CLASSICAL ETHOLOGY

Ethology, the European science of animal behavior, developed

out of zoology and emphasized instinctive behaviors

N. Tinbergen K. Z. Lorenz K. Von Frish

(1907-1988) (1903-1989) (1886-1982)

Ethology has three «fathers», who shared the Nobel prize

for Medicine in 1973 1
 Niko Tinbergen wrote The Study of Instinct (1951), a book filled
with entertaining examples of animal behavior. A year later he
published an influential Scientific American article, "The Curious
Behavior of the Stickleback" (Tinbergen, 1952). For many American
psychologists, this was a first exposure to the concepts of ethology.
He described how the little fish Gasterosteus aculeatus went through
a complex series of behaviors during mating season, each
triggered by a specific stimulus.

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Konrad Lorenz described a famous
phenomenon of animal behavior.
Filial imprinting occurs when
ground-dwelling birds follow the first
object they see after hatching,
usually their mother. Lorenz let baby
greylag geese imprint upon him, and
they followed him all around his
farm.

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KARL VON FRISH described the fascinating communication
system occurring among honey bees (the waggle dance)

Foragers which found a source of
food perform the dance on the
vertical surface of the comb once
they have returned back to the hive.
The round dance is performed when the food source is
less than 80 meters from the hive: «fly out and look
around». The waggle-tail (figure of eight) dance is
performed when the food is farther from the hive «fly
farther away and in a specific direction»

Von Frish described the code

through which bees signal the
specific direction in which the food
can be found. If you are curious to
know how they do, see the next
slide. 4
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CLASSICAL ETHOLOGY CONCEPTS
ETHOGRAM

FIXED ACTION PATTERN (FAP)

SIGN STIMULUS
SUPERNORMAL STIMULUS
INNATE RELEASING MECHANISM

ACTION SPECIFIC ENERGY

VACUUM ACTIVITY
DISPLACEMENT ACTIVITY
REDIRECTED ACTIVITY 6
 Species-typical behaviors are often listed in an
ethogram compiled by close observation

= A complete inventory or descriptive catalogue of the motor

patterns exhibited by members of a particular species.
Development of an ethogram to this day is considered a critical
first step in studying the behavior of any new animal species.

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 Vertical headshaking in the horse
Usually an upward and then downward movement of the
head, generated from the neck
The speed with which this occurs may vary so that it
appears as a nod, a snatch or a rapid flick. The extent of
the movement may also vary from a small flick to a large
arc of movement. May occur as a single spasm or a series.
Common occurrence : It is seen at any pace, including at
rest, although the trot is the most common. It occurs less
commonly at canter where the head movement may more
closely resemble a rotary headshake.
Other names : „headflicking‟, „head tossing‟ (Cook 1992,
Madigan et al. 1995), „head swinging‟, „head throwing‟
(Madigan et al. 1995).
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Est. prevalence : 85–100%
 FIXED ACTION PATTERN (FAP)
= complex motor pattern with all of the following characteristics:
• the behavior sequence is highly stereotyped;
• it is exhibited by all members of a species (of the appropriate
age/sex class);
• it is elicited by a simple, highly specific stimulus;
• it is self-exhausting;
• it occurs in its full-blown form the very first time it is ever
emitted in the animal‟s lifetime;
• once started, the behavior runs to completion without
additional stimulation.

Examples : the gaping response of nestling birds when a parent lands on the nest;
the tongue-flicking behavior with which frogs catch flies; courtship displays; head-
bobbing display by male iguanas; aggressive displays; nestbuilding; egg-retrieval,
pecking at parent‟s beak by baby birds, etc.
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FAP is made by a fixed (cannot be changed) sequence of
movements; it is not learned; the response can be triggered
by stimuli which are not the natural ones; all members of a
same species share the same FAPs

e.g. Egg retrieval in Anser anser (Lorenz and Tinbergen 1938)

Tinbergen and Lorenz collaborated on a classic article describing the egg-

retrieving behavior of the greylag goose. The egg-retrieval movement shows
many characteristics typical of motor programs. 10
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 CHAIN REACTION: SEQUENCES OF FAP
es. Courtship behaviour in the Stickleback

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The egg-retrieval is stereotyped or fixed in form, it is set off by
a highly specific stimulus called a sign stimulus or releaser.
Once triggered, the action runs to completion (endogenous
running-out).

sign stimulus or releaser: It is the simply, yet highly specific

stimulus that triggers (releases) a FAP.

