Pavlovian conditioning of sexual arousal: parametric and background manipulations

Recent studies have shown that a Pavlovian conditioned stimulus (CS) for unconsummated sexual arousal can increase the rate of copulation in the rat. This report includes five experiments examining the effects of parametric manipulations on the conditioned arousal response. Results show that between six and nine trials are necessary for reliable conditioning, but extinction is somewhat slower than acquisition. The function for the CS-US (unconditioned stimulus) interval is quadratic, with a minimum of several minutes required for effective conditioning. In the first three experiments, it appeared that background cues were conditioned as well as the designated CSs, and this was tested explicitly in the last two studies. In one, the effect of background cues was shown by training and testing in different situations; in the second, background cues were shown to be subject to latent inhibition. These results demonstrate the influence of Pavlovian learning in sexual behavior and help to provide the basis for an animal model of the acquisition of deviant sexual arousal in humans.     

Uncertainty, as defined by the contingency between environmental events, and the adrenocortical response of the rat to electric shock

The pituitary-adrenal system is thought to be sensitive to the degree of uncertainty in a situation. In addition, there is some question whether the pituitary-adrenal system can be conditioned in a Pavlovian sense. Three experiments are reported here. The first and third sought to define uncertainty in terms of conditioned stimulus-unconditioned stimulus (CB-US) and US-US contingencies, which vary the amount of information that can be used to predict the occurrence of discrete shocks. The second experiment examined the possibility that the adrenocortical system was subject to the laws of Pavlovian conditioning, by using a conditioned emotional response paradigm. The results showed that the magnitude of the Pituitary-adrenal response varied in a curvilinear manner along the dimension of uncertainty. Very low and very high degrees of uncertainty resulted in greater corticosterone elevations than did moderate levels. No evidence for Pavlovian conditioning of the adrencortical system was found, although behavioral measures showed fear conditioning. The data presented were supportive of the hypothesis that the pituitary-adrenal response reflects the operation of an arousal system.     

Differential effects of forward or simultaneous conditioned stimulus-unconditioned stimulus intervals on the defensive behavior system of the Norway rat (Rattus norvegicus)

To test several predictions derived from a behavior-systems approach, the authors assessed Pavlovian fear conditioning in rats after 30 trials of forward, simultaneous, or unpaired training. Direct evidence of conditioned fear was collected through observation of flight and freezing reactions during presentations of the conditioned stimulus (CS) alone. The authors also tested the CS's potential to reinforce an instrumental escape response in an escape-from-fear paradigm. On the one hand, rats that received forward training showed conditioned freezing, but no conditioned flight was observed. On the other hand, rats that received simultaneous training showed conditioned flight, but no conditioned freezing was observed. Rats that received either forward or simultaneous pairings showed instrumental learning of the escape-from-fear response. Implications for several theories of Pavlovian conditioning are discussed.     

The Adaptive Significance of Sexual Conditioning: Pavlovian Control of Sperm Release

Male quail received Pavlovian conditioning trials that consisted of placement in a distinctive experimental chamber (the conditioned stimulus) paired with the opportunity to copulate with a female (the unconditioned stimulus). For control subjects, exposures to the conditioned and unconditioned stimuli were unpaired. After four and six trials, each subject was placed in the experimental chamber with a probe stimulus that included some of the visual cues of a female's head and neck. Pavlovian conditioning increased how much time subjects spent near the probe stimulus. Conditioned subjects also released greater volumes of semen and greater numbers of spermatozoa than the control subjects. Significant differences were not obtained in serum testosterone levels or in other measures of sperm quality. These results demonstrate that sexual Pavlovian conditioning can affect reflexes involved in sperm release and thereby modulate reproductive fitness.     

Constraints on Pavlovian conditioning of the pigeon: Relative conditioned reinforcing effects of red-light and tone CSs paired with food

Recent Pavlovian conditioning experiments presented all possible CS-US combinations of red-light and tone CSs and food and shock USs to separate groups of pigeons. Pigeons receiving shock USs demonstrated conditioned head raising followed by prancing to both CSs, but CRs were acquired more rapidly to tone than to red light. Although pigeons receiving food USs rapidly acquired a conditioned response of pecking to the red-light CS, there was no evidence of conditioned responding in groups receiving tone-food pairings. This outcome left open the possibility that Pavlovian pairings of tone and food may have resulted in association formation that was not revealed in performance. The present series of experiments attempted to reveal that association, using an indirect method of assessment, conditioned reinforcement. Experiment 1 demonstrated that both red light and tone paired with food became positive conditioned reinforcers, suggesting that an association between tone and food was formed in the same number of Pavlovian conditioning trials that previously failed to yield any direct evidence of conditioning. Experiment 2, which presented fewer conditioning trials, revealed that the tone-food association was formed less rapidly than the red light-food association. Experiment 3 demonstrated that the observed outcomes were not attributable to unconditioned, rather than conditioned, reinforcing effects of the Css.     

