Table Of ContentTowards a Theory of Development
Towards a Theory
of Development
EDITED BY
Alessandro Minelli
Department of Biology, University of Padova, Padova, Italy
Thomas Pradeu
Department of Philosophy, Paris-Sorbonne University, Paris, France
1
Towards a Theory of Development. Edited by Alessandro Minelli and Thomas Pradeu
© Oxford University Press 2014. Published 2014 by Oxford University Press.
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List of contributors
Wallace Arthur School of Natural Sciences, Armin P. Moczek Department of Biology, Indiana
National University of Ireland, Galway, Ireland University, Bloomington, Indiana, USA and
Jonathan Bard Department of Physiology, National Center for Evolutionary Synthesis,
Anatomy, and Genetics, University of Oxford, UK Durham, North Carolina, USA
Roberto Carrer Department of Mathematics, Michel Morange Centre Cavaillès, Ecole Normale
V. Volterra Institute, San Donà di Piave, Italy Supérieure, Paris, France
Giuseppe Fusco Department of Biology, University Stuart A. Newman Department of Cell Biology and
of Padova, Italy Anatomy, New York Medical College, Valhalla,
NY, USA
Scott F. Gilbert Department of Biology,
Swarthmore College, Swarthmore, PA, USA and Spencer V. Nyholm Department of Molecular and
Biotechnology Institute, University of Helsinki, Cell Biology, University of Connecticut, Storrs,
Helsinki, Finland CT, USA
James Griesemer Department of Philosophy, Thomas Pradeu Department of Philosophy,
University of California-Davis, Davis, CA, USA Paris-Sorbonne University, Paris, France
Isaac Hernández Department of Philosophy, Emanuele Serrelli Riccardo Massa Department of
University of Toulouse 2, France Human Sciences, University of Milano Bicocca,
Milano, Italy
Johannes Jaeger Centre de Regulació Genòmica,
Barcelona, Spain James Sharpe Centre de Regulació Genòmica, and
Institució Catalana de Recerca i Estudis Avan-
Jean-Jacques Kupiec Centre Cavaillès, Ecole
çats, Barcelona, Spain
Normale Supérieure, Paris, France
Davide Vecchi Department of Philosophy, Uni-
Lucie Laplane Department of Philosophy,
versidad de Santiago de Chile, Santiago, Chile
University of Paris X, Nanterre, Paris, France
and Instituto de Filosofía y Ciencias de la
Alan C. Love Department of Philosophy,
Complejidad, Santiago, Chile
Minnesota Center for Philosophy of Science,
Michel Vervoort Institut Jacques Monod, Centre
University of Minnesota, Minneapolis, MN, USA
National de la Recherche Scientifique, Université
Margaret J. McFall-Ngai Department of Medical
Paris Diderot, Sorbonne Paris Cité, Paris, France
Microbiology and Immunology, University of
and Institut Universitaire de France, Paris,
Wisconsin-Madison, Madison, WI, USA
France
Alessandro Minelli Department of Biology,
University of Padova, Padova, Italy
vii
Foreword: a biologist’s view
Laws to understand and explain embryonic devel- of development with the intention of moving to-
opment have been in existence for two centuries or wards ‘a theory of development’.
more. These laws, which are based on comparative The ‘theory of development’ proposed by Lud-
embryology, resulted in conclusions of sufficient wig von Bertalanffy in the late 1920s is most well
generalization that they could be termed a law. known from the English translation of his 1928
von Baer proposed the law that general features book Kritische Theorie der Formbildung, published in
of animals appear in development before more 1933 under the English title Modern Theories of De-
specific features and concluded that general fea- velopment: An Introduction to Theoretical Biology. The
tures were those of phyla and specific features, link to theoretical biology shows that Bertalanffy
those of species. Such a ‘law’ does not propose a posited that a theory of development would point
mechanistic explanation for development and, in the way towards a theory of life. As J. Z. Young
von Baer’s case, not even an evolutionary expla- commented in his back cover blurb: ‘This book con-
nation. Phylogenetic analyses support von Baer’s tains a discussion of the nature of living organisms
generalizations but do not provide an explanation and the methods by which they should be studied.’
for them. Recent identification of shared Hox genes, Conrad Hal Waddington’s four-volume Towards
phylotypic stages, and conserved gene pathways a Theoretical Biology (1968–1972) makes the same
identify shared mechanisms but do not provide a claim as do a number of the authors in the present
theory of development. book, although theoretical biology has a much wid-
Haeckel’s Biogenetic Law that ontogeny reca- er (and potentially more diffuse) ambit now than it
pitulates phylogeny proposes that embryos pro- had 80 or even 40 years ago.
vide the way to understand evolution through the von Bertalanffy used embryology to illustrate
mechanism of recapitulation. But the Biogenetic the approaches required for a theory of biology.
