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Reuven Dukas

  Chapter  3:  Hormone  &  Neurobiology     Ultimate  &  Proximate  Perspectives   • Proximate  Perspective   o HOW  a  behavior  is  expressed  in  terms  of  genetic  &  physiological  mechanisms.  Studies  the  immediate   cause  &  effect  of  behavior.  (“WHAT”)   o What  causes  diffs  in  plumage   coloration  b/w  male  &  female  finches?  Diffs  in  foraging  b/w  genders.   Males  actively  search  for  &  ingest  carotenoid -­‐based  foods.  Females  eat  carotenoid -­‐based  foods,  but  do   not  actively  search  for  it.   • Ultimate  Perspective   o WHY  a  behavior  exists.     o Why  do  males,  but  not  females,  actively  search  for  carotenoid-­‐based  foods?  B/c  females  are  attracted  to   males  w/  bright  plumage  as  potential  mates.     o Why  do  females  prefer  males  with  bright  plumage?  Males  appear  to  be  better  able  to  fight  of   pathogens.  Brighter  males  f eed  chicks  more  than  twice  as  often  as  drabber  males.  Brighter  males  forage   better  &  this  trait  may  be  passed  to  offspring.     How  the  Endocrine  System  Integrates  Sensory  Input  &  Output   • Hormone  changes  can  modify  behavior  by  ↑  or  ↓  the  frequency  of  that  behavior,  or  they  might  trigger  the  onset   or  end  of  a  behavior.  Hormones  might  prime  animals  so  that  they  are  more  or  less  likely  to  behave  in  a  specific   way  (i.e.  When  testosterone  levels  are  high,  males  are  primed  for  aggression).     The  Long-­‐term  Effects  of  In  Utero  Exposure  to  Hormones   • A  developing  male  fetus  su rrounded  by  female  fetuses  (2F)  is  exposed  to  lower  levels  of  circulating  testosterone.   When  males  mature,  they   are  less  aggressive  &  less  sexually  active  than  males  sur rounded  by  male  fetuses  (2M).   • 2M  males  have  2x  the  level  of  circulating  testosterone ,  sire  more  offspring,  &  show  less  parental  care.   • 2M  females  have  high  levels  of  circulating  testosterone  &  lower  levels  of  circulating  estradiol.   • 2M  females  are  less  attracted  to  males,  reproduce  later  than  other  females,  &  have  fewer  litters.     Vocalizations  in  Plainfin  Midshipman  Fish   • Type  I  males:  high  gonad-­‐to-­‐body  size  ratio,  generate  short  grunts  when  engaged  in  contests  w/  other  males,   generate  long  “hums”  when  courting  females.  If  female  chooses  T ype  I  male,  they  lay  eggs  on  male’s  nest.   • Type  II  males:  low  gonad-­‐to-­‐body  size  ratio,  do  not  build  nests,  linger  around  the  nests  of  Type  I  males  where   they  dart  in  &  shed  sperm  in  an  attempt  to  fertilize  the  nesting  female,  do  not  “hum”  to  attract   females,  only   grunt  occasionally.   • Type  I  has  larger  sonic  muscles  than  Type  II,  affecting  sound  production.  Neurons  in  sonic  muscle  of  Type  I  fires   ~20%  higher  than  Type  II,  which  explains  diffs  in  vocalization.   • AVT  inhibits  production  of  sound  in  Type  I,   doesn’t  affect  Type  II.   • IT  inhibits  production  of  sound  in  Type  II,  doesn’t  affect  Type  I.     Sleep  &  Predation  in  Mallard  Ducks   • Mallards  sleep  with  one  eye  open  &  one  hemisphere  of  the  brain  awake  (“unihemispheric”  slee   • Mallards  on  the  periphery  of  the  g roup,  who  are  more  susceptible  to  predation,  rely  on  unihemispheric  sleep   more  than  birds  in  center  of  the  group.     Chapter  1:  Principles  of  Animal  Behaviour     Natural  Selection   • Hawaiian  Crickets:  To  sing,  the  male  cricket  rubs  2  wings  together.  Songs  not   only  attract  females,  but  parasites   as  well  (trade-­‐off).  In  order  to  avoid  parasites,  males  modified  their  wings  so  that  they  were  incapable  of   producing  song.  