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BIONB 4240: NEUROETHOLOGY

L02: ROOTS in ETHOLOGYAug. 26, 2011; C. D. Hopkins

Apis melifera, Frisch’s favorite study species.

Tinbergen uses models to evoke egg rolling behavior. He discovers that convex objects work better than cubic objects.

Make stickleback fish (above) respond to models of female that have enlarged abdomens.

Herring gull chicks peck at the red spot on the parent’s bill, or a model, but prefer a red stick with candy-stripe patterns at the sharp tip: a super-normal stimulus.

Lorenz’s psychohydraulic model of motivation explains the changing threshold of a stimulus. Control theory model to the right of the same.

Writing Assignment

W3: Due Monday Aug. 291) How does a reflex differ from a fixed action

pattern? 2) Define “Instinct”2) Define Instinct3) Define IRM (Innate Releasing Mechanism)

First Discussion: Wed. Aug 31

Kaas and Catania (2002) How do features of sensory representations develop? BioEssays 24 (4) 334-343.

Writing (W4)

Carl Hopkins will lead the discussion

Outline

1. The ethological tradition2. Fixed action patterns3. The ethogram4. Innate release mechanisms and sign stimuli 5. Courtship behavior of the common fruit fly 6. Influential (neuro)ethologists

Ethology and the Ethological Tradition

The study of natural behavior of animals, usually conducted in the natural setting. - observe & describe behavior: “ethograms”- Tinbergen’s: goals of Ethology are to understand causation,

ontogeny, function, and evolution of behavior.

Neuroethologists:Observe behavior and make a descriptions.Develop categories when appropriateUse objective descriptors, avoid anthropomorphismsDetermine its adaptive function.Develop hypotheses about mechanism of neural control.

FIXED ACTION PATTERNS

EGG ROLLING in Grey-lag Goose (Lorenz & Tinbergen, 1938)Taxis und Instinkthandlung in der Eirollbewegung der Graugans. Z. Tierpsychol. 2, 1-29.

Waving display in Fiddler crab.(Crane)

TRITONIA: escape swimming(Willows)

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Fixed Action Pattern

Lorenz, K., Tinbergen, N. (1938) Taxis und Instinkthandlung in der Eirollbewegung der Graugans. Zeitschrift für Tierpsychologie 2, 1-29.tr: Taxis and instinctive behaviour pattern in egg-rolling by the Greylag goose. In: Studies in Animal and Human Behaviour, vol 1, Robt Martin transl. Harvard univ. Press (1970).

Greylag goose (Anser anser)

Key Features of FAP

• A ‘complex’ behavior; a sequence or a pattern, generated by the animal (an endogenous pattern)

• Triggered or “released” by relatively complex stimuli (eggs, other round objects)

• Reaction specific exhaustibility: with repetition even• Reaction-specific exhaustibility: with repetition, even though muscles were apparently not fatigued (implies CNS)

• Once released, constant in form, regardless of environmental feedback (goes to completion even when interrupted)

• FAPs might occur after a long resting period without releaser (vacuum activity)

A. fixed action pattern is not a reflexReflexes:

Activated by simple and generalized stimulus (pain, hot, cold, sharp, stretch)

No adaptation or fatigue.Response is graded in intensity according

to stimulus intensity (recruitment of neurons).

No response w/o stimulus.

Flexion reflex is di or tri synaptic; ipsi and contralateral projections

Modified by changes in sensory stimulus.

monosynaptic stretch reflex

Tritonia diomedia

video Electrophysiol video

Frost, W.N.

Escape swimming in Tritonia, a marine mollusc

Willows, 1971 Sci. Am.

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Fiddler Crab (Salmon, Atsaides ;1968)

Claw wave is a fixed action pattern used in display by males for attractive females.

StereotypyThe wave is highly stereotyped in time. Repeated performances by the same individual.

Claw waving in fiddler crab – a method of measurement

Salmon, Hyatt, McCarthy, Costlow (1978)

Variation in performance

Uca baturmys male waving

Uca deichmanni male waving

Uca beebei male waving

The wave is also species-specific and is used for mate attraction in areas where two or more species coexist.

