Factors governing one-trial contextual conditioning

When a rat was placed in a chamber and shortly thereafter received a single footshock, it showed conditional freezing upon re-exposure to that chamber but not a different one (Experiment 1). Experiments 2–4 showed that the probability of this freezing decreased linearly with decreases in the delay between placement in the chamber and shock delivery. With very short delays (e.g., less than 27 sec), there was no freezing. Experiments 2 and 3 demonstrated that a 2-mm pre-exposure to the chamber, 24 h prior to shock delivery, reduced the minimum delay necessary to successfully condition freezing. Experiment 4 demonstrated that shorter delays were successful in conditioning freezing if a salient tone was a component of the contextual stimulus. The changes in freezing caused by delay interval and preexposure did not simply reflect the total time in the context, suggesting that there may be two requirements that place temporal restrictions on the conditioning of the freezing response. One is satisfied by sufficient exposure, whether or not that exposure is contiguous with shock. The second requirement is for a small amount of context exposure that is contiguous with shock.     

The relationship between shock magnitude and passive avoidance learning

Three experiments examined the relationship between shock magnitude and the rate of acquisition of a passive avoidance response. Experiment 1 indicated that the use of a relatively large magnitude of shock can disrupt learning to remain on a platform in the center of an open field to avoid shock. The inferior learning of the group trained with high shock was replicated in Experiment 2, which also demonstrated that avoidance learning can occur rapidly with this level of shock if the platform is located in the corner of the apparatus. To explain this, it was proposed that thigmotactic behavior is responsible for the disruption in avoidance behavior when training is conducted in the center with high-magnitude shock. Finally, Experiment 3, essentially a replication of Experiment 1 except that the platform was placed in the corner of the test compartment, demonstrated a direct relationship between shock magnitude and passive avoidance learning. The results are seen as being consistent with accounts which maintain that avoidance learning can be influenced by the occurrence of species-specific defense reactions.     

Immediate shock,passive avoidance,and potentiated startle: Implications for the unconditioned response to shock

Three experiments were performed to study the immediate-shock freezing deficit, a deficit in freezing in rats that results when electric shock is delivered immediately upon exposure to a novel context. This deficit was accompanied by failures to detect evidence of passive avoidance (Experiment 1) or potentiation of the auditory startle response (Experiment 2). The deficit in freezing was attenuated by preexposure to the shocked context (Experiment 3). The results support the view that fear-related behaviors are activated by signals for shock rather than by shock itself. They also suggest that the immediate-shock freezing deficit is due to a failure to process the to-be-conditioned contextual cues (Fanselow, 1986a, 1990).     

Dominance,the bidirectional hypothesis,and Pavlovian backward conditioning in the US-US paradigm

Based upon considerations raised by Soviet research, the role of relative stimulus intensity, or dominance, in the unconditioned stimulus-unconditioned stimulus (US-US) paradigm was investigated under circumstances presumed favorable to the backward conditioned response (CR). Using the classically conditioned forelimb response of the cat, a brief shock (USD delivered to one forepaw preceded a shock (US2) to the opposite forepaw in paired conditioning fashion; subjects in the control group received explicitly unpaired presentations of the stimuli. Conditioning in both the forward and backward directions was evaluated by the appearance of contralateral CRs on test trials to each of the USs. In Experiment 1, a ratio of the intensities between US1 and US2 of 100:80 was used to create a relative dominance in favor of the backward CR. In addition, to evaluate the suggestion that the appearance of the backward CR is retarded in the Pavlovian paradigm, overtraining was provided to a forward conditioning criterion of 200%. In Experiment 2, the cats were exposed to successive reductions in the intensity of US2 to verify manipulations of dominance reportedly involved in the reactivation of a latent backward CR. Although forward conditioning was readily established to USl, there was no evidence of back-ward conditioning to US2 under any of the conditions.     

