Conditioning copulatory behavior to an artificial object: Efficacy of stimulus fading

Copulatory behavior rarely occurs in response to an arbitrary inanimate object. However, such behavior can provide important information about the stimulus control of copulatory behavior. In the present study, male Japanese quail were administered 15–20 conditioning trials that included exposure to an inanimate object and an opportunity to copulate with a live female quail. For some subjects, the stimulus object was always entirely covered with terrycloth. For other birds, the stimulus object contained a taxidermically prepared head and neck of a quail hen. Fading consisted of gradually covering up the neck and then the head portions of the stimulus object over successive trials. After conditioning, all subjects were tested with the entirely covered object. The fading procedure facilitated the conditioning of copulatory behavior to the entirely artificial object. In subjects that received the fading procedure, learning did not depend on pairings of the stimulus object with copulatory opportunity. The results are discussed with respect to an associative mediation mechanism.     

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.     

Dissociation of conditioned appetitive and consummatory sexual behavior: Satiation and extinction tests

Sexual responses were conditioned in male Japanese quail using the opportunity to copulate with a female as the unconditioned stimulus (US). The conditioned stimulus (CS) was a 3-D object made of a taxidermically prepared female quail head mounted on a terry-cloth body. Both appetitive conditioned responses (approach and proximity to the CS) and consummatory conditioned responses (mount and cloacal contact directed toward the CS) developed when 2-min presentations of the CS were followed immediately by the US, but not when the CS and US were separated by trace intervals of 10 or 20 min (Experiment 1). Postconditioning sexual satiation suppressed conditioned cloacal contact responses more than conditioned approach to the CS (Experiment 2), and “acute” extinction suppressed both conditioned mounting and conditioned cloacal contact responses more than conditioned approach to the CS (Experiment 3). These results demonstrate a functional dissociation between conditioned appetitive and consummatory responses and imply that the motivational and/or associative mechanisms underlying the two types of behavior are distinct.     

Observational conditioning of sexual behavior in the domesticated quail

Three experiments were conducted with male domesticated quail to explore whether sexual responses to a three-dimensional conditioned stimulus (CS) object could be acquired through observation. Observational learning was measured by a savings test in which the observers received exposures to the CS paired with the opportunity to copulate with a female bird (the unconditioned stimulus, or US). In all of the experiments, observing a demonstrator copulate with the CS object and then receive access to the US facilitated the subsequent conditioning of the observers. This facilitation effect was not due to observation of just another male bird (Experiment 1) or observation of a male bird that copulated with the CS object (Experiment 2). Rather, the critical factor was observation of pairings of the CS object with the US. Facilitated sexual conditioning was evident in groups of birds that observed pairings of the CS and US, whether or not they witnessed a demonstrator copulating with the CS object (Experiment 3).     

Sign tracking in domesticated quail with one trial a day: Generality across CS and US parameters

When the conditioned stimulus (CS) is located some distance from the unconditioned stimulus (US), pairings of the CS and US can yield either conditioned approach to the CS (sign tracking) or conditioned approach to the US (goal tracking). However, goal tracking is the more common outcome, and, because of that, goal tracking has come to serve as a “standard” measure of associative learning in several laboratories. In contrast, in previous studies of sexual conditioning with domesticated quail, only sign tracking was observed. In the present study, quail continued to show sign tracking rather than goal tracking whether or not the US was highly localized (Experiment 1), whether food or a sexual US was used (Experiment 2), whether the CS was mobile or immobile (Experiment 3), and whether the CS was 91 or 233 cm from the US compartment (Experiment 4). The present findings encourage caution in the routine use of goal tracking as a measure of learning. The possible mechanisms of goal tracking are discussed in terms of occasion setting and behavior systems theory.     

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.     

