Conditioning with masked stimuli affects the timecourse of skin conductance responses.

In Pavlovian fear conditioning, an aversive unconditional stimulus (UCS) is repeatedly paired with a neutral conditional stimulus (CS). As a consequence, the subject begins to show conditional responses (CRs) to the CS that indicate expectation and fear. There are currently two general models competing to explain the role of subjective awareness in fear conditioning. Proponents of the single-process model assert that a single propositional learning process mediates CR expression and UCS expectancy. Proponents of a dual-process model assert that these behavioral responses are expressions of two independent learning processes. We used backward masking to block perception of our visual CSs and measured the effect of this training on subsequent unmasked performance. In two separate experiments we show a dissociation between CR expression and UCS expectancy following differential delay conditioning with masked CSs. In Experiment I, we show that masked training facilitates CR expression when the same CSs are presented during a subsequent unmasked reacquisition task. In Experiment II we show that masked training retards learning when the CS+ is presented as part of a compound CS during a subsequent unmasked blocking task. Our results suggest that multiple memory systems operate in a parallel, independent manner to encode emotional memories. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

The dopaminergic modulation of fear: Quinpirole impairs the recall of emotional memories in rats.

Past studies examining the contributions of dopamine to fear have produced inconsistent results. The present experiments reevaluated this issue. It was found that systemic pretreatment with the D2 agonist quinpirole before pairing 2 conditioned stimuli (CSs; CS2–CS1) dose dependently blocked the acquisition of second-order fear conditioning. Quinpirole's actions were not due to nonspecific impairments in the ability to perceive the CSs, or form and store an association, because the identical drug pretreatment before pairing the same 2 CSs had no effect on the acquisition of sensory preconditioning. In a separate study, rats were given fear conditioning while untreated and then received extinction sessions while under the influence of quinpirole or its vehicle. Quinpirole pretreatment blocked extinction. Findings suggest that quinpirole decreased fear by blocking the retrieval of a learned association between a CS and unconditioned stimulus (UCS), rather than by devaluing the UCS, which would have resulted from summation of quinpirole's appetitive properties with the aversive properties of fear. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Conditioned diminution of the unconditioned skin conductance response.

During Pavlovian conditioning the expression of a conditioned response typically serves as evidence that an association between a conditioned stimulus (CS) and an unconditioned stimulus (UCS) has been learned. However, learning-related changes in the unconditioned response (UCR) produced by a predictable UCS can also develop. In the present study, we investigated learning-related reductions in the magnitude of the unconditioned skin conductance response (SCR). Healthy volunteers participated in a differential conditioning study in which one tone (CS+) was paired with a loud white-noise UCS and a second tone (CS?) was presented alone. In addition, probe trials that consisted of UCS presentations paired with the CS+ (CS + UCS) and CS? (CS ? UCS), as well as presentations of the UCS alone were included to assess UCR diminution. SCR and participants' expectations of UCS presentation were monitored during conditioning. Greater diminution of the UCR was observed to the UCS when it followed the CS+ compared to when it followed the CS? or was presented alone. Further, UCR amplitude showed an inverse relationship with the participants' ratings of UCS expectancy. However, conditioned UCR diminution was also observed independent of differential UCS expectancies. Our findings demonstrate conditioned diminution of the unconditioned SCR. Further, these findings suggest that although UCR amplitude is modified by conscious expectations of the UCS, conditioned diminution of the UCR can be expressed independent of learning-related changes in these expectations. (PsycINFO Database Record (c) 2011 APA, all rights reserved)     

Slow reacquisition following extinction: Context, encoding, and retrieval mechanisms.

Investigated the slow reacquisition (RAQ) of responding in rats that occurs when the conditioned stimulus/stimuli (CS) and unconditioned stimulus/stimuli (UCS) are paired again after prolonged extinction training. In Exp 1, an extinguished CS acquired less suppression than a novel CS during a final conditioning phase, but more suppression than CSs that had received comparable nonreinforcement without initial conditioning. In Exp 2, CS–UCS pairings resumed in the context of extinction caused the least RAQ of suppression: Pairings in a neutral context produced better RAQ, while return of the CS to the conditioning context caused an immediate renewal of responding to the CS. In Exp 3, a return of the CS to the extinction context after RAQ training caused renewed extinction performance and interfered with performance appropriate to RAQ. This effect was not due to demonstrable inhibitory conditioning of the extinction context. Results suggest that representations of conditioning and extinction (or CS–UCS and CS–no UCS relations) are both retained through extinction and that performance appropriate to either phase can be cued by the corresponding context. RAQ may thus be slow when the context retrieves an extinction memory. Similar mechanisms may also play a role in other Pavlovian interference paradigms. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Pavlovian conditioned inhibition.

