The interaction of hunger and thirst in the rat

Two groups of rats were given a series of trials in an enclosed runway with a food reward at the end, one group being run hungry, the other hungry plus thirsty. Then each group was split into three sub-groups: one run hungry, the second thirsty and the other hungry plus thirsty, in each case without food reward.

It was found that, whereas on the rewarded runs the extra, “irrelevant,” thirst increased running speed, on unrewarded runs it had the opposite effect and slowed up performance. Thus on unrewarded runs the two sub-groups running thirsty, and hungry plus thirsty, ran as slowly as those running hungry. Differences were found not to depend on whether the animals had been hungry or hungry plus thirsty on previous rewarded runs.

The interaction of primary needs therefore depends on the external situation. This can be accounted for in terms of the Pavlovian theories of mutual induction and conditioning, but not in terms of Hull's theory of “irrelevant drives.”     

Schedules of reward shift in the double runway

Eighty food deprived rats received 62 trials in a double runway. On Trials 1-30, reward in the first goal box (GB1) was either always two food pellets or always zero pellets. All subjects received two pellets in the second goal box (GB2). On Trials 31-62 subjects in each preshift group (GB1 reward or GB1 nonreward) were shifted to the opposite GB1 reward level on 0, 25, 50, 75 or 100% of occasions. GB2 reward remained unaltered in all cases. For subjects experiencing reward decrease, second runway (A2) run and goal speeds after nonreward were generally enhanced, both within-group and in comparison with never rewarded controls. No such effect was evident on A2 start speed, nor was there any evidence to suggest that A2 performance after decreased reward was a function of the schedule of decrease. Increased GB1 reward resulted in general within-group impairment of A2 start and run speeds, with no effect on A2 goal performance. However, comparisons of speeds after increased reward with those of always rewarded controls revealed no difference on A2 start or run but indicated impairment of A2 goal performance. With the 50% schedule of reward increase, A2 run speeds after nonreward (the training level) exceeded those of never rewarded controls. Results are discussed with reference to McHose's contrast account of double runway phenomena and Amsel's frustration theory.     

Recall of the goal box in latent learning and latent discrimination

Two experiments showed that in a runway rats recalled incidental features of the goal box which they had previously found at the end of it. They were shown to modify their running speed according to the current associated reward value of the goal box. Retrieval cues for features of the goal box were intra- and extra-maze cues, not the response of running. Two goal boxes differing only in colour could be discriminatively recalled by virtue of their association with two discriminable runways. These experiments disconfound recall of the goal box from place learning as factors in latent learning.     

Double drive learning in rats without previous selective reinforcement

Can rats which have never been selectively satiated by food or water respond appropriately by choosing the relevant reward when they are made hungry or thirsty ? A group of 21 rats was put on a diet of wet mash and trained both hungry and thirsty in a T-maze with water on one side and food on the other. 8 out of 10 of the animals which were made thirsty on the test-run chose correctly in spite of an established preference for the other reward. Only 1 out of the 11 animals made hungry chose correctly. Possible explanations of this result are examined.     

Infantile handling and the extinction of responses acquired under food reward

A comparison was made of the runway behaviour of rats which had been handled from Days 1-21 and their non-handled litter mates. The training began on Day 70 after the animals were habituated to a restricted food schedule for 10 days. The subjects were given six trials each day in the runway and were rewarded with a 0.045 g. Noyes pellet. After 10 days of rewarded training trials, subjects were given 6 extinction trials a day for 10 days. Results showed that handled rats ran faster than non-handled rats during acquisition and during the first 3 days of extinction. The extinction data suggested that the relationship between emotionality and effects of frustrative non-reward should be re-evaluated.     

The non-occurrence of a stimulus as a signal

Two groups of rats were rewarded for pressing a panel following a varying number of bar presses; a signal following the bar press indicated whether or not a panel press would be rewarded if made before the next bar press. For one group the signal indicating reward was a flash of light, for the other it was the non-occurrence of the flash (in the latter case the flash thus signalled non-reward). The first group learned to withold panel presses except when reward was signalled, but the second group did not. This result is related to the “feature positive effect” of Jenkins and Sainsbury (1970), in which pigeons failed to withold pecks at a negative stimulus display when it was the same as the positive display except for the addition of a distinctive feature.     

