Infantile stimulation and its effects on frustration- and fear-motivated behavior in rats

Infant rats that were either removed from the nest each day (handled) or left undisturbed (nonhandled) were, in adulthood, given 72 food-reinforced runway acquisition trials followed by 24 trials of extinction training with or without shock. Handled and nonhandled control animals were given runway training without food reinforcement. Reinforced rats ran faster than nonreinforced rats, and handled rats ran faster than nonhandled rats during the initial trials of runway acquisition irrespective of the reinforcement condition. Nonhadled rats stopped running sooner than handled rats when shock was introduced in the goalbox, but differences between handled and nonhandled rats given extinction training without shock were small. Results of a second experiment showed no differences between handled and non-handled rats in the magnitude of the depression effect after an incentive shift. It was concluded that infantile handling had little effect on frustration-motivated behavior, but did affect fear-motivated behavior.     

The effects of early handling on the partial reinforcement extinction effect and the partial punishment effect in male and female rats

Handled (Day 1-22) and non-handled infantile Wistar rats were tested in maturity for the partial reinforcement extinction effect (PREE) and the partial punishment effect (PPE). In Experiments 1 and 2, mature male and female rats were trained to run in an alley for food reward on a 1-trial/day schedule. In the PREE paradigm (Experiment 1), the partially reinforced group (PRF) received reinforcement on a quasi-random 50% schedule, while the continuously reinforced group (CRF) received reinforcement on every trial. In the test stage, both groups were given extinction training. In the PPE paradigm (Experiment 2), the partially punished (PP) group received, together with continuous reinforcement, shocks on a quasirandom 50% schedule, while the continuously reinforced group was reinforced on every trial. In test, all animals were given both reinforcement and shock on every trial. In Experiment 1, PREE—i.e. increased resistance to extinction in the PRF as compared to the CRF group—was more pronounced in the handled animals. More specifically, no PREE was obtained in the non-handled males, and in the non-handled females the PREE was reduced compared to the handled females. The results of Experiment 2 revealed no effect of handling or sex on PPE, that is, increased resistance to punishment in the PP group as compared to the CRF group was evident in all four conditions. In Experiment 3, handled and non-handled male rats were tested for the PREE using a multi-trial procedure in an operant chamber. PREE was obtained in the handled but not in the non-handled animals. The implications of these results for the differential effects of handling on male and female rats and the distinction between the PREE and PPE paradigms are discussed.     

Infantile handling and the facilitation of discrimination and reversal learning

One group of rats was removed from their home cage and received daily handling from Days 3-21, while the control group of litter mates remained in the cage and did not receive any treatment. On Day 22 all rats were weaned and they were housed in individual cages until they were 60 days old. After two pretraining days, subjects were given daily blocks of one free and three forced trials in the T-maze and were rewarded with food after making the correct response. Following 20 days of training and testing on black-white discrimination, subjects were given 10 days of reversal training with four daily trials. Results indicate that the handled animals showed faster running and a greater number of correct choices than the control rats during both the acquisition and reversal learning phases of the experiment.     

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.     

Response persistence in the rat following intermittent punishment and partial reward: Effects of trial sequence

Thirty-six rats were given 16 days of partial reward training in a runway. During the final 12 days each of the animals received one foot-shock experience each day. One group received shock on an N trial preceding an R trial (P-R), a second group was shocked on N trials not followed by an R trial (R-P), and the third group received shock after completing all daily trials (Control). Following acquisition the rats were split within each group (one half received 24 trials of unpunished extinction and one half continued to receive partial reward but were punished on every trial). During consistent punishment the P-R animals were more persistent than the R-P or Control rats and during unpunished extinction the P-R and Control animals were equal in persistence but both were superior to the R-P animals. The results were discussed in terms of Capaldi's sequential trial theory.     

Effect of interpolated extinction after partial delay of reward training on subsequent reacquisition and extinction

Four groups of albino rats were run four trials a day in a straight runway for 44 days. On the first 15 days, two groups were given continuous immediate reward (IR) and two groups a 50 per cent, schedule of 30-sec. partial delay of reward (PDR). On the next 15 days, one IR group and one PDR group were extinguished, while the other IR and PDR groups remained on their original schedules. In the third phase, all groups received 8 days of training on IR. Finally, all groups were given 6 days of extinction training. In the first extinction, PDR produced greater resistance to extinction than IR. In the second extinction period, the PDR group which had previously been given extinction and the two IR groups extinguished relatively rapidly and at approximately the same rates, while the PDR group which had not been extinguished was significantly more resistant to extinction than the other three groups.     

