Learning Relatively permanent modification of behavior that occurs through practice or experience Learning is not maturation Learning is not always observable Associative learning vs. non-associative learning

Habituation - relatively persistent waning of a response, results from repeated stimulus not followed by any reinforcement, a process of learning to ignore irrelevant stimuli Benefit - save time by protecting animals from responding the irrelevant stimuli, to concentrate on more important stimuli

Delayed conditioning (1/2~2 sec.) Simultaneous conditioning Trace conditioning (no overlap) Backward conditioning Time as a stimulus (zoo animal) e.g. red light, alarm call

Operant conditioning (instrument) Trail-and-error learning Response is emitted by reinforcement, it is rewards which cause animals to respond e.g. chimp fishing termites e.g. search image

Reinforcement - anything that alters the probability of behavior Positive vs. negative reinforcement Schedule of reinforcement: Fixed ratio, variable ratio, fixed interval, variable interval Extinction and spontaneous recovery

Original principle of learning Learning is a unitary trait - General process theory Natural scale of learning ability Equivalence of association - Principle of equipotentiality

Reinforcement is required for learning - Law of effect Association strength - more reinforcement stronger learned response

Rebuttal of principle of learning Can't fix animal to scale Latent learning--association made with neither immediate reinforcement or reward nor particular behavior evident at the time of learning. – e.g. digger wasp, rat in a maze Preparedness

Complex learning Avoidance learning (aversive conditioning) – R* don't come immediately – One trial learning – Long lasting effect Biased learning

Latent learning Insight learning - the animal makes new associations between previously learned tasks in order to solve a new problem Imprinting – Acquired preference and development predisposition

Social learning – local enhancement: locate foraging sites by attending to others – local facilitation: animals feed faster in a group – observational learning: observers modify behavior after demonstrators – imitation: observers match behavioral action and goal

Learning set - the acquisition of a learning strategies, or given a series of problems, an animal will transfer some of what it has learned about solving the first problem to the solution of subsequent problems in a series. Constraints of learning – preparedness – Methods constraints

Cost and Benefit of learning Cost-- – Take time – More neural complexity – more vulnerable until learning is complete – increase investment by parent – can be fooled

Benefit-- – Ability to cope w/ a range of event (high adaptability) – Programming everything is too costly

Development of behavior Question of ontogeny – how behavior changes over lifetime of an individual? Seek to identify the factors influencing the acquisition of behavior

Food of larvae determines if they become queens (behaviorally and morphologically) Environmental sex determination: incubation temperature 2M males more likely to attack stranger at 90 days than 0M males 2M females have larger territory, are more aggressive and less attractive to males than 0M female

Seed storing in marsh tit Hand-reared individuals allowing to store seeds at different developmental stages have larger hippocampus (region ~ spatial learning) than control (no experience)

Effects of environment A variety of environmental cues seem to act as developmental “switches” between behavioral phenotypes e.g. caste switching in bee D1 ~12: young adult, clean nest D13 ~ 20: mid-aged adult, brood & queen care, nest maintenance, food storage D20 ~: old adults, foraging

Interactions within hives can change timing of behavioral switches Many workers of same young age, some remain nursing till later, some become foragers earlier When younger bees add to colony, young residents become precocial foragers When older bees add to colony, young residents do not become precocial foragers

Sequential hermaphroditism Normal pattern = sex changes related to size, but in some species developmental change triggered by social cues In gobies, change may occur based on the size and sexes of new partners (usually smallest partners is female)

In anemonefish, largest females dominate group; when largest female removed, larger males switch to female

Social deprivation in rhesus monkey Total isolation Isolation + cloth or wire surrogate mother Peer group W/ mother only W/ mother, but w/ varying periods of separation at specific age interval Small social group (control) Behavior homeostasis

Methods in behavior development 7 parameters of treatment – Age when treatment is given – Type or quality – Duration or quantity – Age of testing – Type of test – Test for the persistence – Test different strains or species

Testing procedure Longitudinal Cross-sectioned Deprivation, enrichment, alter the quality of the stimuli