IDENTIFYING THE SIGN STIMULUS 13

complex motor stimuli are often set off by highly specific stimuli:
(A) "A male red-winged blackbird copulating with a mount consisting of the tail
feather of a female raised in pre-copulatory position." Even though the tail feather is
on a seed pod, not a female bird, it sets off the sexual response in the male.
(B) "Willow warblers attacking the stuffed head of a cuckoo." Even though the head
is on a stick, clearly not alive and not a real bird, the small warblers nevertheless 14
attack it.
es. The black moustache in the male of northamerican
woodpecker Colaptes auratus is a sign stimulus to trigger
aggression against male conspecific intruders
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 RELEASERS

In one experiment,
Tinbergen fashioned a
series of stickleback
models, ranging from a
very realistic but
colorless model to a
very unrealistic blob
with a red belly. The
male stickleback
ignored the realistic fish
without the red belly but
attacked all the models
with the red bellies.
TINBERGEN, 1948
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One day Tinbergen's lab students noticed all the sticklebacks attacking the
sides of their aquariums near a window. Across the street was a red postal
van. The small patch of red visible through a window was enough to set off
attack responses in the Sticklebacks.
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More examples of SIGN STIMULI or RELEASERS
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 Lorenz showed that the egg-retrieval response was also
triggered by supernormal stimuli : exaggerated sign stimuli
which may trigger a stronger than normal response.

Supernormal stimulus: a model stimulus that elicits a FAP

more effectively than the real thing (e.g. the huge eggs
Tinbergen gave his female greylag geese elicited eggretrieval
more effectively than did their own eggs).
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Tinbergen and Perdeck, 1950
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Intention movement: The incomplete or preparatory
movements that occur at the beginning of a behavior
sequence. In spotted hyenas, for example, a “head wave” is an
intention movement to engage in more serious attack if the
pesky social partner does not leave the aggressor alone. The
head wave is also the initial part of the larger attack sequence.

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Vacuum Activity: Behavioural pattern may appear in the
absence of the appropriate stimulus, as if animals had the
need to exercise their behavioural repertoire.
When the stimulus is not available. The activity may appear in
total absence of any stimulus or in the presence of a usually
not suitable stimulus.

squirrels raised from birth in a metal cage will go through the entire sequence of
nut-burying activities, despite the lack of dirt or a nut in the cage. 24
Redirected activity: A behavior directed at a third party
or an inanimate object when the animal emitting the
behavior is conflicted about whether or not to direct this
behavior at its original social partner in the first place.

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Displacement activity (displacement behavior): A
common behavior occurring entirely out of context when
the animal is experiencing serious conflict between two
opposing tendencies (eg during a fight with a territorial
neighbor, when a male bird is torn between attacking
and fleeing from his opponent, he suddenly starts
grooming or nest-building!).

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Babyishness / Neotenia
Babyishness is another pattern pointed out
by Lorenz. He observed that all baby
animals have a distinctive look. They have
prominent foreheads, large eyes, soft jaws,
fuzzy or furry bodies, and stubby limbs.
This appearance seems to inspire merciful,
parenting behavior in adult animals of
other species, sometimes resulting in
cross-species adoption.

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Babies resemble each other, they share some common features
(Lorenz 1943)

Adults respond positively to these features even if they belong to

different species

Stephen Jay Gould, 1980

MORE TERMINOLOGY FROM CLASSIC ETHOLOGY

Innate releasing mechanism (IRM): An hypothesized locus in

the central nervous system on which sign stimuli act to release
a FAP. The early ethologists suggested that a sign stimulus
would “fit” an IRM much like a key would fit into a lock.

Action specific energy (ASE): Each FAP was believed to

have its own reservoir of action-specific energy that appeared
to increase in quantity as more and more time elapsed since
the FAP was last emitted. Sometimes ASE builds up so much
in the reservoir that the FAP will occur in the absence of any
stimulus (this was called a “vacuum activity”).

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LORENZ’S “HYDRAULIC” MODEL OF MOTIVATION (1950)
Explains both vacuum as well as
displacement activities
Hydraulic drive model of
motivation. Fluid enters a reservoir
through the tap, which represents a
continuing flow of energy from
endogenous sources. The height of the
fluid in the reservoir indicates the build-up
of drive energy. The fluid is held in the
reservoir by a spring-loaded valve, which
represents the inhibitory pressure from
higher-level brain areas. The weighted
scale pan represents the strength of
external releasing factors, which can
increase the likelihood of valve release in
addition to the pressure from the fluid in
the reservoir. When the valve opens,
energy flows through the outlet into a
slanted trough. The holes on the bottom of
the trough coordinate muscle action
pattern responses, where the yellow
numbers indicate the rank of the response
strength. (After Lorenz 1950.)
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 QUESTIONS :

How can you determine for a given species if a certain behaviour is

or is it not a fixed action pattern?

Why does the male stickleback readily attack anything that may
resemble a fertile male intruder, even if this means at time attacking
the wrong object, e.g. the image of the postal van?

Why are supernormal stimuli a very interesting topic of investigation

for ethologists and psychologists?

What is a vacuum activity and how can it be explained?

Give some examples of displacement activities and redirected

activities in humans.

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