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Seyfarth, R. M., & Cheney, D. L. (2002). What are big brains for? Proc. Natl. Acad. Sci. U.S.A., 99(7), 4141–4142.
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Shettleworth, S. J. (1972). Stimulus relevance in the control of drinking and conditioned fear responses in domestic chicks (Gallus gallus). J Comp Physiol Psychol, 80(2), 175–198.
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Shettleworth, S. J. (1985). Foraging, memory, and constraints on learning. Ann N Y Acad Sci, 443, 216–226.
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Shettleworth, S. J. (1993). Varieties of learning and memory in animals. J Exp Psychol Anim Behav Process, 19(1), 5–14.
Abstract: It is often assumed that there is more than one kind of learning--or more than one memory system--each of which is specialized for a different function. Yet, the criteria by which the varieties of learning and memory should be distinguished are seldom clear. Learning and memory phenomena can differ from one another across species or situations (and thus be specialized) in a number of different ways. What is needed is a consistent theoretical approach to the whole range of learning phenomena, and one is explored here. Parallels and contrasts in the study of sensory systems illustrate one way to integrate the study of general mechanisms with an appreciation of species-specific adaptations.
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Shettleworth, S. J. (2005). Taking the best for learning. Behav. Process., 69(2), 147–9; author reply 159–63.
Abstract: Examples of how animals learn when multiple, sometimes redundant, cues are present provide further examples not considered by Hutchinson and Gigerenzer that seem to fit the principle of taking the best. “The best” may the most valid cue in the present circumstances; evolution may also produce species-specific biases to use the most functionally relevant cues.
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Shettleworth, S. J., & Krebs, J. R. (1982). How marsh tits find their hoards: the roles of site preference and spatial memory. J Exp Psychol Anim Behav Process, 8(4), 354–375.
Abstract: Marsh tits (Parus palustris) store single food items in scattered locations and recover them hours or days later. Some properties of the spatial memory involved were analyzed in two laboratory experiments. In the first, marsh tits were offered 97 sites for storing 12 seeds. They recovered a median of 65% of them 2-3 hr later, making only two errors per seed while doing so. Over trials, they used some sites more often than others, but during recovery they were more likely to visit a site of any preference value if they had stored a seed there that day than if they had not. Recovery performance was much worse if the experimenters moved the seeds between storage and recovery. A fixed search strategy that had some of the same average properties as the tits' search behavior also did worse than the real birds. In Experiment 2, any tendency to visit the same sites on successive daily tests in the aviary was placed in opposition to memory for storage sites by allowing the tits to store more seeds 2 hr after storing a first batch. They tended to avoid individual storage sites holding seeds from the first batch. When the tits searched for all the seeds 2 hr later, they tended to recover more seeds from the second batch than from the first, i.e., there was a recency effect.
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Shettleworth, S. J., & Plowright, C. M. (1992). How pigeons estimate rates of prey encounter. J Exp Psychol Anim Behav Process, 18(3), 219–235.
Abstract: Pigeons were trained on operant schedules simulating successive encounters with prey items. When items were encountered on variable-interval schedules, birds were more likely to accept a poor item (long delay to food) the longer they had just searched, as if they were averaging prey density over a short memory window (Experiment 1). Responding as if the immediate future would be like the immediate past was reversed when a short search predicted a long search next time (Experiment 2). Experience with different degrees of environmental predictability appeared to change the length of the memory window (Experiment 3). The results may reflect linear waiting (Higa, Wynne, & Staddon, 1991), but they differ in some respects. The findings have implications for possible mechanisms of adjusting behavior to current reinforcement conditions.
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Shettleworth, S. J., & Sutton, J. E. (2005). Multiple systems for spatial learning: dead reckoning and beacon homing in rats. J Exp Psychol Anim Behav Process, 31(2), 125–141.
