neuron numbers are refined through a primarily selective process, whereas synaptic
patterns are reshaped according to constructive mechanisms. The latter mechanisms
have been likely evolved to exploit influences derived from contextual experience
to favour the development of adaptive function.
4 Experience-Dependent Mechanisms, Neural Development

and the Emergence of Function
A major feature of the last phases of neural development is the appearance of

reactive processes that essentially shift adaptation from species to individuals. Such
processes, however, are accomplished during distinct ontogenetic phases,
characterized by strongly different conditions [27, 46]. Neurogenesis and physio-
logical cell death primarily occur before birth and are influenced by somatic
changes taking place within the same developing organism. On the other hand,
the bulk of synaptogenesis is carried out after birth, while the newborn organism is
actively interacting with the external world. The latter condition exerts a most
dramatic influence on the course and on the outcome of this process.
Higher vertebrates, notably mammals, are born with immature neural circuits,
and this feature is most prominent in primates and humans [45, 57]. This implies
that crucial phases of neural development occur while the organism is exposed to
Fig. 2 External stimuli direct developmental synaptogenesis and adult circuit plasticity. External
stimuli drive plastic modification of neural circuits by inducing neuritic remodelling and directing
the formation of functionally meaningful contacts. The process is regulated by inhibitory cues
present in the CNS microenvironment (represented by the STOP signals), required to prevent
aberrant growth and dysfunction
the external environment rather than sheltered in an egg or in the uterus. The
newborn CNS, and particularly those structures that are more immature at birth
such as the neocortex, is subjected to a wide range of powerful stimuli, which
induce specific patterns of neuronal activation, stimulate neuritic extension and
influence the number and the distribution of newly-formed synapses [63]. This
ability of experience to stimulate neural growth is the crucial event that shifts the
nature of synaptogenesis from a selective process aimed at achieving synaptic
specificity to a constructive one capable of building new functionally meaningful
connections (Fig. 2).
Sensory deprivation experiments, such as dark rearing or exposure to un-
modulated acoustic stimulation [11, 60, 65], show how administration of meaningful
stimuli immediately activates neuronal growth mechanisms, associated with rapid
acquisition of new functional properties. All these examples of experience-dependent
structural remodelling are characterized by a clear prevalence of expansive phenom-
ena, with the formation and strengthening of new synapses, over regressive events and
loss of contacts. Hence, experience drives neuronal growth to create adaptive function.
The evolutionary advantage of this strategy is obvious: each individual organism
capable of exploiting contextual experience to generate appropriate novel responses
will be able to successfully cope with a wide range of unprecedented situations.
Once function is acquired, synaptogenic processes are greatly reduced if not
completely arrested [24]. This decline of neuronal growth properties, that marks the
end of developmental critical periods for the acquisition of experience-dependent
capabilities, has been attributed to a set of concurrent mechanisms. The remodelling
of neural circuits often leads to a substantial segregation of afferent axons, which
impinge upon private target domains, being individual dendrites, single neurons or
discrete anatomical modules. This process of input segregation would progressively
reduce the need and the opportunity for activity-dependent competitive interactions
that sustain synaptogenesis [62]. Hence, growth would be arrested when a stable
connection pattern is achieved and all partners had their share.
168 F. Rossi
In spite of the attractive simplicity of this mechanism, the end of synaptogenic

processes is actually coincident with profound modifications that occur in the
neurons themselves and in the surrounding microenvironment [53]. Within the
nerve cells, growth-associated gene programs are actively suppressed to favour
information processing and signalling function. Coincidentally, the maturation of
glia, namely myelination, and the deposition of the extracellular matrix are
accompanied by the appearance of a variety of growth-inhibitory molecules that
stabilize contacts and hamper further elongation of neuronal processes (Fig. 2).
These phenomena are precisely aimed at restricting growth properties of neural
circuits. As we will see in the next section, synaptogenic properties typical of
juvenile organisms can be restored in the mature CNS by specific manipulations
that boost intrinsic neuronal growth properties or remove environmental inhibition.
The presence of such strict growth control mechanisms, which have been
progressively implemented during the evolution of vertebrates [17, 56], represents
an additional argument favouring the constructive nature of developmental
synaptogenesis. Indeed, a purely selective mechanism is self-limiting and does
not require additional regulatory devices to be terminated. On the contrary, a
constructive mechanism must be actively arrested, either by removing the sustain-
ing stimuli or by dampening growth processes. Experience cannot be prevented or
abolished: the whole ontogenetic process is precisely aimed at making the nervous
system able to cope with external constraints. Therefore, when the development of
neural circuits adopted the constructive strategy driven by experience-dependent
stimulation, a set of growth-inhibitory mechanisms evolved to stabilize meaningful
connections and to restrain neuronal growth once function is achieved. Not surpris-
ingly, the induction of such regulatory molecules is also triggered by experience
[26, 59].
5 Constructive Mechanisms and Plasticity in the Adult
In spite of the clear decline of intrinsic neuronal growth potentialities, after the end of
canonical ontogenesis the nervous system retains a certain degree of ability to modify
his structure and function in response to external stimuli or changes in the environ-
ment. Adaptation in the mature nervous system, which is generally known as plastic-
ity, shares some fundamental features and mechanisms with developmental processes.
The notion of plasticity in the adult CNS was established several decades ago with
the discovery of reactive synaptogenesis and synaptic turnover [9, 50]. Accordingly,
for a long time the adaptive abilities of neural circuits were thought to be exclusively
sustained by changes of connectivity. Recently, however, the demonstration that
neurogenesis persists at least in some regions of the adult mammalian brain has
revealed that functional adaptation can be also carried out by integrating new
neurons in pre-existing circuits.
Compared to neural development, synaptogenic phenomena occurring in the
adult nervous system are considerably restricted in space and time. They involve
both formation and withdrawal of synaptic contacts and, although they usually lead
to moderate changes of synaptic numbers, they obey to reactive mechanisms and
have a clear constructive character. One important difference with juvenile
synaptogenesis is the requirement of active participation [29, 64]. Synaptic
remodelling in immature organisms is usually triggered by the mere exposure to
external stimuli. In contrast, in adulthood plastic changes also require motivation
and active participation of the involved organism. Hence, in mature individuals
adaptation is no more an automatic response to environmental conditions, but
requires an individual volition that determines the nature of the response and
influences its outcome.
Plasticity in the adult is strongly hampered by the presence of the above-
mentioned inhibitory mechanisms that terminate developmental synaptogenesis.
These mechanisms are partially counteracted by the growth promoting effect
exerted by external stimuli [18, 20, 54]. Accordingly, structural plasticity and
functional adaptation in the adult can be conspicuously enhanced by experimental
procedures that activate neuronal growth genes or neutralize inhibitory molecules
of the CNS microenvironment [53]. Nevertheless, whatever effective the simple
manipulation of the molecular devices that control neuritic growth is not sufficient
to induce adaptation. Endurable structural changes associated with significant
functional modifications can only be established if these procedures are combined
with specific environmental stimuli [43]. Hence, growth regulatory mechanisms
exert a purely permissive role by setting the degree of plasticity of neural circuits,
whereas environmental stimulation has a primarily instructive function in deter-
mining the shape of the connectivity that will be formed [53].
These features are consistent with a reactive mechanism that induces structural
remodelling of neural circuits to generate adaptive responses. As for developmental
synaptogenesis, the presence of multiple inhibitory mechanisms is required to
maintain constructive modifications within the limits of adaptive function. Indeed,
there are several examples showing that altered regulatory mechanisms and/or
unusual experience may induce unspecific growth associated with frank pathological
phenomena, such as seizures or dystonia [1, 6, 40]. A selective mechanism may fail
to generate an adaptive response if the required option is not available, but it should
be intrinsically unable to produce abnormal structures and aberrant function. Thus,
plasticity in the adult also follows a constructive strategy and, for this reason, it
must be subjected to inhibitory control.
Adult neurogenesis shares its major functional significance with adult plasticity.
In some CNS structures adaptation is not exclusively sustained by changes of
connectivity, but also involves the integration of newly generated neurons into
pre-existing circuits. As discussed above, developmental neurogenesis comprises a
predictive mechanism that generates excessive amounts of neurons, whose final
number is defined by a reactive mechanism that operates through selection. The
scenario of adult neurogenesis is very different. In both regions of mammalian brain
where new neurons are generated throughout life, the hippocampal dentate gyrus
and the olfactory system, the rate of neuronal generation is clearly influenced by
external stimuli and/or activity-dependent mechanisms [15, 35]. Thus, while the
170 F. Rossi
adult system retains the capacity for generating neurons, the course and outcome of
the process are no more determined by an intrinsically-coded predictive mecha-
nism, but regulated by extrinsic cues according to a reactive strategy.
Many of the newly generated neurons survive only for a short time, suggesting
that survival may depend on selective mechanisms, as for developmental
neurogenesis. However, the number and the specific features of the neurons that
eventually become stably integrated in adult circuits depend on the activity of the
involved network and on specific functional demands [2, 28, 32, 41]. In other
words, integration of the newborn neuron is directly related to the function that is
being established and not to the intrinsic receptive capacity of the system. There-
fore, similar to synaptic remodelling, adult neurogenesis appears to work as a
reactive device obeying to a primarily constructive strategy.
This conclusion is further supported by the observation that neurogenesis, or at
least neurogenic attempts, may be induced in other regions of the CNS by strong
stimulation or pathological conditions [4, 34, 52, 58]. In these instances, non-
neurogenic structures react to extreme environmental constraints by redirecting
the specification of local progenitors towards neuronal lineages. These phenomena
of intraparenchymal neurogenesis are often abortive, because non-neurogenic
regions fail to provide adequate conditions to support the differentiation and
integration of new neurons. Hence, latent neurogenic potentialities may be diffused
in many CNS regions, but actively repressed by local constraints. In any case, adult
neurogenesis appears to be driven by environmental stimuli influencing the mature
tissue, rather than local regulatory cues acting in a primary germinal structure.
Another feature that adult neurogenesis shares with adult plasticity is the pres-
ence of strict inhibitory control. Intrinsic inhibitory control prevents adult neurons
from de-differentiating or re-entering the cell cycle [25]. In addition, environmental
cues regulate the proliferation of progenitors as well as the migration, differentia-
tion and integration of newborn neurons [38]. Thus, successful incorporation of
new neurons in adult networks is restricted to precise phenotypes in defined circuits.
Furthermore, transplantation experiments show that the endogenous ability of the
adult CNS to accommodate donor neurons in functional circuits is limited to a few
types and locations [21, 36]. These inhibitory constraints also appear to be primarily
aimed at preventing aberrant phenomena that may lead to maladaptive function or
behaviour. However, these considerations indicate that adult neurogenesis also has
the main characters of a reactive/constructive process, in which experience-dependent
growth is exploited to modify neural structures so to achieve adaption.
6 Conclusions
The initial phases of neural development are primarily regulated by predictive

mechanisms that have been established by evolution. These processes, which are
highly conserved throughout vertebrate phylogenesis, are designed to develop a
nervous system that is suitable to control the main bodily functions of the organism
and is capable of interacting with the external world. The sensitivity of neural
circuits to external stimuli, however, profoundly influenced the strategy of neural
development. When coping with rather constant phenomena, such as the physio-
logical expansion or retraction of different body parts, suitable adaptation can be
obtained by merely selective mechanisms, which share some features with natural
selection. Hence, neurogenesis starts with the production of surplus neurons and
their final number is adjusted to match actual requirements, which may fluctuate
among individuals, but always remain within predictable ranges. A similar mecha-
nism may also apply to synaptogenesis if the nervous system was designed to be
completely hardwired by intrinsic genetically-determined mechanisms.
Quite surprisingly, however, the exposure of the immature nervous system to the
external environment dramatically changed the ontogenetic strategy. Now, the
ability of coping with a great variety of unpredictable environmental constraints
could not be adequately fulfilled by a selective process. Rather, the expanding
variety of situations favoured the emergence of an alternative mechanism, able to
create unprecedented structure and function to face unprecedented situations. Thus,
evolutionary pressure pushed developmental synaptogenesis, adult plasticity and
even adult neurogenesis to become reactive processes obeying to the rules of
constructive mechanisms. This constructive revolution of neural ontogenesis
induced the appearance of specific regulatory mechanisms, which evolved to
restrain the unchained growth driven by external stimuli within the limits of
adaptive function. These inhibitory cues first appeared in fish and amphibians
[56], but their importance consistently increased during later vertebrate evolution,
in parallel with the increasing complexity of CNS structure and function. Now, they
clearly fulfil the fundamental task of controlling potentially dangerous growth
properties that enable the nervous system of powerful plastic and adaptive
capabilities. However, they also bring with themselves some relevant side effects,
such as the loss of neural regeneration capabilities [16, 17]. In any case, construc-
tive mechanisms, such as those directing adult plasticity and neurogenesis, repre-
sent a most successful phylogenetic invention that greatly increased the individual
ability to cope with increasingly wide ranges of environmental conditions.
Acknowledgements The scientific work of Ferdinando Rossi is supported by grants from

Ministero dellUniversita e della Ricerca Scientifica e Tecnologica (MIUR-PRIN 2007 prog.
nr. 2007F7AJYJ), Compagnia di San Paolo (Neurotransplant Project 2008; GABAGEN Neurosci-
ence project 2009), Regione Piemonte (Project A14/05; Ricerca Sanitaria Finalizzata, 2008,
2009), Ataxia UK; Fondazione Cavaliere del Lavoro Mario Magnetto of Turin.
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Is the Human Brain Unique?
Gerhard Roth
Abstract The human brain is not unique in terms of general structure. It exhibits a
morphology typical of mammals and more specific of primates. Also, humans do
not have the largest brain either in absolute or in relative terms, although they
possess a brain that is seven to eight times larger than expected from general
mammalian brain-body relationship. The size of the human cerebral cortex and of
the prefrontal cortex as the seat of intelligence exhibit a slightly positive allome-
tric growth, i.e. the cortex increases faster in size than the rest of the brain, which is
again typical of mammals. Due to a relatively thick cortex and a relatively high
neuronal packing density, humans have the highest number of cortical neurons
(1215 billions), which is more than the number of cortical neurons found in
cetaceans (whales and dolphins) and elephants with much larger brains up to
10 kg. Furthermore, due to a higher axonal conduction velocity and shorter inter-
neuronal distance, humans have a higher cortical information processing capacity
than these large-brained mammals. The largest differences between humans on the
one hand and all other mammals/vertebrates on the other consist in (1) a strongly
increased growth period of the human brain exposing it to a much higher degree to
education, and (2) the presence of the Broca speech center which is a necessary
prerequisite of syntactical language and developed only recently, i.e., about
100,000 years ago. While these two traits appear to be minor steps in human
biological evolution, they had enormous consequences for human culture and
intelligence.
Humans are proud of their brain and their cognitive abilities, and many of us
including many neuroscientists believe that the alleged uniqueness of human nature
is due to the uniqueness of the human brain. In the following, I will briefly discuss
G. Roth (*)
Brain Research Institute, Bremen, Germany
e-mail: gerhard.roth@uni-bremen.de

