PSY 336: Part 1--Page 1 of 10
PSY 336: Ethology
Ethology is the scientific study of animal behavior especially in a natural
setting. The word “ethology” comes from “ethos” which means the
distinguishing character, habit, manner, or behavior of an organism.
Part 1: Chapters 1, 2, and 3
This is not a new area of study. Before the development of the
supermarket, people had to know animal behavior to get protein and fat. What
distinguishes ethology from the work of hunters, horse trainers, and so forth is
ethology’s scientific methods.
History of the Study of Animal Behavior
I. Prehistoric times
A. hunters and gatherers
B. L. S. B. Leakey (1903-1972), Kenyan anthropologist who worked in
Olduvai Gorge in Tanzania
1. Jane Goodall (chimpanzee)
2. Dian Fossey (mountain gorilla)
3. Birute Galdikas (orangutan)
C. fire to drive animals
II. Classic Greek World
A. dualism--two worlds
1. psychic world--humans are rational, this separates humans from
2. physical world--animals are like machines and cannot think
B. Aristotle (384-322 BCE)
1. used observational method
2. vitalist: living things had a vital force
3. De Anima (On the Soul)
a. postulated phylogenetic development and continuity
between species (Scala naturae)
b. postulated a doctrine of behavior modification (an
elementary S-R theory)
Pliny (23-79), wrote Natural History, and used anthropomorphism
(attribution of human characteristics or personality to nonhumans)
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IV. The natural philosophers
A. Francis Bacon (1561-1626): be empirical, be objective, observe,
experiment, and use inductive reasoning rather than rationalism.
B. René Descartes (1596-1650) dualistic interactionism (two worlds--
psychic world and physical world interact)
1. animals are like machines, their behavior can be predicted
2. humans have a soul and free will
3. but the human body is a machine, it can be studied
As the natural world came to be quantified and measured by the mathematical
method, the human body, as part of the natural world, posed special
considerations. How could one measure emotions? Or quantify the soul? René
Descartes provided the philosophical justification for conceptualizing the
human body in mathematical terms by positing two separate but interacting
aspects that comprise the human body--the res cogitans, thinking substance,
and res extensa, the extended or physical substance. Quite simply, the human
body could now be divided into mind and body. With the advent of Newtonian
physics, the extended or physical world, including the human body, came to be
interpreted through the laws of matter and motion.
C. John Locke (1632-1704)
At birth the human mind is a blank slate (tabula rasa).
D. Immanuel Kant (1724-1804)
1. rejected the concept of a tabula rasa
2. there has to be some native (innate or inborn) ability to
organize what is observed
V. Theory of evolution by natural selection
A. Charles Darwin (1809-1882) was the naturalist on HMS Beagle,
which visited the Galápagos Islands. This got him thinking about earlier
B. Thomas Malthus (1766-1834)
1. Essay on the Principle of Population (1798)
2. population increases geometrically (2, 4, 8, 16, 32, ...) but the
food supply increases arithmetically (2, 4, 6, 8, 10, ...)
C. Sir Charles Lyell (1797-1875) geologist who observed rock strata
showed a succession of fossils that indicated a process of continuous
2. species are not fixed
3. animal breeding had already provided support
D. Herbert Spencer wrote Principles of Psychology in (1855). In it, he
proposed there was an intellectual continuity among animals
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E. Alfred Russell Wallace (1823-1913)
F. Both Darwin and Wallace independently formulated the theory of
evolution by natural selection.
1. Darwin wrote:
a. The Origin of Species (1859)
b. On the Expression of the Emotions in Man and Animals
2. The theory of evolution restored the continuity between
humans and other animals.
VI. Comparative method
A. George John Romanes (1848-1894)
1. was a close friend of Charles Darwin
2. coined the term “comparative psychology”
4. used anecdotal evidence rather than empirical tests
B. C. Lloyd Morgan (1852-1936)
1. Introduction to Comparative Psychology (1894)
2. both observations and empirical method
3. Morgan's canon: “In no case may we interpret an action as the
outcome of the exercise of a higher mental faculty, if it can be
interpreted as the exercise of one which stands lower in the
psychological scale” (Morgan, 1894), the Law of parsimony or
William of Occam's razor
VII. Theories of genetics and inheritance
Gregor Mendel (1822-1884)
A. Physiological psychology
1. Marie-Jean-Pierre Flourens (1794-1867)
2. Karl Lashley searched for the engram
3. Roger W. Sperry won a Nobel prize in 1981
B. Animal psychology (or comparative psychology)
1. Edward L. Thorndike (1874-1949)
a. puzzle box
b. trial-and-error learning (instrumental learning) (aka
c. Law of Effect: “If a response in the presence of a
stimulus is followed by a satisfying event, the association
between the stimulus and the response will be strengthened.
