1. Chapter 5: Learning Individual learning How do animals learn Why?
What?

2. FIGURE 5. 1. Facial learning in wasps
FIGURE 5.1. Facial learning in wasps. Set of images used for learning trials in wasps. To see how trials were run, look at panel A (P. fiscatus faces). A wasp was given the chance to pair one of the images in each row of panel A with a shock, and was then tested to see how quickly it learned to select the image that was not paired with the shock. A similar approach was used for all treatments. Wasps learned the facial images in panel A, but did not learn to pair an image with shock in panels B, C, or D. The “caterpillar treatment” (E) found that wasps did not learn to avoid the caterpillar image associated with negative stimulus, further suggesting that the learning was specific to conspecific facial learning. (Photo credit: Sheehan et al., 2011, Science, 334, 1272–1275)
Sheehan and Tibbetts, 2011

3. Individual Learning
Shettleworth (1998): Learning is a “relatively permanent change in behavior as a result from experience.”
Phenotypic Plasticity: production of different phenotypes as a result of different environmental conditions
(phenotype: the set of observable characteristics of an organism)

4. Dugatkin suggests: Learning is a subset of phenotypic plasticity
Changes in colonial bryozoans as a result of predation: spine production in response to predators in Membranipora membranacea
Dugatkin suggests: Learning is a subset of phenotypic plasticity
FIGURE 5.2. Inducible defenses. In some bryozoans, like Membranipora membranacea, colonies produce spines when predators are present. (A) Spines are shown protruding from a colony as a defense against predators (red arrows point to spines), and (B) of a colony of Membranipora membranacea. (Photo credits: Ken Lucas/Visuals Unlimited; © Sue Daly/naturepl.com)

5. Membranipora membranacea colonies exposed to predator tainted water.
(Harvell, 1991)
FIGURE 5.3. PHENOTYPIC plasticity. When colonies of the bryozoan Membranipora membranacea are exposed to chemical stimuli from a predator, individuals in these colonies grow spines. This graph shows the response to a single “dose” of water conditioned with bryozoan predators. Large colonies produce more spines. (From Harvell, 1991, p. 4)

6. Learning is a subset of phenotypic plasticity; but all phenotypic plasticity is not necessarily learning.
Jablonski, et al, 2006, looked at wing flapping and tail flipping in birds as a learned response. Birds may do this to “flush” insects from trees and the eat them.
This may be learned, or it may be fixed (see Lorenz , “fixed action pattern) or perhaps both.

7. Jablonski found that Painted Redstarts birds increase flapping when under branches in the field; but they also do this in the lab; even when they are not rewarded for the behavior.
“Naïve” birds have the same response as experienced birds.
Thus, increased flapping under branches is an example of phenotypic plasticity (producing different phenotypes under different environmental conditions), but it not learning.

8. Single stimulus learning: the blue stick experiment
Habituation versus Sensitization
FIGURE 5.4. Habituation and sensitization. Numerous times each day, a blue stick is placed in a rat’s cage. If the rat takes less and less notice of the stick, habituation has occurred. If the rat pays more attention to the blue stick over time, sensitization has taken place.

9. FIGURE 5. 5. Habituation as a problem
FIGURE 5.5. Habituation as a problem. In controlled laboratory experiments prey may habituate to the presence of a predator over time.

10. Pavlov’s work 1898 through 1930
Pavlovian Conditioning
Conditioned Stimulus: the blue stick, the bell, the stimulus that initially fails to produce a response
Unconditioned Stimulus: the cat odor, the meat powder, the stimulus that elicits a strong response

11. FIGURE 5.6. Paired stimuli. Five seconds after a blue stick (stimulus 1) is placed in a rat’s cage, the odor of a cat (stimulus 2) is sprayed in as well. The question then becomes: Will the rat pair the blue stick with danger (cat odor)?

12. The Conditioned Response
FIGURE 5.8. Conditioned response. If the rat pairs the blue stick (CS) and the cat odor (US), it will hide under the chips when the blue stick alone is presented. Such hiding represents a conditioned response (CR).

13. FIGURE 5. 9. Second-order conditioning
FIGURE 5.9. Second-order conditioning. The rat learns to respond to a second CS—the yellow light—with the conditioned response.

14. FIGURE 5.10. Overshadowing. The process of overshadowing is shown in two groups of rats.

15. FIGURE Blocking. Learning can be slowed down depending on prior association or lack of association between stimuli.

16. Operant (goal-directed) Response Instrumental Learning
Operant Conditioning
Operant (goal-directed) Response
Instrumental Learning
FIGURE Rats in a Skinner box. To test various theories of animal learning, rats are often placed in “Skinner boxes,” where they have to take an action (here, pressing a button) to get a reward of food or water. (Photo credit: Walter Dawn/Science Photo Library/Photo Researchers, Inc.)

