Gibson, B. M., & Shettleworth, S. J. (2005). Place versus response learning revisited: tests of blocking on the radial maze. Behav Neurosci, 119(2), 567–586.
Abstract: Neurobiological and behavioral research indicates that place learning and response learning occur simultaneously, in parallel. Such findings seem to conflict with theories of associative learning in which different cues compete for learning. The authors conducted place+response training on a radial maze and then tested place learning and response learning separately by reconfiguring the maze in various ways. Consistent with the effects of manipulating place and response systems in the brain (M. G. Packard & J. L. McGaugh, 1996), well-trained rats showed strong place learning and strong response learning. Three experiments using associative blocking paradigms indicated that prior response learning interferes with place learning. Blocking and related tests can be used to better understand how memory systems interact during learning.
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Skov-Rackette, S. I., & Shettleworth, S. J. (2005). What do rats learn about the geometry of object arrays? Tests with exploratory behavior. J Exp Psychol Anim Behav Process, 31(2), 142–154.
Abstract: Six experiments using habituation of exploratory behavior tested whether disoriented rats foraging in a large arena encode the shapes of arrays of objects. Rats did not respond to changes in position of a single object, but they responded to a change in object color and to a change in position of 1 object in a square array, as in previous research (e.g., C. Thinus-Blanc et al., 1987). Rats also responded to an expansion of a square array, suggesting that they encoded sets of interobject distances rather than overall shape. In Experiments 4-6, rats did not respond to changes in sense of a triangular array that maintained interobject distances and angles. Shapes of object arrays are encoded differently from shapes of enclosures.
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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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Shettleworth, S. J. (2003). Memory and hippocampal specialization in food-storing birds: challenges for research on comparative cognition. Brain Behav Evol, 62(2), 108–116.
Abstract: The three-way association among food-storing behavior, spatial memory, and hippocampal enlargement in some species of birds is widely cited as an example of a new 'cognitive ecology' or 'neuroecology.' Whether this relationship is as strong as it first appears and whether it might be evidence for an adaptive specialization of memory and hippocampus in food-storers have recently been the subject of some controversy [Bolhuis and Macphail, 2001; Macphail and Bolhuis, 2001]. These critiques are based on misconceptions about the nature of adaptive specializations in cognition, misconceptions about the uniformity of results to be expected from applying the comparative method to data from a wide range of species, and a narrow view of what kinds of cognitive adaptations are theoretically interesting. New analyses of why food-storers (black-capped chickadees, Poecile Atricapilla) respond preferentially to spatial over color cues when both are relevant in a memory task show that this reflects a relative superiority of spatial memory as compared to memory for color rather than exceptional spatial attention or spatial discrimination ability. New studies of chickadees from more or less harsh winter climates also support the adaptive specialization hypothesis and suggest that within-species comparisons may be especially valuable for unraveling details of the relationships among ecology, memory, and brain in food-storing species.
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Gibson, B. M., & Shettleworth, S. J. (2003). Competition among spatial cues in a naturalistic food-carrying task. Learn Behav, 31(2), 143–159.
Abstract: Rats collected nuts from a container in a large arena in four experiments testing how learning about a beacon or cue at a goal interacts with learning about other spatial cues (place learning). Place learning was quick, with little evidence of competition from the beacon (Experiments 1 and 2). Rats trained to approach a beacon regardless of its location were subsequently impaired when the well-learned beacon was removed and other spatial cues identified the location of the goal (Experiment 3). The competition between beacon and place cues reflected learned irrelevance for place cues (Experiment 4). The findings differ from those of some studies of associative interactions between cue and place learning in other paradigms.
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Shettleworth, S. J. (1985). Handling time and choice in pigeons. J Exp Anal Behav, 44(2), 139–155.
