1. Behavioral Biology Chapter 51

2. Behavior is what an animal does and how it does it. What is Behavior?

3. Ecology: Interaction between organisms and the biotic and abiotic environment Abiotic- non living; e.g., temperature, light, dissolved gas, water Biotic- living; e.g., predators, prey, mates Behavioral Ecology The study of the behavior of organisms within an evolutionary framework. e.g., communication, finding food, protection from predators

4. Mechanistic basis for behavior: E.g., MIGRATION A.Identify the stimuli that triggers the behavior or B.Study the psychological, neuronal, and hormonal changes that underlie the behavior Proximate causes External stimuli- changes in day length and temp Internal stimuli - hormone levels

5. Ultimate causes - address the evolutionary significance for a behavior and why natural selection favors this behavior. Why did a behavior evolve? Is it adaptive? Does it contribute to reproductive success? Example: birds that migrate have a selective advantage over birds that don't/didn't, selected for over time, could be due to long term climate changes, glaciation, disease, taking advantage of food sources, etc.

6. Nature versus Nurture: Revisiting an Old Debate Nature Behaviors are: Innate Hard-wired Instinctual Genetically determined Nurture Behaviors are: Learned Flexible Not genetically determined Behavior results from both genes and environmental factors

7. A.Behavior - What an animal does and how it does it. - some behavior is learned, some behavior is inherited B. To some extent ALL behavior has a Genetic Basis 1.some is totally genetic - which implies heritable 2.some is learned but relies ENTIRELY on genetically based mechanisms C. In general, behavior is a response to some environmental stimulus

8. Innate Behaviors – inherited, instinctive A. programmed by genes; B. highly stereotyped (similar each time in many individuals) C. Four Categories 1.Kinesis 2.Taxis 3.Reflex 4.Fixed Action Pattern

9. 1.Kinesis: "change the speed of random movement in response to environmental stimulus“ 2.Taxis: "a directed movement toward or away from a stimulus; positive and negative taxes 3.Reflex: "movement of a body part in response to stimulus". 4.Fixed Action Pattern (FAP): "stereotyped and often complex series of movements, responses to a specific stimulus", hardwired, however, not purely genetic, may improve with experience a.programmed response to a stimulus b.stimulus of FAP = "releaser", sometimes called "sign stimulus“ c.examples: - courtship behavior - rhythms - daily (circadian); annual (circannual)

10. D. Characteristics of Innate Behaviors - especially FAPs: 1.The behavior is performed correctly the 1 st time without prior experience (no opportunity to learn) 2.Triggered by some external stimulus 3.Once started, run to completion with no further input 4.Breeding crosses produce hybrid behaviors

12. Ethology is the study of how animals behave in their natural habitat. –Karl von Frisch: bee communication –Niko Tinbergen: herring gull experiment; digger wasps –Konrad Lorenz: imprint in geese Classical ethology presaged an evolutionary approach to behavioral biology

13. Karl Von Frisch- communication in bees

14. Herring gull experiment by Niko Tinbergen Releaser Stimuli- stimuli that release FAP E.g., Chick and red dot on gull parents beak triggers feeding response- parent regurgitates food Laysan albatross feeding chick

15. Niko Tinbergen Hypothesis: digger wasps use visual landmarks to keep track of her nests Move pine cones Visual cue is arrangement pattern rather than objects themselves

16. Egg rolling behavior in geese is a Fixed Action Pattern

17. Male three-spined stickleback shows aggression at models with red undersides Life-like model

18. Songbird repertoires provide us with examples. –Why has natural selection favored a multi-song behavior? Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings Fig. 51.5

19. It may be advantageous for males attracting females. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings Fig. 51.6

20. Cost-benefit analysis of foraging behavior. –Foraging is food-obtaining behavior. The optimal foraging theory states that natural selection will benefit animals that maximize their energy intake-to-expenditure ratio. Height of Drop (m) Average Number Of Drops Required to Break Shell Total Flight Height (Number of Drops  Height per Drop) 255110 31339 5630 7535 15460

21. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings Fig. 51.7

22. The behavior is adaptive - signs that natural selection is at work a. survival may depend on behavior, learning not an option (one chance only) b. animals with simple NS may not have capacity to learn - not strictly true, "simple" animals learn c. social interactions dependent on survival require rigid performance of roles mating rituals, termite mounds