Recovery of conditioned suppression after backward pairings

In four conditioned suppression experiments with rats (Rattus norvegicus), backward pairings of a shock unconditioned stimulus (US) and a tone conditioned stimulus (CS) eliminated an already established conditioned response (CR), but there was recovery of the CR if the shock was later withheld. In Experiment 1, there was recovery after backward pairings, regardless of whether the period after the US was normally shock free or not. In Experiment 2, the occurrence of recovery depended on the CS’s being presented closely after the US in response elimination. Levels of recovery were positively correlated with the resistance of the response to elimination during backward pairings (Experiments 3 and 4). Taken together, these data support the notion that recovery after backward pairings is a form of renewal (see, e.g., Bouton, 1991) and is not due toprotection from extinction.     

Overshadowing of situational cues with variable but not fixed intertrial intervals

In two experiments, we examined the conditions under which signaling an unconditioned stimulus (US) with a nominal conditioned stimulus (CS) interferes with the conditioning of situational cues in defensive freezing in the rat. Subjects received footshock USs that were (1) either signaled or unsignaled and (2) either varied or fixed in their temporal location within the conditioning session. Experiment 1, with only one trial per session, yielded no evidence that signaling affected pretrial freezing using either a fixed or variable interval between placement in the context and shock onset. In a test in which no CSs or footshocks were presented, groups that previously had received footshock at a fixed temporal location showed greatest freezing at around that same time. For groups that had received footshocks at various times, freezing declined across the test session. Experiment 2 showed overshadowing of pretrial freezing after more extensive conditioning with many trials per session, but only if the intershock intervals were variable rather than fixed.     

Influence of conspecific stress odors and shock controllability on conditioned defensive burying

In Experiment 1, rats received a session of 80 inescapable tail shocks or no shocks while restrained in a tube. During tests of conditioned defensive burying 24 h later, the bedding of the chamber contained odors from either stressed or nonstressed conspecific donor rats. Following a single prod shock, subjects that had had prior shocks or that were tested with the stress odors spent significantly less time burying the prod, made smaller piles of bedding, and displayed more freezing behavior. The combination of prior shock and stress odors during later testing enhanced these effects. In Experiment 2, a yoked group of rats that was given inescapable shocks, in contrast to a group that had wheel-turn escape training and one that was restrained but not shocked, later showed significantly less burying and more freezing when tested for defensive burying with stress odors present. In both experiments the duration of burying and the heights of piles were positively correlated, and both of these measures were negatively correlated with freezing. The demonstrated capacity of unconditioned stress odors to mediate different degrees of fear, depending upon the controllability of prior shock, is related to other studies of learned helplessness, and the predominance of freezing over burying is discussed in terms of various types of defensive strategies, stimulus-control processes, and the author’s stress-coping-fear-defense (SCFD) theory.     

Effects of US parameters on classical conditioning of cat hindlimb flexion

Unconditioned stimulus (US) intensity and duration were manipulated to determine their effects on cat hindlimb flexion conditioning. Seven consecutive days of acquisition training of a hindlimb flexor response to a tone conditioned stimulus (CS) were followed by 2 days of extinction. Eight animals in each of 12 groups received a 1-, 2-, 3-, or 4-mA, 60-Hz shock delivered to the right hindleg for 25, 50, or 100 msec as the US. Analysis of conditioned-response (CR) frequency indicated that conditioned responding was a positive function of both the intensity and duration of shock, although these variables did not interact with one another. CR latency and amplitude were decreased and increased, respectively, by increases in US intensity. The pattern of results reported here may support a contiguity notion of conditioning, and are discussed in the context of other conditioning preparations.     

Role of shock intensity in the learned helplessness paradigm

Although there have been many studies of the interference effect produced by exposure to inescapable shock, little is known about the role of shock intensity. This experiment factorially manipulated four levels of shock intensity during exposure to inescapable shock and three levels of intensity during the test for interference. Interference occurred at each training shock intensity when training and test shocks were similar. Interference was not obtained when training intensity was high but testing intensity low or medium or when training intensity was low or medium and test intensity was high. These findings pose problems for learned helplessness, learned inactivity, competing motor response, and catecholamine depletion hypotheses of the interference effect in the rat.     