In  order  to  still  attract  mates,  the  song -­‐less  males  will  stay  near  a  handful  of  singing  males.   • Xenophobia:  The  fear  of  strangers  or  those  from  outside  one’s  group.   • Mole  Rat:  Live  in  underground  colonies.  Each  group  1   pair  of  breeders  that  produce  offspring  in  the  colony,   meaning  most  members  are   genetic  relatives.  Some  inhabit  mesic  (moderately  moist)  environments  that  have   mild  resource  limitations,   some  inhabit  arid  (dry)  environments  that  have  intense  limited  resources.  An  arid  &  a   mesic  mole  were  placed  together.  When  the  pair  was  both  males  or  both  females,  aggression  towards  strangers   was  more  pronounced  in  arid  than  mesic  moles   –  identification  of  a  stranger  initiated  aggression.  Natural   selection  favored  stronger  xenophobic  responses  as  a  function  of  both  where  they  live  &  the  sex  of  strangers.     Individual  Learning   • A  relatively  permanent  chang e  in  behavior  as  a  result  of  experience.   • Grasshopper:  2  food  dishes:  (1)  balanced  diet  =  proteins  &  carbs  (2)  deficient  diet  =  protein,  no  carbs.  Diets   supplemented  w/  citral  or  coumarin,  &  was  placed  in  a  brown  pla te  or  a  green  plate.  Learning  treatment  –   balanced  diets  always  paired  w/  1  color  &  odor.  Grasshopper  could  learn  which  dish  contained  balanced  diet.   Random  treatment  –  odor  &  colors  associated  w/  balanced  diet  randomly  changed,  preventing  grasshopper  from   learning  which  dish  contained  balanced  diet.  Results:  Grasshoppers  in  learning  treatment  ate  more  balanced  diet   than  in  random  treatment  &  ∴  had  a  higher  growth  rate.     Cultural  Transmission   • The  transfer  of  info  from  individual  to  individual,  through  teaching  or  social  learnin g.   • Rats:  Observer  &  demonstrator  rat  live  in  same  cage  for  a  few  days,  then  demonstrator  rat  removed  &  taken  to   another  cage  where  it  was  give   one  of  two  new  diets  –  food  flavored  w/  cocoa  or  cinnamon.  Demonstrator  then   st brought  back  to  1  cage  &  allowed  to  interact  w/  observer  for  15min  before  being  removed  again.  After  a  few   days,  observer  rat  given  the  2  bowls  of  food.  Results:  Through  the  use  of  olfactory  cues,  observer  rats  were   influenced  by  the  food  demonstrators  had  eaten  &  were  more  likely  to  eat   that  food.     Direct  Fitness:  Measured  by  the  #  of  viable  offspring  produced.   Indirect  Fitness:  Measured  by  the  ↑  reproductive  success  of  genetic  relatives.   Inclusive  Fitness:  Direct  +  indirect  fitness     Chapter  2:  The  Evolution  of  Behaviour     Natural  Selection   • Traits  ↑  or  ↓  in  frequency  as  a  function  of  how  well  they  suit  organisms  in  their  environment.   • Even  a  fitness  advantage  of  1%  can  affect  natural  selection.   • For  natural  selection  to  occur,  there  must  be:   o Variants  in  the  trait     o Fitness  consequences  of  the  trait   o A  mode  of  inheritance   • For  birds,  approaching  novel  objects  can  be  beneficial  or  dangerous.  The  time  to  approach  a  novel  object  is  the   “approach”  score.   • VARIATION:  Genetic  variation  (mutations),  genetic  recombination,  migration   • FITNESS  CONSEQUENCES:  The  diff  in  reproductive  success  associated  w/  slow  vs.  fast  approach.   • MODE  OF  INHERITANCE:  W/o  a  mode  of  inheritance,  any  fit ness  diffs  that  exist  w/in  1  generation  are   lost.   • TRUNCATION  SELECTION:   o Truncation  selection  experiment:  Measures  heritabil ity  by  allowing  only  those  w/  extreme  forms  of  a   trait  to  breed  &  then  tracking  changes  in  that  trait  across  generations.   