Law is not a theory of development, even though He sought to counter the strongly mechanistic ap-
it too is based on comparative studies leading to proach to biology that dominated research and
a generalization. If anything, it is a law of evolu- interpretation by stressing organization, organi-
tion. Similarly many of the explanatory terms used cism, and a systems approach; von Bertalanffy is
in development— regulative development, pre- acknowledged as the founder of general systems
formation, programmes, fields, epigenetic land- theory. It is no coincidence that his German text
scape, genotype-to-phenotype map, increasing was translated and adapted by the biologist turned
complexity—are metaphors and neither laws nor logician-philosopher and anti-mechanist, Joseph
theories. A theory is a statement of a principle(s) Woodger, whom von Bertalanffy had met in Vienna
derived from and able to explain facts or events in 1926 when both came under the influence of the
and proposes a mechanism(s) that explains the Vienna Circle. For both men, biology would never
facts or events. Laws and metaphors describe be physics, and so no general theory would unite
but do not explain. The 17 chapters in the present the sciences. Woodger was a founder and major
ambitious book set out to evaluate past theories participant in the Theoretical Biology Club, which
ix
x FOREWORD: A BIOLOGIST’S VIEW
first met in 1932 in the grounds of Woodger’s home explaining one aspect of development or as sign-
in Epsom Downs.1 posts to a more general theory.
Any theory is embedded in and arises from ap- The discussions in various chapters that ex-
proaches in vogue at the time. von Bertalanffy’s plicitly or implicitly acknowledge that we have
book was reissued in 1962 with a new Foreword theories of evolution and of heredity but not a
(von Bertalanffy died in 1972) but with no changes theory of development should make us pause
to the text of the 1933 edition. Anti-mechanism, and reflect. The diversity of approaches in Towards
anti-reductionism, organicism, and the develop- a Theory of Development leads one to ask wheth-
ment of theoretical approaches to biological organi- er ‘a’ theory of development is attainable and/
zation, such as cybernetics, information theory, and or whether such a theory should be ‘A theory
general systems theory, remained sufficient for von of ontogeny’ (as discussed in Chapters 1 and 2)
Bertalanffy’s theory of development. or incorporated into ‘A theory of life?’ Given the
Although approaches from theoretical biology multitude of processes and mechanisms operating
remain and are discussed in the present book, during development/ontogeny and the multiple of
times have changed. As well represented in this scales of the biological hierarchy that emerge dur-
volume, today’s theories have different bases, ing development—genes; cells; germ layers; tissues;
reflecting increased knowledge of (and perhaps organs; epigenetic, functional, environmental, and
even understanding of) the processes of devel- symbiotic interactions—it may be a good thing that
opment. For the past half-century, approaches to we do not have a theory of development. That said,
a theory of development have been gene based. Towards a Theory of Development is a great place to
The three editions of the seminal book Gene Ac- find answers to such questions as whether a theory
tivity in Early Development by Eric Davidson pub- of development will:
lished in 1968, 1976, and 1986 have been replaced
(cid:114)(cid:1)(cid:66)(cid:68)(cid:68)(cid:80)(cid:86)(cid:79)(cid:85)(cid:1)(cid:71)(cid:80)(cid:83)(cid:1)(cid:88)(cid:73)(cid:66)(cid:85)(cid:1)(cid:88)(cid:70)(cid:1)(cid:79)(cid:80)(cid:88)(cid:1)(cid:76)(cid:79)(cid:80)(cid:88)(cid:1)(cid:85)(cid:80)(cid:1)(cid:67)(cid:70)(cid:1)(cid:85)(cid:73)(cid:70)(cid:1)(cid:69)(cid:74)(cid:84)(cid:85)(cid:83)(cid:74)(cid:67)-
by Genomic Regulatory Systems: Development and
uted nature of causation including from the ge-
Evolution (2001), reflecting the transition from
nome and environment, and between cells?
genes, to gene cascades/networks, to gene regu-
(cid:114)(cid:1)(cid:74)(cid:79)(cid:68)(cid:80)(cid:83)(cid:81)(cid:80)(cid:83)(cid:66)(cid:85)(cid:70)(cid:13)(cid:1)(cid:80)(cid:83)(cid:1)(cid:74)(cid:79)(cid:69)(cid:70)(cid:70)(cid:69)(cid:1)(cid:67)(cid:70)(cid:1)(cid:67)(cid:66)(cid:84)(cid:70)(cid:69)(cid:1)(cid:80)(cid:79)(cid:13)(cid:1)(cid:70)(cid:81)(cid:74)(cid:72)(cid:70)(cid:79)(cid:70)(cid:85)(cid:74)(cid:68)(cid:1)
latory networks. Gene-based theories need not
control of development?
necessarily conflict with systems-based theories
(cid:114)(cid:1)(cid:70)(cid:79)(cid:73)(cid:66)(cid:79)(cid:68)(cid:70)(cid:1)(cid:85)(cid:73)(cid:70)(cid:1)(cid:83)(cid:80)(cid:77)(cid:70)(cid:1)(cid:80)(cid:71)(cid:1)(cid:79)(cid:66)(cid:85)(cid:86)(cid:83)(cid:66)(cid:77)(cid:1)(cid:84)(cid:70)(cid:77)(cid:70)(cid:68)(cid:85)(cid:74)(cid:80)(cid:79)(cid:1)(cid:80)(cid:79)(cid:1)(cid:69)(cid:70)(cid:87)(cid:70)(cid:77)(cid:80)(cid:81)-
but often are cast as alternative explanations for
ment and speed synthesis of development and
development, reflecting alternate worldviews.
evolution?
