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evolutionary theory:

the view from altenberg

by Massimo Pigliucci

NATURE|Vol 455|18 September 2008


Woodstock? Austrias Konrad Lorenz Institute for

Evolution and Cognition Research is hosting a
much-discussed evolutionary biology meeting.

Beyond genes
Insights from ecology, developmental biology,
and genomics in particular are nudging evolutionary biology away from a focus on population geneticshow the distribution of genes
changes across groups of individualsand
toward an understanding of the molecular
underpinnings of these changes. Better family
trees that give researchers greater confidence
about the relatedness among organisms have
helped promote a credible, comparative
approach to these mechanisms, says invitee
Gnter Wagner, an evolutionary developmental biologist at Yale University.
Some studies, for example, indicate that
development constrains evolution. From the
modern synthesis perspective, Wagner
explains, the body plan is a historical residue
of evolutionary time, the afterglow of the evo-

Postmodern evolution?

h my gosh, says Massimo

Pigliucci, maybe I shouldnt
use that term. Pigliucci,
responding to comments on his
talk about how living things respond to their
environment, and what it means for evolution,
has just let slip the p-word. Later the same day,
Gnter Wagner, an evolutionary theorist at
Yale University in New Haven, Connecticut,
puts up a slide bearing the words Postmodern Synthesis. Pigliucci is moved to make an
editorial suggestion from the floor: Id really
rather we didnt use that term. Wagner says
the slide was intended to be tongue-in-cheek,
but Pigliucci is worried about the impression
the word creates: If theres one thing we dont
want, its for people to get the idea that theres a
bunch of evolutionary theories out there, and
that theyre all equal.
A lot of scientists loathe what they take to
be postmodernisms intellectual relativism,
and shy away from using the word. But doing
so puts Pigliucci in something of a bind. An
evolutionary ecologist at the State University
of New York in Stony Brook, Pigliucci is one
of the conveners of this small meeting on the
future of evolutionary thought taking place
at the Konrad Lorenz Institute for Evolution
and Cognition Research in Altenberg, Austria. The meeting has received a fair amount of
hype in the blogosphere it was dubbed The
Woodstock of Evolution. Its agenda is, pretty
explicitly, to go beyond the modern synthesis
that has held sway in evolutionary theory since

the middle of the twentieth century. And in Darwins ideas about natural selection and Greeveryday speech, it is pretty clear what comes gor Mendels insights into how traits pass from
after the modern.
parents to offspring which many biologists
Whats more, some of this work sounds of the time believed antithetical and fused
as though it fits the term quite nicely. Over them into a mathematical description of the
dinner at the meetings end, Pigliucci expresses genetic makeup of populations and how it
his hope of moving from a gene-centric view changes. That fusion was the modern synthesis.
of causality in evolution to a pluralist, multi- It treats an organisms form, or phenotype, as a
level causality. Postmodernists in the humani- readout of its hereditary information, or genoties call this decentering, and they are all for it. type. Change is explained as one version of a
Over the course of the meeting,
gene being replaced by another.
its fairly clear that the means
Natural selection acts by changThe modern
to this pluralist end are being
ing the frequency of genes in
synthesis is good
sought through mixing and
the next generation according
matching neglected ideas and at modelling the
to the fitness of phenotypes in
old problems from biologys
this one. In this world view, the
survival of the
gene is a black box, its relationpast with the latest experimenfittest,
tal and analytical techniques.
ship to phenotype is a one-way
Apply that sort of bricolage to arrival of the fittest. street, and the environment,
both cellular and external, is a
architecture and you get the
Scott Gilbert
selective filter imposed on the
sort of brutalist-right-angle
readout of the genes, rather
here, classical-column-there,
swirling-titanium-ceiling-above-it-all look than something that can influence an organthat is normally pigeonholed, for better or isms form directly.
worse, as postmodern.
Whats wrong with this picture, say the
would-be extenders at Altenberg and elsewhere, is what it leaves out. Molecular biolEvolution of ideas
Leaving aside the troublesome adjective, what ogy, cell biology and genomics have provided
is the modernism that the Altenburg meeting a much richer picture of how genotypes make
is meant to move beyond or to use Pigliuc- phenotypes. The extenders claim that enough
cis preferred term, extend1? Between about insights have now come from this and other
1920 and 1940, researchers such as the Ameri- research for it to be time to re-examine probcan Sewall Wright and the Englishmen Ron- lems that the modern synthesis doesnt address.
ald Fisher and J. B. S. Haldane took Charles These problems include some of the key turning


