Psychology 118, UCLA

Cathleen Cox, Ph.D, Instructor

TOPICS DISCUSSED IN LECTURE AND/OR INCLUDED IN TEXT
August 3 -12, 2015
PLEASE NOTE: THESE NOTES ARE TO HELP YOU. THEY ARE NOT INTENDED TO BE A COMPLETE
SUMMARY OF THE COURSE MATERIAL COVERED IN LECTURE OR IN THE TEXT (WHICH YOU ARE
RESPONSIBLE FOR). IN SOME AREAS THEY MAY BE INCOMPLETE, AND IN SOME AREAS TOO
DETAILED. And, a few concepts are included that have not been presented in class and are not
covered in your text or readings. These are here to provide additional information but you will
not be tested on such material.
Reasons why the behavior of animals is studied
intrinsic interest
to aid in the maintenance of wildlife
useful in generating hypothesis regarding human behavior
some animals are "simpler" than humans and therefore "easier" to study
some animals are shorter lived than humans so that effects of variables can be more
quickly and/or
comprehensively assessed
(diminishing) belief that it is okay to "sacrifice" animals
Animal behavior is an interdisciplinary field, including ethology, psychology, zoology, molecular
biology, anthropology, neurobiology, genetics
Origins of the discipline, with emphasis on comparing/contrasting the approaches of ethologists
and comparative psychologists
locations in which disciplines originated
how questions studied
where studies conducted
areas of study emphasized
types of questions asked
criticisms of each discipline
Tinbergen posed 4 main questions to pursue in the study of behavior:
How does the behavior influence an animal's ability to survive and reproduce?
How does the animal use its sensory and motor abilities to activate and modify its
behavior, e.g., what are the
immediate causes of behavior?
How does an animals behavior change as it develops, especially in response to
particular
experiences during maturation?
How does an animal's behavior compare with that of other closely related species, and
what
does this tell us about the origins of its behavior and the changes that have occurred
during
the history of the species?
The 1st and 4th questions ask why and take an ultimate approach
The 2nd and 3rd questions ask how and utilize a proximate approach, focusing on mechanisms
The example of changes in foraging behavior in the honeybee in chapter 4 and lecture
nicely illustrates the integration of all Tinbergins 4 approaches to the study of behavior

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Proximate and ultimate causes are interrelated. Genes present today have survived historical
process dominated by the effects of past differences among individuals in their reproductive
success. A living animal's phenotype then exerts proximate influences of the course of it's
development, affecting the nature of the mechanisms that the animal possesses, which in turn
enables it to do certain things.

When studying animal behavior:

Need to avoid anthropocentrism and anthropomorphism
Need to be aware of what the animal perceives
Need to be aware of the investigator's cultural biases
Importance of understanding experimental design and inferences and being able to
critically evaluate
information obtained
Need to consider alternative hypothesis and design tests that will discriminate between
them
Need to consider proximate (how) as well as ultimate (why) causes of behavior
Dugatkin defines behavior as the coordinated responses of the whole living organism to internal
and/or external stimuli and structures the text around 3 foundations of the discipline: natural
selection, individual learning, and cultural transmission. In Chapter 1 he briefly defines these
processes and gives nice examples of each.
Most of the studies in the field of animal behavior are empirical in nature. Conceptual
formulations serve to integrate previously disparate and unconnected areas. Theoretical
approaches utilize mathematical models to which serve to simplify variables and identify
relationships among them so that general predictions that are applicable across species can be
made.
EVOLUTION OF BEHAVIOR
NATURAL SELECTION - must understand it if you are to generate and/or critically evaluate
ultimate explanations of behavior. Should be able to describe observations and inferences made
by Darwin that were key in formulating natural selection.
Darwins formulation was presented in class
Dugatkin further simplifies by identifying 3 prerequisites: variation in the trait, fitness
consequences for the trait, and a mode of inheritance. Also, the resources must be
limited with respect to the trait being studied.
Just a small different in fitness consequences can result in a shift in morphology/behavior
over evolutionary time.

