Animal behavior Exam 2

Learning

An enduring change in the mechanisms of behavior that result from experience and environmental events.

• not directly observable

• selective process where consequence is a punishment or reward

• anatomical and physiological changes underlay learning

Functional definition of learning

A process that allow animals to adjust to their environment through individual experience.

Ex: organisms need to know what to eat, as well as where and when to find it.

Performance

Change is behavior based on opportunity and motivation (not learning). (Effects)

Ex: rats full and no longer respond to lever in a Skinner box

Phenotype Plasticity

The ability of an organism to produce different phenotypes depending on the environment. A broader category that includes learning phenomena

Phenotype

Physical appearance (resulting from genotype plus the environment)

General Learning Phenomena

-Non-associative

-Associative/ Cognitive:

Non-associative

Habituation and Sensitization. Behavior changes that respond to/ require a stimulus.

Ex. come from ethology

Associative/ Cognitive

Classical Conditioning and Instrumental Conditioning. Behavior changes, pairs cues with a reflexive response

Specific Learning Phenomena

Human language, song learning, imprinting

Non-Associative Learning

Reactions to stimulus events in the environment (hard-wired response)

Includes: Habituation and Sensitization

Taxis

Reflexive locomotion and orientation to a stimulus

2 Types: Phototaxis and Geotaxis

Phototaxis

Orient to light

Ex: cockroaches move away from light (negative phototaxis)

Geotaxis

Orient to spatial/ physical position

Ex: Upside down catfish in an aquarium

Ex: mice on a slope to test anxiety and turn to face upward (negative geotaxis)

Orienting Response

Innate response to a novel stimulus

Ex: Infants turn their head and gaze at an unfamiliar visual stimuli

Ex: Dogs cock their head in response to novel stimuli

Ex: slapping on a desk and everyone looks up

Fixed Action Patterns

Species specific stereotypical response; dependent on sign stimuli acting as a "releaser"

Ex: Tinbergen's Stickle Back Fish experiment- males attack red bellies or any red item to elicit a territorial or attack response

Ex: Going into stores and buying bigger and better things vs. the smaller, less nice things

Habituation

Decrease in responsiveness after repeated exposure to a stimulus; ignore repeated stimuli

Ex: Ignore sounds of a highway if you live nearby it

Sensitization

Increase in responsiveness after repeated exposure to a stimulus

Ex: Relapse in drug use- take it in increased effects

Distinction Between Sensitization and Habituation

-Strong stimulus- sensitization predominates

-Weak stimulus- habituation predominates

Dual Process Theory

-Habituation and sensitization are not mutually exclusive; they operate together

-Two neural processes work together; the behavioral outcome depends on which system is most active

-Reduce reactivity to irrelevant stimuli or increase to relevant stimuli

• Ex: prey animals freak out or ignore presence of a predator like a lion- if they are about to become prey (freak out) or are at a watering hole (ignore presence)

Sensitization in Aplysia

Experiment by Erick Kandel (Neuroscience Nobel Prize Winner)

-Examined the neuro-network in Aplysia (who have a ganglia type of nervous system), have simple reflex of behavior.

-Examined aplaysia's gill withdrawl reflexes. Touch the siphon and gill contracts but do it minutes later, it doesn't respond as largely (less sensitized); do it multiple times and it could not react as sensitively for a couple of days

-Short and long term type of sensitization and habituation (ex. touching roly-polies they respond the first time and curl up but after that they don't anymore to each touch)

• No cues involved

Classical Conditioning

Associative learning of stimuli and stimulus responses. Pairing of two stimuli endows one of them with the ability to act as a signal for the other

• Pavlov Conditioning

• associative form of learning

• Basis is reflexive

• Many varieties

  • Ex: appetitive learning and aversive learning

Example of Classic Conditioning

UCS- food

CS- biological stimulus ex: bell

CR- reaction ex: salivating

Give a dog biscuits and drools before the biscuit (CR) but if he drools with the biscuit (UCR)

Appetitive Learning

Unconditioned stimulus is pleasant

Ex: food, finding mates/ sex

Aversive Learning

Unconditioned stimulus is unpleasant

Ex: electric shock, avoid toxins and hazards

Stimulus Contingencies

~Excitatory Conditioning: the presence of one stimuli is followed by another; Example: dog hears a sound and knows it's gonna get food

Excitatory Conditioning

the presence of one stimuli is followed by another

Ex: dog hears a sound and knows it's gonna get food

Inhibitory Conditioning

the presence of one stimuli is not followed by another

Ex: dog hears a sound but is not gonna get food

Example of Human Classical Conditioning

In Japan, people are conditioned to move out of the street at the sound of a bicycle bell ringing. One man videoed him ringing a bicycle bell everywhere even though he wasn't on a bike and people moved to the side.

