Course admin

- sequence of events

Female Male

(1) Appears & gives head up

display

zigzag swim (2)

(3) swims, head up to

male

swims to nest (4)

(5) female follows

shows nest (6)

(7) enters nest

shows tremble thrust (8)

(9) female spawn

- in this example each FAP acts as a releaser for next FAP

- but this is not always what happens with stickleback
Predicted Real

(1) (1)

(2) (2)

(3) (3)

(4) (4)

(5) (5)

(6) (6)

(7) (7)

(8) (8)

(9) (9)
This observation that the ‘fixed’ action pattern was not so ‘fixed’.

- lead to proposition of an

MAP - Model Action Pattern (Barlow 1968)

- spatiotemporal behaviour pattern that is common to members of a species

i.e. all members of a species perform pattern in a similar (or model) fashion

- but there is room for individual variation

A Few Final ideas that have been proposed for behaviour and its expression

(1) supernormal stimuli

(experimental)

- can exaggerate characteristics of a sign stimulus

- elicit behaviour more strongly

- e.g. oyster catcher

- if given choice of normal or very big egg (20x)

- will choose v. big egg

- will try to incubate even while falling off.

(2) Action specific energy

(Loney) - difficult to explain in neurological terms

- Loney observed that a stimulus could release a behaviour more easily if the behaviour had not been released for a while

- action specific energy

- constantly produced in animal’s CNS

- held in check until receive stimulus

But this idea of A.S.E. is consistent with 2 other kinds of behavioural activity

(1) Vacuum Activity

- courtship behaviour of male ring dove

- took isolated male

(2) Displacement Activity

- performance of irrelevant activity

e.g. male stickleback

- in own territory attacks

- in others territory flees

- at boundary - males dig nest

- acc. to Loney & Tinbergen

- as conflicting ASE builds

spills third activity

BEHAVIOUR GENETICS

- it is a truism to say that behaviour is under the control of genes.

- but the problem with studying behavioural genetic

- behaviour far removed from genes

e.g.

But one e.g. that genes are due involved in behaviour

- precisely measurable beh.

Behaviour Genetics

Most beh.

- under influence of gene complexes

- not just a 1 gene effect

- since it is under the influence of many genes

- behaviour expressed as a continuous

often difficult or impossible to detect influence of single gene

The kinds of traits I am talking about

- Quantitative

for a quantitative trait

- need different methods of analysis

- looking at variation (statistically - variance)

- experimentally - can get at genetic controls to variance by standardizing one or other of VE or VG

from this - get Heritability

Heritability - propositions of variability due to genetics.

Five general approaches to study of beh. genetics

(1) single gene mutant

(2) artificial selection experiments

(3) hybridization experiments

(4) parent/offspring regressions

(5) comparative - inbred lines from different species

(1) Single gene mutants

(-single gene but multiple effects!)

e.g. yellow bodied mutant

- males less successful than ++

- why - courtship song - less active

Classical experiment (in book?)

Hygenic behaviour of Honeybees

Nonhygenic strain - susceptible to bacterial disease

Hygenic strain - workers uncap cells

- remove brood

This is a dihybrid

u - for uncapping

r - removal

U - no uncap

R - no removal

Parents

but from back cross

get various combinations of

- no uncapping, no removal UR

- uncapping with no removal uR

(Experimenter - no uncapping removal Ur

uncapped) - uncapped removal ur

(2) Artificial selection

- experimenter chooses from ends of freq. distrib.

e.g. - disruptive selection

(2) Mating speed in Drosophila

- got 3 lines

- fast - 3 mins for 50%

- control - 5 mins

- slow - 80 mins
(C) Hybridization Experiments

- lovebirds - peach-faced - carry strips of material tucked in wing feathers

- Fischers

- carry strip in beak

Hybrids

- try to tuck material into

- fail

(1) doesn’t let go

(2) strips fall out

(3) wrong location

(4) tucking

- eventually were successful only in carrying

- took 3 yrs to learn
(A) HYBRIDIZATION (2)

More precise experiment

Sokolowski (late 80’s early 90’s)

found a gene in fruit flies

‘Rover” Sitter

- hypothesized - single gene covered a very complex behaviour - foraging

rovers - moved continually eating

sitters - moved little

(E) Parent offspring regression

- heritability

- scores parents & offspring for some beh. & compare

- generally get heritability < .6

- activity scores in Drosophila

(F) Comparative Approach

- Temperature selection in deer mice peromyscus

- subspecies live on a gradient

if take offspring

- lab-reared

- show similar preference

- also found genetic based (but weak M)

- in experiment with amount of nest material.
BUT

All the experiments cited so far point at a genetic effect

Some more recent experiments have pointed to the details of how these effects might work

In late 70’s

fruit fly mutant - dunce

role in olfactory learning

- especially good example because scant discrimination - v. important for fruit flies

- expose fruit flies to 2 odours

- shock them for one

- normal flies - learn to avoid odour with shock

- dunce flies don’t

Why?