Species specificity of fiddler crab waves

Data from Salmon & Atsaides, Amer. Zool. 8:623.

Single wave

Pattern of waves

Fixed Action Patterns

• These FAPs are complex, coordinated, stereotyped.• A sequence of movements integrated into a single

action.• Evoked or released by a complex stimulus, but not

guided nor controlled by that stimulus The behavior isguided nor controlled by that stimulus. The behavior is endogenous, and it is released.

• Goes to completion once started.

Ethogram

• Catalog of all fixed action patterns• Description is objective, not anthropomorphic

– rather than “afraid” or “angry” use description of posture and coordination of limbs.

– includes limb and posture events, including temporalincludes limb and posture events, including temporal organization

– avoids prejudging function (rather than intimidation posture, use upright posture, throat puff, or tail slap)

• Catalog should incorporate variation (clusters)• Behavioral repertoire may also include reflexes, highly

variable acts, taxes.• Early ethology: long descriptive list of behaviors

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Common MarmosetBEHAVIOR CODE DEFINITIONAgonismTufts-flick TF rapid back-and-forth movement of ear tuftsFrown FR lower eyebrows, furl brow, and turn down corners of mouth while staringCuff CU swift, superficial blow or scratch performed aggressivelyChase CH pursue partner, with one or both animals exhibiting aggression and/or submission (not play)Fight FI grapple aggressively with partner(s), involving biting, clawing, and wrestlingAttack AT lunge at or pounce on partner aggressively; may or may not result in fightSnap bite SB direct a single short, sharp bite at partnerSubmit SU flatten ear tufts and/or facial grimace (partially open mouth with corners of mouth retracted, exposing lower and sometimes upper teeth) and/or slit eyes (eyelids half closed)Continuous submit CS continuous submit; start scoring after 5 secRetreat RE starting from a stationary position, move at lest one body length away from another animal within 1 sec of the other animal establishing proximity (within 10 cm)

PlayPlay PL two or more animals lunge, grapple, wrestle or chase for at lest 1 sec in absence of aggression or intense submission; play face may or may not be presentSolicit play SP direct play face toward, pounce on, or initiate grapple with partner, in absence of ongoing play with partnerPlay face PF open mouth without retraction of the lipsJoin play JP join ongoing play bout between two or more partnersEnd play EP discontinue all social play for _> 3 secSocial play SO social interactions involving non-aggressive physical contact with other individuals; high activity

Infant-associated behaviorsClimb on ON climb onto any part of partner's body so that all four limbs are on partnerSolicit climb on SC position body directly above infant and/or pull infant onto body; may or may not result in infant climbing onto partner's bodyClimb off OF voluntary climb off partner's body after having all four limbs on partnerPush off/reject PO prevent juvenile from climbing onto body, or rub or otherwise force juvenile off bodyNurse NU have mouth on female's nipple for _> 1 secEnd nursing EN discontinue nursing posture

Other social behaviorsSniff/nuzzle SN orient face against or toward partner, excluding anogenital regionAnogenital inspect AI orient face against or toward anogenital region of partner, or use hands or mouth to investigate anogenital region of partner; includes anogenital groomGroom GR use hands and/or mouth to pick through fur and/or mouth of partner, excluding anogenital regionSexual solicit SS stare at partner with ear tufts flattened and eyes slitMount MO climb on partner's back from behind and grip partner around waist and legs; may be accompanied by pelvic thrustingInitiate huddle IH establish passive, torso-torso body contact with partner, with both animals remaining stationary and in passive contact for at least 3 secLeave huddle LH terminate huddle after at least 3 sec of passive, torso-torso body contact during which both partners remained stationaryObject steal OS take any non-food object from hands or mouth of partnerAttempt object steal AO attempt but fail to take non-food object from hands or mouth of partner

Food-associated behaviorsFood steal ST take any food from hands or mouth of partnerFood steal ST take any food from hands or mouth of partnerAttempt food steal AF attempt but fail to take food from hands or mouth of partnerShare food SH eat from a food source from which partner is simultaneously eating or occupying without removing any food from partner's mouth or handsNew food NF eat from a food source which no other animal is currently holding, eating from, or occupying

Individual behaviorsBristle strut BS arching posture and/or strut locomotion and/or general PiloerectionScentmark SM rub or drag anogenital, suprapubic, or sternal region along substrate, object, or partnerGenital present GP raise tail to expose genitalsObject manipulation OM sniff, bite, chew, gouge, handle, pounce on, grapple with, or otherwise manipulate inanimate object, excluding food items and water bottle, for at least 1 sec

Anolis carolinensis

Cichlids: Baerends and Baerends-Van Roon (1950)

Innate Release Mechanism

What stimuli triggers the F.A.P. ?- experimental approach- use models, dummies- focus on components that evoke natural responses.

