So is the individual the ultimate beneficiary? Not
necessarily. In his classic 1966 book Adaptation and
Natural Selection, the American biologist George Wil-
liams argued that we should recognise adaptations at
the level necessitated by the facts and no higher. But
which level is that? Multilevel selection theory entails
selection operating at many levels, including group
selection in which groups of animals, or tribes, or cul-
tures compete with each other for survival. Arguably,
cultural evolution makes this more likely (Boyd and
Richerson, 2009). If, for example, one human group has
a taboo on eating pork in a region where pigs carry fatal
diseases, that group would have an advantage. In this
case, selection would operate both within the groups
and between them. But group selection is a highly
contentious topic. Powerful advocates include David
Sloan Wilson (Wilson and Sober, 1994) and E. O. Wil-
son (D. S. Wilson and E. O. Wilson, 2008), while Steven
Pinker (2016, p. 878) says it is guaranteed to confuse
because it is too often used to make ‘loose allusions to
the importance of groups in human evolution’.
Against group selection is what is known as ‘selfish
gene theory’ after Dawkins’s 1976 book The Selfish
Gene. On this view, the ultimate beneficiary of natu-
ral selection is neither the species, nor the group, nor
even the individual, but the hereditary information:
the gene. If this seems odd, think about our London
rats. They have genes for numerous physical and
behavioural traits, and natural selection can work
on them all. Although it is the individual rats who
live or die, the net result is changes in the frequency
of different genes in the gene pool. Another way of
putting it is to say that the gene is the ‘replicator’: it is
the information that is copied, either accurately and
frequently, or not. This explains how genes can be
‘selfish’. They are not selfish in the sense of having their
own desires or intentions (they couldn’t; they are just information coded on strands
of DNA); they are not selfish in the sense that they produce selfish behaviour in their
carriers (they produce altruistic behaviours too); but they are selfish in the sense
that they will get copied if they can – regardless of their effect on other genes, on
their own organisms, or on the species as a whole. From this perspective, human
beings (like all other animals) are the ‘lumbering robots’ that have been designed
by natural selection to carry the genes around and protect them (Dawkins, 1976).
There is a danger of seeing every trait as necessarily adaptive (a tendency derided
as ‘panadaptationism’ by palaeontologist Stephen Jay Gould [Gould and Lewon-
tin, 1979]). In fact, many features of organisms are not adaptations, or are far from
optimal if they are. Some are strongly influenced by physical constraints and
random forces, and none is optimally designed because evolution always has to
start from whatever is available and work from there. Some useless traits survive
PRoFILe 10.1
Richard Dawkins (b. 1941)
Born in Nairobi,
Dawkins came to
England with his
family in 1949.
He studied at
the University of
Oxford, where he subsequently became Lecturer in Zo-
ology, Fellow of New College, and then Charles Simonyi
Professor of the Public Understanding of Science until
his retirement in 2008. His first book, The Selfish Gene
(1976), established what came to be called ‘selfish gene
theory’, and was a bestseller for many decades. As a pro-
tagonist in the ‘Darwin Wars’, he battled against Stephen
Jay Gould over the importance of natural selection and
adaptation in evolution (Brown, 1999; Sterelny, 2001).
His book The God Delusion (2007) inspired ‘the new
atheism’, a movement against religious dogma and in-
doctrination whose main proponents, including Dawkins,
were dubbed ‘The four horsemen’. He describes human
beings as mere ‘survival machines’ – the ‘lumbering ro-
bots’ designed to carry our genes around. In promoting
‘Universal Darwinism’, he invented the concept of the
meme as a cultural replicator, and refers to religions as
viruses of the mind. As for consciousness, he thinks it is
‘the most profound mystery facing modern biology’.