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proximate causation
what causes behavior to occur
sensory motor (proximate)
immediate occurrence in behavior thru internal/external stimuli
ex of sensory motor (proximate)
psychology
endocrinology (short-term, activational effects of hormones)
neurobiology (changes/effects of neurotransmitters)
ontogenetics (proximate)
behavior development
ex of ontogenetics
genetics
instincts & learning (experience)
endocrinology (long-term, organizational effects of hormones)
ultimate causation
what causes behavior to evolve
functional (ultimate)
how adaptive behavior (if so)
ex of functional (ultimate)
costs & benefits
fitness (survival, mating success)
phylogenetic (ultimate)
historical pathway that behavior evolved from
ex of phylogenetic (ultimate)
evolutionary origins, transitions & precursors
natural selection
proximate mechanisms are the result of …..
ultimate (evolutionary) proceses
functional hypotheses do ….
not imply that animals understand that animals understand the fitness consequences of their actions
cortical magnification
proportional representation of touch receptors in the somatosensory cortex, as determined by touching different body parts while recording neural activity in the cortex
stimulus filtering is ..
a fundamental and adaptive property of the nervous system
onstant bombardment of potential stimuli necessitates filtering out irrelevant stimuli
mech of stimulus filtering
receptors tuned to respond to particular bandwidths of particular stimuli
interneuron relay receptor signals to ganglia
ganglia integrate and process inputs
ganglia send signals to brain or directly to muscles
hormones come from…
endocrine gland secretions
hormones are
chemical messengers
hormones are transported by
blood
hormones are _____ than neurotransmitters
slower-acting
hormones affect …
tissues with hormone receptors throughout the body
ovaries produce
estradiol and progesterone
testes produce
testosterone
modes of hormone action/effects on behavior
organizational
activational
not a true dichotomy; two modes grade into one another
organizational hormone action
long-term, developmental effects
activational hormone action
short-term, modulatory effects
organizational effects of hormones set the stage for…
later activational effects
chromosomal sex determination
females (XX)
males (XY)
gene on y chromosome codes for protein that causes …
testes instead of ovaries
what mode of hormone action is this?
gonadal hormones trigger sex differences in body and brain development
organizational
what mode of hormone action is this?
gonadal hormones have sex-specific effects on adult behavior
activational
primary function of gonadal hormones
regulate sexual development and reproductive readiness
activational effects of gonadal hormones vary by
species
mech of endocrine disrupting chemicals
mimic or block endogenous hormones
increase or decrease hormone levels
effects of endocrine disrupting chemicals
feminization or de-masculinization of males
masculinization of females
changes in sexual preferences and mating behavior
changes in parental car
sterility
sex change
genotype
set of genes possessed by an individual OR
the alleles possessed by an individual at a specific genetic locus
phenotpe
all characteristics of an individual other than its genotype OR
a specific characteristic of an individual other than its genotype
mendelian trait
a phenotype that varies among individuals because of variation at a single genetic locus
polygenic trait
a phenotype that varies among individuals because of variation at more than one genetic locus
is tongue rolling a mendelian trait
no, there is a strong genetic basis
“gene for trait X” is shorthand for…
“variation in the DNA sequence at this locus influences the likelihood of developing trait X”
the development of most traits is influenced by numerous genes and environmental factors. how can we sort out the relative influence of genetic and environmental factors?
quantitative traits, frequency histograms, and variances
equation for quantitative genetics
Vp = Vg + Ve
where V = variance, G = genetic, E = environmental
equation for quantitative heritability
h² = Vg / Vp
where h² = heritability, G = genetic, P = phenotypic
what kind of measure is h²
population measure
when looking at a graph where offspring mean (y variable) is influenced by parental mean (x variable), h² (heritability) is the …
slope
one solution to common environment problem
cross fostering (ex):
human twin adoption studies
nest-swapping experiments in birds
R = h²S
heritability can be used to predict evolutionary response to selection
h² = R/S
heritability can be estimated from response to selection
what does the S in h² = R/S mean
the selection differential S is the change in the trait mean caused by selection within the parental generation
what does the R in h² = R/S mean
the response to selection R is the change in the trait mean between generations
what is instincts
innate behavior - behaviors that do not have to be learned (and which are usually triggered by specific types of stimuli)
what is learning
process through which experience changes an individual’s behavior
durable and usually adaptive change in an indivudal’s behavior traceable to a specific experience
does not include changes in behavior caused by maturational growth processes, fatigue, or sensory adaptation
ex of innate behavior
gull chick pecking red spot on parent’s bill
what is habituation
animal learns not to respond after repeated encounters with a benign (harmless) stimulus
one of the simplest and most universal forms of learning
adaptive value: conserves energy, time and attention for more important activities
imprinting
forms of learning that depend on exposure to key stimuli during sensitive developmental period and which affects subsequent responses to those stimuli
filial imprinting
(ex) goslings learn the shape/sight/sound of '‘mom” and follow her
sexual imprinting
(ex) goslings later attempt to mate with someoen similar to “mom”
home stream imprinting
(ex) salmon learn the smell of the stream in which they were born; later home back to the same stream to spawn
social learning
catch-all category for forms of learning in which animals in some sense learn from each other
observational learning / imitation
basis of traditions and culture
development of behavior is influenced by
numerous genes and environmental factors