176 G. Roth
some popular claims about the human brain that can be found even in the scientific
literature. These claims are (1) The human brain is anatomically unique; (2)
Humans have the largest brain in absolute terms; (3) Humans have the largest
brain relative to body size; (4) Humans have largest cerebral cortex, particularly
prefrontal cortex; (5) Humans have some brain centers or functions not found in
other animals.
First claim: The human brain is anatomically unique. All tetrapod vertebrates
(land vertebrates, i.e., amphibians, reptiles, birds, mammals) have brains that
despite enormous differences in outer appearance, overall size and relative size of
major parts of the brain are very similar in their general organization and even in
many details [19, 27]. More specifically, all tetrapod brains possess a median,
medial and lateral reticular formation inside the medulla oblongata, a pons and
ventral mesencephalon, including a noradrenergic locus coeruleus, serotonergic
raphe nuclei and a medial ascending reticular activating system. There is a corpus
striatum, a globus pallidus, a nucleus accumbens, a substantia nigra, a basal fore-
brain/septum and an amygdala within the ventral telencephalon, a lateral pallium,
homologous to the olfactory cortex of mammals, and a medial pallium, homologous
to the hippocampal formation (at least Ammons horn and subiculum). This means
that all structures required for attention, declarative memory (or its equivalents in
animals), emotions, motivation, guidance of voluntary actions and evaluation of
actions are present in the non-human tetrapod brain. These structures essentially
have the same connectivity and distribution of transmitters, neuromodulators and
neuropeptides in the different groups of tetrapods including man.
A more difficult problem is the presence of structures homologous to the
mammalian isocortex in the telencephalon of other tetrapods. Amphibians possess
a dorsal pallium, turtles and diapsid reptiles have a dorsal cortex plus a specific
structure called dorsal ventricular ridge (DVR), birds have a hyperpallium and
meso-nidopallium, and these structures are believed by many comparative
neurobiologists to be homologous to parts of the cortex and not to the basal ganglia
of mammals, as previously assumed [15, 17, 20, 23]. However, major differences
exist between these structures with regard to cytoarchitecture and size. In
amphibians, the dorsal pallium is small and unlaminated; in lizards it is relatively
larger, and in turtles and some diapsid reptiles it shows a three-layered structure. In
birds, those parts assumed to be homologous to the mammalian cortex (i.e.,
hyperpallium and meso-nidopallium) are large, but unlaminated. In mammals
with the exception of insectivores and cetaceans the dorsal pallium or isocortex
shows the characteristic six-layered structure. Despite these differences it is safe to
assume that the dorsal pallium and cortex of amphibians and reptiles is at least
homologous to the limbic and associative cortex of mammals, while a primary
sensory and motor cortex appears to be absent. When we compare birds such as
pigeons or parrots with roughly equally intelligent mammals such as dogs, then it
becomes apparent that the same or very similar cognitive functions are performed
by anatomically very different kinds of pallium/cortex (cf. [27, 28]).
Second claim: Humans have the largest brain in absolute terms. This is definitely
wrong, as can be seen from Fig. 1 and Table 1. Humans have large brains (1.4 kg
Is the Human Brain Unique? 177
Toothed whale
Man
Ape
Dog
Hare
Armadillo
Fig. 1 Series of mammalian brains, all drawn to the same scale. Evidently, man has neither the
largest brain nor the most convoluted cortex. Convolution of the cortex as well as of the
cerebellum increases monotonically with an increase in brain size
average weight), which is the largest among extant primates (the extinct Homo
neandertalensis had a somewhat larger brain), but by far not the largest one among
mammals. The largest mammalian brains (and of all animals) are found in elephants
(up to 5.7 kg) and whales (up to 10 kg).
Third claim: Humans have the largest brain relative to body size. This is wrong,
too. While the human brain occupies about 2% of body mass, in very small rodents
relative brain size goes up to 10%. However, again among primates, humans have
178 G. Roth
Table 1 List of brain weights Brain weight in mammals (g)

in mammals
Sperm whale 8,500
Elephant 5,000
Man 1,400
Horse 590
Gorilla 550
Cow 540
Chimpanzee 400
Lion 220
Dog 135
Cat 30
Rat 2
Mouse 0.4
Brain weight (g)

10,000
1000
100
10
0.1
0.001 0.01 0.1 1 10 100 1000 10,000 100,000
Body weight (kg)
Fig. 2 The relationship between brain size and body size in vertebrates. Double-logarithmic
graph. Open circles: bony fishes; open triangles: reptiles; filled triangles: birds; filled circles:
mammals except primates; open squares: primates; encircled open squares: Homo sapiens (After
Jerison [8])
the largest relative brain size. The relationship between brain size and body size is
being discussed for more than 100 years (cf. [8]). It appears that body size is the
single most important factor influencing brain size, i.e., large animals generally
have large brains in absolute terms. However, increase in brain size does not strictly
parallel the increase in body size, but follows only to the power of 0.66 0.75 (i.e.,
2/3 or 3/4, depending on the statistics used; [9]), a phenomenon called negative
brain allometry [8] (Figs. 2 and 3). Consequently, small animals of a given taxon
have relatively larger brains and large animals of this group have relatively smaller
brains. Among mammals, this is reflected by the fact that in very small rodents
Fig. 3 The relationship between brain size and body size in mammals. Data from 20 mammalian
species. Double-logarithmic graph (From Nieuwenhuys et al. [19], modified)
Fig. 4 Brain weight as a percentage of body weight for the same 20 mammalian species as above.
Double-logarithmic graph (From Nieuwenhuys et al. [19], modified)
brains occupy up to 10% of body mass, in pigs 0.1% and in the blue whale, the
largest living animal less than 0.01% (Fig. 4).
In addition, the different groups of vertebrates, while satisfying the principle of
negative brain allometry, exhibit considerable differences in their fundamental
brain-body relationship (Fig. 5). Among tetrapods, mammals and birds generally
180 G. Roth
Brain weight (g)

10,000 Man
1000
Mammals
100
10 Birds
Chondrichthyes
1
Reptiles
0.1
Amphibians
Cyclostomes
0.01
0.001 Osteichthyes
0.001 0.1 10 1000 100,000

Body weight (kg)
Fig. 5 Diagrams showing the relationship between body weight and brain weight in the different
classes of vertebrates. Evidently, these classes differ in their general brain weightbody weight
relationship, with the cyclostomes having the smallest and mammals having the largest relative
brain weights. Remarkably, chondrichthyans (cartilaginous fishes, i.e., sharks and rays) have much
larger relative brains than osteichtyans (bony fishes, above all teleosts). Double-logarithmic graph
(After Jerison [9], modified)
have larger brains relative to body volume or weight than amphibians and reptiles,
and among mammals, primates have relatively larger brains than other orders.
Thus, during the evolution of birds and mammals and more specifically of
cetaceans and primates, genetic and epigenetic systems controlling brain size
have undergone substantial changes in favor of relatively larger brains. These
changes resulted in enlargements of brains beyond that associated with body size
[9, 11].
Thus, contrary to a common belief, humans do not have the largest brain either in
absolute or relative terms. Unless we accept that cetaceans and elephants are more
intelligent than humans and/or have states of consciousness not present in humans,
the absolute or relative size of the human brain per se cannot account for our factual
or alleged superior cognitive abilities. However, among relatively large animals
man stands out with a brain that constitutes 2% of body mass. We can quantify this
fact by determining the so-called encephalization quotient (EQ) which indicates the
ratio between the actual relative brain size of a group of animals to the relative brain
size as expected on the basis of brain allometry determined by body size alone
(Table 2). Calculating the EQ for the human brain, it turns out that it is more than
seven times larger than that of an average mammal and about three times larger than
that of a chimpanzee, if they had the size of a human being [8, 9].
Table 2 Encephalization in Encephalization quotient in mammals

mammals (After [1, 8, 9])
Man 7.4
Dolphin 5.3
Chimpanzee 2.5
Monkey 2.1
Elephant 1.9
Whale 1.8
Marmot 1.7
Fox 1.6
Walrus 1.2
Camel 1.2
Dog 1.2
Squirrel 1.1
Cat 1.0
Horse 0.9
Sheep 0.8
Mouse 0.5
Rat 0.4
Rabbit 0.4
While man stands out in this respect among primates, similar processes must
have taken place among cetaceans. Toothed whales, particularly members of the
family Delphinidae, exhibit EQs that are far superior to all primates except Homo
sapiens [16]. While man has an EQ of about 7, dolphins have EQs up to 5, and the
great apes (except man) have EQs around 2. Thus, humans have a much larger brain
than expected among primates, but even in this respect their brain is by no means
unique, as the example of dolphins shows.
Fourth claim: Humans have the largest cerebral cortex, particularly prefrontal
cortex. There are enormous differences both in absolute and relative brain and
pallial/cortical size among tetrapods and among mammals in particular. For exam-
ple, man has a brain and a cortex that are roughly 3,000 times larger in volume than
those of a mouse. This implies that changes in relative size of cortex are inconspic-
uous, because in mammals cortical size rather strictly follows changes in brain size,
but, again, there are differences within mammalian groups. Apes (including man)
have somewhat larger isocortices than other primates and other mammals, because
their forebrains (telencephalon plus diencephalon) are generally somewhat larger
constituting 74% of the entire brain as opposed to about 60% in other mammals
including mice. At 40% of brain mass the human cortex has the size expected in an
ape [9].
The enormous increase in cortical volume is partly the result of an increase in
brain volume and consequently in cortical surface (which is related to an increase
in brain volume by exactly the power of 2/3; [8]), and partly the result of an increase
in the thickness of the cortex. The cortex is about 0.8 mm thick in mice and
2.54 mm in man. Remarkably, both cetaceans (whales and dolphins) and elephants
have unusually thin cortices with 1.21.6 mm.
182 G. Roth
In mammals, the number of neurons per unit cortical volume decreases with an
increase in cortical thickness and brain size. According to data from Haug [5],
cortical neuronal density (i.e., the number of neurons per cubic millimeter of
cortex) amounts to a maximum of 60,00075,000 in prosimians and monkeys,
while with about 25,000, apes and humans have a much lower cortical neuronal
density. However, with 6,0007,000 neurons, whales and elephants exhibit the
lowest cortical neuronal density found in mammals. These data clearly contradict
the much-cited claim of Rockel and co-workers [25] that in all mammals a standard
vertical cortical column contains the same number of neurons independent of total
cortical volume. After Haug such a column contains 190,000 in monkey, and
50,000 (varying between 30,000 and 100,000 across different cortical areas) in
humans and only 19,000 in elephants and whales and dolphins. This decrease in the
number of cortical neurons per unit volume is a consequence of a roughly equal
increase in the length of axonal and dendritic appendages of neurons, in the number
of glial cells and in the number of small blood vessels. Without such an increase in
glial cells and blood vessels, large isocortices would probably be both architectur-
ally and metabolically impossible. We recognize that the dramatic decrease in
nerve cell packing density is only partly compensated for by an increase in cortical
thickness.
Knowing cortical volume or surface, we can calculate the total number of
cortical neurons in mammals [28]. As shown in Table 3, humans with 1215 billion
neurons (depending on cell density estimates) have the highest number of cortical
neurons, because their cortex is relatively thick and has a medium neuronal density
despite the fact that their cortex is by far not the largest one. Humans are closely
followed by whales and elephants; they have much larger cortices, but these are
much thinner, and the neuronal packing density is much lower. In addition, the
cortex of whales and dolphins shows a different cytoarchitecture, e.g., lacking a
distinct cortical layer IV. This is considered by experts due to secondary loss,
Table 3 Number of cortical Number of neurons in the cortex of mammals

neurons in mammals (From
[28] (from Roth and Dicke, 2005)
No. cortical neurons
Species (millions)
Man 1215,000
Elephant 11,000
Whale 10,500
Chimpanzee 6,200
Dolphin 5,800
Rhesus monkey 480
Cat 300
Dog 160
Opossum 27
Hedgehog 24
Rat 15
Mouse 4
because the ungulate ancestors of cetaceans presumably had a normal six-layered

