Stagewise multidimensional visual discrimination by pigeons

We trained six pigeons in a stagewise Multiple Necessary Cues (MNC) go/no‐go task to document the dynamics of discrimination learning involving increasingly complex visual stimuli. The compound stimuli were composed from four dimensions, each of which could assume either of two extreme values or their intermediate value: Shape, Size, Line Orientation, and Brightness. Starting with a stimulus composed entirely from intermediate values, we replaced those values with each of the two extreme dimensional values in four successive stages, thereby increasing the stimulus set from 2 in Stage 1 to 16 in Stage 4. In each stage, only one combination of values signaled food (S⁺), whereas the remaining combinations did not (S^?s). We calculated the rate of pecking during the first 15 s of each stimulus presentation and, in any given stage, training continued until the rate of responding to all of the S^?s was less than 20% of the rate of responding to the S⁺. All pigeons successfully acquired the final discrimination, suggesting that they attended to all of the dimensions relevant for the discrimination. We also replicated the key results of prior MNC studies: (1) the number of extreme dimensional values in each stage was positively related to the amount of training required for pigeons to acquire the discrimination; (2) attentional tradeoffs were most often observed when three or four dimensions were being trained; and (3) throughout training, the number of dimensional values in the S^?s that differed from the S⁺ was positively related to their discriminability from S⁺.     

Repeated acquisition and discrimination reversal in the squirrel monkey (Saimiri sciureus)

Repeated acquisition and discrimination reversal tasks are often used to examine behavioral relations of, respectively, learning and cognitive flexibility. Surprisingly, despite their frequent use in cognitive neuroscience and behavioral pharmacology, variables that control performance under these two tasks have not been widely studied. The present studies were conducted to directly investigate the controlling variables in nonhuman primates. Squirrel monkeys were trained with a touchscreen variant of the repeated acquisition task in which a novel pair of S⁺/S^? stimuli was presented daily. Subjects learned to discriminate the two stimuli (acquisition) and, subsequently, with the contingencies switched (reversal). Results indicate that rates of both acquisition and reversal learning increased across successive sessions, but that rate of reversal learning remained slower than acquisition learning, i.e., more trials were needed for mastery. Subsequent experiments showed this difference between the rate of learning novel discriminations and reversal was reliable for at least 5 days between acquisition and reversal and notwithstanding the interpolation of additional discriminations. Experimental analysis of the S⁺/S^? elements of the tasks revealed that the difference in the rate of learning could not be attributed to a relatively aversive quality of the S^? or to a relatively appetitive quality of the S⁺, but, rather, to contextual control by the S⁺/S^? stimulus complex. Thus, if either element (S⁺ or S^?) of the stimulus complex was replaced by a novel stimulus, the rate of acquisition approximated that expected with a novel stimulus pair. These results improve our understanding of fundamental features of discrimination acquisition and reversal.     

Facilitation of instrumental behavior by a Pavlovian appetitive conditioned stimulus

Three experiments examined appetitive Pavlovian-instrumental interactions by presenting separately trained conditioned stimuli (CSs) during reinforced instrumental responding in rabbits. Intra-oral reinforcement was used to minimize interference from peripheral responses such as magazine approach. In experiment 1, the rabbits were first trained to perform an instrumental head-raising response for sucrose reward. A conditioned jaw movement response was then established to a 2-sec CS by pairing it with sucrose; a control stimulus was unpaired with sucrose. Instrumental responding maintained by a variable-interval 40-sec schedule was enhanced during 10-sec presentations of the paired, but not the unpaired, CS. Responding on a variable-ratio 15 schedule was unaffected except on trials on which the pre-CS baseline response rate was low; in such cases the paired CS caused a long-lasting acceleration of responding. Noncontingent presentation of the sucrose reinforcer itself briefly suppressed responding but had no long-term effect. In Experiment 2, a CS that had been conditioned at a 10-sec duration produced the same pattern of effects as in the first study, indicating that facilitation resulted from CS presentation rather than from the frustrative effects of non-reinforcement of the CS. In Experiment 3 an inhibitory CS blocked facilitation by the excitatory CS but did not itself affect instrumental responding. These results support the view that Pavlovian processes play a positive role in instrumental performance and suggest that previous findings of suppression by a short-duration CS reflect peripheral interference. The dependence of facilitation on the baseline level of responding is discussed in terms of associative and motivational theories of Pavlovian mediation.     