the proportion of genetic variation in a trait can be inferred from
breeding studies and the response to artificial selection
genetic differences between populations can be inferred from
common garden experiments
what animals are capable of learning can generally be explained in terms of their …
ecology
instincts may be favored over learning when conditions are …
predictable or when there is no opportunity to learn
which behaviors are innate versus learned matters in…
conservation translocation programs
what led darwin to natural selection
effects of selective breeding → also known as artificial selection
the “struggle for existence” → based on malthus (1798)
darwin’s crucial insight
any trait that gives the individual bearing it an edge in the struggle to survive and reproduce would be more likely to be transmitted to future generations
evolution by natural selection requires:
variation among individuals in
heritable traits
that affect survival and reproduction
if these conditions hold (generally do), evolution by natural selection is inevitable
fitness
the relative number of gene copies contributed by an individual to the next generation - usually measured in reproductive success (number of offspring) relative to the population average
adaptation (process)
changes caused by natural selection leading to greater fit between a population and its environment across generations
adaptation (trait)
a product of natural selection
individual selection
caused by differential survival and reproduction of individuals
group selection
caused by differential survival and reproduction of groups (colonies, populations, species, etc)
altruistic behavior
one that helps others but reduces the individual’s own fitness (their chances of surviving and reproducing)
can altruistic behaviors evolve “for the good of the species”
altruism can’t evolve just “for the good of the species” at the individual level, but Wynne-Edwards argued that if groups with altruists do better overall than groups without them, then group selection could allow altruism to evolve.
wynne-edwards view about group selection
group selection usually trumps individual selection
william’s view about group selection
group selection is theoretically possible but usually much weaker than individual selection
reproductive restraint and other forms of true altruism are illusions
conclusion: “group level adaptations do not, in fact, exists”
is it altruistic to protect your own offspring
no, altruism occurs when it negatively affects one’s self (cost and benefit)
is selection at the colony level required to explain eusociality
no, selection at the colony level is not strictly required to explain eurosociality
eusociality
highest level of social organization found in animals; defined by:
cooperative brood care
reproductive division of labor (queen vs workers)
overlapping generations within a colony
what is the multilevel selection theory
explains evolution by considering that natural selection operates simultaneously at multiple levels of biological organization, such as genes, individuals, and groups
adaptationist approach
starts assuming animals are well-adapted to natural environment
comes up with hypothesis to explain why particular trait has evolved through natural selection
collects data to tests predictions made by hypothesis
if data not match predictions, modify hypothesis
adaptive behavior
behavior that increases the fitness of the individual performing it, relative to a nearby alternative
maladaptive behavior
behavior the reduces the fitness of the individual performing it, relative to a nearby alternative
possible reasons for maladaptive behavior
environmental change and evolutionary time lag
parasites exploiting or controlling host behavior
gene flow between populations under different selection regimes
genetic and developmental constraints
genetic correlation between sexes
behavior evolves thru the same processes as ….
other organismal traits (morphology, physiology)
natural selection can operate at …
different levels (gene, individual, group, population, etc)
there is no reason to invoke group-level benefits for…
traits that increase for the individual (copulatory suicide)
the behavior of animals in the wild is usually…
adaptive, not withstanding some interesting exceptions
most instances of maladaptive behavior can be explained by…
recent changes in the environment (evolution takes place on a multi-generation time scale)
what is needed for group selection to work
suicidal populations would have to colonize areas where non-suicidal populations went extinct faster than non-suicidal genotypes take over suicidal poulation
why has copulatory suicide evolved in redback spiders
suicidal males fertilize more eggs → genetic standpoint: copulatory suicide appears to be selfish, not altruistic
evolutionary arms race between predators and prey
prey continually evolving defenses to avoid being eaten
predators are continually evolving counter defenses
economics of prey defenses
net prey profitability = E/T (energy per unit of time)
E = energy content per prey item
T = time required to harvest one prey item
T = S + h
S = search time
h = handing time
stages of predation
Encounter (within detection distance)
Detection (as objects)
Identification (as prey)
Attack
Subjugation (preventing escape)
Consumption
what prey defenses are effective at stage 1: Encounter, to increase “search time”
hiding
inactivity when predators are active
flight before detection
what prey defenses are effective at stage 2: Detection, to increase “search time”
immobility
crypsis (background matching)
background preferences
polymorphism (if predator forms search images)
background-matching movements
communication on ‘private wavelengths’
crypsis (background matching)
color patterns:
resemble sample of visual background (as viewed by predator) are cryptic
contrast with visual background are conspicuous
slight degree of crysis increase detection time
usually have both behavioral and morphological components
cryptic prey often come in multiple forms. what maintains variation"?
polymorphism - existence of multiple forms or morphs within a population
term used if known that variation doesn’t have genetic basis
apostatic selection
type of negative frequency dependent selection
occurs whenever predators don’t attack prey morphs in proportion to their availability but instead prey more on more common morphs
favors polymorphism in prey because rare morphs have survival advantage