Comparative analysis:

Basic concepts

Comparative analysis of learning and cognition

Sponges

Cnidarians

Flatworms

Arthropods

Vertebrates

Mollusks

Nematods

Annelids

•Neurons

•Radial symmetry

•CNS

•Bilateral symmetry

•Ganglionar

nervous

system

•Cranium/Brain

Taxonomic context

Key issues in the comparative analysis of learning and cognition

Issue # 1: distinguish between learning phenomena and learning

mechanisms

Issue # 2: homology of learning mechanisms is demonstrated when

equivalent phenomena are shown to depend on the same

processes at all levels of analysis.

Issue # 3: divergence in learning mechanisms is demonstrated when

species differences in learning phenomena cannot be attributed to

contextual variables.

Key issues in the comparative analysis of learning and cognition

Issue # 1: distinguish between learning phenomena and learning

mechanisms

Issue # 2: homology of learning mechanisms is demonstrated when

equivalent phenomena are shown to depend on the same

processes at all levels of analysis.

Issue # 3: divergence in learning mechanisms is demonstrated when

species differences in learning phenomena cannot be attributed to

contextual variables.

Psychological

Neurobiological

Neurochemical

Cell-molecular

Mechanistic Level Example Modular Representation

S→S learningAPPETITIVE FEAR

Coincidence

detectorsCEREBELLUM

S S S S

AMYGDALA

Synaptic

receptors

SPECIFICITY

NMDA GABA

Second-

messenger

systems ARACHIDONIC ACIDcAMP

Issue # 1: distinguish between learning phenomena and learning mechanisms

Key issues in the comparative analysis of learning and cognition

Issue # 1: distinguish between learning phenomena and learning

mechanisms

Issue # 2: homology of learning mechanisms is demonstrated when

equivalent phenomena are shown to depend on the same

processes at all levels of analysis.

Issue # 3: divergence in learning mechanisms is demonstrated when

species differences in learning phenomena cannot be attributed to

contextual variables.

Coover et al., 1973, J Comp Physiol Psychol, 85, 111-122.

Issue # 2: homology of learning mechanisms is demonstrated when equivalent

phenomena are shown to depend on the same processes at all levels of analysis.

•Lesions in the basolateral

nucleus of the amygdala.

•Sham: simulated lesion.

•Two-way active avoidance

training.

Avoidance learning in rats

Issue # 2: homology of learning mechanisms is demonstrated when equivalent

phenomena are shown to depend on the same processes at all levels of analysis.

Avoidance learning in goldfish

Master-yoked design Impaired by MP but no LP lesions

MP: medial pallium (amygdala homologous)

LP: lateral pallium (hippocampus homologous)

Homology or homoplasy?

Some learning phenomena found in mammals and honeybees

Overlearning extinction effect

Partial delay of reinforcement extinction effect

Overshadowing

Potentiation

Summation

Within-compound associations

Compound uniqueness

Conditional discrimination

Successive negative contrast

Partial reinforcement extinction effect

Positive behavioral contrast

Progressive improvement in spatial reversal learning

Dimentional transfer

Second-order conditioning

Spontaneous recovery in extinction

Latent inhibition

US preexposure effect

Escape and avoidance learning

Bitterman, 1988, Vertebrate-invertebrate comparisons.

Rats Honeybees

>600 Mya

(a planaria-like animal)

Key issues in the comparative analysis of learning and cognition

Issue # 1: distinguish between learning phenomena and learning

mechanisms

Issue # 2: homology of learning mechanisms is demonstrated when

equivalent phenomena are shown to depend on the same

processes at all levels of analysis.

Issue # 3: divergence in learning mechanisms is demonstrated when

species differences in learning phenomena cannot be attributed to

contextual variables.

Determinants of performance

Acquired

behavior

Learning

factors

Contextual

variables

Signal/Response

strength

Memory retrieval

Sensory/Perception

detection

Motivation

Motor control

Issue # 3: divergence in learning mechanisms is demonstrated when species

differences in learning phenomena cannot be attributed to contextual variables.

Two views of divergence

Ecological view:

•Learning mechanisms solve specific ecological problems

•Species vary in their ecology

•Thus, divergence in mechanisms should be widespread

•And associative selectivity should be common

Examples:

•Taste aversion learning seems to violate equipotentiality

•Imprinting and song learning occur during a sensitive period

•Some species show selectivity for a specific sensory dimension

Two views of divergence

General-process view:

•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 lineages

Examples:

•Contextual factors account for many examples of divergence

•Conditioning occurs in all animals with a CNS

•Conditioning may be involved in some otherwise specialized

cases

Morphological divergence in Hawaiian honeycreepers

Do learning

mechanisms diverge

like the beaks of

these honeycreepers?