Reward, novelty and spontaneous alternation

It has been suggested that rats' propensity for “win-shift” behaviour in spatial memory and spontaneous alternation tests reflects a species-specific foraging strategy which leads them to avoid places where they have recently found food. An alternative explanation is that they avoid places which are familiar. In three experiments using a T-maze, we evaluated these accounts by comparing the probability of avoiding or re-entering a recently visited arm, as a function of whether food had or had not been found on the previous visit. Each rat received a series of 16 exposure-test trial pairs over 8 days. Neither alternation nor repetition of the previous choice was differentially reinforced. Experiments I and II forced rats to enter a specific arm before a subsequent choice, and differed in the overall probability of reward; in Experiment III all choices were free. In all three experiments the probability of alternating was greater after nonreward than after reward. This effect occurred more reliably on later tests within a day, little difference appearing on earlier tests. It was concluded that there was no evidence for a spontaneous “win-shift” tendency as such, and that these and other results can be adequately accounted for by a combination of exploratory tendencies (spontaneous alternation) and the conventional effects of reward.     

D-Cycloserine enhances memory consolidation of hippocampus-dependent latent extinction

Adult male Long-Evans rats were trained to run in a straight-alley maze for food reward and subsequently received hippocampus-dependent latent extinction training. Immediately following latent extinction, rats received peripheral injections of the NMDA receptor partial agonist D-cycloserine (DCS, 15 mg/kg), or saline. Twenty-four hours later, rats received four extinction "probe" trials. Relative to saline controls, latencies to reach the goal box on probe trials were significantly higher in rats that had received DCS. These findings indicate that memory consolidation underlying hippocampus-dependent latent extinction, a cognitive form of learning in which the previously rewarded response is not made during extinction training, can be enhanced by NMDA-receptor agonism.     

Effects of preshift small magnitude of reward received during goalbox placements on the negative contrast effect

During preshift, one experimental group of rats was given a large magnitude of food reward following a traversal of a straight alley and during a goalbox placement, while the other experimental group was given a small reward during goalbox placement and a large reward following a run. During postshift, all experimental groups were given a small reward of food following a traversal down the runway and during a goalbox placement. A control group was maintained on small reward during placements and following a traversal throughout the study. Only the group who received preshift large reward during placement and following a runway response ran slower to small reward during postshift than the control group maintained on small reward (negative contrast effect).     

The role of response and reward in spatial memory

Two experiments were conducted to determine the role of the response and of reward in spatial working memory. Rats were initially trained on a four-arm maze to run to the end of each arm for a single pellet of food. On subsequent tests, rats were first placed at the end of one, two, or three arms. In Experiment 1, the arms on which the rat was placed (“placed arms”) had food which the rat was allowed to eat, whereas in Experiment 2 these placed arms did not have food. Following the placements the rat was allowed to choose among the four arms; only unplaced arms contained food. Two measures indicated that the response made a slight but reliable contribution to spatial memory. (a) When a rewarded arm was still available, choice accuracy after placements was less than choice accuracy on tests in which no placements had occurred; this difference diminished over test days. (b) When all four arms had been chosen once, the rats were more likely to go back to a placed arm rather than an unplaced arm. No influence of the presence or absence of food on the placed arms was found. These data demonstrate that the response of running down an arm, but not the reward outcome at the end, had a small influence on the memorability of a visit. Overall, above chance performance in the spatial working memory task was maintained without either running to the arm or obtaining food on it.     

Memory for feeding in rats' spatial and visual choice behaviour

Four experiments with mazes examined the effects of feeding upon rats' subsequent choice between spatially or visually distinct alternatives. It is concluded that rats can recall both whether any food remained at the cessation of feeding (nondepletion) and whether or not feeding was interrupted; both features can be associated with the place where they occurred. The memory of nondepleted food evokes an unlearned tendency to return to the place where it was present (“win-stay” behaviour). The memory of interruption transiently had a similar effect in naive animals but eventually exerts a more nonspecific influence, which facilitates not only win-stay learning but its opposite, win-shift. The nondepletion effect was also obtained when the alternatives were defined by a visual cue (brightness) rather than spatial location. The determinants of staying and shifting are discussed in terms of reward memory and exploration.     

Effects of expectancies of different reward magnitudes in transfer from noncontingent pairings to instrumental performance

In two experiments, rats received noncontingent pairings of two stimuli with food reward, one paired with small reward and the other with large reward, and received bar press training with large reward or with small reward. When the noncontingent stimuli (NS) were presented for test during subsequent rewarded bar pressing and during early extinction of bar pressing, responding for each group was faster in the presence of the NS which was paired with the same reward magnitude that group received in bar press training than to the NS which had been paired with a different reward magnitude. As extinction progressed, all groups responded more slowly in the presence of the NS which had been paired with the large reward than in the presence of the NS which had been paired with small reward. These results were interpreted as indicating that responding in the presence of an NS depends on: (i) whether the reward expectancy elicited by the NS has been conditioned to the instrumental response, and (ii) the relationship between the reward expected in the presence of the NS and that received in test.     