o Step  1:  Measure  the  approach  score  of  every  bird  >12  months0  of  age  ( x ).   o Step  2:  Cut  off  the  pop  variation  in  approach  scores  by  allow ing  only  individuals  w/  approach  scores  >   some  value  to  breed.  Calculate  the  mean  approach  score  of  individuals  that  we  have  allowed  to  breed   (x ).  Selection  differential  (S)  =  the  diff  b/w  x  and  x .  It’s  the  max  amount  we  could  expect  natural   1 0 1 selection  to  change  approach  scores.   o Step  3:  Raise  offspring  produced  by  generation  1  until  they  reach  12  months.  Then  measure  their  mean   approach  scores  (2 ).  Response  to  selection  (R)  =  the 0  diff 2  b/w  x  and  x  –  measures  how  much  truncation   selection  has  changed  approach  scores  across  generations  1  &  2.   Heritability  =  R/S.   • PARENT-­‐OFFSPRING  REGR2SSION:   2 o Measures  h . N  When  h N  is  high,  the  behavioral  variation  in  the  offspring  should  map  into  the  behavioral   variation  observed  in  parents.  The  greater  the  role  of  environ mental  variance  plays  in  determining   variance  in  behaviour,  the  lower  the  heritability  of  that  behaviour.   o Group  size  of  swallows  is  similar  to  group  size  of  parents.  This  was  true  for  offspring  that  bred  at  the   same  site  as  parents,  as  well  as  for  offsprin g  that  emigrated  elsewhere,  suggesting  that  the  correlation   b/w  parent  &  offspring  was  not  a  function  of  living  in  the  same  place.   ∴  preferable  group  size  may  be   heritable.   o ½  the  young  from  large  colonies  were  removed   &  the  young  from  small  colonies  were  placed  in  their   stead.  Likewise,  ½  the  young  in  the  small  colonies  were  replaced  by  young  from  large  colonies.  Led  to   offspring  from  both  large  &  small  colonies  ( cross-­‐fostering  experiment).  +  correlation  w/  the  group-­‐size   preferences  of  their  genetic  parents.   –  correlation  uncovered  when  group-­‐size  preferences  of  offspring   &  foster  parents  were  compared.   ∴  young  displayed  same  group-­‐size  preference  as  their  genetic   parents.     Antipredator  Behaviour  is  Guppies   • Upstream  =  low  predation  from  small  fish.  Downstream  =  severe  predation  from  large  fish.   • Little  gene  flow  b/w  high  &  low  predation  sites,  so  natural  selection  should  favor  diff  traits  in  each  site.   • High  predation:  Guppies  mature  faster,  produce  smaller  offspring,  &   energy  used  for  reproduction.  Why?  B/c   predators  are  large  &  can  eat  a  guppy  no  matter  how  big  they  get  ∴  producing  many  smaller  fish  is  favored.   Larger  group  size.  Inspect  predators  more  cautiously  &  more  often.   • Low  predation  sites:  Only  small  predators.  If  guppies  can  get  past  a  certai n  size  threshold,  they  are  safe  ∴  they   produce  fewer,  larger  offspring.     Kinship  and  Naked  Mole  Rat  Behaviour   • Display  eusociality  (an  extreme  form  of  sociality  prese nt  in  many  social  insect  groups).   • Reproductive  division  of  labor,  overlapping  generations,  communal  care  of  young   • Within-­‐colony  aggression  exists  but  cooperative  behaviour  are  more  common.   • Non-­‐reproductive  rats  (shorter  life  span)  undertake  a  wide  variety  of  cooperative  behaviou rs  (i.e.  digging  tunnels,   grooming  queen,  defending  against  predators).     • Kinship  theory  suggests  that  the  more  genetically  related  individuals  are,  the  more  cooperation  they  will  show .     Phylogenetic  Trees   • Depicts  the  evolutionary  history  of  a  group  of  species.   • Homology:  a  trait  shared  by  2  or  more  species  b/c  they  shared  a  common  ancestor.   • Homoplasy:  a  trait  resulting  from  natural  selection  acting  independently  on  each  species  ( convergent  evolution).       