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Logical Argument on Why Genes Influence Behavior
genes code for proteins used in structure or regulation, and proteins influence behavior
Genes are effected by ____
the environment (social & Ecological) and behavior
Behavior is effected by ____
genes and the environment (social & ecological)
Environment is effected by ____
behaviour
Evidence for Genetic Basis of Behaviour
Inbreeding, Artificial Selection, Hybridization, Molecular Genetics
Inbreeding
selective mating of closely related individuals. Reduces heterozygosity, different alleles become fixed in different inbred lines
used to explore the effect of one variable
Adam Miklosi
studied paradise fish larvae, research on antipredator behavior. Two inbred strains studied after 30 generations, larvae showed different predator recognition behaviors. Behavioral differences attributed to genetic factors
Heritability
a statistical measure of the extent to which a trait is influenced by genes. Ranges from 0 to 1 (0.5 indicates 50% of observed variation attributed
Artificial Selection
humans decide which animals will reproduce, choice is based on desired behavioural traits (physical or behavioral).
EX: domestic Dogs
Tolman
“bright” vs “dull” rats study:
Bright rats were good at navigation, dull rats were not
mated lines of Bright and Dull mice
found that “brightness” was not transferable
Lynch
studied normal mice vs. genetically altered mice that couldn’t build nests. Found that both genetics and environment impacted behaviors
Hybridization
Cross individuals with different expressions of the same underlying behavioral trait
eg: agressive x submissive
Complete dominance
the phenotype of the heterozygote is indistinguishable from that of the dominant heterozygote
incomplete dominance
the phenotype of the heterozygote is intermediate to the two homozygotes
Marla B. Sokoloskwi
Studied foraging behavior in fruit fly larvae. Two strains of foraging behavior: Rovers (move more in a patch) and sitters (move less), by crossing the two strains found that 75% of F2 generation was rovers, rover is a dominant allel.
Calling Behaviour in crickets
males produce call to attract females and are species specific. In the lab hybrids can be produced: hybrid males produce calls that are intermediate to parentals in frequency and duration, hybrid females preferentially orient to intermediate calls → shows genetic control with incomplete dominance
Molecular Genetics
enhances our understanding of the relationship between genes and behavior. Revolutionized in the last quarter century
mutagens
chemical, biological, or physical agents that alter DNA sequences
Quinn conditioned behavior in fruit flies
Studied test tubes of flies and odors, one odor paired with electric shock, flies were then averse to the stimuli. Shows associative learning
Recombinant DNA
precise experimental method for determining the effect of a particular gene on behavior. Once a gene is identified and located, it can be cloned and inserted into another organism’s genome, its effects can then be observed.
Circadian Rhythms in fruit flies
fruit flies show a 24 hour rythm, but some individuals are arrythmic viia a mutation. Gene therapy restores the 24 hour rythm
Social Behaviour in Voles
differences in the response to AVP are due to differences in the distribution of V1a receptors, small changes in these receptors’ genes will have a large effect on social behaviour.
