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bio 472 midterm

Terms

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What are PROXIMATE questions?
-'how' questions
-deals with the mechanisms of any function
-how they know, how they do (physiological), etc.
What are ULTIMATE questions?
-'why' questions
-deals with the survival value or the function of the behaviour
-eg: why N.Tenbergen removes eggs from the nest
Give an example of a PROXIMATE situation.
-wasps needing special marks in order to find home
-that's HOW they find it
Give an example of a ULTIMATE situation.
-N.Tenbergen removes egg shells from it's nest
-does it BECAUSE it reduces predation
What are the 3 foundations to Animal Behaviour?
1) Ethology
-founders: Karl vonFrisch, Konrad Lounz and Niko Tinbergen
-evolution a prominent theme
-emphasizes study of animals in natural conditions
-focus on proximate & ultimate caustion

2) Comparative Psychology
-compared humans to animals
-few subjects (eg: rats)
-lab based
-focus on how behaviour is acquired by learning and conditioning animals as 'blank states'

3) Neurobiology
-arose from human medicine, anatomy and physiology
-function of nervous system
-roles of hormones in behaviour
-mechanistic (how behaviour is controlled or enacted)
What are FAP's?
-Fixed Action Potentials
-early work emphasized innate behaviours
-FAP's triggered by simple stimuli (sign stimluli or releasers - stim that causes FAPs)
-lead to the concept of the Innate Releasing Mechanism (IRM) [ie: yawning triggers yawning]
What are the 5 Aims of Ethology?
To answer questions about...
1)Form
-what body postures and movements?

2)Function
-what is the immediate consequence of the behaviour

3)Causation
-what are the internal (eg: hunger levels) and external (eg: presence of food) factors that bring about the occurrence of a behaviour at a certain time

4) Acquisition
-how does the animal come to have the behaviour? (eg: experience, innate, evolution, culture, developement)

5) Biological significance
-why is the behaviour present, and what does it do for the animal
What are some potential biases/limitations to Ethology?
1) Human Social Conditioning
-cultural belief influences what we see and how we interpret it

2)Human Sensory Perception
-many other animals do not percieve the same stimuli we do

3)Ethical and Legal Concerns
-may constrain where, what, when and why animals are studied
-all research must be approved by a scientific board
-Federal Regulation by 'Canadian Council on Animal Care'
What is Hypothesis Testing?
-make an educated guess/inference about the behaviour (what, how, why) called a WORKING HYPOTHESIS
-set out to confirm or reject the hypothesis by making testable predictions "if->then"

**a hypothesis can be disproven; but never proven in the strict sense... one is never certain this is the one absolutely correct explanation. Therefor we can never say "This proves".
What is MODELLING? What's the pupose?
-a mathematical formulation of a natural phenomenon (eg: graphs)
-purpose
i)to clarify the hypothesis
ii)clarify assumptions
iii)to simplify the system so we can test the critical variables
T/F

Predators should always eat prey, regardless of size.
F

-if a predator encounters large prey, ALWAYS EAT IT
-if a predator encounters small prey, only eat it if (Es/hs)>(El/[sl+hl])

E=energy of animal (small/large)
h=handling time of animal
s=searching time
How would you study CONVERGENCE in regards to evolution of behaviour with emphasis on the COMPARATIVE approach?
-look at behaviour as potentially analogous traits among unrelated organisms and atempt to explain those traits by searching for correlated environment factors
-eg: eggshell removal
How would you study DIVERGENCE in regards to evolution of behaviour with emphasis on the COMPARATIVE approach?
-look at groups of closely related animals and attempts to reveal evolutionary relationships through studying behaviours
-eg: balloon flies tricking females with decoy so that he can hump her before she eats him
How would you study CORRELATIONAL/OBSERVATIONAL APPROACHES in regards to evolution of behaviour with emphasis on the COMPARATIVE approach?
-see if there's a relationship between 2 variables
-however, correlation does not equal causation
How would you study EXPERIMENTAL APPROACHES in regards to evolution of behaviour with emphasis on the COMPARATIVE approach?
-a repeatable, deliberate manipulation of a phenomenon when one or a few factors of interest are permitted to vary while the others are controlled or held constant
-it is important to observe without interfering
What is an ETHOGRAM? What are the number of BEHAVIOURS dependant on?
-an inventory of behaviors
-number of behaviours depends on:
1)how many behaviours the animal has
2)how rare particular acts are
3)how much time you observe
4)how you classify/subdivide behaviours