Phylogeny  &  Parental  Care   • Ray-­‐finned  fish:  Maternal  care  tended  to  evolve  after  species  moved  from  external  to  internal  fertilization.   External  fertilization  =  paternal  care,  intense  male  coloration  &  nest  construction.  Male  coloration  was  also   associated  with  systems  in  which  females  fertilized  their  eggs  internally.  In  external  breeders,  parent al  care  was   found  in  25%  of  families  studied.  In  internal  breeders,  parental  care  was  found  in  90%  of  families  studied.       Chapter  5:  Learning     Classical  Conditioning:     • The  experimental  pairing  of  a   conditioned  stimulus  (CS)  &  an  unconditioned  stimulus  (US).   • CS:  A  stimulus  that  initially  fails  to  elicit  a  particular  response,  but  comes   does  when  associated  w/  US   • US:  A  stimulus  that  elicits  a  response  in  the  absence  of  training   • Conditioned  Response  (CR):  The  learned  response  to  a  CS   • Appetitive  Stimulus:  Any  stimulus  that’s  considered  pleasant,  rewarding   • Aversive  Stimulus:  Any  stimulus  that’s  associated  w/  an  unpleasant  experience  (i.e.  shock)   • Excitatory  Conditioning:  When  a  CS  leads  to  an  action.   • Inhibitory  Conditioning:  When  a  CS  inhibits  behaviour.   • Second-­‐order  Conditioning:  Once  a  CR  has  been  learned  by  pairing  US  &  CS1,  a  new  stimulus  is  presented  before   CS1,  &  if  the  new  stimulus  itself  eventually  elicits  the  CR,  then  the  new  stimulus  becomes  CS2.   • Overshadowing:  A  situation  in  which  the  learned  response  to  an  US  is  stronger  when  it’s  presented  alone  vs.   when  it’s  paired  w/  a  US2.   • Blocking:  When  an  association  b/w  an  US1  &  a  response  prevents  an  individual  from  responding  to  US2  or  causes   the  individual  to  respond  less  strongly  to  the  US2.   • Group  1:  blue  stick  (CS1)  &  cat  odor  (US).   • Group  2:  yellow  light  (CS2)  is  always  presented  w/  blue  stick,  just  before  cat  odor   • Rats  from  both  groups  are  tested  in  response  to  blue  stick  alone.  If  yellow  light  overshadows  blue  sti ck,  group  2   will  respond  less  strongly  to  blue  stick  than  group  1.   • Group  3:  1  trained  to  associate  blue  stick  w/  odor,  then  yellow  light  presented  at  the  same  time  as  blue  stick,  &   then  odor.   • Blocking  occurs  when  group  3  responds  less  strongly  to  yellow   light  (when  presented  alone)  than  group  2.  It’s  as  if   initially  learning  to  associate  blue  stick  w/  odor  blocked  the  ability  to  pair  yellow  light  w/  odor.     Instrumental  Conditioning   • Learning  that  occurs  when  a  response  made  by  an  animal  is  reinforced  by  re ward/punishment.  An  animal  must   undertake  some  action/response  in  order  for  the  conditioning  process  to  produce  learning.   • Law  of  Effect:  If  a  response  in  the  presence  of  a  stimulus  is  followed  by  a  satisfying  event,  the  association  b/w  the   stimulus  &  the  response  will  be  strengthened.   • Operant  Response:  A  learned  action  that  an  animal  makes  to  change  its  environment.     Within-­‐Species  Studies   • Optimal  forgetting  in   Stomatopods:  Males  &  females  share  a  nest  cavity  for  a  few  days  before  they  breed,   &  they   actively  repel  all  intruders  from  this  area.  Shortly  after  mating,  males  leave  breeding  cavity   &  go  search  for  a  new   mate.  Males  &  females  remember  each  other  after  male  has  left  for  4  weeks.  Females  guard  brood  for  4  weeks,   after  which  brood  leaves  cavity  for  good.  Males  not  aggressive  toward  female  during  4  weeks,  but   likely   afterwards.     Between-­‐Populations  Studies   • Learning,  foraging,  &  group  living  in  Doves   o Prediction:  Animals  in  groups  should  learn  more  quickly  than  individual  animals.   