Spontaneous recovery in cross-motivational transfer (counterconditioning)

In two experiments with rat subjects, we examined the effects of a retention interval on performance in two conditioning paradigms in which a conditioned stimulus (CS) was associated with different unconditioned stimuli (USs) in successive phases of the experiment- Experiment 1 was designed to examine aversive-appetitive transfer, in which the CS is associated with shock and then food; Experiment 2 was designed to examine appetitive-aversive transfer, in which the CS is associated with food and then shock. Aversive and appetitive conditioned responses (freezing and head-jerk responding, respectively) were scored from videotape. In both experiments, a 28-day retention interval following the end of Phase 2 caused a recovery of the Phase 1 response and a resuppression of the Phase 2 response. The results suggest that the original association is not destroyed when the CS is associated with a new US in Phase 2. They also suggest that both retroactive and proactive interference effects may result-from interference with performance output rather than a disruption or loss of what is learned during or stored from the target phase.     

General transfer across sensory modalities survives reductions in the original conditioned reflex in the rabbit

Two experiments demonstrated that transfer of training between CSs from different sensory modalities survived substantial reductions in responding to the first CS. In both experiments, animals received three stages of training. Stage 1 entailed CS-US training with a CS from one modality (e.g., light), and Stage 3 entailed CS-US training with a CS from another modality (e.g., tone). The experiments differed in treatment during Stage 2. In Experiment 1, animals either remained in their home cages or received unreinforced exposures to the first CS, which extinguished the original CR. In Experiment 2, the animals received either continued CS-US training or exposure to the CS and US but at a long interval (2,800 msec), which eliminated the original CR. As the baseline for detection of transfer effects, each experimental group had a control group that received Stage 1 training with a 2,800-msec CS-US interval, which produced minimal CR acquisition. The results of both experiments revealed substantial positive transfer across CS modalities regardless of the treatment during Stage 2. The transfer did not appear immediately on test presentations of the second CS in Stage 3. Rather, the transfer appeared as an enhancement in the rate of CR acquisition after reinforced training with the second CS had commenced. The results are discussed with respect to stimulus generalization, neutralization of background stimuli, and learning processes superordinate to specific associations.     

Second-order conditioning of the rabbit’s nictitating membrane response as a function of the CS2-CS1 and CS1-US intervals

Three experiments investigated conditioning of the rabbit’s nictitating membrane response (NMR) in a second-order conditioning procedure which intermixed first-order trials (CS1-US) and second-order trials (CS2-CS1) from the outset of training. Experiment 1 provided a controlled demonstration that substantial levels of second-order conditioning can be obtained with the NMR preparation. Experiment 2 showed that the level of CR acquisition to CS2 was an inverse function of the CS2-CS1 interval over the values of 400, 800, and 2,400 msec. Experiment 3 found that CR acquisition to CS2 and CS1 in second-order conditioning varied in a parallel fashion across CS1-US intervals. Similarly, Experiment 3A found that the level of CR acquisition to the two components of a serial compound (CSA-CSB-US) varied in a parallel fashion as a function of the CSB-US interval. The results of the CS2-CS1 and CS1-US interval manipulations were all predictable from the known CS-US interval effects in NMR conditioning with a single CS. The present results are discussed with regard to their implications for accounts of serial compound conditioning and second-order conditioning.     

Activity during prior shock determines subsequent shock-elicited fighting in the rat

Two experiments investigated the relationship between activity during shock and the magnitude of subsequent impairment of shock-elicited fighting in the rat. Different levels of intra-shock activity were engendered in two ways. In Experiment 1, differing temporal forms of inescapable shock were employed to produce markedly different levels of activity. In Experiment 2, a passive-escape procedure was used to explicitly reinforce nonmovement during shock relative to a yoked, inescapable shock control. Results indicated that relative to the performance of subjects not previously shocked, fighting impairment was produced only by those prior treatments that promoted reduced intrashock activity. Since one of the prior shock treatments involved inescapable shock but the other did not, these findings may be viewed as strong support for the notion that behavior during shock, rather than uncontrollability, is the critical determinant of the observed impairment effects. There was some suggestion in both studies that shock treatments that resulted in sustained or increased intrashock activity tended to produce augmentation of fighting. Both inhibitory and facilitative effects of prior shock exposure are discussed in terms of an interacting response theory of shock treatment effects.     