**Remember to consider all domains when trying to describe the behaviour, or else you may end up with an incomplete or subjective description**

**try to avoid assigning functions to behaviours when naming them, even if the function seems quite obvious (eg: instead of saying "begging", say "gaping")
According to Drummond, how do you describe, define and name an ETHOGRAM?
Describing:
-report the properties that comprise a behaviour or behaviour pattern
-the decription should allow the reader to envision it exactly

Define:
-specify the properties necessary to identify this behaviour as distinct from all others

Naming:
-apply a name, however, avoid interpreting the behaviour or assaying a function to it
What are the 5 Domains that Contribute to Defining a Behaviour?
1)Location in space:
-put the animal in relation to a specific component of the environment (eg: "Lecturing" is at the front of the blackboard)

2)Orientation in the Environment:
-the placement of various structures of he animal in relation to other parts of the environment (includes directing body parts towards objects)
-eg: when lecturing, her face directed to the class

3)Topography of the Animal:
-the core of descriptions
-eg: movement of limbs
-remember, some movements may be static

4)Intrinsic Properties (internal control):
-internal physiological processes
-eg: releases of an electric signal, flashing of fireflies, release of pheromones, etc.

5)Impacts on the Physical Environment:
-displacement, consumption, destruction, etc. of organic or inorganic things (including other animals)
-results can be caused by mechanical, chemical, electric means, etc
-not all consequences of a behaviour are part of the behaviour (eg:walking will kick up dust... not relevant to descriptions)
Apply Darwin's principle of Natural Selection in genetic terms. 4
1)All organisms have genes that code for proteins. These proteins regulate the nervous system and the morphology of an animal and so determine it's behaviour

2)Within a population, alternate forms of genes (alleles) cause slight differences in behaviours between individuals

3)There is a competition betwen alleles for a particular site (locus) on chromosomes

4)Any allele that can make more surviving copies of itself will, over time, replace other alleles in the population. Natural selection is the differential survival of alternate alleles.

**Individuals are viewed as temporary vehicles by which genes replicate**

**Note that selection of genes is caused by the differential survival and reproduction of individuals (via their phenotypes therefore genes that promote survival or reproduction are favoured**
Define FITNESS
-the relative performance of alleles compared to other alleles
In general, are animals more concerned with Individual Selfishness or Group Advantage?
-individual selfishness
-groups evlove because a selfish ass developed it
-animals aren't really concerned with the species as a whole, they are only worried about their own reproduction
-therefor, animals selected for will have their offspring advance, thus strengthening the group
In Wynne Edwards opinion, what is the leading cause to extinction?
-a pop'n that overexploits its resources would go extinct
-controls evolve to ensure a pop'n/spp limits its rate of consumption (eg: restriction of birthrates)
What are 2 Theoretical Problems with Group Selection?
1) lifespan of the individual is shorter than the groups. Strength of selection on the individual will be greater then the selection on the group

2)Groups must be isolated from immigrants from other groups. Each group is susceptible to invasion by a selfish mutant
What are 2 LIFE HISTORY TRADE OFFS?
1) Quantity vs. Quality of offspring

2)Parental survival (Current vs. Future Reproduction)
-adult motrablity increases as clutch size increases
-terminal reproductive investment (older age classes put more effort in because of shorter expected future opportunities)
What is the OPTIMALITY THEORY? 2 types
Simple Optimum:
-the best behaviour for an animal to do given free choice

Competitive behaviour:
-the best behavior given a constraint (eg: competition from others)
-could consider how constraints or tradeoffs involving optimum behaviour such as predation
-(eg: foraging lapwings take smaller worms if kleptoparasitic gulls are present)
What 5 factors should be considered when determining if optimality is adaptive or non-adaptive? ie: what are neutral traits, phylogenetic constraints, physiological constraints, evolutionary time lags and rules of thumb?
1)Neutral traits (eg: driving on the right side of the road

2)Phylogenetic constraint
-similar evolutionary history may effect which behaviours are present (eg: be good for a mouse to fly away from a hawk... but mice don't have wings)

3)Physiological constraints
-speed of nerve transmissions, muscle contractions, etc.

