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).     

Blocking of the sexual conditioning of differentially effective conditioned stimulus objects

The blocking phenomenon was investigated in the sexual response system of male Japanese quail. Access to a live female quail served as the unconditioned stimulus (US). The same audiovisual cue served as the pretrained stimulus in all of the experiments. Following asymptotic conditioning of the audiovisual cue, a second conditioned stimulus (CS2) was added. In Experiment 1, CS2 was a rectangular wood block that had little or no resemblance to a female quail and could not support copulatory behavior. In Experiment 2, CS2 was a terrycloth object that had no quail parts but could support copulatory behavior, and, in Experiment 3, CS2 was a terrycloth object that had a taxidermically prepared head of a female quail added. The terrycloth-only object supported more rapid conditioning than did the wood block, but the blocking effect was obtained with both kinds of stimuli. Approach responding to the terrycloth + head object required pairing it with copulatory opportunity, and the terrycloth + head object supported at least as rapid conditioning as did the terrycloth-only object. However, responding to the terrycloth + head object was not blocked by the pretrained audiovisual cue. These results indicate that the blocking effect occurs in sexual conditioning even with stimulus objects that can support copulation. However, the addition of species-typical head cues to an object makes that object such a powerful stimulus that conditioned approach responding to it cannot be blocked by a previously conditioned arbitrary audiovisual cue.     

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.     

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.     

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.     

Conditioned avoidance responses survive contingency degradation in the garden slug,Lehmannia valentiana

Joint presentations of a conditioned stimulus (CS) and an unconditioned stimulus (US) strengthen the contingency between them, whereas presentations of one stimulus without the other degrade this contingency. For example, the CS can be presented without the US either before conditioning (CS–no US and then CS–US; latent inhibition) or after conditioning (CS–US and then CS–no US; extinction). In vertebrate subjects and several invertebrate species, a time lapse usually results in a return of the conditioned response, or spontaneous recovery. However, in land mollusks, spontaneous recovery from extinction has only recently been reported, and response recovery after latent inhibition has not been reported. In two experiments, using conditioned aversion to a food odor as a measure of learning and memory retention, we observed contingency degradation via latent inhibition (Experiment 1) and extinction (Experiment 2) in the common garden slug, Lehmannia valentiana. In both situations, the contingency degradation procedure successfully attenuated conditioned responding, and delaying testing by several days resulted in recovery of the conditioned response. This suggests that the CS–US association survived the degradation manipulation and was retained over an interval of several days.     

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.     

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.     

Conditioned inhibitory effects of discriminated Pavlovian training with food in rats depend on interactions of search modes, related repertoires, and response measures

Like other accounts of conditioned inhibition, behavior systems predicts (and Experiment 1 shows) that during summation and retardation tests, presentation of a negative conditioned stimulus (a CS−) created by discriminative Pavlovian food conditioning will interfere with a focal search response, such as nosing in the feeder. Unlike most other views, behavior systems predicts (and Experiment 2 shows) that the same CS− can potentiate a general search response, like attending to a moving artificial prey stimulus. Contacting the prey stimulus in extinction increased over baseline when a CS- but not a CS Novel preceded it. Experiment 3 showed this effect was not due to unconditioned qualities of the CS−. It appears that the effects of a discriminative CS-depend on the interaction of the training contingency with search modes related to the unconditioned stimulus (US), their perceptual-motor repertoires and environmental support, and the choice of response measure.     

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.     

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.     

Two components of responding in Pavlovian lick suppression

The present research examined the temporal distribution of responding in a lick suppression paradigm. In Experiment 1, rats were trained with either a 30- or a 120-s conditioned stimulus (CS), which was followed either by a footshock (unconditioned stimulus [US]) or nothing. Licking during the CS was suppressed only in the former condition. Suppression was more pronounced early in the CS. In Experiment 2, rats were exposed to two 30-s or two 120-s CSs, with delivery of the shock being contingent on CS1 for half of the animals and on CS2 for the other half. For both the paired and the unpaired conditions, suppression at the beginning of CS1 was observed for all the groups. By discounting the possibility of generalization between CS1 and CS2, it appears that this initial suppression was not a conditioned response to the CS, but an unconditioned one due to mere exposure to the shock US.     

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.     

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).     

Pre-exposure enhances recovery of conditioned responding after extinction

Four experiments used a within-subjects design with rats to study the effects of preexposure on the restoration of fear responses (freezing) to an extinguished conditioned stimulus (CS). In each experiment, rats were preexposed to one CS (A), but not to another (B), and then were exposed to pairings of each of these CSs with an aversive unconditioned stimulus (US). In each experiment, there was less freezing to A than to B across extinction, showing a latent inhibitory effect of preexposure. There was no differential recovery to A and B following either a US reexposure (Experiment 1) or a delay interval (Experiment 2). However, when a delay interval included US reexposure, there was greater recovery to the preexposed CS, A, than to the nonpreexposed CS, B (Experiments 1, 3, and 4). These results suggest that the effects of US reexposure and delay combine to affect recovery from the depressive effects of CS-alone exposure. The results are consistent with the view that US reexposure produces better mediated conditioning of CSs that are strongly associated with the context. The results may additionally reflect an effect of preexposure on the learning produced by extinction.     