cortex.
In addition to a much higher neuronal density, the human cortex, when com-
pared with that of the large-brained cetaceans, exhibits a much higher axonal
conduction velocity, which is due to a much thicker myelin sheath of cortical
axons [2, 3, 26, 30]. In combination with a shorter distance between pyramidal
cells this suggests that primates including humans possess a much faster velocity of
information processing than cetaceans.
Recently, Kaas [13, 14] argued that the number of cortical areas increased
dramatically from about 20 such areas in the hypothetical insectivore-like ancestor
to more than 60 in primates. However, what has increased according to Kaas
was the number of functionally intermediate areas, but neither the primary nor the
highly associative areas. Kaas is right to warn about the danger of greatly
underestimating the number of functionally different cortical areas in small-brained
mammals.
Available data suggest that contrary to common belief the associative cortex
has increased roughly in proportion to an increase in brain and cortical size. This
apparently is the case for the prefrontal cortex, which is regarded by many
neuroscientists and neurophilosophers as the true seat of consciousness.
Anatomically, the prefrontal cortex is defined as the cortical area with major
(though not exclusive) input from the mediodorsal thalamic nucleus [24, 29].
Using this definition, it turns out that the PFC has increased with an increase in
cortical and overall brain volume within groups of mammals, but here again we find
an additional increase in relative PFC size with an increase in absolute brain size
across mammalian orders: in rats, PFC constitutes 6.5%, in dogs, 8.7%, in cows
9.8% and in man 10.6% of brain mass [10]. What follows is that the human PFC has
exactly the size expected according to primate brain allometry. Of course, cetaceans
as well as elephants have prefrontal cortices which are much larger in absolute
terms than the human PFC, but what they do with this massive highest brain
center, remains a mystery so far.
Fifth claim: Humans have unique cortical structures and functions. We have not
yet found anything in brain anatomy that would explain the factual or alleged
uniqueness of the human brain and of humans regarding cognition and conscious-
ness. Given the fact that Homo sapiens has an absolutely and relatively large brain
and cortex, it appears to be the animal with the highest number of cortical neurons
and/or synapses. Remarkable, however, is the strong increase in relative (and
absolute) brain size in hominid evolution during the last 34 million years. While
in the Great apes as well as in the australopithecines that did not belong to our
ancestors, brain size increases with body size to a power of 0.33, in the hominin
lineage leading to Homo sapiens it increased to a power of 1.73, i.e. in a positively
allometric fashion, which means that brain size increased faster than body size
(Fig. 6). However, the reasons for this phenomenon are completely unclear.
What remains is the question whether there are any anatomical or physiological
specializations in the human cortex that could be correlated with the unique
cognitive abilities attributed to man. As to the general cytoarchitecture of the
184 G. Roth
Hominins
1250 Homo sapiens
Homo erectus
1000
3
1.7
Log endocranial volume (cm3)
=
pe
Slo
750
Homo habilis
Australopithecines
0. 33
Slope =
500 Australopithecus boisei
Australopithecus robustus 34 Gorilla
= 0.
S l ope
Australopithecus africanus
Great apes
Orangutan
Chimpanzee
350
Bonobo
30 40 50 75 100
Log body weight (kg)
Fig. 6 Increase in endocranial volume in the Great apes, Australopithecines and in Hominids.
Double-logarithmic graph (After Pilbeam and Gould [21], modified)
human cortex, it is indistinguishable from that of other primates and most other
mammals. Likewise, no differences have been discovered so far between humans
and non-human mammals with respect to short-term or long-term plasticity of
cortical neurons, the action of neuromodulators etc. Only two traits have been
discovered that could drastically distinguish the human cortex from that of other
primates, viz., (1) differences in growth rate and length of growth period and (2) the
presence of the Broca speech center.
As to (1), maturation of the brain is more or less completed at 2 years after birth
in prosimians and 67 years in monkeys and non-human apes, but the human brain
still continues to mature until the age of 20, which is much longer than in any other
primate [7, 21]. A critical phase in the development of the human brain seems to
occur around the age of 2.5 years. At this time, major anatomical rearrangements in
the associative cortex have come to a stop, and the period of fine-wiring appears to
start, particularly in layer 3 of the prefrontal cortex [18]. As mentioned above, at
this time, human children take off cognitively compared to non-human primates.
Without any doubt, the drastically prolonged period of brain development

constitutes one important basis for an increased capability of learning and memory
formation.
The other trait concerns the presence of the Broca speech center in the frontal
lobe responsible for temporal aspects of language including syntax, along with the
Wernicke speech center in the temporal lobe which is responsible for the meaning
of words and sentences (while meaning is likewise dependent on syntax and
grammar). It is to date unclear whether these speech centers are true evolutionary
novelties. All mammals studied so far have a center for intraspecific communica-
tion within the temporal lobe (mostly left side) which may be homologous to the
Wernicke center for semantics. It has been reported that destruction of these areas
leads to deficits in intraspecific vocal communication [6]. In addition, it has long
been argued that the posterior part (A 44) of the Broca speech center in humans and
the ventral premotor area of non-human primates probably are homologous [22].
The ventral premotor area controls the movement of forelimbs, face and mouth,
which is likewise the case for the posterior portion of the Broca area.
According to a number of primatologists, non-human primates lack a direct
connection between the motor cortex and the nucleus ambiguous, where the
laryngeal motor neurons are situated. In man, bilateral destruction of the facial
motor cortex abolishes the capacity to produce learned vocalization including
speech or humming a melody, while a similar destruction in monkeys has no
such consequences [12]. According to a number of experts, the evolutionary basis
for human language was an emotionally driven stereotyped language typical of
non-human primates. During hominin evolution, the cortex gained control over this
system such that beyond the initiation of hard-wired, innate sounds a flexible
production of sounds and their sequences became possible [4, 12]. Such an inter-
pretation, however, contrasts with evidence of a high degree of sound learning in
monkeys [31] and the mentioned consequences of destruction of left-hemispheric,
Wernicke-like temporal areas in all mammals.
Non-human primates including the great apes are strongly limited even in non-
vocal speech based on the use of sign language or symbols, and these limitations
seem to concern mostly syntax. Accordingly, anything concerning language in the
human brain developed relatively recently or underwent substantial modifications,
and it was probably the Broca center rather than the Wernicke center. Such an
assumption is consistent with the fact that the most clear-cut differences between
humans and non-human primates concern syntactical complexity of language.
Thus, during hominin evolution a reorganization of the frontal-prefrontal cortex
appears to have taken place such that the facial and oral motor cortices and the
related subcortical speech centers came under the control of a kind of cortex that is
specialized in all aspects of temporal sequence of events including the sequence of
action [4]. This evolutionary process appears to have taken place relatively
recently, i.e. 100,00080,000 years ago.
186 G. Roth
Conclusions
It turns out that the human brain is not unique in terms of general structure; rather, it
exhibits the bauplan typical of mammals and more specific of primates. Also,
humans do not have the largest brain either in absolute or in relative terms, although
they possess a brain that is 78 times larger than expected from general mammalian
brain allometry. The size of the human cerebral cortex and of the prefrontal cortex
exhibit a slightly positive allometric growth, which is again typical of mammals.
However, due to a relatively thick cortex and a medium neuronal packing density
found in hominids, humans have the highest number of cortical neurons (about 12
billions) found mammals (and animals), which, however, is only slightly above the
numbers found in cetateans and elephants. The fact that axonal conduction velocity
and neuronal packing density is much higher in humans compared to these large-
brained mammals, this may lead to a much faster cortical information processing as
a basis for human intelligence [27].
The greatest differences between humans on the one hand and all other
mammals/vertebrates on the other consist in (1) a strongly increased growth period
of the human brain exposing it to a much higher degree to education, and (2) the
presence of the Broca speech center which is a necessary prerequisite of syntactical
language. While these two traits appear to be minor steps in human biological
evolution, they had enormous consequences for human culture.
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Aristotle and the Chicken: Animacy
and the Origins of Beliefs
Giorgio Vallortigara
Abstract Mechanisms seem to be available at birth in the vertebrate brains to

distinguish the domain of inanimate objects (for inferring physical causality) and
the domain of animated objects (for inferring social causality). These include
responses to biological motion, self-propelled motion and face-like stimuli in
animals so different as newly-hatched domestic chicks and human newborns.
Detecting the presence and understanding the intentions of other agents is crucial
in order to survive and reproduce. Mechanisms to detect animacy (and agency)
have been argued to underwent a sort of hypertrophic development in our species,
likely because of the demands and the consequent complexities of our social life.
There has been a long road from the primitive animacy detectors that we can see
operating even in simple brains to the intricacies of agency attribution and theory of
mind of human beings. Nonetheless, the origins of beliefs in supernatural things
seem to be deeply rooted in the natural history of animacy detection.
1 A Perceptual Life Detector in the Brain?
In The Descent of Man (Chap. 3) Darwin [17] wrote:

The tendency in savages to imagine that natural objects and agencies are animated by
spiritual or living essences, is perhaps illustrated by a little fact which I once noticed: my
dog, a full-grown and very sensible animal, was lying on the lawn during a hot and still day;
but at a little distance a slight breeze occasionally moved an open parasol (. . .) every time
that the parasol slightly moved, the dog growled fiercely and barked. He must, I think, have
reasoned to himself in a rapid and unconscious manner, that movement without any
apparent cause indicated the presence of some strange living agent.
G. Vallortigara (*)
Faculty of Cognitive Sciences, Director of the Animal Cognition and Neuroscience Lab (CIMeC),
University of Trento, Trento, Italy
e-mail: giorgio.vallortigara@unitn.it

190 G. Vallortigara
Research in modern cognitive neuroscience picked up Darwins implicit sugges-

tion that there maybe primitive neural pathways that ensure a bias to attend toward or
preferentially process, sensory cues about other living things [13, 30] and in
particular members of the same species.
I became interested in what has been called the perceptual life detector in the brain
[24] in an indirect way. While studying filial imprinting in domestic chicks the
process by which the young of some precocial species learn to recognize an object,
usually the mother hen and siblings, by simply being exposed to it for a short period
of time during a critical period soon after hatching [6, 21] I became curious about
the role of motion in the imprinting process. Ethology textbooks state that imprinting
is more likely to be obtained and more strong using motion rather than stationary
stimuli. I wondered, however, whether any type of motion would be identically
effective or whether animals would be in some ways prepared to be sensitive to
certain types of movement. The broody hen shows a very peculiar pattern of
movement and I decided to check whether, before any imprinting has occurred,
newly-hatched chicks would show a tendency to selectively approach such a type
of motion. To this aim, together with my collaborators, I take advantage of point-light
displays [23] that prevent use of two-dimensional information about the shape. An
example is shown in Fig. 1. The picture is obtained by locating 13 point of lights on
the joints of a digitalized image of a moving hen. When a single motionless frame is
observed it is difficult to extract any structure from this very impoverished stimulus.
However, when point of lights are set into motion the shape of the animal and the type
of action it is involved in become effortlessly and immediately available to human
observers. When we tested chicks using these stimuli, however, it turned out that my
initial conjecture was wrong [41]. Newly-hatched, completely visually nave chicks
did indeed show a preference but it was not specific of their own species. Faced with a
Fig. 1 A newly hatched chick confronted with a point-light display of a moving hen (see text for
details)
Aristotle and the Chicken: Animacy and the Origins of Beliefs 191
choice between point-light displays depicting a moving hen versus random motion or
rigid motion (the latter obtained by rotating around the vertical axis a single frame of
points of light of the moving hen) chicks showed a preference for the moving hen.
However, when presented with a scrambled hen, in which points of light were spatially
dislocated at random, though retaining their local motion signals, the walking hen was
chosen as much as the scrambled hen; though both the walking hen and the scrambled
hen were preferred to the rigid and random motion. This finding suggests that chicks
preferentially approach semi-rigid motion, the type of motion which is exhibited by
vertebrate animals. In semi-rigid motion some points maintain a fixed distance from
each other (e.g., two points placed close on the same limb) but can nonetheless vary
their distance with respect to other points (e.g. with respect to points located on the
torso). Such a pattern of semi-rigid motion is shared by the walking and the scrambled
hen, even though the latter does not match any existing biological creature. As a
control for this hypothesis we used the motion of a point-light cat, a species that can
predate on young chicks. As predicted, chicks did not exhibit any preference between
the walking hen and the walking cat point-light sequence, though they did prefer the
walking cat to the random and to the rigid motion point-light sequence.
2 Of Chicks and Faces
The predisposition found for certain kinds of movements shares characteristics in

common with those earlier demonstrated for the head and neck region of a hen to
artificial objects [25]. In 1988 neurobiologist Gabriel Horn in Cambridge, and, at
for a time, his graduate student Mark Johnson, carried out some groundbreaking
studies showing that, contrary to widely held beliefs, filial imprinting seems to
consist of two separate processes: an inborn predisposition of the young bird to
attend to visual stimuli that resemble a broody hen, and a learning mechanism
(which would be guided and supported by the innate predisposition) to learn by
exposition about the specific, unique characteristic of a particular mother hen [25].
Subsequently, Johnson moved to studies on human infant face preferences and
together with John Morton published a series of seminal papers (see for a review
[26]) arguing that a similar two-mechanism device would be available to human
neonates for face recognition. According to their model, infants are born with some
information about the structure of faces. This structural information, termed
Conspec, guides the preference for facelike patterns found in newborn infants.
Conspec is contrasted with a device termed Conlern, which is responsible for
learning about the visual characteristics of conspecifics.
Subsequent work, however, has cast some doubt as to the precise nature of the
Conspec mechanism, in particular it has been argued that contrary to Morton and
Johnsons theory, newborn infants preferences for faces would be a secondary
effect determined by non-specific biases due to constraints imposed by the imma-
ture visual system of the child. In particular, Turati et al. [39] provided evidence
that the preference for face-like stimuli would be determined by an Up-Down
192 G. Vallortigara
bias that would direct babys attention toward any configuration presenting more
elements in the upper part (a top-heavy configuration). However, differently from
chicks, human newborns cannot be completely prevented from being exposed to
faces. Thus, it is unclear as to whether part of the controversy would depend on an
effect of early learning. We have recently reconsidered the issue and our results
suggest that chicks have indeed an inborn preference to approach face-like stimuli,
resembling their head region (e.g., they preferred the stimulus on the left in the pairs of
pictures shown in Fig. 2; [33].). We also showed that both newly hatched chicks and
human newborns demonstrate similar preferences for face stimuli over spatial fre-
quency matched structured noise ([32]; see Fig. 3), providing strong converging
evidence that vertebrates have a domain-relevant bias toward faces shortly after
hatching or birth. Similarly to the preference for biological movement, the preferences
for faces is not species-specific (chicks, for instance, respond to face-like
characteristics of a predator like a polecat; [25]).
Fig. 2 Schematic face-like