Conversely, if the response is followed by an aversive event,
the association will be weakened” (Dugatkin, p. 130).
2. Ivan Pavlov
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3. Robert M. Yerkes (1876-1956)
a. studied many species
b. founded a primate center
c. Army Alpha and Beta
4. Frank Beach
a. APA president in 1950
b. too many rat studies
5. Hodos and Campbell (1969)--psychology needs an evolutionary
1. Jacques Loeb (1859-1924)
a. tropism (forced movement)
b. mechanistic point of view
2. Herbert Spencer Jennings
a. disagreed with Loeb
b. said behavior was variable and modifiable
3. John B. Watson (1878-1958)
a. S-R psychology
b. tabula rasa
c. Behavior: An Introduction to Comparative Psychology
4. B. F. Skinner (1904-1990)
a. operant conditioning
b. laws of learning
A. Early ethology
Behavior can be studied with an evolutionary point of view just like
anatomy and physiology.
1. Charles O. Whitman (1842-1910) used the display patterns of
birds to classify them.
2. Jacob J. von Uexkull (1864-1944) Umwelt (sensory-perceptual
4. Konrad Lorenz (1903-1989)
a. Nobel prize in 1973
b. fixed action pattern (FAP)
c. sign stimulus
d. supernormal sign stimulus
e. innate releasing mechanism (IRM)
f. chain of reactions, e.g., courtship in the three-spined
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g. theories, e.g., psycho-hydraulic model
h. action specific energy
i. vacuum activity
j. displacement activity
k. critical periods (imprinting)
5. Niko Tinbergen (1907-1988)
a. Nobel prize in 1973
b. Four Questions (on page 6)
1) causation (or immediate stimuli)
3) survival function
c. hierarchical model
1) IRM (innate releasing mechanism)
2) after a sign stimulus is presented, it is identified
by the IRM which removes a block and allows an FAP
6. Karl von Frisch (1886-1982)
a. Nobel prize in 1973
B. Comparisons in 1950:
biological background psychological backgrnd
many species “white rat”
field work laboratory setting
innate behavior learned behavior
evolutionary not evolutionary
There is an interaction between what is innate and the environment,
which allows life and learning.
D. Changing terms:
1. instinctive behavior
to: species-specific behavior
to: species-typical behavior
2. fixed action pattern
to: modal action pattern
3. innate as absolute
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to: innate as relative
E. isolation experiment
F. breeding experiment
G. Hess: “innate” pecking behavior of gull chicks gets better with
practice (is learned)
H. Brelands: “instinctive drift” nonreinforced innate behavior
interfered with “learned” behavior
I. Seligman: there is a continuum of preparedness to learn new
associations from prepared (learn quickly) to contraprepared (takes
many trials or may not learn at all)
J. Garcia has shown that the internal state of nausea can easily be
paired with internal cues, such as tastes or odors, but not with
external cues, such as sounds or lights.
K. Today we say that genes and environment interact in the
development of every behavior.
X. Sociobiology (also called behavioral ecology)
A. applies the principles of evolutionary theory to the study of social
B. How could helping another individual raise its young (altruism)
C. William D. Hamilton wrote in 1964 that evolutionary success is the
result of your inclusive fitness. Inclusive fitness is your surviving
offspring (direct fitness) plus offspring of kin (indirect fitness).
D. Edward O. Wilson wrote Sociobiology: The New Synthesis (1975)
Principles of Animal Behavior, pages 2-25
I. Early art shows animal behavior
Fig1.2 Fig1.3 (parallel walk) Fig1.4
II. Types of Questions and Levels of Analysis
Niko Tinbergen (1907-1988)
Four Questions (on page 6)
causation (or immediate stimuli)
III. Three Foundations
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A. Foundation 1 --Natural Selection
B. Foundation 2 -- Individual Learning
C. Foundation 3 -- Cultural Transmission through social learning
IV. Conceptual, Theoretical, and Empirical Approaches
A. Conceptual Approaches
B. Theoretical Approaches
C. Empirical Approaches
The Evolution of Behavior, pages 26-71
I. Artificial Selection
Controlled by humans
II. Natural Selection
A. Selective Advantage of a Trait
If one allele (variant form of a gene) gives an advantage over other
alleles, it will increase in frequency over generations.
This changes the genotype (the genetic information of the
organism) over many generations.
B. How Natural Selection Operates
Ernst Mayr (1977) proposed that Darwin and Wallace thought like this:
Fact 1: All species are capable of overproducing.
Fact 2: Populations of species tend to remain stable.
Fact 3: Resources are limited
Inference 1: There is a struggle for existence among individuals
Fact 4: Individuals are unique.
Fact 5: Individual differences can be inherited.
Inference 2: Differential survival, or natural selection, occurs.
Inference 3: Through many generations--evolution.