CS: bell ringing

UCS: bicycle coming towards you

CR: move at the sound of the bell

UCR: move out of the way of the bicycle as they ring the bell

Second-Order Conditioning

Conditioned Stimulus (CS) serves as an unconditioned stimulus (US) for a new cue

CS{1} --> US

CS{2} --> CS{1}

Ex: light predicts the bell which predicts the food, until food associated with the light

Compound Conditioning

Presentation of 2 conditioned stimuli (at the same time); biologically prepared

Includes overshadowing, latent inhibition, and blocking

Ex: CS{1} (light); CS{2} (tone)

Ex: CS relevancy (belongingness)

Overshadowing

Compound stimulus in which one is more salient

Latent Inhibition

Preexposure to CS results in slower learning; possibly due to irrelevance of cue or context learning

Blocking

-Leon Kamin showed that a previously learned CS1–US association prevents learning that CS2 also predicts US

-CS{2} in combination with CS{1}; compound CS

-Conditioning only occurs if cue is both useful and non-redundant predictor

-US has to be surprising to produce conditioning

Ex: Phase 1- Light CS, shock UCS

Phase 2- Tone CS combined with Light CS, shock UCS

Phase 3- Tone CS --> little or no CR (learning is blocked)

Light CS --> Big CR

Taste Aversion Learning

-A functional and adaptive aspect of classical conditioning

-one trial learning that a novel taste is a powerful cue for illness

Garcia-Koelling taste-aversion study: Avoidance with tone/ light (CS1) and saccharine flavor water (CS2) + shock (US1) and Lithium Chloride (US2)

-Taste cue, better for learning to predict illness; audiovisual cue, better predictor of shock.

Example of Taste Aversion

Not going to a restaurant after getting sick or not drinking a type of alcohol after a hangover; or give coyotes LiCl laced sheep carcasses to deter them from the flock by making them sick

Sexual Conditioning

-A functional and adaptive aspect of classical conditioning

Michael Domjan

Examined quail mating, cues predicting access to mating opportunities increase mating success in Japanese quail, cues allow for increased reproductive success

3 Components of Instrumental Learning

-Stimulus (SD)

[A discriminative stimulus is a cue, not a US or CS; a signal for when response will lead to consequence]

-Response (R)

[Voluntary-creation of new behavior defined by what is required to obtain reinforcer or avoid punisher]

-Consequence (SR)

[Reinforcer {strengthens} or punisher {weakener}]

Instrumental Learning

Selective forcer for behavior; probability of behavior increasing or decreasing, associative learning

Types of Instrumental Learning

-Positive & Appetitive

-Positive & Aversive

-Negative & Appetitive

-Negative & Aversive

Positive & Appetitive

Positive reinforcement; SD --> R --> SR; R increases, example: food reward, water, sex, etc

Positive & Aversive

Punishment; SD --> R --> SR; R decreases, positive contingency, example: spanking

Negative & Appetitive

Omission; SD --> R --> no SR; R decreases. Negative contingency, behavior doesn't follow, punishing by what the individual wants doesn't occur, example: time out from play, dunce cap

Negative & Aversive

Negative reinforcement; SD --> R --> no SR; R increases (escape/avoidance), prevents aversive behavior, examples: seat belt warning signal, choke chains, electric shock, and avoiding mimicry

Example of Operant Conditioning Reinforcements

Positive Reinforcement: Sheldon gives Penny chocolates to stop annoying behaviors while watching television

Negative Reinforcement: spray with water from a spray bottle

Example of Operant Conditioning

Rats in a skinner box. Floor is electrified (negative reinforcement). Rat pushes lever to receive food into the box to increase the pushing lever response. Prime first with free pellets. Increased response to push lever with food reward (positive reinforcement)

Thorndike's Law of Effect

First attempt to explain reinforcement (i.e. the first question).

