Chapter 4

The development of behavior:

A focus on the environment
(2nd lecture)
 Konrad Lorenz was the first person to hypothesize that
learning could be genetically determined

He postulated that animals were predisposed to learn about

essential features of their environment, at the most relevant point
in their ontogeny

He asked how young birds learn to identify and follow

their mothers. Through this work, he developed the
concept of filial imprinting--i.e., a process by which
an animal forms a strong attachment to a specific
animal or object during a critical period in
development
 Filial imprinting
Moving stimulus to which
bird is being imprinted Young bird

Circular arena used to test for Illustration of the sensitive period

filial imprinting in young birds for filial imprinting in mallard ducklings.
The critical period was the time during
which the imprinting process was most
effective (i.e., the 13-16 hr time interval).
 Locomotion-fear dichotomy in developing birds:
its relationship to filial imprinting

Percentage of animals showing fear

Sensitive Age (hours) Sensitive
period for period for
ducklings chicks

Critical period occurs during the peak of the sensitive period,

when fear is declining and locomotor ability is maximized.
Imprinting is a proximate mechanism for helping young
learn to identify and follow their mother.

Is the imprinting phenomenon specific to the young bird’s

genetic mother?

If not, then why don’t birds frequently imprint on

inappropriate animals or objects in nature?

Konrad Lorenz being

followed by geese
Is imprinting limited to birds? that had imprinted
upon him.
No. Here twin orphan lambs of
the Soay breed were fostered
to a Blackface ewe by tying half
of her dead lamb’s skin over
each of the orphans.
Do other animals have other mechanisms for learning to
distinguish kin from non-kin?

Nests of Polistes wasps contain odors

that adhere to the bodies of the larvae
reared in them, providing a proximate
cue for kin recognition in these insects.

Note the larvae with their bodies pressed

up against the cell walls.

Any wasp that lands on the nest, which

possesses an unfamiliar odor, will be
attacked fiercely by the adult wasps.
 Evidence for kin discrimination in Belding’s ground squirrels

Sisters reared apart display significantly less aggression towards one

another than nonsibling females reared apart; a weaker version of the same
result is apparent for males
Evidence that golden hamsters can distinguish kin from strangers
The investigators used females that were ovulating, and thus most likely to
be interested in the odors of males. Further the unfamiliar kin and nonkin were
reared with foster parents

The females preferred to mate

with unfamiliar nonkin over both
unfamiliar kin and familiar non-kin. Further, they flank-marked most often
after smelling the odor of an unfamiliar nonkin. This marking behavior
deposits individually specific odors onto substrates, and is thought to
warn off close relatives at mating time.
So, how do we explain the behavior of the female hamsters?

At the ultimate level?

At the proximate level?

 Proximate explanation:
The results with the hamsters (and Belding’s ground squirrels) provides
support for phenotype matching (or the “armpit effect”)—that is, a self-matching
mechanism for assessing the degree of relatedness of conspecifics

Background information
1. Each individual has a unique odor signature
2. The difference in odor signature between individuals decreases with
degree of relatedness (i.e., you smell more like your sister than your
third cousin twice removed)
3. The major histocompatibility complex (MHC), a highly variable family of
genes involved in your secondary immune response, plays a critical role
in this process. Each individual’s unique combination of MHC alleles
confers a unique odor signature.

Thus, according to the phenotype matching mechanism, an animal learns

its own odor during development, and then assesses its degree of
relatedness to a stranger by comparing its own odor with that of the
stranger’s. The more similar the odor, the more related the individual.
 In addition to learning to discriminate kin from nonkin,
developing individuals must learn a variety of other
features of its environment

e.g.,
location of home
location of food
friends vs. foe
safe vs. unsafe foods

Key point:
each species has a genetically determined aptitude for learning
specific types of information
Spatial learning in the
digger wasp

Niko Tinbergen’s experiments

on nest-locating behavior in the
digger wasp illustrate the use of
landmark orientation
 Spatial learning in a vertebrate: retrieval of stored food by
black-capped chickadees

Birds spent more time searching parts of the cage where they stored food
24 hr previously (hoard sites) that they did during their initial exposure to
those sites. They also made many more visits to hoard sites, evidently
because they remembered having stored food there.
Note: food was removed before birds were allowed to search for it
 The bird species with the greatest
reliance on locating hoard sites to
endure the winter are the ones with
the most prodigious long-term spatial Clark’s
memory nutcrackers

Note that this spatial memory does not

generalize to nonspatial tasks

Birds had to remember location of a circle Birds had to remember color of a circle