Adaptive Behaviour
evolutionary framework for studying behaviour for non-adaptive and maladaptive traits via observation, comparative and experimental methods
Darwin’s Definition of Evolution by Natural Selection
a trait must exhibit natural variation
some of that variation must be heritable
must be a struggle for existence
different variants of the trait should confer different probabilities of survival
Adaptive Basis of Behaviour
behaviour has a genetic basis that is subject to natural selection
→assume behaviour is adaptive
Adaptation
a trait that allows its bearer to survive and reproduce better than competing individuals that lack the trait
Fitness
a measure of an individual’s relative success in propagation its genes into the next generation via the production of decedent kin that survive to reproduce
inclusive fitness
a measure of an individual’s relative success in propagating it’s genes into the next generation via the production of descendant and non-descendant kin that survive to reproduce → W.D. Hamilton
cooperative Courtship in wild turkeys
Less fit males team up with dominant males to improve mating chances by defending territory. Displays inclusive fitness as helping kin (brothers) boosts the shared genetic shared genetic success
Non-adaptive traits
a characteristic of an organism that does not improve its ability to survive or reproduce
maladaptive traits
a trait that is more harmful than helpful, preventing an organism from adapting effectively
non-adaptive traits and maladaptive traits are expressed due to ____
domestication
context-dependence
developmental anomaly
disease/pathology
parasitism (induced changes in behaviour)
Gene flow causing aridland spider maladaptive behaviour
? Gene flow from wetter regions introduces traits that are not suited for dry environments. These imported traits reduce survival or reproduction in arid conditions → gene flow can spread non-benifical traits when environments differ
Non-adaptive or maladaptive traits are velated via:
selectively neutral behaviours
conflict between natural and sexual selection (irish elk antlers were too big for traversal between trees)
linkage between loci
pleiotropic gene effects
evolutionary time lags
Coevolution
reciprocal adaptations in interacting species due to natural selection imposed on the other species →evolutionary arms race
Red Queen Hypothesis
Van-Valen: animals must constantly evolve to maintain the same state of adaptation relative to their environment
adaptive infanticide
Maladaptive in most cases, however it is also:
Resource Exploitation: reduced competition for resources
Resource Competition: decreases number of competitors
Parental Survival: increases parents chances of survival and future reproduction
Sexual Selection: allows mother to quickly become sexually receptive, enabling mating with a new male
3 empirical approaches to understanding the Adaptive Basis of Behaviour
Observational, Comparative, Experimental
Observational Approach
an empirical approach to understanding the adaptive basis of behaviour wherein, animals are observed in their natural setting to gain insight into the adaptive basis of behaviour
Tinbergen’s black-headed gull work
eggs shells in nest attract predators that eat chicks. The black headed gull will remove shells from the nest 30 minutes after → the delay appears to be maladaptive, however the delay is to reduce neighboring gulls from eating chicks (a wet chick takes seconds to consume, but a dry one takes minutes)
Comparative Approach
an empirical approach to understanding the adaptive basis of behaviour wherein members of the same species are studied in different environmental conditions
Barash’s study of social behaviour of marmots
studied several species of marmot social systems and growing season. Found that with increasing elevation and decreasing growing season length selects for increased sociality
Problem with comparative Approach
based on correlation
another variable may underline increased social behaviour (eg: predation rather than food scarcity)
coloniality and sociality impart antipredator benifits
Experimental Approach
experimenter proposes a hypothesis for the adaptive function of a behaviour. Involves manipulate some aspect of the animal or its environment to test that hypothesis
adaptive head colour in Gouldian Finches
has 3 head colours that coincide with domination in social interactions. Red dominated black, red + black dominate yellow. When head colour was manipulated it had no effect on dominance
Optimality Theory
accounts for costs and benefits of a trait. Evolved traits represent an optimal balance between benefits and costs
optimum point on a Graphical Model of Optimality
when benefit is high and cost is low
common measurements of Optimality
season reproductive success, as well as time/energy budgets
Territory
a fixed area defended against rivals for a period of time. Related to food, shelter, mates, and may change in breeding or non-breeding seasons
Benefits of holding a territory
guaranteed access to resources
enhanced acquisition of mates
reduced transmission of disease
reduced predation
Costs of holding a territory
increased energy/time expenditure
enhanced risk of injury or death (competition and predator risk via displays)
lost opportunity to exploit shifting resources
Resource attributes influence the economics of territoriality
resource abundance
resource distribution (space and time)
degree of competition for resources
Resource Abundance
a resource attributes influence the economics of territoriality that involves the correlation between less resources being available the less of a likelihood that a species will be territorial
Resource Distribution
a resource attributes influence the economics of territoriality that involves the incidence of territoriality increases with clumping of resources
Competition for Resources
a resource attributes influence the economics of territoriality that involves the increase of competition increases costs of defense, decreasing the incidence of territoriality
Relationship between Optimality and Territoriality
diminished benefits/increased costs may prevent the formation of territory however individuals can maximize net gain by optimizing their location/size/time that they hold a territory
Optimization in Rufous Hummingbirds
Defends territories around patches of flowers over the course of its winter migration. Accomplishes maximum energy gain via trial-and-error adjustment of territory size.
