Context excitation and modulation of conditioned sexual behavior

Three experiments were conducted to investigate direct and modulatory influences of context in the conditioned sexual behavior of male Japanese quail. A preference test procedure was used to assess the acquisition of contextual excitation. In Experiment 1, following direct context-unconditioned stimulus (US) pairings, male quail shifted their contextual preference from an initially preferred context to one in which they received copulatory opportunity with a female quail (US). Unpaired control group subjects did not demonstrate this shift in preference. This place preference procedure was used in Experiments 2 and 3 to assess contextual excitation when context was trained in the presence of a discrete conditioned stimulus (CS). Experiment 2 provided evidence that context can modulate responding to a discrete CS. In Experiment 3, we varied the spatial contiguity between the context and the US. Some subjects received the US directly in the training context, whereas other subjects received the US in an alternate context. Contextual excitation was evident only in subjects that received the former. Thus, there is a dissociation between the modulatory and excitatory properties of context in sexual conditioning that may depend on the context-US spatial contiguity.     

Stimulus competition in the absence of compound conditioning

Most associative theories have assumed that stimulus competition occurs only between conditioned stimuli (CSs) that are trained in compound. The present research investigated the possibility of competition between two CSs that were individually paired to the same unconditioned stimulus (US). We used human subjects in an anticipatory suppression analogue to Pavlovian conditioning. Experiment 1 showed that X+ training followed by A+ training resulted in impaired responding to X. This did not occur when A+ training preceded X+ training. Experiment 2 replicated the basic effect and showed that it did not occur when the Phase 2 training consisted of A? instead of A+ nor when the A+ pairings occurred in a second context. Experiment 3 showed that A+ pairings occurring in a second context could still produce the effect when X was tested in the context in which the A+ pairings had occurred, but not when X was tested in a context different from that used for A+ training. Collectively, these results suggest that individually trained CSs may compete with each other when one of those CSs is more strongly activated by the test context than the other one.     

Temporal coding in conditioned inhibition: Retardation tests

Two lick suppression experiments with rats were conducted in order to determine the nature of the temporal information that is encoded as a result of Pavlovian conditioned inhibition training (conditioned stimulus {CS} A→unconditioned stimulus {US}/AX→noUS). After inhibition training, the conditioned inhibitor (X) was paired with the US in order to measure inhibition, as assessed through retarded behavioral control by CS X. Three temporal relationships were manipulated: the A-US interval, the X-A interval of inhibition training, and the X-US interval of the retardation test pairings. Retardation was greatest when the X-US temporal relationship matched the time at which the US was expected (but not delivered) on the X-A inhibition training trials. Thus, the present experiments provide evidence with retardation tests that, during conditioned inhibition training, subjects encode the temporal location of the omitted US relative to the inhibitory CS. These findings complement those of previous studies using summation tests of conditioned inhibition (Barnet & Miller, 1996; Denniston, Blaisdell, á Miller, 1998; Denniston, Cole, & Miller, 1998).     

Pavlovian conditioning in multiple contexts: Competition between contexts for comparator status

Water-deprived rats were used to investigate the effects of training a CS in more than one context on conditioned lick suppression. In each experiment, partial reinforcement of the CS was intermingled with unsignaled presentations of the US. In Experiment 1, subjects were either trained in one context alone, trained consecutively in two contexts (such that all training in one context occurred prior to any training in the second context), or trained alternately in two contexts. Following training, the first context, the second context, or neither context was extinguished. Testing of the CS occurred in a third (neutral) context. To the extent that either training context became established as a comparator stimulus for the CS, the comparator hypothesis (Miller & Matzel, 1988) predicts an increase in excitatory responding to the CS following extinction of that context. Subjects trained in a single context exhibited appreciable fear of the CS only when the CS’s training context had been extinguished. Additionally, subjects trained consecutively in the two contexts showed increased fear of the CS following extinction of the second, but not the first training context (i.e., a recency effect). Subjects trained alternately in the two contexts showed no increased fear of the CS as a result of either context alone being extinguished. In Experiment 2, subjects trained alternately in two contexts showed increased fear of the CS only when both training contexts were extinguished, suggesting that both training contexts had become comparator stimuli. These data indicate that multiple training contexts can either compete or act synergistic-ally in modulating responding to a Pavlovian trained CS as a function of the order of training in the different contexts.     

Divergence of conditioned eyeblink and conditioned fear in backward Pavlovian training

Two experiments of Pavlovian conditioning with rabbits evaluated the effects of initiating or continuing a conditioned stimulus (CS) after a paraorbital unconditioned stimulus (US). In Experiment 1, backward pairings, in which a CS came on after the US, produced a CS that appeared inhibitory on a measure of eyeblink conditioning but excitatory on a potentiated-startle measure of conditioned fear. In Experiment 2, extending the duration of a CS that came on prior to the US, so that it continued after the US, decreased eyeblink conditioned responses, whereas it increased conditioned fear. The data from the two experiments confirm and extend those of Tait and Saladin (1986), supporting the suppositions of AESOP (Wagner & Brandon, 1989) that conditioned eyeblink and conditioned fear can be dissociated under various temporal relationships between the CS and US.     

