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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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Sole, L. M., Shettleworth, S. J., & Bennett, P. J. (2003). Uncertainty in pigeons. Psychon Bull Rev, 10(3), 738–745.
Abstract: Pigeons classified a display of illuminated pixels on a touchscreen as sparse or dense. Correct responses were reinforced with six food pellets; incorrect responses were unreinforced. On some trials an uncertain response option was available. Pecking it was always reinforced with an intermediate number of pellets. Like monkeys and people in related experiments, the birds chose the uncertain response most often when the stimulus presented was difficult to classify correctly, but in other respects their behavior was not functionally similar to human behavior based on conscious uncertainty or to the behavior of monkeys in comparable experiments. Our data were well described by a signal detection model that assumed that the birds were maximizing perceived reward in a consistent way across all the experimental conditions.
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Jones, J. E., Antoniadis, E., Shettleworth, S. J., & Kamil, A. C. (2002). A comparative study of geometric rule learning by nutcrackers (Nucifraga columbiana), pigeons (Columba livia), and jackdaws (Corvus monedula). J Comp Psychol, 116(4), 350–356.
Abstract: Three avian species, a seed-caching corvid (Clark's nutcrackers; Nucifraga columbiana), a non-seed-caching corvid (jackdaws; Corvus monedula), and a non-seed-caching columbid (pigeons; Columba livia), were tested for ability to learn to find a goal halfway between 2 landmarks when distance between the landmarks varied during training. All 3 species learned, but jackdaws took much longer than either pigeons or nutcrackers. The nutcrackers searched more accurately than either pigeons or jackdaws. Both nutcrackers and pigeons showed good transfer to novel landmark arrays in which interlandmark distances were novel, but inconclusive results were obtained from jackdaws. Species differences in this spatial task appear quantitative rather than qualitative and are associated with differences in natural history rather than phylogeny.
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Shettleworth, S. J., & Westwood, R. P. (2002). Divided attention, memory, and spatial discrimination in food-storing and nonstoring birds, black-capped chickadees (Poecile atricapilla) and dark-eyed juncos (Junco hyemalis). J Exp Psychol Anim Behav Process, 28(3), 227–241.
Abstract: Food-storing birds, black-capped chickadees (Poecile atricapilla), and nonstoring birds, dark-eyed juncos (Junco hyemalis), matched color or location on a touch screen. Both species showed a divided attention effect for color but not for location (Experiment 1). Chickadees performed better on location than on color with retention intervals up to 40 s, but juncos did not (Experiment 2). Increasing sample-distractor distance improved performance similarly in both species. Multidimensional scaling revealed that both use a Euclidean metric of spatial similarity (Experiment 3). When choosing between the location and color of a remembered item, food storers choose location more than do nonstorers. These results explain this effect by differences in memory for location relative to color, not division of attention or spatial discrimination ability.
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Shettleworth, S. J., & Juergensen, M. R. (1980). Reinforcement and the organization of behavior in golden hamsters: brain stimulation reinforcement for seven action patterns. J Exp Psychol Anim Behav Process, 6(4), 352–375.
Abstract: Golden hamsters were reinforced with intracranial electrical stimulation of the lateral hypothalamus (ICS) for spending time engaging in one of seven topographically defined action patterns (APs). The stimulation used as reinforcer elicited hoarding and/or feeding and supported high rates of bar pressing. In Experiment 1, hamsters were reinforced successively for digging, open rearing, and face washing. Digging increased most in time spent, and face washing increased least. Experiments 2-5 examined these effects further and also showed that “scrabbling,” like digging, was performed a large proportion of the time, almost without interruption, for contingent ICS but that scratching the body with a hindleg and scent-marking showed relatively little effect of contingent ICS, the latter even in an environment that facilitated marking. In Experiment 6, naive hamsters received ICS not contingent on behavior every 30 sec (fixed-time 30-sec schedule). Terminal behaviors that developed on this schedule were APs that were easy to reinforce in the other experiments, but a facultative behavior, face washing, was one not so readily reinforced. Experiment 7 confirmed a novel prediction from Experiment 6--that wall rearing, a terminal AP, would be performed at a high level for contingent ICS. All together, the results point to both motivational factors and associative factors being involved in the considerable differences in performance among different reinforced activities.
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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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Mrosovsky, N., & Shettleworth, S. J. (1968). Wavelength preferences and brightness cues in the water finding behaviour of sea turtles. Behaviour, 32(4), 211–257.
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Plotnik, J. M., de Waal, F. B. M., & Reiss, D. (2006). Self-recognition in an Asian elephant. Proc. Natl. Acad. Sci. U.S.A., 103(45), 17053–17057.
Abstract: Considered an indicator of self-awareness, mirror self-recognition (MSR) has long seemed limited to humans and apes. In both phylogeny and human ontogeny, MSR is thought to correlate with higher forms of empathy and altruistic behavior. Apart from humans and apes, dolphins and elephants are also known for such capacities. After the recent discovery of MSR in dolphins (Tursiops truncatus), elephants thus were the next logical candidate species. We exposed three Asian elephants (Elephas maximus) to a large mirror to investigate their responses. Animals that possess MSR typically progress through four stages of behavior when facing a mirror: (i) social responses, (ii) physical inspection (e.g., looking behind the mirror), (iii) repetitive mirror-testing behavior, and (iv) realization of seeing themselves. Visible marks and invisible sham-marks were applied to the elephants' heads to test whether they would pass the litmus “mark test” for MSR in which an individual spontaneously uses a mirror to touch an otherwise imperceptible mark on its own body. Here, we report a successful MSR elephant study and report striking parallels in the progression of responses to mirrors among apes, dolphins, and elephants. These parallels suggest convergent cognitive evolution most likely related to complex sociality and cooperation.
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Broom, M. (2002). A unified model of dominance hierarchy formation and maintenance. J. Theor. Biol., 219(1), 63–72.
Abstract: In many different species it is common for animals to spend large portions of their lives in groups. Such groups need to divide available resources amongst the individuals they contain and this is often achieved by means of a dominance hierarchy. Sometimes hierarchies are stable over a long period of time and new individuals slot into pre-determined positions, but there are many situations where this is not so and a hierarchy is formed out of a group of individuals meeting for the first time. There are several different models both of the formation of such dominance hierarchies and of already existing hierarchies. These models often treat the two phases as entirely separate, whereas in reality, if there is a genuine formation phase to the hierarchy, behaviour in this phase will be governed by the rewards available, which in turn depends upon how the hierarchy operates once it has been formed. This paper describes a method of unifying models of these two distinct phases, assuming that the hierarchy formed is stable. In particular a framework is introduced which allows a variety of different models of each of the two parts to be used in conjunction with each other, thus enabling a wide range of situations to be modelled. Some examples are given to show how this works in practice.
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