1. Animal Behavior
Chapter 51

2. Why study behavior?
Ethology— the study of behavior

3. Ethologist Niko Tinbergen, four questions should be asked about 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 mechanisms?
How does the behavior aid survival and reproduction?
What is the behavior’s evolutionary history?

4. What is behavior? Behavior
Defined as observable and coordinated responses to environmental stimuli.
Result of genetic & environmental factors
Innate Behaviors
inherited, “instinctive”
automatic & consistent
“Built-in”, no “learning curve”
Despite different environments, all individuals exhibit the behavior
Ex. early survival, reproduction, kinesis, taxis

5. Evolutionary perspective
Learned behaviors
Modified by experience
Variable, changeable
change with experience & environment
Flexible with a complex & changing environment
Ability to learn is inherited, but the behavior develops during animal’s lifetime

6. Levels of Behavioral Analysis
Proximate causation
“how” explanations
Environmental stimuli that trigger a behavior
Genetic, physiological, and anatomical mechanisms underlying a behavior
Ultimate causation
evolutionary significance
“why” explanations
Proximate cause questions
Male songbirds sing during the breeding season as a response to a high level of testosterone which binds to hormone receptors in the brain & triggers the production of song.
Ultimate cause questions
The male sings to defend territory from other males & to attract a female with which to reproduce. This is the evolutionary explanation for the male’s vocalization.
The red–crowned cranes, like many animals, breed in spring and early summer. A proximate question about the timing of breeding by this species might be, “How does day length influence breeding by red–crowned cranes”? A reasonable hypothesis for the proximate cause of this behavior is that breeding is triggered by the effect of increased day length on an animal’s production of and responses to particular hormones. Indeed, experiments with various animals demonstrate that lengthening daily exposure to light produces neural and hormonal changes that induce behavior associated with reproduction, such as singing and nest building in birds.
In contrast to proximate questions, ultimate questions address the evolutionary significance of a behavior. Ultimate questions take such forms as, Why did natural selection favor this behavior and not a different one? Hypotheses addressing “why” questions propose that the behavior increases fitness in some particular way. A reasonable hypothesis for why the red–crowned crane reproduces in spring and early summer is that breeding is most productive at that time of year. For instance, at that time, parent birds can find ample food for rapidly growing offspring, providing an advantage in reproductive success compared to birds that breed in other seasons.
Courtship behavior in cranes
 what…how… & why questions

7. Ex. Most people like fatty foods
What is the proximate cause?
Because it tastes good
What is the ultimate cause?
Because it is high in calories which are the stuff the body needs to function

8. Innate Behaviors-developmentally fixed
Taxis
Kinesis
Migration
FAP
Animal Signals & Communication

9. Innate Behaviors-Taxis
Automatic response toward or away from a stimulus.
Automatic movement toward (positive taxis) or from away (negative taxis) a stimulus
phototaxis
Chemotaxis
Examples:
Trout are and swim upstream so as not to be swept away.

10. Innate Behaviors- Kinesis
A change in the activity rate in response to a stimulus.
Randomly directed, unlike taxis.
Example:
Sow bugs are more active in dry areas and less in humid ones; this keeps them in moist environments.

11. Innate behavior: Fixed action patterns (FAP)
Unlearned
Usually carried to completion once started
Triggered by a sign stimulus
male sticklebacks exhibit aggressive territoriality

12. Fixed Action Patterns (FAP)
Digger wasp
Human babies always smile at a mask with two dark spots for eyes.
Do humans exhibit Fixed Action Patterns? This question was addressed by Irenaeus Eibl-Eibesfeldt and Hans Hass who worked at the Max-Planck-Institute for Behavioural Physiology in Germany. They created a Film Archive of Human Ethology of unstaged and minimally disturbed social behaviour.
They filmed people across a wide range of cultures with a right-angle reflex lens camera i.e. the subjects did not realize that they were being filmed because the camera lens did not appear to be pointing at them! Eibl-Eibesfeldt has identified and recorded on film, several human Fixed Action Patterns or human 'universals' e.g. smiling and the "eyebrow-flash"
Eibl-Eibesfeldt took these pictures of a Himba woman from Namibia (SW-Africa). She shows a rapid brow raising (between the second and third still images) which coincides with raising her eyelids. Because all the cultures he examined showed this behaviour, Eibl-Eibesfeldt concluded that it was a human 'universal' or Fixed Action Pattern.
Some Sphex wasps drop a paralyzed insect near the opening of the nest. Before taking provisions into the nest, the sphex first inspects the nest, leaving the prey outside. During the sphex's inspection of the nest an experimenter can move the prey a few inches away from the opening of the nest. When the sphex emerges from the nest ready to drag in the prey, it finds the prey missing. The sphex quickly locates the moved prey, but now its behavioral "program" has been reset. After dragging the prey back to the opening of the nest, once again the sphex is compelled to inspect the nest, so the prey is again dropped and left outside during another stereotypical inspection of the nest. This iteration can be repeated again and again, with the sphex never seeming to notice what is going on, never able to escape from its genetically-programmed sequence of behaviors. Douglas Hofstadter and Daniel Dennett have used this mechanistic behavior as an example of how seemingly thoughtful behavior can actually be quite mindless, the opposite of human behavioral flexibility that we experience as free will
Do humans exhibit Fixed Action Patterns?
The “eyebrow-flash”

