1. Learning A “relatively permanent change in behavior as a result of experience” - what is learning? - which behaviors are learned? - why learn or when will learning evolve? - evolution of learning and forgetting

2. A simple experimental example Can fish learn about predation risk in a body of water through chemical cues? 1) They react with an appropriate anti-predatory behavior 2) They retain this reaction for a period of time

3. ++ DW PO TSE Farm raised juvenile brown trout

4. Short-term effects

5. 4-days later

6. 21-days later A “relatively permanent change in behavior as a result of experience”

7. Eagle alarm Leopard alarm Snake alarm Socially acquired predator avoidance

8. Infants N=24 Raptors Non-raptors Goshawk Snake Bateleur Tawny Fish Eagle Eagle Eagle Vulture Bee-eater Juveniles N=53 Raptors Non-raptors Goshawk Snake Bateleur Martial Crowned Tawny Fish Harrier Owl Eagle Eagle Eagle Eagle Eagle Vulture Stork Bustard Adults N=55 Raptors Non-raptors Goshawk Snake Martial Crowned Tawny Fish Hawk Owl Eagle Eagle Eagle Eagle Eagle Eagle Vulture < 5 6-10 11-15 >15 How do infants come to recognize the association between an alarm call and 1-2 predators?

9. 100% 64% 14% 3% 4% Probability of adult alarm calling after infant alarm call by avian species – Reinforcers?

10. 3/4 4/5 5/6 6/7 100 Responses of infants to playbacks of adult alarm calls Number of infants responding correctly incorrectly

11. Vervet eagle alarm Starling eagle alarm Vervet eagle alarm Vervet leopard alarm Starling eagle alarm Vervet leopard alarm habituation Vervets habituate to call (learn it’s a false alarm) Learning of vervet eagle alarm extends to starling raptor alarm But learning does not extend between classes of predators

12. Learning curves associated with locating and extracting nectar as a function of experience

13. detections

14. Simple recognition learning – In this example, birds learn who their neighbors are and associate them to a particular place – the territory boundary songs per min “Dear-enemy” effect

15. Learning is a proximal cause of animal behavior because learning itself is a strategy/trait subject to natural selection Under this perspective we can redefine learning as a tool that allows individuals to adjust their behavior to the local state of their world – i.e., to the set of local spatial, temporal, and social circumstances

16. Learning is a proximal cause of animal behavior because learning itself is a strategy/trait subject to natural selection Under this perspective we can redefine learning as a tool that allows individuals to adjust their behavior to the local state of their world – i.e., to the set of local spatial, temporal, and social circumstances. - Where is food and what make-up is it? - Who are my territorial neighbors – do I recognize them? - What predators are nearby and what are their intentions?

17. Learning is a proximal cause of animal behavior because learning itself is a strategy/trait subject to natural selection Under this perspective we can redefine learning as a tool that allows individuals to adjust their behavior to the local state of their world – i.e., to local spatial, temporal, social, and causal relationships Where is food and what make-up is it? Who are my territorial neighbors – do I recognize them? What predators are nearby and what are their intentions? - recognizing kin from non-kin - social hierarchies: recognizing strangers from non-strangers - remembering past interactions (winners and losers) - recognizing high quality mates

18. Within-lifetime Predictability Low High Between-generation Predictability learn ignore experience ignore experience ignore experience Dave Stephen’s Model Low experience in the past is not useful High it is useful If offspring’s environment is nearly identical, fixed genetic transmission is favored over costly learning experience is useful within generation, and learning anew is favored each generation Evolution of Learning

19. Summary: - Learning is a likely element of proximal causation for most of the behaviors discussed in the remainder of the class - More importantly, the entire learning process is under evolutionary selection pressure - We can broadly understand the evolution of learning in response to the predictability of an organism’s environment in its own lifetime and beyond or we can ask detailed questions about the evolution of learning under specific contexts with a cost-benefit analysis ….

20. e.g. members of each new generation may find food in different places, but there is always value in being able to learn where food is learn Low between-generation predictability High within-lifetime predictability The conditions that bring about learning should be reliable correlates of the state of the world the animal needs to adjust to This differs from classical views of learning that assumed: (1) The ability to learn is an unadulterated good (2) Learning abilities are human-like: general and unlimited Dave Stephen’s Model

21. Brd Parasitism Tailorbird feeding a Plaintive Cuckoo

22. Brown-headed Cowbird