Temporal encoding in trace conditioning

Conditioned lick suppression in rats was used to explore the role of timing in trace conditioning. In Experiment 1, two groups of rats were exposed to pairings of a CS (CS1) with a US, under conditions in which the interstimulus interval (ISI) that separated CS1 offset and US onset was either 0 or 5 sec. Two additional groups were also exposed to the same CS1→US pairings with either a 0 or a 5-sec ISI, and then received “backward” second-order conditioning in which CS1 was immediately followed by a novel CS2 (i.e., CS1→CS2). A trace conditioning deficit was observed in that the CS1 conditioned with the 5-sec gap supported less excitatory responding than the CS1 conditioned with the 0-sec gap. However, CS2 elicited more conditioned responding in the group trained with the 5-sec CS1-US gap than in the group trained with the 0-sec CS1-US gap. Thus, the CS1-US interval had inverse effects on first- and second-order conditioned responding. Experiment 2 was conducted as a sensory preconditioning analogue to Experiment 1. In Experiment 2, rats received the CS1?CS2 pairings prior to the CS1→US pairings (in which CS1 was again conditioned with either a 0 or a 5-sec ISI). Experiment 2 showed a dissociation between first- and second-order conditioned responding similar to that observed in Experiment 1. These outcomes are not compatible with the view that differences in responding to CSs conditioned with different ISIs are mediated exclusively by differences in associative value. The results are discussed in the framework of the temporal coding hypothesis, according to which temporal relationships between events are encoded in elementary associations.     

The role of signaled periods of nonreinforcement in responding on a random schedule in autoshaping

Little responding develops to a conditioned stimulus (CS) that is placed in a random relation to an unconditioned stimulus (US). However, if the USs not preceded by that CS are themselves signaled by another stimulus, then the CS does come to elicit responding. This result has been attributed (e.g., by Durlach, 1983) to the signal’s blocking of conditioning to background cues that otherwise would prevent conditioning of the CS. However, Goddard and Jenkins (1987) have suggested the alternative that signaling the USs promotes responding due to the adventitious creation of periods of signaled nonreinforcement. Two experiments were conducted to assess this alternative, involving an autoshaping preparation in pigeons. In Experiment 1, little responding to a keylight CS presented in a random relation to a food US occurred, despite the explicit presentation of a discrete noise signaling periods of no food in the intertrial interval (ITI). Experiment 2 was designed to replicate the procedure of Goddard and Jenkins, in which an auditory stimulus extended throughout the ITI of a random schedule, terminating only prior to extra USs and during the CS. Contrary to their findings, little responding developed to the target CS. However, responding did develop when the sound-free period occurred only prior to the extra USs. These results offer little support for the hypothesis that signaled periods of nonreinforcement promote responding on random schedules. However, they are consistent with the view that signaling of ITI USs acts by preventing conditioning of potentially competitive background cues.     

Blocking and enhancement of fear conditioning by appetitive CSs

Prior research on Pavlovian-to-instrumental transfer has shown that when a CS previously associated with shock (AvCS+) is presented contingent upon a choice response to a discriminative stimulus for food reinforcement, it facilitates discrimination learning. Conversely, a response-contingent CS previously associated with the absence of shock (AvCS?) retards discrimination learning. To evaluate whether these findings reflect across-reinforcement blocking and enhancement effects, two experiments investigated the effects of appetitively conditioned stimuli on fear conditioning to a novel stimulus that was serially compounded with the appetitive CS during conditioned-emotional-response (CER) training. Although there were no differential effects of the appetitive CSs in CER acquisition, Experiment 1, using a relatively weak shock US, showed that a CS previously associated with food (ApCS+) retarded CER extinction to the novel stimulus, in evidence of enhanced fear conditioning to that stimulus. In addition, Experiment 2, using a stronger shock US, showed that a CS previously associated with the absence of food (ApCS?) facilitated CER extinction to the novel stimulus, in evidence of weaker fear conditioning to that stimulus. These results parallel traditional blocking effects and indicate not only that an ApCS+ and an ApCS? are functionally similar to AvCSs of opposite sign, but that their functional similarity is mediated by common central emotional states.     

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.     