13. Fixed Action Patterns Examples
Male stickleback fish will show aggression toward any shape that has a red area.
attack on red belly stimulus
court on swollen belly stimulus

14. Complex Innate behaviors
Migration
“migratory restlessness” seen in birds bred & raised in captivity
Migration is learned, but how to learn them is innate
Celestial navigation- by sun, stars, Earth magnetic fields
Sandpiper
Monarch
migration
Bird migration, a behavior that is largely under genetic control. Each spring, migrating western sandpipers (Calidris mauri), such as those shown here, migrate from their wintering grounds, which may be as far south as Peru, to their breeding grounds in Alaska. In the autumn, they return to the wintering grounds.
Bobolink
Golden plover
ancient fly-ways

15. Examples of Migration
Monarch butterflies from our area migrate to Mexico in the fall and the next generation flies north in the spring.
Snow geese fly 2,700 km from James Bay, Canada to Louisiana nonstop in 60 hours.
The green sea turtle migrates 2,200 km across the Atlantic ocean from the east coast of South America to Ascension Island (10 km across) every 2-3 years where the adult females lay eggs.

16. Biological Rhythms & Clocks
Circadian Rhythms
“Sleep, wake cycle”
Humans~about 24 hours long
Drosophila the normal clock is 24.2 hrs
Mutant flies have cycles of 19 & 28 hrs.
Diurnal —active during the day
Nocturnal —active during hours of darkness
Crepuscular —having greatest activity during twilight hours or at dawn or both

17. Biological Rhythms & Clocks
Lunar cycles
Ex. courtship in fiddler crabs occurs during the new and full moon

18. Biological Rhythms & Clocks
What controls the biological clock?
No single mechanism—an interaction of a number of biochemical processes
The pineal gland is thought to play a role in the timing system of rats, birds and some other vertebrates.
Hormone? Melatonin
In mammals, regions of the hypothalamus are involved.

19. Biological Rhythms & Clocks
How much is internal and how much is governed by external clues?
Answer: There is usually a strong endogenous (internal) component, but an exogenous (external) cue is necessary to keep the behavior properly timed in the real world.

20. Animal Signals & Communication
signal- behavior that causes a change in another animal’s behavior
Communication- transmission and reception of signals
Does not have to be purposeful
Ex. Honeybees show complex communication with symbolic language
Honeybee Dance Video

21. Social interaction requires communication
Pheromones
chemical signal that stimulates a response from other individuals
Ex. Female moths secrete chemicals which attract males.
Cheetahs and other cats mark their territory with urine, feces, and anal gland secretions.
alarm pheromones
sex pheromones

22. Communication: Auditory
Faster than chemical and also effective both night and day.
Can be modified by loudness, pattern, duration, and repetition.
Ex. Male crickets have calls and birds may have one song for courting, another for distress, and still another for marking territories.
Humpback whales have complicated songs.
Nonhuman primates have up to 40 different vocalizations.

23. Communication by song Bird song Insect song
species identification & mating ritual
mixed learned & innate
critical learning period
Insect song
mating ritual & song
innate, genetically controlled

24. Communication: Tactile
Occurs when one animal touches another.
Ex. A male leopard nuzzles a female’s neck to calm her and to stimulate her willingness to mate.

25. Honey bee communication
dance to communicate location of food source
waggle dance
Discovered by Karl von Frisch in the 1940s. Waggle dance indicates both distance to and direction of food.
View Waggle Dance AVI file: waggledance180x135.avi

26. Communication: Visual
Most often used by species that are active during the day.
Contests between males involve threat postures and possibly prevent fighting.
Ex. Male birds often put on courtship dances to attract females.
Much human communication is nonverbal such as smiling, blushing, body posture.