-A reinforcer is a stimulus that produces a "satisfying state of affairs," but did not explain why it is reinforcing- calling a reinforcer a "satisfier" is merely giving it a label.

-Establishes an association between the response (R) and the stimulus (S) in the presence of which the response is performed (reinforcer is satisfying (increase R) or unsatisfying (decrease R)).

Edward Thorndike

Experimented in animal intelligence with putting cats in puzzle boxes (escape by pushing the peddle). At first the cat did behaviors that led to no positive outcome, but increased trials increased efficiency. This led to him creating his Law of Effect.

Clark Hull Drive Reduction Theory

• Proposed that what makes something an effect reinforcer is its ability to return the organism to the homeostasis point.

• Motivation is a drive reduction brought about by a physiological need.

• Food is an effect reinforcer because it reduces the hunger drive.

-Primary reinforcers: stimuli that are capable of reducing biological drives, strengthening response (reduction in motivation).

-Secondary reinforcers: stimuli associated with drive reduction thru classical conditioning (smell of food) [cue associated with primary reinforcer].

• Thus we have acquired drives and any response that decreases the intensity of acquired drives is reinforcing. Ex of secondary reinforcers: clicker training, money

Reinforcer

Anything that reduces drive

David Premack

-What makes eating different than pressing a lever for a rat (reinforcers as responses)?

-Proposed that the opportunity to perform a higher probability response will serve as a reinforcer for the lower probability response.

• Ex: a food deprived rat is more likely to eat than press a lever if given free-access to both activities.

• Doesn't require a drive reduction mechanism to explain reinforcement but look at the baseline probabilities.

• Ex: clean the dishes before going outside to play

Example of Premack Principle

1. Rat is water deprived

2. Rat drinks more than it runs

3. Drinking reinforces running

and vice versa

1. Rat is not water deprived

2. Rat runs more than it drinks

3. Running reinforces drinking

Instinctive Drift

Instinctual behaviors compete with learned behavior. A discriminative stimulus can become associated with an US that produces instinctual responses. These instinctual responses become competing responses to the learned response. And the evolutionary behavior of an animal will take precedence over learned behavior.

Misbehavior of Organism (1961)

-Example of Instinctive Drift

• Breland and Breland (were Skinner's students) who worked as animal trainers, realized can't train behavior, and wrote this

-Evidence that contradicts general principles of learning assumed in behaviorism

-jab at Skinner's "Behavior of Organism"

• Ex: motor response by pigs being trained for a commercial; they chewed on coins but wouldn't give it back because they like to root things around

Aversive control

-Escape learning is the performance of a response to terminate an ongoing aversive stimulus

• Un-signaled responses

Avoidance learning

The performance of an escape response in order to prevent the aversive stimulus

• Signaled stimulus or warning signal

• Basis for evolution of aposematic

Two forms of avoidance

– Passive avoidance
– Active avoidance

• Shuttle box studies

How does a CS influence the strength of an Operant Response

Generally regarded as two separate processes; Need to understand how they work in combination.

CS elicit a reflexive response that might interfere with or modulate an operant response.

Instrumental- reflexive, automatic response; CS within are processes that need to be processed together

Two ways that Respondent Conditioning influence Operant Behavior

-Motivational Effects: changing motivational support for behavior, increase response, ex: going to a happy place

-Response-Cueing Effects: cues that are associated with an operant response, ex: encourage animal behavior by cueing that food is coming

Comparative Learning

-Where psychology and evolutionary biology overlap. Includes Ecological View (Bio.) and General Process View (Psych.)

-Distinguishing between mechanism and learning phenomena (divergence, homology, homoplasy)

-Divergence in learning mechanisms is demonstrated when species differences in learning phenomena cannot be attributed to non-learning contextual variables (e.g., memory, motor control, motivation).