Reminder-induced attenuation of the effect of relative stimulus validity

The roles of deficient acquisition and deficient expression of learned information in the effect of relative stimulus validity were examined using rats in a conditioned lick suppression paradigm. Recovery from the effect without further pairings of the conditioned stimulus (CS) and the unconditioned stimulus (US) would favor an interpretation of the relative validity effect based on a latent CS-US association as distinct from a failure to acquire the CS-US association. As a potential recovery manipulation, “reminder” treatments, consisting of the US alone (Experiment 1) or the CS alone (Experiment 2), were administered following relative validity training. In both cases, subjects for which the CS target was of low relative predictive validity exhibited enhanced responding relative to appropriate controls. Additionally, Experiment 2 showed that the amelioration of the relative validity deficit was stimulus specific. Thus, the results of these experiments support previous suggestions that the performance deficit resulting from low relative stimulus validity is due, at least in part, to a failure to express acquired information (Cole, Barnet, & Miller, 1995a).     

Overshadowing and CS duration: counteraction and a reexamination of the role of within-compound associations in cue competition

In the present experiments, we examined the role of within-compound associations in the interaction of the overshadowing procedure with conditioned stimulus (CS) duration, using a conditioned suppression procedure with rats. In Experiment 1, we found that, with elemental reinforced training, conditioned suppression to the target stimulus decreased as CS duration increased (i.e., the CS duration effect), whereas, with compound reinforced training (i.e., the overshadowing procedure), conditioned suppression to the target stimulus increased as CS duration increased. In subsequent experiments, we replicated these findings with sensory preconditioning and demonstrated that extinction of the overshadowing stimulus results in retrospective revaluation with short CSs and in mediated extinction with long CSs. These results highlight the role of the duration of the stimulus in behavioral control. Moreover, these results illuminate one cause (the CS duration) of whether retrospective revaluation or mediated extinction will be observed.     

Bidirectional associations

Associations are usually assumed to have only a predictive directionality, from Event 1 (E1) to Event 2 (E2). However, this unidirectionality, if it really exists, could be specific to the use of conditioned responses as the index of learning (i.e., conditioned responses are ordinarily elicited only when a conditioned stimulus [CS] predicts an unconditioned stimulus [US]). The present research used neutral stimuli, devoid of CS-US directionality, in order to test whether the underlying associative mechanism was unidirectional in nature. We used colors and figures as E1s and E2s. In Experiment 1, human subjects saw the events in the E1→E2 direction during training. In theconsistent group, the same E1 was always followed by the same E2; in theinconsistent group, E1s and E2s were paired randomly. In a subsequent test phase, we presented an E2 and subjects had to judge whether a particular E1 was associated with it. The judgments of the consistent group were higher than those of the inconsistent group, thereby suggesting that the association that the consistent subjects had learned was bidirectional. Experiments 2 and 3 confirmed the results of Experiment 1 while controlling for some alternative interpretations based on the representation-mediated formation of associations.     

Stimulus preexposure speeds or slows subsequent acquisition of associative learning depending on learning test procedures and response measure

Prior exposure to a conditioned stimulus (CS) typically results in latent inhibition—slower acquisition of associative learning about that stimulus in subsequent training. Here, we found that CS preexposure had different effects on the appetitive conditioning of rats with a sucrose unconditioned stimulus (US) depending on training test procedures, the similarity of preexposure and training procedures, and the choice of response measure. Preexposure to a visual or an auditory stimulus produced facilitation of acquisition of food-cup-directed responding when both of those cues were (separately) paired with sucrose delivery in the training test (Experiments 1 and 3). By contrast, the same preexposure procedure resulted in latent inhibition of food-cup learning if the second stimulus in the test phase was of the same modality as the preexposed stimulus (Experiment 2). In Experiment 3, latent inhibition was enhanced if both phases included a single CS or both phases included both auditory and visual CSs, compared to treatments in which only one CS was presented in one phase but two CSs were presented in the other phase. In Experiment 4, preexposure of an auditory cue slowed subsequent learning about it if the context was salient but enhanced learning if the context was of weaker salience. Finally, a measure of general activity revealed latent inhibition after preexposure in all conditions in all 4 experiments. We discuss the results within several classes of latent inhibition theories, none of which provides a comprehensive account.