stimuli used for experiments
with visually-inexperienced
chicks (see text for details)
Fig. 3 Example of the face (left) and noise (right) stimuli used in the study comparing human
newborns and chicks preferences (see text for details)
When considered together, these findings seem to fit a general scheme for
cognitive development of recognition of other animals based on the interaction
between two separate and independent systems. The first of these systems would
direct the attention of the young animal toward the appropriate class of objects to
learn about, in the absence of any prior specific experience (e.g., in the case of
motion those objects that move semi-rigidly). The second system would be
concerned with learning about the peculiar characteristics of the objects to which
attention has been directed by the first system. Given that in a natural environment it
is more likely that the newly-hatched chick would encounter a mother hen rather
than a cat, a developing predisposition to pay attention to objects showing the
characteristic motion of vertebrates would assure the highest probability to learn
(by way of the imprinting mechanism) about the specific pattern of motion of the
mother-hen. Could these findings relative to biological motion be generalized to the
human species as well? The answer seems to be positive.
An inborn predisposition to attend to biological motion has long been theorized
for the human species, but had so far not been demonstrated. In particular, no
preference for biological motion was reported for human infants of less than
3 months of age [18]. Recently, however, Simion et al. [34] tested 2-day-old babies
discrimination after familiarization and their spontaneous preferences for biological
vs. nonbiological point-light animations, using the same type of stimuli (i.e.
walking hens) used in our research with newly-hatched chicks. Newborns were
shown to be able to discriminate between the two different patterns of motion and,
when first exposed to them, selectively preferred to look at the biological motion
display. This preference was also orientation-dependent: newborns looked longer
at upright displays than upside-down displays (the same result was previously
reported for newly-hatched visually nave chicks, [40]). Overall, these parallel
results in the two species strikingly support the hypothesis that detection of bio-
logical motion is an intrinsic capacity of the vertebrate visual system, which is
presumably part of an evolutionarily ancient and non-species-specific system
predisposing animals to preferentially attend to other animals.
3 Aristotle and the Chicken
Recent research on human infants has provided important evidence for early
sensitivity to causal agency (animacy) and intention (for a review see Biro et al.
[8]). I wanted to investigate whether evidence for an inborn preference for objects
conveying an impression of causal agency or intention could be observed in
inexperienced animals.
In the Physics Aristotle wrote:
Of the proper subjects of motion some are moved by themselves and others by something not
themselves, and some have a movement natural to themselves and others have a movement
forced upon them which is not natural to them. Thus the self-moved has a natural motion.
Take, for instance, any animal: the animal moves itself, and we call every movement natural,
the principle of which is internal to the body in motion. ( vol. V, p. 307) [1]
194 G. Vallortigara
Now, let us suppose to present newly-hatched chicks with a video showing two
objects, of different colours (Fig. 4a), on a stage, initially shown at rest. Then one
object is shown to move slowly until contact with the other object, thus producing
(to a human observer at least) the classical Michottes [29] launching effect (see
[31] for evidence that non-human animals are also sensitive to Michottes perceived
causality). The second object would thus move along a straight trajectory for the
same length as the previous one, stopping before exiting from the stage. After this
exposure phase, chicks would be tested in a free-choice task for spontaneous
preference for the two different objects. Complete balancing for use of different
colours and left-right direction of movement would be ensured during exposure and
at test. If chicks do attribute a notion of animacy to the object that starts moving and
contacts the other object, then a preference for such an object would be observed in
free-choice preference tests, irrespective of its colour and direction of movement.
When tested for their preferences for objects A and B (see Fig. 4b leftmost picture),
we found that chicks showed indeed a preference for object A, the self-propelled
object playing the agentive role during the exposure phase [28].
In order to check whether the perceived causality was crucial, we ran another
experiment in which the order of the displacements was swapped temporally: thus
object B moved first and object A started its movement only after object B had
stopped (Fig. 4b centre picture). In this animation sequence any physical causality
between the movements of the two objects would be disrupted (no contact between
A and B), whereas distances travelled and perceptual features of the two objects
would be identical to those of the launching effect. Both objects would thus appear
to be self-propelled. At test, we found no significant preference for either object,
showing that the results of the first experiment could not, therefore, be due to a
preference for the stimulus that moved first in the animation, since no preference for
stimulus B (which moved first) was apparent in this case. However, given that
in the latter experiment any physical contact was removed, it was necessary to
check whether chicks preferences in the first experiment could be accounted
for in terms of which object applied physical contact over the other object,
which perhaps may have acted as a cue of agency. To determine this, chicks
were imprinted onto a non-causal physical animation. The stimulus sequence was
identical to the launching effect used in the first experiment except for the
presence of a 3 s-delay between the time of contact and the motion of B. In
human subjects, the presence of such a delay is known to abolish any impression
of physical causality: object B would appear in this case as being self-propelled as
was object A. We found that chicks similarly showed in this case no preference
for either stimulus.
It remained unclear, however, whether the choice shown by chicks in the first
experiment was due to a preference for the self-propelled stimulus or to a prefer-
ence for the object which was the cause of the motion sequence. To determine
this, we exposed chicks to a video animation identical to the one used in the first
experiment except for the presence of two opaque screens, one of which occluded
Fig. 4 (a) A chick underwent presentation of the launching effect (see text) (b) Schematic
representation of the animation sequences used during exposure. In Experiment 1 (left-most
sequence), the classical launching effect is reproduced: To humans the sequence appears as object
A being a self-propelled agent hitting object B and causing its movement. In Experiment
2 (central sequence) the order of movement was swapped: The sequence should appear as the
independent motion of two objects, both self-propelled. In Experiment 3 (not shown) chicks were
imprinted onto a sequence identical to that used in Experiment 1 except for the presence of a 3 s-
delay between the objects contacting one another and the start of motion of object B, which would
abolish any impression of physical causality. In Experiment 4 (right-most sequence) no cues were
available as to the nature of motion of object A because of the presence of occluding screens,
though the causality of the launching effect would be preserved
the object at the beginning and one at the end of the motion sequence (see Fig. 4b
rightmost picture). In this way no cues was available about the self/not-self
propelled nature of object A, although it continued to be perceivable as the cause
196 G. Vallortigara
of motion of object B. We found that chicks did not show any significant preference
for either object.
The results of our experiments are thus consistent with Aristotles hypothesis:
only when one of the two objects appeared as being self-propelled and the other did
not, did a preference emerge, as a choice for the self-propelled stimulus. Detection
of self-propelled motion is likely to be part of a primitive neural pathway that
ensures a bias to attend toward, or preferentially process, sensory information about
other living entities, using sensory cues of animacy.
4 From Intuitive Psychology to Intuitive Physics
Evidence has been collected suggesting that human infants are born with a concep-
tion of objects as spatially bounded entities that exist continuously in time and
move continuously in space, maintaining their internal unity and their external
boundaries [14, 36, 37]. This is not limited to the human species, however. Other
organisms as well seem to possess the rudiments of what has been dubbed intui-
tive (or nave) physics. For instance, similar to human infants [3, 4, 14, 15, 35,
36], non-human primates [12] and dogs [27] represent physical objects and reason
about the motion of physical objects in accordance with the basic constraint of
solidity of material bodies. Also, rooks [7] and chimpanzees [11] appear to under-
stand the basic rule that contact is required for support, suggesting that they have
some understanding of the fact that the physical world is governed by unobservable
forces such as gravity.
In seminal experiments by Baillargeon et al. [4] 3.5-month old infants habituated
to a opaque screen that rotated 180 , upwards and then 180 downwards from a
horizontal surface. When subsequently shown an object which was occluded as the
screen rotated upward, infants showed surprise at the sight of the impossible event
in which the screen continued to rotate the full 180 (as during habituation) despite
the apparent presence of the object, whereas they were not surprised when the
screen stopped at the angle where it met the object. This suggests that 3.5- month
old infants form representations of hidden objects and make the inference that a
solid object cannot move through the space occupied by another solid object. But of
course such a knowledge about the properties of physical object may be the
outcome of observing the behaviour of the objects or of acting upon them. Alterna-
tively, it could be that basic concepts of intuitive physics are present at birth and do
not require specific experiences with objects. Recently, we investigate this issue
using our animal model Chiandetti and Vallortigara [16]. Chicks were reared singly
with a small object that became their social partner. They were then accustomed to
rejoin such an imprinting object when it was made to move and disappear behind
either one of two identical opaque screens. At test, after disappearance of the
imprinting object, chicks were faced with two screens of different slants which
could or could not, be compatible with the presence of the imprinting object hidden
beneath them. Chicks consistently chose the screen of slant compatible with the
presence of the object beneath it. When chicks were imprinted on an object of a
smaller size, which could be occluded beneath the screen closest to the horizontal
surface, no preference for either screen was observed. Preventing chicks from
touching and pecking at the imprinting object before testing did not affect the
results, suggesting that the physical constraint according to which a solid object
cannot occupy the space of another solid object is a biological predisposition of
their brains.
5 Animacy Detection and Origins of Beliefs
Several authors (e.g. [2, 5, 9, 10, 20]) have pointed out the implications for our
species of the clear-cut divide that our brains operates between the domain of
inanimate objects (for inferring physical causality) and that of animated objects
(for inferring social causality). In particular the fact that the mechanisms to detect
animacy (and agency) underwent a sort of hypertrophic development in our species
[10], likely because of the demands and the consequent complexities of our social
life [22].
We may appeal to natural causes in order to account for an event. Why did that
apple fall from the tree? Because the wind shook the branch of the tree, causing the
apple to fall. Alternatively, we may explain phenomena by appealing to agents,
entities who act on the basis of their beliefs and desires. Why did the apple fall from
the tree? Because Mary shook the tree, causing the apple to fall down, because she
wanted to eat the apple.
We evolved in an environment containing many agents prey, predators,
conspecific friends and rivals, potential mates and so on. Detecting the presence
and understanding the intentions of other agents is crucial to members of our
species, in order to survive and reproduce. Thus we evolved to be very sensitive
to signals of animacy and agentivity overly sensitive in fact. As stressed by
Barrett [5] humans evolved to have (or, perhaps more plausibly, to be) hyper-active
agency detectors.
Hyper-active agency detectors would explain the human tendency to believe in
the existence of invisible agents, such as spirits, ghosts, angels or gods. As put
forwards by Bloom [9], if body and souls are thought of as separate, there can be
bodies without souls (stones, tables, corpse, zombies) and souls without bodies
(angels, ghosts, demons), opening the possibility to the belief that we ourselves can
survive the death of our bodies.
There has been of course a long road from the primitive animacy detectors that
we can see operating even in simple brains to the intricacies of agency attribution
and theory of mind of human beings. Nonetheless, the origins of beliefs in super-
natural things [19, 20, 38] and of our intuitive dualism [9] seem to be deeply rooted
in natural history.
198 G. Vallortigara
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Evolution: Remarks on the History of a Concept
Adopted by Darwin
Volker Gerhardt
Abstract Evolution is a term invented by the Philosopher Gottfried Wilhelm

Leibniz at the beginning of eighteenth century. Referring to the latin verbum
evolvere, evolution means the step by step development of the organism from
the ovum up to the grown up exemplar. This conception includes the thesis, that all
organisms were totally preformed in the first real exemplar of living being at all.
This thesis was sharply criticized by Immanuel Kant. He saw that there would be
neither learning nor variation in history of life when Leibniz should be right. But in
his political and cultural philosophy Kant contrasted evolution against revolu-
tion and gave it the meaning of a continual historical development. It was this
understanding that influenced Darwin in choosing evolution for his description of
the process of natural development. So one of the fundamental concepts of the
modern science of nature is derived from philosophy, but not in its biological sense
in the tradition of Leibniz, but in the political dimension as Kant pointed out.
1 The Rediscovery of Nature in Culture
There was a time when it was thought that society could subsist in and of itself.
Nature was taken to be the ultimately indispensable substratum for everything that
happened, but it was not thought to have any significance for the assessment of
human action. It seemed that knowledge of nature could only put to technical use
and was thus considered to be deficient, like technology itself. It was only accorded
any significance as a fact, something we could supposedly only refer to positivis-
tically and that was only thought to have instrumental significance.
Human action, in contrast, was thought to depend only on what was considered
societal and historical and could be criticized in terms of political notions.
V. Gerhardt (*)
Institut fur Philosophie, Humboldt-Universitat zu Berlin, Berlin, Germany
e-mail: volker.gerhardt@philosophie.hu-berlin.de

202 V. Gerhardt
That we have overcome this way of thinking is due not least of all to Charles
Darwin. There are obviously other factors that played a role, such as our realization
of the scarcity of natural resources, the susceptibility to epidemic diseases or the
increasing threat of natural catastrophes with the growth of civilization. But is
essentially thanks to the theory of evolution that today we see nature, technology,
society and culture as a differentiated but unified and interconnected whole.
As I hope to demonstrate, this relates not just to the theory of evolution but to the
very concept. For the concept itself opens a historical dimension that binds together
nature, society and culture. But what the theory of evolution accomplished was to
historicize nature, both in its process and in its products.
2 The Historical Constitution of Life
The theory of evolution was not strictly necessary for us to see the interconnected-
ness of nature, history, society and culture. In its earliest beginnings philosophy
started from the assumption of the unity of the cosmos, taking it as self-evident at
first and then later explicitly establishing the point. The ancient materialists as well
as Plato and Aristotle had already drawn inferences from the conception of an
origin to a theory of development. In the modern era these ideas were worked out in
greater detail, primarily under the influence of Newtonian theory, and by the
beginning of the eighteenth century they had been elaborated into a natural history.
The young Kant had even explained why one could only treat of a theory of the
heavens within the framework of a general natural history that encompassed the
development of the earth and even of the life on earth.
Biology also began with the assumption of a unified origin of all life and saw no
reason to exempt humanity and the human mind from the events of life. This is even
rooted in the very method of biological classification: for the researcher seeking to
identify and organize genera and species, the individuals are only examples
illustrating what is significant for the character of the species.
In fact, as much as biology deals with the individual organism, it is ultimately
oriented towards the general forms of life, of which the individuals are merely
representatives.
If we turn this around and say that the forms of life are themselves always
representations of individually inherited traits, we can see how close biology is to
the central problem of the social and cultural sciences, since it deals with the
general characteristics of individual phenomena of life that not only outlast
the existence of their individual instantiations but shape and guide them. And if
this includes social formations such as pairs, packs, herds, swarms, and colonies, it
becomes clear that as soon as biologists turn to humans (a mammalian species),
they have to take into account our social, moral, religious, and political
accomplishments as well.
Thus methodologically biology occupies the same territory as the social and
cultural sciences. That the cultural accomplishments of humanity including
Evolution: Remarks on the History of a Concept Adopted by Darwin 203
technology, stemming from the appropriation of individual natural processes when-

ever conducive to life belong to one reality, cannot, in any case, be a controversial
proposition within philosophy.
Nonetheless, Darwins theory of evolution brought greater depth and clarity to
this insight. It advanced our knowledge of the interconnectedness of nature, life,
technology, history, society and culture. More than that: from its very beginning it
involved questions of cultural evolution. In what follows I hope to make this clear
by means of a small detail in the history of the concept.
3 The Origin of the Concept in Leibniz
In } 74 of his Monadology, the founder of the Prussian Academy of Sciences,

