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The Ultimate Benefits of Learning
Kathy Baker and Vicky A. Melfi

3.1 Introduction whether it is selected (survival value and
evolution), whilst proximate explanations are
One of the first theories or concepts univer- concerned with the physiological mecha-
sity undergraduates studying an animal nisms which enable a trait to be performed
behaviour degree come across in their aca- (causation and ontogeny).
demic careers is likely to be Tinbergen’s four Proximate explanations for learning are
‘questions’ relating to animal behaviour. discussed to some degree in other chapters
Proposed in his (1963) article ‘On aims and within this book, with an emphasis on ontol-
methods of ethology’, the four fundamental ogy rather than causation, i.e. how learning
questions, sometimes considered to be prob- develops within an animal’s lifetime, which is
lems, and sometimes termed whys, of animal explored in discussions about the processes
behaviour have had a lasting appeal and and methods of learning (e.g. Chapters 1 and
application within biology and changed little 4). In the current chapter we will focus on the
in over 50 years since their first inception ultimate explanations for learning; how learn-
(Bateson and Laland 2013). Tinbergen iden- ing can effect survival of an individual and
tified four fundamentally different questions evolution of a species. This focus will also
that can be asked to explain animal behav- explore why certain learned behaviours
iour which were: how does the behaviour evolve in certain species and not others? We
contribute to the animals’ survival, what is it will begin by conducting a brief review of
for (survival/function); how has the behav- current literature on the survival value of dif-
iour changed over the animals’ lifetime, how ferent learned behaviours such as predator
did it develop (ontogeny/development); how recognition and highlight case studies where
has the behaviour evolved over time within behaviours with a learned component have
the species, how did it evolve (evolution/phy- determined the extent to which the species
logeny); and how is the behaviour caused has succeeded, or failed, during reintroduc-
physiologically, how does it work (causation/ tion attempts. We will then explore the less
mechanism). The four questions can be obvious benefits of the learning process itself,
broadly split into two categories; proximate such as enhanced brain development, which
and ultimate explanations for behaviour may not have immediate or obvious benefits
(Mayr 1961). Ultimate explanations consider to inclusive fitness but no doubt offers poten-
the fitness consequences of a trait and thus tial survival/reproduction advantages.

Zoo Animal Learning and Training, First Edition. Edited by Vicky A. Melfi, Nicole R. Dorey, and Samantha J. Ward.
© 2020 John Wiley & Sons Ltd. Published 2020 by John Wiley & Sons Ltd.