Animal Behavior Midterm 1

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Georges Romanes

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developed the systematic comparative method

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Kino Tinbergen, Konrad Lorenz, Karl von Frisch

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thought of behavior as instinctual- developed fixed action pattern way of thought: innate response triggered by a stimulus (ethological thinking)

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Konrad Lorenz

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studied imprinting in baby geese - discovered imprinting

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Niko Tinbergen

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noticed male three-spined sticklebacks responded aggressively to the color red

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Karl von Frisch

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discovered the meaning of dancing honeybees

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Psychological approach to animal behavior

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different from ethological because it was learned behavior, not innate

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Pavlov

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showed conditioned response with dogs

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BF Skinner

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saw behavior as learned and shaped by experience - operant conditioning

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WD hamilton

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contributed a genetic approach to behavioral evolution, hamilton's rule about kin selection,

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evolutionary biology approach to behavior

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Hamilton, Wilson, Maynard Smith

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EO Wilson

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invented sociobiology (understanding social systems across taxa), island biogeography,

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John Maynard Smith

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introduced game theory to evo bio, strategy of one individual depends on what the strategy of other individuals is

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Nature vs Nurture debate

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is it learned (psychology) or inherited (ethology)?

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behavioral ecology, what is it and what is it's approach?

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ecological and evolutionary processes can explain the occurrence and adaptive significance of behaviors. examines the costs and benefits of a particular behavior and its impact on reproductive success

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Proximate cause

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an immediate cause based on operation of mechanisms (How?) ie muscles etc - bluegill sunfish - breeding is triggered by increase day length = THE STIMULUS

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Ultimate cause

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The evolutionary reason why something is the way it is? (Why) - bluegill sunfish - breeding is most successful when water temperatures and food is optimal

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Tinbergen's four questions

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CDEF
1. What is the Cause?
2. How did it Develop within the lifetime?
3. How did it Evolve?
4. What is it's Function?

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Observation: different populations of sparrows sing different songs
Begs what question?

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Are these differences learned (environmental) or inherited (genetic)?

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How do you examine selection on a behavior?

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1. Why does the behavior occur?
2. What is the advantage of learning the behavior?
3. Why would within population variation exist?
4. Why would between-population variation exist?

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Natural selection

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trait value is positively associated with reproductive success

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Random drift

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No association between trait value and reproductive success

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Darwin said natural selection requires what?

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1. Variation in a trait
2. Trait must be heritable
3. Differential reproductive success
4. Variation in fitness is correlated with the trait

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How do we predict expected patterns of evolution by natural selection?

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1. population genetics
2. quantitative genetics

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Population genetics

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how do allele frequencies change through time as a result of the differences in survival of the possible genotypes?

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how do you calculate p and q?

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Front 26

why is quantative genetics different from population genetics?

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most behavior patterns are behavior is influenced by many genes (polygenic), so simple mendelian style population genetics doesn't really work (assumes 1 gene per trait).

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what is quantative genetics based on?

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it is based on the idea that the traits we observe are influenced by the environment, many genes, and interactions of the two

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how is phenotypic variation defined by quantative genetics

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Vp = Ve + Vg; environmental influence on variation plus genetic influence on variation

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what is heritability?

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the proportion of the phenotypic variation that is genetic

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what is additive genetic variation?

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the cumulative effect of the individual genes

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what is narrow sense heritability and how is it estimated?

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h^2 = Va/Vp = the cumulative effect of genes / the phenotypic variation = the proportion of the phenotypic variation that is genetic.

regression of offspring on parents can help to demonstrate if a trait is heritable, but it is not conclusive (environmental?)

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what is one way to predict evolutionary change using heritability?

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response R, selection S
R = heritability h^2 * selection;

heritability h^2 = R/S ;the proportion of the differences between parents that is transmitted

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Describe the difference between stabilizing, directional, and disruptive selection?

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How do you study the evolution of behavior?

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1. measure fitness (and heritability of a trait)
2. observe changes in wild populations over time
3. artificial selection in the lab
4. map traits onto phylogenies (comparative), to determine the characteristics of ancestral species

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norm of reaction

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all of the phenotypes that can originate from a phenotype (depending on the developmental context)

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Hamilton's rule

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when br - c > 0, be altruistic and select kin

c = the degree of modification to the actor's fittness
b = the degree of modification to the recipient's fittness
r = the degree of genetic similarity between the actor and the recipient

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Genetic approach

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1. gene knockouts create artificial genetic variation
2. can also study pre-existing genetic variation - figure out the genetic basis of different polymorphisms

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what is a QTL?

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a region in the genome that houses natural genetic variation in a quantitative trait

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what is QTL mapping?

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the study of the alleles which occur in a locus and the phenotypes that they produce
1. using genetic markers genetic variation is IDd; must also measure phenotypic variation
2. use stats to map which loci affect the phenotypic trait

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What are the goals of QLT mapping?

