Gregory
B. Pollock (1989, p. 205) construed the criticism against
Wynne-Edwards as follows:
“Critique
of Wynne-Edwards’ view on population regulation and sociality
suppose a population of discrete, mutually exclusive groups essential
to his thought. Yet both his past and present work focus on
continually distributed, philopartric populations; his critics have
argued the untenability of a position never his own.”
As
I understood the criticism of Maynard Smith (1964), he did not claim
that Wynne-Edwards (1962, 1963) assumed strict isolation of breeding
groups, but that the sort of group selection he supposed would
require such isolation in order to prevent cheats from undermining
the group benefit.*
One frequently repeated criticism against Maynard Smith (1964) is that his haystack model was biased against group selection:
One frequently repeated criticism against Maynard Smith (1964) is that his haystack model was biased against group selection:
“While
the model was admittedly extreme, it became the sine qua non
of group selection for many [citations omitted]. Groups were assigned
the same indivisible integrity as individuals. Conjectured group
advantageous traits such as altruism (i. e., foregoing some degree of
personal reproduction to aid others) were, by definition, expressed
homogeneously within groups; intra-group heterogeneity was always
eliminated through intra-group selection [citations omitted]. The
appearance of a single mutant cheater, foregoing altruism for an
immediate reproductive advantage in its group, then necessarily
converts a group into a collection of selfish individuals. Group
selection would require that groups with cheaters be eliminated as
fast as they appear; that is, that the rate of group extinction is
identical to the individual mutation rate for cheating, a not quite
absurd possibility in nature [citation omitted].” (Pollock 1989, p.
207)
Though
Maynard Smith (1964, p. 110) admits that his model is “greatly
oversimplified”
the
passages before the model suggest that he tried to bias it towards
what he thought would favour group selection. He lists the conditions
supposedly necessary for group selection as:
“(1) Groups must
for a time be reproductively isolated, because a is eliminated
from mixed groups.” [Where a is a genotype refraining
from breeding at high population density.]
(2) Groups must be
started by one or a few founders, since otherwise groups consisting
entirely of a individuals would never come into existence.
(3) When a group of
A individuals outstrips its food supply, it must not
immediately encroach on the food supply of neighboring a
groups, for if it did so, the advantage of a groups would
disappear.” (Maynard Smith 1964, p. 109f)
Wynne-Edwards’s
scenario lacked, as far as I can tell, a mechanism to prevent cheats
from undermining the group benefit. Clarifying such mechanisms were
later contributions to multilevel selection theory associated with
David Sloan Wilson and others.
Now,
there are the opposing claims that (1.) Wynne-Edwards was wrong because he
did not provide a mechanism against cheats, and that (2.) the critics were
wrong because later research showed that such mechanisms were
possible after all.
Given that Maynard Smith's (1964) criticism was retrospective in so far as he already was in the know of Hamilton's seminal publication on social behaviour and inclusive fitness theory, the above opposition seems to amount to a choice between two Whiggish alternatives. Either retrospectively falsify or vindicate Wynne-Edwards. The falsification is based on a distinction between group and kin selection that Wynne-Edwards did/could not make. And the vindication would be based on still later distinctions of different models of multilevel selection that Maynard Smith did/could not make.
Given that Maynard Smith's (1964) criticism was retrospective in so far as he already was in the know of Hamilton's seminal publication on social behaviour and inclusive fitness theory, the above opposition seems to amount to a choice between two Whiggish alternatives. Either retrospectively falsify or vindicate Wynne-Edwards. The falsification is based on a distinction between group and kin selection that Wynne-Edwards did/could not make. And the vindication would be based on still later distinctions of different models of multilevel selection that Maynard Smith did/could not make.
* I do not deal with the criticism of Williams (1966, p. 92-124), here, because it is a piece of advocacy against group selection mixing arguments of simplicity (Occham's razor), just-so stories for individual selection, new terms (organic vs. biotic adaptation), fossil evidence for species selection (differential extinction), and general ideas about the practical irrelevance of group selection. As Williams (1966, p. vi) states himself: “Reasons are advanced for doubting, a priori, the effectiveness of such group selection.”
P.S.: Other posts on this blog concerning the history of the group selection controversy are:
Wright (1945) on group selection
Hamilton on group/kin selection
Darwin on group/kin selection
What did Wynne-Edwards (1962) actually propose?
Wynne-Edwards regarded groups as a third level of selection - not second
References
Maynard Smith J
(1964) Group selection and kin selection: a rejoinder. Nature 201:
1145-1147. [reprinted in Williams (1971) Group selection. Aldine
Transactions]
Pollock GB (1989)
Suspending disbelief—of Wynne-Edwards and his reception. J. Evol.
Biol. 2: 205-221.
Williams GC
(1966/1996) Adaptation and natural selection. Princeton University
Press.
Wynne-Edwards VC
(1962) Animal dispersion. Oliver and Boyd, Edinburgh.
Wynne-Edwards VC
(1963) Intergroup selection in the evolution of social systems.
Nature 200: 623-626. [reprinted in Williams (1971) Group selection.
Aldine Transactions]