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.     

Determinants of tonic and phasic reactions in transswitching

Forty-eight college students were assigned randomly to four groups in a 2 × 2 factorial arrangement of phasic conditional stimuli (samevs. different) and tonic conditional stimuli (samevs. different) to receive 2 days of classical conditioning with a transswitching procedure. Tonic stimuli were a 5-minute projected white triangle or circle; phasic stimuli were a 5-second red or green square superimposed over the tonic stimuli. There were six tonic stimulus segments each day, separated by 20-second periods of no stimulus, three containing six trials of the phasic stimulus paired with shock and three containing six trials of the phasic stimulus alone, in the counterbalanced order. Tonic responding at the onset of the tonic stimuli or during brief periods following its onset were recorded, along with phasic responses to the phasic stimuli. Responses included magnitude of skin conductance responses, frequency of unelicited skin conductance responses, and tonic heart rate. Both skin conductance measures of responding to the tonic stimuli differentiated significantly between positive and negative tonic segments during Day 2, but only in the group with two different tonic stimuli and one phasic stimulus (“standard” transswitching). This supported the hypothesis that tonic stimulus differentiation would be absent when two different phasic stimuli were present. The heart rate data did not support this hypothesis, showing tonic differentiation in both groups with two tonic stimuli. Phasic differentiation controlled by the different phasic stimuli was observed on Day 1; on Day 2, phasic differentiation was present only in the group with two tonic and one phasic stimuli and the group with one tonic and two phasic stimuli. The results were interpreted to mean that temporal contiguity between the tonic stimuli and shock or no shock is not sufficient to establish tonic response differentiation in transswitching.     

Discriminative conditioning alters food preferences in the leech, Haemopis marmorata

The feeding behavior of the carnivorous leech, Haemopis marmorata, was aversively trained in a discriminative classical conditioning task. Two conditioned stimuli were used: One consisted of a food (chicken or liver) paired with an unconditioned stimulus of quinidine (bitter chemical); the other consisted of the alternate food presented in an unpaired relationship with the quinidine. Training consisted of alternating exposures to the two conditioned stimuli. Testing consisted of the simultaneous presentation of a conditioned stimulus food and a neutral food, beef. The percentages of responding to the conditioned stimuli were tabulated. Haemopis could discriminate between the conditioned stimuli. As a result of pairing a food with quinidine, the leeches selectively reduced their preference for that paired food, while they did not alter their preference for the unpaired food.     

Dopamine D1 and D2 receptors in the nucleus accumbens core and shell mediate Pavlovian-instrumental transfer

Pavlovian stimuli previously paired with food can markedly elevate the rate of food-reinforced instrumental responding. This effect, termed Pavlovian-instrumental transfer (PIT), depends both on general activating and specific cueing properties of Pavlovian stimuli. Recent evidence suggests that the general activating properties of Pavlovian stimuli are mediated by mesoaccumbens dopamine systems; however, the role of NAC dopamine D1 and D2 receptors is still unknown. Here we examined the effects of a selective dopamine D1 and D2 receptor blockade in the shell and core subregion of the NAC on general PIT. Rats were trained to press a single lever for food, and the effect of a single Pavlovian stimulus previously associated with the same food on performance of that lever was measured in extinction. Results reveal that PIT, that is, the increase in instrumental responding during presentation of the Pavlovian stimulus, was reduced by microinjections of the dopamine D1 receptor antagonist SCH-23390 and, less pronounced, by microinjections of the dopamine D2 receptor antagonist raclopride into the NAC core or shell, respectively. Our data suggest that dopamine D1 and D2 receptors in the NAC core and shell mediate the general activating effects of Pavlovian stimuli on instrumental behavior.     