Homology of learning mechanisms

Demonstrated when equivalent phenomena are shown to depend on the same processes at all levels of analysis

Homoplasy of learning

Demonstrated when learning phenomena evolve independently under similar selective pressure (convergence, parallel, reversal)

Some Learning Phenomena Found in Mammals and Honeybees

-Overshadowing

-Within-compound association

-Conditional discrimination

-Successive negative contrast

-Partial reinforcement extinction effect

-Second-order conditioning

-Spontaneous recovery in extinction

-US pre-exposure effect

-Escape and avoidance learning

• show ecological and general process views

Two-Factor theory of avoidance learning

-Mower (1947) proposed that avoidance learning
involved two fundamental learning processes

• Classical conditioning of fear via aversive (US) paired with (CS)

• Instrumentals conditioning of avoidance behavior via negative reinforcement

-Kamin (1957) showed that avoidance depends on the termination of the warning signal (CS)

Ecological View (of Comparative Learning)

Learning mechanisms solve specific ecological problems. Species vary in their ecology thus, divergence in mechanisms should be widespread and associative selectively should be common. Examples: taste aversion learning, imprinting, and song learning occur during a sensitive period.

General Process View (of Comparative Learning)

Most ecological niches involve causality, space, and time; Learning mechanisms deal with such common dimensions thus, divergence in mechanisms should be rare and similar phenomena should appear in very different linages. Examples: contextual factors account for many examples of divergence; conditioning occurs in all animals with a CNS

Models of Evolution of Learning

-Assumption in models: costs of learning, learning defined as heritable trait (genetic basis).

-Environmental stability: low and high variability in environments favor a fixed genetic rule with low cost rate. Medium variability environments favor learning over transmission of fixed genetic rule.

Ex: organism living in a different daily environment vs. living in a daily same environment- if everything is different, then it's not predictable and learning wouldn't take place, but if everything is the same, then there would be physical cost

Stephen's Model (1991)

Within lifetime vs between generation predictability.

-2 forms of environmental predictability

-Learning is supported by high within-life time and low between-generation predictability

-Due to confabulation due to forms of variation; can't learn in the other quadrants due to learning costs

-Anytime you add a psychological concept, there will be a physical cost

Learning About Predator

Learning to avoid predator increases survival directly and indirectly.

• Trade-offs between foraging and mating behavior with risk of predation.

Ex: Chiver's Damselfly learning (1996)

• Predators (pike) were fed 1 of 3 diets (minnow, damselflies, and mealworms- control)

• Damselfly larva exposed to tank water in which specific diets were consumed.

• Foraging was reduced as a function of learning predator diet cues.

Avoidance learning by damselflies, chemical transfer cues alter damselfly behavior due to water exposure

Learning About Mates

-Males and females learn about mate pair-related cues (odor cues in Mongolian gerbils)

-Parental investment and learning about mates (Hollis and Domjan, 1988)

Learning depends on respective amount of parental investment (positive correlation between learning and investment level)

If males and females equally invest in off-spring both learn mate-related cues

Females that provide high parental investment are valuable to males, thus males undergo selective pressure to learn about female cues

Males need to learn about females in order to survive, but females will always find a mate

Learning about Familial Relationships

Social recognition in which animals respond selectively to their relatives. Kin recognition learning has adaptive value for cooperative and altruistic behavior.

Ex: Kin-Recognition in birds. Siblings assist in rearing of younger offspring of parents. Playback study of churr calls in long-tailed tits (Contact call is learned, show preference for familiar churr call, cross foster study confirms learning) (Hatchwell and Sharp)

Learning about Aggression

Intrinsic aggression: inherent fighting ability. Extrinsic aggression: winning and losing effects (change in probability of future aggressive acts)

Ex: Hollis and the blue gourami (very aggressive fish)

Tested cues associated with territorial defense in gourami fish. Increased behavior. Predicted winner and loser effects (instigated by ability to learn or not learn).

Paired males together, gave one light cues to predict a territorial, aggressive encounter making him more likely to win that fight over the non-cued. Increased cues= Increased wins. Take winners with new losers and they'll continue to win.

Animal Cognition

esoteric, animals think and have their own mind, consciousness, and higher order mental processes

Comparative Cognition

Looking for divergences, homologies, homoplasies, etc

Emergence of the cognitive approach:

-Failure of S-R model (Behaviorism) to explain complex behavior (input or output, doesn't measure in between)

-Investigation of mental processes in other areas of psychology

-New technologies/method of research in brain function

-Fundamental concept of cognition is mental representation (i.e., coding of information)

-Mental activity is bottom-up or top-down

-Animals form perceptions, mental presentations Ex. think about something sad but don't have to make you sad

Behaviorist Folly

-Rejects the argument that consciousness is beyond the realm of science; animals appear to have intentions, beliefs, and self-awareness

Ex: injury feigning in kill deer and nesting plovers (fake a broken wing to fool predators, know what the predator is thinking?)