23. Learning - Learned Behavior: Five Categories A. Imprinting 1. a strong association learned during a specific developmental period a. "sensitive period" or "critical period" b. imprinting of baby geese on mother - Lorenz baby geese imprint on mother within hours of hatching will imprint on any object during that period 2. learning a releaser for an innate FAP

24. Goose imprinting by Conrad Lorenz Geese imprint on the first moving (with sound) object that they see after birth There is a selection of a specific period of time (critical period) for social attachment and mate recognition in geese (to ensure geese imprint on the same species)

25. Imprinting in conservation biology: Need to minimize/eliminate human presence while raising California Condors

26. B. Habituation 1. decline in response to a harmless, repeated stimulus filter - prevents animal from wasting energy/attention on irrelevant stimuli adaptive

27. C. Conditioning - laboratory setting 1. classical conditioning animals make associations - Pavlov's dog associates bell with food, begins to salivate, can be extinguished and later followed by recovery (unconditioned stimulus - meat, unconditioned response - salivation, conditioned stimulus - bell, conditioned response - salivation) a.animal learns to perform an "old" response to a new stimulus b.Pavlov's dog - place dried meat powder in dog mouth - salivation - associate with bell - salivation to bell c.Stimulus first, behavior second (but of course there is an expectation of reward second)

29. 2. operant conditioning a. perform behavior to receive reward or avoid punishment b. Skinner Box - levers, reward - self training elaborate protocols c. behavior first, reward second (but of course there is a stimulus, if only a thought of reward)

30. Operant conditioning. –This is called trial-and-error learning - an animal learns to associate one of its own behaviors with a reward or a punishment.

31. D. Trial and Error Learning - nature setting 1. natural operant conditioning 2. modify responses to specific stimuli (releasers) - making both more adaptive 3. modify releaser to specific FAP - making both more adaptive Observational learning - social imitation

32. Octopus opening jar with crab Trial & Error & Observational Learning

33. Play as a behavior has no apparent external goal, but may facilitate social development or practice of certain behaviors and provide exercise. Practice and exercise may explain the ultimate bases of play

34. E. Insight, reasoning 1.manipulating concepts in the mind to arrive at adaptive behavior 2.mental trial-and-error 3.internal memory stores are used as additional sensory/information source All examples of tool-using: chickadees/tits and opening milk bottles Egyptian Vulture - uses rocks Cocos Finch - uses splinters of wood North American Gulls, Northwestern Crow - smash clams on sandy beaches

35. Cognition is the ability of an animal’s nervous system to perceive, store, process, and use information gathered by sensory receptors. The study of cognition connects nervous system function with behavior

36. Kinesis and taxis. –These are the simplest mechanisms of movement. Kinesis is a change in activity rate in response to a stimulus. –For example, sowbugs are more active in dry areas and less active in humid areas. Taxis is an automatic, oriented movement to or away from a stimulus. –For example, phototaxis, chemotaxis, and geotaxis. Animals use various cognitive mecha- nisms during movement through space Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings

37. Use of landmarks within a familiar area. –Some organisms move in response to a recognized object or environmental cue, the object is the landmark. Cognitive maps. –Some animals form cognitive maps (internal codes of spatial relationships of objects in the environment). Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings

38. Migration Behavior. –Migration is the regular movement of animals over relatively long distances. –Piloting: an animal moves from one familiar landmark to another until it reaches its destination. Whale Migration Routes

39. Orientation: animals can detect directions and travel in particular paths until reaching destination. –Navigation is the most complex, and involves determining one’s present location relative to other locations in addition to detecting compass directions. –Cues for these behaviors include the earth’s magnetic field, the sun, and the stars.