Modulatory mechanisms in Pavlovian conditioning

Pavlovian conditioning has traditionally been thought to involve the acquisition of excitatory and inhibitory associations between a conditioned stimulus (CS) and an unconditioned stimulus (US). Recent research, however, has encouraged the view that Pavlovian learning may also encompass a higher order modulatory mechanism, in which animals use information about another stimulus to control responding to the CS. Positive modulators signal a positive relationship between the CS and the US, whereas negative modulators signal that the CS-US relationship is not in force. In both cases, the modulatory control appears to be orthogonal to the modulator’s direct associations with the US. This article reviews and evaluates the literature on this Pavlovian modulatory mechanism.     

Massive preexposure and preexposure in multiple contexts attenuate the context specificity of latent inhibition

Latent inhibition, which refers to attenuated responding to a conditioned stimulus (CS) after CS-unconditioned stimulus (CS-US) pairings as a result of CS-alone presentations prior to the pairings, is often attenuated if preexposure and conditioning occur in different contexts (i.e., it is context specific). Here we report two conditioned lick suppression experiments, using rat subjects, that examined whether manipulations known to attenuate the context specificity of extinction could also eliminate the context specificity of latent inhibition. Context specificity of latent inhibition was eliminated when the CS was preexposed in multiple contexts (Experiment 1) and when the CS was massively pre-exposed in the training context alone (Experiment 2). These results and their practical implications are discussed in the framework of contemporary theories of latent inhibition.     

CS-duration and partial-reinforcement effects counteract overshadowing in select situations

Two experiments used rats in a conditioned lick suppression preparation to investigate how the conditioned stimulus (CS)-duration and partial-reinforcement effects (i.e., weakened responding due to conditioning with a CS of longer duration and presenting nonreinforced CSs intermingled with CS—unconditioned stimulus [US] pairings, respectively) interact with overshadowing. Experiment 1 found that when overshadowing treatment was combined with either extended CS duration or partial reinforcement, the response deficit was weaker than when either of these three treatments was administered alone. In Experiment 2, the generality of the findings in Experiment 1 was investigated by replicating it with various US—US intervals. This time counteraction was observed only when both the absolute duration of total CS exposure and the US—US interval were short. The results support neither the view that the ratio between the total CS exposure and total time in the context determines the CS-duration and the partial-reinforcement effects nor the view that these two effects arise from a loss of effectiveness of the excitatory CS—US association during CS-alone exposures in partial reinforcement or early periods of CS exposure with long CSs.     

Latent inhibition as a performance deficit resulting from CS—context associations

Treatments that attenuate latent inhibition (LI) were examined using conditioned suppression in rats. In Experiment 1, retarded conditioned responding was produced by nonreinforced exposure to the CS prior to the CS-US pairings used to assess retardation (i.e., conventional LI). In Experiment la, retarded conditioned responding was induced by preexposure to pairings of the CS and a weak US prior to retardation-test pairings of the CS with a strong US (i.e., Hall-Pearce [1979] LI). Both types of LI were attenuated by extensive exposure to the training context (i.e., context extinction) following the CS-US pairings of the retardation test. Experiment 2 examined the specificity of the attenuated LI effect observed in Experiment 1. After preexposure to two different CSs in two different contexts, each CS was paired with a US in its respective preexposure context. One of the two contexts was then extinguished. This attenuated LI to a greater degree for the CS that had been trained in the extinguished context. Experiment 3 differentiated the roles in LI of CS-context associations and context-US associations. Following preexposure to the CS in the training context, LI was reduced by further exposure to the CS outside the training context. This observation was interpreted as implicating the CS-context association as a factor in LI. Thus, the results of these experiments suggest that LI is a performance deficit mediated by unusually strong CS-context associations. Implications for Wagner’s (1981) SOP model and Miller and Matzel’s (1988) comparator hypothesis are discussed.     

Recovery of the rabbit’s conditioned nictitating membrane response without direct reinforcement after extinction

Three experiments demonstrated that, following the extinction of an established conditioned stimulus (CSA—e.g., tone), the pairing of a novel, cross-modal stimulus (CSB—e.g., light) with the unconditioned stimulus (US) results in strong recovery of responding to the extinguished CSA. Experiment 1 demonstrated that the recovery of responding to CSA is not the result of US reinstatement but is attributable to pairings of CSB with the US. Experiment 2 demonstrated that the recovery of responding is specific to CSA and is not the result of cross-modal generalization. Experiment 3 revealed that a large number of CSB-US pairings in Stage 1 significantly reduced the amount of recovery to CSA during subsequent CSB-US trials. Experiment 3 also provided unexpected evidence of cross-modal secondary extinction. The extinction and subsequent recovery of responding seen in the present experiments is discussed with respect to possible contributions from contextual associations, CS processing, US processing, conditioned response expression, and layered excitatory associations.     

