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Chapter 35 Behavioral Adaptations to the Environment http://kevinspear.com/category/computers/ Animal Behavior Vocabulary Ethology = study of animal behavior Behavior = action performed in response to stimulus What Is Behavior? • Behavior – Is what an animal does and how it does it – Includes muscular and non-muscular activity Dorsal fin Anal fin Figure 51.2 Stimulus = something that elicits a response External Stimulus = Something outside the animal e.g. sound, sight, smell, etc, presence of another animal Internal Stimulus = something inside the animal e.g. hunger, fatigue, feeling cold, hormones Proximate and Ultimate Questions • Proximate, or “how,” questions about behavior – Focus on the environmental stimuli that trigger a behavior as well as internal mechanisms – Focus on the genetic, physiological, and anatomical mechanisms underlying a behavioral act • Ultimate, or “why,” questions about behavior – Addresses the evolutionary significance of a behavior This explanation of an animal's behavior is based on evolution. It requires that behavioral traits, like physical ones, are genetically heritable, and then explains behavior using an explanation of why this specific behavioral trait was favored by evolutionary mechanisms such as natural selection. Proximate and Ultimate Questions In philosophy a proximate cause is an event which is closest to, or immediately responsible for causing, some observed result. This exists in contrast to a higher-level ultimate cause (or distal cause) which is usually thought of as the "real" reason something occurred. Example: Why did the ship sink? Proximate cause: Because it was holed beneath the waterline, water entered the hull and the ship became denser than the water which supported it, so it couldn't stay afloat. Ultimate cause: Because the ship hit a rock which tore open the hole in the ship's hull. In most situations, an ultimate cause may itself be a proximate cause for a further ultimate cause. Hence we can continue the above example as follows: Example: Why did the ship hit the rock? Proximate cause: Because the ship failed to change course to avoid it. Ultimate cause: Because the ship was under autopilot and the autopilot's data was inaccurate. Separating proximate from ultimate causations frequently leads to better understandings of the events and systems concerned. Behaviors are categorized as follows: 1. Instinct is behavior that is innate, or inherited. In mammals, care for offspring by female parents is innate. “in most mammals, care for offspring by female parents is innate” 2. Fixed action patterns (FAP) are innate behaviors that follow a regular, unvarying pattern. 3. Imprinting is an innate program for acquiring a specific behavior only if an appropriate stimulus is experienced during a critical period (a limited time interval during the life of the animal). Once acquired, the behavior is irreversible. Fixed Action Patterns • A fixed action pattern (FAP) – Is a sequence of unlearned, innate behaviors that is unchangeable – Once initiated, is usually carried to completion A FAP is triggered by an external sensory stimulus Known as a sign stimulus • In male stickleback fish, the stimulus for attack behavior – Is the red underside of an intruder (a) A male three-spined stickleback fish shows its red underside. Figure 51.3a • When presented with unrealistic models – As long as some red is present, the attack behavior occurs (b) The realistic model at the top, without a red underside, produces no aggressive response in a male three-spined stickleback fish. The other models, with red undersides, produce strong responses. Figure 51.3b • Proximate and ultimate causes for the FAP attack behavior in male stickleback fish BEHAVIOR: A male stickleback fish attacks other male sticklebacks that invade its nesting territory. PROXIMATE CAUSE: The red belly of the intruding male acts as a sign stimulus that releases aggression in a male stickleback. Figure 51.4 ULTIMATE CAUSE: By chasing away other male sticklebacks, a male decreases the chance that eggs laid in his nesting territory will be fertilized by another male. Imprinting • Imprinting is a type of behavior – That includes both learning and innate components and is generally irreversible http://www.familycourtchronicles.com/philosophy/attachment/ • There are proximate and ultimate causes for this type of behavior BEHAVIOR: Young geese follow and imprint on their mother. PROXIMATE CAUSE: During an early, critical developmental stage, the young geese observe their mother moving away from them and calling. ULTIMATE CAUSE: On average, geese that follow and imprint on their mother receive more care and learn necessary skills, and thus have a greater chance of surviving than those that do not follow their mother. Figure 51.5 4. Associative learning (association) occurs when an animal recognizes (learns) that two or more events are connected. A form of associative learning called classical conditioning occurs when an animal performs a behavior in response to a substitute stimulus rather than the normal stimulus. 5. Trial-and-error learning (or operant conditioning) is another form of associative learning. 6. Habituation is a learned behavior that allows the animal to disregard meaningless stimuli. 7. Observational learning occurs when animals copy the behavior of another animal without having experienced any prior positive reinforcement with the behavior. About half a century ago, a juvenile Japanese macaque developed sweet-potato washing on the island of Koshima. The habit spread to the rest of the population. None of these monkeys is still alive today, but their descendants are still washing potatoes. 