Influence of conspecific and predatory Stressors and their associated odors on defensive burying and freezing responses

In Experiment 1, male rats were exposed to either aggressive (i.e., alpha) or nonaggressive conspecific colonies and tested 24 h later, with or without alpha odors, for freezing behavior and burying of a wall prod that had been the source of a brief electric shock. The results indicated that prior defeat experience and the presence of alpha odors alone during testing had no significant effects, but the combination of prior defeat and alpha-odor testing significantly decreased burying and increased freezing behavior. In Experiment 2, we examined the effects of noncontact exposure to a cat, as a predatory Stressor, during subsequent prod-shock tests involving the presence or absence of cat odors. Exposure to a cat failed to disrupt later prod burying and did not produce freezing. However, the presence of cat odors during testing significantly reduced the amount of defensive burying,without resulting in an increment in freezing. In Experiment 3, rats were given 1, 5, or 30 inescapable preshocks in the presence of either cat odors or a hedonically neutral citronella odor and were tested 24 h later for prod burying and freezing with or without these odors. Both the cat and the citronella odors resulted in a significant reduction in burying and an increase in freezing for rats given 5 and 30 preshocks and tested in the presence of these respective conditioned odors. For the groups that were given 5 preshocks, preshock and later testing in the presence of cat odors resulted in significantly less prod burying and more freezing than for rats that were preshocked and tested in the presence of citronella. The findings of these three ethoexperimental studies are discussed in terms of the learned-helplessness theory, the stress-coping-fear-defense (SCFD) theory, and the concept of selective CS-US associability.     

Conditions that potentiate the effects of electroconvulsive shock administered 24 hours after avoidance training

Three experiments assessed the conditions that potentiate effects of an electroconvulsive shock (ECS) administered 24 h after avoidance training. Stimuli present immediately prior to the ECS were systematically varied. In Experiment 1, which employed a passive avoidance task, the primary determinant of whether the ECS disrupted retention was whether the situational cues present at the time of ECS delivery were those associated with the initial training experience: ECS disrupted performance only when it was administered in the original training apparatus, regardless of whether or not a footshock was presented immediately prior to ECS. In Experiment 2, which employed an active, shuttlebox avoidance task, both the situational cues from the training apparatus and a footshock were necessary to potentiate the disruptive effects of the ECS. Experiment 3 revealed that ECS effects on performance of the active avoidance task can also be potentiated by a combination of apparatus cues and the warning signal used in initial training. These results are interpreted as indicating that informational functions of stimuli present when an ECS is administered are important determinants of the effects of the ECS.     

The US-preexposure effect in honeybees

Signaled avoidance was studied in individual honeybees that visited the laboratory regularly to take sucrose solution from a target set on the sill of an open window. During feeding, substrate vibration or airstream was used to signal a brief shock that could be avoided by breaking off contact with the food for a few seconds. Aversive conditioning of the context was measured in terms of return time (the time between successive visits). In Experiment 1, experience with unsignaled shock was found to lengthen return time—which experience with signaled shock did not—and to impair performance in subsequent avoidance training with signaled shock (the US-preexposure effect). In Experiment 2, experience with unsignaled shock given after signaled avoidance training lengthened return time but had no effect on response to the signal in a subsequent extinction test. These results closely resemble the results obtained in analogous experiments with vertebrates.     

Effects of handling and context preexposure on the immediate shock deficit

Theimmediate shock deficit refers to the failure of a shock to become associated with contextual stimuli when the shock is presented simultaneously with the rat’s placement in a context. The basic procedure consists of a presentation of the shock as soon as the animal is placed in an observation chamber. Handling of the animal, which immediately precedes the shock, and the novelty of the chamber in which the immediate shock is delivered are potential variables that might be responsible for this associative deficit. In Experiment 1, handling reduced context conditioning but was not responsible for the immediate shock deficit. Experiment 2 revealed that the novelty of the chamber was not a significant factor. These results discount the possibility that handling and the novelty of the chamber are responsible for the deficit produced by the immediate shock. It is suggested that immediate shock could be employed as a control procedure for the study of context conditioning.     

