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Ethology is the scientific study of animal behavior, and a sub-topic of

zoology.

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Proximate vs. Ultimate Causes

• Proximate causes-what mechanism enables the organism to exhibit the behavior?

• Ultimate causes-in terms of evolution, how and why did that behavior come to be?

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For example, if a zebra is drinking at a water hole, and all of a

sudden it hears another zebra nearby make an alarm call, it may

stop drinking immediately and start running away instead.

The “proximate cause” is the immediate trigger for a behavior.

• The proximate cause of the zebra running away would be the alarm call. • But the ultimate cause, or real (evolved) reason why the zebra is running is

survival. It is running away because it wants to survive. The alarm call is not the source of danger, but the alarm call alerts the zebra that danger, such as a lion, may be nearby and the lion can threaten the zebra's chance to survive.

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Another example: The mouthless jellyfish-Mastigias paupua migrates upward in the water column during the day, and descends at night.

• Proximate Explanation- – when the sun rises, the jellyfish switch from infrequent, undirected pulsing of their bells to a

clear taxis, directing themselves toward the rising sun and pulsing continuously • This directed movement brings them to the surface, where they drift into the sunlight.

– Although they lack eyes, jellyfish are sensitive to light and are able to orient themselves toward it • This is because electrically coupled, photosensitive neurons control their swimming

behavior and can direct the top of the animal toward the sun.

• Ultimate explanation-• Although they are animals, Mastigias sp. do not eat.

– They are dependent upon photosynthetic algae called zooxanthelle which are imbedded in their tissues.

• Over millions of years, jellyfish that behaved in such a way as to maximize the photosynthesis of their symbionts left more offspring than others, and ultimately evolved coordinated muscle contractions to ensure that their symbionts had enough light.

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Innate vs. Learned Behavior• Innate behavior is developmentally fixed. Despite

differing environments, all individuals that can exhibit the behavior do exhibit the behavior. – This does not mean the environment does not play a role,

innate behaviors are triggered by a stimulus which must occur within the context of the animals environment

• Learned behaviors are modified by experience. – There is no hard-and-fast distinction between the two, many

learned behaviors have a strongly pre-determined chronology (I.e., language learning) and many innate behaviors are improved by experience (I.e., parasitoid foraging).

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TAXIS vs KINESIS• A taxis is an innate behavioral response by an

organism to a directional stimulus or gradient of stimulus intensity.

• A taxis differs from a tropism (“turning response”, often growth towards or away from a stimulus) in that the organism has motility and demonstrates guided movement towards or away from the stimulus source.

• It is sometimes distinguished from a kinesis, a non-directional change in activity in response to a stimulus.– Example: For example, flagellate protozoans of the genus

Euglena move towards a light source. Here the directional stimulus is light, and the orientation movement is towards the light. This reaction or behavior is a positive one to light and specifically termed "positive phototaxis”.

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TAXIS vs KINESIS

• Kinesis, like a taxis, is a movement or activity of a cell or an organism in response to a stimulus.

• However, unlike taxis, the response to the stimulus provided (such as gas exposure, light intensity or ambient temperature) is non-directional.

• • Example: Pillbugs are decreasingly active as humidity

increases.

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Fixed Action Patterns

• A FAP is a sequence of innate behavioral acts that is essentially unchangeable and usually conducted to completion once it is started. – It is triggered by a sensory environmental stimulus

called a sign stimulus (or releaser). – Because the animal responds to the sign stimulus,

rather than the environment as a whole, the animal may be tricked by signals that are out of their usual context.

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Fixed Action PatternsExample of a FAP:• Many mating dances, commonly carried out

by birds, are examples of fixed action patterns. In these cases, the sign stimulus is typically the presence of the female.

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Fixed Action Patterns• Another example of fixed

action patterns is aggression towards other males during mating season in the red-bellied stickleback.

• A series of experiments carried out by Niko Tinbergen showed that the aggressive behavior of the males is a FAP triggered by anything red, the sign stimulus.

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Fixed Action Patterns• Another well known case is the classic

experiments by Tinbergen and Konrad Lorenz on the Graylag Goose.

• Like similar waterfowl, it will roll a displaced egg near its nest back to the others with its beak.

• The sight of the displaced egg triggers this mechanism.