Abstract: According to optimal foraging theory, animals should prefer food items with the highest ratios of energy intake to handling time. When single items have negligible handling times, one large item should be preferred to a collection of small ones of equivalent total weight. However, when pigeons were offered such a choice on equal concurrent variable-interval schedules in a shuttlebox, they preferred the side offering many small items per reinforcement to that offering one or a few relatively large items. This preference was still evident on concurrent fixed-cumulative-duration schedules in which choosing the alternative with longer handling time substantially lowered the rate of food intake.
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Shettleworth, S. J. (1978). Reinforcement and the organization of behavior in golden hamsters: Pavlovian conditioning with food and shock unconditioned stimuli. J Exp Psychol Anim Behav Process, 4(2), 152–169.
Abstract: The effects of Pavlovian conditioned stimuli (CSs) for food or shock on a variety of behaviors of golden hamsters were observed in three experiments. The aim was to see whether previously reported differences among the behaviors produced by food reinforcement and punishment procedures could be accounted for by differential effects of Pavlovian conditioning on the behaviors. There was some correspondence between the behaviors observed to the CSs and the previously reported effects of instrumental training. However, the Pavlovian conditioned responses (CRs) alone would not have predicted the effects of instrumental training. Moreover, CRs depended to some extent on the context in which training and testing occurred. These findings, together with others in the literature, suggest that the results of Pavlovian conditioning procedures may not unambiguously predict what system of behaviors will be most readily modified by instrumental training with a given reinforcer.
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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. (2001). Animal cognition and animal behaviour. Anim. Behav., 61(2), 277–286.
Abstract: Cognitive processes such as perception, learning, memory and decision making play an important role in mate choice, foraging and many other behaviours. In this review, I summarize a few key ideas about animal cognition developed in a recent book (Shettleworth 1998, Cognition, Evolution and Behaviour) and briefly review some areas in which interdisciplinary research on animal cognition is currently proving especially productive. Cognition, broadly defined, includes all ways in which animals take in information through the senses, process, retain and decide to act on it. Studying animal cognition does not entail any particular position on whether or to what degree animals are conscious. Neither does it entail rejecting behaviourism in that one of the greatest challenges in studing animal cognition is to formulate clear behavioural criteria for inferring specific mental processes. Tests of whether or not apparently goal-directed behaviour is controlled by a representation of its goal, episodic-like memory in birds, and deceptive behaviour in monkeys provide examples. Functional modelling has been integrated with analyses of cognitive mechanisms in a number of areas, including studies of communication, models of how predator learning and attention affect the evolution of conspicuous and cryptic prey, tests of the relationship betweeen ecological demands on spatial cognition and brain evolution, and in research on social learning. Rather than a `new field' of cognitive ecology, such interdisciplinary research on animal cognition exemplifies a revival of interest in proximate mechanisms of behaviour.
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Ratcliffe, J. M., Fenton, M. B., & Shettleworth, S. J. (2006). Behavioral flexibility positively correlated with relative brain volume in predatory bats. Brain Behav Evol, 67(3), 165–176.
Abstract: We investigated the potential relationships between foraging strategies and relative brain and brain region volumes in predatory (animal-eating) echolocating bats. The species we considered represent the ancestral state for the order and approximately 70% of living bat species. The two dominant foraging strategies used by echolocating predatory bats are substrate-gleaning (taking prey from surfaces) and aerial hawking (taking airborne prey). We used species-specific behavioral, morphological, and ecological data to classify each of 59 predatory species as one of the following: (1) ground gleaning, (2) behaviorally flexible (i.e., known to both glean and hawk prey), (3) clutter tolerant aerial hawking, or (4) open-space aerial hawking. In analyses using both species level data and phylogenetically independent contrasts, relative brain size was larger in behaviorally flexible species. Further, relative neocortex volume was significantly reduced in bats that aerially hawk prey primarily in open spaces. Conversely, our foraging behavior index did not account for variability in hippocampus and inferior colliculus volume and we discuss these results in the context of past research.
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