Reward schedule effects in extinction: Intertrial interval, memory, and memory retrieval

Rats were trained in a runway such that partial reward occurred on Trial 1 of the day and consistent reward on subsequent massed trials (Group PRT1), or consistent reward occurred on Trial 1 of the day and partial reward on subsequent massed trials (Group PRTM). Under spaced (24-hr) extinction, Group PRT1 was more resistant to extinction than Group PRTM and under massed (1-min) extinction, Group PRTM was more resistant to extinction than Group PRT1. These findings suggest that (a) distinctive stimuli are associated with Trial 1 of the day and with subsequent massed trials, (b) these distinctive stimuli function as retrieval cues for memories, memory retrieval being independent of intertrial interval, and (c) behavior in extinction is controlled by a stimulus compound consisting of the memory of nonreward plus stimuli which accompany the memory of nonreward on rewarded acquisition trials.     

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.     

Dependence of the overtraining extinction effect on attention to runway cues

Six groups of rats were given 60 extinction trials in a runway following 30, 90 or 120 acquisition trials. For half the groups, a hurdle was interposed in the runway during both acquisition and extinction. For rats tested with the hurdle, the 30-trial group extinguished more readily than the other two groups. With the hurdle absent, overtrained rats extinguished more readily. This was interpreted as supporting the hypothesis that overtraining transfers control of responding from external to internal cues, and that internally controlled running is relatively more susceptible to extinction.     

The partial punishment effect following minimal acquisition training: Sodium amobarbital and the stimulus properties of early punished trials

In a runway investigation, six groups of rats received limited runway training such that partial punishment, partial reinforcement, or continuous reinforcement was accompanied by sodium amobarbital or saline. Following an interpolated phase of continuous reinforcement without injections, all groups were given punished extinction. The entire experiment was conducted under widely spaced conditions (ITI = 24 hr). It was found that partial punishment increased resistance to punished extinction relative to partially and continuously reinforced controls when acquisition was given under saline. When partial punishment training was accompanied by amobarbital this effect was eliminated. The drug was observed to have no effect on the punished extinction performance of the partial reinforcement and continuous groups, respectively. Moreover, the partial reinforcement effect (PRE) did not generalize to punished extinction. These data provide information concerning the difference between the stimuli associated with the early trials of punishment and nonreward and indicate that the former but not the latter contain emotional elements.     

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.     

Punishment persistence and sequence: Some effects of P-length and N-length in the rat

In a runway investigation, two groups of rats received partial punishment training with P-lengths of 1, 2, and 3 (Group P123) and P-lengths of 1 (Group P1), respectively. Two additional groups received partial reinforcement with N-lengths of 1, 2, and 3 (Group N123) and N-length of 1 (Group N1). An additional group was given unpunished continuous reinforcement in the runway, but received control shocks in a separate apparatus. Following training all subjects received punished extinction (shock plus nonreward). The results indicated that P-length increased resistance to punished extinction, however N-length did not have the corresponding effect. In addition, partial reinforcement did not increase resistance to punished extinction relative to continuous reinforcement. These results were interpreted within a sequential-theoretical framework.     

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.     

A test for order relevance in a three-element serial learning task

Rats were trained in a straight runway on a simple three-element series of differing reward quantities. The first trial of the series ended with a two-pellet reward, and the second and third trials ended with a 12-pellet reward and a nonreward, respectively (2-12-0). All animals developed accurate anticipation of the terminal nonreward in the 2-12-0 series before it was rearranged for two test days during which the elements appeared in the order 12-2-0. The rats' anticipation of the terminal nonreward did not transfer to the reordered series, a result taken to mean that anticipation had been based upon interitem associations among memorial representations of the differing reward quantities. A second transfer test to 0-0-0, given after anticipation was reestablished to the 2-12-0 series, gave evidence that the ordinal position of the series elements was also a source of anticipation; the animals continued to run relatively slowly on the third trial in extinction. In neither transfer test was there evidence that the rats employed the strategy of enumerating rewarded trials, because counting is an order-irrelevant process.     

Effects of anxiolytic drugs on the partial reinforcement extinction effect in runway and skinner box

The partial reinforcement extinction effect (PREE) in the runway is reduced by anxiolytics in a non-state-dependent manner when a 24 h inter-trial interval is used, but there is some doubt as to the nature of the drug effects when shorter intervals are used. Experiment 1 repeated a study by Gray (1969), in which ambiguous results were obtained using eight trials/day. It demonstrated that the anxiolytic barbiturate, sodium amylobarbitone, given both in acquisition and extinction does not reduce the PREE. It confirmed Gray's observation that the PREE is abolished if the drug is given in acquisition but not in extinction. This suggests that a 24 h inter-trial interval is one critical factor in non-state-dependent reduction of the PREE. Experiments 2 and 3 tested the effects of the anxiolytic benzodiazepine, chlordiazepoxide, on the PREE with a 24 h inter-trial interval in the Skinner box. The basic task was a single FR5 sequence terminating in delivery of 10 (Experiment 2) or 20 (Experiment 3) reward pellets each day. There were 10 acquisition trials and partially reinforced rats received either three (Experiment 2) or four (Experiment 3) non-rewarded trials. The drug abolished the PREE in Experiment 2 and effectively reversed it in Experiment 3. These results confirm previous work with this drug in the runway (Feldon and Gray, 1981) and extend them to a very different experimental situation. These results support the idea that the PREE depends on different processes with different acquisition parameters; and that when a 24 h inter-trial interval is used the PREE is largely produced by some general process, probably the counterconditioning of conditioned frustration, which is sensitive to anxiolytic drugs. They also demonstrate very clearly the paradoxical effects of the anxiolytic drugs when given in both acquisition and extinction: they generally increase resistance to extinction in continuously reinforced animals, but block the increase in resistance (the PREE) produced by behavioural schedules.     