# of offspring that survive to reproduce relative to others in the

population
or
Lifetime reproductive success, usually measured in relative terms
Fitness -

In the introductory chapter kin selection is used to illustrate a conceptual advance in the field of
animal behavior. In the evolution chapter eusociality is introduced and the use of DNA
fingerprinting/molecular genetics in identifying the degree of relationship between individuals is
introduced.

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Gene - basic units of heredity

Chromosome - structure in cell nucleus in which genes are embedded; the genes/alleles on a
chromosome provide the code or "blue print" according to which proteins are
manufactured. These instructions analogous to "Read Only Memory" in a computer, e.g.
the instructions are copied but not modified. Most species have diploid chromosomes.
Genotype - complete set of genes of a particular individual
Gene Pool - hypothetical collection of all genes of all reproductive individuals within a population
Phenotype - physical and behavioral characteristics an organism possesses which result from
the interaction of the genes and the environment; natural selection operates on
the
phenotype rather than genotype
Alleles - alternate forms of a gene at a particular locus
Homozygous/Heterozygous - If an individual is heterozygous at homologous loci, the phenotypic
expression of the alleles may show that a particular allele is dominant, recessive, or
intermediate. Alternatively, the combination may result in overdominance, e.g. hybrid
vigor.
If and only if a behavior has a heritable component can it be shaped by natural selection.
Ways in which genes influence the expression of behavior in a proximate sense. Review the
relationship between DNA/genes and the development/expression of traits/characteristics.
Typically behavior is the product of interactions between genes and environment
Can only infer what differences in behavior result from genetic variation (substituting one allele
for another) when environment kept constant.
Genetic variation in a population can result from mutation, genetic recombination, and/or
conspecifics from a different population entering the population, e.g., migrating
An empirical approach to studying the effects of environment is to keep genes constant and
vary the environment, for example as can be done in cross-fostering studies.
An empirical approach to studying the effects of genes on behavior is to keep the environment
constant and vary the degree of genetic similarity (cross-fostering as well as twin
studies: identical vs fraternal, separations, etc.)
Other approaches that illustrate the effect of genes on behavior include:
single gene effects - examples & reasons so infrequently seen (hint: polygeny)
S/s alleles in ruffs and male mating type, coloration, size
sickle cell anemia: a lethal condition which is only expressed when individuals are
homozygous for recessive sickle cell allele; individuals who are heterozygous,
with a normal allele and a sickle cell allele have greater resistance to malaria than
those who are homozygous for the normal allele.
Avpr1a allele variation and degree of male parental care in prairie voles

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Knocking-out particular genes - examples given in text include Neofem2 gene in

termites, FOXP2 in zebra
finches
artificial selection.
Pleiotropy: alleles often influence more than one thing!
Polygeny: several genes at different loci may contribute to the development/expression of a
behavior . When searching for the genes underlying polygenic traits/characters investigators
are looking for quantitative trait loci/QTLs
Heritability (in the narrow sense): a statistical measure that represents the proportion of
phenotypic variance in a study population that can be attributed to genetic differences among
individuals over total phenotypic variance which results from both environmental and genetic
differences.
Truncation selection experiments as well as parent-offspring regression can be used to measure
heritability in the narrow sense
Intelligence and its heritability
Some traits are relatively plastic such as male morphology in plainfin midshipman fish, others
are not, need to look at adaptive significance of flexibility ..........
The most parsimonious explanation of the evolutionary process is that changes occur in small steps. When considering
the evolution of a modern trait from a previous trait, the pattern that takes the fewest transitions/changes between
intermediate steps is the most likely.
the cumulative effects of small changes may be major changes
changes are layered, that is, new changes are made on top of previous changes
over time both the form and function of characters may change
phylogenetic tree/cladogram
Phylogenetic tree: depiction of the evolutionary history of a group of related organisms with branching tree-like
diagrams
Historical/Comparative approaches are useful in inferring evolutionary pathways leading to the development of
particular structures and behaviors including signals used in communication
Homologous: Two phenotypic traits of similar or different character expressed by different species are homologous if
they are derived from the same ancestral characteristic.
Example: wings of a bat, flippers of a dolphin, arms of a human
Analogous: Two phenotypic traits of similar character expressed by different species are analogous if they are derived
from different ancestral characteristics, but now serve a similar purpose or character. Dugatkin uses the term
homoplasy.
Example: wings of a bat and wings of a butterfly
Example: Pandas thumb is not homologous to a primates thumb, but it is used for a similar purpose, that is, it
is an analogous structure. It is actually derived from the wrists radial sesamoid bone and is well adapted for the
purpose it now serves.
Sometimes different forces act on closely related species which may result in divergent evolution, or similar selective
pressures act on very different species leading to convergent evolution.
Divergent evolution: The evolution of differences among closely related species because of differing selection pressures
in their different environments.