4)Evolutionary Time Lags
-environment is constantly changing, therefore what is optimal also changes over time
-co evolution and arms races influence how behavriour evolves: ASYMMETRIC (eg: attack/defense); Symmetric (eg: competition to be bigger)

5)Rules of Thumb
-animals may use approximations to make decisions
-as long as mistakes aren't too grequent, there is little selection to evolve more elaborate mechanisms
-eg: prey choice of shrews. Normally profitability of meal worms peices increase with size except between 2-4 segments when shrews must pick up the segs and chew then open. If geiven a choice between 2 segments vs 4, they should choose 4... However, they still preferred 4.
Define ALTRUISM
-acting to increase one individuals lifetime number of offspring at a cost to one's own survival and reproduction
Give an example of Altruism.
Belding Ground Squirrels
-give alarm calls when they see a predator
-increases the probability that the caller will be attacked
-benefits the receiver (increases probability of escape)
According to WD Hamilton, why would animals be ALTRUISTIC?
-related individuals have genes in common
-parents share 50% of genes with their offspring
-therefor their coefficient of relatedness (r) is r=0.5
According to Maynard Smith, why would animals be ALTRUISTIC?
-'Kin Selection' - how behaviours are selected by promoting the survival of relatives
- Direct Fitness + Indirect Finess = Inclusive Fitness
What is HAMILTON'S RULE?
-'helping' between animals should be favored when

rB-c > 0

Where:
B = benefit = extra offspring produced by relatives as a result of help
c = cost to self
r = degree of relatedness
What is COOPERATIVE BREEDING and KINSHIP?
-individual takes care of another which is not directly their own (keep in mind degrees of relatedness)
-individuals should judge costs and benefits for their own unique situation (Hamilton's Rule)
Do HELPERS benefit?
-they rear some (partly) related relatives (r=0.3-0.6). But on their own, could rear approx 1 offspring
-so, helpers usually have no options of finding their own territory or mate. As soon as a breeder dies, it's place is taken over by a helper
Give 3 reasons why a HELPER should stay?
1)Increases ones own survival
2)No option of own territory
3)Increases chance of inheriting territory
Give 4 reasons why a HELPER should help?
1)Increases survival of helper's parents
2)Increases production of relatives
3)Increases territory size
4)Gain experience in rearing offspring
Give an example of HELPERS helping.
Social Hymenoptera (bees, wasps, ants, etc)
-only the queen lays eggs, workers forego reproduction and help the queen reproduce. An example of extreme altruism
-males are haploid; females are diploid
-queen is equally related (r=0.5) to her sons and daughters
-workers: more related to sisters (r=0.75) than to own offspring (r=0.5)
-workers can maximize fitness by helping their mother produce more sisters
How do individuals recognize kin? 4
1)Recognition Alleles = Green Beard Effect
-linked alleles cause altruistic behaviour when a prominent physical trait is displayed
-recognition is triggered without learning
-theoretical; hard to model it in real life

2)Rules of Thumb
-treat anyone at home as kin
-brood parasites exploit this rule

3)Learning from association in early life
-eg: aggression in ground squirrels
-siblings reared together vs non siblings reared together
-siblings reared apart vs non siblings reared apart
-aggression is equal and low for siblings reared together
-aggession is higher when raised apart, but slightly lower among relatives

4)Phenotype matching
-most cooperative with individuals she is most phenotypically most similar to
-eg: odours, chemoreception, visual. etc.
Define RECIPROCITY
-you help me, and I'll help you
-when the benefit of an altruistic act to the recipient is greater thatn the cost to the actor, thenas long as the help is reciprocated, both participants will gain
-possibility of cheating (see Jan.18 HO)
-so is it evolutionarily possible to maintain?
How can RECIPROCITY be initially established?
*not well understood from modelling perspective
1)Maybe first evolves in clusters of kin (therefore kin selection operates and initially gets co op going)

2)More likely to evolve in small, fixed pop'n 'clusters' where cooperators had higher prob of interacting with eachother