Reversal from blocking in humans as a result of posttraining extinction of the blocking stimulus

In a blocking procedure, conditioned stimulus (CS) A is paired with the unconditioned stimulus (US) in Phase 1, and a compound of CSs A and X is then paired with the US in Phase 2. The usual result of such a treatment is that X elicits less conditioned responding than if the A-US pairings of Phase 1 had not occurred. Obtaining blocking with human participants has proven difficult, especially if a behavioral task is used or if the control group experiences reinforcement of a CS different from the blocking CS in Phase 1. In the present series, in which human participants and a behavioral measure of learning were used, we provide evidence of blocking, using the above described control condition. Most important, we demonstrate that extinction of the blocking CS (A) following blocking treatment reverses the blocking deficit (i.e., increases responding to X). These results are at odds with traditional associative theories of learning, but they support current associative theories that predict that posttraining manipulations of the competing stimulus can result in a reversal of stimulus competition phenomena.     

Facilitation of appetitive conditioning with naturalistic conditioned stimuli: CS and US factors

Adding limited female cues to a conditioned stimulus (CS) facilitates conditioned male sexual responding. In two experiments, we examined the mechanisms of this facilitation effect. The color of the female cues on the CS was varied in Experiment 1. Similarity between the CS plumage color and the color of the live female (the unconditioned stimulus [US]) could only partially account for the results. The extent to which the facilitation effect represents a specialization of sexual behavior was examined in Experiment 2 by comparing conditioning with either food or copulation as the US. The CSs with female cues elicited more approach and grab responses regardless of which US was used. However, uniquely sexual conditioned responses (mounts and cloacal contacts) were enhanced only when sexual reinforcement served as the US. These findings suggest that the facilitation effect of female cues represents a general feature of appetitive behavior systems.     

Context sensitivity of conditioned suppression following preexposure to the conditioned stimulus

Conditioned suppression in rats is often unaffected when the context (or set of background stimuli) is changed following conditioning. This suggests that responding to the conditioned stimulus (CS) can be relatively insensitive to the context in which the CS is presented. In two experiments, we examined whether sensitivity to contextual stimuli is affected by preexposure to the CS. In Experiment 1, when the CS was novel at the outset of conditioning, conditioned suppression was not affected when the context was changed following conditioning. However, when the CS had been preexposed, responding was weaker when extinction occurred outside of the conditioning context. In Experiment 2, responding was again sensitive to the test context, regardless of whether preexposure occurred in the conditioning context or in an alternate context. These results suggest that the extent to which responding is sensitive to context can depend on the conditioning history of the CS.     

Conditioning of simultaneous and serial feature-positive discriminations

Three experiments with rat subjects examined the development of simultaneous and serial feature-positive discriminations in appetitive conditioning. In Experiment 1, reinforced presentations of a simultaneous light-tone compound were intermixed with nonreinforced presentations of either the light or the tone. The compound stimulus acquired conditioned behaviors of a form characteristic of the predictive feature alone; the element common to reinforced and nonreinforced trials did not evoke conditioned behavior. In Experiment 2, reinforced presentations of a serial light-trace-tone compound were intermixed with nonreinforced tone-alone presentations. The light feature stimulus acquired conditioned behaviors characteristic of visual CSs. The tone stimulus, common to reinforced and nonreinforced trials, evoked conditioned behaviors characteristic of auditory CSs, but only when preceded by the light. In Experiment 3, variations in the interval between the light and tone on reinforced trials had little effect on responding to the light CS but substantially altered the pattern of responding to the tone CS. These results suggested that simultaneous and serial feature-positive discriminations may be solved differently. Performance in simultaneous feature-positive discriminations may be determined solely by associations between the feature stimulus and the reinforcer, but performance in serial discriminations may also involve the acquisition of a conditional cue function to the feature.     