Retardation of conditioned excitation following operational inhibitory blocking

Water-deprived adult rats were used in a conditioned-suppression-of-licking procedure to determine the effect of inhibitory training with a novel stimulus trained in simultaneous compound with a previously established conditioned inhibitor. This procedure constitutes an inhibitory analogue to the excitatory blocking procedure in classical conditioning. The conditioned-inhibition training consisted of either explicitly unpaired CS and US presentations or negative contingency training, in which the likelihood of the US was greater in the absence than in the presence of the CS, but the CS and the US were occasionally paired. To assess conditioned inhibition, a retardation test was used, and comparable retardation was obtained for subjects that were administered the blocking treatment and control subjects given similar conditioned-inhibition training with a compound stimulus in which the nontarget element was not previously established as a conditioned inhibitor.     

Latent inhibition of the conditioning context: Further evidence of contextual potentiation of retrieval in the absence of appreciable context-US associations

Previous research in our laboratory has found superior performance when classically conditioned responses are observed in the training context as opposed to outside it, even when direct context-US associations have been minimized by either the choice of conditioning parameters or extinction to the context. The present experiment used latent inhibition of the conditioning context as an alternative method of examining contextual cue effects in the absence of appreciable direct context-US associations. Water-deprived rats received tone-footshock pairings in one of two distinctly different apparatuses, but all were tested in a common apparatus. Animals conditioned in the test enclosure displayed more lick suppression than those conditioned outside the test enclosure. Other animals tested without the tone present also exhibited more suppression if conditioning had occurred in the test context rather than outside it, indicating that direct associations between the conditioning context and shock had been formed. However, when formation of direct associations to the conditioning context was attenuated in additional animals through extensive preexposure to the context prior to conditioning, more suppression to the tone was still seen when conditioning had occurred in the test context rather than outside it. These results add support to the position that the training context augments recall even when direct associations between the context and the US are attenuated. The phenomenon is discussed in terms of facilitated retrieval of nominal CS-US associations, configural retrieval cues, and conditional discriminations.     

Some determinants of second-order conditioning

In a Pavlovian conditioning situation, an initially neutral stimulus may be made excitatory by nonreinforced presentations in compound with an established conditioned excitor [i.e., second-order conditioning (SOC)]. The established excitor may be either a punctate cue or the training context. In four conditioned suppression experiments using rats, we investigated whether SOC phenomena parallel other cue interaction effects. In Experiment 1, we found that the response potential of a target stimulus was directly related to the intertrial interval when SOC was mediated by a punctate cue, and inversely related to the intertrial interval when SOC was mediated by the training context. Experiment 2 demonstrated that punctate- and context-mediated SOC are oppositely affected by posttraining context extinction, and Experiments 3 and 4 demonstrated that context- and punctate-mediated SOC are differentially affected by conditioned stimulus (Experiment 3) and unconditioned stimulus (Experiment 4) preexposure treatments. These findings parallel phenomena in conditioned inhibition and cue competition situations.     

Summation of responding to CSs and an excitatory test context

The influence of test-trial delay of CS onset in obtaining response summation of an excitatory CS and an independently conditioned context was investigated. Water-deprived rats were given tone-shock and click-shock pairings in the training context and unsignaled footshocks in the test context. Durations of lick suppression in response to the tone (Test 1) and the click (Test 2) were then assessed in the test context. Licking was more suppressed when CS onset occurred early in the test session (e.g., 0 sec) than when it occurred later in the test session (e.g., 300 sec). The results from control groups that had received shock in an irrelevant context rather than the test context indicated that this effect was due to fear of the test context rather than diffuse, nonassociative fear. With onset of the clicks early in the test session on the second test day, response summation of the test context with the clicks was observed. This suggests that appreciable spontaneous recovery had occurred from any test-context extinction that took place on the first test day. We conclude that early onset of the CS on the test trial favors response summation of CSs and test context.     

