Acquisition of the classically conditioned eyeblink response in humans over the life span

Human subjects ranging in age from 18 to 85 years underwent classical conditioning of the eyeblink response to a tone conditioned stimulus (CS) and an air-puff unconditioned stimulus (UCS). There was a decline in percentage of conditioned responses with age. This decline was most noticeable in subjects over age 50. These conditioning deficits were not due to age-related changes in sensitivity to the tone CS or the air-puff UCS, nor could the conditioning deficits be attributed to an age-related decline in general cognitive abilities or to changes in spontaneous blink rates. The results are discussed in terms of using the classically conditioned eyeblink in humans in conjunction with the classically conditioned nictitating membrane response in rabbits as a model system for studying the neurobiology of age-related conditioning deficits.     

Pavlovian second-order conditioned analgesia

Three experiments with rat subjects assessed conditioned analgesia in a Pavlovian second-order conditioning procedure by using inhibition of responding to thermal stimulation as an index of pain sensitivity. In Experiment 1, rats receiving second-order conditioning showed longer response latencies during a test of pain sensitivity in the presence of the second-order conditioned stimulus (CS) than rats receiving appropriate control procedures. Experiment 2 found that extinction of the first-order CS had no effect on established second-order conditioned analgesia. Experiment 3 evaluated the effects of post second-order conditioning pairings of morphine and the shock unconditioned stimulus (US). Rats receiving paired morphine-shock presentations showed significantly shorter response latencies during a hot-plate test of pain sensitivity in the presence of the second-order CS than did groups of rats receiving various control procedures; second-order analgesia was attenuated. These data extend the associative account of conditioned analgesia to second-order conditioning situations and are discussed in terms of the mediation of both first- and second-order analgesia by an association between the CS and a representation or expectancy of the US, which may directly activate endogenous pain inhibition systems.     

An analysis of second-order autoshaping

Three mechanisms can explain second-order conditioning: (1) The second-order conditioned stimulus (CS2) could activate a representation of the first-order conditioned stimulus (CS1), thereby provoking the conditioned response (CR); The CS2 could enter into an excitatory association with either (2) the representation governing the CR, or (3) with a representation of the reinforcer evoked by the CS1. A series of experiments using second-order autoshaping with birds was performed to examine these possibilities. Following second-order autoshaping, birds’ responding to the CS2 was found to be unaffected by extinction of the CS1, a result interpreted as showing mechanism (1) to be unimportant and implicating either one or both of the other mechanisms. The CS2 was also shown to provide the birds with specific sensory information about the reinforcer, a pattern of results uniquely predicted by mechanism (3). Implications for the understanding of second-order conditioning and for the SOP model (Wagner, 1981) of associative learning are discussed.     

Autoshaping in the rat: The effects of localizable visual and auditory signals for food

Two experiments investigated autoshaping in rats to localizable visual and auditory conditioned stimuli predicting response-independent food. In Experiment 1 considerable conditioned-stimulus approach behavior was generated by a localizable visual conditioned stimulus that was situated approximately 35 cm from the food tray. Using the same apparatus in Experiment 2 we found that the conditioned-stimulus approach was generated only to a visual conditioned stimulus and not to a localizable auditory conditioned stimulus even though subjects (1) could discriminate presentations of the auditory conditioned stimulus, (2) had associated it with food, (3) could localize it, and (4) would approach the auditory stimulus if this behavior constituted an instrumental response to food. The predominant conditioned responses to the auditory stimuli were goal tracking (entering the food tray) and orienting towards the food-paired conditioned stimulus by head turning and rearing and turning. These results imply that rats do not invariably approach a localizable appetitive Pavlovian conditioned stimulus but that stimulus-approach responses depend on the nature and modality of the conditioned stimulus.     

Cardiac and respiratory conditioning, differentiation, and extinction in the pigeon

Paired light and foot shock in 12 pigeons rapidly developed acceleratory heart and respiratory conditioned responses. Sensitization and stimulus control birds did not condition. When a different colored stimulus light that was not reinforced was mixed in a random series with the reinforced light, rapid differentiation of both heart and respiratory responses occurred. In most cases neither heart nor respiratory conditioned responses could be extinguished by 100 non-reinforced presentations of the conditioned stimulus.     