-How consciousness evolved is problematic because natural selection is thought to act only on the behavior (what they do) not on the subjective state (what they experience)

Emergence of the cognitive approach (Darwin’s argument):

"It is highly probable that with mankind the intellectual faculties have been mainly and gradually perfected though natural selection.... Undoubtedly it would be interesting to trace the development of the separate faculty from the state in which it exists in lower animals to that which exists in man. (1874) [based on common ancestry, if we can show it then animals can too]

Behaviorist Folly

"First step is to measure whatever can be easily measured. This is okay as far as it goes. The second step is to disregard that which can't be measured or give it an arbitrary quantitative value. This artificial and misleading. The third step is to presume that what can't be measured easily isn't very important. This is blindness. The fourth step is to say what can't be easily measured really doesn't exist. That is suicide." -- Daniel Lovich (in Smith, 1972).

• only want to measure what's easy and ignore what isn't; form of blindness (blind in that animals don't have minds); saying cognition doesn't exist

Consciousness and Animal Cognition

Perceptual vs reflective consciousness

-"umvelts" perceptual world of animals (Romanes- everything has a consciousness (umvelt))

-are animals aware (reflective) of their own cognitions? (metacognition- do you know what you know and don't know, thinking doesn't imply consciousness, animals have a mind for their niche)

Consciousness implies thinking, but not the other way around

Argues not why do animals have a mind, but why do animals have this particular mind.

Functional similarities of behaviors/cognitions is a better approach than assuming the subjective mental states of animals.

Cambridge Declaration on Consciousness (2012)

We declare the following: "The absence of a neocortex does not appear to preclude an organism from experiencing affective states. Convergent evidence indicates that non-human animals have the neuroanatomical, neurochemical, and neurophysiological substrates of conscious states along with the capacity to exhibit intentional behaviors. Consequently, the weight of evidence indicates that humans are not unique in possessing the neurological substrates that generate consciousness. Non-human animals, including all mammals and birds, and many other creatures, including octopuses, also possess these neurological substrates."

-revamp what's appropriate for investing cognition; animals have same capacities as humans, animals have their own consciousness; won't accept outdated data, no neocortex doesn't mean that animals don't have emotional & intellectual states

Metacognition

Thinking about thinking

-how well do you know what you know or knowing how well you remember something

-hierarchal executive control processes that oversees lower-level cognition

-self-awareness of subjective states due to uncertainty

-declarative consciousness (because humans are conscious of their states and knowing and can declare them to each other)

Example of metacognition

Ex: Duration-discrimination test—offering rats rewards for classifying a signal as either short or long (Crystal & Foote, 2007)

-Tone presentation short vs. long

-Options:

• Correct choice receives large reward

• Wrong choice receives no reward

• Opt out choice receives small reward

-Discrimination that are difficult were opted out thus suggesting rats know what they don't know

-showed rats how what they knew when easy to discriminate auditory cue, and opted out when they didn't know as it got hard to tell, minimized no reward from happening; evidence of thinking even if small neocortex

Self-Awareness

If you are self-aware then you are in a position to use your experience to model the existence of comparable experiences in others.

Ex: when you see someone who is carrying a large box approach a door, you might be prompted to walk ahead of them and hold the door open. This is based on a set of inferences you make as a consequence of having found yourself in comparable situations

Ex: Gordon Gallup (1970) Mirror test for self-recognition in primates (capacity for self awareness which is developed not born with)

• Monkeys respond socially to self reflection

• Gorillas fail test

• Dolphins, Chimpanzees, Orangutan pass test

• Humans 18 -24 months old

-do animals recognize themselves in a mirror? -control- anesthetize, non toxic dye unseen, wake up to mirror, do they examine the mark?

Memory

Mental time travel (chronesthesia) is the awareness of past and future and the use of that information in the present.

Two types: Semantic and Episodic

Semantic Memory

A form of long term memory that involves memory of *facts and non-declarative procedural skills and abilities (not a form of mental time travel)

Episodic Memory

A form of long term memory that involves what, where, and when; uses autobiographical details.