Gottfried Wilhelm Leibniz, speaks of a problem that had already troubled
philosophers before his time, and tries to resolve it with recourse to the latest
research results involving the microscope:
Scientists have had great difficulties over the origin of forms, entelechies or souls. But now
that meticulous research has been carried out on plants, insects, and animals, it has been
recognized that naturally organic bodies (les corpes organique de la nature) are never the
product of gas or rotting, but always of seeds (semences), which undoubtedly contain some
sort of preformation. The conclusion has been drawn that, not only does the organic body
already exist before conception, but also a soul in this body in a word, the animal itself
(lanimal seule). The only function of conception is to precipitate a major transformation,
so that the animal becomes an animal of a different species. Even outside the process of
generation, something similar is observed when maggots become flies, or caterpillars
become butterflies
We can see this proposed solution, published in 1714, as an equally metaphysical

and biological reaction to the increasing interest within the sciences in questions of
the anatomy and physiology of the human body as well as in the problems of life per
se. Thus it is not surprising that it is precisely this passage from the Monadology that
generated a great amount of interest and played a not insignificant role in developing
the problems that contributed, in the course of the eighteenth century, to the
founding of that science that soon came to be known as biology.
These questions revolve above all around what Leibniz called developpemens,
developments.1 This primarily concerns the thesis of preformation advanced in
the text, that is, the claim that the peculiar characteristics of a living being are
already determined at the time of its formation.
We can see the range of variation that Leibniz allows for when he refers to
the metamorphosis of the maggot to the fly and the caterpillar to the butterfly. At the
same time he believes that the traits that are decisive for heredity and way of life are
1
Leibniz contrasts developpement with enveloppment, the degeneration or decrease in the living
forces. Developpement unwraps that which had previously been enveloped.
204 V. Gerhardt
originally inherent in the organism at the time of its formation and unfold them-
selves into their characteristic form (such as that of a horse, a cow or a human)
during growth and maturation. Development is just a process of unfolding of that
which God put into the world in the form of countless monads, a process that recurs
anew in each individual.
4 On the Sense of the Metaphor of Evolution
Leibniz had already used the term evolution to describe this ontogenesis of the
individual living being from the microscopic elementary traits that it came
equipped with, and the term began to circulate in the eighteenth century, first in
Latin treatises and then increasingly in the French.2 Some time before the French
Revolution other European national languages began to follow suit.
In Latin Evolution was already an artificial coinage, derived from evolvere,
which had originally meant to unwrap, to unfold and was generally used in
reference to the unrolling of a scroll. The opposite is envelopment, to enfold or
envelop. It means death. Developpement or evolution unwrap that which had
already been placed inside. This is the original sense of the word that the theory
of evolution takes its name from.3
This etymology brings out the process of enlargement as well as that of becom-
ing visible and knowable, aspects that are a prerequisite for any adequate under-
standing of the concept of evolution. This persists today in the concept of the
phenotype, which brings the genotype into view. Moreover, the unfolding (of a leaf)
or unwinding (of a scroll) has a certain proximity to reading, which can be almost
taken literally today since the progress of evolution can essentially be read off of
the genetic sequences.
Thus the term evolution refers first of all to the unwrapping of the inherent traits
of a living creature from its embryo to the matured living individual. But the
preformation thesis at the same time extends the term to include the sequence of
generations, since the embryo only comes into existence through the merging of
ovum and spermium and bears the traits of at least one parent. Thus it was thought
there had to be a line of continuity leading back to the first parents driven from
Paradise.
Yet things didnt end with this first sense of the term. Rather, the explanatory
power Leibniz hoped to achieve for the generation of individuals met with a decisive
critique in the emerging biology in the late eighteenth century and in the philosoph-
ical theory of life with which Kant brought new life to the thought of his age.
2
Wolfgang Wieland: Entwicklung/Evolution, in: Geschichtliche Grundbegriffe, eds. Brunner
et. al., vol. 2, Stuttgart 1992.
3
Evolvere also had the meaning of driving out and displacing, both of which recur in the later
theory of evolution, namely in the survival of the fittest and in the process of selection.
Thus it came about that Darwin did not name his theory of inheritance after the
term that Leibniz had meant biologically in his Monadology, but rather took up a
concept of evolution with a philosophical application to culture and history. Hence
the prospect of a cultural evolution is inherent in the use of the concept prior to the
emergence of Darwins theory.
5 Kants Critique of the Concept of Evolution
In } 81 of his Critique of Judgment, a book that offers, in its first part, an aesthetics
based on the experience of life, and in its second part a comprehensive theory of life
compatible with the causal explanations of natural science, Immanuel Kant
distinguishes between educt and product. He sees product the way we see it today:
as a new creation of nature with its particular origin and its individual character.
Every technical production can be taken as a product, but every living creature can
also be seen as a product, as something newly created, as we still do today.
On Kants account we are only justified in this so long as we do not do so on the
basis of the theory of evolution founded by Leibniz.4 As Kant shows in his
detailed examination of its central theses, the theory actually does not allow
anything new. According to this theory of evolution, the only thing that develops
is the seed that had already been inherent in the sequence of generations from the
very beginning. This theory of evolution should really be called a theory of
involution a theory of nesting that in each individual only acknowledges that
which must already have existed in the origin of life.
If this origin is to have any efficacy, the theorist of evolution who follows
Leibniz has to have recourse to something supernatural, a hyperphysics, in
order to explain how the seed containing in advance each living creature can
have come into nature in the first place at least according to Kant.
Thus Kant sees a contradiction between the preformation theorem of evolution-
ary theory and the facts of life, a contradiction that cannot be reconciled under the
conditions of modern physics. For life brings forth something new with each new-
born creature. For Kant the metaphysics of evolution is not able to acknowledge the
innovative productivity of all that is alive.
6 Three Knock-Down Objections
The general objection from any thinker who insists on the unity of nature is
reinforced by three points that we could call the life-world objection, the phenome-
nological objection and the biological objection.
4
Kant. Kritik der Urteilskraft, } 8, Akademie-Ausgabe 5, 423.
206 V. Gerhardt
First Kant notes that the hypothesis of involution makes copulation . . . a mere
formality.5 If the individual with all of its traits is already contained in one cell,
then the addition of a second cell can hardly be more than an external impulse.
Maybe it serves as food? at any rate this hypothesis has some plausibility if we
assume, as Kant does, charitably as it seems, that the preformed embryo is located
in the male sperm cell and the nutriments in the egg. The proponents of Leibniz
position, as Kant ironically notes, proceed from the contrary assumption and thus
have to explain how the male sperm could play any role whatsoever.
Kants second objection is founded on the pervasive interdependency of nature
throughout all its processes and notes how the involution hypothesis of the
Leibnizian theory of evolution interrupts precisely these elementary intercon-
nections: if Leibniz were right, there could be no interaction between the embryo
and its environment. A great number of supernatural arrangements would be
necessary to preserve the embryo formed in the beginning of the world from any
harm that might come from the surrounding forces. Thus the influence of
hyperphysics in the world would not just be limited to its beginning but would be
required again and again.
The third objection is less theoretically elaborate but not any less decisive: if
Leibniz theory of evolution were correct, there could not be any hybrids.
Hybrids could absolutely not be accommodated with the system of preformation.
In this system the male seed would serve as nourishment for the embryo, but would
not exercise any purposive formative power, which is impossible according to the
premises of the theory of evolution.6
Following these objections, each of which is a knock-down argument, Kant opts
for a theory that sees living creatures as true products of nature, which in their
individual development are subject to the influence of ongoing forces. On this theory
living creatures are generated in accordance with the principle of an original
organization, for which he assumes the productive potency of a formative
impulse.7 Kant outlines a theory of self-organization that is still current today, a
theory that sets out the natural conditions for his ethical theory of self-determination
though I cannot discuss this in any more depth here.8
5
Ibid.
6
Ibid., 424.
7
Ibid. Kant is probably drawing on the second edition of Blumenbachs text Uber den
Bildungstrieb und das Zeugungsgeschafte, G ottingen 1789. For more on the historical and
systematic context of these considerations see: Siegfried Roth, Kant und die Biologie seiner
Zeit, in: O. Hoffe (ed.), Immanuel Kant: Kritik der Urteilskraft (Klassiker auslegen vol. 33),
Berlin 2008, 275 288.
8
See V. Gerhardt, Selbstbestimmung. Das Prinzip der Individualitat, Stuttgart 1999.
7 Evolution as an Alternative to Revolution
If history followed a rational course founded on human insight, the concept of

evolution would have ended its career in the year 1790. Kant shows that the concept
as Leibniz understood it is unsuited to comprehend the essential aspect of life,
namely that something new continually emerges with and in each individual.
But of course the surprising thing is that its career only really took off at that
moment and with Kants help. For he not only continued using the concept after
having shown its unsuitability in the context of biology, but opened up a new field of
application for the concept in his theory of history and culture.
He did this by transferring the concept to the developmental dynamics of human
civilization. And it is, in fact, a quite significant question how we should conceive
the continuous development of law, if its conditions, namely freedom, equality
and the autonomy of individuals, are to be preserved throughout. Kant believes
that only the republican principle of law and the monopoly on violence necessarily
joined with it can give the future of humanity a self-determining form. Education,
morality and religion are indispensable, but they cannot do very much to bring
about progress in human culture that at the same time allows for the validity of the
law of each present era, if nothing ensures the continuity and stability in this
development.9
But securing a future in freedom, equality, and autonomy, requires the continuous
security of the principles of law and a form of the state that is capable of ongoing self-
improvement. This also requires that the state reform itself from time to time and,
attempting evolution instead of revolution, progress perpetually toward the better.10
Here we have the new use of the concept it refers to a perpetual progress
within the course of a history determined by humanity itself. Strictly speaking it
describes the political maxim to avoid revolutions. And it is only in transferring this
normative conception to the course of historical developments that evolution is
made into a descriptive term used to represent an internally cohesive historical
course without break and without revolutionary upheaval.
8 Revolutions in Nature
Kant had widely-read precursors for this normative political and historically
descriptive use of the concept of evolution in his students Herder and Erhard.11
Both draw on considerations from early Kant and inspired their teacher, who had
9
Kant, Der Streit der Fakultaten (1798), 2. section 10; Akademie-Ausgabe 7, 92 f.
10
Ibid., 93.
11
A few pages before this quotation Kant acknowledges his student Erhard. He speaks of the
gradual development of the moral traits of humanity, which are more and more capable of taking
208 V. Gerhardt
since advanced in age; for in his early Cosmology and his first lectures on Physical
Geography he presented the formation of the cosmos and the development of the
earth has an ongoing event of natural history though without using the concept of
evolution.
This historical use of the concept of evolution in application to politics, society
and culture persists even today. Evolution is the opposite of revolution
although there has also long since emerged an understanding of revolutions as
something that can occur over longer periods of time, such as the industrial
revolution or the scientific, technological, or so-called sexual revolution.
Incidentally, Kant was already familiar with an understanding of revolutions as
spanning longer stretches of time, as shown by a remark in the Critique of
Judgment. He is particularly close to Darwin in this remark: after the great eras
of the formation of the solar systems with their cooling planets, the longer periods
of earths history saw recurring spurts in the development of life. First the seas are
populated, then life colonizes the swamps and finally living creatures succeed in
conquering the land. Kant calls these cataclysms in the developmental history of
life the oldest revolutions, of which only traces survive, which the
archeologist of nature is to follow. In this context the earth is apostrophized as
universal mother, as a womb and as the motherly womb.12
Eight years later Kant would have been able to say that these revolutions of
nature are not contrary to the thought of an evolution of life on earth. This is of
course an evolution that cannot be understood according to Leibnizs proposal, but
rather only according to Kants own model, which can be transferred from an
understanding of continuous societal developments oriented around principles to
an understanding of nature even though there are no legal principles and
constitutions in nature. But there are law-like regularities that belong to the
mechanics of nature and the self-organization of life.
In transferring the concept of evolution, which initially describes a political
maxim for the peacable achievement of reforms, to the development of culture,
Kant strips away the biological significance that Leibniz had meant the term to
have. Yet by way of a detour through a speculation about the entirety of the
dynamics of society, wherein Kant sees a continuation of nature with other
means, he gives back to the concept its connection to life. Within his critical system,
society, politics and culture do not just stand alongside nature unrelatedly. Rather
the ongoing progress of civilization should bring together nature, society, politics
and culture into an alignment that is not just retrospectively recognized but also
prospectively hoped for. This is possible if humanity makes the antagonism of
part in the destiny of the race: This even is the phenomenon of, not of revolution, but (as Erhard
expresses it) a phenomenon of the evolution of a constitution in accordance with natural law.
(Akademie Ausgabe 7, 87) The reference to the text of his enthusiastic adherent Johann Benjamin

Erhard, Uber das Recht des Volks zu einer Revolution, Jena 1795, shows how new this use of the
concept of evolution for historical and cultural theory still is for Kant.
12
Kritik der Urteilskraft } 80; 5, 419.
nature productive in itself and proves capable of self-determined development of

their natural traits in regulated competition and legally organized opposition.
9 Darwin as the Heir of the Full-Fledged Concept
Once we see how Kant strives to bring together a dynamically conceived nature
with the unfolding of human culture and to unite this in turn with the development
of law, it would be quite erroneous to say that he removes the concept of evolution
from nature and gives it over entirely to self-determined political movement.
Rather, in Kants critical philosophy the concept is available as a means to
interpret all events that change over time. And my concluding thesis is that Darwin
assumed this concept of evolution, though hesitantly, to describe his theory.
What is noteworthy about this conceptual history is that Darwin began with a
concept that was quite loaded with connotations from politics, the philosophy of
history and cultural theory rather than using, say, a biologically restricted term. The
Origin of the Species and The Descent of Man take up the concept of evolution in its
most comprehensive sense and from the outset bring together the problems of the
natural history of life with all aspects of plant, animal and human behavior.
Consequently it should not come as a surprise that Darwin came to write about
the problems of the emergence of human expression and of the feelings, of empathy
and even the development of morality, and did so from within the logic of his own
conception.
Darwin worked on a description of the transitions between nature, society and
human culture with the same intensity with which he sought to understand the
selective and diversifying mechanisms in the development of the species. For him
life does not stop short of the phenomena of history. He is also able to give a place
in the life of humanity to philanthropy and religion.
For this reason his theory obliges anyone who takes it seriously not to break life
down into an organic substratum and a purportedly intellectually aloof superstruc-
ture, but rather instead to uncover the dynamic unity between the mechanical and
the organic forces.
This is also the source of the obligation not to let the sciences themselves come
asunder into contrary cultures. Instead they must recognize that they arose from the
necessity of self-guidance in human culture and thus have to assume the responsi-
bility for their self-organization in coordination with societal forces.
The most important argument for the unity of the sciences follows from the fact
that evolutionary theory can prepare us like no other to see how that which we call
intention, need, meaning, sense, consciousness or mind arose from the
natural development of life. Darwin enables us to finally give the functions of
reason their position and role in life rather than having to derive them from their
own premises (in the always unsatisfying form of self-confirmation).
Thus reason does not have to keep repeating why it holds itself to be so important
if it can see how it became necessary and under what conditions it is in fact
210 V. Gerhardt
indispensable. I believe that evolutionary theory can liberate reason from the
burden of its thousands of years of self-confirmation and lead it back to the
conditions that preceded it that are themselves not yet rational. But they might
show us why reason is neither so unreasonable nor so irrelevant as it can seem from
the perspective of its critics.
I cannot think of any other problem that the natural sciences and the humanities
should take a greater interest in. For it is in the natural elucidation of the origin and
the potential achievements of reason and consciousness that both fields have the
chance to shed light on themselves as well and to clarify why they not only emerged
from the same impulses of curiosity, knowledge, and rational guidance, but continue
to depend on one another.
10 Addendum
Darwins results also began the mediation of the dispute between Kant and Leibniz.
For Darwins development of the theory of evolution showed that our genes contain
those enduring instruments of control of life that demonstrate a surprising con-
stancy despite their mutability through mutations. The genetic control of life
secures both continuity and variability.
This result is not just interesting for the history of philosophy. It should also
appease the Berlin-Brandenburg Academy of Sciences. For Leibniz and Kant were
both members of its precursor, the Prussian Academy of Sciences. Darwin
reconciled them.
An Evolving Research Programme:
The Structure of Evolutionary Theory
from a Lakatosian Perspective
Telmo Pievani
Abstract The main topic of the paper is a discussion of the ways through which the
theory of evolution remakes itself, changes and grows, keeping alive and
reinforcing its Darwinian explanatory core. The theory shows a 150 years old
history of theoretical and empirical extensions and revisions, without any apparent
radical change of paradigm and without a rival Research Programme able to
replace it. The ongoing transition from the Modern Synthesis (MS) to a so-called
Extended Evolutionary Synthesis (ES) is here interpreted through the Methodol-
ogy of Scientific Research Programmes, proposed by the epistemologist Imre
Lakatos and updated. The current situation in evolutionary biology could be
represented by a progressive shift of the Darwinian research programme, moving
from the quite rigid theoretical framework of the standard version of Modern
Synthesis (gradualism, extrapolationism, adaptationism) to the more inclusive and
pluralistic core and protective belt of the Extended Synthesis. Promising and
advanced researches like those concerning evolutionary developmental biology
(Evo-Devo), epigenetics, multiple ways of speciation and the role of structural
internal constraints find in this perspective a realistic interpretation as theoretical
and empirical novelties with huge implications, nevertheless not incoherent with an
extended Neo-Darwinian explanatory core. A Neo-Lakatosian approach seems
useful when we discuss the extension of evolutionary models in non biological
fields, avoiding the application of just metaphorical forms of ultra-Darwinism.
This analysis in terms of a rational and continuous dynamics of growth of biological
thought seems much needed also for a critical examination of some popular and
radicalized controversies about the health of a no better defined Darwinism or
Neo-Darwinism.
T. Pievani (*)
Philosophy of Science, University of Milano-Bicocca, Milan, Italy
e-mail: telmo.pievani@unimib.it