17. Why do animal’s learn? Adaptation Natural Selection Do animals forget?
FIGURE How long to remember? Imagine a bee foraging at a nectar-producing flower. While it might be beneficial for the bee to remember the flower’s location, it might not be to remember specific nectar content, as that shifts within and between days.

18. Extinction is the weakening and ultimately the ending of the paired association of stimuli and response in learning experiments… forgetting
FIGURE Stomatopod threat. Male and female stomatopod crustaceans share a cavity for a few days before they breed. Although the males leave the breeding cavity soon after mating, they tend to remember their former mates and to be less aggressive toward them during the weeks that their brood remains in the cavity. This is a photo of a male Gonodactylus smithii in a threat position. (Photo credit: Roy Caldwell)

19. Learning in populations
FIGURE Zenaida doves. Zenaida doves from populations where individuals live in groups appear to be better at learning foraging tasks than individuals from populations where doves are territorial. (Photo credit: Jean-Philippe Paris)

20. FIGURE 5. 16. Group living and learning
FIGURE Group living and learning. More birds from the group-living population surpassed the “learning criteria” for foraging tasks than did birds that had lived alone (territorial population). (From Carlier and Lefebvre, 1996, p. 1203)

21. Group learning and antipredator response in three-spines sticklebacks
FIGURE Learning differences across populations. The number of fish that learned to avoid areas associated with predation. (A) Fish descended from individuals from low-predation sites. (B) Fish descended from individuals from high-predation sites. (From Huntingford and Wright, 1992)

22. Does Natural Selection favor the ability to learn?
Evolution of Learning
Does Natural Selection favor the ability to learn?
Is there accost to learning and can we select for the ability (genetic basis)?
FIGURE A trade-off between learning and life span. “Learning index scores” for normal (control) fruit flies and fruit flies from a line selected for artificially prolonged life spans. The difference between these groups suggests a trade-off between long life and the ability to learn. (From Burger et al., 2008)

23. Evolutionary tradeoffs and Environmental Stability
FIGURE 5.19 Stephens’s model for the evolution of learning. The key variables in this model are within-lifetime environmental predictability and between-generation environmental predictability.

24. What can animal’s learn? About predators About mates
About animal relationships
About aggression
FIGURE 5.20a. Learning and response to predators. (A) A hellbender salamander. (B) Hellbenders that were given the opportunity to pair the alarm secretion and the odor of a predator moved quickly when exposed to the odor of the predator alone. (Photo credit: Robert J. Erwin/Photo Researchers)

25. FIGURE 5. 20b. Learning and response to predators
FIGURE 5.20b. Learning and response to predators. (A) A hellbender salamander. (B) Hellbenders that were given the opportunity to pair the alarm secretion and the odor of a predator moved quickly when exposed to the odor of the predator alone. (Photo credit: Robert J. Erwin/Photo Researchers)

26. FIGURE Damselfly. Larval damselflies learn about predation threat through chemical cues. An adult damselfly is shown here. (Photo credit: Kim Taylor/natureplace.com)

27. FIGURE 5. 22. Chemically mediated changes
FIGURE Chemically mediated changes. Numerous aspects of damselfly behavior, including the frequency of feeding bites, head bends, and moves, changed as a function of whether the damselflies were exposed to chemical stimuli from a pike predator that had eaten mealworms, minnows, or damselflies. (From Chivers et al., 1996)

28. Learning about mates
Mating systems, parental investment and selection for learning
FIGURE 5.23a. Parental investment and learning ability. (A) Parental investment is shared in blue gourami, and differences in learning about pairmates between the sexes is small. (B) In Japanese quail, the females care for the young, as shown by this female at the nest with her eggs, and there is no parental investment by males. In this species, males show greater learning about mates abilities than females. (Photo credits: © Wil Meinderts/FotoNatura/Minden Pictures; public domain)

29. Learning about kin: Helpers at the nest and indirect fitness
FIGURE Learning who is kin. Young long-tailed tits (Aegithalos caudatus) often become helpers at the nests of their close genetic relatives, helping build nests and forage for food to feed the chicks. As helpers, they accrue indirect fitness benefits by contributing to the survival of their close genetic kin. (Photo credit: Andrew MacColl)

30. Learning may shape aggressive behavior
FIGURE Pavlovian learning in fish. Males that had learned to associate a light with the presence of another male were more aggressive when the light cue was present than were males that did not associate the light with the presence of another male. (From Hollis, 1984)

31. FIGURE 5. 27. Winners and losers
FIGURE Winners and losers. (A) Males that won in contest 1 were more likely to win in contest 2 (WW) than were males that had lost in contest 1 (LW). (B) Males that lost in contest 1 were more likely to lose in contest 2 (LL). (From Hollis et al., 1995, p. 129)