The efficacy of cue exposure with response prevention in extinguishing drug and alcohol cue reactivity

It has been suggested that drug cue‐elicited urges and psychophysiological reactions are the results of Pavlovian conditioning processes and that it should be possible to extinguish these responses with cue exposure with response prevention. It has already been shown that subjective cue‐elicited urges can be extinguished, but it is unclear whether this is also true for cue‐elicited psychophysiological arousal. This was tested in the present study in a heterogeneous sample of drug and alcohol dependent patients. It was found that cue‐elicited urges can indeed be extinguished. However, such a clear pattern of extinguished cue reactivity was not found for the psychophysiological measures. Furthermore, the extinction of drug urges was not specific for cue exposure treatment. It is concluded that cue‐elicited psychophysiological arousal does not underlie subjective cue reactivity and may not reflect Pavlovian conditioned drug responding. Copyright © 2006 John Wiley & Sons, Ltd.     

Effect of telencephalon ablation on the reinforcing and eliciting properties of species-specific events in Betta splendens

In male Betta splendens, aggressive behavior is drastically attenuated following telencephalon ablation. Because instrumental training and Pavlovian conditioning experiments with intact fish have suggested that associative factors may play an important role in the performance of agonistic behaviors, the effect of ablation on instrumental learning and Pavlovian conditioning was studied. In Experiment 1, ablation had no effect on the learning of the instrumental tunnel-swimming response reinforced by mirror presentation (i.e., viewing a conspecific), although the mirror presentations in yoked-control groups elicited fewer responses in ablates than in normal and sham-operated control fish. Yoked controls further established that instrumental responding was maintained by the reinforcement contingency and was not merely the result of increased motor activity. Experiment 2 studied Pavlovian conditioning of the components of the agonistic display. Unconditioned fin erection, gill erection, and tail beating (i.e., unconditioned responses, URs) to the mirror US all were less frequent in ablates than in normals or shams. Of these, only gill cover erection showed evidence of true conditioning (i.e., conditioned responses; CRs) in which responses to the conditioned stimulus (CS) are due to the pairings of CS and US (unconditioned stimulus). However, ablates suffered no impairment of conditioned gill erections. Ablates performed fewer fin erections to the CS; however, fin erection responses were not due to CS-US pairings but were attributable to pseudoconditioning. These results are related to hypotheses postulating the involvement of learning mechanisms in ablation-produced deficits and normal aggressive behavior.     

Pavlovian conditioning of Hermissenda: current cellular, molecular, and circuit perspectives

The less-complex central nervous system of many invertebrates make them attractive for not only the molecular analysis of the associative learning and memory, but also in determining how neural circuits are modified by learning to generate changes in behavior. The nudibranch mollusk Hermissenda crassicornis is a preparation that has contributed to an understanding of cellular and molecular mechanisms of Pavlovian conditioning. Identified neurons in the conditioned stimulus (CS) pathway have been studied in detail using biophysical, biochemical, and molecular techniques. These studies have resulted in the identification and characterization of specific membrane conductances contributing to enhanced excitability and synaptic facilitation in the CS pathway of conditioned animals. Second-messenger systems activated by the CS and US have been examined, and proteins that are regulated by one-trial and multi-trial Pavlovian conditioning have been identified in the CS pathway. The recent progress that has been made in the identification of the neural circuitry supporting the unconditioned response (UR) and conditioned response (CR) now provides for the opportunity to understand how Pavlovian conditioning is expressed in behavior.     

Skin conductance response conditioning with CS intensities equal to and greater than UCS intensity

With .2-sec bursts of white noise as both conditioned stimulus (CS) and unconditioned stimulus (UCS), conditioning of first-interval skin conductance responses was obtained when the intensity of the CS equaled and exceeded that of the UCS. There was no evidence that second-interval response conditioning occurred. Nonspecific response frequencies were also affected by the variations in stimulus intensity, this raising some question about typical controls employed in SCR conditioning. There was some evidence that second interval responses were suppressed by the intense CS values. It was concluded that the existence of simple conditioning with a CS/UCS intensity ratio equal to or greater than unity was contrary to the Pavlovian proposition that a CS must be biologically less salient than the UCS in order for conditioning to occur. It was noted, however, that the suppression of second-interval responses might indicate that anticipatory CRs which are not confounded with orienting reflexes are prevented from exhibiting a conditioning effect when a high CS/UCS intensity ratio is employed.     

