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Chapter 51 Behavioral Biology What is Behavior? • • • • • Movement Noise Learning Secretions Behavior: what an animal does and how it does it Proximate and Ultimate Causes • Proximate questions are mechanistic, concerned with the environmental stimuli, if any, that trigger a behavior, as well as the genetic and physiological mechanisms underlying a behavioral act • Ultimate questions address the evolutionary significance of a behavior Proximate and Ultimate Causes • Ex: Magnolia warbler breeds in spring and early summer – Proximate: breeding is triggered by the effect of increased day length on an animal’s photoreceptors – Ultimate: why did natural selection favor this behavior and not a different one? Nature Versus Nurture • It is not an “either-or” debate • It includes both when it comes to behavior • There have been genes linked to certain behavior – Dg2 gene in fruit flies • Gene influences the level of a protein that functions in signaling within cells – One allele causes low levels of the protein, and the fly is lazy – One allele causes high levels of the protein, and the fly buzzes around like crazy – Genetic claims for human behaviors is still being investigated Innate Behavior • Developmentally fixed • Behaviors that are genetically programmed without any environmental influence – All individuals exhibit virtually the same behavior despite environmental differences within and outside their bodies during development and throughout life • Ex: blind newborn birds open their mouths and chirp to get parents to feed them Ethology • Originated in the 1930’s with naturalists that tried to understand how a variety of animals behave in their natural habitat – Was the “mother” of modern behavioral biology • von Frisch, Lorenz, Tinbergen won Nobel Prize in 1973 – Focused on proximate mechanisms, but with an eye toward the genetic links to behavior and the adaptive nature of behavior Ethology • Fixed Action Pattern (FAP): a sequence of behavioral acts that is essentially unchangeable and usually carried to completion once initiated – Triggered by an external sensory stimulus known as a sign stimulus • Usually some feature of another species – Some moths fold their wings and drop instantly in response to the ultrasonic signals sent out by predatory bats – Male Three-Spined Stickleback Fish: attacks other males that invades its territory just at the sight of the red belly of the intruder » Can fish using red lures » Red triggers either aggressive or sexual behavior in many species that have color vision – Mayflies swarm and mate on the water surface and detect the water by the light reflected from it » Roads can produce the same reflection – FAP’s triggered by simple cues prevent an animal from wasting time processing or integrating a wide variety of inputs Behavioral Ecology • Research field that views behavior as an evolutionary adaptation to the natural ecological conditions of animals • Natural selection will favor behavioral patterns that enhance survival and reproductive success Behavioral Ecology • Songbird Repertoires – Many songbirds have a repertoire of songs that are different across species – Increases fitness because the older a male gets, the more songs he knows and females prefer males with more tunes to sing • So they don’t get too bored Behavioral Ecology • Cost-Benefit Analysis of Foraging Behavior – Natural selection refines behaviors that enhance the efficiency of feeding – Food obtaining behaviors include recognizing, searching for, capturing, and eating food • Foraging encompasses all of these behaviors – Optimal foraging theory views foraging behavior as a compromise between feeding costs and feeding benefits – Crow Experiment • Opportunistic feeders that avail themselves a variety of foods • In British Columbia (where they did the experiment) crows pick up crustaceans off the beach with their beak, fly upward and drop them on a rock to break their shells and will continue until the shell breaks – The higher they fly, the less they have to retry » Energy Cost • Predicted the crow would fly to a height that provided the most food relative to the amount of total energy required to break the shell – Prediction was proven true – Bluegill Sunfish Experiment • Feed on Daphnia, generally selecting larger individuals that provide the most energy • Smaller prey will be selected if larger prey is too far away • Predicts that the proportion of small to large prey eaten will also vary with the overall density of prey – Foraging behavior tends to minimize the risk of the predator becoming prey while foraging Learning • Modification of behavior resulting from specific experiences • Most innate behaviors improve with performance as animals learn to carry them out more efficiently • Ex: Alarm calls of vervet monkeys – Give distinct alarms when they see leopards (bark), eagles (doublecough), or snakes (chutter) – Behave in the appropriate way depending on