Human Aversive Conditioning Research Articles

Neurophysiological and Autonomic Dynamics of Threat Processing During Sustained Social Fear Generalization.

Survival in dynamic environments requires that organisms learn to predict danger from situational cues. One key facet of threat prediction is generalization from a predictive cue to similar cues, ensuring that a cue-outcome contingency is applied beyond the original learning environment. Generalization has been observed in laboratory studies of aversive conditioning: behavioral and physiological processes generalize responses from a stimulus paired with threat (the CS+) to unpaired stimuli, with response magnitudes varying with CS+ similarity. In contrast, work focusing on sensory responses in visual cortex has found a sharpening pattern, in which responses to stimuli closely resembling the CS+ are maximally suppressed, potentially reflecting lateral inhibitory interactions with the CS+ representation. Originally demonstrated with simple visual cues, changes in visuocortical tuning have also been observed in threat generalization learning across facial identities. It is unclear to what extent these visuocortical changes represent transient or sustained effects and if generalization learning requires prior conditioning to the CS+. The present study addressed these questions using EEG and pupillometry in an aversive generalization paradigm involving hundreds of trials using a gradient of facial identities. Visuocortical ssVEP sharpening occurred after dozens of trials of generalization learning without prior differential conditioning, but diminished as learning continued. By contrast, generalization of alpha power suppression, pupil dilation, and self-reported valence and arousal was seen throughout the experiment. Findings are consistent with threat processing models emphasizing the role of changing visucocortical and attentional dynamics when forming, curating, and shaping fear memories as observers continue learning about stimulus-outcome contingencies.

Human Sensory Cortex Contributes to the Long-Term Storage of Aversive Conditioning.

Growing animal data evince a critical role of the sensory cortex in the long-term storage of aversive conditioning, following acquisition and consolidation in the amygdala. Whether and how this function is conserved in the human sensory cortex is nonetheless unclear. We interrogated this question in a human aversive conditioning study using multidimensional assessments of conditioning and long-term (15 d) retention. Conditioned stimuli (CSs; Gabor patches) were calibrated to differentially activate the parvocellular (P) and magnocellular (M) visual pathways, further elucidating cortical versus subcortical mechanisms. Full-blown conditioning and long-term retention emerged for M-biased CS (vs limited effects for P-biased CS), especially among anxious individuals, in all four dimensions assessed: threat appraisal (threat ratings), physiological arousal (skin conductance response), perceptual learning [discrimination sensitivity (d') and response speed], and cortical plasticity [visual evoked potentials (VEPs) and cortical current density]. Interestingly, while behavioral, physiological, and VEP effects were comparable at immediate and delayed assessments, the cortical substrates evolved markedly over time, transferring from high-order cortices [inferotemporal/fusiform cortex and orbitofrontal cortex (OFC)] immediately to the primary and secondary visual cortex after the delay. In sum, the contrast between P- and M-biased conditioning confirms privileged conditioning acquisition via the subcortical pathway while the immediate cortical plasticity lends credence to the triadic amygdala-OFC-fusiform network thought to underlie threat processing. Importantly, long-term retention of conditioning in the basic sensory cortices supports the conserved role of the human sensory cortex in the long-term storage of aversive conditioning.SIGNIFICANCE STATEMENT A growing network of neural substrates has been identified in threat learning and memory. The sensory cortex plays a key role in long-term threat memory in animals, but such a function in humans remains unclear. To explore this problem, we conducted multidimensional assessments of immediate and delayed (15 d) effects of human aversive conditioning. Behavioral, physiological, and scalp electrophysiological data demonstrated conditioning effects and long-term retention. High-density EEG intracranial source analysis further revealed the cortical underpinnings, implicating high-order cortices immediately and primary and secondary visual cortices after the long delay. Therefore, while high-order cortices support aversive conditioning acquisition (i.e., threat learning), the human sensory cortex (akin to the animal homolog) underpins long-term storage of conditioning (i.e., long-term threat memory).