Hypothesis (1) - can detect

- not true

dunce - problem with memory formation

- dunce flies can learn - but memory fades v. quickly

What does dunce gene do?

- codes for cyclic AMP phosphod

- breaks down c-AMP

- if look at cells of dunce flies

- reduced cAMP - phosphod

incr. cAMP

and if take normal flies & inhibit c-AMP-ase

- get poor memory

a link between cAMP & learning

Another mutant

- rutabaga - poor learning & memory

- level of cAMP - depends on 2 enzymes

(1) - cAMP-pdase - lowers cAMP

(2) - adenylylcyclase - raises - cAMP level by breaking down ATP

- rutabaga - defective adenyl cyclase

not activated by learning

concluded that rutabaga and dunce can’t learn for different reasons

flies need “s in cAMP to learn

rutabaga - no because cAMP not produced

dunce - no because cAMP not broken down

Final Step in This

- cAMP binds to another enzyme

protein Kmase A (PKA)

PKA - activates CREB gene which codes for cAMP response binding protein

activates other genes

growth of cell connectors

- similar pattern is seen in Aplysia, mice

HORMONES & BEHAVIOUR

Hormone - any chemical messenger

- produced in endocrine glands

- action on target tissue

- usually at some distance from endocrine glands

- action of hormone can be very specific

i.e. not all cells have receptors for all hormones

- those that do - target cells

What are Endocrine glands

- duct less - if have ducts - exocrine

- rich blood supply

- hormones - secreted into blood

- hormones - travel to every cell

- hormones - receptors - very specific

AMONG Invertebrates

- wide variety of endocrine systems

but just to give one e.g.

grasshopper (and most insects)

(1) neurosecretory cells - brain

(2) corpora cardiaca - brain

(3) corpora allata - beside esophagus

(4) prothoracic gland - neck

(5) gonads

one e.g. of how this works

tobacco horn worm - Manduca septa - green caterpillar

- all arthropods must molt to ground

- to moult

- series of rhythmic contractions - post ant

60-80 mins

- serves to loosen cuticle

Moulting - 2 separate behav’s

(1) peristatic waves - more pronounced

- cause arching

(2) withdrawal of legs

This entire process under hormonal control

e.g. Eclosion hormone (EH)

- triggers above behaviour through a series of interactions with other hormones
This works at a complex series of interactions


Among Vertebrates

can be divided into 2 categories

(1) hypothalamo-pituitary system

- ventral side of brain

- very close connection between CNS and endocrine system

pituitary & hypothalamus

- 2 connections

Posterior Pt - mainly nervous

connects from hypothalamus

Anterior Pit - linked via blood system

- in effect - neural signals received by hypothalamus

affect post. pituitary

neurosecretion by anterior pit

(2) Endocrine glands

What do they do?

Gland Hormone Effect

pineal melatonin - annual reprod. cycle

post pit. oxytocin - milk ejection, birth H2O balance

vasopressor

Ant.pot. luteinizing horm (LH) - corpora lutea formation

- progesterone secr.

- androgen secr.

Follicle stim Horm (FSH) - follicle dev. (female)

- ovulation (with LH & estrogen)

- spermatogenesis

Prolactin - milk secretion

- parental beh (birds)

Adren. Corticotrophic - regulates adrenal glands

hormone (ACTH)

intermediate Melanophore stim. Horm - colour

pituitary (MSH)

Adrenals steroids - water balance

(Cortex) - metabolism

- electrolyte balance

- blood sugar

- stress reaction

pancreas insulin - blood sugar

testes Androgens - testes dev

- spermatogenesis

- 2o sex char.

- sexual activity

ovary/placenta estrogens - uterine growth

- mammary gland dev.

- sexual activity

progestogens - gestation

How do we find out what a hormone does

(1) Surgical intervention

- surgical removal of endocrine gland

- should decrease beh.

- replace hormone

- should reinstate behav.

D. Crews (70s)

- Anolis lizards - v. territorial

- several displays involved

e.g. head bobbing with develop

- when 2 males meet

head bobbing submissive gesture from intruder

- intruder leaves

- courtship - v. similar

- Crews - removed testes & then replaced androgens

can conclude that androgens have eff.

but interesting wrinkle

- if put castrated male in home area

- sam level of aggressiveness

- if put in new cage

- got drop in activity

This shows that there is an influence of social factors

- e.g. resident status

(2) Correlational Studies

- less invasive

- correlate beh. & hormone level

Wingfield (1984)

- examined birds (song sparrow) under natural conditions

- caught males regularly - ident. with leg band

- took blood sample

- also observed male behav.