3)Cooperating is stable if cheaters can't get away withough retaliation
-repeat encounters
-individual recognition
-long term memory
Give an example of RECIPROCITY.
Regurgitation of blood of Vampire Bats
-some bats suck at hunting, therefore bats regurgitate blood to feed other, hoping 1 night that they'll get paid for it
Define MUTUALISM.
-different than reciprocity because there is no temptation to cheat
-both players get immediate benefits by cooperation
-eg: wolves pack hunting
Give an example of MUTUALISM.
Wolves and Lions hunting in packs
-cooperation means larger prey, then they share it
-male lions take over pride
-larger coalitions tended to be related males and then only a few males mated (kin sel)
-smaller groups - unrelated males - then mating distributed more equally
Define MANIPULATION
-AKA Pseudoaltruism
-some behaviours seem cooperative but actually are not.
-eg: brood parasites, cuckoos. Parent hosts seem altruistic, but they are just being manipulated
What are the 4 kinds of ALTRUISM?
1) Kin Selection
2) Reciprocity
3) Mutualism
4) Manipulation (not really though)
What must occur in order for behaviour to evolve?
1) Must be behavioral alternatives in the pop'n (variation)

2) Must be heritable (genetic origin)

3) Some behavioural alternatives must confer greater reproductive success
What is considered when dealing with Inheritance of Behaviours?
Nature vs. Nurture
-is behaviour inborn or is it learned?

eg: honeybee nest cleaning is genetic
eg: golden hamster photoperiod is genetic
What are POLYGENIC TRAITS?
-traits that are controlled by multiple genes
-ie: bee's cleaning hive. Genes control what they do
How would you investigate POLYGENIC TRAITS? 3
1) Controlled Breeding and Hybridization
eg: foraging behaviour of garter snakes
-1)Coastal-terrestrial pop'n eat slugs
-2)Inland-lake pop'n eats frogs and fish
-naive snakes, 73% of coastal will eat slugs if presented with one, but only 35% of inland snkaes will
-controlled breeding, hybrids, intermediate prey choice responses
-this indicates that prey selection is at least in part controlled by genetic differences between the populations

2)Knock Out Experiment
-use mutagens to knock out the function of certain genes, then observe any impacts on behaviours
eg: knocked shit off Drosophilia, got stuck after humping, made dunce and amnesiac flies too

3)Artificial Selection
-choose parents with most extreme values for traits
-if polygenic, should expect offspring to show intermediate traits
Define HERITABILITY?
-the proportion of variation of a characteristic that is attributable to genes (varies 0-1)
-the greater the genetic variance, the greater the potential for natural selection and artificial selection to act
-estimates can be confused by environment genotype interactions
-estimates of heridity may be confused by genotype, environmental interactions
Give an example of HERITABILITY.
-Mallards may build nests out of different grasses in different location, but this may result from local availability of plants, therefore the choice of plant has 0 heritability
-many characters may have gentic and environmental components
How is HERITABILITY investigated?
1) Correlation of trait between parents and offspring

2) Selective breeding of extreme phenotypes over several generations

eg: switching the eggs of 3 different nests
In terms of behaviour developement, what is MATURATION?
-normal changes during growth and developement not correlated with experience but which give the appearance of learning
Give an example of BEHAVIOURAL MATURATION.
Colour preference of frogs
-frightened frogs avoid green and prefer blue (b/c blue = water = safety)
-...but tadpoles prefer green (green = shelter = safety)
-...do they learn to avoid green??? NO!!!
-the changers correspond with changes in sensory cells and pigments in the eye with age

******************************
Chick foodpecking
-practicing vs no practicing makes no difference
Define LEARNING
-a specific change in behaviour as a result of experience with an external event or a series of events in an individuals life
What aspect of learning did PSYCHOLOGISTS study?
-looked at permanancy of changes
What aspect of learning did PHYSIOLOGISTS study?
-looked at changs in neural substrates
What aspect of learning did ETHOLOGISTS study?
-looked at how changes are adaptive
What are the 9 TYPES OF LEARNING?
1)Habituation
2)Sensitization
3)Classical Conditioning
4)Operant Conditioning
5)Conditioned Taste Aversion
6)Latent Learning
7)Observational Learning
8)Insight Learning
9)Imprinting
What is HABITUATION?
-a type of learning
-gradual fading of an unlearend response when a stimulus proes to be irrelevant
-an adaptive way of dealing with lots of irrelevant inputs (ie: are you feeling your clothes?)
-animals seem to habituate to all stimuli except for pain
-most primitive and universal form of learning
What is SENSITIZATION?
-a type of learning
-increase in responsiveness to a stimulus
-involves a relevant stimulus (eg: visitors to birds nests are ignored at first, but with repeated visits, birds get more alarmed)
What is CLASSICAL CONDITIONING?
-a type of learning
-animals become conditioned (learn) to respond to a new stimulus with a response that previously occurred to a different stimulus
-eg: Pavlov's dogs