Tests of two explanations of response elimination by noncontingent reinforcement

Two experiments are reported on the elimination of autoshaped keypecking in pigeons by introducing added feedings that have the effect of removing the contingency between keylight (CS) and feedings. In Experiment 1, added feedings were signaled, by an already conditioned stimulus, in one group but not in another. Contrary to the Rescorla-Wagner theory, but consistent with scalar expectancy theory, responding in these groups declined at equal rates. In Experiment 2, following acquisition, some groups were exposed to repeated sessions in which they received feedings alone prior to receiving both CS and feedings noncontingently. Although prior exposure to feedings did reduce the total amount of responding in subsequent noncontingent sessions, it did not, contrary to scalar expectancy theory, reduce the initial level of responding to the CS. It is suggested that a differential between reinforcing conditions in the CS as compared with neighboring non-CS periods is more fundamental to conditioning than is acknowledged in the Rescorla-Wagner theory or in scalar expectancy theory.     

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.     

Stimulus-reinforcer interactions in Pavlovian conditioning of pigeons: Implications for selective associations

In a Pavlovian procedure, groups of pigeons were presented with a compound auditory-visual stimulus that terminated with either response-independent electric shock or food. In a subsequent test, the tone CS was dominant in aversive conditioning, reliably eliciting conditioned head raising and prancing. The red light CS was dominant in appetitive conditioning, reliably eliciting pecking. This result was replicated in a second experiment, in which trials were widely spaced. Pour additional groups of pigeons received pairings of the separate element CSs with the USs. Red light, but not tone, was an effective CS in appetitive conditioning, whereas tone, but not red light, was effective in aversive conditioning. There was no discriminative responding in zero-contingency control groups. Several theoretical accounts of these data are discussed.     

Competition between ethanol-induced reward and aversion in place conditioning

Previous place conditioning studies in mice have shown that injection of ethanol immediately before a conditioned stimulus (CS+) produces conditioned preference, whereas injection of ethanol immediately after CS+ produces conditioned aversion. In the present experiments, we examined the learning that occurs when ethanol is injected in “ambiguous“ procedures that provide the opportunity for both types of conditioning. When ethanol was given midway through the CS (Experiments 1 and 2) or both before and after the CS (Experiment 3), the direction of place conditioning was the same as when mice were exposed only to whichever contingency occurred first (a primacy effect). That is, injection of ethanol in the middle of the CS conditioned aversion, whereas injection both before and after the CS conditioned preference. Because these results support the idea that ethanol elicits both aversive and rewarding effects, they are most consistent with conditioning theories that conceptualize unconditioned stimuli (USs) as events that can activate multiple representational components.     

Changes in the retrievability of associations to elements of the compound CS determine the expression of overshadowing

In two experiments, rats were presented with a taste conditioned stimulus (CS) alone, an odor CS alone, or an odor-taste compound followed by lithium chloride injection. When tested 1 day following conditioning, there was evidence that the odor cue overshadowed conditioning to the taste; however, there was no indication of overshadowing following a longer (21-day) retention interval, despite undiminished strength of the aversion in animals conditioned with only the single element (taste). The overshadowing observed at the 1-day retention interval was not reciprocal. Rats conditioned with the odor CS alone or with the compound CS expressed odor aversions of comparable strength—that is, no overshadowing. However, in contrast to the taste aversion, overshadowing of conditioning to the odor by taste was evident following a 21-day retention interval. Rather than reflecting a failure of the overshadowed stimulus to acquire associative strength, these data suggest that overshadowing may be expressed, or not expressed, as a result of changes in the relative retrievability of learned associations over time.     

Second-order sexual conditioning in male Japanese quail (Coturnix japonica)

Second-order conditioning of social approach to a female conspecific in male Japanese quail was investigated in four experiments. Subjects that received paired first- and second-order trials acquired second-order conditioning in both Experiments 1 and 2. In contrast, subjects that received paired first-order but unpaired second-order trials, and subjects that received unpaired first-order but paired second-order trials, did not acquire second-order conditioning. In Experiment 3, subjects for whom the first-order conditioned stimulus was presented in extinction showed second-order conditioning comparable to that shown by subjects in a control group that did not receive the extinction procedure. In Experiment 4, subjects approached a second-order stimulus less when sexually satiated than when sexually deprived. These findings suggest that second-order sexual conditioning in quail is mediated by an association of the second-order stimulus with a representation of the unconditioned stimulus.