212 T. Pievani
1 What Makes Biology Unique (and What It Does Not)
The elements of uniqueness and epistemological autonomy of evolutionary biology

(or, better, of the set of evolutionary fields), firstly outlined by Ernst Mayr [33],
could be updated today in the following lines:
The apparent absence of universal laws, due to the abundance of descriptions
and singular existential assumptions;
The historical nature of the explanations (narrative dimension, historical contin-
gency, the role of chance, unrepeatable events, irreversible processes);
The diffusion of objects of study with features of uniqueness, ambiguity in
definitions, unlikely to be categorized and ascribed to an unambiguous set of
phenomena;
The nested multiplicity of spatial-temporal levels of analysis and the stratified
hierarchy of non-reducible patterns of explanations (like in the case of emergent
properties between different levels of organization, or in the case of different
spatial and temporal focalizations between macroevolution and microevolution,
with the need to integrate non reducible patterns emerging at the nested levels of
genes, cells, organisms, populations, species, ecological systems and so on).
Nevertheless, it is interesting to observe that now three other elements of
uniqueness (in a negative sense, with respect to the epistemological status
of physical sciences used as a model) are no longer considered as crucial, because
of the ways through which the theory of evolution is able to deal with evidences
today [57]:
The alleged impossibility of falsification of evolutionary hypotheses, avoided
thanks to the strongly convergent evidences coming from very heterogeneous
fields like paleontology, comparative anatomy, molecular biology, cladistics,
paleo-ecology and others; using such convergent proofs, the evolutionary
reconstructions could be compared and discussed in terms of parsimony (like
in the case of phylogenies) and explanatory power, selecting alternative models
case by case;
The alleged absence of experimental methodologies based on the repeatability of
experiments and the modulation of parameters in laboratory (researchers can
simulate and experiment selective pressures acting on populations of
microorganisms, observing in laboratory dozen of thousands of generations
with their mutations and rates of genetic change; evolutionary developmental
biology and synthetic biology are pushing ahead the experimental status of
evolutionary researches, with the possibility of tuning the molecular parameters
of living beings, and in other fields like cognitive ethology and evolutionary
psychology it is time for an extensive use of comparative behavioral experiments
in different experimental situations);
The alleged incapacity to produce genuine scientific predictions: this is not the
case anymore, when several evolutionary disciplines have predictive models
An Evolving Research Programme 213
testable both in ecological fields, and even now in the wild [23], and in labora-
tory with micro-evolutionary experiments.
The relatively recent adoption of classical experimental procedures cannot
justify, anyway, a reduction of the evolutionary biology to the epistemological status
of mathematical and physical sciences (status itself evolving). The mimicry with
physics is not realistic when we have to deal with historical, hierarchical and singular
processes. Not only past evolutionary events, but also current phenomena like
speciation, are unlikely to be observed and reproduced in laboratory. It seems more
convenient to outline a specific epistemological status for evolutionary researches
different, and certainly not weaker than those of other disciplines where reproduc-
ibility and classic experimental verifications represent a subset of the methodologies
that evolutionists should adopt, including the consilience of heterogeneous evidences
(like in the case of the total evidence approach to phylogenies), comparative proofs,
a multiplicity of coherent observations, historical reconstructions based on direct and
indirect evidences coming from several local disciplines. Only this articulated set of
methodologies seems able to manage such a complex and interdisciplinary empirical
basis. So,
The modern theory of evolution explains a huge amount of single, verifiable
facts, in the past and present, and frames of verifiable facts;
These facts are logically connected and produce coherent frames of corroborated
evidences;
These connections of explained facts allow to formulate risky predictions and
retro-dictions, both verifiable or falsifiable with new facts, enlarging progres-
sively the empirical content of the theory through a process of criticism and
growth of knowledge.
What could we say about the structure of a scientific theory able to do that? The
elements of uniqueness, above described, usually expose the reconstructions of
the history of biological thought to quite appealing radical interpretations of the
ongoing theoretical changes, as if we had more theories of evolution or, even,
more alternative languages on the field. This is the case of the call for the concept
of paradigm, in Thomas Kuhns sense [28], in the description of the history of the
different stages of the theory of evolution after Darwin. According to Mayr [33],
paradigm is too strong a concept in order to understand the theoretical fluidity and
the heterogeneous basis of empirical facts that we find in natural history. Like
Massimo Pigliucci recently noted [53], there is nothing in the field at the moment
that could suggest the typical features of a paradigm shift between incommensu-
rable explanations and conceptual languages.
If we would say that the current theory of evolution is a paradigm, actually we
do not see the accumulation of serious anomalies and the dogmatic crystallization
that should precede a paradigmatic crisis. What is typical of the field is, on the
contrary, the fact that new problems and apparent exceptions can be resolved and
understood mainly through integrative explanations, different modulations of the
empirical domain of application of already established patterns of explanation, new
214 T. Pievani
calculations of the relative frequency of a pattern with respect to another. It means

that the dynamics of growth and evolution of the theory is based on processes of
theoretical extension and empirical enlargement of an elastic set of explanations
already consolidated but constantly needing adjustments and integrations [2].
Something completely different from a revolutionary overthrow.
Does it mean, conversely, that we are in the middle of a static and prolonged
period of normal science, with just marginal scientific puzzles to be solved? Not
properly. An articulated set of explanations that brings together heterogeneous
facts, absorbs new facts, updates its theoretical toolkit when it is healthy, or
accumulates anomalies and auxiliary hypotheses when it is sick and it plods
along the experimental novelties, was defined neither simply a theory, nor a
paradigm, but a scientific research programme by epistemologist Imre Lakatos
[29]. It means that the evolutionary explanation is submitted to continuous changes,
even very deep sometimes, and appears like an open and busy yard, not like an old,
traditional building. So, the theory of evolution is evolving, or better, the evolu-
tionary scientific programme is evolving. But how exactly? Along which lines?
What is under potential falsification here is not a single concept or the content of
a single theory, but a succession of theories and integrated models: in other words
the rational dynamics inside the process of updating and redefinition of a coherent
whole of explanations, included the strategies of defense against contrary evidences
and apparent contradictions [29, 37]. The process of criticism and growth of the
evolutionary knowledge after Darwin seems a continuous one made by
extensions, revisions, and even reversions to originally Darwinian insights [9]
and not a discontinuous, paradigmatic one. Without any underestimation of the
importance of external, social and psychological, factors in science, the pivotal
course of the conceptual change is an internal, logical one, depending on new
corroborated evidences and theoretical advancements.
2 How Many Darwinisms?
A flexible structure of the evolutionary scientific programme suits with the standard
pattern of the history of biological thought very well, intended as a succession of
stages of integration, revision and expansion. Synthetically:
The originally Darwinian explanatory frame, structured with: (a) a wide descrip-
tive apparatus (evolution as a matter of fact, common descent with
modifications, the tree of life); (b) an integrated set of multiple explanatory
factors, such as individual and non-directed variations, natural and sexual
selection (the variation-selection core), and Lamarckian residuals (where
each concept has had afterwards a different fate [31, 32]); (c) some meaningful
risky predictions and retro-dictions, with different fates as well (about the depth
of time, the pervasive gradualism, the hypotheses concerning the evolution of
complex structures, the role of geographic isolation); (d) a powerful cultural
revolution, but technically not a paradigm shift, being rather a transition from
pre-scientific views like natural theology and the entry of the first general
scientific programme in evolutionary fields (adopting already circulating idea,
but in a new and wholly naturalistic explanatory frame).
The original Neo-Darwinism (term coined by George J. Romanes, a pluralistic
Darwinian, interested in ethology and evolution of the animal minds, and used
also by scholars like Alfred R. Wallace and Asa Gray), at the end of the
nineteenth century reinforces the Darwinian core, removes the Lamarckian
residuals and establishes with August Weissman the bases for the conceptual
separation of hereditary biological materials and their bearers, giving to natural
selection a quite exclusive explanatory priority.
The Evolutionary Modern Synthesis (MS) preceded by a phase of eclipse of
Darwinism (according to Julian Huxley, 1942, and then [3]) during the re-naissance
and rediscovery of Mendelian genetics and the birth of macro-mutational and
saltationistic theories of biological change could be represented as a kind of
powerful Neo-Darwinism of second generation, with the fusion of Mendelism
and population genetics, the first mathematical models for the changes in the
frequencies of genetic variants in populations, and a genetic theory of natural
selection; at that time, the convergence of two traditions of research created the
first global evolutionary scientific programme, with a strongly coherent theoretical
frame (the variation-selection core), a great expansion of predictive power, entire
new fields of evidences for the Darwinian core of the theory of evolution; at the
same time, the synthetic power of MS was reached through a crystallization around
few methodological assumptions (in some cases so strong and normative, espe-
cially in worldwide handbooks, that they seemed quite paradigmatic), like the
extrapolation of any macro-evolutionary phenomenon from micro-evolutionary
processes, gradualism and uniformitarism, a diffused adaptationism [20], the
underestimation of evidences (with exceptions in single authors [35]) coming
from embryology, ecology and human evolution [16, 45].
Since the sixties of twentieth century, we see a new rapid extension of the
empirical basis of evolution, together with technological advices, because of
the molecular revolution, the acceleration of genomics and post-genomics, the
new detailed phylogenies, the role of the evolutionary developmental biology
(Evo-Devo), the growth of epigenetics, and there is a general consensus about
the fact that we are in front of a contemporary Neo-Darwinism, the third
generation, quite different from the previous ones.
Disciplines less highlighted in MS, embryology and ecology, are acquiring their
deserved, lost centrality. But mostly, MS was subjected to deep theoretical
challenges, like neutralism, Punctuated Equilibria, and now Evo-Devo itself. The
assimilation of these challenges and the metabolism of such a huge amount of new
data were much slower and harder than any other passage in the past decades, and
claimed a profound change in the structure of the evolutionary scientific
programme. So what kind of Neo-Darwinism do we have today in the field? Do
we need a completely new Evolutionary Synthesis or will some superficial restyling
216 T. Pievani
be enough? Is the new structure of contemporary Neo-Darwinism already stabilized

or in a phase of transmutation? The Lakatosian methodology of the Scientific
Research Programmes (SRP) seems the right candidate in order to understand the
ongoing transition.
With respect to the alleged incompleteness of MS frequently addressed in
contemporary debates, it seems quite clear that MS showed a great capacity of
inclusion [2], through processes of theoretical assimilation that enlarged its empiri-
cal basis and prevented logical and explanatory objections. But, in a first phase, this
assimilation produced the need for a more pluralistic way to understand the
rhythms, the units and the levels of evolution.
In the case of Punctuated Equilibria theory [15], after decades of debates the
general consensus around the mechanisms of speciation is that we need a multiplic-
ity of processes and modes of birth of new species (punctuated in some ecological
circumstances and gradual in others), a multiplicity of possible rates of speciation,
and a multiplicity of levels of change (from an ecological and a genealogical point
of view) to be considered [8]. So the main methodological stance today is a
calculation of the relative frequencies of one pattern (punctuationism) with respect
to another (gradualism and trends) [44], and not a radical alternative between two
incompatible patterns.
What we see is precisely a balance between points of breaking of past methodo-
logical stances inside MS (like phyletic gradualism) and points of theoretical
continuity. In the case of Punctuated Equilibria, the points of breaking are:
punctuations are not due to imperfections of the geological record; speciation is
not only anagenetic, but frequently cladogenetic; speciation is connected with
major episodic evolutionary changes; the wide diffusion of apparent stasis in
natural histories. The points of continuity with MS are: the evolutionary
mechanisms in action during the speciation are Darwinian; gradual trends (plurality
of patterns) are not excluded; and mostly, punctuation and stasis stand at the level of
geological scale of species life, so they do not clash with normal mechanisms of
change at the level of populations of organisms [20].
A quite similar process of assimilation of new evidences with a consistent
internal theoretical accommodation is at the core of the history of another challenge
to MS: neutralism [26] and contemporary weak-neutralism. In the frame of a
renewed MS the evidence of a huge amount of variations and sequences inside
the genome with no adaptive and selective origin is accepted with the cost of a
robust quantitative and mathematical integration in the models. Neutralistic
patterns based on drifts and on structural internal mechanisms, non selective at
the level of organisms, show that natural selection is not the exclusive factor of
genomic change, but an important one among others. Like in the case
of punctuationism, we need a calculation case by case of the relative frequencies
of selective patterns and drift patterns when we look inside the structure of the
genome.
A quite different example is the challenge posed by the discovery of the
crucial role of macro-evolutionary patterns in evolution (like turnover pulses of
species, rapid adaptive radiations, mass-extinctions), because it breaks the strong
methodological assumption that every macro-evolutionary phenomenon should be

