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Animal Behavior


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Bluegill Sunfish (Sperm competition) 

Paternal, sneaker, and satellite (F faker) M forms. Hereditary and environment determine strategies.
Adaption (3)

hereditary trait that has:

  1. spread b/c of natural selection
  2. is spreading relative to other traits (b/c of ns)
  3. conferred/confers greater fitness than other traits
"A hereditary trait that exists because of
Adaptive Behaviors
positive (benefit-cost>0)
Adoption (Gulls)
Foster gulls more likely to accept closely related chicks. Chicks choose nests where the have a good chance.
Territory and reproductive success. Need big leaves, best to be alone on stem.
Apprentice Birds
Mutualism, with long delay. Inherit territory and Fs.
High ranking M monopolize F better; but F spend time w/ low-ranked M!
Bats and maternal care
F bats can recognize offspring, feed them most.
  • Dance
  • Behavior not mandated by genes
  • von Frisch
  • i.e. Nurse to forager
Belding's Ground Squirrels
"Kiss" to smell kin. Alarm calls from F favor F kin (GENETIC)
Biological Clock/Circadian Pacemaker

Neurons or cell groups that generate circadian rhythm. No cues needed to run, but cues needed to synchronize the clock.

Temperature compensation (period independent of body temp)

Mechanism is negative feedback w/ delay 

Males co-opt aggresive Skraa call (fitness proxy) to attract F
Brood Parasitism
eggs laid by female into nest of another. chicks/eggs look the same... or chicks are bigger (get fed better!)
Brood-Parasitism: Co-evolution
Cuckoos must look like eggs, look like chicks, and sound like chicks to fool fairy-wren
M feats of strength!
Central Nervous System

CNS generates/controls animal behavior

Nature/Instinct and Nature/Learning

evaluation, interneurons, descision 

Challanges w/ Human Behavior
  • Circus animals (genes+environment!)
  • high variation
  • no close relatives (vervets?)
  • effect of culture (sex roles)
  • most evolution coevolution... with humans!
  • objectivity impossible
  • predict
Charles Darwin
Origin of Species (1859)
Chase-away Selection

F not interested in trait (confers decreased fitness) so M increasingly elaborate display of trait

M display has high costs and low benefits 

F lays orange eggs and puts them in her mouth. M orange anal spots
Circadian Rhythm
Innate clock w/ period of about 24 hrs
Group with common ancestor.
Code Breaking:
Blue butterfly + Ants + Parasitic Wasps
Compasses (5)
  1. Sun position
  2. Winds
  3. Ocean currents
  4. Celestial ('cept North Pole)
  5. Polarization of Sky
*bold = require time compensation
Conditions for Infanticide (4)
  1. F ovulate soon after death
  2. Competition for F by M
  3. F smaller/weaker
  4. M can indentify offspring
Edward O. Wilson: Interconnection b/w all aspects of knowledge ("molecules to motivation")
Correlational studies
compare blood hormone levels w/ behavior
Cortical Magnification
critical inputs get more space
Cost-Benefit of abandoning chicks
  1. Chance to make more chicks (reproductive rate and length of life)
  2. Availability of new nest sites
Whales sing, hamsters make a smell, manakins dance.
Crested auklet
Crest=dominance. F chooses. EVEN IN AUKLETS WITHOUT CRESTS! Same as swordtail.
Cuckoo begging
call same as FOUR warblers.... parents bring 4X as much food.
Daniel Lehrman
Influence of hormones on dove reproduction (ENVIRONMENT [NURTURE])
Developmental Homeostasis vs. Dev. Switch
  • Genes code for developmental sensitivity
Less competiton for food, less inbreeding. One sex only. Birds usually F, mammals usually M.
Divergent vs. Convergent Evolution

Divergent: Diff selection -> diff behavior (even though common ancestor)

Convergent: Same selection -> analogous behavior (even w/out recent common ancestor) 

Donald Griffin
Bats and ultrasound
Ebo Gwinner
Circannual Rhythms (bird migration and navigation)
to set biological clock by zeitgeber (synchronize/regulate behavior)

Systematic study of animal behavior, animal behavior is innate, instinctive, fixed (GENETICS [NATURE])

  • Releaser/Sign Stimulus
  • FAP
  • Stimulus filtering (feature extraction) 
Eusocial Behavior

Groups that contain specialized non-reproducing castes that work for the reproductive members of the society. Helpers are obligate altruists.

Also... if helper has low probability or reproductive success

Relatedness requir

Evolution of intraspecific communication
  • COEVOLUTION: simultaneous evolution of transmitter, receiver, andcode (Bat/Moth, Wasp/Orchid)
  • SENSORY EXPLOITATION: transmitter and receiver exist, co-opt for new code and message
Evolutionarily Stable Strategy
Strategy used by all that can't be replaced by mutation`
  • Inherited
  • Can be modified by learning
  • not ALWAYS activated by sign stimulus
  • Specific behaviors variable
  • multiple FAPs possible from same SS
Female Firefly

Decodes M signal, fakes flashes, eats M


Finding Food
Leaves and poison, seeds in thorns, bald eagle, minke whale
Fisher's comment
2 brothers=8 cousins

measured by number of offspring/gene propagation (= reproductive success)

Proxies are indirect measures (eggs, color of feathers, song...) 