Terminology:
-bell is the conditioned stimulus (CS)
-food is the unconditioned stimulus (US)
-salivation in response to food is the unconditioned response (UR)
-salivation with the bell is the conditional response (CR)
What is OPERANT CONDITIONING?
-a type of learning

eg: cat in box must push a lever to escape
-at first, cats reached through slots, scratched walls, etc
-eventually hit lever to escape
-subsequent trials, activity is concentrated around the lever
-it eventually learns and escapes whenever
-AKA 'Reinforcement of Desired Behaviour' (like training dogs)

-associate action with reward
What is CONDITIONED TASTE AVERSION?
-a type of learning
-learning to avoid certain foods if animals become sick
-helps to avoid poisoning
-may be a sub category of operant conditioning; but only takes 1 trial and effect is long lasting
-considered a management tool to control activities in the wild (eg: allow coyotes to feed from lamb carcasses laced with distasteful chemicals... learn to avoids lambs as prey items)
What is LATENT LEARNING?
-a type of learning
-experience gained is used to modigy behaviour later on
-learning without reinforcing a stimulus
-eg: rats allowed to wonder around a maze with no food reward... later, they will perform better than naive rats in a maze with food
What is OBSERVATIONAL LEARNING?
-AKA SOCIAL LEARNING
-AKA CULTURAL TRANSMISSION
-for young individuals, probably big advantage to copying adults of the same species
-therefore avoid energy and time costs of trial and error learning

Used for:
-food choice
-tool use
-movement patterns
-predator avoidance
-mate choice
-etc.
What is INSIGHT LEARNING?
-a type of learning
-an animal finds a solution without any previous experience to it, trial and error not the cause
-mental modelling and therefore capacity for reasoning
What is IMPRINTING? What 4 things can you imprint from?
-a type of learning
-occurs early in life during a critical period
-is retained for a long time (life)

-used to learn the properties of a stimulus towards which the animal will direct behaviour
i)Filial imprinting
-young imprint from parents
ii)Sexual imprinting
-learn from crew who to mate with
-has implications for conservation efforts
iii)Kin
-learn odours of individuals in their burrow
iV)inanimate objects
-food type... snakes imprint on prey types
-habitat... salmon imprint on stream chemistry
What is PLAY?
-motor activity performed after birth which appears purposels, in which motor patterns from other context may be used
-when it occurs, more dominant in young then in old
-may take 5-20% of the energy budget (not used for growth or metabolism)
-also risks of injury and predation
Why do animals PLAY? 3
These are Hypotheses...
1)Play helps to practice skills used later in life
-eg: with cheetahs, there were some correlations with play and predator attack skills but direct links were weak
-another problem... not all play resembles adult behaviour
-another problem... why bother with 'playful' practice and not real practice?

2)Play generates new and novel skills that may be useful later
-form of latent learning

3)Play helps with socialization and helps developement of communication
How is FOOD CHOICE learned by OBSERVATIONAL LEARNING?
-Steinberger placed poison bati in wild rat colony. Numbers initially declined, but then increased again
-a few rats did not eat enough poison to die, they became taste averse
-offspring didn't even taste the baits
-Steinberger thought the adults were placing warning pheramones on the baits
-later shown that jubeniles copied where and what adults ate (learning)
-young mammals may also learn food preferences throught he scent of mom's milk
-they also learend by smelling the breath of other rats
How is FOOD GATHERING TECHNIQUE learned by OBSERVATIONAL LEARNING? 4
1)Learn a specific task or method (maybe via observing)

2)Learning
-eg: chimps use grass blades to eat ants; Galapagos finches used cactus spines to probe bark crevices

3)Learne Route to Foraging site
-don't need obvious landmarks to get to site

4)Public information
-refine a behavior (not learning a specific task)

-learning about environmental variability to help in food searching
i)use which direction other group embers go to forage, later follow the same direction
ii)indicate the profitability of a food patch
-B2 allowed a sampling period on the food cups, B1 (naive) allowed to forage alongside B2, if B1 and B2 give up, it also quickly gave up
How is MATE CHOICE learned by OBSERVATIONAL LEARNING?
-females will want to mate with males that mated with other females