• If the egg is taken away, the animal continues with the behavior, pulling its head back as if an imaginary egg is still being maneuvered by the underside of its beak.

• However, it will also attempt to move other egg shaped objects, such as a golf ball, door knob, or even an egg too large to have possibly been laid by the goose itself.

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All behaviors are influenced by both genes and the environment

• The extent to which genes control the manifestation of a behavior differs from one behavior to the next. – Example: Alcoholism can actually be broken into two

behaviors; teen-onset alcoholism and adult-onset alcoholism.

– Cross-fostering studies using children placed in foster homes indicate that teen onset alcoholism has a very strong genetic basis.

– The same studies indicate that adult-onset alcoholism has a stronger environmental basis.

– All genes, including those whose expression underlies an innate behavior, require an environment to be expressed.

– If the same study had been conducted in Saudi Arabia, there may have been no alcoholics at all (strong cultural reasons not to drink, and alcohol less readily available).

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Example of Genetic Variation for a Behavior-Roving vs. Sitting

• Some behavioral differences are known to be caused by variation at a single gene. – In the fruit fly, Drosophila melanogaster, larvae

exhibit two different behaviors. • “Rovers” crawl around constantly, moving several

centimeters after feeding in a particular location. • “Sitters” basically stay put. • The two different behaviors are found in natural

populations, as well as in laboratory strains. • By isolating rovers and sitters, it was possible to breed

pure strains for the two behaviors.

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Learning

• Learning is pervasive in the behavior of birds and mammals, but other animals are capable of learning as well. – Example; learning in the sea anemone Stomphia sp.

Chemostimulation, paired with mechanical pressure, will cause a sea anemone to detach and swim off. With repeated trials, pressure alone will cause the anemone to detach itself. YouTube video

Learning can be defined as a more or less permanent change in the behavior or potential behavior of an animal based upon experience.

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Simple non-associative learning• Habituation is the relatively persistent waning of a response

to a stimulus when the expected result does not occur. – Sensory Adaptation is sometimes called habituation, it is the

phenomenon where environmental noise is filtered out by the nervous system over time and the animal no longer responds to it.

• Sensitization is the growing sensitivity of an animal to repeated stimuli. In essence, it is the opposite of sensory adaptation-it occurs when the stimulus is rewarded or is of adaptive significance. – A search image is an image an animal becomes especially adept at

recognizing, as a result of repeatedly searching for it and being rewarded for finding it.

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Associative Learning

• These are processes by which an animal comes to associate events in its environment with consequences or objects with stimuli.

• Associative learning is the process by which an association between two stimuli or a behavior and a stimulus is learned. The two forms of associative learning are classical and operant conditioning.

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Classical (Pavlovian) conditioning involves strengthening of the stimulus-outcome association.

• Classical conditioning (also Pavlovian conditioning or respondent conditioning) is a form of learning in which one stimulus, the conditioned stimulus or CS, comes to signal the occurrence of a second stimulus, the unconditioned stimulus or US.

• The US is usually a biologically significant stimulus such as food or pain that elicits a response from the start; this is called the unconditioned response or UR.

• The CS usually produces no particular response at first, but after conditioning it elicits the conditioned response or CR.

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Ex: R. Menzel- associative learning in honeybees

• Bees were trained to enter restraining tubes. – Sugar water was offered as a CS – the orienting response is an extension of

the proboscis by the bee. • A variety of smells and colors were offered as

US. • Bees could learn to associate a variety of

smells and colors with sugar water-i.e., they would eventually extend their proboscis in the absence of the sugar water

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Operant conditioning is the use of consequences (reinforcement/punishment ) to modify the occurrence

and form of behavior.

• In Operant conditioning, the behavior emitted by the organism is strengthened or weakened by its consequences (e.g. reward or punishment).– 1. Reinforcing a voluntary behavior. Positive

reinforcement: A child receives a 'gold star' at school for behaving well.

– 2. Negative reinforcement: A child does his or her homework to stop her parents from nagging. Punishment: Grounding a child for behaving inappropriately

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EX: Operant conditioning• EX: Skinner Box. An operant conditioning chamber permits

experimenters to study behavior conditioning (training) by teaching a subject animal to perform certain actions (like pressing a lever) in response to specific stimuli, like a light or sound signal.

• When the subject correctly performs the behavior, the chamber mechanism delivers food or another reward.

