Instrumental learning as a result of omission training on wheel running

The present experiment sought to provide unequivocal evidence of instrumental learning under omission training. Hungry rats received free food reinforcement while spontaneously running in a wheel. For an omission group, running postponed or cancelled reinforcers in the presence of a discriminative stimulus (SD) requiring subjects to reduce responding to earn food. Background food presentations were then yoked to reinforcement delivered in the presence of the discriminative stimulus. For a control group, which received the same stimulus presentations, reinforcement delivery was yoked to the experimental group at all times. The procedure allowed both within- and between-subject comparisons between omission and response-independent schedules. The response-reinforcer delay under the omission contingency was adjusted so as to equate reinforcement frequency in the presence and absence of the SD. As the SD was not correlated differentially with reinforcement and the running response did not involve approach or withdrawal to the site of food delivery, the successful discrimination performance observed in this experiment cannot be accounted for by appeal to implicit classical conditioning. Instead, it is suggested that decreased running in the presence of the discriminative stimulus was based on the animals' veridical representation of the negative contingency between the response and reinforcement.     

Autoshaping, random control, and omission training in the rat

The role of the stimulus-reinforcer contingency in the development and maintenance of lever contact responding was studied in hooded rats. In Experiment I, three groups of experimentally naive rats were trained either on autoshaping, omission training, or a random-control procedure. Subjects trained by the autoshaping procedure responded more consistently than did either random-control or omission-trained subjects. The probability of at least one lever contact per trial was slightly higher in subjects trained by the omission procedure than by the random-control procedure. However, these differences were not maintained during extended training, nor were they evident in total lever-contact frequencies. When omission and random-control subjects were switched to the autoshaping condition, lever contacts increased in all animals, but a pronounced retardation was observed in omission subjects relative to the random-control subjects. In addition, subjects originally exposed to the random-control procedure, and later switched to autoshaping, acquired more rapidly than naive subjects that were exposed only on the autoshaping procedure. In Experiment II, subjects originally trained by an autoshaping procedure were exposed either to an omission, a random-control, or an extinction procedure. No differences were observed among the groups either in the rate at which lever contacts decreased or in the frequency of lever contacts at the end of training. These data implicate prior experience in the interpretation of omission-training effects and suggest limitations in the influence of stimulus-reinforcer relations in autoshaping.     

Acquisition of lever-press responding in rats with delayed reinforcement: A comparison of three procedures

The present study examined the acquisition of lever pressing in rats under three procedures in which food delivery was delayed by 4, 8, and 16 seconds relative to the response. Under the nonresetting delay procedure, food followed the response selected for reinforcement after a specified interval elapsed; responses during this interval had no programmed effect. Under the resetting procedure, the response selected for reinforcement initiated an interval to food delivery that was reset by each subsequent response. Under the stacked delay procedure, every response programmed delivery of food t seconds after its occurrence. Two control groups were studied, one that received food immediately after each lever press and another that never received food. With the exception of the group that did not receive food, responding was established with every procedure at every delay value without autoshaping or shaping. Although responding was established under the resetting delay procedure, response rates were generally not as high as under the other two procedures. These findings support the results of other recent investigations in demonstrating that a response not previously reinforced can be brought to strength by delayed reinforcement in the absence of explicit training.     

Autoshaping a leverpress in rats with lateral, medial, or complete septal lesions

Rats with either control operations or lateral, medial, or complete septal lesions received 600 trials of leverpress training using an autoshaping procedure, i.e., food delivery followed a 10 s illuminated lever presentation, or occurred immediately after a leverpress. Rats with complete septal lesions acquired the leverpress faster than controls and had more food-tray entries per minute during the first 100 trials than the other groups. Rats with lateral or medial septal lesions had leverpress and food-tray entries equivalent to controls. The facilitation of autoshaping a leverpress may partially be explained by the general increase in motor reactivity to stimuli found following septal lesions.     

Renewal of operant performance formerly eliminated by omission or noncontingency training upon return to the acquisition context

Renewal of operant performance formerly eliminated by omission or noncontingency training was explored in two experiments with rats. When pressing a lever was trained with food reinforcement in one context (A) and then eliminated in a second context (B), responding was renewed by returning the rats to the original context (A). This ABA renewal effect was demonstrated in Experiment 1 when the elimination training was an omission procedure (delivery of food for withholding responding) and in Experiment 2 when it was a noncontingency procedure (delivery of food irrespective of responding). Because omission training (differential reinforcement of other behavior) and noncontingency training have been used in applied settings as effective procedures to reduce undesired human behaviors, the clinical implications of our findings for the relapse of undesirable behavior were discussed.     