Event-related potential components as measures of aversive conditioning in humans.

For more than 60 years, the gold standard for assessing aversive conditioning in humans has been the skin conductance response (SCR), which arises from the activation of the peripheral nervous system. Although the SCR has been proven useful, it has some properties that impact the kinds of questions it can be used to answer. In particular, the SCR is slow, reaching a peak 4-5 s after stimulus onset, and it decreases in amplitude after a few trials (habituation). The present study asked whether the late positive potential (LPP) of the ERP waveform could be a useful complementary method for assessing aversive conditioning in humans. The SCR and LPP were measured in an aversive conditioning paradigm consisting of three blocks in which one color was paired with a loud noise (CS+) and other colors were not paired with the noise (CS-). Participants also reported the perceived likelihood of being exposed to the noise for each color. Both SCR and LPP were significantly larger on CS+ trials than on CS- trials. However, SCR decreased steeply after the first conditioning block, whereas LPP and self-reports were stable over blocks. These results indicate that the LPP can be used to assess aversive conditioning and has several useful properties: (a) it is a direct response of the central nervous system, (b) it is fast, with an onset latency of 300 ms, (c) it does not habituate over time.

Appetitive vs. Aversive conditioning in humans.

In classical conditioning, an initially neutral stimulus (conditioned stimulus, CS) becomes associated with a biologically salient event (unconditioned stimulus, US), which might be pain (aversive conditioning) or food (appetitive conditioning). After a few associations, the CS is able to initiate either defensive or consummatory responses, respectively. Contrary to aversive conditioning, appetitive conditioning is rarely investigated in humans, although its importance for normal and pathological behaviors (e.g., obesity, addiction) is undeniable. The present study intents to translate animal findings on appetitive conditioning to humans using food as an US. Thirty-three participants were investigated between 8 and 10 am without breakfast in order to assure that they felt hungry. During two acquisition phases, one geometrical shape (avCS+) predicted an aversive US (painful electric shock), another shape (appCS+) predicted an appetitive US (chocolate or salty pretzel according to the participants' preference), and a third shape (CS–) predicted neither US. In a extinction phase, these three shapes plus a novel shape (NEW) were presented again without US delivery. Valence and arousal ratings as well as startle and skin conductance (SCR) responses were collected as learning indices. We found successful aversive and appetitive conditioning. On the one hand, the avCS+ was rated as more negative and more arousing than the CS– and induced startle potentiation and enhanced SCR. On the other hand, the appCS+ was rated more positive than the CS– and induced startle attenuation and larger SCR. In summary, we successfully confirmed animal findings in (hungry) humans by demonstrating appetitive learning and normal aversive learning.

The scent of salience — Is there olfactory-trigeminal conditioning in humans?

Pavlovian fear conditioning has been thoroughly studied in the visual, auditory and somatosensory domain, but evidence is scarce with regard to the chemosensory modality. Under the assumption that Pavlovian conditioning relies on the supra-modal mechanism of salience attribution, the present study was set out to attest the existence of chemosensory aversive conditioning in humans as a specific instance of salience attribution. fMRI was performed in 29 healthy subjects during a differential aversive conditioning paradigm. Two odors (rose, vanillin) served as conditioned stimuli (CS), one of which (CS+) was intermittently coupled with intranasally administered CO2. On the neural level, a robust differential response to the CS+ emerged in frontal, temporal, occipito-parietal and subcortical brain regions, including the amygdala. These changes were paralleled by the development of a CS+-specific connectivity profile of the anterior midcingulate cortex (aMCC), which is a key structure for processing salience information in order to guide adaptive response selection. Increased coupling could be found between key nodes of the salience network (anterior insula, neo-cerebellum) and sensorimotor areas, representing putative input and output structures of the aMCC for exerting adaptive motor control. In contrast, behavioral and skin conductance responses did not show significant effects of conditioning, which has been attributed to contingency unawareness. These findings imply substantial similarities of conditioning involving chemosensory and other sensory modalities, and suggest that salience attribution and adaptive control represent a general, modality-independent principle underlying Pavlovian conditioning.