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Using Models Super-Normal Stimulus

normal

lacks red

super normal

Questions about Innate Recognition Cues

TINBERGEN, N. 1953. The Herring Gull's world. Collins, London. xvi q-255 pp.

TINBERGEN, N . 1958. Curious naturalists. New York. 280 pp.

Tinbergen, N., and A. C. PERDECK. 1950. On the stimulus situation releasing thebegging response in the newly hatchedbegging response in the newly hatched Herring Gull chick (Larus argentatusargentatus Pont.). Behav., 3: 1-39.

Hailman, J. P. 1967. The ontogeny of an instinct: the pecking response in chicks of the laughing gull (Larus atricilla L.) and related species. Behaviour Supplement XV. E.J. Brill, Leiden. 196 pp.

Hawk Goose Recognition

Vacuum Acts

In after a long period of absence of an appropriate releaser the animal may perform a fixed action pattern without a stimulus. Vacuum activity.

Motivation

ACTION-SPECIFICENERGY

FIXED ACTIONPATTERNS RELEASER

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Drive theory and Haplochromis burtoniLater, Heiligenberg showed that the cichlid, H. burtoni did not have an increase in aggressiveness in the absence of an aggressive opponent.

Attack Behavior in CichlidsStudied with an isolated male, exposed to a model male for 1/2 minute. Attack rate is measured on smaller fish in the same tank. Attacks increase markedly after exposure to the model.

Heiligeberg et al., 1972

Attack Rate after Stimulus

The visual appearance of the model can be manipulated by changing the color pattern of the model. The dark eye bar is a strong stimulus for increasing aggression.

The orange spot reduces the attack rate. Attack Rate after Stimulus

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The chirp rate of a cricket is similarly elevated for 10’s of seconds after an external chirp. Note the build up of chirp rate after 18 stimulus chirps separated by 10 s.

Displacement Activity

Inappropriate to stimulus.

Given when appropriate response is prevented or when two stimuli are in conflictprevented, or when two stimuli are in conflict.

Courtship in Fliesvideo

R. J. Greenspan and R. Ferveur (2000): Courtship in Drosophila. Ann. Rev. Genetics34:205 Males respond to the odor,

and to contact chemicals (taste). Males use sound to court the female.

Sex difference in odors of cuticular hydrocarbons produced by Drosophila.

Olfactory receptors are distributed on the antenna, and on the maxillary and labial palps

sex differences in chemosensory areas on the forelegs

Chemoreceptors are GPCR proteins

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The chemoreceptive circuitry of the male and female also differs. Light green neurons show cells the come from the antennae and the palps. Axons project into the Antennal lobe and from there into the Lateral Protocerebrum and mushroom bodies.

Male Song

pulse song of the male

0.5 s

Hearing in Drosophila

Female aristaJohnston’s Organ

Song Evolution

Ethology’s Contribution to Neuroethology

1) Ethologists focused attention on natural behavior, usually in the natural environment.

2) Ethologists recognized the importance of endogenous motor programs that produced recognizable, rather stereotyped behaviors. These were called “fixed action patterns”.

3) Ethologists proceeded to describe repertoires of behavior: the ethogram. Many ethograms were produced.

4) FAPs appeared to be “released” by specific stimuli, or models of key features of stimuli (= sign stimuli). The study of sign stimuli led to understanding of the sensory processing.

5) Motivation or drive varies. Lorenz’s psychohydraulic model, although useful is not supported by quantitative data.

6) Drosophila courtship is a model system of behavioral neurobiology that can be studied using genetic, neurobiological and behavioral tools.

The EndThe End