Professional Documents
Culture Documents
-1
Review
PAUL E. GRIFFITHS*
Department of History and Philosophy of Science
University of Pittsburgh
Pittsburgh
PA 15206
USA
Research on the effects of niche construction is one of the most exciting recent
developments in evolutionary theory. The idea that organisms shape their
environments as well as being shaped by them is familiar to many philosophers
from two famous papers by Richard Lewontin from the early 1980s (Lewontin
1982, 1983). There is a deal of difference, however, between a challenge to
orthodox evolutionary theory, however compelling, and a substantial program
of research that can claim to have expanded evolutionary theory so as to meet
that challenge. In numerous publications over the last decade and a half the
three authors of this work have succeeded in giving real substance to the idea of
niche construction by embodying it in a series of population genetic models
and demonstrating its potential effect on evolutionary dynamics.
Niche construction is a challenge to what Lewontin called the ‘lock and key’
model of adaptation. In this metaphoric vision of evolution the adaptations of
organisms are solutions (keys) to the problems posed by the environment
(locks). Organisms are adapted to their ways of life because they were made to
fit those ways of life. In place of this traditional metaphor of adaptation as ‘fit’,
Lewontin suggested a metaphor of mutual construction. Organisms and their
ecological niches are co-constructing and co-defining. Organism’s both physi-
cally shape their environments and determine which factors in the external
environment are relevant to their evolution, thus assembling such factors into
what we describe as their niche. Organisms are adapted to their ways of life
because organisms and their way of life were made for (and by) each other. For
example, beavers are well-adapted to life in the lakes which are created by their
own dams. Eucalypt forests are well-adapted to the intense fire regime that is
sustained by their own habits of growth and decay. Humans are well-adapted
to survive and reproduce in the context of human material and ideational
cultures.
*Present address: Philosophy program, SHPRC, University of Queensland, Brisbane, QLD 4072,
Australia (e-mail: pauleg@pitt.edu)
1
This review has benefited considerably from discussions with Karola Stotz and John Norton. The
discussion of thermodynamics draws on (Earman and Norton 1999a).
12
The introductory chapter of this book recalls two pairs of equations that
Lewontin used to contrast the traditional model of adaptation as ‘fit’ and the
new model of adaptation as construction. The first pair of equations express
the conventional picture. Change in organisms over time is a function of the
state of those organisms and their environment at each previous instant. The
environment acts on the existing state of organisms by selecting those best
fitted to the environment. The environment itself changes over time too, but as
the second equation shows these changes are not a function of what organisms
are doing at each previous instant.
dO=dt ¼ f(O,E) ð1Þ
evolution. The authors embed their account of biology within the still more
general framework of thermodynamics. When organisms are viewed as a dis-
tinctive kind of thermodynamic system, they argue, it becomes clear that niche
construction is a universal feature of living systems and that at the most ab-
stract level, ecological relationships can be conceived of as organisms inter-
acting thermodynamically with the environment. Most of chapter four of the
book is devoted to elucidating this idea and the results of that discussion are
appealed to regularly in later chapters. Despite this extensive treatment I do
not think that the authors succeed in establishing the implications of ther-
modynamics for biology to which they later appeal.
Odling-Smee, Laland and Feldman are not the first biologists to try to locate
biology within the more general framework of thermodynamics. The best
known attempt is probably Daniel Brooks and Edward Wiley’s Evolution as
Entropy: Toward a Unified Theory of Biology (Brooks and Wiley 1988), al-
though many other authors have made contributions in this field (Weber et al.
1988). Like most of the earlier treatments, the present authors start with the
uncontroversial claim that living systems are dissipative systems – they main-
tain a highly ordered internal state by drawing on external sources of energy
and exporting entropy into their surroundings. The authors identify this export
of entropy with niche construction and set out to give a general, thermody-
namic characterisation of niche construction: ‘some properties of niche con-
struction should be general, if only because, regardless of species, all organisms
have to live at their environments’ expense, and they can do that only by acting
on and perturbing their environments’ (p. 170). If niche construction is to
evolve, it must be profitable to the organisms that engage in it, yielding more
energy that it consumes. Organisms, they argue, are ‘Maxwell’s demons’ which
on average decrease their own entropy at the expense of their surroundings. It
follows that:
Niche construction in its interactions with natural selection must there-
fore be regulated and controlled by information. Specifically, the niche-
constructing activities of organisms must be at least partly governed by
and directed by semantic information, or knowledge, supplied by the
evolutionary process as a result of natural selection… and typically
encoded in DNA (p. 177).
While I think that an important part of this thesis can be salvaged, I will
argue here (i) that the argument from accepting this thermodynamic charac-
terisation of organisms to the conclusion that their activities must be guided by
information is much less straightforward than the authors suppose, (ii) that
semantic information is, in any case, entirely the wrong sort of information to
make the necessary connections to thermodynamics, and (iii) that locating
semantic information primarily in DNA sequence sacrifices many of the
potential insights of the niche-construction framework.