-this means that there are few males with huge reproductive success, and many with little
-tends to lead to runaway selection for certain male traits and linked to female selection
In general, is copying good or bad?
-bad
-a population with a minority of copiers is most stable
-otherwise, if copying is too common, copiers copy copiers, not accurate 'choosing' individuals
When should animals choose to COPY?
1) When learning is costly (eg: predators)

2) When the environment is fairly stable. Otherwise learned info becomes irrelevant
If animals choose to copy, who should they copy?
1) Copy the successful individual
2)Copy the older individual
3)Copy the ones better than you
What is PHYLOGENY OF LEARNING?
-natural selection shapes learning for each animal according to animals ecological needs
-not necessarily for 'human like' learning
-learning has a basic genetic basis too "innate vs. learned"
When should learning evolve?
1) If environments don't change, then a fixed genetic (innate) method may be more efficient

2) If environment is totally unpredictable, than learning is also of no value because past experience doesn't help in the future

3) Learning in intermediate environments
-learning should be selected when individuals face new conditions in each generation (gives flexibility). But not so random within a lifetime so that learning is useful

**See Jan.30 for table**
What is PHYSIOLOGY OF LEARNING?
-like HABITUATION
-touching the siphon of a sea slug causes the gill to withdraw, but a slug touched repeatedly habituates and gills don't withdraw
-sensory neurons fire, but action potentials result in less neurotransmitter being released from the presynaptic cell
-transmitter release is modulated by calcium
-less transmitter because less calcium influx
What 3 factors should be considered when OPTIMIZING FORAGING?
1)What to eat?

2)How Long to stay in patch

3)Predation risks.
When optimizing foraging, WHAT SHOULD AN ANIMAL EAT? What are the 2 types of eaters?
-want to maximize (net energy/time) while foraging

2 General Types
A) Energy Maximizer: try to eat as much as possible (eg: spider)
B) Time Minimizers: try to meet a certain energy requirement as fast as possible

Net Energy = energy in (search time and handling effort and time) - energy out (energy content and digestability)
What is RISK SENSITIVE FORAGING?
-basic idea is, should you stay with an ok but known amount, or should you go for the big reward but risk not getting anything?
-Weibe explained this with sausages......

-to buffer risks: store as fat but risk predation costs if too fat; cache food (store food for another day)
When optimizing foraging, HOW LONG SHOULD AN INDIVIDUAL STAY IN A PATCH? ie: How do you get diminshing returns in a patch?
Diminishing Returns in a Patch b/c:
1)Largest prey eaten first
2)Most vulnerable prey eaten first
3)Prey #'s get depleted so they become harder to find
4)Prey hide themselves
5)'Loading Effect' (eg: pelicans pouch fillis with fish so difficult to hold more)

**Cycle of: search->encounter->exploit
What is the MARGINAL VALUE THEOREM?
-it plots energy intake in the patch vs travel time and time on an offset x axis.
-the intersect between the tangent and graph line can be correlated to find the GUT (Give Up Time)
-slope of the tangent line is (energy load)/(travel time + foraging time) = rate of energy gain in the patch
What are 4 RULES OF THUMB in MOVING BETWEEN PATCHES?
1)Number Rule
-leave after catching 'n' prey

2)Leave after spending 't' time units in patch

3)Giving up rule
-leave after 't' time units of unsuccessful searching

4)Rate Rule
-leave when capture efficiency drops to critical value 'r'
What effect does PREDATION RISK & FORAGING have on feeding habits?
-affects what is eaten and where, also behaviour
-lots of animals consider prey risk heavily and will change feeing aspects (ie: find cover)
-larger groups increase vigilence levels, spot predators earlier, more time to escape
-individual vigilence decreases with group size (inverse relation)
What is SEARCH IMAGING?
-animals learn certain stimulus characteristics of food they encounter. This improves their chance of finding similar food items in the future
-may also involve stimulus filtering or selective perception
-has been debated: improvements in capture, can also be explained by other mechanisms
What is AREA RESTRICTED SEARCHING?
-if prey is patchy, forager should slow down and turn to more often encountering prey items
How do NICHE SCALES affect foraging?
-foraging can be influenced by competitors
-also influences food type and therefor foraging efficiency

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