o 16  group  +  16  territorial  doves.  All  had  to  learn  to  pull  a  ring  that  opened  a  drawer  w/  food.  Group -­‐living   doves  learned  faster.  The  more  difficult  the  learning  task,  the  more  pronounced  the  b/w -­‐pop  diff.   • Learning  &  antipredator  behaviour  in  Sticklebacks   o Raised  fish  derived  from  high-­‐predation  &  low-­‐predation.  Individuals  from  both  pops  were  equally  adept   at  learning  that  food  would  come  to  1  side  of  tank.  After  having  food,  fish  subjected  to  a  simulated   heron  attack  on  the  side  of  the  tank  w/  the  food.  Both  pops  lea rned  to  avoid  danger  side  of  tank,  but   high  predation  learned  faster.  Natural  selection  on  learning  &  antipredator  behaviour.     Learning  About  Your  Mate   • Mongolian  Gerbils :  Rely  on  chemical  communication  for  social  in teractions.  Presented  males  w/  odor  (mint,   lemon)  &  then  gave  them  access  to  mates.  Presented  males  in  another  group  (control)  w/  odor,  but  didn’t  give     them  access  to  mates.  Males  w/  odor  +  mate  learned  quickly  to  approach  the  area  w/  odor  to  access  female,   males  in  control  group  made  no  associat ion.     Learning  About  Familial  Relationships   –  Fig  5.25   • Long-­‐tailed  Tits:  Young  breed  independently  asap,  but  most  nests  fall  victim  to  predation.  At  that  point,  breeders   become  helpers  at  the  nest  of  close  genetic  relatives.  “Churr”  call  develops  before  yo ung  birds  leave  nest.  Used   for  short-­‐range  communication.  Individuals  showed  strong  preference  for  calls  given  by  their  kin.  Why  is  churr   call  learned?  (1)  The  calls  of  foster  siblings  raised  together  were  as  similar  as  the  calls  of  biological  siblings  rai sed   together  (2)  the  calls  of  biological  siblings  raised  apart  were  dissimilar  as  the  calls  of  unrelated  tits  in  nature  (3)   Songs  of  foster  parents  &  their  foster  offspring  very  similar,  while  songs  of  biological  parents   &  their  offspring   very  diff  when  those  offspring  were  raised  by  foster  parents.     Learning  About  Aggression  –  Fig  5.26&5.27   • Blue  Gouramis:  Can  be  trained  to  associate  light  w/  the  presence  (+)  or  absence  ( -­‐)  of  intruder  male  in  territory.   After  presentation  of  light,  fish  that  learned  that  a  light  meant  presence  of  another  fish  were  much  more   aggressive  towards  intruder.  ½  of  fish  trained  to  associate  red  light  w/  intruder.  ½  of  fish  (control)  presented  w/   red  light  6hrs  before  intruder  –  didn’t  make  association.  Contest  1:  After  red  light,  t rained  +  control  males  pitted   against  stch  other  until  a  winner  &  loser  emerged.  Trained  males  more  likely  to  win.  Contest  2:  stnners  +  losers   from  1  contest  paired  w/  new  intruders  3  days  later.  No  red  light  shown.  All  fish  that  won  1  contest  had  won   2  contest  (trained).  Trained  males  that  lost  1  contest  were  likely  to  lose  2  contest.     Molecular  Genetics  of  Learning  in  Rats   • High  avoidance  (HA)  +  low  avoidance  (LA)  lines  descended  from  single  pop  of  rats.  Each  generation  tested  on  its   tendency  to  avoid  auditory  +  visual  cues  associated  w/  shock.  HA=best  at  avoiding  shock,  LA=worst  at  avoiding   shock.  LA  showed  higher  levels  of  anxiety.  4  genes  showed  greater  expression  in  hippocampus  of  HA,  4  genes   expressed  in  greater  quantities  in  LA.   ∴  genes  may  control  complex  traits  like  avoidance  learning.   • Corticosterone  plays  role  in  stress  responses  &  learning.  Binds  to  receptor   Mineralocorticoid .   • Fear  test  then  spatial  test  –  placed  rats  in  water  maze  &  measured  ability  to  find  &  remember  location  of   submerged  escape  platforms.  HA  took  longer  than  LA  to  learn  to  swim.  HA  had  higher  corticosterone  levels  than   LA.  HA  had  fewer  mineralocorticoid  receptors  resulting  in  reduced  ability  to  bind  corticosterone,  indirectly   leading  to  an  ↑  of  stress  hormones.       Chapter  6:  Cultural  Transmission     Cultural  Transmission   • The  transfer  of  info  from  individual  to  individual,  through  teaching  or  social  learning.   • Local  Enhancement:  An  individual’s  drawn  to  a  particular  area  b/c  it’s  observed  another  individual  in  that  location   • Social  Facilitation:  When  the  presence  of  a  model  facilitates  learning  on  the  part  of  an  observer.   • Capuchin  Monkeys:  What  factors  affect  monkey  probability  of  eating  novel  food?  Treatment  1  (control):  a  lone   monkey  was  tested  on  its  tendency  to  tr y  new  food  (veggies  that  were  color  dyed).  Treatment  2:  monkey  +  novel   food  on  1  side  of  cage,  group  of  monkeys  was  on  other  side  of  cage  w/  no  food.  Treatment  3:  monkey  +  novel   food  on  one  side  of  cage,  group  of  monkeys  +  familiar  food  on  other  side.  Here ,  lone  monkey  saw  the  group   eating  food,  but  not  the  novel  food.  Evidence  for  local  enhancement:  (1)  lone  monkey  in  treatment  3  was  more   likely  to  be  eating  food  than  monkey  in  treatment  1  (2)  lone  monkey  ate  more  food  when  it  saw  other  monkeys   eat  more  food.  No  evidence  for  social  facilitation.     Social  Learning   • Imitation:  The  acquisition  of  a  novel  response  through  observation  of  a  demonstrator  making  that  response.   o Budgerigars:  Birds  that  observed  model  that  used  its  beak  to  gain  access  to  food  were  more  likely  to  use   beaks  &  vice  verse  when  using  foot.  Birds  learned  a  novel   response  from  watching  others.   o Correspondence  Problem:  Individual  1  attempts  to  imitate  individual  2,  it  can  only  see  2’s  movements   not  the  muscle  activation  underlying  movement.  How  does  1  know  what  to  do  to  perform  movement?     • Copying:  Observer  repeats  action  of  demonstrator.  Copier  is   rewarded  for  whatever  behaviour  is  copied.   o Guppies:  Color  matched  males  placed  in  side  chambers.  Model  female  placed  near  one  male.  Observer   female  saw  model  choose  a  male  to  mate  with.  Result:  Observer  chose  same  mate  as  model.   o Mice  &  Flies:  Mice  not  affected  by  presence  of  fly.  Mouse  hides  under  debris  when  bitten  by  fly.   Observer  buries  hides  under  debris  when  exposed  to  fly.     Rise  &  Fall  of  Tradition   • Meerkats:  7/9  groups:  demonstrator  trained  to  show  a  preference  for  1  of  the  2  colour  &  shape  coded  landmarks.   2/9  groups:  no  demonstrators  trained.  Meerkats  preferred  landmark  t hat  demonstrator  had  been  trained  to   prefer.  Preference  lasted  for  a  few  days,  then  slowly  dissipated.  Overtime  meerkats  explored  other  landmark.   Once  they  learned  themselves  that  the  other  landmark  was  just  as  good,  they  spent  time  at  both  landmarks.     • Social  learning  produces  tradition,  but  individual  learning  leads  to  its  demise.     Teaching   • Meerkats:  Young  pups  incapable  of  catching  prey.  Pups  assisted  by  older  foragers  (“helpers”).  Helpers   incapacitate  scorpions  by  removing  stingers  &  present  them  (dead)  as  food  to  pups.  As  pups  got  older,  helpers   presented  them  more  &  more  live  scorpions.  Helpers  respond  to  begging  calls  of  pups  even  when  pups  can’t  be   seen.  Hearing  call  from  old  pup,  helpers  bring  live  scorpions.  