Reappraisal of Threat Value: Loss of Blocking in Human Aversive Conditioning

Non-specificity of fear is a core aspect of what makes anxiety disorders so impairing: Fear does not remain specific to a single stimulus paired with danger, but generalizes to a broad set of stimuli, resulting in a snowballing of threat signals. The blocking procedure can provide a valuable laboratory model for gaining insight into such threat appraisal and generalization processes. We report two experiments in which we induced selective threat appraisal by using a blocking procedure in human aversive conditioning. We subsequently assessed to what extent such selective threat appraisal is sensitive to different kinds of interference. Results illustrate that the maintenance of selective threat appraisal is not guaranteed: Stimuli present during an aversive conditioning event that are initially tagged with a low threat value, can come to be tagged with a higher threat value later on, without additional experience with these stimuli. We argue that such interference in selective threat appraisal might be one of the mechanisms underlying the pathogenesis of non-specific fear.

Expectancy bias in a selective conditioning procedure: Trait anxiety increases the threat value of a blocked stimulus

Background and ObjectivesIn a blocking procedure, a single conditioned stimulus (CS) is paired with an unconditioned stimulus (US), such as electric shock, in the first stage. During the subsequent stage, the CS is presented together with a second CS and this compound is followed by the same US. Fear conditioning studies in non-human animals have demonstrated that fear responding to the added second CS typically remains low, despite its being paired with the US. Accordingly, the blocking procedure is well suited as a laboratory model for studying (deficits in) selective threat appraisal. The present study tested the relation between trait anxiety and blocking in human aversive conditioning. MethodsHealthy participants filled in a trait anxiety questionnaire and underwent blocking treatment in the human aversive conditioning paradigm. Threat appraisal was measured through shock expectancy ratings and skin conductance. ResultsAs hypothesized, trait anxiety was positively associated with shock expectancy ratings to the blocked stimulus. LimitationsIn skin conductance responding, no significant effects of stimulus type could be detected during blocking training or testing. The current study does not allow strong claims to be made regarding the theoretical process underlying the expectancy bias we observed. ConclusionsThe observed shock expectancy bias might be one of the mechanisms leading to non-specific fear in individuals at risk for developing anxiety disorders. A deficit in blocking, or a deficit in selective threat appraisal at the more general level, indeed results in fear becoming non-specific and disconnected from the most likely causes or predictors of danger.

Neural Systems Underlying Aversive Conditioning in Humans with Primary and Secondary Reinforcers

Money is a secondary reinforcer commonly used across a range of disciplines in experimental paradigms investigating reward learning and decision-making. The effectiveness of monetary reinforcers during aversive learning and associated neural basis, however, remains a topic of debate. Specifically, it is unclear if the initial acquisition of aversive representations of monetary losses depends on similar neural systems as more traditional aversive conditioning that involves primary reinforcers. This study contrasts the efficacy of a biologically defined primary reinforcer (shock) and a socially defined secondary reinforcer (money) during aversive learning and its associated neural circuitry. During a two-part experiment, participants first played a gambling game where wins and losses were based on performance to gain an experimental bank. Participants were then exposed to two separate aversive conditioning sessions. In one session, a primary reinforcer (mild shock) served as an unconditioned stimulus (US) and was paired with one of two colored squares, the conditioned stimuli (CS+ and CS−, respectively). In another session, a secondary reinforcer (loss of money) served as the US and was paired with one of two different CS. Skin conductance responses were greater for CS+ compared to CS− trials irrespective of type of reinforcer. Neuroimaging results revealed that the striatum, a region typically linked with reward-related processing, was found to be involved in the acquisition of aversive conditioned response irrespective of reinforcer type. In contrast, the amygdala was involved during aversive conditioning with primary reinforcers, as suggested by both an exploratory fMRI analysis and a follow-up case study with a patient with bilateral amygdala damage. Taken together, these results suggest that learning about potential monetary losses may depend on reinforcement learning related systems, rather than on typical structures involved in more biologically based fears.