extrapolated by the uniform accumulation of micro-evolutionary processes. The
integration needed seems more and more profound. Other recent fruitful fields of
researches threaten the capacity of inclusion of MS, because they touch fundamen-
tal ribs of the theoretical architecture of MS or in some cases add entire new
domains of experimental evidences that claim for a powerful theoretical updating:
The discovery of families of genes and hierarchies of genes, with a still up to
now underestimated complexity of the genetic regulation, changes the idea itself
of the genome, the machinery of the mutations and the phenotypic effects, the
definition of the concept of gene; so, the raw material of any evolutionary
process is no longer so raw;
Evo-Devo suggests a crucial role for the constraints to variation, for the internal
developmental constraints, for systems innovations, functional cooptations,
changes with a modular logics [5, 18, 34, 36];
The field of epigenetics enlarges the range of the sources of variation and
inheritance [24];
The phenotypic and developmental plasticity modifies the relationships between
genomes, phenotypes and ecological niches [51, 62];
The niche construction hypothesis [40] is a new, constructivist, way to see the
active role of the organism in evolution and the reciprocal modifications of
organisms and niches; this idea fits with the Developmental Systems Theory
[42, 43];
The concept of evolvability is evolving itself [27, 52];
The generation of order and structural complexity through mechanisms of
biological self-organization is real and deserves attention [25].
For each of these lines of researches we need more data and a careful consideration
of the real theoretical impact. Anyway, the capacity of painless assimilation of scientific
novelties by MS seems to be progressively declining. The problem is no longer of
partial incompleteness, but the adequacy of the whole conceptual structure of the
theory [20]. Maybe we need a new kind of Neo-Darwinism, revised and extended.
Using the methodology of Lakatosian SRP, we argue here that the transition in progress
from the MS to the so called Evolutionary Extended Synthesis (ES) [55] could be
represented as a shift from a previous evolutionary research programme (ERP1), turned
to be regressive, and a new evolutionary research programme (ERP2), with an
extended Neo-Darwinian core and a protective belt of new assumptions and auxiliary
hypotheses with a pluralistic and integrative explanatory approach.
3 Core and Protective Belt of a Scientific Research Programme
According to Imre Lakatos, the growth of the scientific knowledge is a process quite
independent from the mind of a single scientist: the product of the discovery
becomes autonomous from the mental activity that created it, like a growing living
218 T. Pievani
organism with proper laws of development. So the history of science could have
internal rational reconstructions, with an acceptable degree of objectivity, a logical,
continuous, internal dynamics of change through criticisms, new assumptions,
extensions and revisions. As we have seen, the term theory appears inadequate
to depict the current state of the evolutionary programme of researches in different
fields and with several integrated patterns of explanations (better than separated
concepts, like in [32]). How does a corpus of explanations with these
characteristics evolve and transmute?
In Lakatos the continuity of the history of science is given by the transformations
of scientific research programmes, that are sets of models, concepts and hypotheses
delimited by the choices of a community of scientists: firstly, they establish a
complex of theoretical postulates and corroborated explanations which are no
more subjected, in the practice of research and publications, to falsification. This
operative and methodological choice gives birth to the core of the RP. Of course,
the core is un-falsifiable exclusively from a methodological point of view, not in
principle: the scientists decide (according to an inductive conjectural principle)
that a core of explanations is confidently corroborated and reliable, and go ahead.
The choice, in this sophisticated fallibilism, is pragmatic, provisional and risky,
and any rival RP has the duty to hit and weaken the core. The inductive reliability of
the core has two criteria: the additional empirical content (predictions of new facts);
the corroboration of the additional empirical content (the predictions indeed
increase the empirical content of the RP).
Though the corroboration will never be ultimate in science, the pragmatic
preference for theories scientifically healthy is rational because they are conjectur-
ally reliable. The core contains also the influent metaphysics of the RP, driving the
researches through the great iconographies and metaphors of the RP, the philosoph-
ical views, the initial assumptions. The core produces the heuristic rules that
indicate the ways to be taken (positive heuristic) and the ways to be avoided
(negative heuristic). These aspects of the core reveal a kuhnian mood and a
defensive tenacity, but the structure of the core can evolve continuously, depending
on the anomalies emerging and on the fight with rival RPs, nevertheless always
keeping its physiognomy and peculiarities. So the defensive tenacity is not dog-
matic and blind, but articulated in a continuous requalification of the core through
the heuristic.
The followers of a RP cannot insert everything in the core, because its formula-
tion should respect a historical criterion (a long process of proofs and refutations,
trials and errors [30]) that is, a series of successive, refutable models, able to
predict new facts and a normative criterion, according to which the responsibility
of the choice is in charge with the scientists of the RP, and the success of the RP will
be evaluated by its empirical results. Then, with their creativity, scientists surround
the core with a protective belt, made by auxiliary hypotheses that protect the core
and nourish its evolution. The protective belt is made also by observational theories,
initial conditions, open problems, provisional hypotheses, other conjectures, and
must tolerate the external attacks by rival RPs thanks to continuous modifications,
adaptations, refutations and auxiliary hypotheses able to absorb anomalies and
predict new facts. The crucial difference is that the protective belt is by definition
fallible, unceasingly mutable, and its specific contents are not vital for the survival
of the whole RP.
In its early phases, the protective belt is composed specifically [38, 39] by
assumptions of negligibility (neutralization of facts considered non essential for
the study of the phenomena explained by the RP), assumptions of domain (the
external application of the RP to other kinds of phenomena, keeping the RP under
control) and heuristic assumptions (defining the problems that the RP does not
intent to deal with, the anomalies considered non essential, the risky, provisional
ad-hoc hypotheses). If the heuristic is fruitful and acts on the protective belt
modifying the hypotheses in a way that powers the RP, if it makes it more realistic
with respect to new observations, and increases the empirical content of the set of
explanations, the whole RP is defined as progressive. If the RP loses empirical
content, accumulates anomalies and ad-hoc hypotheses, needs continuous
emendations, the heuristic is in crisis and the RP is defined as regressive. Such
a regressive RP will be sooner or later substituted by a rival RP, with a different
explanatory core, able to cover all the empirical content of the previous RP. The
ultimate refusal of a RP, like its initial foundation, is a free, operative, fallible
choice of the scientists. By principle, any attempt of extreme defense of a RP is
legitimate, as long as it respects the criteria of a critical rationalism and the
adherence to evidences, like a nucleus of standards of evaluation adopted by the
scientific community.
In front of an anomaly, the scientist pins the blame initially on a marginal part of
the protective belt (or on some initial conditions, parameters, auxiliary
assumptions), protecting the core as long as possible (like in the case of the
Darwinian ad-hoc hypothesis concerning the imperfection of the geological record,
in this way protecting the gradualist assumption involved in the core-mechanism of
natural selection). This defensive strategy could be successful: the anomaly is
solved, the heuristic is reliable, the empirical content grows. Or provisionally
successful: the patch apparently works, but it is weakening the protective belt. Or
clearly unsuccessful, showing a pseudo-scientific attachment to non refutable and
useless ad hoc hypotheses.
Avoiding both a strict and a-historical fallibilism and by the way any methodo-
logical anarchism or external subjectivism or radical change of paradigms, the
methodology of RP provides a possible rational criterion for understanding
the continuous growth of knowledge in a field of researches, in order to compare
the explanatory power of rival RPs, and evaluate the state of health of a long time
dominant RP as well. The shifts of RPs could be rationally evaluated: if the
modification of one or more auxiliary hypotheses with the aim to restore
the matching between the core and reluctant empirical evidences has turned out
to be a new version of the RP, supported by new facts, by an additional corroborated
empirical content and by more successful predictions, then the shift of the RP has
been progressive (that is, we have had a growth of knowledge).
On the contrary, if any modification of a previous version of the RP has only the
aim of saving the phenomena and generates a new theoretical system not
220 T. Pievani
supported by more independent evidences, then the shift is regressive and the RP
is facing a crisis. When a RP accumulates anomalies and is no longer able to
accommodate itself in a succession of models and theories that increase the
empirical content, the scientists should decide to abandon it and move to an
alternative frame, shifting the RS toward a new version (with a different theoretical
structure, even if compatible with the previous core) or choosing a rival RP already
available in the field. This decision is itself risky and subjected to falsification,
depending on the experimental results of the new RP.
4 The Progressive Shift Between MS (ERP1) and ES (ERP2)
What is the case of MS and ES in such dynamics? Using the methodology of RP, we
could say that:
The core of the Evolutionary Modern Synthesis seems to have acquired today a
defined dimension, as long as we consider its boundaries not too much immuta-
ble and impervious to change [47, 49];
There is no alternative approach able to catch the continuous dynamics of
change inside the evolutionary research programme, its meaning and its internal
debates;
There is no evidence of a possible substitution of a dominant RP with a rival one,
with a radical refusal of a whole previous tradition of researches, but there is in
the field the typical scenario of a nonstop succession of shifts, extensions and
revisions of the contemporary Neo-Darwinian RP.
Furthermore, we know that MS has never been a monolith, but a meeting of
languages, disciplines and tradition of researches: at least the tradition of popula-
tion geneticists and that of naturalists, with respectively micro-evolutionary and
macro-evolutionary perspectives. Nevertheless, MS tended to crystallize its con-
sensus around some rigid methodological principles, intended as operationally no
more under falsification (as it is usual in any RP according to Lakatos), like phyletic
gradualism and genetic extrapolationism, defending a version of the RP based on a
gene-centered and pan-selectionist view [12]. The contemporary followers of this
ultra-Darwinian interpretation of the RP, with a great success in popularization,
have even more oversimplified the core in a rhetorical fashion, speaking about an
alleged universal Darwinian algorithm based on the research and development
activities of the supreme engineer, the natural selection. But it is interesting to note
the gap between this standard and popular view of evolution and the diversified
reality of the experimental field, monitored in the major International publications.
If we want a realistic epistemological representation of what is going on in the
field, we should recognize a received view of the RP, in these terms:
The core of the RP named MS is the Neo-Darwinism as the genetic theory of
natural selection;
The protective belt is made by three major methodological postulates: phyletic

gradualism; extrapolationism from microevolution; and strict functionalism or
adaptationism; so we have a prevalence of assumptions of negligibility (strong in
the cases of embryology, ecology and paleontology) and heuristic assumptions
(like in the case of the underestimation of structural internal factors in
evolution);
The deriving positive and negative heuristic is based on the programmatic idea
of a universal, basic Darwinian logics, able for decades to gather a huge amount
of new facts and prediction, with more and more sophisticated mathematical
models.
The MS so conceived shows also weak assumptions of domain in the belt (the
application of the theory is supposed to be large, and widely extra-biological): the
most important extensions of evolutionary models outside the strictly biological
territory come from the application of this oversimplified version of the RP (like in
the case of the strongly adaptationist approach in early evolutionary psychology
[4, 11]). But the claims of extended applicability clash with the growing difficulty
to absorb in this standard frame the reluctant lines of researches above mentioned.
So, the structure of the forthcoming theory of evolution could be represented as a
profound and substantial reformation of this RP, without the substitution with
another, but with a progressive shift. In other words, we have at the same time a
serious conceptual novelty (the global structure of the RP changes because of
discoveries not predictable by the previous programme) and a compatibility in
the core with the previous RP [54, 60]. It is neither a superficial maquillage on
marginal points of a hardened structure, nor a radical break with complete substitu-
tion of RP: rather, a steady and irreversible transformation of the architecture of the
previous RP. Particularly, the lines of the reformation seem to be two:
An extension of the central Neo-Darwinian core, through theoretical and
experimental updating, with in some cases the unexpected reversion to some
pluralistic insights of Charles Darwin himself;
A quite complete substitution of the protective belt, from methodological mono-
factorial postulates to multi-factorial postulates and integrations of patterns.
This transition has progressive features and could represent the passage from
MS (the Evolutionary Research Programme 1 ERP1) to ES (the Evolutionary
Research Programme 2 ERP2).
4.1 An Extended Neo-Darwinian Core
In this epistemological hypothesis, we see in detail that the core of the ES is today
an extended Darwinism, with two interacting hierarchies of nested levels: a
genealogical hierarchy of nested levels of transmission of genetic materials
(organisms, populations like demes, species, monophyletic taxa); an ecological
222 T. Pievani
hierarchy of nested levels of transfers of matter and energy (organisms, populations

like avatars, local ecosystems, regional ecosystems) [14]. The extended core
includes:
(A) The descriptive domain of evolution intended as factual evidence, that is the
common descent with modifications, the postulate of the continuity of any
evolutionary change, the phylogenetic unit of all living beings, the role of
organisms as basic units of evolution.
(B) The explanatory domain of plural, integrated patterns of evolutionary change
(we use the term patterns instead of models, because the semantic view, with
its emphasis on families of models as formal description of scientific
theorizing, is clearly inadequate for the evolutionary RP, where models are
useful tools in some fields but not the primary entities of the structure of the
theory itself [10, 17]); defining patterns the repeated schemes (epistemo-
logical and ontological) of historical real events, emerging from scientific
observations, sufficiently general to be considered as law-like regularities
in evolution [13], we see specifically:
(B1) Variational patterns: multiple sources of non directed, inheritable varia-
tion, genetic and epigenetic, included the possible inheritance of ecolog-
ical niches; the material basis for evolution and for natural selection is
going to be diversified;
(B2) Selective patterns: Darwinian natural selection and sexual selection,
artificial selection, kin selection, in some cases group selection [58],
briefly multilevel selection [41]; included here all the trade-offs between
these selective processes, and peculiar competitive patterns like meiotic
drive and spermatic competitions;
(B3) Neutralist patterns: the non adaptive domain of the extended core, with
three main sub-patterns: genetic drifts; nearly-neutralist mechanisms in
the genomes, re-assortments and systemic mutations; non selective struc-
tural effects due to genetic constraints (genetic webs, limits to variation),
developmental constraints (Evo-Devo), and physical-chemical constraints
(self-organization);
(B4) Macro-evolutionary patterns: the ecological side of the evolutionary
hierarchy, with all the external, influential macro-evolutionary factors,
able to produce repeated schemes of events like, in a decreasing order of
impact on biological populations: mass extinctions; turnover pulses of
species; rapid adaptive radiations; major evolutionary transitions due to
processes of symbio-genesis [13, 49].
According to this kind of pluralistic core in ERP2, the evolutionary explanation