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Convergent evolution: The independent acquisition over time of similar characteristics in two or more unrelated species
subject to similar selection pressures.
Adaptation: A characteristic that confers higher fitness on individuals than any other existing alternative exhibited by
members of a species in a particular environment Adaptations are typically the result of the process of natural selection.
Constraints on Adaptive Perfection/Reasons why some behaviors in some species may not be adaptive

Failure of appropriate mutations to occur recent changes in environmental conditions render present form of a
behavior/characteristic maladaptive. This can result from a number of situations:
- The behavior/character trait evolved under previous conditions that favored those who carried the trait.
Under current conditions that trait has a maladaptive effect but has not yet been replaced by a more
adaptive mutation.
- The behavior/character did not appear under previous conditions, but under the current change in
conditions the trait has emerged and is maladaptive. There has not been sufficient time for natural selection
to act upon those bearing the trait.
- The behavior/character is constrained by previously evolved structures, e.g., phylogenetic inertia

Pleiotropy the allele underlying a less than optimal behavior may also be responsible for a positively selected
character

Co-evolution - evolutionary stability may never be reached as each species is changing in response to selection
pressure imposed by other species (predator/prey, host/parasite relationships).

PROXIMATE APPROACHES
Research on the coloration of male house finches is used to illustrate the utility of both
proximate and ultimate approaches
Hormones, neurobiology, molecular genetics and development are 4 different research areas
used to investigate the mechanisms involved in the expression of behavior
ENDOCRINE SYSTEM/HORMONES
Example:Sex differences stem from the sex determining chromosomal mechanisms
which are responsible for the development of testes/ovaries which produce different
hormones. Hormones present early in developmental may have an organizing effect and
act as developmental switch which determines the subsequent path of development. At
a later age the same hormones may have an activating effect. The HPA axis & stress as
well as the role of vasopressin and oxytoxin in sociality & parental care is covered in the
text.
NERVOUS SYSTEM
Neurons rapidly receive and relay messages within an organism. In species living today
neurons that serve specific functions are clustered. Over evolutionary time there has
been a tendency for nervous systems to become centralized. Areas of study include
how variations in the nervous systems of different but related species impact learning
and how differences within a species impact behavior.
Sleep/unihemispheric sleep is an adaptation used in a number of species to minimize
potential predation. How this is achieved is studied by neurobiologists
MOLECULAR GENETICS