11 JULY 2008

VOL 321


Published by AAAS


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This summer a group of high-profile researchers met in

Altenberg, Austria, to try and plot the future course of
evolutionary theory. John Whitfield was there.


Massimo Pigliucci is no Jimi Hendrix. This passing genes from parent to offspring, as the
soft-spoken evolutionary biologist from Stony environment, chemical modification of DNA,
Brook University in New York state looks and other factors come into play as well. Organnothing like that radical hard-rock musician isms vary not only in how they adapt to changwhose dramatic guitar solos helped revolution- ing conditions but also in how they evolve.
ize rock nroll. But to Suzan Mazur, a veteran
Evolution is much more nuanced than the
journalist who occasionally covers science, founders of the modern synthesis fully appreciPigliucci is the headliner this week at a small ated, says Pigliucci. That doesnt mean that the
meeting she believes will be the equivalent of overall theory of evolution is wrong, as some
Woodstock for evolutionary biology. The invi- intelligent design proponents have tried to
tation-only conference, being held in assert using Mazurs story as support, but rather
Altenberg, Austria, promises to be far more that the modern synthesis needs to better incortransforming for the world than the 1969 porate modern science and the data revealed by
music festival, Mazur wrote online in March it. More than genes pass on information from
for, an independent
news publication in New Zealand.
That hyperbole has reverberated throughout the evolutionary
biology community, putting
Pigliucci and the 15 other participants at the forefront of a debate
over whether ideas about evolution need updating. The mere
mention of the Altenberg 16, as
Mazur dubbed the group, causes
some evolutionary biologists to
roll their eyes. Its a joke, says
Jerry Coyne of the University of Daring duo. Massimo Pigliucci (right) and Gerd Mller want to
Chicago in Illinois. I dont think update the modern synthesis.
theres anything that needs fixing. Mazurs attention, Pigliucci admits, one generation to the next, for example, and
frankly caused me embarrassment.
development seems to help shape evolutions
Yet Pigliucci and others argue that the so- course. Many things need fixing, emphasizes
called modern synthesis, which has guided evo- one invited speaker, Eva Jablonka of Tel Aviv
lutionary thought and research for about University in Israel. I think that a new evolu70 years, needs freshening up. A lot has hap- tionary synthesis is long overdue.
pened in the past half-century. DNAs structure
was revealed, genomes were sequenced, and Modern tradition
developmental biologists turned their sights on The modern synthesis essentially represents a
evolutionary questions. Researchers have come marriage of the 19th century concept of evoto realize that heredity is not simply a matter of lution with Mendelian genetics, which was

rediscovered at the beginning of the 20th century; the birth of population genetics in the
1920s added to the intellectual mix. By the
1940s, biologists had worked out a set of ideas
that put natural selection and adaptation at
evolutions core. Julian Huxleys 1942 book,
Evolution: The modern synthesis, brought
together this work for a broad audience.
Simply put, the modern synthesis holds
that organisms have a repertoire of traits that
are passed down through the generations.
Mutations in genes alter those traits bit by bit,
and if conditions are such that those alterations make an individual more fit, then the
altered trait becomes more common over
time. This process is called natural selection.
In some cases, the new feature can replace an
old one; in other instances, natural selection
also leads to speciation.
However, several concepts have arisen
since then that make the modern synthesis
seem too simplistic to some, Pigliucci
among them. In a 2007 Evolution paper, he
called for the development of an extended
evolutionary synthesis. His plea coincided
with a similar one made that year by Gerd
Mller, a theoretical biologist at the University of Vienna. Together, with support from
the Konrad Lorenz Institute for Evolution
and Cognition Research in Altenberg, they
organized this weeks conference, inviting
many who share the view that the modern
synthesis is incomplete. Whats happening
now in evolutionary theory is as exciting and
foundational as during the early days, says
David Wilson of Binghamton University in
New York, another attendee.

