Finding a goal on dry land and in the water: differential effects of disorientation on spatial learning.

Two previous studies, Martin et al. (J. Exp. Psychol. Anim. Behav. Process. 23 (1997) 183) and Dudchenko et al. (J. Exp. Psychol. Anim. Behav. Process. 23 (1997) 194), report that, compared to non-disoriented controls, rats disoriented before testing were disrupted in their ability to learn the location of a goal on a dry radial-arm maze task, but that both groups learned at the same rate in the Morris water maze. However, the radial-arm maze task was much more difficult than the water maze. In the current set of experiments, we examined the performance of control and disoriented rats on more comparable dry land and water maze tasks. Compared to non-disoriented rats, rats that were disoriented before testing were significantly impaired in locating a goal in a circular dry arena, but not a water tank. The results constrain theoretical explanations for the differential effects of disorientation on different spatial tasks.

Hippocampus and memory in a food-storing and in a nonstoring bird species.

Food-storing birds maintain in memory a large and constantly changing catalog of the locations of stored food. The hippocampus of food-storing black-capped chickadees (Parus atricapillus) is proportionally larger than that of nonstoring dark-eyed juncos (Junco hyemalis). Chickadees perform better than do juncos in an operant test of spatial non-matching-to-sample (SNMTS), and chickadees are more resistant to interference in this paradigm. Hippocampal lesions attenuate performance in SNMTS and increase interference. In tests of continuous spatial alternation (CSA), juncos perform better than chickadees. CSA performance also declines following hippocampal lesions. By itself, sensitivity of a given task to hippocampal damage does not predict the direction of memory differences between storing and nonstoring species.

Hippocampal lesions impair memory for location but not color in passerine birds.

The effects of hippocampal complex lesions on memory for location and color were assessed in black-capped chickadees (Parus atricapillus) and dark-eyed juncos (Junco hyemalis) in operant tests of matching to sample. Before surgery, most birds were more accurate on tests of memory for location than on tests of memory for color. Damage to the hippocampal complex caused a decline in memory for location, whereas memory for color was not affected in the same birds. This dissociation indicates that the avian hippocampus plays an important role in spatial cognition and suggests that this brain structure may play no role in working memory generally.

Pigeons do not complete partly occluded figures.

One of the most common obstacles to object perception is the fact that objects often occlude parts of themselves and parts of other objects. Perceptual completion has been studied extensively in humans, and researchers have shown that humans do complete partly occluded objects. In an effort to understand more about the mechanisms underlying completion, recent research has extended the study of perceptual completion to other mammalian species. Monkeys and mice also seem to complete two-dimensional representations of partly occluded objects. The present study addresses the question of whether this capacity generalizes to a nonmammalian species, the pigeon (Columba livia). The results point to a limit of the generalizability of perceptual completion: pigeons do not complete partly occluded figures.

Effects of photoperiod on food-storing and the hippocampus in birds.

Birds that store food have a relatively large hippocampus compared to non-storing species. The hippocampus shows seasonal differences in neurogenesis and volume in black-capped chikadees (Parus atricapillus) taken from the wild at different times of year. We compared hippocampal volumes in black-capped chickadees captured at the same time but differing in food-storing behaviour because of manipulations of photoperiod in the laboratory. Differences in food-storing behaviour were not accompanied by differences in the volume of the hippocampus. Hippocampal volumes also did not differ between two groups of a non-food-storing control species, house sparrows (Passer domesticus), exposed to the same conditions as the chickadees.

Hippocampal volume and food-storing behavior are related in parids.

The size of the hippocampus has been previously shown to reflect species differences and sex differences in reliance on spatial memory to locate ecologically important resources, such as food and mates. Black-capped chickadees (Parus atricapillus) cached more food than did either Mexican chickadees (P. sclateri) or bridled titmice (P. wollweberi) in two tests of food storing, one conducted in an aviary and another in smaller home cages. Black-capped chickadees were also found to have a larger hippocampus, relative to the size of the telencephalon, than the other two species. Differences in the frequency of food storing behavior among the three species have probably produced differences in the use of hippocampus-dependent memory and spatial information processing to recover stored food, resulting in graded selection for size of the hippocampus.

Commentary: What are behavior systems and what use are they?