Electrodermal conditioning to potentially phobic stimuli: Effects of instructed extinction

It was hypothesized that electrodermal responses to potentially phobic stimuli “conditioned” through verbal threats about an aversive UCS should be equally resistant to instructed extinction as the responses obtained by actual CS-UCS pairings. Two groups of human subjects were exposed to pictures of either snakes and spiders (phobic CSs), or circles and triangles (neutral CSs) in a differential Pavlovian conditioning paradigm. Changes in skin conductance were recorded. Half of the subjects in each group were threatened with a shock-UCS while the other half were given shock-reinforced CS presentations. At the onset of extinction, all subjects in each of the four groups were informed that no more UCSs were to be delivered, and the shock electrodes were removed. All groups showed evidence of conditioning during acquisition. During extinction there was an immediate drop in responding in the two neutral groups, whereas the two phobic groups showed reliable evidence of resistance to extinction, with no differences between the threatened- and the CS-UCS-group. The observed resistance to extinction found in the phobic groups implies a similarity to the irrationality of real-life phobias. Furthermore, the data are in accordance with analysis of electrodermal fear-conditioning as a case of prepared learning.     

Conditioning in human opiate addicts

Eight volunteers maintained on daily methadone participated in a classical conditioning procedure to determine which if any of the elements of narcotic withdrawal could be conditioned. The unconditioned stimulus was the injection of a small dose of naloxone. The unconditioned response was a brief precipitated withdrawal syndrome. The conditioning stimulus was a tone, odor, and injection of saline. Conditioning was successful in the pilot study in 5 of 8 subjects. The conditioned response consisted of tearing, yawning, lacrimation, systolic blood pressure increase, respiratory irregularities and subjective feelings of narcotic withdrawal sickness (nausea, muscle aches, chills). A second group of 8 subjects showed, in addition to the above, evidence of conditioning of heart rate, respiratory rate and skin temperature decrease. These laboratory findings support the clinical reports of a conditioned withdrawal syndrome and suggest ways to improve treatment results by detecting and extinguishing or modifying conditioned responses.     

Blocking of inhibitory conditioning within a serial conditioned stimulus-conditioned inhibitor compound: Maintenance of acquired behavior without an unconditioned stimulus

Well-trained classically conditioned stimuli, presented unreinforced, were protected from extinction when they were followed by a signal of the omission of the reinforcer (conditioned inhibitor Konorskian type) in eight cats. An aversive classical conditioning paradigm with shock as the reinforcer was used. Of several behavioral (leg flexion, vocalization) and organismic arousal (heart rate, respiration rate, respiration amplitude) measures of conditioned responses, the respiration amplitude changes were found to be most informative for the continuous assessment of elicited arousal of low and medium intensity. In all subjects conditioned stimuli presented during extinction in serial compound with the conditioned inhibitor elicited larger responses than did conditioned stimuli presented alone during extinction. The mechanism of protection from extinction in a paradigm in which the elicitor of learned behavior occurs prior to the conditioned inhibitor provides the organism with the mechanism for the maintenance of learned behavior in the absence of a reinforcer.     

Brain asymmetry and autonomic conditioning: Sensitization control

In the present experiment we report effects of cerebral asymmetry, or laterality, during classical conditioning to facial emotional stimuli. Twenty-five female subjects were presented with slides of a happy face in one visual half-field (VHF), and simultaneously a slide of an angry face in the other VHF, followed by shock as the unconditioned stimulus (UCS). To control for effects of sensitization, a new stimulus, never associated with the UCS, was introduced in the extinction phase. Dependent measures were phasic heart rate responses (HR) and skin conductance responses (SCR). The HR results showed a significant right hemisphere effect for the CS-UCS association, that was not attributable to UCS sensitization. No significant effects were found for the SCRs. The basic HR finding was a right hemisphere superiority for learning of a conditioned association.     