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The effects of an allele can be visualized using molecular genetics. The text presents the
approach in investigating vole sociality, UV vision and song learning. The diminishing
response/habituation of zenk mRNA to song playback of longer duration seen at the
molecular level appears to underlie behavioral habituation in zebra finches.
DEVELOPMENT/ONTOGENY/EPIGENESIS = the process of going through the stages that lead to
maturity
Development involves complex interactions between genotype and environment. The
development of any trait is the result of an interaction between the genotype of a
developing organism and its environment (both abiotic and biotic), which consists not
only of the food it receives and all metabolic products produced by its cells (the material
environment), but also its sensory experiences (the experiential environment). The
value of genetic information lies in the ability of genes to respond to signals from the
environment by altering their activity, leading to changes in the gene products available
to the developing organism. Environment includes in utero hormonal exposure as well
as the timing, type, and level of sensory and social surroundings following birth/hatching.
When a gene is expressed or turned on it produces particular enzymes. In turn, the
enzymes produced may act to turn on different genes.
Some traits/characters are plastic while others are fixed. Need to assess the
evolutionary consequences of flexibility in particular traits.
A species with distinct alternative phenotypes is said to be polyphenic. Polyphenisms can
be induced by
Social factors
Presence of predators
Food available
In a particular phenotype the order in which behaviors are first seen is quite predictable.
Some changes behavior look like they are a result of practice or learning but actually
reflect maturation
RESEARCH IN DEVELOPMENT includes assessing the effects of deprivation as well as
enrichment as well as the consequences of maturation and practice. In some cases
sensitive or critical periods have been identified which can dramatically affect
subsequent abilities (vision in cats, ovipositing behavior in wasps). Deprivation may also
impact social behavior including song learning (in birds), social relationships, maternal
care, mate preferences and kin recognition.
One classic phenomenon that involves a critical period is imprinting; two forms of
imprinting have been identified, filial and sexual. The strength of the imprinting has
been shown to increase with the effort expended in following the animal or object the
youngster is focusing on.
Despite the complexity of some behaviors and the variety of environmental conditions in
which these behaviors may develop, there are few individuals which fail to develop fully
functional behavior. Indeed, no one develops under exactly the same genetic and/or
environmental conditions. The capacity of the developmental process to buffer itself
against potentially disruptive genetic and experiential influences is called developmental
homeostasis.
LEARNING
Phenotypic plasticity: ability of organism to produce different phenotypes as a function of

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environmental conditions
includes having different morphs
includes having different behaviors which may result from learning
Learning is a relatively permanent change in behavior as a result of experience
Types

of experience that can lead to learning

Single stimulus
Stimulus paired with stimulus
Behavior followed by consequence, e.g. response-reinforcement

Learning in a single stimulus situation

arbitrary item placed in rat box, rat may look
If over time become more likely to look when placed in box we have sensitization
If over time become less likely to look when placed in box we have habituation
Both sensitization and habituation are changes in behavior and, if they are relatively
permanent, they are forms of learning
Learning in a 2 stimulus situation
US elicits UR
CS paired with US and results in CR which is Pavlovian/classical conditioning
If US is positive/pleasant it is an appetitive stimulus,
If US in negative/unpleasant it is an aversive stimulus
If US predicts occurrence of a 2nd event the relationship between events is
positive and excitatory conditioning may occur
If US predicts that a 2nd event will not occur there is a negative relationship
between events and inhibitory conditioning may occur
2nd order conditioning can result in blocking, overshadowing and/or latent inhibition
Learning when behavior is followed by consequence, e.g. response-reinforcement is known as
operant or instrumental conditioning
Behavior followed by + or reward
Thorrndike/Law of effect
Superstitious behavior
Hard to know what is going on in uncontrolled situation (but, thinking about what
learning processes could be involved is useful)
Initially but incorrectly thought that any animal could learn any association. Is important to
think about a species situation/habitat/survival in the wild and how selection would be expected
to act
Garcia used light/noise paired with water as well as taste paired with water as cues to
study learning
Rats could learn to avoid consuming water paired with taste when followed by
radiation or nausea/toxin but could not form this association when water was
paired with noise/light
Rats could learn to avoid shock when water paired with noise/light but not when
paired with taste
Taste aversion learning
Facial recognition (wasps)
How long should what is learned be remembered? Extinction curves