27. Learned Behavior: Modified behaviors by experience
Habituation
Imprinting
Associative learning
Classical conditioning
Operant conditioning
Cognition

28. Learning: Habituation
Loss of response to stimulus
“Cry-wolf” effect
Decrease in response to repeated occurrences of stimulus
enables animals to disregard unimportant stimuli
ex: falling leaves not triggering fear response in baby birds

29. Innate & Learning: Imprinting
Learning to form social attachments at a specific sensitive (critical ) period
both learning & innate components
Konrad Lorenz- spent time with geese hours after hatched
But how do the young know on whom—or what—to imprint? How do young geese know that they should follow the mother goose? The tendency to respond is innate in the birds; the outside world provides the imprinting stimulus, something to which the response will be directed. Experiments with many species of waterfowl indicate that they have no innate recognition of “mother.” They respond to and identify with the first object they encounter that has certain key characteristics. In classic experiments done in the 1930s, Konrad Lorenz showed that the most important imprinting stimulus in graylag geese is movement of an object away from the young. When incubator–hatched goslings spent their first few hours with Lorenz rather than with a goose, they imprinted on him, and from then on, they steadfastly followed him and showed no recognition of their biological mother or other adults of their own species. Again, there are both proximate and ultimate explanations

30. Conservation
Conservation biologists have taken advantage of imprinting by young whooping cranes as a means to teach the birds a migration route. A pilot wearing a crane suit in an Ultralight plane acts as a surrogate parent.
Wattled crane conservation
teaching cranes to migrate
Cranes also imprint as hatchlings, creating both problems and opportunities in captive rearing programs designed to save endangered crane species. For instance, a group of 77 endangered whooping cranes hatched and raised by sandhill cranes imprinted on the sandhill foster parents; none of these whooping cranes ever formed a mating pair–bond with another whooping crane. As a consequence, captive breeding programs now isolate young cranes and expose them to the sights and sounds of members of their own species. But imprinting can also be used to aid crane conservation
Young whooping cranes imprinted on humans in “crane suits” have been taught to follow these “parents” flying ultralight aircraft along new migration routes. And importantly, such cranes have formed mating pair–bonds with other whooping cranes.

31. Associative learning
Learning to associate a stimulus with a consequence
Operant conditioning
BF Skinner
Trial & error learning
Associate behavior with reward or punishment
Big Bang Theory Operant Conditioning of Penny
Classical conditioning
Pavlovian conditioning
Associate stimuli with reward or punishment
Classical Conditioning a 3 year old

32. Operant Conditioning
Gradual strengthening of stimulus-response connections.
Examples:
Teaching an animal a trick by rewarding correct behavior with affection or food.
B.F. Skinner’s experiments
Rats connect sugar treat with pressing a lever.

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

34. Critical period Sensitive phase for optimal imprinting
some behavior must be learned during a receptive time period
As a brood parasite, the Cuckoo never learn the song of their species as a nestling. Song development is totally innate.

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

36. Learning: Cognition
process of knowing that may include awareness, reasoning, recollection, and judgment
For example, honeybees can distinguish “same” from “different”

37. Learning: Problem Solving
Process of devising a strategy to overcome an obstacle
For example, chimpanzees can stack boxes in order to reach suspended food
problem-solving

38. Genetic components of behavior evolve through natural selection
Concept 51.4: Selection for individual survival and reproductive success can explain most behaviors
Genetic components of behavior evolve through natural selection
Behavior can affect fitness by influencing foraging and mate choice

39. Foraging Behavior
Natural selection refines behaviors that enhance the efficiency of feeding
Foraging, or food-obtaining behavior, includes recognizing, searching for, capturing, and eating food items
Natural selection favors different foraging behavior depending on the density of the population

40. Optimal Foraging Model
Optimal foraging model views foraging behavior as a compromise between benefits of nutrition and costs of obtaining food
The costs of obtaining food include energy expenditure and the risk of being eaten while foraging
Natural selection should favor foraging behavior that minimizes the costs and maximizes the benefits

41. Mating Behavior and Mate Choice
Mating behavior includes seeking or attracting mates, choosing among potential mates, and competing for mates
Mating behavior results from a type of natural selection called sexual selection
Needs of the young are an important factor constraining evolution of mating systems
Consider bird species where chicks need a continuous supply of food
A male maximizes his reproductive success by staying with his mate, and caring for his chicks (monogamy)

42. Sexual Selection and Mate Choice
In intersexual selection, members of one sex choose mates on the basis of certain traits
Intrasexual selection involves competition between members of the same sex for mates