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1. How many genes influence the expression of a quantitative trait?
2. What is the level of influence of each gene?
3. where are the genes located on the chromosome?
4. What is the function of each gene?

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What is a candidate gene?

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"any gene that has been identified in one organism that is hypothesized to influence a similar phenotype in another organism"

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What does the candidate gene approach allow?

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It allows for the use of extensive information from well known species to understand less well known species

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Microarray

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examines the expression of many genes simultaneously; correlations of level of gene expression and behavior patterns can help to determine which genes affect specific behaviors

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Cichlids from African lake:
observations:
males have two phenotypes that are reversible and under social control

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many genes are probably responsible - using a microarray can help to narrow the selection

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Development of behavior

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behavior will vary across the lifetime of an individual.
1. depends on environment

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Developmental homeostasis

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"the capacity of developmental mechanisms within individuals to produce adaptive traits, despite potentially disruptive effects of mutant genes and suboptimal environmental conditions

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Polyphenism

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variation in environmental condtions leads to different phenotypes = proximate cause
ultimate cause = different phenotypes produced by different environmental conditions are more reproductively fit in the context of their environment

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the concept of optimization

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organisms through their behavior will maximize a certain valued linked more or less to fittness

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Phenotypic vs comparative vs genetic approach

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phenotypic: valid for populations in which the trait in question is fixed
comparative: reconstruction of evolutionary scenarios
genetic: can serve as a basis for a phenotypic approach and then

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Proximate mechanisms

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1. neurobiology
2. sensory cues and navigation
3. temporal patterns
4. hormonal influences

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monogamous vs polygamous prarie voles

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monogamous (pair bonded) have higher levels of oxytocin and AVP than polygamous. more receptors for oxytocin are present in monagamous brains

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why would understanding the proximate mechanism help to understand the evolution of a behavior?

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you need to know how these mechanisms affect fitness

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navigation

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examples.
experimental manipulation of magnetic fields affects the orientation of sea turtles and birds

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temporal patterns of behavior

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daily cycles (circadian rhythms), lunar patterns, annual variation (circannual rhythms)

cues can be internal or external, ie biological clock that can be adjusted by external cues

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side-blotched lizard

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three types of males exist; one is more aggressive and has more testosterone than the other two, but also lower survival - evolutionary tradeoff

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What does ecology examine?

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The distribution and abundance of organisms and the interrelationship between organisms and their environment.

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What does population ecology focus on?

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understanding changes in species abundance - population dynamics

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What does community ecology examine?

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understanding how interactions between species affect patterns of species diversity and abundance

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the general form for population dynamics

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N(t+1) = (B-D) N(t); B-D is the rate

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Effect of environment on behavior: abiotic factors

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Temperature, light, rain, oxygen, refugia, act as direct cues through their effect on growth and surfifal

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Effect of environment on behavior: biotic factors

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Food/resource abundance, competition within and between species, predation/parasites/disease

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How does behavior affect ecology?

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Distribution: Territoriality, dispersal, migration, habitat selection

Abundance: Survival, reproduction

Interaction with other species: competition, being food, finding food, mutualism

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What are the four types of studies?

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Theoretical, observational, experimental, and comparative

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What makes a good research design?

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1. Estimation of treatment effects
2. estimation of random variation
3. absence of bias
4. precision and accuracy in measurements
5. applicability
6. simplicity n execution and analysis

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type 1 error

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reject null and null is true

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type 2 error

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do not reject null and null is false

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types of behavioral measurements

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1. latency
2. frequency
3. duration
4. intensity
5. sequence

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issues to consider for behavior

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1. effect of observer
2. experimenter bias
3. anthropomorphism

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The two main types of models

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Genetic models: tries to predict evolutionary process
Phenotypic models: tries to predict the outcome of evolution

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Optimality theory

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there exists a value for a particular trait that optimizes the organism's fitness/ ex territory size:
small territory has less food but is easier to defend, large territory has more food but is harder to defend - evolutionary tradeoffs are everything

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Where do local maxima occur?

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A maximum occurs where dF(x)/dx=0 AND d2F(x)/dx<0

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what is game theory, when can it be used?

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developed as an economic theory, it tries to predict the optimal behavior, given that fitness depends on the actions of others in the population

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Evolutionarily stable strategy

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the strategy by which nobody else can do better (this is not the optimal solution - it maximizes the individual speed based on what others are doing ie, a checkout line at a supermarket - you have to wait on line, but it would be faster to skip the line)

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game theory notation
E(A,B) means what?