Pavlovian conditioned stimulus effects upon instrumental choice behavior are reinforcer specific

In a transfer-of-control experiment with rats, Pavlovian CSs were tested for the specificity of their effects. The instrumental behavior consisted of a discriminative, conditional two-lever choice task in which qualitatively different appetitive reinforcers were contingent upon the two correct choices. In a Pavlovian phase, subjects experienced conditioning to establish either a CS⁺ or CS^? for one reinforcer or a CS⁺ or CS^? for the other reinforcer. Finally, in a test, these CSs were presented when there was the opportunity to make choice responses. The CS⁺s evoked choices of the lever which had eventuated in the reinforcer that had served as the Pavlovian US, while the CS^?s showed only a slight tendency to evoke the other choice responses. When the CSs were compounded with the original S^Ds, the CS⁺s had little effect upon the vigor of responding while the CS^?s reduced the vigor of responding to the S^D for the reinforcer that was the same as the US used in establishing the CS^?. The results are discussed in terms of associative mediational theory and the reinforcer specificity of Pavlovian conditioned excitation and inhibition.     

Effects of long- and variable-duration signals for food on activity,instrumental responding,and eating

Incentive-motivation theories typically assume that the conditioning of appetitive motivation involves the same parameters as Pavlovian salivary conditioning. In contrast, the Soltysik-Konorski model asserts that drive is inhibited by stimuli closely associated with food (salivary CSs) and augmented by stimuli more loosely associated with food (long and variable CS-US interval). Experiment 1 examined this latter proposition. Sixty-four rats were given extensive exposure to each of four environmental CSs, two while hungry and two while satiated. Within each deprivation condition, food was given 30–300 sec after placement of the rats in one environment, and was not given in the other environment. Performance on three separate measures—activity, lever-pressing, and food consumption—was higher in the environments previously associated with food. Experiment 2 examined the effects of discrete stimuli presented in advance of eating; in accord with the results of Experiment 1, food consumption was greater after a stimulus (1- to 9-min duration) previously paired with food than after no stimulus or after a stimulus unpaired with food. The overall results indicate (a) that stimuli associated with food become capable of facilitating a variety of food-directed behaviors, possibly via the conditioning of a common appetitive system, and (b) that a close association between the stimuli and food is not essential for such conditing to occur.     

Asymmetrical stimulus generalization following differential fear conditioning

Rodent ultrasonic vocalizations (USVs) are ethologically critical social signals. Rats emit 22 kHz USVs and 50 kHz USVs, respectively, in conjunction with negative and positive affective states. Little is known about what controls emotional reactivity to these social signals. Using male Sprague–Dawley rats, we examined unconditional and conditional freezing behavior in response to the following auditory stimuli: three 22 kHz USVs, a discontinuous tone whose frequency and on–off pattern matched one of the USVs, a continuous tone with the same or lower frequencies, a 4 kHz discontinuous tone with an on–off pattern matched to one of the USVs, and a 50 kHz USV. There were no differences among these stimuli in terms of the unconditional elicitation of freezing behavior. Thus, the stimuli were equally neutral before conditioning. During differential fear conditioning, one of these stimuli (the CS⁺) always co-terminated with a footshock unconditional stimulus (US) and another stimulus (the CS⁻) was explicitly unpaired with the US. There were no significant differences among these cues in CS⁺-elicited freezing behavior. Thus, the stimuli were equally salient or effective as cues in supporting fear conditioning. When the CS⁺ was a 22 kHz USV or a similar stimulus, rats discriminated based on the principal frequency and/or the temporal pattern of the stimulus. However, when these same stimuli served as the CS⁻, discrimination failed due to generalization from the CS⁺. Thus, the stimuli differed markedly in the specificity of conditioning. This strikingly asymmetrical stimulus generalization is a novel bias in discrimination.     