Pavlovian processes in the motivational control of instrumental performance

Hungry rats were rewarded for pressing a lever on a multiple schedule. During one component the reward was a sucrose solution, whereas food pellets acted as the reward during another component. Lever pressing was never rewarded during the third component. When the drive state was switched from hunger to thirst and the animals tested in extinction, they pressed more in the presence of the component stimulus that had been associated with the sucrose reward during training. A similar effect was observed during the extinction test of a second study in which the component stimuli had signalled non-contingent presentations of either the sucrose or pellet rewards in the absence of the lever. This suggests that the instrumental irrelevant incentive effect observed in the first experiment was due, at least in part, to the Pavlovian relationship between the component stimuli and the reinforcers during training. In fact, when the size of the effects controlled by purely Pavlovian and supposedly instrumental contingencies was compared directly in the final study, no difference could be detected.     

Ordinal Position Learning and Remote Anticipation

Rats were runway trained on each of two 3-trial series of reward outcomes. The series are labeled XNY and ZNN, for which X represents a trial that was rewarded with Noyes pellets and N represents a trial that ended with no reward. Units of distinctively flavored breakfast cereals served as reward on trials labeled Y and Z. One group (Floor) had each series occur with a correlated runway floor, either smooth and black or rough and white. For a second group (Memory), the floor cue was uncorrelated with series. Animals in both groups learned to approach the goal rapidly on the 1st trials of the 2 series and slowly on the 2nd trials, but only Group Floor learned to differentiate the 3rd trials of the series. These results recommend a view of serial learning that emphasizes the role played by information about the ordinal position of series items.     

Resistance to satiation: Reinforcing effects of food and eating under satiation

It has been shown previously that rats which have learned a response when hungry will continue to make that response when tested satiated, a phenomenon labeled resistance to satiation. Here we showed that rats which were previously trained hungry will learn a new response for the opportunity to consume pellets in a new situation when tested satiated. In four experiments various groups received each of the components of the training given when rats learn an instrumental response when hungry. Rats were placed in the goalbox of a straight alley and given food pellets when hungry or were hungry only in their home cages prior to running a straight alley in the satiated test in Experiment 1. In Experiments 2, 3, and 4 learning of a differential conditioning problem for pellets in S+ (nonreward in S?) was measured in the satiated test. Groups given pellets in their home cages when hungry with or without alley exposure learned to run more rapidly in S+ than in S? in the satiated test phase. The tendency to eat pellets in the apparatus and the reinforcing effect of eating the pellets was larger for rats which ate the pellets when hungry in their home cage than for rats which ate the pellets when satiated in their home cage. Being hungry in the home cage with no pellets was not sufficient to produce eating or running for pellets in the satiated test, indicating that any inherent reinforcing effect of the pellets is not sufficient to produce eating or running, and that incomplete satiation cannot account for the learning. These data indicate that a reinforcing effect of eating pellets under satiation is an important determiner of resistance to satiation.     

Ordinal position learning and remote anticipation

Rats were runway trained on each of two 3-trial series of reward outcomes. The series are labeled XNY and ZNN, for which X represents a trial that was rewarded with Noyes pellets and N represents a trial that ended with no reward. Units of distinctively flavored breakfast cereals served as reward on trials labeled Y and Z. One group (Floor) had each series occur with a correlated runway floor, either smooth and black or rough and white. For a second group (Memory), the floor cue was uncorrelated with series. Animals in both groups learned to approach the goal rapidly on the 1st trials of the 2 series and slowly on the 2nd trials, but only Group Floor learned to differentiate the 3rd trials of the series. These results recommend a view of serial learning that emphasizes the role played by information about the ordinal position of series items.     