Blocking, unblocking, and overexpectation in autoshaping with pigeons

Three experiments used pigeons in an autoshaping procedure and a single-subject design to examine compound stimulus control in classical conditioning. Experiment 1 examined the blocking effect, and Experiment 2 examined the unblocking effect. In both experiments, response-independent food was first delivered intermittently in the presence of one distinctively colored houselight but not another. Then, conventional autoshaping trials were carried out in the presence of each houselight. In Experiment 1, the keylight readily elicited responding in the presence of the houselight that had been negatively correlated with food, but not in the presence of the houselight that had been positively correlated with food. In Experiment 2, the keylight readily elicited responding in the presence of the houselight positively correlated with food, but only when the amount of food used on the autoshaping trials was either greater or less than that previously delivered in the presence of the houselight. Experiment 3 examined the overexpectation effect. Conventional autoshaping trials were first carried out by presenting each of two keylights individually. Then, additional autoshaping trials were carried out by presenting the two keylights as a compound, with either the same amount of food or a greater amount of food per trial. Finally, the keylights were retested by again presenting them individually. The number of responses per trial elicited by the keylights decreased when the amount of food used in compound trials was the same as that used in individual trials. However, the number of responses per trial remained approximately the same when the amount of food used in compound trials was greater than that used in individual trials. Taken together, the results of the three experiments demonstrate (a) the generality of the blocking, unblocking, and overexpectation effects by virtue of their extension to appetitive unconditioned stimuli; (b) the suitability of pigeons as subjects and autoshaping as a procedure for studying classical conditioning; and (c) the appropriateness of single-subject designs.     

Cue competition between elementary trained stimuli: US miscuing, interference, and US omission

The present series of experiments was designed to assess whether rule-based accounts of Pavlovian learning can account for cue competition effects observed after elemental training. All experiments involved initial differential conditioning training with A-US and B alone presentations. Miscuing refers to the fact that responding to A is impaired after one B-US presentation whereas interference is the impairment of responding to A after presentation of C-US pairings. Omission refers to the effects on B of A alone presentations. Experiments 1-2a provided clear evidence for miscuing whereas interference was not found after 1, 5 or 10 C-US pairings. Moreover, Experiments 3 and 3a found only weak evidence for interference in an A-US, B ∣ C-US, D ∣ A design used previously to show the effect. Experiments 4 and 5 failed to find any effect of US omission after one or five omission trials. The present results indicate that miscuing is more robust than is the interference effect. Moreover, the asymmetrical effects of US miscuing and US omission are difficult to accommodate within rule-based accounts of Pavlovian conditioning.     

Choice as a dependent measure in autoshaping: sensitivity to frequency and duration of food presentation.

Previous investigations have shown that rate, latency, and percentage of trials with at least one response are somewhat insensitive measures of the strength of autoshaped responding. In the present studies, these measures were contrasted with the allocation of responding during simultaneous choice tests, a measure of response strength frequently used in operant paradigms. In two experiments, nine pigeons were exposed to a forward pairing autoshaping procedure. Training sessions consisted of the successive presentation of three stimuli, each followed by food on either 100%, 50%, or 0% of the trials. Choice testing involved the simultaneous presentation of the three stimuli. In Experiment I, all pigeons consistently directed their initial choice responses and the majority of subsequent responses to the stimulus always followed by food, despite the fact that during training sessions the response rates of most birds were highest in the presence of the stimulus followed by food on 50% of the trials. In Experiment II, rate, latency, and percentage of trials with at least one response did not change appreciably as a function of duration of feeder presentations. However, choice responding was lawfully affected by duration of feeder presentations. These data suggest that choice is perhaps a more sensitive measure of the strength of autoshaped responding than other, more commonly employed, indices.     

Automaintenance in guinea pigs: effects of feeding regimen and omission training

Behavior maintained by stimulus-reinforcer pairings was examined. Guinea pigs maintained at 85 per cent of free-feeding weights reliably contacted a retractable lever presented before delivery of a single piece of guinea-pig chow or a 45-milligram guinea-pig pellet. When animals were given free access to one food and received the second food preceded by the lever, contact responses persisted. Such responses seldom occurred when a single food was freely available and was also delivered after lever presentation. Introduction of an omission training (negative automaintenance) procedure, in which lever contacts resulted in lever retraction and prevented food delivery, strongly reduced lever contacts. Observation indicated that mouthing the food cup, instead of the lever, became the prominent behavior during the prefood stimulus under the omission training procedure.     