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Comparison of different populations of same species living under different ecological conditions
Conditions under which the ability to learn would be advantageous. Stephenson s model
predicts that ability to learn will be favored/selected in species where individuals inhabit
relatively predictable surroundings during their lifetimes but that predictability in surroundings
is low between generations
Learning can prove very valuable in migration/homing, foraging, (spatial memory for where food
is to be found/hidden), predator avoidance, kin recognition, species identification, locating
sexual partners, winner/loser effects.
Stress hormones negatively impact learning in rats.
In rats selected for high and low avoidance of shock Zhang explored the hippocampus for
differential expression of 7500 genes. He found the eight genes were differentially expressed in
the high vs low avoidance lines and uses this finding to support the idea that avoidance learning
is polygenic and each contributing gene contributes just a small amount to the phenotypic
expression of avoidance.
CULTURAL TRANSMISSION
Transfer and acquisition of information from individual to individual through social learning or
teaching
Information can be transferred within and across generations
Patterns of behavior within groups can change quickly much more quickly than would occur by
natural selection
As a result of cultural transmission different populations of same species may behave
differently/have different customs
Cultural transmission is well documented in primates\
Japanese macaques potato and grain washing, stone play
Capuchins - Citrus/leaf wiping, reduces insect bites
Chimps vocalizations & gestures & tool use
Cultural transmission occurs in social situations, but not all changes in activity in social
situations are a result of cultural transmission
In social situations there are models/demonstrators/tutors and observers but not all
changes in the observers behavior are the result of social learning
Local enhancement: a model may be engaged in a particular activity and a particular
location; observer is drawn to the location & engages in the same activity but doesnt
learn it from model p.171. In glossary Dugatkin is less detailed a phenomena in which
an individual is drawn to a particular area because observed another individual at the
location. To summarize this somewhat disparate information, we will say that local
enhancement does not involve social learning but may or may not involve self-learning
Social facilitation: presence of model, regardless of models behavior, facilitates learning
by observer
Cultural transmission/social learning: occurs when observer learns a particular behavior from
model

Copying

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Observer repeats what model did

Copier is often rewarded
What is repeated need not be a novel activity
Need not involve learning a new topographical manipulation
Documented for osprey/foraging, rats/novel foods preferred, guppies/mate
preference
Imitation
Acquisition of a new behavior & behavior must involve making new topographic
manipulation
Goal must be reached
Documented in parakeets learning to lift cover off a dish, pigeon learning to
get rewards by executing particular patterns of movement

Mirror neurons likely underlie imitation

The same set of neurons, mirror neurons, fire when a monkey performs a simple
goal directed interaction and when he sees another perform that action
The neurons may encode templates for specific actions and comprehend the
same actions when performed by another
Are spread across pre-motor and parietal cortices
mirror neurons play a role in under-standing an action
evidence: they fire when behavior heard rather than seen
evidence: they fire when monkey cannot see action but has sufficient clues to
create a mental
representation of it and they do fire when the monkey can see the experimenter
moving hand to pick up and object but not when they see the same hand motion
without the final pickup of the object. When the final action blocked by a screen
but the monkey knew whether or not there was a object behind the screen the
neurons still fired when the monkey viewed the hand moving towards the point
which was where the monkeys vision was blocked, but not as strongly when there
was no object behind screen
Appear to direct the immediate comprehension of others actions
Appear to provide the ability to simulate anothers topographic actions, e.g.,
imitate
Known to exist in humans, non-human primates and birds

Do animals actively teach how to behave?

Both natural selection and cultural transmission influence behavior over generations
On the Galapagos both natural selection and cultural transmission act to minimize
chances of interbreeding between the medium ground finch and cactus finch
In the laboratory mate choice in guppies has been explored when natural selection and
cultural transmission exert opposing forces
Both primates and bird species that have relatively large brains show a higher frequency of
innovative behavior, innovative behavior that functions well has a high likelihood being cultural
transmitted. A survey of more than 600 bird releases shows that the species which were more
innovative in the feeding techniques were more likely to be successful in the habitat into which
they were introduced.