Maintenance-feeding effort affects instrumental performance

The effort required of rats to obtain food, employing standard maintenance-feeding procedures, can affect the effort subsequently expended in an instrumental-learning task. Rats received 43 food-rewarded runway trials followed by access to food (a) on the home-cage floor, or (b) from a hopper attached to the home-cage's wiremesh front wall. Hopper feeding involved the greater effort since the rats had to gnaw the food pellets through the wire mesh. Following 9 or 27 days of maintenance feeding, the rats were returned to the runway for one rewarded trial and 16 extinction trials. The hopper-fed rats ran faster than floor-fed rats on the rewarded test trial. In extinction, long-term maintenance feeding produced faster subsequent running times by the hopper-fed rats than the floor-fed rats. Short-term maintenance feeding, on the other hand, produced slower running times by the hopper-fed rats than the floor-fed rats. The results are consistent with previous findings of transfer of effort across behaviours and suggest that the effort required for reinforcement in the maintenance environment can systematically influence the effectiveness of various experimental treatments.     

Transfer of persistence to the acquisition of a new behaviour

Two experiments tested whether the degree of effort required for the reinforcement of one behaviour would affect the acquisition of a second behaviour. In the first experiment, rats were placed in a conditioning chamber and: (a) were required to press a lever for food pellets on a fixed ratio schedule, (b) received free presentation of the pellets, or (c) did not receive pellets. Next, all rats were rewarded for a new behaviour, round trips across the length of a runway. As predicted, the fixed-ratio group had the greatest shuttle rate. In the second experiment, two groups were required to press a lever, and the number of presses per pellet was varied. For two other groups not required to press the lever, the amount of food presented per approach to the feeder was varied. The greater required number of lever presses and the lesser number of pellets per approach to the feeder produced the higher subsequent shuttle rates. Two alternative explanations were compared: the degree of accustomed effort per reinforcer becomes a learned component of behaviour, or high effort increases the habituation of frustration-produced disruptive responses.     

Elasmobranch cognitive ability: using electroreceptive foraging behaviour to demonstrate learning, habituation and memory in a benthic shark

Top predators inhabiting a dynamic environment, such as coastal waters, should theoretically possess sufficient cognitive ability to allow successful foraging despite unpredictable sensory stimuli. The cognition-related hunting abilities of marine mammals have been widely demonstrated. Having been historically underestimated, teleost cognitive abilities have also now been significantly demonstrated. Conversely, the abilities of elasmobranchs have received little attention, despite many species possessing relatively large brains comparable to some mammals. The need to determine what, if any, cognitive ability these globally distributed, apex predators are endowed with has been highlighted recently by questions arising from environmental assessments, specifically whether they are able to learn to distinguish between anthropogenic electric fields and prey bioelectric fields. We therefore used electroreceptive foraging behaviour in a model species, Scyliorhinus canicula (small-spotted catshark), to determine cognitive ability by analysing whether elasmobranchs are able to learn to improve foraging efficiency and remember learned behavioural adaptations. Positive reinforcement, operant conditioning was used to study catshark foraging behaviour towards artificial, prey-type electric fields (Efields). Catsharks rewarded with food for responding to Efields throughout experimental weeks were compared with catsharks that were not rewarded for responding in order to assess behavioural adaptation via learning ability. Experiments were repeated after a 3-week interval with previously rewarded catsharks this time receiving no reward and vice versa to assess memory ability. Positive reinforcement markedly and rapidly altered catshark foraging behaviour. Rewarded catsharks exhibited significantly more interest in the electrical stimulus than unrewarded catsharks. Furthermore, they improved their foraging efficiency over time by learning to locate and bite the electrodes to gain food more quickly. In contrast, unrewarded catsharks showed some habituation, whereby their responses to the electrodes abated and eventually entirely ceased, though they generally showed no changes in most foraging parameters. Behavioural adaptations were not retained after the interval suggesting learned behaviour was not memorised beyond the interval. Sequences of individual catshark search paths clearly illustrated learning and habituation behavioural adaptation. This study demonstrated learning and habituation occurring after few foraging events and a memory window of between 12 h and 3 weeks. These cognitive abilities are discussed in relation to diet, habitat, ecology and anthropogenic Efield sources.     

The order of continuous,partial and nonreward trials and resistance to extinction

In the first experiment rats were given either 16 or 48 nonrewarded or continuously rewarded trials prior to 24 continuously or partially rewarded trials, followed by extinction. Increased resistance to extinction was found for increasing numbers of nonrewarded trials when they were followed by partial reward, but not when followed by continuous reward. Similarly, more continuously rewarded trials followed by partial reward tended to increase resistance to extinction. Because of the theoretical importance of the effect of continuous reward followed by partial, a second experiment was performed where the range of the number of continuously rewarded trials was extended to 0, 48, and 96. Contrary to many theoretical predictions, resistance to extinction increased as a function of increasing amounts of continuous reward.