Backward conditioning as an extinction procedure

In two conditioned suppression experiments, rats received Pavlovian forward defense conditioning in which tonal conditioned stimuli (CSs) terminated with the onset of scrambled grid shock unconditioned stimuli (USs). After this experience, the rats then received a Pavlovian backward conditioning procedure in which the same USs now terminated with the onset of the same CSs. Although the two experiments differed greatly in terms of CS and US parameters, number of forward and backward pairings, and in terms of the general techniques used to establish and measure the Pavlovian conditioned response (CR), the results of both experiments agreed in showing that backward conditioning can indeed weaken a CR based on forward pairings. The results also show that, under some conditions, the backward procedure can be at least as effective in weakening an established CR as the traditional CS-alone extinction procedure; but, under other conditions, the backward procedure is less effective and leads to more spontaneous recovery than the CS-alone procedure.     

The Elusiveness of Evaluative Conditioning

Evaluative conditioning (EC) is an important variant of Pavlovian conditioning in which the outcome is a change in affective response to the conditioned stimulus (CS). It is the best extant account, with evidence, for affective change in humans. Good laboratory models are available. This paper reviews a set of findings which suggest that the actual occurrence of EC, and its magnitude, varies widely in both real world and laboratory situations. Attention to known parameters of Pavlovian conditioning may account for some, but not all, of the failures and successes. Six empirical studies on humans are described; two document frequent failures of EC to occur in real world situations where the Pavlovian conditions for development of animal phobias or taste aversions are present, two are real-world experiments in which no evidence for EC is obtained, and two are laboratory failures to produce EC, by pairing neutral odor CSs with a variety of unconditioned stimuli (USs). We suggest that there are important, not understood factors, that modulate the appearance of EC, and that for both theoretical and applied reasons, these factors should be identified.     

Further evidence for the summation of latent inhibition and overshadowing in rats’ conditioned taste aversion

Repeated exposures to a target taste (X) attenuated subsequent development of rats’ conditioned aversion to X (latent inhibition effect). Presentation of another taste (A) after X in conditioning (serial X-A compound conditioning) also attenuated conditioned X aversion compared with conditioning without A (overshadowing). Furthermore, the latent inhibition and overshadowing effects summed to show the least conditioned aversion in the rats given both the target preexposures and the serial X-A compound conditioning treatment. These results question the validity of the comparator hypothesis as an explanation for Pavlovian conditioning of rats’ conditioned taste aversion.     

Learned fear of "unseen" faces after Pavlovian, observational, and instructed fear

This study compared fear learning acquired through direct experience (Pavlovian conditioning) and fear learning acquired without direct experience via either observation or verbal instruction. We examined whether these three types of learning yielded differential responses to conditioned stimuli (CS+) that were presented unmasked (available to explicit awareness) or masked (not available to explicit awareness). In the Pavlovian group, the CS+ was paired with a mild shock, whereas the observational-learning group learned through observing the emotional expression of a confederate receiving shocks paired with the CS+. The instructed-learning group was told that the CS+ predicted a shock. The three groups demonstrated similar levels of learning as measured by the skin conductance response to unmasked stimuli. As in previous studies, participants also displayed a significant learning response to masked stimuli following Pavlovian conditioning. However, whereas the observational-learning group also showed this effect, the instructed-learning group did not.     

The role of nucleus accumbens dopamine in outcome encoding in instrumental and Pavlovian conditioning

Considerable evidence suggests that dopamine in the core subregion of the nucleus accumbens is not only involved in Pavlovian conditioning but also supports instrumental performance. However, it is largely unknown whether NAc dopamine is required for outcome encoding which plays an important role both in Pavlovian stimulus-outcome learning and instrumental action-outcome learning. Therefore, we tested rats with 6-hydroxydopamine (6-OHDA) induced dopamine depletion of the NAc core for their sensitivity to outcome devaluation in a Pavlovian and an instrumental task. Results indicate that 6-OHDA-lesioned animals were sensitive to outcome devaluation in an instrumental task. This finding provides support to the notion that NAc core dopamine may not be crucial in encoding action-outcome associations. However, during instrumental conditioning lever pressing rates in 6-OHDA-lesioned animals were markedly lower which could reflect an impaired behavioral activation. By contrast, after outcome-specific devaluation in a Pavlovian task, performance in 6-OHDA-lesioned animals was impaired, i.e. their magazine-directed responding was non-selectively reduced. One possibility to explain non-selective responding is that NAc core DA depletion impaired the ability of conditioned stimuli to activate the memory of the current value of the reinforcer.     

Reinstatement of extinguished conditioned responses and negative stimulus valence as a pathway to return of fear in humans

The present study investigated reinstatement of conditioned responses in humans by using a differential Pavlovian conditioning procedure. Evidence for reinstatement was established in a direct (fear rating) and in an indirect measure (secondary reaction time task) of conditioning. Moreover, the amount of reinstatement in the secondary reaction time task was significantly correlated with the difference in valence between the conditioned stimulus (CS)+ and the CS-after extinction. These data provide clear evidence for reinstatement and for the role of negative stimulus valence in the return of conditioned responding after extinction.     