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the expected fitness gain to an individual adopting behavior A if the other individual is adopting behavior B

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Evolutionarily stable strategy

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a behavior is an ESS if no other strategy will have greater fitness when there is a population of individuals adopting the ESS

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When is strategy A an ESS compared to B

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when E(A,A) > E(B,A)

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expected fitness of a behavior A, Wa=

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Wo + pE(A,A) + (1-p)E(A,B), where Wo is the baseline, p is the proportion that exhibit behavior A, and B is the alternative behavior

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when does a stable coexistance occur?

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when both game theory strategies have equal fittness, in the hawk dove example, p = V/C

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according to game theory, the fitness of a behavior is dependent on what

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it depends on the frequencies of the other behaviors in the population

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How do you study foraging?

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Direct observation, stomach contents, chemical signatures

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what does the Optimal Foraging Theory account for?

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accounts for two measures of fitness; food intake (rate or total), and survival probability

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Marginal value theorem

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within the category of optimal foraging, the marginal value theorem tries estimate how much time an animal should spend in a patch

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Assumptions of marginal value theorem

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optimal foraging
1. assume they are taking food to their young
2. assume they have to travel back and forth
3. as they catch prey they become less efficient

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Marginal value theorem rate of gain = slope of the line =

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food gain/(travel time + search time)

as search time in a patch increases, the food gain rate increases until it reaches an optimum "marginal value," at which time it would be good to stop searching in this patch and start in another one

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Assumptions of the marginal value theorem

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predator has all the information necessary for a rational decision, knows average travel time, and expected quality of patches, prey are distributed randomly, predator cannot do better than random search

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Prey selection

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o Variables
• Amundance/search time for each food item (S)
• Handling time to process and digest (h)
• Quality of food (energy) E
• Assume animals maximize their rate of food intake
• Profitability E/h
• So if E1/h1 > E2/h2 just eat item 1 right?
• If there are two food items that differ in E, when should an animal ignore the lower energy gain item
• Gain for lesser item E2/h2
• Gain if searching for better item E1/(h1 + S1)
• When does E2/h2 < E1/(h1+s1)
o Assumes the items are in the same patch and that they sometimes encounter the lesser quality food item while searching for the higher quality food item
• (h1 +S1)/ h2 < E1/E2 = specialize on item 1
• (h1 +S1)/ h2 > E1/E2 = eat item 2 when they encounter it, but keep searching for 1

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Why is predation important?

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Because it changes the allele frequencies in the prey; with a mutation that even slightly decreases predation risk, that mutation is selected for and becomes prevalent in the population very rapidly

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list 5 Anti-predator behaviors

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1. decrease detection
2. escape more (ie larger numbers)
3. avoid predators (bluegill sunfish avoid the habitat with predators, red eyed tree frog escape hatching)
4. warn conspecifics (warning calls)
5. warn the predators (toxins and warnings, even deception when you are not actually dangerous to eat)
6. share the risk: attacks per individual decreases with group size, thus its selected for - hamilton
7. confuse the predators: schooling fish are harder to see and catch, zebra stripes maybe?
8. attack the predators with mobbing behavior, pygmy owl call can induce mobbing effect in birds
9.

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convergent evolution

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when the same behavior or trait arises in evolutionarily distinct lineages, vs divergent evolution, when you have similar ancestry but different traits/behaviors

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what behaviors does predation effect?

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possibilities = mate choice, foraging, paternal care, territorality, etc... everything..

different populations of the same species under different predatory pressures can differ in many traits - selection acts quickly

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Habitat selection; biotic vs abiotic factors

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abiotic factors; temperature, rainfall, oxygen, determines range
biotic factors such as disease, predation, conspecifics can also influence habitat selection -

pretty much everything is a factor in habitat selection

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Ideal free distribution

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Ideal: assume animals have perfect knowledge
free: assumes patches are not defended

if either of these assumptions is violated then predictions will deviate

gain per patch = resources / number of individuals

the ratio of the abundance of resources will predict the relative distribution of organisms - use as a baseline model

ducks example with bread
sticklebacks with different quantities of food in different parts of the tank

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Individuals rewards in a particularly selected habitat decrease with

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increasing competition. • The change from selecting rich habitat to low quality habitat can be caused by influence of competition in the higher quality habitat

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what are the reasons for dispersal

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chance to locate new mates, chance to find superior territory, no extreme inbreeding

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what are the reasons for not dispersing (remaining at the natal site)

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reduced exposure to predators, lower energetic expenses, no extreme outbreeding, no loss of familiar resources

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why is inbreeding bad?

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chances of getting deleterious alleles that are identical by descent are much higher

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why is outbreeding bad?

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can result in low or no fitness ie mule

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what makes male belding ground squirrels disperse? answer tinbergen's four questions

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AB CDEF
Cause: hormones
Development: body mass
Evoluton: strong male bias in dispersal is selected for
Function: reduce chances of incest

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migration patterns

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magnetic, smell, echolocation, landmarks, celestial, orientation to the sun,