Foraging and Defense (RESOURCE POLYMORPHISM)
Sunbirds don't defend when lots of nectar around.
elimination ends nurturing behavior in mice
Free-running period (tau)
Time for animal to go through 1 cycle
Fry v. Eggs
Fish can "smell" if fry are another fish's.
Fundamentals of Science (6)
  1. Observation
  2. Hypothesis
  3. Prediction
  4. Experiment
  5. Theory
  6. Repeatability
Genetic Compatability
George C. Williams
Modern evolutionary biology. Darwinian approach to humans, evolutionary psychology. TRADEOFFS

Necks for dominance and sexual competition


Good Genes Hypothesis

M starling: song indicates healthy immune system

Peacock: offspring grow faster w/ good tail

Bowerbirds: parasites, body size, feather growth. Bower signals for young F, body for old, 

Group Selection

when some populations have a greater chance of survival... but cheaters win!

kin selection is more likely (group of altruists beats any single cheater)

Groups are safer
Orange food. M are orange.
Hangingfly, Redback Spider (Nuptial Gifts)
M bring food gifts, Spider IS food gift
Isolation and monkeys :(
Health Effects of Promiscuity
Primates have increased WBC with multiple sexual contacts
Ravens. Yelling attracts conspecifics to food. Single ravens call in help to defend against mating pair!
Helping Behavior (5)
  1. Mutualism--all benefit
  2. Reciprocity--delayed direct fitness gain
  3. Faultative Altruism--temporary loss, delayed gain
  4. Obligate Altruims--only indirect fitness gain
  5. Altruism--direct fitness loss, indirect gain
Hierarchy and Local Control
Brain prevents expression of incompatible behaviors
Home Range v Habitat v Territory

HOME RANGE: geographical area animal occupies

HABITAT: HR environment

TERRITORY: Area defended 

Honest Signals
Some birds (hunger), toads, red deer, antlered flies
  • Slow acting
  • Peptides or steroids
  • act in brain neuron... modify behavior
  • Secreted by glands and CNS, distributed by blood
Hume's Guillotine

What is =/= what ought to be

description cannot lead to prescription 

Hunting and Escaping
Orchid is predator, killer whales ambush sea lions
Lots of eusocial behavior, because relatedness varies from 1 to .5! (haplodipoids)
Illegitimate Receivers

Bats and Frogs... Great tits (high pitch can't be heard) and hawks...

Can detect signals of prey 

Inclusive Fitness
Direct Fitness PLUS Kin Selection [Indirect fitness] (use coefficient of relatedness)
Decision, activation, pattern
Intrasex Competition
compete for mates
Intrasex Selection
limiting sex chooses behavior and morphology
J.B. Watson
Behaviorism: all animal activity result of conditioning
Jacana (Polyandry)

F defend harem of up to 4 M. M incubate and care for eggs.


Phalarope: F abandons M and eggs 


John Maynard Smith
Game theory
Jurgen Aschoff
Human circadian rhythms
Karl von Frisch
Bees and dancing.
Kenneth Roeder
Evasive tactics (cricket) and mating
Butterflies and peppered moths, adaptive coloration
Kleptoparasitism & Frequency-dependent Selection

gulls, birds, terns, spiders steal from other species

ROSEATE TERNS STEAL FROM CONSPECIFIC. Fitness changes with frequency of expression.

Konrad Lorenz
Ethology, imprinting
Lazuli Birds

Dominant (bright) M tolerate dull M.

Dull M gets territory and mate.

Adults has EPC :)


Communication among bees (sound).
want many rocks and smaller territories.
Long-ta9ped widowbird in Kenya
tail result of sexual selection
Maps (6)
  1. Visual landmarks
  2. Magnetic field
  3. Infrasound
  4. Odor/taste
  5. Salinity of water
  6. Humidity

Mark Konishi
w/ Marler, Owls and "multiplication"
Mice (Progesterone)
Infanticide by M
M mimics food of F by waving legs
Mobbing (Gulls)
Ground Gulls mob, but not cliff gulls.
Monogamy (Splendid Fairy Wrens, possums)

1 mate.

Common in birds, rare in mammals.

Social: live together and raise young together.

Sexual: mating restricted.