The contributions of this symposium on behavior systems are summarized and evaluated by considering two questions: (1) What is a behavior system? (2) What use to the learning theorist are behavior systems? Two examples of behavior systems from the classical ethological literature are compared with the behavior systems discussed in the symposium, and some similarities and differences in the type of analyses used are discussed. Analysis of the pre-organized species-typical behavior systems relevant to the unconditioned stimuli or reinforcers in learning experiments can contribute performance rules and better understanding of the conditions and contents of learning. The organization of behavior systems can also provide important clues to the neural circuitry underlying behavior, and a behavior systems approach can raise novel questions concerning learning and behavioral development. Possible future directions for the behavior systems approach are briefly discussed.

Varieties of learning and memory in animals.

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.

How pigeons estimate rates of prey encounter.

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.

Detection of cryptic prey: search image or search rate?

Animals' improvement in capturing cryptic prey with experience has long been attributed to a perceptual mechanism, the specific search image. Detection could also be improved by adjusting rate of search. In a series of studies using both naturalistic and operant search tasks, pigeons searched for wheat, dyed to produce 1 conspicuous and 2 equally cryptic prey types. Contrary to the predictions of the search-rate hypothesis, pigeons given a choice between the 2 cryptic types took the type experienced most recently. However, experience with 1 cryptic type improved accuracy on the other cryptic type, a result inconsistent with a search image specific to 1 prey type. Search image may better be thought of as priming of attention to those features of the prey type that best distinguish the prey from the background.

The role of shifting in choice behavior of pigeons on a two-armed bandit.

Pigeons (Columba livia) were confronted with a problem in choice known as the two-armed bandit (TAB): two concurrent discrete trials random-ratio schedules in which the good side changes randomly from day to day. In the first experiment a substantial proportion of the birds' choices were contrary to those predicted by immediate maximization. Under a variety of parameter settings the pigeons chose the bad side more often than expected by immediate maximization. The hypothesis was advanced that shifting, that is, the tendency to avoid the most recently visited location, was responsible for limiting the number of choices on the good side of the TAB. Experiments 2-4 examined this hypothesis. The first of these experiments compared the accuracy on win-shift vs win-stay trials, and lose-shift vs lose-stay. When the response keys were at opposite sides of the operant chamber (keys-far), as in the first TAB experiment, there was a significant tendency to shift. This tendency was not present when the keys were close together (keys-close), in Experiment 3. Experiment 4 compared TAB performance in the original keys-far situation with that in the keys-close chamber. The proportion of choices on the good side of the TAB increased faster and attained a higher level for the keys-close group, and the proportion of choices which were shifts from one side to the other was smaller. A higher proportion of individual choices in the keys-close condition could be accounted for by immediate maximization.

Individual differences in choice of food items by pigeons.

Pigeons (Columba livia ) select food items idiosyncratically when feeding on grains (Brown, 1969; Moon & Zeigler, 1979; Giraldeau & Lefebvre, 1985). In three experiments pigeons under various conditions of food restriction were offered artificial "grains", pellets of pigeon food that differed only in size, to see whether individual differences in preference would still be observed. When 300-mg ("large") and 20-mg ("small") pellets were available simultaneously there were still wide individual differences in choice, but when encountering the same items successively pigeons took nearly all the items offered.

Handling time and choice in pigeons.

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.

How marsh tits find their hoards: the roles of site preference and spatial memory.

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.

Reinforcement and the organization of behavior in golden hamsters: brain stimulation reinforcement for seven action patterns.

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.

Reinforcement and the organization of behavior in golden hamsters: Pavlovian conditioning with food and shock unconditioned stimuli.

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.

Behavioral adaptation to fixed-interval and fixed-time food delivery in golden hamsters.

Food-deprived golden hamsters in a large enclosure received food every 30 sec contingent on lever pressing, or free while their behavior was continuously recorded in terms of an exhaustive classification of motor patterns. As with other species in other situations, behavior became organized into two main classes. One (terminal behaviors) increased in probability throughout interfood intervals; the other (interim behaviors) peaked earlier in interfood intervals. Which class an activity belonged to was independent of whether food was contingent on lever pressing. When food was omitted on some of the intervals (thwarting), the terminal activities began sooner in the next interval, and different interim activities changed in different ways. The interim activities did not appear to be schedule-induced in the usual sense. Rather, the hamsters left the area of the feeder when food was not due and engaged in activities they would normally perform in the experimental environment.