- explanation

- first peak - rel. to aggressive territory def. & courtship

- second peak - more problematic

- defending females - sexually receptive

- why no peak for second brood?

- speculated - male is feeding

- aggressive beh. would interfere

(3) Hormone substitution

- does artificial administration of hormone

e.g. in several spp. of rodents & birds

if inject androgens into

(1) intact males outside breeding season

(2) castrated males

(3) young males

(4) females

all will show male sexual behav.

There are a number of other techniques

Immunocytochemistry

Autoradiography stem from above

Genetic manipulations

Hormone & Behaviour

Behaviour

- much beh. is under the control of hormones

- either in long term or short term

but having said this

- their “short-term” effect is still much longer than nervous control

in general the effect is

It has been hypothesized that there are 2 ways hormones act

(1) modifier of behaviour

- alters responsiveness

e.g. female mosquitoes

- ecdysone

- on biting beh

-ecdysone - secreted by ovaries

- female need blood to dev. ovaries

- if remove ovaries

- biting continues

- if inject ovarectomized female with ecdysone

- biting stops

(2) Release of behaviour

- presence of hormone act as a sign stimulus

- releases series of motor patterns

in moths

- “eclosion hormone”

- responsible for behaviour of coming our from pupa

- produced in brain & corpora cardiaca

neurosecretory

- its secretion is genetically “timed”

- when it is secreted

“turns on” a complete sequence of behaviours - that results in movements that lead to eclosion

Among vertebrates

- somewhat more complex

- more endocrine glands are involved

Their system

- see from this that p neurosecretory

Remember - another wrinkle

- endocrine system - slower but hormones can reach many target tissues

Effects of hormones

(1) Influence on releasers

e.g. - 2o sexual char. - with commune

- by gonadotropins or sex hormones

- e.g. - red belly of male stickleback

- sexual swelling in female chimps

These changes can be

(A) once in a lifetime

- puberty in humans

(B) annual - mating colouration

(C) periodically - sexual swelling

- estrous cycle

(2) Influence on receptors

- perception of transmitted message

e.g. Prolactin - incr. sensitivity of brood patch in birds

- brooding

(3) Activation of neuron groups on CNS

e.g. eclosion beh. of moths

- activation of nerves from 3rd 6th segment

(4) Effects on brain structures

- hormones can have effects at sensitive periods in dev.

e.g. female canaries ingest testosterone

- dev. of HVc region to look like a male

Final e.g. is reverse (in a way)

(5) Hormone release controlled by beh.

e.g. 1 - in male red deer

infl. onset of estrous

Gp. A - female exposed to norectomy roaring male

Gp. B - female exposed to tape of roaring

Gp. C - female isolated

e.g. (2) ring doves

- fairly complex system (about a 6-7 week cycle)

- start by increasing the photoperiod
photoperiod
FIGURE

Finally an example with a simple behaviour that summarizes a lot of the proximal mech.

sea hare - Aplysia - egg laying
When about to lay eggs

stops moving & eating

increase heart & resp.
muscles on reprod. tract contract
eggs appear - egg string
FAP grabs string in upper lip
FAP waves head - stereotype
FAP pulls string & coats with mucous
FAP attaches to rock

FIGURE

NEW PART OF COURSE

Ontogeny of Behaviour

Beh. of animal

- from birth to sexual maturity

Kinds of changes

(1) complete repertoire develops over time

(2) beh. of young animals

- special adaptations - suckling

(3) Seasonal changes in particular pattern

- e.g. parental care

Study of ontogeny of behaviour

(1) description of developmental

(2) interaction of internal & external factors

Historically - all traits categorized as learned vs innate

Features of an innate beh. pattern

- genetically pre-determined motor pattern

- fully functional first time this performed

- usually elicited by a single use

- inherited

Early ethologists - most beh. if innate

e.g.’s -

- birds - aggressive resp. to coral snake

- young, inexperienced hand reared

- motmot - prey to coral snake

- coral snake red/yellow rings

- o-pecks

- any combination of ring absence, presence or colour

- pecking

Learned behaviour

- behaviour pattern dependent mostly on env. or

- environmentally determined

- psychologist - argue most beh. is learned

These are early view

- now look at it - both genome & env. important - and they interact

can separate into distinct categories same as tenta

Now - think of difference between learned and innate

- degree to with a beh. appears in its complete form the first time it appears

- additional problem

(1) innate beh. modified by experience

(2) learned - often genetically biased

Recognizing beh. patterns

3 classical criteria

(1) characterized by stereotyping

- great constancy

(2) i.e. appears to be same in all members of species

(3) cannot be influenced by environments called a FIXED ACTION PATTERN

(FAP) - of temporally & spatially arranged sequences of muscle contraction which (Loney) produce biologically appropriated stereotyped movement patterns