becomes a way to integrate, case by case, different patterns, irreducible one to
another, but compatible one with another, and all needed to cover the heteroge-
neous empirical basis of evolutionary phenomena. The ecological hierarchy
perturbs at different degrees the genealogical one, and the genealogical one
accumulates changes and produces diversity. This multiple core (and the super-
pattern of the relationships between the ecological and the genealogical levels of
evolution) is a candidate framework for the theoretical unification of evolutionary
biology [14]. The first patterns (excluded epigenetic inheritance in modern sense),
the second ones (excluded kin selection) and the structural ones are already
Darwinian in their first formulations. The other ones are not originally Darwinian
but compatible and integrative. This is an integrated framework where genealogical
levels and ecological levels coexist and interact, but the Darwinian unit, the
organism (at the same time, genetic replicator and ecologic interactor), and the
Darwinian mechanism, natural selection (as differential survival and reproduction),
remain the fundamental junction of the two hierarchies of levels of change (see the
sloshing bucket model of evolution, in [14]).
In this epistemological frame, we clearly understand the dynamics of assimila-
tion and accommodation, and the mix of points of breaking and point of continuity,
in the theoretical challenges above mentioned of Punctuated Equilibria and Neu-
tralism. As restriction of validity, these patterns of the core of ERP2 keep their
explanatory power in natural history until ca. 600 million years ago, that is the
explosive beginnings of organized and individualized Metazoans. For earlier eras,
the patterns could be intended as corroborated explanatory hypotheses, waiting for
more information about the environmental conditions and the kinds of organisms
involved.
This theoretical nucleus represents also an interesting way to depict the unique-
ness of the nature of explanation in evolutionary fields. The probabilistic
generalizations and correlations of evolutionary biology have causal explanations
that do not depend exclusively on the context of the scientific questions (like in
some recent forms of relativistic contextualism), but reveal a deeper ecological-
genealogical structure founded on the complementarity of three kinds of
explanations. The explanandum of the theory of evolution is variational (the
diversity of species and variants on Earth, their genealogical relationships, their
transformations) and adaptive (any trait contributing to fitness, the complexity of
structures and behaviors). The architecture of types of explanations is built on three
pillars: (1) functional and selective explanations of the remote causes of emergence
of a trait; (2) structural explanations of the relevance of internal constraints, and
their trade-offs with external pressures; (3) historical and contingent explanations
of chains of causal singular events, and clashes of independent chains of causes,
a priori unpredictable but intelligible a posteriori. Starting from these three
typologies, we can construct a matrix of relationships between the kinds of
explanations: 1 + 2 in the cases of exaptations and trade-off between functions
and structures, analogies and homologies; 1 + 3 in the cases of interferences
between natural selection and Neutralistic processes and macro-evolutionary pro-
cesses; 2 + 3 when Neutralistic processes create structural constraints; 1 + 2 + 3
when we need total evidence reconstructions of evolutionary pathways.
224 T. Pievani
4.2 A Pluralistic Protective Belt
Secondly, the protective belt of the RP becomes more and more pluralistic and
flexible, with a robust diversification of models concerning the rhythms, the units,
the levels and modes of evolution.
Particularly:
A plurality of rates of evolution and speciation: gradual trends, stasis,
punctuations, plural modes of speciation and changing relative frequencies [13];
A plurality of units of evolution: group selection; multilevel selection; niche
construction, endo-symbiosis and units organism-niche [41]; where the selection
acts (genomes, organisms, developmental systems, organisms plus niches) and
how (trade-offs between competition and cooperation);
A plurality of adaptive dynamics and interactions between structures and
functions: frequency of exaptations in natural history; Evo-Devo; modularity;
A complexity of relationships between levels of evolution: phenotypic plasticity;
evolvability.
In the methodology of RP, the protective belt is subjected to continuous
modifications due to the updating of evidences in surrounding conditions. The
equilibrium between the weights of explanatory power in different patterns is
instable. But what is crucial is that a refutation in the belt, or a radical change in
the perspective of specific debate, does not mean necessarily a problem for the
structure of the core. The belt is the magmatic, ever-changing, constantly refutable
zone of the RP, exposed to external influences and contingencies of the researches.
It could be also possible that an auxiliary pattern in the belt becomes so
important and so diffuse in current evolutionary explanations that it deserves to
be part of the core itself. In absence of strong contrary evidences, and having a good
theoretical stability, it becomes part of the core, like in the case of genetic drifts and
Neutralistic phenomena. So we have an extension of the core, always submitted to
the falsification logics: if you have more patterns in the nucleus, a rival RP has more
possibilities to reach a refutation of some assumptions or to outline a contradiction
between explanations. For that reason, a RP should be not too much minimal and
not too much extended, preserving its structural coherence and avoiding inappro-
priate extensions of the explanandum.
In the passage between ERP1 and ERP2 the heuristic has changed its sign, being
driven now by a programmatic pluralism, in definitions (like species and gene)
and in explanations (with frequencies of integrated patterns), able to predict a lot of
new facts in several evolutionary disciplines. We observe in literature that the new
heuristic satisfies the criteria of progressivity. The assumptions of negligibility
become less influent and the heuristic assumptions highlight the necessity of the
integrations of factors. Conversely, the assumptions of domain and applicability
become more stringent, stressing the opportunity to tune possibly other RPs for
specific adjacent fields (like the evolution of culture [6, 56]).
5 The Multiple Advantages of the Current Neo-Darwinian

Pluralism
The change of sign in the heuristic of ERP2 carries a more general epistemological
transition. The standards of evaluation of hypotheses and theories are changing: the
proofs through convergence and consilience of data become more and more
important; we see the birth of originally interdisciplinary lines of researches;
functions, structures and singular histories, mixed together, recover the original
pluralism of the evolutionary explanations.
A new theoretical challenge (as it was the Punctuated Equilibria theory, with its
early revolutionary phase and later considerations and pondering about its real
theoretical impact) is not necessarily concerning a marginal epiphenomenon
surrounding a monolithic paradigm or on the contrary a radical crisis of its core.
Otherwise, it could mean a profound extension and revision of its structure,
remaining nevertheless compatible with other components and patterns of it [59].
The result is a structure of the theory of evolution, intended as a RP, more articulated
in a pluralistic frame, more realistic in its assumptions about the currently available
evidences [19, 50], with drastic revision of previous restrictive concept (regarding
the universality of some patterns) hardened in the protective belt of ERP1 (or
MS).
The same case could be traced in the history of the idea of functional
cooptation in Darwin, then pre-adaptation in Ernst Mayr and MS, then in the
more radical sense of exaptation [22, 61] and spandrels by structural non-
aptations [21]. It is clear in current literature that we do not need a conflation
between standard adaptations and exaptations, but an extended taxonomy of
fitness [20] made by three typologies of processes: classical Darwinian adaptations
by natural selection; the functional shift, by natural selection, from a previous
function to a secondary one; spandrels and other side effects with no adaptive
reasons in their beginning, possibly co-opted by natural selection in new external
conditions [46]. Also in this case we have points of breaking with ERP1 (not always
the current utility corresponds to the historical origin) and points of continuity
(natural selection acts, but finding trade-offs with internal constraints of organisms).
So the advantage is double. Firstly, through these epistemological processes of
assimilation and accommodation, we can better appreciate the conceptual tools
involved in the most promising lines of evolutionary research today: for an excel-
lent case, mixing genes, developments and ecological conditions, and considering
all the patterns of evolution (mutations, natural selection, drifts, migration,
hybridization, speciation), see [23]. Secondly, we can understand why one of the
most important scientists involved in the developmental evolutionary biology,
Alessandro Minelli, being conscious of the huge theoretical impact of Evo-Devo
on ERP1 in terms of conservation of genes families, developmental constraints,
limits to variation, modularity of change, multiple effects, evolvability neverthe-
less wrote that clearly Evo-Devo does not offer a significant challenge to the Neo-
Darwinian paradigm [35]. Evo-Devo likewise extends the conceptual frame of
226 T. Pievani
Neo-Darwinism and rectifies some previous interpretations (like gene-centrism and

adaptationism) in its surrounding assumptions. The structure of ERP is evolving
towards a more pluralistic version [1].
Furthermore, not less important, this Neo-Lakatosian approach offers some
precious advantages in public debates (frequently hot) concerning evolution and
its debates. The fact that in some points Darwin was wrong, of course, and MS was
inadequate loses a great part of its striking dramatic power. Any RP is an evolving
system, made by patterns, concepts, models, hypotheses, structured in a core (what
is crucial for the survival of the RP) and a protective belt (where nothing is crucial
for the fate of the RP). So a single refutation and an internal contradiction are not
necessarily the end of the world for the theory of evolution, and we need to see the
whole architecture of the RP and how it is rationally changing. A RP is never
falsified by a single observation or anomaly, it could be defeated by a set of new
evidences, new theories, organized in a rival RP [30].
The dynamics of criticism and growth of knowledge is continuous and rational,
with a verifiable internal logics of proofs and refutations, so all the debate around
the alleged dogmatism of the Darwinians appears like a non sense [48]. Every
aspect of the evolution of ERP is under control and under potential falsification,
even if some parts of the core (the descriptive domain and the major patterns of the
explanatory domain, above described) are so much corroborated that they appear
just like plain evidences without any reasonable doubt [7]. For the same reasons, we
can solve another debate concerning the alleged scenarios beyond Darwin, or the
supposed ultimate crisis of Neo-Darwinism: the challenges (frequently interesting
by themselves) evocated as proofs of the imminent death of the theory are quite
always debates concerning the protective belt of the RP. So, nothing ultimate, and
moreover criticisms without a rival RP.
But mostly, the methodology here exposed is very useful in the definition of
what is the burden of proof in these debates. If someone wants to demonstrate that
another theory of evolution is on the field (like in all the cases in which someone
says that now there would be more theories of evolution), he has to prove that:
(1) there is a rival ERP able to explain all the empirical basis of the current one; (2)
this ERP is able to explain something more (it has an additional empirical content,
predictions of new facts, on the basis of other corroborated facts); and it is able to do
1 and 2 using patterns, concepts and principles not reducible to those of the current
one. It is a scheme of rational evaluation (and sophisticated falsificationism) clear
and very fruitful for comparisons between ERPs. Only if we see these three
conditions we can say that a rival RP has defeated a previously dominant one.
We should grant anyone the benefit of doubt, even if the challenge seems to have
nothing to do with science, but of course there is nothing like that rival RP in the
field at the moment.
These and other cases show how the application of the methodology of ERP
could not only represent the ongoing transition between MS and ES very well, but
could also offer valid arguments and epistemological tools of rational analysis in
the never-ending debates that enliven, inside and outside the scientific community,
the developments of the Neo-Darwinian research programme.
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The Theory of Evolution and Its Impact

The Theory of Evolution

ISBN 978-88-470-1973-7 e-ISBN 978-88-470-1974-4

# Springer-Verlag Italia 2012

Cover illustration: Nanni Valentini, Cratere, 1978-80. Private collection.

Printed on acid-free paper

Springer is part of Springer ScienceBusiness Media (www.springer.com)

Introduction: The Sand Walk (on the Darwins Steps) . . . . . . . . . . . . . . . . . . . . . 1

Idola Tribus: Lamarck, Politics and Religion in the Early

Darwinism Past and Present: Is It Past Its Sell-by Date? . . . . . . . . . . . . . . . 41

Evolutionary Theory and Philosophical Darwinism . . . . . . . . . . . . . . . . . . . . . . . . 53

Struggle for Existence: Selection, Retention and Extinction

The Theory of Evolution and Cultural Anthropology . . . . . . . . . . . . . . . . . . . . . 83

The Concept of Evolution in Linguistics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103

Theory of Evolution and Genetics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119

Genes, Evolution and the Development of the Embryo . . . . . . . . . . . . . . . . . . . 131

Evolutionary Mechanisms and Neural Adaptation: Selective Versus

Is the Human Brain Unique? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 175

Evolution: Remarks on the History of a Concept

An Evolving Research Programme: The Structure of Evolutionary

Abstract To understand the status of Theory of Evolution, highly multidisciplinary

A. Fasolo (ed.), The Theory of Evolution and Its Impact, 1

This is even more relevant, if we step from biological sciences to humanities.

1 Roots and Buds of Evolutionary Theory

In history of science, for instance, notwithstanding a few crucial contributions, the

2 The Mankind Affair

A key-corner between biology and society is the language. Manfred Bierwisch

3 Development, a Persistent Problem of Evolutionary Theory

4 Brain Evolution and Plasticity

Overcoming the traditional dichotomy opposing neural selectionism to construc-

of the nervous system requires complex neural networks bearing precise

6 Cognition and Reasoning

Moving from comparative neuroanatomy to modern cognitive neuroscience,

A. Fasolo (ed.), The Theory of Evolution and Its Impact, 11

(and later) debates on natural selection: many saw it as a plausible mechanism to

He had no chance to be listened to in a world moving towards disciplinary

1 Lamarck Versus Institutional Science

consequence, France was rather poor in periodical publications, since only a

2 The Politics and Religion of Science

3 Authors and Audiences

Historians of science have traditionally shown a remarkable reluctance to accept the

Meckel published in German an essay, Fragments sur lhistoire du

as to the French one. It was the entrepreneurial genius of Jean-Baptiste Marie

1. Adelon NP (1821) Systeme analytique des connaissances positives de lhomme. Revue

29. Duvernoy GL (1849) Ovologie. Dictionnaire universel dhistoire naturelle 9:281353

A. Fasolo (ed.), The Theory of Evolution and Its Impact, 41

2 The Consilience of Inductions

Clearly influenced by the rationalist strain in philosophy (Whewell was much

In the case of instinct we get the explanation through selection of a beautiful

Embryology, as we draw to a close, was a particular point of triumph for Darwin,

4 From Then to Now

5 The Consilience Today

1 Philosophical Darwinism: A Short History

Four decades of controversy concerning evolution had elapsed, and Darwins

A. Fasolo (ed.), The Theory of Evolution and Its Impact, 53

evolutionary logic, according to Deweys subtle analysis, expelled from biology,

2 Spencer and Darwin on Evolution

These early surveys of the historical-philosophical aspects of the evolution theory

historian of anthropology, emphasized Spencers priority and originality, at least as

In spite of this afterthought, Spencer remained a staunch follower of Lamarck.

rubbish on the nature of reproduction [20]. When Spencers First Principles

Spencer s semantics is misleading: what he meant in speaking of natural

We may conclude that the long-debated problem of reciprocal influence is, in a

Furthermore, Darwin first introduced Spencers favourite term evolution in the