Spatial contiguity facilitates Pavlovian second-order conditioning

Two experiments investigated the effects of spatial contiguity upon the formation of second-order conditioning in pigeon subjects. Experiment 1 used an autoshaping procedure to pair two visual stimuli, S2 and S1, after S1 had previously been paired with food. The resulting second-order conditioning of S2 was superior when both stimuli appeared on the same response key within a trial, compared with their appearing on different keys. Experiment 2 found a similar importance of spatial contiguity between S2 and S1 in a conditioned suppression paradigm. In addition, consistently presenting S2 and S1 in the same spatial location produced superior conditioning compared with varying their spatial relation from trial to trial. The design of these experiments was such as to imply that spatial contiguity facilitates performance by improving the formation of association rather than by promoting stimulus generalization or pseudoconditioning. Moreover, the observation of a facilitative effect of spatial contiguity between S1 and S2 in two different paradigms that use qualitatively different unconditioned stimuli and evoke different responses implies some generality for these findings. Consequently, these results suggest that spatially contiguous stimuli are especially associable in Pavlovian conditioning paradigms.     

Excitatory strength of expressive faces: effects of happy and fear expressions and context on the extinction of a conditioned fear response

In a recent study, Orr and Lanzetta (1984) showed that the excitatory properties of fear facial expressions previously described (Lanzetta & Orr, 1981; Orr & Lanzetta, 1980) do not depend on associative mechanisms; even in the absence of reinforcement, fear faces intensify the emotional reaction to a previously conditioned stimulus and disrupt extinction of an acquired fear response. In conjunction with the findings on acquisition, the failure to obtain extinction suggests that fear faces have some of the functional properties of "prepared" (fear-relevant) stimuli. In the present study we compared the magnitude of conditioned fear responses to happy and fear faces when a potent danger signal, the shock electrodes, are attached or unattached. If fear faces are functionally analogous to prepared stimuli, then, even in the absence of veridical support for an expectation of shock, they should retain excitatory strength, whereas happy faces should not. The results are consistent with this view of fear expressions. In the absence of reinforcement, and with shock electrodes removed, conditioned fear responses and basal levels of arousal were of greater magnitude for the fear-face condition than for the happy-face condition.     

Extinction of conditioned sexual responses in male Japanese quail (Coturnix japonica): role of species-typical cues

The authors examined how a conditioned stimulus (CS) that included species-typical cues affected the acquisition and extinction of conditioned sexual responses in male quail (Coturnix japonica). Some subjects were conditioned with a CS that supported sexual responses and included a taxidermic head of a female quail. Others were conditioned with a similar CS that lacked species-typical cues. Pairing the CSs with access to live females increased CS-directed behavior, with the head CS eliciting significantly more responding than the no-head CS. Responding to the head CS persisted during the 42-day, 126-trial extinction phase; responses to the no-head CS extinguished. Responding declined when the cues were removed or the subjects were sexually satiated. Possible functions and mechanisms of these effects are discussed.     

Conditioned reinforcement: Experimental and theoretical issues

The concept of conditioned reinforcement has received decreased attention in learning textbooks over the past decade, in part because of criticisms of its validity by major behavior theorists and in part because its explanatory function in a variety of different conditioning procedures has become uncertain. Critical data from the major procedures that have been used to investigate the concept (second-order schedules, chain schedules, concurrent chains, observing responses, delay-of-reinforcement procedures) are reviewed, along with the major issues of interpretation. Although the role played by conditioned reinforcement in some procedures remains unresolved, the results taken together leave little doubt that the underlying idea of conditioned value is a critical component of behavior theory that is necessary to explain many different types of data. Other processes (marking, bridging) may also operate to produce effects similar to those of conditioned reinforcement, but these clearly cannot explain the full domain of experimental effects ascribed to conditioned reinforcement and should be regarded as complements to the concept rather than theoretical competitors. Examples of practical and theoretical applications of the concept of conditioned reinforcement are also considered.     