Dynamic causal modelling of anticipatory skin conductance responses

Anticipatory skin conductance responses [SCRs] are a widely used measure of aversive conditioning in humans. Here, we describe a dynamic causal model [DCM] of how anticipatory, evoked, and spontaneous skin conductance changes are generated by sudomotor nerve activity. Inversion of this model, using variational Bayes, provides a means of inferring the most likely sympathetic nerve activity, given observed skin conductance responses. In two fear conditioning experiments, we demonstrate the predictive validity of the DCM by showing it has greater sensitivity to the effects of conditioning, relative to alternative (conventional) response estimates. Furthermore, we establish face validity by showing that trial-by-trial estimates of anticipatory sudomotor activity are better predicted by formal learning models, relative to response estimates from peak-scoring approaches. The model furnishes a potentially powerful approach to characterising SCR that exploits knowledge about how these signals are generated.

Dopamine-induced changes in neural network patterns supporting aversive conditioning

The aim of the present paper is to assess the effects of altered dopamine (DA) transmission on the functional connectivity among brain regions mediating aversive conditioning in humans. To this aim, we analyzed a previous published data set from a double-blind design combined with functional magnetic resonance imaging (fMRI) recordings in which healthy volunteers were randomly assigned to one of three drug groups: amphetamine (an indirect DA agonist), haloperidol (DA D2 receptor antagonist), and placebo. Participants were exposed to an aversive classical conditioning paradigm using cutaneous electrical stimulation as the unconditioned stimulus (US), and visual cues as the conditioned stimuli (CS) where one colour (CS+) was followed by the US in 33% of the trials and another colour (CS−) had no consequences. All participants reported awareness of stimulus contingencies. Group analysis of fMRI data revealed that the left ventral striatum (VS) and amygdala activated in response to the CS+ in all the three groups. Because of their activation patterns and documented involvement in aversive conditioning, both regions were used as seeds in the functional connectivity analysis. To constrain the functional networks obtained to relate to the conditioned response, we also correlated seed activity with the Galvanic Skin Response (GSR). In the placebo group, the right ventral tegmental area/substantia nigra (VTA/SN), bilateral caudate, right parahippocampal gyrus, left inferior parietal lobule (IPL), bilateral postcentral gyrus, bilateral middle frontal (BA 46), orbitofrontal, and ventromedial prefrontal cortices (PFC, BA 10/11) correlated with the VS and amygdala seeds in response to the CS+ compared to the CS−. Enhancing dopamine transmission via amphetamine was associated with reduced task differences and significant functional connectivity for both CS+ and CS− conditions between the left VS seed and regions modulated by DA, such as the left VTA/SN, right caudate, left amygdala, left middle frontal gyrus (BA 46), and bilateral ventromedial PFC (BA 10). Blocking dopamine transmission via haloperidol was associated with significant functional connectivity across an alternate network of regions including the left amygdala seed and the right insula, the left ACC (BA 24/32), bilateral IPL (BA 40), precuneus (BA 7), post-central gyrus, middle frontal gyrus (BA 46), and supplementary motor area (SMA, BA 6) to the CS+ versus the CS−. These data provide insight into the distinct effects of DA agents on the functional connectivity between striatal, limbic, and prefrontal areas.