-several studies attempt to demonstrate this memory in animals

-animals have semantic but what about episodic?

Example of Episodic Memory

Nicola Clayton investigated food caching behavior in scrub jays (scrub jays bury worms and nuts in sand filled ice cube tray compartments using visual and auditory spatial cues) a divergence concept

Experimental design of Nicola Clayton’s experiment

Two stages

-Phase 1: cache food

-Phase 2: food recovery (either 4 hrs or 124 hrs)

Results: Jays tend to worms food 4 hrs later (before the worms spoil) and tend to recover nuts 124 hrs later (after worms are spoiled). Also if a thief is nearby they'll cache the food in the quieter sand vs. the loud gravel

Evidence: for what (food type), where (cache location), and when (length time before recovery phase); shows animals may possess rudimentary episodic memory

Using and Understanding Tools

The use of an external object as a functional extension of the mouth, beak, hand or claw in attainment of and immediate goal (van Lawick-Goodall 1970) ex. opposable thumbs

Folk Physics

The implicit/ innate understanding of how and why tools work

Basic Questions of Using and Understanding Tools

How is a tool acquired?

What do animals understand about tools?

Can animals construct tools?

Also involves development of motor skills to use tool properly

Trap Tube Case Study in Tool Learning

Visalberghi and Trinca (1989)

Test what the animal knows about the tool

Results show approx. 50% success rate

• Distance-based association rule: learning the position of the food relative to the trap

Seed (2006)

Improved trap tube design - test of conceptual understanding

-Involved a trap tube, with a stick, a monkey or crow decides which end to get the food out of, involves distance- related cues.

How does tool use develop?

Perceptual and motor skills, without causal understanding.

3 kinds of learning:

1. Instrumental learning- Animals tend to improve with experience

2. Imitation-True imitation is rare, but many observational learning studies involve learning tool use. Or do they learn during trial and error

3. Insight (aha moment)- sudden comprehension of problem

Kohler (1959) test of insight in chimps

-give chimps boxes and sticks to reach a suspended banana, insight "aha" is gained spontaneously after unsuccessful attempts

Ex: Epstien (1984) S-R learning, challenged Kohler

-elements of solution learned then combined spontaneously (automatic chaining)

• pigeons learned to push a box and climb on a box to reach a banana (toy) for food

• tests required combining the two actions without any insight (instrumental learning)

Evolution of Language

Robin Dunbar (1993) suggested that rapid evolution of human intelligence is a consequence of language comprehension.

-humans have long complex language, animals don't; increased language increased neocortex size

-language did not emerge suddenly (suggesting animals posses rudimentary language), but it did lead to rapid cultural and intellectual changes in humans. [social intelligence is dependent on size of neocortex]

• language is hypothesized to be a more efficient means for establishing social intelligence than other social behavior such as social grooming

-larger animal group= larger neocortex

-positive correlation between sociality and brain size

-Conclusions:

• Knowing the neocortex ratio for humans, an extrapolation indicated that a typical group size for humans is 150 individuals

• Extrapolating again, humans should spend about 40% of their time grooming

  • Too much time, it would be inefficient because it would disrupt other tasks

• Language may have evolved as a more efficient device than grooming to establish social bonds

  • Easier to chit chat (groom)

Example of the Evolution of Language

Symbolic Thinking in animals (involves the ability to form mental representations)

Irene Pepperberg tested the ability to learn category concepts in African Grey Parrot (Alex)

-Verbal label are a form of referential language

-Alex demonstrated concepts of material, color, shape, same vs different, and numerical concepts

Language in Nonhuman Primates

-Washoe: a chimpanzee raised in human environment: First chimp taught American Sign language

-Koko: a gorilla trained to use modified sign language: Koko can reliably sign 500 words in GSL (Gorilla Sign Language)

-Nim Chimpsky: a chimpanzee trained to use sign language : Critical of language studies

-Kanzi: a bonobo chimpanzee: Artificial language on keyboard (lexigrams)

Characteristics of Cultural Transmission

Information passed from individual to individual and from one generation to the next. Unlike individual forms of learning, shows rapid learning within population and produces important changes in a few generations; evolutionary/ selection mechanism

Mechanisms of Cultural Transmission

Social learning, imitation, teaching, and social facilitation