Pavlovian processes do mediate control of the “advance” response strategy

Two parallel experiments used pigeons in a transfer of control design to determine the basis of the transfer of the use of the “advance strategy” across successive discriminations (W. K. Honig & H. Lindsay, Learning and Motivation, 1975, 6, 157–178). Pigeons were first trained under instrumental contingencies to use an “advance” response to maximize their exposure to S^D and minimize their exposure to S^Δ. Then orthogonal stimuli (key lights in Experiment 1 and diffuse illuminations in Experiment 2) were independently established through separate discriminative Pavlovian procedures as CS+ and CS? or as CS^o's. Later in special tests, it was demonstrated that the associative values of the Pavlovian CS's were a major factor governing the use of the advance strategy: only pigeons exposed to $\text{CS+}\text{CS?}$ used the advance response to regulate their exposure to either CS. Although these experiments showed that the necessary and sufficient conditions for utilization of the advance strategy are the Pavlovian associative values of the discriminative stimuli, opportunity for differential “operant” responding during the discriminative stimuli seems to be a contributor to the optimal use of the advance response across successive discriminations.     

Pavlovian heart rate and jaw movement conditioning in the rabbit: effects of medial prefrontal lesions

An experiment was conducted in which jaw movements (JM) and heart rate (HR) were concomitantly assessed in rabbits during simple Pavlovian conditioning. A 2-s 1200-Hz tone was the conditioned stimulus (CS) and an intraoral 1-cc pulse of 0.5 M sucrose-water solution was the unconditioned stimulus (US). Sham and medial prefrontal (mPFC)-lesioned animals received paired CS/US training with a 70- to 75-dB CS and were compared with sham- and mPFC-lesioned animals that received explicitly unpaired CS/US presentations. The percentages of JM CRs were significantly greater in the paired than the unpaired groups, but mPFC lesions had no effect on this measure. Conditioned HR decelerations occurred only in the paired groups and then only during the first session of training. Moreover, these CS-evoked cardiac decelerations were somewhat attenuated by the mPFC lesion. CS-evoked HR accelerations, which were significantly greater in unpaired than in paired animals, occurred during the four subsequent sessions. These results suggest that a CS-evoked cardioinhibitory process, mediated by the mPFC, is engendered by Pavlovian appetitive conditioning, as has been previously demonstrated for aversive conditioning. However, during JM conditioning these inhibitory changes are quickly replaced by tachycardia, possibly related to increased nonspecific somatomotor activity, since the tachycardia was somewhat greater in the unpaired animals.     

Complex Visual Learning by Rats

Rats’ learning about visual patterns was studied in a computerized Y-maze where wide-angle stimuli were viewed from a distance. Many patterns were available; some were spatially complex and others were more homogeneous figures. Experiment 1 used a discrimination paradigm in which a single S⁺could be paired with any one of 15 different S⁻s. Hooded rats learned successively six such discrimination problems. Their learning rate improved across the series, and comparison with controls suggested that the learning-set did not merely reflect simple habituation. Experiments 2 and 3 employed Dark Agouti rats, again learning many discrimination problems. Each problem comprised a constant stimulus which was paired with stimuli which varied in trial-unique fashion. The version in which the constant stimulus was nonrewarded (S⁻) and the varying stimuli rewarded was performed better than the converse, constant S⁺and varying nonrewarded, reflecting rats’ preference for relatively unfamiliar stimuli. In the constant S⁻task, rats showed substantial within-problem learning when three novel problems were given per day for 20 trials each. Rats are capable of rapid learning about complex visual displays if we engage their natural dispositions to use vision for distal stimuli and to approach relatively unfamiliar cues.     

The development of an attentional bias for angry faces following Pavlovian fear conditioning

Although it is well documented that fear responses develop following aversive Pavlovian conditioning, it is unclear whether fear learning also manifests in the form of attentional biases for fear-related stimuli. Boschen, Parker, and Neumann (Boschen, M. J., Parker, I., & Neumann, D. L. (2007). Changes in implicit associations do not occur simultaneously to Pavlovian conditioning of physiological anxiety responses. Journal of Anxiety Disorders, 21, 788-803.) showed that despite the acquisition of differential skin conductance conditioned responses to angry faces paired (CS+) and unpaired (CS−) with an aversive shock, development of implicit associations was not subsequently observed on the Implicit Association Test. In the present study, participants (N = 76) were assigned either to a Shock or NoShock group and completed a similar aversive Pavlovian conditioning procedure with angry face CS+ and CS− stimuli. Participants next completed a visual probe task in which the angry face CS+ and CS− stimuli were paired with angry face control stimuli and neutral faces. Results confirmed that differential fear conditioning was observed in the Shock group but not in the NoShock group, and that the Shock group subsequently showed a selective attentional bias for the angry face CS+ compared with the CS− and control stimuli during the visual probe task. The findings confirm the interplay between learning-based mechanisms and cognitive processes, such as attentional biases, in models of fear acquisition and have implications for treatment of the anxiety disorders.     