Conditioned inhibition training of the competing cue after compound conditioning does not reduce cue competition

In each of two experiments, we studied Pavlovian fear conditioning (as assessed by barpress conditioned suppression) in 32 albino rats. Following a two-stage cue-competition procedure (A+ then AX+), we subjected the competing cue (A) to conditioned inhibition training (B+, BA?) before testing the target cue (X). Conditioned inhibition training was designed to weaken the putative A-unconditioned stimulus (US) association, perhaps changing it to an A-no-US association. Performance-deficit theories of cue competition, such as comparator theory and retrieval-interference theory, predict that such procedures should weaken cue competition, causing Conditioned Stimulus X (CS X) to evoke strong responding. The same prediction can be deduced from recent acquisition-focused models (Dickinson &; Burke, 1996; Van Hamme &; Wasserman, 1994). In opposition to this prediction, however, we found in both experiments that conditioned inhibition training had no detectable effect on cue competition even though it successfully abolished conditioned responding to CS A. In Experiment 2, moreover, we found evidence against the hypothesis that the weak response to CS X was due to generalization decrement rather than to cue competition. Results favor early learning-deficit theories of cue competition over performance-deficit theories and over the recent acquisition-focused models.     

Retention of conditioned autonomic and behavioral responses in preweanling rats: Forgetting and reinstatement

Preweanling rats, 16 days of age, responded to an olfactory conditioned stimulus (CS) paired with a shock unconditioned stimulus (US) with increases in heart rate and behavioral activation. In two experiments this finding was replicated and, in addition, it was found that the form of these conditioned responses (CRs) changed after a retention interval. When tested 24 h after CS-US pairings, the subjects displayed a decrease in heart rate accompanied by CS-elicited freezing. Giving two unsignaled shocks prior to the delayed test effectively reinstated the tachycardia and behavioral arousal CRs. The results are discussed in terms of contextual influences on the form of the CR and how changes in the magnitude of context fear may alter responding to an olfactory CS.     

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.     

Contribution of conditioned opioid analgesia to the shock-induced associative US-preexposure deficit

The modulatory effect of conditioned opioid analgesia on learning in the US-preexposure paradigm was examined in three experiments using water-deprived rats. In Experiment 1, it was found that tailflick latencies increased immediately after the rats were exposed to a context in which footshock had previously been administered. Prolonged nonreinforced exposure to the context attenuated this analgesia. Experiment 2 tested the possibility that the effectiveness of CS-US pairings in an excitatory context might be reduced by a conditioned analgesic response that lessens the perceived intensity of the US. Administration of the opiate antagonist naloxone prior to CS-US pairings in the excitatory context reduced the US-preexposure deficit—that is, the retarded response to the CS—but did not eliminate it, suggesting that part of the observed deficit resulted from conditioned activation of the endogenous opioid system. In Experiment 3, it was found that exposure to the excitatory context immediately prior to a CS-US pairing in an associatively neutral context resulted in a conditioned response deficit, indicating that the analgesia elicited by the excitatory context was sufficient to reduce US effectiveness. In combination with other recent reports, these results suggest that the associative deficit resulting from preexposure to a shock US may, in certain instances, represent the sum of several different associative processes.     

One-trial simultaneous and backward fear conditioning as reflected in conditioned suppression of licking in rats

In three experiments, groups of albino rats received one strictly simultaneous pairing of a 4-sec auditory conditioned stimulus (CS) and a 4-sec 1-mA shock unconditioned stimulus (US). Other groups received a backward pairing, in which the US began before the CS, or a forward pairing, in which the CS began before the US. Control groups received only the US or received both the CS and the US but widely separated in time. Later, the CS was presented while the rats licked a drinking tube for water, and CS-elicited suppression of licking was taken as an index of the Pavlovian conditioned response (CR). It was found that groups receiving a single forward or a single simultaneous pairing suppressed more than groups that had received a backward pairing; and the backward groups, in turn, suppressed more than the control groups. It appears, then, that excitatory fear conditioning, as reflected in conditioned suppression of licking in rats, can be produced in a single trial by both backward and simultaneous conditioning procedures.     

Contrasting AAC and ABC renewal: the role of context associations

Rats were used in a lick suppression preparation to assess the contribution of conditioned-stimulus (CS)–context and context–unconditioned-stimulus (US) associations to experimental extinction. Experiment 1 investigated whether strengthening the CS–acquisition context association enhances extinction by determining whether stronger extinction is observed when CS-alone trials (i.e., extinction treatment) are administered in the acquisition context (AAC renewal), relative to a context that is neutral with respect to the US (ABC renewal). Less recovery of responding to the CS was observed in the former than in the latter case, extending the finding that AAC renewal is weaker than ABC renewal to our lick suppression preparation. Experiment 2 assessed the contribution of the acquisition context–US association to extinction of a CS by examining the effect of postextinction exposure to the acquisition context on responding to the extinguished CS. This manipulation enhanced responding to the extinguished CS in AAC, but not ABC, renewal. Experiment 3 addressed the contribution of the CS–acquisition context association by examining the potential of a neutral stimulus, presented in compound with the target CS during extinction treatment, to overshadow the CS–acquisition context association. This manipulation enhanced responding to the extinguished CS in AAC, but not ABC, renewal. The results stress the important role of contextual association in extinction and renewal.