First, what does it mean to say that organisms are Maxwell’s demons?
Maxwell’s ‘demon’ is a creature who knows the positions and velocities of
15
can now go back to evolutionary biology and use this list of properties to
indicate the universal characteristics we should find in all niche-con-
structing organisms (p. 175).
According to the authors there are six of these properties. Fool’s demons,
and thus organisms, must (1) have a goal; (2) discriminate states of their
environment; (3) anticipate those states; and in order to do this must (4) be
‘instructed by knowledge’ (p. 173, their emphases), which in turn implies pos-
sessing a memory. They must (5) be designed in a way that allows them to
accomplish these things and (6) be supplied with an external source of energy.
The last two properties are uncontroversial. Six follows directly from the
second law of thermodynamics. Five is supported by all the arguments that
usually lead us to attribute complex adaptations to natural selection rather
than chance. It is perhaps also uncontroversial that any Fool’s demon must be
a goal-directed system in some broad, cybernetic sense since its operation can
be represented as an attempt to minimise some physical quantity. It is prop-
erties 2–4 about which I am sceptical. The idea that all Fool’s demons must be
information processing systems in any non-trivial sense is far from obvious.
Physicists have described many Fool’s demons. One of the earliest suggestions
was to replace Maxwell’s original demon and trapdoor with a one-way valve
delicate enough to be opened by fast moving molecules. Another involved a
nano-scale ratchet mechanism operated by Brownian motion. One recent
proposal implements something like the one-way valve using an arrangement
of velocity dependent force-fields. These systems can be described as ‘pro-
cessing information’, but only in the trivial sense that the planets ‘compute’
their course around the sun by ‘integrating’ the gravitational forces to which
they are subject. The only reason to describe them in this fanciful way would be
if it gave access to some theorems of information theory that would throw light
on why the second law of thermodynamics holds. But the well-known proofs
that any information processing system that is designed to implement a per-
petual motion machine of the second kind will be defeated by the entropic costs
of information processing are derived from the second law and not the other
way around (Earman and Norton 1999b). There is an active research program
which aims to derive the second law from an information-theoretic treatment
of thermodynamics, but the viability of that research program remains
controversial.
I have suggested that thermodynamics does not provide strong support for
the view that information processing is amongst the universal characteristics of
niche constructing organisms. But suppose that I am wrong about this. If the
present authors could show that organisms must use information to sustain
themselves far from thermal equilibrium, how could they get from there to the
view that this information is ‘‘supplied by the evolutionary process as a result
of natural selection … and typically encoded in DNA’’ (p. 177). The problem is
that, while there is an intellectually respectable way to treat genes as an
information conduit by which the effects of selection on past generations are
17
2
Viruses are, of course, a near exception which is one reason why they are necessarily parasitic on
systems with such a metabolism.
19
that the book’s focus is so much on how organisms construct the selective niche
of their descendants, rather that on how they construct their developmental
niche. The authors recognise that ‘‘Niche construction … contributes to both
the building of the next generation and to changes in the niche-constructing
organism’s own selective environments’’ (p. 381) and it is clear from earlier
work that they have a substantial interest in the former process. Nevertheless,
they devote only three of this book’s four hundred and seventy pages to po-
tential links between the niche-construction framework and developmental
biology. There are substantial issues that urgently need to be addressed con-
cerning the relationship between niche construction and recent work on the
ecological context of development (Gilbert 2001). Such work would rapidly
reveal the need to extend the framework of this book to include the extra-
genetic channels through which parents influence the phenotypes of their
offspring. It is indisputable that organisms recreate in their offspring the dis-
sipative structures that allow them to maintain themselves far from thermal
equilibrium. That they are able to do this owes a great deal to natural selection.
However, as Daniel Lehrman wrote many years ago, ‘‘Natural selection acts to
select genomes that, in a normal developmental environment, will guide
development into organisms with the relevant adaptive characteristics. But the
path of development from the zygote stage to the phenotypic adult is devious,
and includes many developmental processes, including, in some cases, various
aspects of experience’’ (Lehrman 1970). One lesson of niche construction re-
search is surely that those aspects of experience, as much as the genes with
which they interact, have evolutionary explanations.
References
Lewontin R.C. 1983. Gene, organism & environment. In: Bendall D.S. (ed), Evolution: From
Molecules to Man. Cambridge University Press, Cambridge, pp. 273–285.
Sterelny K., Dickison M. and Smith K. 1996. The extended replicator. Biol. Philos. 11: 377–403.
Weber B.H., Depew D.J. and Smith J.D. (eds) 1988. Entropy, Information and Evolution: New
Perspectives on Physical and Biological Evolution. MIT Press, Cambridge, MA.