Within-session Changes In Responding During Autoshaping And Automaintenance Procedures

Four pigeons were exposed to autoshaping procedures in which an 8-second light on a response key was followed by food. Pecks on the key had no scheduled consequences. Subjects were also exposed to negative automaintenance procedures in which a peck on the illuminated key canceled the following food. The intertrial interval varied from an average of 7 seconds to an average of 232 seconds in different conditions. Rate of responding usually changed within sessions during autoshaping. Responding also changed within sessions for the 1 subject that responded during negative automaintenance. The within-session patterns of responding were flatter, peaked later, and were more symmetrical around the middle of the sessions at lower rates of food presentation, regardless of whether subjects responded on autoshaping, negative automaintenance, or previously reported variable-interval schedules. These results imply that similar variables produce within-session changes in responding during both classical (Pavlovian) and operant conditioning procedures.     

Aversive and compensatory classical conditioning with diazepam as conditional stimulus

The purpose of the first experiment was to investigate whether diazepam could acquire anxiogenic properties by signalling an aversive event. Rats were trained in an operant chamber in the pentylenetetrazol (PTZ) model of anxiety. Thereafter the animals were divided into groups that received classical diazepam conditioning (Group 1), and conditioning of diazepam + tone (Group 2). In the test phase diazepam was injected prior to placement in the operant chamber. Group 2 selected the PTZ-appropriate lever more often than the other groups, indicating that the tone induced anxiety, and diazepam did not. Tones and shock may therefore be more easily associated than diazepam and shock. The second experiment investigated this. Rats were trained the same way as in the first experiment. Thereafter the experimental group received injections of a small dose of diazepam prior to a second injection of a large dose of diazepam. The hypothesis was that a compensatory anxiogenic conditional response to diazepam's anxiolytic effect should be elicited by the small dose. There were no differences between the groups in lever selection, indicating that a compensatory anxiogenic response was not elicited.     

Overshadowing of a stimulus-reinforcer association by an instrumental response

In two experiments, rats were first exposed to pairings of a clicker and food; they were subsequently, in order to measure the effectiveness of the clicker as a conditioned reinforcer, given the opportunity to press a lever which turned the clicker on. For one group of animals the food originally delivered in the presence of the clicker had been contingent on their performance of an instrumental response (running in a running wheel); for a second the contingency between clicker and food had been purely classical. Although the actual correlation between clicker and food was identical for the two groups, the clicker was a less effective conditioned reinforcer for the first group than for the second. In a third experiment, all animals were initially required to run to obtain food in the presence of the clicker, but one group received additional trials on which food was delivered contingent on running in the absence of the clicker. This group showed less tendency to lever press for the clicker than a second group that had received free food on trials when the clicker was not presented. The results of all three experiments suggest that conditioning to the clicker could be overshadowed if the occurrence of food was more reliably predicted by the execution of an instrumental running response; they thus support the view that instrumental conditioning depends on the establishment of an association between response and reinforcer similar to the association between stimulus and reinforcer underlying classical conditioning.     

Autoshaping in the rat: Effects of omission on the form of the response

Two experiments were conducted to investigate the effect of an omission contingency on behavior related to and characterizing autoshaped lever contacts in the rat. In Experiment I an omission contingency imposed on autoshaped lever contacts forceful enough to produce a press (.078N) resulted in a significant decrease in lever presses, but had no effect on frequency of lever touches (contacts of insufficient force to produce a press) or rate of food tray entry during lever presentation. In contrast, rats which received a similar number of lever-food pairings, but whose behavior had no programmed consequences (yoked control subjects), showed an increase in lever press rate, a significant decrease in rate of food tray entry, and no change in rate of lever-touches. In Experiment II, the effect of a similar omission contingency on the topography of lever contact responses was investigated. Prior to omission training subjects contacted the lever primarily by pawing it. Following omission training this behavior was suppressed, with a subsequent increase in lever contacts characterized as nosing. Yoked control subjects showed no significant changes in lever contact topography. The results indicate that (1) an omission contingency does not simply eliminate wholesale those topographies which incur the contingency but produces subtle adaptive changes in lever contact topography; and (2) the nature of the autoshaped response in the rat does not appear to be rigid enough to depend solely upon the nature of the unconditioned stimulus or the conditioned stimulus, but can also be determined by the relationships existing between the animal's behavior and these stimuli.     

