Chapter 51 Animal Behavior

1. Chapter 51Animal Behavior

2. You Must Know: How behaviors are the result of natural selection How innate and learned behaviors increase survival and reproductive fitness How organisms use communication to increase fitness The role of altruism and inclusive fitness in kin selection

3. Learning Targets

4. Introduction • Ethology: study of animal behavior • Behavior: what an animal does and how it does it • Both genetic & environmental factors • Essential for survival and reproduction • Subject to natural selection over time

5. How do animals behaviors develop?Scientists ask this question to try and determine what functions certain animal behaviors accomplish and how they arose in the first place. Niko Tinbergen was a pioneer in the field animal behavior and developed the following questions to try and understand any animal's behavior: • What stimulus elicits the behavior, and what physiological mechanisms mediate the response? • How does the animal's experience during growth and development influence the response? • How does the behavior aid survival and reproduction? • What is the behavior's evolutionary history? These question fall into 2 categories…..

6. Understanding behavior • 1. Proximate cause: “how” a behavior occurs or is modified • 2. Ultimate cause: “why” a behavior in context of natural selection • These questions define behavioral ecology: the study of ecological and evolutionary basis for animal behavior A courting pair of East Asian red-crowned cranes.

7. Behavior Requires Communication • Particularly among animals (who tend to have the most interesting behaviors) • Communication takes many forms. It always involves a signal transmitted and received • Signal: stimulus that causes a change in behavior; basis of animal communication 1. Chemical Signals • Pheromones - chemicals emitted by members of one species that affect other members of the species (eg. Queen bee, fruit fly, fish, termites, trees, humans) • Example Minnows in a tank responding to an alarm substance (from the skin of a predatory fish) 2. Symbolic Signals • Info encode in abstracted symbols (calls, gestures) • Example: honeybee workers communicate the location of nectars sources to hive mates via “waggle dance”

8. (a) Minnows are widely dispersed in an aquarium before an alarm substance is introduced. (b) Within seconds of the alarm substance being introduced, minnows aggregate near thebottom of the aquarium and reduce their movement. Figure 51.9a, b • When a minnow or catfish is injured • An alarm substance in the fish’s skin disperses in the water, inducing a fright response among fish in the area

9. Honeybee dance language known as waggle dance • Used to inform other bees about distance and direction of travel to food sources • Watch video below (54 seconds). Must know waggle dance https://www.youtube.com/watch?v=-7ijI-g4jHg

10. Volley period EXPERIMENT Charles Henry, Lucía Martínez, and ent Holsinger crossed males and females of Chrysoperla plorabunda and Chrysoperla johnsoni, two morphologically identical species of lacewings (insects) that sing different courtship songs. SONOGRAMSChrysoperla plorabunda parent Vibration volleys Standard repeating unit crossed with Chrysoperla johnsoni parent Volley period Standard repeating unit The researchers recorded and compared the songs of the male and female parents with those of the hybrid offspring that had been raised in isolation from other lacewings. Auditory Communication • Experiments with various insects • Have shown that courtship songs are under genetic control

11. F1 hybrids, typical phenotype Volley period RESULTS The F1 hybrid offspring sing a song in which the length of the standard repeating unit is similar to that sung by the Chrysoperla plorabunda parent, but the volley period, that is, the interval between vibration volleys, is more similar to that of the Chrysoperla johnsoni parent. CONCLUSION The results of this experiment indicate that the songs sung by Chrysoperla plorabunda and Chrysoperla johnsoni are under genetic control. Standard repeating unit

12. A. Signals…. • Visual signals – eg. Warning flash of white of a mockingbird's wing • Tactile(touch) – eg. Male fruit fly taps female fly Courtship behavior of fruit flies

13. 2 types of behaviors • 1. Innate – entirely under genetic controls, developmentally fixed • 2. Learned- requires experience

14. Simple Innate behavior • Fixed action patterns (FAPs):sequence of unlearned acts that are unchangeable and usually carried to completion • Triggered by sign stimulus, a sensory environment stimulus • Ensures that activities essential to survival are performed correctly without practice • Eg. goose & egg, click on link below 10 seconds • https://www.youtube.com/watch?v=vUNZv-ByPkU

15. Fixed Action Patterns READ!! • 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.

16. Fixed Action Patterns Example 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.

17. 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.

18. 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 3 spine males is a FAP triggered by an object red, underside the sign stimulus. • Watch video below 1 min https://www.youtube.com/watch?v=ZfcGZCGdGVE&feature=related

20. 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. 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.

