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BIONB 4240: NEUROETHOLOGY L02: ROOTS in ETHOLOGY Writing Assignment Aug. 26, 2011; C. D. Hopkins W3: Due Monday Aug. 29

Apis melifera, Frisch’s 1) How does a reflex differ from a fixed action favorite study species. pattern? 2) Define “Instinct”

Tinbergen uses models to evoke 3) Define IRM (Innate Releasing Mechanism) egg rolling behavior. He discovers Herring gull chicks peck at the red spot on the parent’s bill, or a model, but prefer that convex objects work better a red stick with candy-stripe patterns at than cubic objects. the sharp tip: a super-normal stimulus.

Make stickleback fish (above) respond to models of female that have enlarged Lorenz’s psychohydraulic model of motivation explains the changing abdomens. threshold of a stimulus. Control theory model to the right of the same.

First Discussion: Wed. Aug 31 Outline

Kaas and Catania (2002) How do features of sensory 1. The ethological tradition representations develop? BioEssays 24 (4) 334-343. 2. Fixed action patterns 3. The ethogram Writing (W4) 4. Innate release mechanisms and sign stimuli 5. Courtship behavior of the common fruit fly Carl Hopkins will lead the discussion 6. Influential (neuro)ethologists

Ethology and the Ethological Tradition FIXED ACTION PATTERNS

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.

EGG ROLLING in Grey-lag Goose (Lorenz & Tinbergen, 1938) Neuroethologists: Waving display Taxis und Instinkthandlung in der in Fiddler crab. Observe behavior and make a descriptions. Eirollbewegung der Graugans. Z. (Crane) Develop categories when appropriate Tierpsychol. 2, 1-29. Use objective descriptors, avoid anthropomorphisms Determine its adaptive function. Develop hypotheses about mechanism of neural control. TRITONIA: escape swimming (Willows)

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Fixed Action Pattern 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 Greylag goose (Anser anser) 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 Lorenz, K., Tinbergen, N. (1938) Taxis und Instinkthandlung in der Eirollbewegung der Graugans. Zeitschrift für Tierpsychologie 2, 1-29. releaser (vacuum activity) 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).

A. fixed action pattern is not a reflex Tritonia diomedia Reflexes: Flexion reflex is di or tri Activated by simple and generalized synaptic; ipsi and stimulus (pain, hot, cold, sharp, stretch) contralateral No adaptation or fatigue. projections Response is graded in intensity according to stimulus intensity (recruitment of neurons). No response w/o stimulus. Modified by changes in sensory stimulus.

monosynaptic stretch reflex

Escape swimming in Tritonia, a marine mollusc

video Electrophysiol video

Frost, W.N. Willows, 1971 Sci. Am.

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Stereotypy The wave is highly stereotyped in time. Fiddler Crab (Salmon, Repeated performances by the same Atsaides ;1968) individual.

Claw wave is a fixed action pattern used in display by males for Claw waving in fiddler attractive females. crab – a method of measurement

Variation in performance

Salmon, Hyatt, McCarthy, Costlow (1978)

Species specificity of fiddler crab waves Uca baturmys male waving

Data from Salmon Uca deichmanni male waving & Atsaides, Amer. Zool. 8:623. Uca beebei male waving

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

Pattern of waves

Fixed Action Patterns Ethogram

• These FAPs are complex, coordinated, stereotyped. • Catalog of all fixed action patterns • A sequence of movements integrated into a single • Description is objective, not anthropomorphic action. – rather than “afraid” or “angry” use description of posture and • Evoked or released by a complex stimulus, but not coordination of limbs. guided nor controlled by that stimulus. The behavior is – includes limb and posture events, including temporal organization endogenous, and it is released. – avoids prejudging function (rather than intimidation posture, use • Goes to completion once started. 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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BEHAVIOR CODE DEFINITION Agonism Tufts-flick TF rapid back-and-forth movement of ear tufts Common Marmoset Frown FR lower eyebrows, furl brow, and turn down corners of mouth while staring Cuff CU swift, superficial blow or scratch performed aggressively Chase 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 wrestling Attack AT lunge at or pounce on partner aggressively; may or may not result in fight Snap bite SB direct a single short, sharp bite at partner Submit 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 sec Retreat 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)

Play Play 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 present Solicit play SP direct play face toward, pounce on, or initiate grapple with partner, in absence of ongoing play with partner Play face PF open mouth without retraction of the lips Join play JP join ongoing play bout between two or more partners End play EP discontinue all social play for _> 3 sec Social play SO social interactions involving non-aggressive physical contact with other individuals; high activity

Infant-associated behaviors Climb on ON climb onto any part of partner's body so that all four limbs are on partner Solicit 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 body Climb off OF voluntary climb off partner's body after having all four limbs on partner Push off/reject PO prevent juvenile from climbing onto body, or rub or otherwise force juvenile off body Nurse NU have mouth on female's nipple for _> 1 sec End nursing EN discontinue nursing posture

Other social behaviors Sniff/nuzzle SN orient face against or toward partner, excluding anogenital region Anolis carolinensis Anogenital inspect AI orient face against or toward anogenital region of partner, or use hands or mouth to investigate anogenital region of partner; includes anogenital groom Groom GR use hands and/or mouth to pick through fur and/or mouth of partner, excluding anogenital region Sexual solicit SS stare at partner with ear tufts flattened and eyes slit Mount MO climb on partner's back from behind and grip partner around waist and legs; may be accompanied by pelvic thrusting Cichlids: Baerends and Baerends-Van Roon (1950) Initiate huddle IH establish passive, torso-torso body contact with partner, with both animals remaining stationary and in passive contact for at least 3 sec Leave huddle LH terminate huddle after at least 3 sec of passive, torso-torso body contact during which both partners remained stationary Object steal OS take any non-food object from hands or mouth of partner Attempt object steal AO attempt but fail to take non-food object from hands or mouth of partner

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

Individual behaviors Bristle strut BS arching posture and/or strut locomotion and/or general Piloerection Scentmark SM rub or drag anogenital, suprapubic, or sternal region along substrate, object, or partner Genital present GP raise tail to expose genitals Object 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

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 Hawk Goose Recognition

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 the begging response in the newly hatched Herring Gull chick (Larus argentatus argentatus 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.

Vacuum Acts Motivation

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

FIXED ACTION RELEASER PATTERNS

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Attack Behavior in Cichlids Studied with an isolated male, exposed to a model male for 1/2 minute. Attack Drive theory and Haplochromis burtoni rate is measured on smaller fish in the same tank. Attacks increase markedly after exposure to the model. Later, Heiligenberg showed that the cichlid, H. burtoni did not have an increase in aggressiveness in the absence of an aggressive opponent.

Heiligeberg et al., 1972

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

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 conflict.

R. J. Greenspan and R. Ferveur (2000): Courtship in Drosophila. Ann. Rev. Genetics 34:205 Males respond to the odor, and to contact chemicals Courtship in Flies (taste). Males use sound to video court the female.

Sex difference in odors of cuticular Olfactory receptors are distributed on the antenna, hydrocarbons produced by Drosophila. 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 Male Song Protocerebrum and mushroom bodies.

pulse song of the male

0.5 s

Hearing in Drosophila Song Evolution

Female arista Johnston’s Organ

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 The End 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.

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