Neural, electrodermal and behavioral response patterns in contingency aware and unaware subjects during a picture–picture conditioning paradigm

One way of investigating affective learning is the use of aversive pictures as unconditioned stimuli (UCS) in conditioning paradigms. In the last decades, there has been a heated debate on the influence of contingency awareness on conditioned responses (CRs). Only a few studies found CRs in contingency unaware subjects whereas other studies only reported conditioned reactions in contingency aware participants. However, as a shortcoming, most studies employing picture–picture paradigms only investigated one response level (e.g. changes in subjective ratings). Further, changes in brain activity have so far been neglected in this field of research. The aim of the present study was to investigate different response levels with respect to contingency awareness: brain activity measured by functional magnetic resonance imaging (fMRI), skin conductance responses (SCRs) and valence ratings. A neutral geometric shape (conditioned stimulus, CS+) was followed by aversive pictures, whereas another shape (CS−) preceded neutral pictures. Unaware participants showed CRs in brain activity (e.g. the insula). Generally more activity was observed in the fear network (e.g. the amygdala, the lateral orbitofrontal cortex) in aware participants and in the nucleus accumbens (NAcc). Investigation of SCRs and valence ratings revealed that only aware participants showed conditioned reactions. Our results point toward dissociations between response levels (e.g. brain activity) not affected by contingency awareness and more cognitive response levels (e.g. subjective ratings and SCRs) which are affected by contingency awareness. As a unique finding in human aversive conditioning, we discuss the role of the nucleus accumbens as well as practical implications for affective learning models.

Temporal Difference Modeling of the Blood-Oxygen Level Dependent Response During Aversive Conditioning in Humans: Effects of Dopaminergic Modulation

The prediction error (PE) hypothesized by the temporal difference model has been shown to correlate with the phasic activity of dopamine neurons during reward learning and the blood-oxygen level dependent (BOLD) response during reward and aversive conditioning tasks. We hypothesized that dopamine would modulate the PE related signal in aversive conditioning and that haloperidol would reduce PE related activity, while an acute dose of amphetamine would increase PE related activity in the ventral striatum. Healthy participants took an acute dose of amphetamine, haloperidol, or placebo. We used functional magnetic resonance imaging (fMRI) to measure the BOLD signal while they carried out an aversive conditioning task, using cutaneous electrical stimulation as the unconditioned stimulus (US) and yellow and blue circles as conditioned stimulus (CS+ and CS-, respectively). Prediction error related BOLD activity was seen only in the ventral striatum in the placebo subjects. The subjects given amphetamine showed a wider network of PE related BOLD activity, including the ventral striatum, globus pallidus, putamen, insula, anterior cingulate, and substantia nigra/ventral tegmental area. Haloperidol subjects did not show PE related activity in any of these regions. Our results provide the first demonstration that the modulation of dopamine transmission affects both the physiological correlates and PE related BOLD activity during aversive learning.

Contingency awareness in human aversive conditioning involves the middle frontal gyrus

In contrast to the wealth of data describing the neural mechanisms underlying classical conditioning, we know remarkably little about the mechanisms involved in acquisition of explicit contingency awareness. Subjects variably acquire contingency awareness in classical conditioning paradigms, in which they are able to describe the temporal relationship between a conditioned cue and its outcome. Previous studies have implicated the hippocampus and prefrontal cortex in the acquisition of explicit knowledge, although their specific roles remain unclear. We used functional magnetic resonance imaging to track the trial-by-trial acquisition of explicit knowledge in a concurrent trace and delay conditioning paradigm. We show that activity in bilateral middle frontal gyrus and parahippocampal gyrus correlates with the accuracy of explicit contingency awareness on each trial. In contrast, amygdala activation correlates with conditioned responses indexed by skin conductance responses (SCRs). These results demonstrate that brain regions known to be involved in other aspects of learning and memory also play a specific role, reflecting on each trial the acquisition and representation of contingency awareness.