Towards mouse models of perseveration: a heritable component in extinction of operant behavior in fourteen standard and recombinant inbred mouse lines

Extinction of instrumental responses is an essential skill for adaptive behavior such as foraging. So far, only few studies have focused on extinction following appetitive conditioning in mice. We studied extinction of appetitive operant lever-press behavior in six standard inbred mouse strains (A/J, C3H/HeJ, C57BL/6J, DBA/2J, BALB/cByJ and NOD/Ltj) and eight recombinant inbred mouse lines. From the response rates at the end of operant and extinction training we computed an extinction index, with higher values indicating better capability to omit behavioral responding in absence of reward. This index varied highly across the mouse lines tested, and the variability was partially due to a significant heritable component of 12.6%.To further characterize the relationship between operant learning and extinction, we calculated the slope of the time course of extinction across sessions. While many strains showed a considerable capacity to omit responding when lever pressing was no longer rewarded, we found a few lines showing an abnormally high perseveration in lever press behavior, showing no decay in response scores over extinction sessions.No correlation was found between operant and extinction response scores, suggesting that appetitive operant learning and extinction learning are dissociable, a finding in line with previous studies indicating that these forms of learning are dependent on different brain areas. These data shed light on the heritable basis of extinction learning and may help develop animal models of addictive habits and other perseverative disorders, such as compulsive food seeking and eating.     

Time-dependent changes in conditioned suppression

Time-dependent changes in a response following aversive conditioning were investigated using a conditioned suppression procedure in a within-subjects design. Four groups of pigeons received Pavlovian conditioning “off the baseline”, immediately followed by an operant task. During the Pavlovian phase, two groups received a forward pairing of a tone with shock, one group received a backward pairing, and one group received a truly random pairing. One of the forward pairing groups also received a delay between the Pavlovian and operant phases. For all groups, key pecking was reinforced on a variable-interval schedule during the operant phase. Testing sessions were identical to training sessions, except that the tone used during Pavlovian conditioning was presented either 0, 15, 30, 45, of 60 minutes after the operant phase began. Testing sessions in which the Pavlovian phase was omitted were also included. The results showed suppression to change as a function of the retention interval, with maximum suppression occurring at intermediate intervals. This U-shaped function was obtained for 11 of the 12 pigeons in the forward-pairing groups and for three of the five in the truly random group. Pigeons in the background pairing group did not show changes in suppression as a function of the retention interval.     

Discriminative properties of shock-correlated reinforcement in an operant context

A two-choice discrete operant procedure was devised for the study of shock-correlated reinforcement effects in rats. In the presence of one auditory stimulus, responding on one response lever was reinforced with food; with another auditory stimulus, responding on a second lever was reinforced. It was found that discrimination performance of one group, relative to appropriate control groups, was facilitated when electric shock was correlated with reinforcement on one lever and not on the other. Further, relative discrimination levels were found to be higher on the lever correlated with the shock than on the alternate lever. The significance of the results for operant within-S studies and for a mediational theory of shock-correlated reinforcement was discussed.     

Overshadowing of a context aversion by a novel incentive in operant conditioning

Two experiments examined the processes underlying the suppression of instrumental behaviours by lithium in rats, as reported by Meachum (1988 and this issue). Experiment 1 examined whether presenting a novel sucrose solution prior to lithium chloride administration would overshadow aversion learning to either the stimuli of the operant chamber or to familiar food pellets. After lever pressing had been established, and in the absence of responding, animals received free deliveries of a novel sucrose solution, familiar food pellets, or both, or they were exposed to only the cues of the operant chamber, prior to lithium injections. Lever pressing for food pellets was then assessed. It was found that the animals receiving the novel sucrose, either alone or with the familiar food pellets, pressed more for pellets than either the group receiving only food pellets or the group exposed to only the context. In addition, there was no appreciable difference in the response rates between the context-only group and the group that received the familiar food pellets. These outcomes were interpreted in terms of the novel sucrose overshadowing aversion learning to the context. Experiment 2 investigated whether in fact aversive contextual conditioning could be obtained using the present parameters. This was accomplished by directly manipulating the contexts. In this experiment animals were trained to lever press in two distinctive contexts. Subsequently, one context was paired with the novel sucrose, and the second was experienced in the absence of reinforcement prior to toxicosis. During a subsequent non-reinforced test it was found that responding in the context paired with the novel sucrose was considerably higher than responding in the context that was experienced alone. These findings stand in contrast to the taste-mediated contextual potentiation observed when a consumatory response is used to assess aversive contextual conditioning.     