Abstract: Rats homed with food in a large lighted arena. Without visual cues, they used dead reckoning. When a beacon indicated the home, rats could also use the beacon. Homing did not differ in 2 groups of rats, 1 provided with the beacon and 1 without it; tests without the beacon gave no evidence that beacon learning overshadowed dead reckoning (Experiment 1). When the beacon was at the home for 1 group and in random locations for another, there was again no evidence of cue competition (Experiment 2). Dead reckoning experience did not block acquisition of beacon homing (Experiment 3). Beacon learning and dead reckoning do not compete for predictive value but acquire information in parallel and are used hierarchically.
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Shier, D. M., & Owings, D. H. (2007). Effects of social learning on predator training and postrelease survival in juvenile black-tailed prairie dogs, Cynomys ludovicianus. Anim. Behav., 73(4), 567–577.
Abstract: We examined how social context and experience affected development of antipredator behaviour and subsequent postrelease survival in the black-tailed prairie dog. Captive-reared juveniles were initially exposed to four stimulus animals: a ferret, a rattlesnake, a hawk and a cottontail control (pretraining tests). Subjects were then trained with or without an adult female demonstrator. Training involved exposure to each stimulus animal two to three times over 5 weeks. After training, each juvenile was retested with each stimulus animal (post-training tests). During pretraining tests, juveniles responded differentially to the stimulus animals. They were least active with the snake, fled the most in tests with the hawk, and were less vigilant with the ferret than with the snake. Following training, juveniles trained with experienced adults were more wary with all three predators than juveniles trained without an experienced adult present. We then compared the antipredator behaviour of captive-reared juveniles trained with experienced adult females with that of wild-reared juveniles of the same age. For all behavioural measures except shelter use, wild-experienced animals differentiated more strongly among predator types than did captive-trained juveniles. One year after reintroduction, survivorship of juveniles trained with experienced adults was higher than that of juveniles trained without experienced adults, but did not differ from that of wild-reared juveniles. These findings provide the first evidence that social transmission of antipredator behaviour during training can enhance long-term survival following release and that as long as a social training regime is used, predator avoidance training can emulate experience acquired in the wild.
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Simpson, B. S. (2002). Neonatal foal handling. Appl. Anim. Behav. Sci., 78(2-4), 303–317.
Abstract: Recent interest has focused on the advantage of intensively handling young foals as a means of producing more tractable horses, accustomed to humans and receptive to training. To date, the effect of this intensive handling, dubbed “imprint training” in the popular literature, has not been tested. The present study compares seven foals handled intensively as neonates with eight untreated controls. The handling protocol started from 2-8 h after birth and continued daily for a total of 5 days. The protocol consisted of a series of stimuli and experiences that were each repeated until the foal no longer resisted or reacted negatively. Subsequently, foals were tested before weaning, at 4 months of age. Results indicated that handled foals (HF) ranked higher than control foals (CF) in subjective ratings of calmness (*P<0.0117) and friendliness (*P<0.0001) and in several specific handling tasks (venipuncture *P<0.0220; placing in stock *P<0.0128). Although, in approach tests all foals but one allowed approach of a person to 4 m, significantly more HF approached the person than CF (P<0.0080). In stimulus tests, foals were presented specific stimuli to which they had been tested as neonates. Two of eight CF were too unruly and dangerous to test. Of foals that could be tested, CF required significantly more time to hook-up a heart rate monitor (**P<0.0055). Split-plot analysis indicated that HF had lower heart rates to initial left-sided stimuli, presented first, than CF (*P<0.0421). In response to right-sided stimuli, heart rate scores of CF were not significantly different from HF (P<0.2259), suggesting reduced reactivity over time due to a learning effect. Behavioral responses to specific stimuli did not differ between CF and HF, suggesting that neonatal handling has a general rather than specific effect on subsequent behavior. Cortisol concentrations were measured before and after testing and the difference calculated. All foals had higher post-testing levels than pre-testing levels. There was a significant difference between HF and CF, indicating greater reactivity among the CF (*P<0.050). In general, the results indicated that foals handled as neonates were more tractable and less reactive. Specific neonatal handling tasks, such as sticking a finger up the foal's nose or patting the bottom of the foot, seemed to have no beneficial effect on related tasks such as passing a nasogastric tube or tapping with a farrier's hammer at 4 months of age. Mechanisms for the observed effect of neonatal handling require further investigation.
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