- released by external stimuli

- but presence of stimulus not necessary for completion

sign stimulus IRM FAP

IRM - innate releasing mechanism

- internal filter/triggering complex

- produces FAP when perceives correct stimuli

e.g. egg retrieval in greylag goose

Later - this concept as a model action pattern (MAP)

- stereotyping is weak, learned component - high degree of constancy.

from this - there is now a view that most beh. patterns will change over time

Can change to

(1) Experience

- learning - e.g. each response in grill chicks

Hailman

(2) Maturation

- ontogenetic changes due to non-learning

- changes in CNS or hormonal systems

- full development or perfection of a beh. pattern - with no practice

behaviour improves without opportunity to perform

e.g. puppies - peeing

- male and female - same squat

- androgens male cocks leg

so what we appear to have here

Final points (before discussion of learning)

Why have innate responses?

- why not just have simple learning patterns

(1) if relationship between stimuli & adaptive response is PREDICTABLE

- innate beh. is adv.

(2) If result of making an error is dangerous or fatal

- e.g. reaction to predator (coral snake e.g.)

Advantages in

(1) antipredator beh.

(2) prey beh. (food recog.)

(3) courtship patterns - spp. differs

(4) response of young to parents

(5) beh. in short-lived spp. (Social insects - without overlapping genes


LEARNING

(A) DEFINITION - all processes that lead to adaptive champs in individual behav. as a result of experience under a particular set of environmental conditions.

- as a process requires

i) acquisition of information with sense organs

ii) storage of information as a memory trace

iii) recall of information when needed

- can’t measure directly

- measure how beh. has changed as a result of experience

(B) Sensitive phases in Learning

(formerly called critical period)

- through an animal’s life - specific ages (genetically detected)

- time when it is maximally responsive to stimuli and has max. learning ability

- often v. early in life - first few hrs. or days.

- physiological basis

- poorly known - obviously some developmental process in CNS but after that...

e.g. in many birds

- embryo can acquire info. in egg

- chickens - if play sounds into incubator

- will prefer familiar ones

- quillemots - burrow nesting seabird

- dense colonies

- acoustic comm - between embryo or parent

- learns (in egg) to recog. call of parent

- useful in dense colony.

(C) Constraints on Learning

- learning capacity of a species

dep. on two criteria

(1) phylogenetic development

- increase in learning capacity as - go up phylogenetic tree

- corresponds to cephalization

- not always

(2) environmental conditions in which lives

- diff in learning abilities - rel. to ecol. conditions
e.g. gulls vs Kittiwakes

ground nesting cliff nesting

young wander young stay put
parents learn quick parents never learn to

to recognize own recognize young

- just nest location
Can conclude

(A) strong selection pressure to develop specialized learning abilities

(B) genetically pre-determined range of learning abilities

(4) LEARNING PROCESSES

learning (ie. how animals learn has been classified into several categories (very arbitrary)

(1) habituation

- stimulus specific fatigue

- decrease in intensity of a response over time

- which repeated stimulus

neither positive or negative consequences

- - in effect, negative learning process

- can be adaptive

- animal ceases to respond to irrelevant stimuli

(2) Classical (Pavlovian) Conditioning

- previously neutral stimulus

releases a behavioural response as a result of positive or negative reinforcement

- neutral stimuli becomes conditioned stimulus

- reaction elicited conditioned responses

Classical experiment - Pavlov

- salivary reflex in dogs

UCS - unconditioned stimulus - meat

CS - condition stem - light

CR - condition resp.

can’t be maintained

- response decreases over time

- extinction

-must periodically

- give UCS to strengthen CR

these kinds of responses are rare in

- don’t get ‘pure’ pairing of CS & UCS

- one e.g. of it though

- prey avoidance in birds

- monarch

- birds quickly distasteful with colour pattern

UCS CS

(3) Instrumental Learning or Operant Conditioning

- imagine a hungry animal

- active, searching for food

- performs a variety of behaviours

- now suppose one is followed by reinforcement

(e.g. food)

- repeat this association of number of times

- animal associates action X with food

- carried our in mazes and skinner boxes

in nature

- animals forage in areas where got food before

(4) Invitational (“observational” learning)

- copy behaviour of conspecific

- e.g. facial expressions

- acquisition of new ability.

(5) Insight Learning (Cognitive Learning)

characterized by

(1) animal comprehends new problem set = spontaneously

(2) consider necessary spatial and temporal behav. sequence in advance

(3) performs sequence correctly first time to solve problems

- demonstration of it difficult