Autoshaping the pigeon's gape response: acquisition and topography as a function of reinforcer type and magnitude.

The pigeon's key-pecking response is experimentally dissociable into transport (head movement) and gape (jaw movement) components. During conditioning of the key-pecking response, both components come under the control of the conditioned stimulus. To study the acquisition of gape conditioned responses and to clarify the contribution of unconditioned stimulus (reinforcer) variables to the form of the response, gape and key-contact responses were recorded during an autoshaping procedure and reinforcer properties were systematically varied. One group of 8 pigeons was food deprived and subgroups of 2 birds each were exposed to four different pellet sizes as reinforcers, each reinforcer signaled by a keylight conditioned stimulus. A second group was water deprived and received water reinforcers paired with the conditioned stimulus. Water- or food-deprived control groups received appropriate water or food reinforcers that were randomly delivered with respect to the keylight stimulus. Acquisition of the conditioned gape response frequently preceded key-contact responses, and gape conditioned responses were generally elicited at higher rates than were key contacts. The form of the conditioned gape was similar to, but not identical with, the form of the unconditioned gape. The gape component is a critical topographical feature of the conditioned key peck, a sensitive measure of conditioning during autoshaping, and an important source of the observed similarities in the form of conditioned and consummatory responses.     

Hippocampal associative cellular responses: dissociation with behavioral responses revealed by a transfer-of-control technique

Multiunit activity was recorded in the CA3 field of the dorsal hippocampus in freely moving rats during classical conditioning and subsequent presentation of the CS on operant baselines for food reward as well as shock avoidance. Rats were first trained in a nonsignaled bar-pressing-dependent shock omission task and in a food-motivated lever-pressing task (60-s VI). Five sessions with presentations of a previously habituated tone as a CS paired with footshock as a US were then given. Testing was carried out by presenting the CS alone while behavioral responses were maintained by reinforcement in both instrumental tasks on alternate sessions. As expected, the CS induced a marked suppression of lever pressing for food reward and a marked enhancement of bar-pressing for shock avoidance. The analysis of the frequency of multiunit discharges to the CS revealed that the hippocampal cellular responses established during classical conditioning were maintained while two different behavioral responses were exhibited to the CS. The results showed that the associative response of hippocampal neurons may be dissociated from the Pavlovian conditioned responses the CS elicits. They support the hypothesis that hippocampal cellular responses represent a neural index of the acquired CS-US associative representation.     

The role of response-contingent incentives in lithium chloride-mediated suppression of an operant response

Three experiments investigated what role a novel incentive plays in the development of operant response suppression mediated by lithium chloride. In all experiments animals were trained to press two levers under concurrent schedules of reinforcement. In Experiment 1 responding on one lever delivered a familiar incentive (food pellets), whereas responding on an alternative lever delivered a novel incentive (sucrose solution) prior to lithium chloride injections. If lithium was administered immediately after the instrumental session, the action associated with the novel, but not with the familiar, incentive was suppressed. By comparison, in a control group for which responding on both levers led to the familiar incentive, both actions were suppressed. Experiment 2 examined whether the novelty, rather than the sensory properties, of the incentive is crucial for observing performance suppression. It was found that animals familiarized with the “target” incentive were insensitive to aversion conditioning by lithium, in that there was no difference in response rates between the action that delivered the familiar incentive from that which earned the “target”. In contrast, if animals were unfamiliar with the “target” incentive at the time of aversion conditioning, they suppressed responding on the lever that was associated with the novel incentive but did not suppress responding on the lever associated with the familiar incentive. Experiment 3 investigated the mechanism underlying instrumental performance suppression. After the completion of concurrent lever press training, novel sucrose was introduced in conjunction with the pellets for responding on one lever; responding on the other lever continued to deliver only familiar pellets. Lithium injections were then administered either immediately following the sessions or several hours after the sessions. It was found that the rate of responding on the lever associated with the contingent delivery of sucrose was suppressed below that of the pellet-alone action. By comparison, if lithium injections were administered several hours following the session, an elevation in responding on the sucrose-plus-pellet lever was observed. The outcomes of all three experiments demonstrate not only that the novelty of an incentive is important in obtaining performance suppression, but also that a novel incentive can punish instrumental responding if it has been associated with toxicosis.