Questions in the study of foraging behavior ----------------------------1. How do animals select prey? 2. When should animals leave one feeding site and go on to another? 3. How should animals divide their time between food-gathering and other activities? 10 _ LOAD _ 7 4 _ _ 1 | | Travel Time | | | Searching Time | TIME 10 _ LOAD _ 7 4_ Y1 _ 1 | | Travel Time | X1 | | Searching Time | TIME 10 _ Load/Time is maximized at X2,Y2 LOAD _ 7 Y2 4_ _ 1 | | Travel Time | | X2 | Searching Time | TIME 10 _ LOAD _ 7 4 Long optimum _ Short optimum _ 1 | | Travel Time | X1 | X2 | Searching Time | TIME _ Load _ _ _ | | | | | | Round trip travel time STEPS IN ANALYSIS 1. Determine relevant variables through observation 2. Establish (you think) how these variables interact 3. Make predictions 4. Test Crows feeding on whelks Select the largest whelks available Fly roughly 5.5 m high to drop the whelk on the rocks below If whelk doesn’t break open, select the same whelk to drop again Patches Problem: When to leave a patch? Constraints that affect “optimal” foraging behavior 1. Physiological constraints 2. Motivational constraints 3. Ecological constraints 4. Life history constraints Intake_ of aquatic _ plants (gm) Energy constraint _ Sodium constraint _ Rumen constraint | | | | | | Intake of terrestrial plants (gm) Constraints that affect “optimal” foraging behavior 1. Physiological constraints 2. Motivational constraints 3. Ecological constraints 4. Life history constraints Food Choice (reward x probability) reward probability 2 pellets x 1.0 average yield of 2 0 pellets x 4 pellets x 0.5 0.5 average yield of 2 Testing Conditions Food Choice (reward x probability) Starved for 1 hour 2 pellets x 1.0 = avg. 2 Starved for 4 hours (0 pellets x 0.5) + (4 pellets x 0.5) = avg. 2 Testing Conditions Food Choice (reward x probability) Starved for 1 hour 2 pellets x 1.0 = avg. 2 Starved for 4 hours (0 pellets x 0.5) + (4 pellets x 0.5) = avg. 2 Constraints that affect “optimal” foraging behavior 1. Physiological constraints 2. Motivational constraints 3. Ecological constraints 4. Life history constraints Constraints that affect “optimal” foraging behavior 1. Physiological constraints 2. Motivational constraints 3. Ecological constraints 4. Life history constraints Life History Evolution The study of how individuals allocate, throughout life, time and energy to various fundamental activities, such as growth and reproduction Life Histories: An inherent trade-off Investment in any one activity limits an animal’s ability to invest in others. As applied to reproduction, a parent’s dilemma: investment in any one offspring limits an animal’s ability to invest in others. Components of Life Histories: Where the trade-offs occur • Growth and Development • Reproduce Early or Delay • Clutch Size vs. Clutch Number • Offspring Size and Offspring Number • Offspring Size and Parental Care Life histories: the major questions Why do organisms age and die? How many offspring should an individual produce in a given time? How big should each offspring be? Life history traits – characteristics of an individual that influence survival and reproduction Age at maturity House Mouse African elephant 2 months 11 - 20 years Salmon 3-6 years Life history traits – characteristics of an individual that influence survival and reproduction Number of offspring produced House Mouse African elephant 5-8 young every month 1 calf every 3-8 years Salmon 1,500 to 8,000 eggs once Life history traits – characteristics of an individual that influence survival and reproduction Number of reproductive events House Mouse ~6-12 African elephant ~3 - 10 Salmon 1 Life history traits – characteristics of an individual that influence survival and reproduction Lifespan House Mouse ~2 years African elephant 60 - 70 years Salmon 3-6 years Reproductive strategies K-strategists: •long lived •produce few offspring •parental care Population is controlled by density-dependent limiting factors - e.g. food Reproductive strategies r-strategists: •small •short life •no parental care •many offspring Gypsy moth caterpillars Population is controlled by densityindependent limiting factors: weather, pond drying 12-Arm Radial Maze 2.1m _ 80 % _ Corr. 40 _ _ | | | .25 .50 1 | | 2 4 Delay (hours) | 8 12 5 4 9 10 8 11 7 1 3 6 2 12 Food-storing Birds Clark’s nutcracker: 33,000 seeds, 7,500 sites Pinon jays: 22,000 seeds, clumped Scrub jays: 6,000 seeds Cache retrieval in corvids % correct, 1st 4 choices 100 Nutcracker 90 Pinyon jay 80 70 Scrub jay 60 Mexican jay 50 0 2 4 6 8 10 12 Block (avg of 5 trials) Kamil et al. 1994 _ % 80 correct _ 60 40 _ _ 20 Nutcrackers Pinyon jays Scrub jays Mexican jays | | | | | | | | | | | 50 100 150 200 250 300 Retention interval (min) | | Meadow Vole: Polygynous Female ranges Male range Prairie Vole: Monogamous ? ? Olson et al. 1995 Strict behaviorism: Any stimulus can, through conditioning, be associated with any response or reinforcer Learning is a general process phenomenon: All associations are learned equally easily All responses are reinforced equally easily Biological constraints on learning Saccharine taste + lights + noise BECAME SICK SHOCKED (Garcia & Koelling 1966) Biological constraints on learning Saccharine taste + lights + noise BECAME SICK Avoided saccharine, But no fear of light or noise SHOCKED Fear reaction to light and noise, but no aversion to saccharine (Garcia & Koelling 1966) Summary: Economic decisions Increasing evidence that animals make “calculations” when foraging -- make adaptive “choices” among alternative foods -- estimate past rates of return and compare them with current rates Summary: Memory 1. Natural selection has shaped the minds and behaviors of animals so that they optimize (as near as possible) the exploitation of their environment -- species differences in memory -- memory of a very specific sort -- species differences in the brain structures that support memory -- sex differences, too -- differences in the kind of associations that are formed