Second-order conditioning of Pavlovian conditioned inhibition

Two experiments are reported which investigated the conditioning of inhibition to a neutral stimulus as a result of its repeated pairing with a previouslyconditioned inhibitor. Both experiments employed a conditioned suppression technique with rat subjects. Experiment 1 detected the second-order inhibition through the retarded acquisition of concurrently administered excitatory conditioning. Experiment 2 found similar retardation in the acquisition of excitatory fear conditioning following previous second-order conditioning of inhibition to the stimulus. Implications are discussed for theories of the nature of inhibition and for second-order conditioning as an assessment technique.     

Postnatal development of heart rate patterns elicited by an aversive CS and US in cats

Heart rate and motor responses were recorded in cats of different ages during classical conditioning. A deceleratory-acceleratory heart rate pattern observed during the CS-US interval in one and four-week-old kittens is an alpha conditioned response, a potentiated original response to the CS. At eight weeks of age two new distinct patterns of pure acceleration or pure deceleration are acquired during conditioning and in the absence of motor learning. At 12 weeks of age and in adult subjects, heart rate patterns during the CS-US interval become more complex and conditioned motor responses can be observed. A covariance of HR acceleration and motor responses during the CS-US interval is absent in eight-week-old subjects, but quite high in 12-week-old subjects and adult cats. The data are interpreted as suggesting separate elicitatory mechanisms of HR and motor responses which may show synchrony later in ontogeny.     

Classical skin conductance response conditioning: effects of intermittent reinforcement and information about schedule contingencies.

Skin conductance responses were differentially conditioned using reinforcement schedules of 25%, 50%, 75%, and 100%, manipulated between subjects. Half of the subjects were informed about schedule contingencies, and half were uninformed. The interstimulus interval was 6 sec. Discrimination of first-interval responses (1.0-3.5 sec after conditioned stimulus [CS] onset) by informed subjects did not vary with the ratio variable, but that by uninformed subjects improved with increasing reinforcement ratio because of diminished response levels to the nonreinforced CS (CS-). Discrimination of second-interval responses (3.6-7.0 sec after CS onset) improved as a function of increasing reinforcement ratio because of elevated response levels to the reinforced CS (CS+), but the effect was not persistent across trials in informed subjects. Performance in the first and second intervals did not reflect sequential increments and decrements as a function of reinforced and nonreinforced trials. Third-interval responses (7.1-9.9 sec after CS on nonreinforced trials) were not affected by schedule manipulations, but unconditioned responses diminished with increasing reinforcement ratio. Information about schedule contingencies led to superior discrimination of first-, second-, and third-interval responses and to suppression of unconditioned responses.     

Acquisition of instrumental conditioned reinforcement is resistant to the devaluation of the unconditioned stimulus

The associative mechanisms responsible for the efficacy of Pavlovian stimuli during first- and second-order conditioning have been extensively studied, but little is known about the representations underlying instrumental conditioned reinforcement. The present study investigated the associative structure underlying conditioned reinforcement, by employing an unconditioned stimulus (US) devaluation procedure on a commonly used instrumental task: the acquisition of a new response with conditioned reinforcement. Whilst US-directed behaviour was abolished following devaluation, the conditioned stimulus acting as a conditioned reinforcer supported the acquisition of instrumental responding. In this preparation then, the conditioned reinforcer appears to be impervious to devaluation of its associated US, suggesting that the underlying representation maintaining behaviour is independent of the current value of the US and may reflect the activation of a central appetitive motivational state.     

The representation of the reinforcer and the force of the pigeon's keypeck in first and second-order conditioning

