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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., & 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., & 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., & 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., & 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. (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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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. (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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Sharon, L., & Crowell-Davis, S. L. (2007). Sexual behavior of mares. Horm Behav, 52(1), 12–17.
Abstract: The mare is seasonally polyestrus, having an anovulatory period during the short light days of late fall and early winter, and beginning to ovulate as the days become longer during the winter. The complete estrus cycle is typically about 3 weeks, with 5 to 7 days of estrus and approximately 2 weeks of diestrus. When a mare lives within the natural social structure of the horse, i.e. a family band with several adult mares and one or more stallions, estrus is characterized by repeatedly approaching the stallion, frequent urination, deviating the tail away from the perineum, and standing still with the hind limbs spread apart. Diestrus is characterized by avoidance of an approaching stallion, and aggression toward the stallion, such as squealing, striking, and kicking, if he persists in attempting to court the diestrus mare. However, mares and stallions with long-term social relationships will often rest together, graze together and groom each other, all without sexual interactions. Hormonally, estrous behavior in the mare is initiated by estradiol that is secreted by the follicle, while estrous behavior is suppressed by progesterone, secreted by the corpus luteum. Mares are unusual among the ungulates in that they periodically exhibit estrous behavior during the anovulatory period. This is probably due to the release of estrogenic steroids secreted by the adrenal cortex. The display of sexual behavior by the mare throughout the year is thought to facilitate maintenance of the horse's social structure, in which the male remains with a group of females year round, in contrast with most ungulates in which the females and males only come together during the mating season.
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Schmidt, A., Möstl, E., Wehnert, C., Aurich, J., Müller, J., & Aurich, C. (2010). Cortisol release and heart rate variability in horses during road transport. Horm. Behav., 57(2), 209–215.
Abstract: Based on plasma cortisol concentrations it is widely accepted that transport is stressful to horses. So far, cortisol release during transport has not been evaluated in depth by non-invasive techniques such as analysis of salivary cortisol and faecal cortisol metabolites. Transport also causes changes in heart rate and heart rate variability (HRV). In this study, salivary cortisol, faecal cortisol metabolites, heart rate and HRV in horses transported by road for short (one and 3.5 h) and medium duration (8 h) were determined. With the onset of transport, salivary cortisol increased immediately but highest concentrations were measured towards the end of transport (4.1 ± 1.6, 4.5 ± 2.6, 6.5 ± 1.8 ng/ml in horses transported for one, 3.5 and 8 h, respectively). Faecal cortisol metabolite concentrations did not change during transport, but 1 day after transport for 3.5 and 8 h had increased significantly (p < 0.01), reflecting intestinal passage time. Compared to salivary cortisol, changes in faecal cortisol metabolites were less pronounced. Heart rate increased and beat-to-beat (RR) interval decreased (p < 0.05) with the onset of transport. Standard deviation of heart rate increased while root mean square of successive RR differences (RMSSD) decreased in horses transported for 3.5 (from 74 ± 5 to 45 ± 6 ms) and 8 h (from 89.7 ± 7 to 59 ± 7 ms), indicating a reduction in vagal tone. In conclusion, transport of horses over short and medium distances leads to increased cortisol release and changes in heart rate and HRV indicative of stress. The degree of these changes is related to the duration of transport. Salivary cortisol is a sensitive parameter to detect transient changes in cortisol release.
Keywords: Horse; Transport; Cortisol; Heart rate variability
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