8. Insight occurs when an animal, exposed to a new situation and without any prior relevant experience, performs a behavior that generates a desirable outcome. Wolfgang Köhler The use of tools to gain access to food – Problem Space Types of Behavior 1. Foraging = feeding food e.g. locate, obtain & consume 2. Parental Care = ensuring survival of young e.g. carrying, nursing, cuddling, holding young 3. Courtship = attracting a mate e.g. courtship displays, bright feathers, songs, other vocalizations, head butting, fighting 4. Reproductive = e.g. mating, giving birth 5. Offensive/Defensive = aggression, submissive behavior, defense from aggressors e.g. hiding, fighting, escape, threatening 6. Territorial = protect a resource for exclusive use e.g. scenting / marking, birdsong, protection of mate / offspring, space, food or water source 7. Social = work to create alliances, help the group e.g. grooming, babysitting, defense of musk ox young, play, play fight 8. Migratory = movement to a more suitable environment as seasons change e.g. dry vs. rainy seasons, winter vs. summer range for browsers & birds e.g. salmon upriver to spawn, artic tern – 1800 m from Arctic to Antarctic, 2x/year 9. Communication = signaling between one animal & another •Greeting e.g. sniff, hug, kiss “bite” •Aggression e. g. charge, bite, hit, fight, etc. •Non-aggression e.g. patting, head butting, stroking •Grooming (bonding, alliances,keeping clean) •Group hunt ( e.g. hyenas, lions) Vocalizations e.g. bark, growl, snort, howl, hoot, chirp, whinny, alarm sound, other language Non-verbal signaling e.g. body, head, ear, & / or tail position showing teeth, smiling, looking away, looking directly at, gesturing, thumping, beating chest, raising hackles or hood, drumming, tail slap, snort, scenting, pheromones, sign language 10. Curiosity = investigating new stimulus in environment e.g. approach, sniff, chew, bite, mount 11. Elimination = defecation, urination 12. Resting = apparent inaction e.g. lying down, sitting, basking, sleeping, loafing 13. Play Purpose = training for life (defense, hunting, etc) Cognition = Knowing, including awareness and judgement. It is difficult to determine if animals are aware of themselves and their surroundings. The method used, behaviorism, does not test for cognitive functions. Behaviorism = A mechanistic approach which describes behavior in terms of stimulus and response. Cognitive ethnologists think cognitive ability arises through natural selection and forms a phylogenetic continuum stretching into evolutionary history. Cognitive ethology = A view that sees conscious thinking as an inherent part of animal behavior. • Behavioral traits can evolve by natural selection • Because of the influence of genes on behavior – Natural selection can result in the evolution of behavioral traits in populations http://misslorisbiohome.com/we/evolution/thenatsel.html • Natural selection favors behaviors that increase survival and reproductive success • The genetic components of behavior – Evolve through natural selection • Behavior can affect fitness – Through its influence on foraging and mate choice Biston betularia f. typica, the white-bodied peppered moth http://en.labs.wikimedia.org/wiki/Animal_Behavior/Evolution In 1955, H.B. D. Kettlewell published his study on pepper moths: Selection Experiments on Industrial Melanism in the Lepidoptera. Kettlewell hypothesized that the dotted whitish form of the peppered moth were more likely to be eaten than the melanic form because they could be easily detected against the soot covered trees. Howlett and Majerus further examined this hypothesis in their study: The Understanding of Industrial Melanism in the Peppered Moth, published in 1987. They tested it by pinning 50 of both forms of the peppered moths on pale and dark tree trunks. Calculations: Moths in Polluted (dark) Woodland Dotted Whitish Form: (30/50) x 100 = 60% Melanic Form: (20/50) x 100 = 40% 60% - 40% = 20% The Dotted Whitish form is predated on 20% more than the melanic form in the polluted woodland. Moths in Non-polluted (pale) Woodland Dotted Whitish Form: (15/50) x 100 = 30% Melanic Form: approx. (30/50) x 100 = 60% 60% - 30% = 30% The Melanic form is predated on 30% more than the dotted whitish form in the nonpolluted woodland. Foraging Behavior • Optimal foraging theory – Views foraging behavior as a compromise between the benefits of nutrition and the costs of obtaining food The Foraging Paths of the Male Grain Beetle and a Web User http://www.websiteoptimization.com/speed/tweak/information-foraging/theory.html Energy Costs and Benefits • Reto Zach – Conducted a cost-benefit analysis of feeding behavior in crows • The crows eat molluscs called whelks – But must drop them from the air to crack the shells Zach's first experiment, he selected a collection of whelks and sorted them according to size (small, medium, and large). He then dropped these whelks from various heights until they broke, and recorded how many drops at each height were required to break each whelk. A graph of his results is shown below. http://www-rohan.sdsu.edu/~jmahaffy/courses/f00/math122/lectures/optimization/opt.html • Zach determined that the optimal flight height in foraging behavior – Correlated with a fewer number of drops, indicating a trade-off between energy gained (food) and energy expended 125 60 40 Average number of drops 30 Total flight height 20 Drop height preferred by crows 10 0 2 Figure 51.22 3 Height of drop (m) 5 7 Total flight height (number of drops drop height) Average number of drops 50 15 100 75 50 25 • In bluegill sunfish – Prey selection behavior is related to prey density Small prey at middle distance Small prey at close distance Large prey at far distance Low prey density Small prey Medium prey Large prey High prey density 14% 33% 33% 33% 35% 50% Percentage available Small prey Medium prey Large prey 33% 33% 33% 100% Predicted percentage in diet Small prey Medium prey Large prey Figure 51.23 32.5% 32.5% 35% 2% 40% 57% Observed percentage in diet Social Learning • Social learning – Forms the roots of culture • Culture can be defined as a system of information transfer through observation or teaching – That influences the behavior of individuals in a population http://folksonomy.co/?permalink=1612 • No other species – Comes close to matching the social learning and cultural transmission that occurs among humans Figure 51.38 Evolution and Human Culture • Human culture – Is related to evolutionary theory in the distinct discipline of sociobiology • Human behavior, like that of other species – Is the result of interactions between genes and environment • However, our social and cultural institutions – May provide the only feature in which there is no continuum between humans and other animals The End