Or are they as

conservative as the eyes

or the feathers of these

honeycreepers?

Case study:

Surprising nonreward

Comparative analysis of learning and cognition

Osteichthyes

(Goldfish)

Amphibia

(Toads)

Aves

(Pigeons)

Reptilia

(Turtles)

Mammalia

(Rats)

Phylogenetic relationships among fish, amphibians, and reptiles

Classic learning theory

Thorndike, 1898, Psychol Rev

Strengthening-weakening mechanism

Stimulus Response … Incentive

Stimulus Response … Nothing

Thorndike, 1911, Animal ingelligence

Emotional (and cognitive) factors

Tinklepaugh, 1928, Calif Pub Psychol

Drawing by Katsuo & Chiharu Tomita

Elliott, 1928, Univ Cal Pub Psychol

Successive negative contrast (SNC)

SNC

Sunflower seeds

Bran mash

Sunflower seeds

(for both groups)

Preshift Postshift

Group 1-9 10-16

Downshifted (9) Bran mash (6) Sunflower seeds

Unshifted (9) Sunflower seeds (6) Sunflower seeds

Consummatory successive negative contrast (cSNC)

Preshift Postshift

Group 1-10 11-15

32-4 32% 4%

4-4 4% 4%

Papini, 2006, Jap J Anim Psychol

(a) Toads

1.0

1.4

1.8

2.2

0 5 10 15 20 25

Trials

Me

an

Lo

g L

ate

ncy

S

L

L-S

(b) Turtles

1.0

1.4

1.8

2.2

0 5 10 15 20 25 30 35 40

Trials

Me

an

Lo

g L

ate

ncy

S

L

L-S

(c) Pigeons

0.0

1.0

2.0

3.0

0 5 10 15

4-Trial Blocks

ln I

niti

al L

ate

ncy

S

L

L-S

(a) Toads

1.0

1.4

1.8

2.2

0 5 10 15 20 25

Trials

Me

an

Lo

g L

ate

ncy

S

L

L-S

(b) Turtles

1.0

1.4

1.8

2.2

0 5 10 15 20 25 30 35 40

Trials

Me

an

Lo

g L

ate

ncy

S

L

L-S

(c) Pigeons

0.0

1.0

2.0

3.0

0 5 10 15

4-Trial Blocks

ln I

nitia

l L

ate

ncy

S

L

L-S

Amphibians (Toads)

SNC

Reversed SNC

(a) Toads

1.0

1.4

1.8

2.2

0 5 10 15 20 25

Trials

Me

an

Lo

g L

ate

ncy

S

L

L-S

(b) Turtles

1.0

1.4

1.8

2.2

0 5 10 15 20 25 30 35 40

Trials

Me

an

Lo

g L

ate

ncy

S

L

L-S

(c) Pigeons

0.0

1.0

2.0

3.0

0 5 10 15

4-Trial Blocks

ln I

nitia

l L

ate

ncy

S

L

L-S

0

2

4

6

8

10

0 4 8 12 16

Trials

Num

ber

of

Err

ors

S

L-S

Mammals (Rats) Birds (Pigeons) Reptiles (Turtles)

0.0

0.5

1.0

1.5

2.0

2.5

0 5 10 15 20 25

3-Trial BlocksM

ea

n L

og

La

ten

cy

S

L

L-S

Bony Fish (Goldfish)

Surprising nonreward

• Cues predict a larger or more preferred reward than that actually occurring.

• Surprising nonrewardpromotes two kinds of learning:

•Allocentric learning: about a change in the environment (cognitive).

•Egocentric learning: about the organism’s reaction to that change (emotional).

Papini, 2003, Brain Behav Evol

Osteichthyes

(Goldfish)

Amphibia

(Toads)

Aves

(Pigeons)

Reptilia

(Turtles)

Mammalia

(Rats)

Phylogenetic relationships among fish, amphibians, and reptiles

?

???? ?

Osteichthyes

(Goldfish)

Amphibia

(Toads)

Aves

(Pigeons)

Reptilia

(Turtles)

Mammalia

(Rats)

Phylogenetic relationships among fish, amphibians, and reptiles

Allocentric learning[Reversed SNC]

Allocentric

+

Egocentric learning[SNC]