40. Besides humans, are animals aware of themselves? Some would argue that certain behaviors are a result of conscious processing. The study of consciousness poses a unique challenge for scientists

41. Social behavior is any kind of interaction between two or more animals, usually of the same species. Sociobiology places social behavior in an evolutionary context Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings

42. Social Behavior in Vertebrates A. Predator Avoidance Behavior -mimicry - schooling B. Reproductive Behavior -competition -territoriality - displays C. Parental Behavior D. Communication E. Cooperative Behavior -warning alarms

43. Sometimes cooperation occurs. Competitive social behaviors often represent contests for resources Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings Fig. 51.18

44. Agonistic behavior is a contest involving threats. –Submissive behavior. –Ritual: the use of symbolic activity. –Generally, no harm is done. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings Fig. 51.19

45. Reconciliation behavior often happens between conflicting individuals. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings Fig. 51.20

46. Dominance hierarchies involve a ranking of individuals in a group (a “pecking order”). –Alpha, beta rankings exist. The alpha organisms control the behavior of others. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings

47. Territoriality is behavior where an individual defends a particular area, called the territory. –Territories are typically used for feeding, mating, and rearing young and are fixed in location.

48. –Drawbacks are that territoriality uses a great deal of an individual’s energy. In addition, an individual might be defending a territory and die or miss a reproductive opportunity. –Spraying behavior is where an individual marks its territory. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings Fig. 51.22

49. Courtship behavior consists of patterns that lead to copulation and consists of a series of displays and movements by the male or female. Natural selection favors mating behavior that maximizes the quantity or quality of mating partners Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings

50. Fig. 51.23

51. Parental investment refers to the time and resources expended for raising of offspring. –It is generally lower in males –Females usually invest more time into parenting (fecundity, egg size, care of offspring) –Females are usually more discriminating in terms of the males with whom they choose to mate. Females look for more fit males (i.e., better genes), the ultimate cause of the choice. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings

52. Mating systems differ among species. –Promiscuous: no strong bond pairs between males and females. –Monogamous: one male mating with one female. –Polygamous: an individual of one sex mating with several of the other sex. Polygyny is a specific example of polygamy, where a single male mates with many females. Polyandry occurs in some species where one female mates with several females. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings

53. Certainty of paternity can influence mating systems and parental care. –If the male is unsure if offspring are his, parental investment is likely to be lower. –Exceptions do exist. Male sea spider cares for eggs

54. Mimicry monarch viceroy Bastian Mimicry- palatable species resembles an unpalatable one Mullerian- two or more unpalatable species resembles one another monarch queen

55. blenny (Aspidontus taeniatus) mimics Hawaiian cleaner wrasse shortnose wrasse mimics Potter's angel which sports a defensive spine Mimicry

56. Schooling Behavior

57. Defining animal signals and communication. –A signal is a behavior that causes a change in the behavior of another animal. –The transmission of, reception of, and response to signals make up communication. –Examples include the following: Displays such as singing, and howling. Information can be transmitted in other ways, such as chemical, tactile, electrical. Social interactions depend on diverse modes of communication Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings

58. –Pheromones are chemicals released by an individual that bring about mating and other behaviors. Examples include bees and ants. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings Fig. 51.26

59. Most social behaviors are selfish, so how do we account for behaviors that help others? –Altruism is defined as behavior that might decrease individual fitness, but increase the fitness of others. The concept of inclusive fitness can account for most altruistic behavior Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings Fig. 51.28

60. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings Fig. 51.29

61. –Inclusive fitness: How can a naked mole rat enhance its fitness by helping other members of the population? How is altruistic behavior maintained by evolution? If related individuals help each other, they are in affect helping keep their own genes in the population. Inclusive fitness is defined as the affect an individual has on proliferating its own genes by reproducing and helping relatives raise offspring. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings

62. –Hamilton’s Rule and kin selection. William Hamilton proposed a quantitative measure for predicting when natural selection would favor altruistic acts. Hamilton’s rule states that natural selection favors altruistic acts. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings

63. The three key variables are as follows: –B is the benefit to the recipient –C is the cost to the altruist –r is the coefficient of relatedness, which 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 Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings Fig. 51.30

64. –The rule is as follows: rB > C The more closely related two individuals are, the greater the value of altruism. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings

65. –Kin selection is the mechanism of inclusive fitness, where individuals help relatives raise young. –Reciprocal altruism, where an individual aids other unrelated individuals without any benefit, is rare, but sometimes seen in primates (often in humans).

66. Sociobiology connects evolutionary theory to human culture Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings Fig. 51.32