Short-term memory for "surprising" versus "expected" unconditioned stimuli in Pavlovian conditioning.

Rabbits were trained via a Pavlovian eyelid-conditioning regime to respond differentially to a conditioned stimulus (CSR) depending on whether or not that stimulus was preceded several seconds earlier by a preparatory stimulus consisting of an electric shock otherwise employed as an unconditioned stimulus (UCS). Following training, differential responding to CSR varied systematically with the interval between the preparatory stimulus and CSR, consistent with assumptions concerning decay of information in short-term memory. The major finding, however, involved the test effects of announcing the preparatory stimulus by either a conditioned stimulus with which it had otherwise been paired (CS+) or a conditioned stimulus with which it had otherwise been systematically unpaired (CS-). The greater differential responding to the following CSR in the latter case is taken to support the notion that a "surprising" UCS is more likely to be retained (rehearsed) in short-term memory than is an "expected" UCS.     

Automated assessment of conditioning parameters for context and cued fear in mice

A behavioral technique often used to evaluate the cognitive performance of rats and mice is the fear conditioning paradigm. During conditioned fear experiments, freezing responses shown by rodents after exposure to environmental stimuli previously paired to an aversive experience provide a behavioral index of the animal's associative abilities. The present study examined the ability of a computer-controlled automated Freeze Monitor system for recording immobility behavior in mice. The sensitivity of the automated procedure to detect group differences caused by the application of various training protocols was also evaluated. Statistical analyses revealed significant positive correlations between immobility scores obtained with the automated apparatus and hand-scored data collected by a continuous or a time-sampling method. Behavioral patterns recorded by the computerized system were very similar to those obtained by the hand-scoring methods adopted. In particular, during context testing, exposure to environmental stimuli previously paired with a mild foot shock (unconditioned stimulus [US]) evoked increased immobility behavior in mice conditioned with the US compared with levels of immobility displayed by mice previously confined to the same contextual stimuli without receiving the US. Moreover, although during conditioned stimulus (CS) testing, mice previously exposed to the US displayed high levels of immobility when confined to environmental cues much different from those paired with the US (contextual fear generalization), both hand-scored and automated results revealed the effect of CS–US pairing (increased immobility) only in mice trained to associate the two stimuli (paired group) but not in mice exposed to both CS and US separated by a 40-sec time interval (unpaired group) or in mice receiving only the US (US group) during conditioning sessions. Overall, the results show associative conditioning measured in an automated apparatus and highlight the utility of obtaining both latency as well as beam interruption parameters.     

The role of predrug signals in morphine analgesic tolerance: support for a Pavlovian conditioning model of tolerance

According to a model of morphine tolerance, which emphasizes Pavlovian conditioning principles, tolerance results from an association between predrug environmental cues and the systemic effects of the drug. To assess this model, groups of rats were administered morphine on either three or nine occasions, with a complex environmental stimulus either paired or not paired with each injection. Control groups had equivalent experience with the environmental cue and injection procedure, but the injected substance was physiological saline. Subsequently, the analgesic effect of the opiate was tested in all subjects following administration of the drug in conjunction with the environmental cue. As expected on the basis of the conditioning model of tolerance, subjects with a pretest history of paired morphine administrations displayed analgesic tolerance, but subjects with a pretest history of unpaired administration displayed no evidence of such tolerance. The results suggest that prior demonstrations that the display of morphine tolerance is specific to the drug administration environment may be readily interpreted by a conditioning analysis of tolerance.     