Frequency Independent Models
optima calculations are independent of frequency of behavior seen in other individuals. Recognize that fitness payoff of a given behavior is contingent upon what others are doing
Game Theory
models in which the optimum strategy is contingent on the frequency of behavior of others. Individuals ‘play’ against each other and “winning” is equated to fitness
Game Theory in Crickets
Two types of crickets: callers (produces signalling sound to attract mates) and satellites (hover around callers and intercept females). Callers have a higher success in mating but a higher risk of predation, while satellites have a lower success in mating but a lower risk of predation → strategies will reach equilibrium
Evolutionary Stable Strategy (ESS)
a set of strategies that, once adopted by a critical proportion of the population, cannot be replaced by other strategies
Nash Equilibrium
a stable state in a game where no player can improve their outcome by changing their own strategy if all other players' strategies remain the same
Origin of an Evolutionary Stable Strategy
each individual consistently plays one of the possible strategists so that the relative proportion of pure strategists in the population remain stable. Then each individual varies its strategy, playing each with a certain frequency.
ESS in Male Side-blotched lizzards
3 genetically determined colour polymorphs that each display a different reproductive strategy with same fitness payoff. The predominant colour fluctuates annually (Orange, then blue, then yellow then back to orange)
payoff matrix
formally states fitness payoffs to individuals playing all possible strategies
Hawks versus Doves
Hawks are aggressive; continue to fight until seriously injured or opponent retreats
Doves show aggressive displays, but always retreat rather than fight
Payoff Matrix Equations (A + B payoff relationship)
Payoff Matrix: A vs. B
A (Opponent) | B (Opponent) | |
|---|---|---|
A | (Resource value - injury cost) / 2 | Resource cost (if A wins) |
B | 0 (if B loses) | (Resource value / 2) - Display cost |
then tally from left to right row for fitness payoff
Payoff Matrix Equations (Proportion of A and B)
A → p[(resource value - injury cost)/2] + (1-p)[resource value]
B → p[0] + (1-p)[(resource value / 2) - display cost)
Where p is the proportion of A in a population
Asymmetries in Game Theory Models
Resource Hoarding Potential
Value of the resource
Arbitrary Asymmetries
Resource Hoarding Potential
an animal's inherent ability to win a physical fight if one were to occur
Game Theory Model Asymmetry in Funnel-Web spiders
females engage in territorial battles over productive web sites and employs conditional strategy: if an intruder is larger than the resident, the resident will exhibit aggressive displays but never fight. If the intruder is smaller, then the resident will be aggressive
Winner Effect
where wining a fight increases the probability of future wins
Loser Effect
Where losing a fight decreases the probability of future wins
Game Theory Asymmetry in Resource Value
One player has private information about the resource's value that the other lacks
Game Theory Asymmetry in Bowl and Doily Spider
A virgin female lays 40 eggs after fertilization, after 5 minutes there are 10 unfertilized eggs so the occupying male aggressivley defends against rivals. After 7 minutes there are only 4 unfertilized eggs so the male switches to a dove strategy (aggressive displays but not fighting)
→ shows Asymmetry in resource value
Game Theory: Arbitrary Asymmetries
asymmetries that are not connected to RHP or resource value. Rules or social conventions for setting disputes amoung conspecifics. Eg: prior ownership
Prior Ownership as an Arbitrary Asymmetry in Hamadryas Baboons
In mating: a male is perceived as “owner” of female if allowed prior association. This is respected even if intruding male in dominant.
In feeding: if no prior ownership of item is given dominant wins, if prior access is given the dominant does not challenge
Value of refined game theory models
enhance understanding of behaviour
organize empirical findings
generate testable hypothesis