III. Behavioral Genetics
A. Mendel's laws
B. Locating Genes for Polygenic Traits
C. Dissecting Behavioral Variation
IV. The Modern Theoretical Framework for Animal Behavior
A. Sociobiology and Selfish Genes
“Any gene that codes for a trait that increases the fitness of its bearer
above and beyond that of others in the population will increase in
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frequency. So natural selection often, but not always, produces genes that
appear to be selfish” (Dugatkin, 2009, p. 44).
B. Antipredator Behavior in Guppies
A. Adaptation leads to the highest fitness among a specified set of
behaviors in a specified environment.
B. An adaptive trait is an inherited characteristic that increased in a
population (usually through natural selection) because it helped solve the
problem of survival or reproduction during the time it emerged.
C. Brood parasitism in wood ducks appeared to be maladaptive when
there are many man-made nest boxes in the environment.
VI. Genetic Techniques to Test Hypotheses in Animal Behavior
A. Kinship and Naked Mole Rat Behavior
1. Show eusociality
a) show reproductive division of labor where some castes
reproduce and other castes do not
b) there is an overlap in generations where older
generations care for younger generations
c) there is communal care of young
2. There is high genetic relatedness (r = .81) among individuals
within the same colony.
3. Kinship theory states that the more highly related individuals
are, the more we expect to see cooperation and altruistic behavior.
B. Coalition Formation
1. Small coalitions of male lions are sometimes made up of
relatives and sometimes not.
2. Large coalitions of male lions are always composed of
VII. Phylogeny and the Study of Animal Behavior
A. Phylogeny--evolutionary history through common descent
B. Phylogenetic Trees
Homology--a trait shared by species because of a common ancestor
Homoplasy--a trait that is not due to a common ancestor, e.g.,
wings of birds, bats, and insects, which are analogies that are produced
by convergent evolution.
The direction of historical change (polarity [or which came first]) in
a trait must be determined.
A parsimonious analysis should be used. (Use Occam's razor to cut
off unnecessary assumptions.)
C. Phylogeny and parental care in fish
D. Phylogeny, Mating Systems, and Male Aggression
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1. Males may benefit from being more aggressive when sexually
receptive females are present and less aggressive otherwise.
2. Nasonia wasps show high male-male aggression was the
ancestral state and within-host mating is a more recent
Proximate Factors, pages 72-117
I. Ultimate and Proximate Perspectives
A. A perspective is considered ultimate if it concerns how something
may have evolved. It might be a question that begins with, “Why is it
B. A perspective is considered proximate, if it answers a question
about “How is it that…?" or “What is it that…?" It operates within the
lifetime of an organism.
II. Hormones and Proximate Causation
A. The Long-Term Effects of In-Utero Exposure to Hormones
B. Stress Hormones and Spatial Memory in Rats
1. water maze
2. Corticosterone's effects on spatial memory are often
displayed about 30 minutes after stress is induced.
III. Neurobiological Underpinnings of Behavior
A. The Nervous Impulse
B. Neurobiology and Learning in Voles
1. Male meadow voles are polygamous, have a home range that
is 10 times that of females, and have better spatial skills than
2. Prairie voles are monogamous, have a home range that is
about the same size as those of females, and have spatial skills
about equal to that of females.
3. The size of the hippocampus in polygamous meadow voles
was larger in males than in females.
4. The size of the hippocampus in monogamous voles was the
same in both sexes.
5. Male meadow voles have more dendritic spines than females
in the prefrontal and parietal cortex that is associated with spatial
C. Sleep and Predation in Mallard Ducks
Mallards can sleep with one eye open and one hemisphere of the
brain awake while the other hemisphere is asleep. This is more
likely to occur on the edge of the flock.
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IV. Molecular Genetics and Animal Behavior
A. Ultraviolet Vision in Birds
B. Song Acquisition in Birds
V. Foraging in Honeybees: An Integrated Proximate Analysis
A. Mushroom Bodies and Honeybee Foraging
1. Parts of the brains of honeybees, called mushroom bodies,
are associated with spatial navigation and foraging.
2. Mushroom bodies were larger in foraging honeybees than in
bees that stayed in the hive.
B. Genes, mRNA, and Honeybee Foraging
1. The period (per) gene influences circadian rhythms.
2. Foraging bees have significantly higher levels of per mRNA
than younger, non-foraging bees.
C. Hormones and Honeybee Foraging
1. In honeybees, juvenile hormone is associated with foraging.
2. Removing the corpus allatum removes the source of the
3. This reduced foraging in honeybees.
4. Adding artificial juvenile hormone increased foraging.
5. Octopamine modulates learning and memory in honeybees.
6. Foraging bees have more octopamine in their brains than do
7. Increasing octopamine increased flight activity for foraging,
but not for removal of corpses.
8. Bees given octopamine and exposed to new, larger brood
increase their foraging behavior but not caring-for-brood behavior.