21. Fixed Action Patterns (FAP) Digger wasp Do humans exhibit Fixed Action Patterns? The “eyebrow-flash”

22. Complex Innate Behavior • While there is a limit on the amount of behavioral complexity that can be genetically programmed, there are many example of complex innate behaviors • Example: Mating ritual in drosophila

23. 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.

24. Innate: Directed Movements • Kinesis: movement, a simple change in activity or turning rate in response to a stimulus. Response to stimuli is nondirectional • Taxis: change in directions, automatic movement toward or away from a stimulus, oriented movement +/- from stimulus(eg. phototaxis, chemotaxis, geotaxis) Kinesis increases the chance that a sow bug will encounter and stay in a moist environment. Another ex: scattering cockroaches when lights are turned on Positive rheotaxis keeps trout facing into the current, the direction from which most food comes.

25. 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”.

26. Chemotaxis- movement by a cell or organism in reaction to a chemical stimuli • Positive Chemotaxis- movement toward a chemical stimuli • Negative Chemotaxis- movement away from a stimuli • Geotaxis- movement away or toward gravitational force

27. Plant Behaviors (always innate) • Responses to environment • Governed by hormonal signals • Phototaxis – movement towards light • Photoperiodism – response to different times of day

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

29. Simple non-associative learning • Habituation is the relatively persistent waning of a response to a stimulus when the expected result does not occur. • Sensory Adaptationis 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.

30. 2. Learned behaviors: behaviors that are modified based on specific experiences5 TYPES of Learning behavior:

31. Types of Learning • Imprinting: learning + innate components • Limited to sensitive periodin life, generally irreversible • Young animals go through a critical period where after they follow the organism present during the period • Example • Lorenz’ imprinting in greylag young geese and other birds

32. 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.

33. Captive breeding programs for endangered species must provide proper imprinting models Pilot wearing crane suit acts as a surrogate parent to teach young whooping cranes a migration route

34. 2. Spatial Learning- The establishment of memories that reflect the physical structure of the environment • Cognitive Map: internal representation of spatial relationship among objects in an animal’s surroundings Birds use spatial maps to relocate nut caches

35. Example: Association of physical markers (pine cone ring: with nest location in Digger wasps. Some organisms move in response to a recognized object or environmental cue, a landmark. Experiment Nest RESULTS • use of landmarks- animal orients behavior patterns to objects in the environment No nest Nest

36. 3. Cognition: process of knowing that involves awareness, reasoning, recollection, judgment • Problem-solving behavior relies on cognition

37. Example: Cognition • An experiment that demonstrates support for the hypothesis that honeybees can remember and distinguish same from different. Honeybees learned in color maze that choosing the same two color gives them a reward. When moved into a pattern maze, the bees chose the same 2 patterns in a row

38. 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

39. 4. Associative Learning: ability to associate one stimulus with another (eg. monarchs = foul taste) Example: A bluejay learns to associate eating a monarch butterfly with subsequent vomiting Connecting one environmental feature with another

40. Associative Learning • 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.

41. 4. Associate learning continued.. A. Classical conditioning: arbitrary stimulus associated with particular outcome (eg. Pavlov’s dogs: salivate with ringing bell)

42. 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.

43. Classical conditioning • Ivan Pavlov’s dogs • connect reflex behavior (salivating at sight of food) to associated stimulus (ringing bell)

44. B. Operant conditioning: another type of associative learning • Trial-and-error learning • Associate its own behavior with reward or punishment

45. B. F. Skinner Operant conditioning • Skinner box mouse learns to associate behavior (pressing lever) with reward (food pellet)

46. 1. Reinforcing a voluntary behavior. Positive reinforcement: A child receives a 'gold star' at school for behaving well, reward • 2. Negative reinforcement: A child does his or her homework to stop her parents from nagging. Punishment: Grounding a child for behaving inappropriately

47. 5. Social learning: learning by observing others Vervet monkeys learning correct use of alarm calls.

48. Behavior Evolves • 1. Behavior and Genetics • 2. Behavior and the Environment • 3. Behavior and Fitness • 4. Altruism

49. 1. Behavior and Genetics • What is the relationship between an organism’s genetics and its behaviors. • There is always a relationship. Even if it is just for the ability to learn • 2 Examples • A. Insect Calls • B. Migration

50. B. Migration • Regular, long-distance change in location • Environmental cues: sun, stars, earth’s magnetic field, landmarks