418 Animal Memory and Cognition
animal’s coming, with training, to salivate to the tone.
Pavlovian learning of this sort appears to be common in a va-
riety of species (see, e.g., Wasserman & Berglan, 1998). Pre-
sumably this is because learning such Pavlovian relationships
is important to survival in many species. That is, it is hardly
uncommon for events of biological significance to be reliably
signaled by other events—a rustle of leaves indicating an ap-
proaching predator, circling vultures indicating a dead animal
and a potential meal, and so on. If this is so, then as expected,
learning Pavlovian relations is common to a wide variety of
species.
If, as indicated, regularities in the environment play an im-
portant role in shaping an animal’s cognitive capacities, then
it becomes incumbent upon investigations to obtain knowl-
edge about regularities and behavior in that environment. We
have already seen how our understanding of spatial learning
in animals and humans has benefited from such an approach.
To suggest another familiar example, we have seen how our
understanding of language has benefited from examining
real-life situations in which children exposed to a pidgin
develop on their own, within a single generation, a more
complex form of language called a Creole. As another exam-
ple, observations of chimpanzees in the wild suggest that
they possess a high level of cognitive capacity that might oth-
erwise go unnoticed. Field observation of 14 different groups
of widely separated chimpanzees suggests that they may be
characterized as possessing a culture (Whiten & Boesch,
1999). By culture it is meant that the various groups of chim-
panzees engage in complex behaviors that are obviously
learned and, in addition, are passed on from one generation to
the next.
An outstanding example of such complex behavior is ter-
mite fishing, in which chimpanzees insert thin, flexible strips
of bark into termite mounds to extract the insects, which they
eat. This behavior is practiced by 8 of the 14 groups of chim-
panzees observed. However, it is practiced differently by dif-
ferent groups. For example, once the insects have swarmed up
the stick, 3 of the 8 groups pull the stick through their fists and
eat the gathered prey. This technique is used by some other
groups of chimpanzees who also pull the stick through their
mouths. To cite another complex behavior, 3 of the 14 groups
of chimpanzees use small sticks to remove bone marrow in-
side the bones of monkeys they have killed. The point here, at
least as suggested by Whiten and Boesch (1999), is not
merely that termite fishing and removing bone marrow with
sticks are complex learned behaviors, but that they are passed
on from one generation to the next, and so, in the opinion of
some, indicate that chimpanzees possess a culture. Whiten
and Boesch (1999) indicate that still other animals may pos-
sess a culture, as (for example) populations of whales that
sing in different dialects and hunt in different ways. Are the
sorts of behaviors engaged by chimpanzees and possibly
other animals, such as whales, really manifestations of cul-
ture, on par in some respects with human culture as suggested
by Whiten and Boesch (1999)—or is it something simpler?
It may be, for example, that termite fishing is composed
of many component behaviors, gradually and accidentally
learned over time by one or two members of a chimpanzee
group, which are then acquired by other members according
to the same principles that explain other forms of instrumen-
tal learning. Laboratory studies that, for example, compare
learning by imitation in chimpanzees with such learning in a
variety of other animals, including humans and birds, can
help determine whether ascribing culture to chimpanzees is
valid and useful. The plain implication of the previous analy-
sis is that experimental psychologists, if they are to better un-
derstand learning and cognition, must supplement laboratory
studies of learning and cognition with real-life field studies of
animals dealing with problems of evolutionary adaptation in
their environments.
Although the strategy just outlined is already being
followed in some cases, unfortunately, the practice is less
widespread than is desirable. However, it does seem to be
growing. For example, the behavior system approach as-
sumes that in Pavlovian conditioning, an unconditioned stim-
ulus(UCS) such as food or shock activates an underlying
organization of behavior appropriate to that UCS (for the
food UCS, the feeding system, for shock, the defensive sys-
tem). According to this view, a conditioned stimulus(CS) will
come to elicit responses that are components of the behavior
system activated by the UCS. Thus, not only must preorga-
nized response systems be considered but also a particular
species’ sensitivity to different stimuli and its underlying mo-
tivational system. The usefulness of this approach has been
demonstrated in a variety of laboratory experiments that
were cognizant of the animals’ species-typical behaviors. As
merely one example, Timberlake and Washburne (1989)
examined predation in seven different species of rodents.
Characteristic differences in prey-capture of crickets by the
rodents were related to species-typical behavior (e.g., more
carnivorous species captured prey more quickly and reli-
ably), and the response of each species to a CS, a rolling ball
bearing, resembled that particular species’ typical response to
actual prey.
Felial imprinting (e.g., baby ducklings’ following a
parent) is another example of how an understanding of
species-typical behavior may be profitably used to conduct
and illuminate laboratory studies. Some of the so-called
“attributes” of imprinting based on field studies have been
shown by laboratory studies not to be entirely correct. For