Biology Reference
In-Depth Information
Further reading
Hamilton's original papers on inclusive fitness theory have been collected together into
a single volume (Hamilton, 1996), which also provides illuminating autobiographical
notes for each of the papers. Dawkins (1976) provides a very readable popularization of
the gene's eye view, whilst Dawkins (1979) gives a lucid discussion of twelve
misunderstandings of kin selection, many of which are still made today. Grafen (1991)
provides a more technical review of inclusive fitness theory and how it can be tested.
Grafen (1985) is the classic text on the concept of relatedness. We discuss a number of
altruistic and mutually beneficial behaviours in more detail in the following two
chapters. The confusion that may arise from redefining terms such as altruism is
reviewed in West et al . (2007a).
There is sometimes an overemphasis - both conceptually and empirically - on the
importance of the relatedness term ( r ) in Hamilton's rule and a corresponding neglect of
the benefit ( B ) and cost ( C ) terms. To some extent, this is the case because genetic
similarity can be measured more easily than components of fitness (Box 11.2). However,
focusing too strongly on r can lead to misunderstanding and confusion, because variation
in B and C is equally important. An excellent example of where variation in C has clear
consequences for when individuals cooperate is provided by Field et al .'s (2006) study of
queuing for reproductive dominance in social groups of the hairy-faced hover wasp.
Gorrell et al . (2010) use Hamilton's rule to show that indirect fitness benefits explain
adoption in asocial red squirrels.
Further examples of greenbeard genes include: the csa gene in the slime mould
Dictyostelium discoideum, which causes individuals to adhere to each other in aggregation
streams, and cooperatively form fruiting bodies, whilst excluding non-carriers of the gene
(Queller et al ., 2003); and the FLO1 gene in the yeast Saccharomyces cerevisiae , which
causes individuals to adhere to each other in groups that are better defended from stressful
environments (Smukalla et al ., 2008). The evolutionary dynamics of greenbeards and
other known biological examples are reviewed in Gardner and West (2010).
One way in which individuals could behave less selfishly when interacting with
relatives, is by exploiting a resource more prudently and efficiently (analogous to the
discussion of producing rather than scrounging in Chapter 5). Frank (1996) provides
an overview of how this could explain variation in the damage that parasites cause to
their hosts, termed parasite virulence. Specifically, if the parasites infecting a host are
highly related, then they have a common interest that favours prudent exploitation of
the host over time (and hence lower virulence), to maximize the total amount of
resources that can be acquired. Empirical support for the prediction that a lower
relatedness between the parasites infecting a host favours a higher virulence comes
from Herre's (1993) comparative study on fig wasp nematodes and Boots and Mealors
(2007) experimental study in a virus of moths.
Inglis et al . (2009) provide an example of how more specific predictions for spiteful traits
can be made and tested experimentally. They show how the advantage of spiteful traits,
such as bacteriocin production, will vary depending upon aspects of the population
structure, including the proportion of individuals in a patch that are clonemates.
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