Cortisol and DHEA-S are associated with startle potentiation during aversive conditioning in humans

Fear conditioning reliably increases the startle reflex and stress hormones, yet very little is known about the effect of stress hormones on fear-potentiated startle. Cortisol and the sulfate ester of dehydroepiandrosterone (DHEA-S) are involved in stress and anxiety. Evidence suggests that low cortisol/DHEA-S ratio has a buffering effect on stress and anxiety in preclinical and clinical studies, suggesting that there may be a relationship between fear-potentiated startle and cortisol and DHEA-S activity. The aim of the study was to examine whether there is a relationship between cortisol/DHEA-S ratio and fear-potentiated startle. Thirty healthy subjects participated in a differential aversive conditioning experiment during which one of two stimuli (CS+) was paired with a shock, and the other was not (CS-). Conditioned responses were assessed with the startle reflex, defined as startle potentiation during CS+ compared to CS-. DHEA-S and cortisol levels were assayed from blood samples collected in both a baseline and an aversive conditioning session. Subjective state anxiety, arousal, and valence were assessed at various times during testing. Fear-potentiated startle was larger in individuals with high compared to low cortisol/DHEA-S ratio. Multiple regression analyses revealed that fear-potentiated startle was positively associated with cortisol and negatively associated with DHEA-S. There was no significant correlation between DHEA-S and cortisol levels. These data suggest that cortisol and DHEA-S are involved in fear conditioning.

Reinstatement of fear responses in human aversive conditioning

The treatment of choice for a number of anxiety disorders is exposure therapy. However, successful reduction of fear through exposure is sometimes followed by a (partial) return of symptoms of fear (return of fear, ROF; Clin. Psychol. Rev. 9 (1989) 147). Several possible learning mechanisms have been suggested to explain ROF (e.g. mechanisms related to spontaneous recovery, renewal, reacquisition and reinstatement). The present study focuses on reinstatement, which refers to the observation that mere US-only presentations can ‘reinstate’ previously extinguished fear responses. Although animal research has repeatedly demonstrated this phenomenon, little is known about fear reinstatement in humans. The present study employed a differential aversive conditioning procedure: after acquisition and a subsequent extinction procedure, a series of four unpredicted US-only trials was scheduled in the reinstatement group. The control group did not receive additional US presentations. A significant reinstatement effect was observed for US-expectancy ratings and fear ratings in the reinstatement group, but not in the control group. No differences were observed in a reaction time measure of resource allocation to the conditioned stimuli. These findings constitute a first demonstration of reinstatement of conditioned fear responses in humans. Implications for exposure treatment and suggestions for future research are discussed.

Slow potentials, event-related potentials, "gamma-band" activity, and motor responses during aversive conditioning in humans.

We examined slow potentials, transient event-related potentials, and oscillatory-like responses in the electroencephalogram during aversive conditioning in humans, in order to determine what is happening in the neocortex when behavioral adaptations are learned. Pictures of an angry and a happy human face served as reinforced (CS+) and unreinforced (CS-) conditioned stimuli, respectively, in one group, and either the reversed condition or two discriminably different neutral faces in two other groups (total n = 48 subjects). The unconditioned stimulus (US) was intracutaneous shock delivered to the left hand 5 s after CS+ onset. The electroencephalographic (EEG) activity was recorded from Fz, Cz, Pz, C3, and C4, electromyographic (EMG) activity from bilateral forearm and corrugator muscles, and skin conductance from the right hand. During acquisition a negative slow potential developed after CS+ (not CS-), which was more pronounced when a neutral face served as CS+. Early (iCNV, initial contingent negative variation) and late (tCNV, terminal contingent negative variation) components of the slow-potential response were positively related to the magnitude of conditioned EMG responses. Differentiation of tCNV was larger when neutral faces signaled the US; iCNV persisted during extinction when a happy face served as CS+. Late-occurring event-related potentials (ERPs) elicited by the US diminished over conditioning, whereas short-latency US components and ERPs elicited by CS events did not. Fourier analysis revealed oscillatory ("gamma-band") activity between 30 and 40 Hz, which persisted up to 3 s after US delivery and diminished as conditioning progressed. Our findings indicate that learning is expressed in neocortical structures at the earliest stages of conditioning. The functional roles of the three types of EEG response in learning are discussed.

