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Different forms of cooperative behaviours can be
observed in human as well as non-human animal societies
(e.g. cooperative breeding, cooperative hunting,
alliance formation). What makes these cooperative
behaviours so attractive for scientific research is the
element of instability. Why should individuals invest
without a guarantee of net returns? In the proposed
study I plan to investigate a specific aspect of
cooperation namely the sensitivity towards unequal
reward or effort distribution. For a long time, inequity
avoidance has been thought to be a uniquely human
feature, until it has been shown in primate species and
recently, also in dogs. Until the present day all
studies focused on mammalian species. I plan to
investigate sensitivity to unequal reward structure for
the first time in a bird species, the carrion crow (Corvus
corone corone). Tests will be conducted on a group
of 8 captive carrion crows at the Konrad-Lorenz
Forschungstelle (KLF) in Upper Austria. In the last two
years these birds were trained to exchange a token
(non-edible item) for food with a human experimenter.
This procedure has been applied in primate species to
answer questions like, how individuals deal with delay
of reward (Dufour et al. 2007), differences in food
quality (Drapier et al. 2005) or unequal reward (Brosnan
& De Waal 2003; Brosnan et al. 2005). To my knowledge,
the crows at the KLF are the first bird species capable
to perform in exchange tasks. I plan to replicate tests
conducted in primates on inequity avoidance starting in
November 2009. For comparative reasons, I will follow
the methods used by Brosnan and co-workers (2003; 2005):
A focal subject and a model individual will be tested
together in 4 experimental conditions: (1.) equity test:
both subjects will receive a low-value food reward for
exchanging; (2.) inequity test: the test subject
witnesses the model individual exchanging for a high
value reward, afterwards the focal subject itself,
receives a low-value reward for the exchange; (3.)
effort control: the focal subject watches the model
individual receiving high quality reward without
exchanging, then the focal subject has to exchange for
the reward; (4.) food control test: the high-value
reward is presented but not given to the model,
afterwards the focal subject has to exchange for the
reward. For each individual and in each condition I will
measure the frequency of refusals to exchange and the
latency to exchange. In a final step, starting in April
2010, I will investigate how social relationships affect
inequity aversion. It has been previously shown in
chimpanzees that tolerance for inequity increases with
social closeness (Brosnan et al. 2005). I will test all
individuals in the inequity test once with an affiliated
and once with a non-affiliated social partner. To be
sensitive for other individuals efforts and payoffs
compared to the own ones is thought to be a crucial
cognitive skill for cooperation but until now has only
be shown in mammalian species. I expect carrion crows,
to be sensitive to inequity and to respond with refusal
to exchange when being unequally gratificated. If they
are also sensitive to the effort an individual is
investing, they should also respond with refusal in the
effort control condition, when another individual
receives gratification without the effort of exchanging
whereas the focal individual has to exchange to receive
the reward. Additionally, I also expect the identity of
the model individual to influence the tolerance to
inequity. When tested with affiliated individuals focal
subjects should refuse less frequent compared to being
tested with non-affiliated individuals. The objective of
the proposed study is to examine a form of fairness
(inequity aversion) in a bird species. In a broader
sense, however, my goal is to initiate a research
program to investigate carrion crows cooperative skills,
aiming at a better understanding of the evolution of
cooperative behaviour and its cognitive underpinnings. |