Associations in Pavlovian conditioned inhibition

Two experiments with rat subjects used a variety of transfer tests to examine the associations learned when Pavlovian inhibition is established by an A+, AX− paradigm. Experiment 1 found in a conditioned suppression situation that inhibition conditioned to X with one exciter (A) readily transferred to another exciter (B) which had been paired with the same shock US. Transfer occurred even when the response to A had been extinguished prior to testing with B. However, X did not inhibit a general activity response produced by a B which had been subsequently paired with a food US. Experiment 2 employed a Pavlovian conditioning situation in which A and B, when separately paired with the same food US, evoked dissimilar responses. Nevertheless, an inhibitor trained in an A+, AX− paradigm successfully inhibited the different response evoked by B. However, such an X did not inhibit the behaviors acquired by A or B when they were subsequently paired with a shock US. The transfer of Pavlovian inhibition across conditioned stimuli and responses but not across unconditioned stimuli is consistent with the notion that a conditioned inhibitor acts to prevent activation of a US representation which would normally be activated by conditioned exciters.     

Learning with arbitrary versus ecological conditioned stimuli: Evidence from sexual conditioning

Laboratory investigations of Pavlovian conditioning typically involve the association of an arbitrary conditioned stimulus (CS) with an unconditioned stimulus (US) that has no inherent relation to the CS. However, arbitrary CSs are unlikely to become conditioned outside the laboratory, because they do not occur often enough with the US to result in an association. Learning under natural circumstances is likely only if the CS has a preexisting relation to the US. Recent studies of sexual conditioning have shown that in contrast to an arbitrary CS, an ecologically relevant CS is resistant to blocking, extinction, and increases in the CS-US interval and results in sensitized responding and stronger second-order conditioning. Although the mechanisms of these effects are not fully understood, these findings have shown that signature learning phenomena are significantly altered when the kinds of stimuli that are likely to become conditioned under natural circumstances are used. The implications of these findings for an ecological approach to the study of learning are discussed.     

Pavlovian eyeblink conditioning in combat veterans with and without post-traumatic stress disorder

Several recent studies have investigated relationships between post-traumatic stress disorder (PTSD) and learning and memory problems. These reports have found in general that not only does PTSD affect trauma-related memories, but when patients with PTSD are compared with similar trauma patients without PTSD, general memory impairments have been found. The present paper reports a study in which associative learning, using Pavlovian eyeblink conditioning, was investigated in combat veterans with and without chronic PTSD, using interstimulus intervals of 500 and 1000 msec in two separate experiments. Although several recent reports suggest that larger-magnitude autonomic conditioned responses occur in patients with PTSD during Pavlovian conditoning, the present study found evidence of impaired Pavlovian eyeblink conditioning in combat veterans with and without PTSD, compared to non-combat veterans. Although these data suggest that combat leads to an impaired associative learning process regardless of whether PTSD is apparent, a group of community-dwelling combat veterans not under medical treatment showed normal conditioning, suggesting that variables other than prior combat must also be involved.     

Effect of posttrial episodes on conditioning in compound conditioned stimuli

The effect of signaled and nonsignaled posttrial episodes (PTEs) on conditioning to the target CS of a preceding compound CS-US trial was examined in two conditioned emotional response (CER) experiments with rats. Experiment 1 revealed that a nonsignaled PTE, a second shock US, interfered with conditioning to the preceding target CS and that the interference effect of the PTE was greater when it occurred 5 sec as opposed to 1,800 sec after the preceding compound CS-US trial. Experiment 2 revealed that if the shock PTE was signaled by a CS, its ability to interfere with conditioning was reduced. This second study also revealed that omission of an "expected" US interfered with conditioning to the target CS. The implications of these data for information processing models of Pavlovian conditioning are discussed.     

Pavlovian Eyeblink Conditioning in Combat Veterans With and Without Post-Traumatic Stress Disorder

Several recent studies have investigated relationships between post-traumatic stress disorder (PTSD) and learning and memory problems. These reports have found in general that not only does PTSD affect trauma-related memories, but when patients with PTSD are compared with similar trauma patients without PTSD, general memory impairments have been found. The present paper reports a study in which associative learning, using Pavlovian eyeblink conditioning, was investigated in combat veterans with and without chronic PTSD, using interstimulus intervals of 500 and 1000 msec in two separate experiments. Although several recent reports suggest that larger-magnitude autonomic conditioned responses occur in patients with PTSD during Pavlovian conditioning, the present study found evidence of impaired Pavlovian eyeblink conditioning in combat veterans with and without PTSD, compared to non-combat veterans. Although these data suggest that combat leads to an impaired associative learning process regardless of whether PTSD is apparent, a group of community-dwelling combat veterans not under medical treatment showed normal conditioning, suggesting that variables other than prior combat must also be involved.