• In some cases, the mechanism delivers a punishment for incorrect or missing responses.

• With this apparatus, experimenters perform studies in conditioning and training through reward/punishment mechanisms.

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Skinner Box (B.F. Skinner)

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QUIZ: http://www.ar.cc.mn.us/biederman/courses/p1110/conditioning2.htm

•Classical and Operant Conditioning Examples:

• 1. Every time someone flushes a toilet in the apartment building, the shower becomes very hot and causes the person to jump back. Over time, the person begins to jump back automatically after hearing the flush, before the water temperature changes.

• Answer to Example 1• 2. Your father gives you a credit card at the end of your first year in college because you did so well. As a result, your grades continue to get better in

your second year.• Answer to Example 2• 3. Your car has a red, flashing light that blinks annoyingly if you start the car without buckling the seat belt. You become less likely to start the car

without buckling the seat belt.• Answer to Example 3• 4. You eat a new food and then get sick because of the flu. However, you develop a dislike for the food and feel nauseated whenever you smell it.• Answer to Example 4• 5. An individual receives frequent injections of drugs, which are administered in a small examination room at a clinic. The drug itself causes increased

heart rate but after several trips to the clinic, simply being in a small room causes an increased heart rate.• Answer to Example 5• 6. A lion in a circus learns to stand up on a chair and jump through a hoop to receive a food treat. • Answer to Example 6• 7. A professor has a policy of exempting students from the final exam if they maintain perfect attendance during the quarter. His students’

attendance increases dramatically.• Answer to Example 7• 8. You check the coin return slot on a pay telephone and find a quarter. You find yourself checking other telephones over the next few days.• Answer to Example 8• 9. Your hands are cold so you put your gloves on. In the future, you are more likely to put gloves on when it’s cold.• Answer to Example 9• 10. John Watson conducted an experiment with a boy named Albert in which he paired a white rat with a loud, startling noise. Albert now becomes

startled at the sight of the white rat.• Answer to Example 10

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Imprinting

• Imprinting occurs when an animal learns to make a particular response to a particular object. – Process is very rapid. – Can be altered, but the usual

pattern is an instantaneous and irreversible alteration of behavior.

• Konrad Lorenz

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Cognition

• The extent to which animals ”think” is unknown, although it is becoming increasingly obvious that the human mind differs from those of other animals largely by the extent of our cognitive ability, rather than the simple rule humans think, animals don’t. – Some hallmarks of cognitive ability

• ability to classify • ability to recognize attributes of objects • ability to count • ability to remember the locations of objects in space • ability to use language

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Some animals for which cognitive ability has been studied

• Parrots-classification, recognition of attributes, and language

• Bees, Chimps, Gorillas, Various Monkeys, Hyenas-Language

• Ants, Bees, Ravens, Various Other Birds-Spatial Mapping.

• Ex: honeybees “waggle dance” NatGeo video

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PLAY

• Most mammals, some birds, and possibly some other animals (including elephant-nosed fish) exhibit play behavior. – Play is an inexact term that covers a range of behaviors

exhibited by young animals. – 1) social play-such as wrestling and chasing other animals – 2) exercise play-such as running, hanging from tree limbs,

etc.. – 3) object oriented play-involves manipulating a novel object – The most obvious thing these behaviors have in common is

that they have no immediate adaptive value.

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Functions of Play

• It is frequently hypothesized that the function of play behaviors is to practice adult activities in a context where the consequences of failure are not serious or fatal. – Ex: kittens playing with toys,

wrestling with each other• Establish their relative positions

in the dominance hierarchy

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Other Kinds of Behavior• kinesis (moves more/less in response to stimulus) and taxis (moves to/away

from stimulus) • cognitive maps- animal uses cognition to take shortest trip, to plan shortest

route, or to remember location of objects. • use of landmarks- animal orients behavior patterns to objects in the

environment • migration- seasonal movement of individuals or whole populations • dormancy- seasonal or environmentally induced slowdown of metabolism • antipredator behavior/ predatory behavior • courtship • social behaviors

– agonistic behaviors-aggressive interactions – cooperative foraging – communication

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References, credits:

• University of Illinois, Chicago http://www.uic.edu/classes/bios/bios101/zanimalbehavior/tsld001.htm

• http://www.biology.eku.edu/RITCHISO/behavecolsyl.htm