Evolutionary Theory 1.0:


Common descent
Natural selection

Missing a theory of heredity

(after having flirted with Lamarckism
and blending inheritance)

Evolutionary Theory 1.1:


Rejection of Lamarckism
Separation of soma and germ


Still missing a theory of heredity...

Evolutionary Theory 2.0:

the beginning of the Modern Synthesis




Compatibility between Mendelism

and statistical genetics
Theories of selection and random
drift: birth of population genetics

Evolutionary Theory 2.1:

the mature Modern Synthesis
Variation in natural populations
Species concepts, speciation processes
Compatibility of gradualism with paleontology
Applicability of Darwinism to variety of mating
and genetic systems in plants






The major tenets of the evolutionary synthesis were that populations

contain genetic variation that arises by random mutation and
recombination; that populations evolve by changes in gene frequency
brought about by random genetic drift, gene flow, and especially natural
selection; that most adaptive genetic variants have individually slight
phenotypic effects so that phenotypic changes are gradual; that
diversification comes about by speciation, which normally entails the
gradual evolution of reproductive isolation among populations; and that
these processes, continued for sufficiently long, give rise to changes of such
great magnitude as to warrant the designation of higher taxonomic levels.

Doug Futuyma

Do we need Evolutionary Theory 3.0?

Toward an Extended Synthesis

[the Modern Synthesis] is strictly a theory of genes,

yet the phenomenon that has to be explained in
evolution is that of the transmutation of form.
(Karl Popper)

How do we factor in development?

Is evolution always gradual?
Is selection the only organizing principle?
What are the targets of selection?
Is there a discontinuity between micro- and
Is the question of inheritance settled?
Where do evolutionary novelties come from?
Oh, and what about ecology?

The view from Altenberg:

taking evo-bio seriously as a historical science,
the role of contingency

John Beatty

The view from Altenberg:

MS-based population genetic theory is limited

Sergey Gavrilets

The view from Altenberg:

multi-level selection theory
is here to stay

!z = cov (W, Z) + E cov (w, z)

(Price 1972)
David S. Wilson



The view from Altenberg:

understanding the Genotype > Phenotype map
Greg Wray

Michael Purugganan

Genomic networks
(less emphasis on
role of
individual genes)

Gunter Wagner

Evolvability and
evolutionary Robustness

The view from Altenberg:

epigenetic and other inheritances
Eva Jablonka

Epigenetic (methylation, iRNA,
histone conformation)
Behavioral (mimicking)
Cultural (traditions, memes)

Eors Szathmary

The view from Altenberg:

niche construction & inheritance

John Odling-Smee

The view from Altenberg: innovation, facilitated

variation and the role of physico-chemistry

Gerd Muller

Marc Kirschner

Stuart Newman

The view from Altenberg:

tempo and mode of macro-evolution
Yours Truly

David Jablonski

plasticity can catalyze
fast phenotypic shifts via
phenotypic / genetic

The predictive power of

conventional short-term observations
is not strong over larger temporal
and spatial scales ... Evolutionary
novelties originate non-randomly in
space and time ... evolutionary
stasis ... species selection

The view from Altenberg:

implications of and for philosophy of science

Werner Kallebaut

Alan Love

Is this a paradigm shift?

Thomas Kuhn

Agency: Where natural selection acts (so-called

units of selection problem).
Efficacy: The relative power of natural selection in
comparison to other evolutionary mechanisms.
Scope: The degree to which natural selection can
be extrapolated to macroevolutionary processes.


An (extended) synthesis of what?


networks theory

epigenetic theory
Mendelism common
plasticity &
multilevel contingency
selection theory
evolvability &