the call – Infant’s calls are indiscriminate, give an “eagle call” when they see any bird – As adults, they only give a call when they see one of two species of eagles that are proven vervet eaters – Social confirmation speeds up the process – Have the innate behavior to make a call upon seeing danger, but learn to fine-tune it with age Learning versus Maturation • Maturation: ongoing developmental changes in neuromuscular systems – We think of birds “learning to fly” but some have been raised in an environment where they could not flap their wings, and then when they reached an age where flying is normal, they picked it up immediately Habituation • Simple type of learning that involves a loss of responsiveness to stimuli that convey little or not information – Hydra retracts upon touch, but stops responding to repeated stimuli – “Little Boy Who Cried Wolf” • May increase fitness by allowing an animals’ nervous system to focus on stimuli that signal food, mates, or real danger instead of wasting time or energy on other irrelevant stimuli Imprinting • Learning that is limited to a specific time period in an animals life and that is generally irreversible • Mother-offspring bonding: if bonding fails to happen, the parent will not initiate care of the infant • Lorenz’s Geese Experiment – Half a clutch with mother and half with Lorenz • Lorenz’s clutch showed no recognition to mother or others of the same species – Some initiated courtship with humans as adults – Imprinting stimulus • Found that anything that moved, and especially made sound was “Mother” – A ticking clock was once used to imprint on Imprinting • Sensitive Period: a limited phase in an animal’s development when learning of particular behaviors can take place – Geese imprinting is the first 2 days of life – Young birds must imprint to know parents, and parents must imprint to know young – Two species of finches were reared separately and initially imprinted with their own species (first sensitive period) • When they were becoming sexually mature, they were put with members of the opposite species and sexually imprinted on them (second period) • Were placed back with their own species and were reluctant to mate, but when placed with other species they mated, even if it was up to 8 years since they had seen the other species Bird Song • First 50 days they just listen to others’ songs and form a template • Sings tentative notes called a subsong – Hears its own singing and compares it to others and modifies it as needed • It then crystallizes its adult song Human Sensitive Periods • Foreign languages are learned most easily up until the teen years Learn to Associate • Associative Learning: ability of many animals to learn to associate one stimulus with another – Pavlov • Classical conditioning: involves learning to associate an arbitrary stimulus with a reward or punishment • Operant Conditioning: trial-and-error learning – Learns to associate one of its own behaviors with a reward or punishment and then tends to repeat or avoid that behavior – Skinner • Mouse and lever Playing • No apparent internal goal but involves movements closely related to goal-directed behaviors – Stalk-and-attack playing • The “Practice-Hypothesis” suggests that play is a type of learning that allows animals to perfect behaviors needed in functional circumstances • The “Exercise Hypothesis” suggests that play is adaptive because it keeps the muscular and cardiovascular systems in top condition Animal Cognition • Cognition: ability of an animal’s nervous system to perceive, store, process, and use information gathered by sensory receptors • Cognitive ethology: examines the connection between an animals’ nervous system and its behavior – Animal consciousness and awareness – What calculations does a dog make to determine where a Frisbee will land? Kinesis and Taxis • Kinesis: simple change in activity or turning rate in response to a stimulus – Roly Polies become more active in dry areas and less active in humid areas, a behavior that keeps these in moist environments • Don’t move toward or away, buy slow down in favorable environments • Taxis: automatic, oriented movement toward or away from some stimulus Use of Landmarks • Go towards a stimulus that is an arbitrary landmark the animal must learn • Honeybees: – Keep track of the nectar supplies from their local flowers and concentrate their foraging on the flowers that are most nectar-rich • Use landmarks along the way • Know where they are going and make a “bee-line” because they learned certain landmarks as young Migration Behavior • Migration: regular movement over relatively long distances • Generally make one round trip between two regions each year • Notable examples: birds, whales, butterflies, and oceangoing fish – Golden Plover: from arctic breeding grounds to South America, or even Hawaii • Use one of three mechanisms: 1. Piloting: animal moves from one familiar landmark to another until it reaches its destination • Short-distance migrations 2. Orientation: detects compass directions and travels in a particular straight line path for a certain distance until it reaches its destination 3. Navigation: animal determines its present location relative to other locations in addition to detecting compass direction • Cues: – Earth’s magnetic field – Sun – Star • Some fix on the North Star or use an internal clock Social Behavior • Social Behavior: any kind of interaction between two or more animals, usually of the same species – Most sexually reproducing species must be social for part of their life cycle in order to reproduce • Sociobiology: applies evolutionary theory to the study and interpretation of social behavior Competitive Social Behaviors • Sometimes involves cooperation, as when a group carries out behavior more efficiently than is possible for a single individual • Although they may be cooperating, each individual acts in a way that will maximize its own benefits, even if this is at a cost to the other participant Agonistic Behavior • A contest involving both threatening and submissive behavior determines which competitor gains access to some resource • Sometimes involves a test of strength, more often the engage in threat displays that make them look large and fierce – Severity depends on the amount, type, and availability of the resource • Male ground squirrels kill others for females because they are only in heat for a few hours each year • Usually only occurs with individuals of the same species, and can be completely symbolic in nature • Reconciliation Behavior: a kind of friendly relation between the conflicting individuals following the conflict itself Dominance Hierarchies • Chickens that are unfamiliar with each other will sit and peck at each other until the clear dominant one is found – Pecking Order Territoriality • Territory: an area that an individual defends, usually excluding other members of its own species • Typically used for feeding, mating, and rearing young • Established and defended by agnostic behavior • Range in size from very large (hundreds of square kilometers) to very small Courtship • Behavior patterns that lead up to copulation or gamete release • Consists of a series of displays and movements, either by the male, female, or both • Enables animals to identify potential mates of the same species Parental Investment • The time and resources an individual must spend to produce and nurture offspring • The mate who has the highest parental investment is generally the more choosy of the two when it comes to picking a mate Mating Systems • Promiscuous: no strong pair-bonds or lasting relationships • Monogamous: one male mating with one female • Polygamous: an individual of one sex mating with several of the other sex – Most often one male and several females • Polygyny – Sometimes it is one female and several males • Polyandry • Parental investment has a lot to do with mating systems – Birds who need a lot of attention require both parents foraging for food for them • Monogamy – Animals who are relatively independent from birth can make it on their own • Polygamy Defining Signals and Communication • Signal: behavior that causes a change in behavior in another animal • Transmission of, reception of, and response to signals makes up communication – Visual, auditory, chemical, tactile, and electrical signals Pheromones • Odors emit chemical signals for communication – Trail of Ants • Bees http://youtu.be/-7ijI-g4jHg – Maintain the social order of honeybee colonies – Convey the location of good food sources – Dances • Tells what and where new food resources are, and how far away Nature is Selfish • Most animals act in a way that benefits them, and they do not care if it harms others • Altruism: unselfish behavior – Reduce their individual fitness and increase the fitness of the recipient of the behavior – Ex: Belding Ground Squirrel’s cry – Ex: bee societies where workers are sterile and sting intruders – Ex: Mole Rat colonies have one reproducing female who mates with three males Inclusive Fitness • Describes the total effect an individual has on proliferating its genes by producing its own offspring and by providing aid that enables other close relatives to increase the production of their offspring – Prairie Dogs Hamilton’s Rule • Benefit to the recipient (B) • Cost to the altruist (C) • Coefficient of relatedness (r) – Equals the probability that a particular gene present in one individual will also be inherited from a common parent or ancestor in a second individual • B is the average number of EXTRA offspring that the beneficiary of an altruistic act produces • C is the average number of FEWER offspring the altruist produces • For natural selection to favor an altruistic act, the benefit to the recipient multiplied by the coefficient of relatedness must exceed the cost to the altruist • Kin selection: natural selection that favors altruistic behavior by enhancing reproductive success of relatives