Monogamy by isolation (M), pregnancy (M), female enforced, mutual mate guarding, fitness-induced M monogamy&nbs

Motor Neurons + Muscle Glands
Nectar v. Nest (Bees)
More trips for nectar (scout influence increases) Less for next (scout influence decreases)
NEURON Structure & Function
  1. Dendrites and Cell body [Soma] (Input)
  2. Axon: conducts action potential (electrical nerve impulses). Terminals synapse with dendrites on Traget Cell
  3. Chemical neurotransmitters
Niko Tinbergen
Student of Lorenz, Genetics + Environment,
Operational Sex Ratio

OSR = #M/#F

 (Reproductively available)

>1 normal. F choice

<1 reversed. M choice 

Panda Principle
Evolution makes do with whatever elements are available.
Parental discrimination

size, sound, red gape (carotinoids = healthy immune system)

coots feed youngest (brightest) first 

Parental Investment

M: Frog-tadpoles on back, Katydid-edible sparnatophore

F: Wasp w/ cicada, eared grebe carry young 

Classical conditioning
per gene mutation

in drosophila (Konopka and Benzer), changes circadian period

Clock mechanism is negative feedback w/ delay 

Peter Marler

Song learning in birds.

-PROXIMATE: Hearing, Development, Social Envir... WHITE-CROWNED SPARROWS [Baptista: can learn other species' songs]

-ULTIMATE: Genotype has major effect 


Circadian rhythms in drosophila
Secreted protein. high during day, inhibits activity (Gating effect)... CIRCADIAN OUTPUT (during "window" light induces hormone secretion)
gene has many effects on dev/behavior, so behavior can arise w/out selection
F has many M partners (sex role reversal)
M w/ many F. Normal pattern in mammals.
Polygyny (Forms)

F defense

Resource defense (territory)


Scramble (search for F)

Explosive Breeding Assemblage (worms) 


Environmentally induced differences

  1. Food Induced (ants)
  2. Socially (fish)
  3. Predator (aphids)
  4. Seasonal (butterflies)
Proximate Causes of Behavior
  • Neuroethology
  • Sensory
  • Motor
  • Central Nervous Systems
  • How does behavior develop w/in lifetime?
  • What physical mechanisms underlie behavior?
Quails (Testosterone)
After copulation, testosterone in M -> estradiol
Resource Polymorphism
Hereditary and environmental factors, phenotypes w/ different food resources. (Cichlids w/ left or right mouth)

Fly w/ sex role reversal. F not choosy about mates. Honest eggs signal.

R. LONGICAUDA scams M for food. 

Sarah Hrdy
Langur infanticide.
Sedge Warbler
M song size honest signal of M paternal effort.
Sensory Neurons
INPUT: filtering, processing, feature extraction
Sex definitions/roles (George Williams)

F: large gamete. M: small

Masculine sex syndrome: cheap gametes, low parental investment, not choosy 'bout mates, compete for mates, aggressive, sexual displays

FSS: opposite ("cryptic appearance and behavior") 

Sexual Dimorphism
Differing 2y characteristics b/w sexes. Physical AND behavioral
Sexual Imprinting (Tits)
Stronger in Great Tits than Blue Tits
Sexual Selection

Femine sex choose phenotype of masculine sex. Form of natural selection.

Leads to sexual dimorphism. Rapid effects.

Operant Conditioning: Reward/Punishment
Social Behaviors

Agonistic (aggression/threat and conciliation/retreat)



Song Neurobiology (Input)
NCM: song discrimination
Song Neurobiology (Learning)

HVC: learning, recognition?

Area X: Song plasticity

RA: Song plasticity

IMAN: learning 

ZENK GENE (zebra finch) 

Song Neurobiology (Output)


RA: Output control

NXllts: motor nucleus 

Song Pitch (lo or hi?)

LO: dense forests

HI: open areas (woodlands) 

Sparrows (Estrogen)
Estrogen increases territoriality in M
Sponge Isopods

Alpha M: Defend harem

Beta: Mimic F

Gamma: Sneakers

Three morphs have same reproductive success. Are ESS.

St. Peter's Fish
Both M and F mouth brooding. Reduces ability to mate for F, opportunity for M.
Stephen J. Gould
"punctuated equilibrium"
Stotting (Springbok) + Pushups (Lizard)

Cheetah abandons chase. 

Honest signal.

Survival Behavior
Poison, Camouflage, Mobbing, Mimicry
Talking to Strangers
fish love songs, beetle flashes
Talking with Strangers

Honey Guide Bird and Humans

Vervet Monkeys 

Gradual effects, may be associated w/ territorial behavior
mazes, IQ tests for kids
Tiger Salamanders

Normal or CANNIBAL

-More C when larvae are unrelated 

Ultimate Causes of Behavior
  • Evolution
  • Natural Selection
  • Phylogenetic history
  • Adaption
  • How does behavior promote reproductive success?
  • Has behavior arisen over many generations?
Voles (Mating)
  • Prairie are monogamous, meadow polygynous.
  • Vasopressin (hormone) binds to V1aR receptor in pallidum induces bonding behavior.
  • Increase density V1aR -> more bonding.



W.H. Thorpe
Sonogram for bird song, chaffinches learn local dialects!
Warblers (Mate Guarding)
Guard F after fucking. Depends on M density.
M carry eggs on back.
William D. Hamilton, Hamilton's Rule

Inclusive fitness. Altruism.

Gene for altruism spreads iff rbB>rcC (B=indirect fitness gain, C= direct fitness loss) 

"Time giver" sets (entrains) clocks to "local time". External cues (light, temp, social).