Pavlovian conditioning to contexts that precede the place where shock occurs as measured by freezing and activity suppression in mice

Three experiments were conducted to demonstrate that the place where an organism has been, before the organism is moved to a place with aversive consequences, can also become aversive through classical conditioning. In Experiment 1, two groups of 8 mice were exposed to three different contexts in succession, with a single shock occurring in the third context. The distal context was a putative 3-min conditioned stimulus (CS) for freezing; the second context was a delay manipulation; and the unconditioned stimulus (US) occurred in the proximal context. The group delayed for 15 sec showed significantly more freezing to the distal CS context than did the group delayed for 3 h. In a second experiment, conditioning to the distal context was demonstrated with a discrimination procedure for 8 more mice by using two different distal contexts as CS+ and CS? for the proximal context with shock. On CS+ days, 3 min of exposure to the distal context was followed within 5 sec by placement in the proximal box where shock occurred, whereas on CS? days, exposure to a second distal context was followed immediately by return to the home cage. Very strong differences in freezing between the CS+ and CS? distal contexts were found in all 8 mice after 14 days of conditioning. In a third experiment, the discriminative procedure was repeated for 9 more mice, with two changes. More objective stabilometertype activity measures were substituted for observed freezing, and, in addition to the CS+ and CS? distal context trials, each mouse was also exposed to a third discriminative distal context, which was followed by 15 min in a delay chamber followed by shock in the proximal context. This discrimination procedure with the activity suppression measure again resulted in significant differences between the contexts. The CS+ context and the context followed by a 15-min delay did not differ, but both of them differed from the CS? context.     

Transfer across CS-US intervals and sensory modalities in classical conditioning of the rabbit

Two experiments investigated transfer of the rabbit’s conditioned nictitating membrane response (NMR) from shorter to longer CS-US intervals in conjunction with a change in CS modality, for example, light to tone. In Experiment 1, three experimental groups received initial training with a 400-msec CS-US interval, which produced substantial CR acquisition, and three control groups received initial training with a 2,800-msec CS-US interval, which produced minimal CR acquisition. Subsequently, the experimental and control groups received training with an 800-, 1,800-, or 2,800-msec CS-US trace interval. At the same time, the modality of the CS was changed from tone to light (or vice versa). Experiment 2 contained three groups that received initial exposure to a 400-msec CS-US interval, a 2,800-msec CS-US interval, or just the experimental chambers. Subsequently, all three groups received training with an 800-msec CS-US interval in a different CS modality. The results of both experiments revealed substantial positive transfer across CS modalities from the 400-msec CS-US interval to the 800-msec CS-US interval. There was also significant transfer to the 1,800-msec but not the 2,800-msec CS-US interval. The transfer did not appear immediately on test presentations of the second CS. Rather, the transfer appeared as an enhancement in the rate of CR acquisition after reinforced training with the second CS had commenced. The results are discussed with respect to mechanisms of transfer and facilitation of trace conditioning.     

Shuttlebox avoidance to intense white noise: Acquisition and the Kamin effect in rats

Experiment I demonstrated shuttlebox avoidance conditioning using intense white noise as a UCS. Ten rats were given 25 trials a day for 6 days. Escape latencies declined and avoidance responses increased over trial blocks. Experiment II provided support for a functional similarity between shock as a UCS and intense noise as a UCS by demonstrating the Kamin effect following incomplete shuttlebox training to noise. Separate groups of rats were given 25 trials followed by an additional 25 trials either 0, 1, 4, or 24 h later. The U-shaped Kamin effect was evident in the avoidance measure. A similar but inverted U-shaped function was obtained for the escape latency measure. Escape latencies were longer on retraining than on original training at 1 h but not at 0, 4, or 24 h after original training.