Tonic and phasic electrodermal measures of human aversive conditioning with long duration stimuli.

Two experiments investigated phasic and tonic electrodermal responding to long, variable-duration stimuli in aversive conditioning procedures. Experiment I demonstrated reliable differential conditioning on both phasic (first interval response, FIR) and tonic (change in skin conductance level, delta SCL; spontaneous fluctuations) measures, using 10-40-s slides as conditioned stimuli (CSs) and electric shock as the unconditioned stimulus (UCS). Experiment 2 examined the sensitivity of phasic and tonic measures to UCS aversiveness. Both FIR and delta SCL were greater to a conditioned stimulus paired with shock than one paired with an innocuous reaction-time warning tone. Responding was reversed when the reinforcement contingencies were reversed. It was suggested that, with long duration CSs and an aversive UCS, phasic measures to stimulus onset, such as FIR, may reflect both stimulus significance and threat appraisal, whereas tonic measures reflect primarily arousal or anxiety arising from anticipation of the aversive unconditioned stimulus.

Thalamic negativity associated with the endogenous late positive component of cerebral evoked potentials (P300): Recordings using discriminative aversive conditioning in humans and cats

Electrical potentials within the thalamus associated with the endogenous late positive component (LPC or P300) of cerebral evoked potentials were investigated in humans and cats. Essentially identical discrimination tasks using classical aversive conditioning were employed for recording LPC in both species. In humans, a negative wave within the same latency range as that of LPC was noted, with dependency on stimulusrelevance, at all thalamic and mesencephalic recording sites which included the nucl. centrum medianum, nucl. parafascicularis and periaqueductal gray. Similar negative waves were also observed over wide areas of the thalamus of cats including specific as well as non-specific thalamic nuclei. The negative waves seen in the thalamus were generally higher in amplitude than those seen in the white matter and became unclear when the electrodes penetrated the ventral border of the thalamus.

The effectiveness of electric shock and foul odor as unconditioned stimuli in classical aversive conditioning

In a partial replication of an earlier study, eighty undergraduate volunteers were assigned to the eight cells of a three between- and two within-group experimental design. One factor was the nature of the UCS—faradic shock or a carefully-timed blast of pyridine vapor; the second factor was the UCS contingency—the UCS being contingent upon the CS or randomly interspersed; the third factor was the nature of the CS—all CSs were compounds of colored, flavored liquids, with the discriminable dimension being either color or taste. The two within-subjects factors were the successive presentation of either the CS+ of the CS− over trials in a standard classical conditioning format. The outcome measures were sip-size, order of preference, and semantic differential ratings. It was found that the foul odor UCS resulted in no aversive conditioning with either color or taste cues. Where shock was the UCS, color, but not taste, became aversive. While lending no direct support to cue-appropriateness concepts, the results reveal the complexity of cue utilization in human aversive conditioning.

Application of a drive and an associative theory to the effects of CR-contingent increments and decrements in unconditioned stimulus intensity

An analysis of data from classical aversive conditioning of the nictitating membrane of rabbits with the two-phase model was made in order to test predictions from Spence’s drive theory and the Rescorla-Wagner associative theory. While overall group means were ordered as expected from these theories, the interrelationships among estimates of parameters in the two-phase model showed that neither can account for more detailed analyses of the data. It was shown, for example, that the inferior performance of subjects receiving an avoidance treatment was not, as theoretically predicted, a result of the CR-contingent omission of the US leading to a decrement in response probability. Similarly, it was shown that increments in response probability followed trials on which the CR occurred even when CR-contingent US intensity was lower than that on non-CR trials, an effect which neither theory can predict since both are response independent. These two outcomes are opposite to what has been obtained in human aversive conditioning, which means that the two theories, while accounting for the behavior of one species, are of limited generality across species.