Hearing  call  from  yo ung  pup,  helpers  bring  dead   scorpions.  Other  evidence  for  teaching:  (1)  spent  significant  time  monitoring  pups  after  presenting  them  w/  food;   (2)  retrieved  prey  when  pups  lost  their  food;  (3)  on  occasion,  further  modified  scorpion  (removing  stinger,  killin g   scorpion,  etc.)  after  it  was  lost  but  later  retrieved  by  pups;  (4)  nudged  pups  that  were  reluctant  to  eat  scorpions.   • Almost  all  instances  of  teaching  focus  on  parent/offspring  relationship.     Modes  of  Cultural  Transmission   • Vertical  cultural  transmission:  Info  passed  directly  from  parent  to  offspring.   o Dolphins:  “Beaching”  involves  a  dolphin  surging  out  of  the  water  &  onto  a  beach  to  catch  fish,  then   quickly  returning  to  water  (a  rare  behaviour).  Another  form  of  beaching     -­‐  where  1-­‐6  dolphins  isolate  a   school  of  fish  &  herd  fish  toward  land,  creating  a  wave  to  send  them  onto  the  land,  then  surge  out  of  the   water  to  capture  the  fish.  Both  dangerous  as  dolphin  can  get  stranded.  Only  calves  born  to  beaching   mothers  were  beachers   –  they  learn  this  behaviour  from  their  mothers.   • Oblique  cultural  transmission:  Info  is  passed  across  generations,  not  from  parent  to  offspring.   o Snakes  &  Monkeys:  Exposure  to  adult  models  showing  fear  in  presence  of  snake  led  to  observers   showing  similar  fear  responses.  When  observers  saw  a  model  that  had  been  trained  to  display  fear  in   presence  of  neutral  object  (i.e.  flowers),  the  observers  didn’t  display  fear.   • Horizontal  cultural  transmission:  Info  is  passed  across  individuals  of  the  same  age.     o Guppies:  More  difficult  to  train  fish  to  take  longer  path  when  both  long  &  short  paths  were  present.   Once  long  &  short-­‐path  groups  were  trained,  original  members  from  each  group  were  slowly  removed  &   replaced  w/  new  individuals  that  didn’t  know  either  path.  New  fish  used  path  to  which  the  original  fish   had  been  trained.     The  Interaction  of  Genetic  &  Cultural  Transmission   • Grants’  Fishes:  Songs  of  sons  resemble  the  songs  of  their  paternal  grandfathers,  but  no t  of  the  maternal   grandfathers.  This  is  b/c  the  song  is  cultu rally  transmitted  to  the  father  &  then  transmitted  to  the  son.  Cultural   transmission  of  song  allows  females  to  recognize  males  of  their  own  species.  Females  tend  to  avoid  males  who   sing  songs  that  are  similar  to  her  father’s  song   –  prevents  inbreeding.   • Guppy  Mate  Choice:  Whether  or  not  females  copy  a  model’s  mate  choice  is  affected  by  a  threshold  diff  in  the   amount  of  orange  color  in  the  male.  If  the  male’s  orange  colour  is  beneath  this  threshold,  the  effects  of  copying   are  predominant.  But  if  the  orange  color  is  abo ve  this  threshold,  no  copying.     Cultural  Transmission  &  Brain  Size   • Bird  species  in  which  individuals  had  high  brain  size/body  ratio  were  more  likely  to  survive  &  thrive  after  intro  to   novel  environments  than  were  species  w/  lower  brain  size/body  ratios.   • When  large-­‐brained  species  were  introduced  to  novel  environments,  they   ↑  their  rate  of  innovation  (i.e.  new   foraging  technique)  which  in  turn  ↑  their  probability  of  success.       Chapter  11:  Foraging     Search  Image  Theory:  A  representation  of  prey  that  predators  form  over  time  (becomes  more  detailed  w/  experience) .   Optimal  Foraging  Theory:   A  family  of  mathematical  models  developed  to  predict  animal  foraging  behaviour.     What  to  Eat   • Energy  intake  from  prey  can
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