CHAPTER 12

EARLY SOCIAL LEARNING AND

BEHAVIOR

Learning Phenomena

Nonassociative Associative/Cognitive

•Classical conditioning

•Instrumental conditioning

•Rule learning

•Social learning

•Habituation

•Short term

•Long term?

•Sensitization

•Short term

•Long term?

General Specific

•Human language

•Song learning

•Imprinting

Tentative classification of learning phenomena

Imprinting in precocial birds

Konrad Lorenz

Imprinting

Filial: following response,

preference for imprinted object.

Sexual: mating preference.

Chemical: site preference,

usually for egg laying.

A form of recognition learning characterized because

it is available only during an early sensitive period.

Types of imprinting

Filial imprinting

Sensitive period and reversibilityIs imprinting learning a general or a restricted capacity?

Factors influencing imprinting

Stimulus factorsWhich stimuli are more effective? What elements of a stimulus do birds respond to?

Learning factorsDoes conditioning play a role in imprinting?

Brain mechanismsIs there a specific brain area concerned with imprinting?

Motivational factorsAre chicks motivated to respond for an imprinted object the same way they respond for

food or other rewards?

Naturalistic stimuli

• Artificial models (A, B) can be effective,

especially if movement is involved.

• However, natural models, such as a

stuffed animal (C), are especially

effective.

• But what is the element that controls

imprinting in a natural model?

• Experiments in which a

stuffed animal is used as a

model show what are the

elements in the

configuration that are

effective.

• Amazingly, any of these

scrambled versions of the

stuffed animal work about

the same.

• The key element in the

configuration is the

presence of the neck and

head.

• Chick also respond well to a

model of a natural predator

if it contains the head and

neck of the animal.

Head and neck

Sensitive period

• (A) A circular apparatus that moves a model at a constant speed. A duckling placed in the apparatus tracks the

models exhibiting the “following response.” Different models can be used or ducklings of different ages.

• (B) Using ducklings of different ages the shape of the sensitive period can be assessed.

• Using this procedure, the peak of sensitivity is around 13-16 h after hatching.

• Younger and older ducklings exhibit a weaker following response.

(A) (B)

Model

duck Duckling

Po

sit

ive R

esp

on

se (

%)

Age (hours)

Learning Phenomena

Nonassociative Associative/Cognitive

•Classical conditioning

•Instrumental conditioning

•Rule learning

•Social learning

•Habituation

•Short term

•Long term?

•Sensitization

•Short term

•Long term?

General Specific

•Human language

•Song learning

•Imprinting

Tentative classification of learning phenomena

Attachment

Individual recognitionMother and offspring respond selectively to each other

PreferenceMother and offspring are motivated to minimize the

distance from each other

Separation anxietyPlacing mother and offspring in different locations leads to

signs of distress (e.g., vocalizations, stress hormones)

Attachment

Mother-infant attachment in monkeys

https://www.youtube.com/watch?v=OrNBEhzjg8I

Duration: 6:07 min

Video

Pioneering research by Harry Harlow (1950s-70s)

0

5

10

15

20

0 30 60 90 120 150 180

Age (Days)

Ho

urs

in

Pro

xim

ity to

Clo

th M

oth

er

Cloth mother/Cloth fed

Wire mother/Cloth fed

Cloth mother/Wire fed

Cloth mother/Wire fed

• Rhesus monkeys prefer to be

close to the cloth mother,

even when the wire mother

provides nourishment.

• The cloth provides a kind of

contact comfort.Wire Cloth

Early social isolation in monkeys

0 100 200 300 400

Basal

Separation-

Reunion

Infant Removal

Mother Removal

Plasma Cortisol (mg/100 ml)

Mothers

Infants

Separation anxiety

• True attachment leads to a distress response when the mother and offspring are separated physically.

• This study shows the glucocorticoid response (plasma levels of the stress hormone cortisol).

• Stress-hormone levels in the mother are elevated when the infant is removed from the habitat.

• But infants respond even more intensely to separation from the mother, whether because the infant is

removed or the mother is removed.

• Basal hormonal levels are restored after a reunion.

Importance of play: therapeutic monkeys

• Several procedures were tested to determine whether the effects of early isolation could be reversed.

• Moving models and dogs were used, but their effects were only partial. Dogs initiated a substantial number of

interactions, but isolated monkeys still grew up with significant deficiencies in social behavior.

•

Learning Phenomena

Nonassociative Associative/Cognitive

•Classical conditioning

•Instrumental conditioning

•Rule learning

•Social learning

•Habituation

•Short term

•Long term?

•Sensitization

•Short term

•Long term?

General Specific

•Human language

•Song learning

•Imprinting

Tentative classification of learning phenomena

Vocal learning in birds and humans

Common features:

*Learning from adults

*Sensitive periods for early learning

*Same production system

*Dialects

Hatching(summer)

Nesting period

Juvenile period(fall and winter)

(spring)

First breeding season

Adult lifeSensorimotor learning phase

•Crystallized or adult song

•Plastic song

•Subsong

•No song production

•Acquisition of an auditory memory

•Flexible sensitive period

•Initial social interactions

•Exposure to song

Sensory learning phase

Age-dependent plasticity

Swamp Sparrow Song Sparrow

Normal song

Isolate song: Hearing intact

Isolate song: Deafened before singing

Age-dependent plasticity: effects of early song deprivation

0

5

10

0

5

10

0

5

10

Fre

qu

en

cy (

kH

z)

0 0.5 1.0 1.5 2.0

Time (s)

(a)

(b)

(c)

Dialects of the rufous-collared sparrow (Zonotrichia capensis)

Theme Trill

0.5 s

5 k

Hz

Marin

Berkeley

Sunset Beach

San Francisco

Bay Area

Dialects of the white-crowned sparrow (Zonotrichia leucophrys)

Dialects of the ortolan bunting (Emberiza hortulana)

The ortolan bunting breeds in Europe and Asia from Lake Baikal to Spain.

In Asia, it occurs mainly between 48° and 55° north, whereas in Europe it

extends beyond the Arctic Circle. In Germany its centre of distribution is in

the northeast, although there are some small breeding areas in northern

Bavaria, north Rhine-Westphalia and lower Saxony.

The ortolan bunting has many song dialects. In a dialect region there is

only little variation in song. Conrads made up a dialect atlas by comparing

different populations of ortolan bunting in central Europe. In this area 12

dialect classes were found . The songs have a total length of one to two

seconds and consist of two to three (seldom four) different parts. The

beginning (part A) is a row of two to six syllables. Each syllable shows

more or less strong changes in frequency and duration of the sound

elements. Three-part songs have a part M, which normally is of constant

frequency. In combination with M the ending (E) is determinant in

classifying the regional allocation. It is often a tremolo-like and soft sound.

Dialects of the ortolan bunting (Emberiza hortulana)

Song(input)

Vocal behavior(output)

Basilar membrane

Cochlear nucleus

DLM

NOPosterior

telencephalon:

song production

Anterior

telencephalon:

song acquisition

HVC RA

Nucleus of the

Hypoglossal nerve

Syringeal muscles

LMAN

N. XII

N. VIII

Area X

Brain mechanisms of song learning

Thalamus

See full names for brain

areas in book (p. 590).

Size of brain nuclei

involved in song

learning

Quality of song repertoire

(a proxy for cognitive

abilities related to

foraging efficiency)

Female sexual

preferenceEarly

nutrition

Early

stress

Feeding

efficiency during

the plastic song

phase

Endocannabinoid brain levels

Sexual selection contributing

to the evolution of a large

telencephalon and higher

cognition in songbirds

Integrating proximate and ultimate causes