The aim of this study was to determine whether reinforcer-specific conditioned responding would occur in a situation in which responding was not thought to be mediated by a representation that encodes information about the specific properties of the reinforcer. The force of the pigeon's keypeck was monitored during first and second-order conditioning with either food or water as the unconditioned stimulus (US). Each pigeon was trained with four different stimuli: a first-order cue predicting that responding would be reinforced with grain (S1f), a first-order cue predicting water as the reinforcer (S1w), a second-order stimulus predicting the S1f (S2f), and a second-order cue predicting the S1w (S2w). Following conditioning, the pigeons were selectively satiated with one of the two reinforcers and presented with the first and second-order cues in an extinction test. At the end of training, the pigeon's keypecks were less forceful to the S1w than to the S1f. There was not, however, a reliable difference between the force of the pecks to the S2f and the S2w. These force differences are consistent with the conclusion that the topography of the keypecks was systematically related to the nature of the primary reinforcer during first but not during second-order conditioning. The results from the selective satiation test are difficult to interpret. There was no evidence to indicate that second-order responding was mediated by a detailed representation of the primary US, but a detailed representation of the reinforcer may have been mediating first-order responding. Taken together, these findings are consistent with the view that a representation of the reinforcer is an important determinant of the topography of conditioned responding.     

Cognitive factors in the concurrent differential conditioning of eyelid and skin conductance responses

The objectives of the study were to (1) determine if differential conditioning of eyelid responses ocurs only among subjects who accurately report knowledge of the CS-UCS relations, (2) assess whether differentially conditioned eyelid responding occurs only after the initial accurate interrial report, and (3) explore characteristics of differentially conditioned skin conductance responses (SCRs) obtained in an eyelid conditioning paradigm, with special attention to whether eyelid and SCR conditioning are similarly related to the subjects’ knowledge of the stimulus relations. Fifty-one male subjects were administered 80 differential eyelid conditioning trials, the CSs consisting of the 1,200-msec illumination of slides containing either grammatically correct or grammatically incorrect phrases. Significant differential SCR and eyelid conditioning was obtained for both V- and C-form eyelid responders, but only for subjects who accurately reported the CS-UCS relations. The initial appearance of SCR and eyelid differentiation was related within the trial sequence to subjects’ recognition of the stimulus relations. Eyelid and SCR conditioning were related similarly to knowledge of the stimulus relations, the exception being that subjects who recognized the stimulus relations late in the trial sequence did not develop reliable eyelid differentiation.     

Hemispheric processing differences revealed by differential conditioning and reaction time performance.

Two different experimental procedures were used to examine (a) information-processing differences between two groups of subjects (Cs versus Vs) identified by the form of their conditioned eyeblinks; (b) information-processing differences between the right and left cerebral hemispheres; and (c) parallels between hypothesized C-V differences and right-left hemisphere differences. In the first experiment, the evocative command words BLINK and DON'T BLINK served as positive and negative conditioned stimuli. It was found that Vs gave more conditioned eyeblinks than Cs and that differential eyelid conditioning of Vs more than Cs was influenced by the semantic content of the stimuli. More importantly, the conditioning performance of Cs was more influenced by the semantic attributes of the stimuli when they were presented directly to the right visual field (left hemisphere) than when they were presented directly to the left visual field (right hemisphere). In contrast, the conditioning performance of Vs was equally influenced by the semantic attributes regardless of which hemisphere received direct stimulation. A second experiment was designed to determine whether such hemisphere-of-presentation differences for Cs versus Vs could also be obtained in a very different task. Subjects classified as Cs or Vs during a differential eyelid conditioning task then performed two same-different reaction time (RT) tasks that required discrimination of complex polygons in one case and the names of letters in another. On each RT trial both stimuli of a pair appeared briefly either in the center, left, or right visual field. For both Cs and Vs RTs to complex polygon pairs averaged 20 msec faster on left visual field trials than on right visual field trials, consistent with current hypotheses about right-hemisphere specialization for visuospatial processing. In contrast, the results for letter pairs generally confirmed the C-V differences found in Experiment 1. That is, the right visual field (left-hemisphere) advantage for these verbal stimuli was once again larger for Cs than for Vs. The present results suggest that the two groups of subjects (Cs versus Vs) differ qualitatively in the modes of information processing that they typically employ. The results also suggest that these different modes of processing are related to aspects of cerebral hemisphere organization and that even right-handed individuals may differ from each other in the extent to which each cerebral hemisphere is mobilized for a given experimental task. Such individual differences must be incorporated into both models of classical eyelid conditioning and models of cerebral hemisphere specialization.