Explanation and potential explication of these
(cid:114)(cid:1)(cid:73)(cid:66)(cid:87)(cid:70)(cid:1)(cid:85)(cid:80)(cid:1)(cid:85)(cid:66)(cid:76)(cid:70)(cid:1)(cid:84)(cid:85)(cid:80)(cid:68)(cid:66)(cid:84)(cid:85)(cid:74)(cid:68)(cid:74)(cid:85)(cid:90)(cid:1)(cid:80)(cid:71)(cid:1)(cid:69)(cid:70)(cid:87)(cid:70)(cid:77)(cid:80)(cid:81)(cid:78)(cid:70)(cid:79)(cid:85)(cid:1)(cid:74)(cid:79)(cid:85)(cid:80)(cid:1)(cid:66)(cid:68)-
tensions are an important aspect of the present
count?
volume.
(cid:114)(cid:1)(cid:66)(cid:68)(cid:68)(cid:80)(cid:86)(cid:79)(cid:85)(cid:1)(cid:71)(cid:80)(cid:83)(cid:1)(cid:68)(cid:80)(cid:79)(cid:87)(cid:70)(cid:83)(cid:72)(cid:70)(cid:79)(cid:85)(cid:1)(cid:70)(cid:87)(cid:80)(cid:77)(cid:86)(cid:85)(cid:74)(cid:80)(cid:79)(cid:32)
I can hear the murmurs through the ether: but
(cid:114)(cid:1)(cid:70)(cid:79)(cid:73)(cid:66)(cid:79)(cid:68)(cid:70)(cid:1)(cid:80)(cid:86)(cid:83)(cid:1)(cid:86)(cid:79)(cid:69)(cid:70)(cid:83)(cid:84)(cid:85)(cid:66)(cid:79)(cid:69)(cid:74)(cid:79)(cid:72)(cid:1)(cid:80)(cid:71)(cid:1)(cid:73)(cid:80)(cid:78)(cid:80)(cid:77)(cid:80)(cid:72)(cid:90)(cid:1)(cid:66)(cid:79)(cid:69)(cid:1)
surely any theory of development must be gene
variation?
based? Not according to theories discussed herein,
(cid:114)(cid:1)(cid:81)(cid:83)(cid:80)(cid:87)(cid:74)(cid:69)(cid:70)(cid:1)(cid:66)(cid:1)(cid:71)(cid:83)(cid:66)(cid:78)(cid:70)(cid:88)(cid:80)(cid:83)(cid:76)(cid:1)(cid:71)(cid:80)(cid:83)(cid:1)(cid:68)(cid:80)(cid:79)(cid:84)(cid:85)(cid:83)(cid:66)(cid:74)(cid:79)(cid:85)(cid:1)(cid:66)(cid:79)(cid:69)(cid:1)(cid:69)(cid:70)(cid:70)(cid:81)(cid:1)(cid:73)(cid:80)-
which include emphases on morphogenetic fields,
mology on the one hand and plasticity, adaptabil-
cell differentiation, natural selection, disparity,
ity and evolvability on the other? or
plasticity, stem cells, dynamical systems theory,
(cid:114)(cid:1)(cid:67)(cid:70)(cid:1)(cid:66)(cid:1)(cid:85)(cid:73)(cid:70)(cid:80)(cid:83)(cid:90)(cid:1)(cid:67)(cid:66)(cid:84)(cid:70)(cid:69)(cid:1)(cid:80)(cid:79)(cid:1)(cid:68)(cid:80)(cid:79)(cid:68)(cid:70)(cid:81)(cid:85)(cid:84)(cid:1)(cid:79)(cid:80)(cid:85)(cid:1)(cid:90)(cid:70)(cid:85)(cid:1)(cid:69)(cid:83)(cid:70)(cid:66)(cid:78)(cid:85)(cid:1)(cid:80)(cid:71)(cid:32)
and more. These theories are presented either as
Brian K. Hall
1 Present at that meeting were Joseph Woodger, Conrad Hal November, 2013
Waddington, Joseph Needham, Dorothy Needham, J. D. Bernal, Halifax
and Dorothy Wrinch. Except during World War II, the club met
annually until 1952.