Extinction of a running response as a function of reinforcement magnitude

Four groups of rats were trained on different sucrose solutions in a straight runway. Terminal running speed was a monotonic function of reinforcement magnitude. After training each group was subdivided, one subgroup being extinguished under spaced, the other under massed conditions. In spaced extinction the animals trained on non extreme reward magnitudes showed most resistance to extinction. It was concluded that resistance to extinction is an inverted U-shaped function of reinforcement magnitude found in training. The massed extinction trials were conducted with a very short inter-trial interval. The animals showed an immediate drop in running speed followed by a gradual recovery and a subsequent decline. The number of trials taken to reach the peak recovery speed was a function of reinforcement magnitude found in training. Results on both massed and spaced extinction trials were interpreted in terms of the facilitatory and inhibitory effects of momentary and conditioned frustration.     

Effect of neonatal handling on adult rat spatial learning and memory following acute stress

Brief neonatal handling permanently alters hypothalamic-pituitary-adrenal axis function resulting in increased ability to cope with stress. Since stress is known to affect cognitive abilities, in the present study we investigated the effect of brief (15 min) handling on learning and memory in the Morris water maze, following exposure to an acute restraint stress either before training or recall. Exposure of non-handled rats to the acute stress prior to training resulted in quicker learning of the task, than in the absence of the stressor. When acute stress preceded acquisition, male handled rats showed an overall better learning performance, and both sexes of handled animals were less impaired in the subsequent memory trial, compared to the respective non-handled. In addition, the number of neurons immunoreactive for GR was higher in all areas of Ammon's horn of the handled rats during the recall. In contrast, the number of neurons immunoreactive for MR was higher in the CA1 and CA2 areas of the non-handled males. When the acute restraint stress was applied prior to the memory test, neonatal handling was not effective in preventing mnemonic impairment, as all animal groups showed a similar deficit in recall. In this case, no difference between handled and non-handled rats was observed in the number of GR positive neurons in the CA2 and CA3 hippocampal areas during the memory test. These results indicate that early experience interacts with sex and acute stress exposure in adulthood to affect performance in the water maze. Hippocampal corticosterone receptors may play a role in determining the final outcome.     

Spontaneous alternation and pattern of reinforcement

In Experiment I, rats were given one pair of trials per day in a T-maze; group RN were rewarded on the first trial only, NR were rewarded on the second trial only, while RR were rewarded on both trials. Alternation was approximately the same for all groups, and there was little decline in alternation over 28 days. There was, however, a difference in the choice behaviour on trials when rats failed to alternate, NR and RR, but not RN, choosing according to a position preference. These results are incompatible with several earlier theories of spontaneous alternation, and the hypothesis was put forward that alternation is initially part of an exploratory tendency, but that, when the maze becomes explored, the exploratory tendency dies down, and alternation only continues when rats have been reinforced (primarily or secondarily) for alternating. This hypothesis was tested in a second experiment, similar to the first but in which one group of rats was not rewarded on either trial; as predicted, this group showed significantly more decline in alternation than rewarded rats.     

Effect of effort and latent extinction on resistance to extinction

Sixty-four male Wistar rats were given acquisition training in an enclosed straight-alley runway which could be adjusted for angles of inclination. The 2×2×2 design involved two angles of inclination in acquisition (0° and 40°) and two angles of inclination in extinction (0° and 40°). Between acquisition and extinction, half the subjects were exposed to a latent extinction procedure and half served as controls. Number of responses in a 30-min extinction session was an inverse function of effort required in extinction. Additionally, latent extinction procedures resulted in reduced resistance to extinction, but only when the effort conditions of acquisition and extinction were constant. When the effort conditions of acquisition and extinction were dissimilar, latent extinction procedures resulted in increased resistance to extinction. The results raise questions about the nature of the learning which occurs during latent extinction training.     

The operation of an acquired drive in satiated rats

A series of four short experiments indicates that the behaviour of satiated rats in a runway, at the end of which they have previously been rewarded, differs significantly from the behaviour of satiated rats without previous reward in the experimental situation. The former group reach the end box more quickly after having been put in the starting box of the runway and if provided with food in the end box proceed to eat it, although they have just refused similar pieces of food in their home cages. This is shown not to be due to defective satiation or the operation of fear in the control group. When runs and feeding in the end box are separated during the training period, the previously rewarded group still shows a more vigorous response on satiated trials, indicating that it is a reward expectancy about the goal box rather than a running habit which has become “functionally autonomous”, acting as a situation-specific drive.