Modeling Hierarchical Versus Random Exposure Schedules in Pavlovian Fear Extinction: No Evidence for Differential Fear Outcomes

In exposure therapy, the client can either be confronted with the fear-eliciting situations in a hierarchical way or in a random way. In the current study we developed a procedure to investigate the effects of hierarchical versus random exposure on long-term fear responding in the laboratory. Using a fear conditioning procedure, one stimulus (CS +) was paired with an electric shock (US), whereas another stimulus was not paired with the shock (CS-). The next day, participants underwent extinction training including presentations of the CS-, CS + and a series of morphed stimuli between the CS- and CS +. In the hierarchical extinction condition (HE; N = 32), participants were first presented with the CS-, subsequently with the morph most similar to the CS-, then with the morph most similar to that one, and so forth, until reaching the CS +. In the random extinction condition (RE; N = 32), the same stimuli were presented but in a random order. Fear responding to the CS +, CS- and a new generalization stimulus (GS) was measured on the third day. Higher expectancy violation, t(62) = -2.67, p = .01, physiological arousal, t(62) = -2.08, p = .04, and variability in US-expectancy ratings, t(62) = -2.25, p = .03, were observed in the RE condition compared to the HE condition, suggesting the validity of this novel procedure. However, no differences between the RE and HE condition were found in fear responding as tested one day later, F(1, 62) < 1. In conclusion, we did not find evidence for differential long-term fear responding in modeling hierarchical versus random exposure in Pavlovian fear extinction.     

Constraint, alcoholism, and electrodermal response in aversive classical conditioning and mismatch novelty paradigms

This study investigated whether low levels of the personality trait of constraint and early-onset alcoholism would be associated with deficits in aversive conditioning and smaller responses to novelty in a stimulus mismatch protocol. Personality traits (constraint and socialization) and skin conductance responses (SCRs) during conditioning and novelty paradigms were assessed in alcoholics (n=41) and non-alcoholics (n=32). The conditioning protocol involved measuring SCRs after conditioned stimuli (CS+: tones) paired with shock, CS− tones unpaired with shock, and CS+ probes unpaired with shock. The mismatch protocol involved measuring SCRs to auditory stimuli consisting of a series of 5 pure tones of the same pitch followed a shorter white noise stimulus (the novel stimulus). Contrary to the hypothesis, alcoholics did not differ from non-alcoholics in SCRs to CS+ probes or on the mismatch measure (SCR novel tone—SCR to 5^th tone). Higher levels of constraint and self-reports of fear during conditioning were associated with smaller responses to both the CS+ probes and the CS− tones as well as the mismatch measure within non-alcoholics, but not within alcoholics. In alcoholics, low constraint was associated with greater habituation to CS+ probes, and poor differential conditioning on measures of change across trials in SCR to CS+ probes and CS− stimuli. The results suggest that different processes influence levels of constraint in non-alcoholics and alcoholics. The data indicate that low constraint in non-alcoholics is associated with allocating fewer processing resources to potentially significant stimuli, rather than being associated with a specific deficit in aversive conditioning per se.     

Developmental changes in eyeblink conditioning and neuronal activity in the pontine nuclei

Neuronal activity was recorded in the pontine nuclei of developing rats during eyeblink conditioning on postnatal days 17–18 (P17–P18) or P24–P25. A pretraining session consisted of unpaired presentations of a 300-msec tone conditioned stimulus (CS) and a 10-msec periorbital shock unconditioned stimulus (US). Five paired training sessions followed the unpaired session, consisting of 100 trials of the CS paired with the US. The rats trained on P24–P25 exhibited significantly more conditioned responses (CRs) than the rats trained on P17–P18, although both groups produced CRs by the end of training. Ontogenetic increases in pre-CS and stimulus-elicited activity in the pontine nuclei were observed during the pretraining session and after paired training. The activity of pontine units was greater on trials with CRs relative to trials without CRs in rats trained on P24–P25, but almost no CR-related modulation was observed in the pontine units of rats trained on P17–P18. The findings indicate that pontine neuronal responses to the CS and modulation of pontine activity by the cerebellum and red nucleus undergo substantial postnatal maturation. The developmental changes in pontine neuronal activity might play a significant role in the ontogeny of eyeblink conditioning.