43. Mate Choice by Females
Female choice is a type of intersexual competition
Females can drive sexual selection by choosing males with specific behaviors or features of anatomy
For example, female stalk-eyed flies choose males with relatively long eyestalks
Ornaments, such as long eyestalks, often correlate with health and vitality

44. Male Competition for Mates
Male competition for mates is a source of intrasexual selection that can reduce variation among males
Such competition may involve agonistic behavior, an often ritualized contest that determines which competitor gains access to a resource

45. Concept 51.5: Inclusive fitness can account for the evolution of altruistic social behavior
Some animals are largely solitary and join with a member of the opposite sex only for mating; others pair, bond and cooperate in the raising of offspring.
Still others form a society in which members are organized in a cooperative manner.
Society— a group of individuals of the same species that cooperate in an adaptive manner; e.g. bee hive, flock of birds, wolf pack, school of fish.

46. Social behaviors Interactions between individuals
develop as evolutionary adaptations
Result in survival and reproductive success
Agonistic behaviors
Dominance hierarchy
Cooperation
Altruistic behavior

47. Social behaviors Agonistic behaviors Threatening & Submissive rituals
symbolic, usually no harm done
ex: territoriality, competitor aggression
View Lifewire territoriality video: “lizards cost of defending-lifewire.swf”
Review setting up a behavior experiment:

48. Social behaviors Dominance hierarchy social ranking within a group
Pecking order

49. Dominance Hierarchies
A higher ranking animal has greater access to resources than a lower ranking animal.
Decided by confrontation during which one animal gives way to another.
Once established, little or no time is wasted in fighting.
Dominant male mate more often with the females.

50. Territoriality-- Protecting an area against other individuals.
Male songbirds sing to signify their territories and other males know to stay away. The song also alerts females to presence of a male.
May be adaptive—tends to reduce conflict, to control population growth, and to ensure the most efficient use of resources by spacing organisms throughout a habitat.

51. Social behaviors Cooperation working together in coordination
Pack of African dogs hunting wildebeest cooperatively
White pelicans “herding” school of fish

52. Social behaviors Altruistic behavior (selflessness)
Reduces individual fitness but increases fitness of others in a population
kin selection
increasing survival of close relatives passes these genes on to the next generation
How can this be of adaptive value?
Belding ground squirrel

53. Examples of Altruistic Behaviors
Inclusive fitness- total effect an individual has on proliferating its genes by producing offspring and helping close relatives produce offspring
A female chimp often mates with several males in a group because they are all related and share genes in common.

54. Reciprocal Altruism
Altruistic behavior toward unrelated individuals can be adaptive if the aided individual returns the favor in the future
This type of altruism is called reciprocal altruism
Reciprocal altruism is limited to species with stable social groups where individuals meet repeatedly and those who don’t reciprocate are punished
helping out a friend, and expecting something back in return

55. Social Behavior & Reproduction
Natural selection has favored mechanisms that promote successful reproduction.
Behavior is thus adaptive—behavioral traits can evolve.
Sexual competition among males has contributed to the evolution of large size, brilliant breeding colors, antlers, etc.
Known as sexual selection and the traits are called secondary sexual characteristics.

56. Social Behavior & Reproduction
Courtship rituals may be long and elaborate.
May serve as a sign signal to trigger nest building and ovulation.
Male spiders offer food.
Female praying mantis eats head of male

57. Behavior: Nature or Nurture?
Behavior is also shaped by environmental influences (nurture).
Studies of human twins have been used to help determine to what extent behavior is inherited.
Identical twins come from a single zygote. Fraternal twins are derived from two different fertilized eggs.
Fraternal twins, even when raised together, do not have similar behaviors.

58. Identical twins raised separately are sometimes very similar; for example, the German twins raised in Germany and the Caribbean both liked sweet liqueurs, stored rubber bands on their wrists, read magazines from back to front, dipped buttered toast in their coffee, and had similar personalities.
Data seems to show that about 50% in human personality traits are due to polygenic inheritance and 50% are due to environmental influences.

59. You should now be able to:
State Tinbergen’s four questions and identify each as a proximate or ultimate causation
Distinguish between the following pairs of terms: kinesis and taxis, classical and operant conditioning, intersexual and intrasexual
Suggest a proximate and an ultimate cause for imprinting in newly hatched geese
Explain how associative learning may help a predator avoid toxic prey

60. Describe optimal foraging theory
Describe how the certainty of paternity may influence the development of mating systems
Define altruistic behavior and relate the coefficient of relatedness to the concept of altruism
Distinguish between kin selection and reciprocal altruism