Examined the notion of conditioned inhibition and suggests a definition in terms of the learned ability of a stimulus to control a response tendency opposed to excitation. 2 techniques of measuring inhibition are outlined: (1) the summation procedure in which an inhibitor reduces the response that would normally be elicited by another stimulus, and (2) the retardation of acquisition procedure in which an inhibitor is retarded in the acquisition of an excitatory CR. Examples of the use of these procedures are given for a variety of UCS modalities. Several possible operations for generating conditioned inhibitors are reviewed: extinction following excitatory conditioning, discriminative conditioning, arrangement of a negative correlation between CS and a UCS, use of an extended CS-UCS interval, and presentation of a stimulus in conjunction with UCS termination. These operations suggest that conditioned inhibitors are not generated either by simple extinction procedures or by pairing a stimulus with UCS termination. By contrast, for both salivary and fear conditioning the other procedures do appear to generate inhibitors. Most of the procedures generating conditioned inhibitors can be described as arranging a negatively correlated CS and UCS. (2 p. ref.) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Arousal-related associative response characteristics of dorsal lateral geniculate nucleus neurons during acoustic Pavlovian fear conditioning.

This research determined whether fear-conditioned, acoustic stimuli induce thalamic arousal reflected in associative responses in dorsal lateral geniculate nucleus (dLGN) neurons. Rabbits received a Pavlovian discriminative fear conditioning procedure in which one tone conditioned stimulus (CS+) was always paired with an aversive unconditioned stimulus (UCS) and another tone (CS–) was never paired with the UCS. Responses of single dLGN neurons to random CS+ and CS– presentations were then recorded. Nine of 15 recorded neurons demonstrated significantly greater firing during the CS+ versus the CS–. Their spontaneous activity demonstrated tonic firing during increased neocortical arousal and burst firing during decreased neocortical arousal. The results demonstrate that dLGN neurons show associative responses to fear-conditioned, acoustic stimuli and present a model for investigating the neural circuits by which such stimuli affect sensory processing at the thalamic level. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Effects of amygdala central nucleus lesions on blocking and unblocking.

The effects of lesions of the amygdala central nucleus (CN) on blocking and unblocking of appetitive Pavlovian conditioning were examined in 2 experiments with rats. In both lesioned and unlesioned rats, prior pairing of one CS with a food unconditioned stimulus/stimuli (UCS) blocked the acquisition of conditioning to a 2nd CS when a compound of both stimuli was paired with that same UCS. If the value of the UCS was increased or decreased when the 2nd CS was added, unlesioned rats acquired substantial conditioning to the 2nd cue (unblocking). Unblocking occurred in lesioned rats only when the UCS value was increased. In both lesioned and unlesioned rats, unblocking was prevented if the compound cue was paired with the original UCS prior to the change in UCS value. These data suggest that the CN is involved in increasing attention to signals for significant events but not in tuning out redundant cues. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

NMDA receptor antagonist MK-801 blocks learning of conditioned stimulus-unconditioned stimulus contiguity but not fear of conditioned stimulus in goldfish (Carassius auratus L.).

The blockade of learning of Pavlovian fear conditioning by the N-methyl-D-aspartic acid (NMDA)-receptor antagonist MK-801 was examined in 166 goldfish. In previously untrained fish, MK-801 blocked learning of a light-off or a tone conditioned stimulus (CS) paired with an electrical shock unconditioned stimulus (UCS). Pretraining on the light-off CS did not affect the rate of learning of the tone CS but protected the tone learning from disruption by MK-801. Switching from the light-off to the tone CS changed the identity of the CS but not its temporal contiguity with the UCS. Pretraining consisting of pseudoconditioning of the light-off CS did not protect subsequent tone learning from blockade by MK-801. Thus, the NMDA receptor functions are necessary for learning related to the temporal contiguity of the CS and UCS but not to the identity of the CS as a cue to the occurrence of the fearful effects of the UCS. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Simultaneous and backward fear conditioning in the rat.

Examined simultaneous and backward Pavlovian conditioning paradigms using a UCS event which was longer in duration than the CS. 3 experiments with male Sprague-Dawley rats (N = 96) paired a 4-sec electric shock with a 2-sec tone-light stimulus under conditions in which the onset of the stimulus occurred 0, .25, 1.0, 2.0, 4.0, or 4.5 sec. after the onset of the shock. Relative to nonpaired control procedures, response-contingent presentations of the CSs in these paradigms significantly suppressed a food-rewarded free operant, indicating that these temporal relationships can produce excitatory associative conditioning. It is suggested that the distinctions between "forward," "simultaneous," and "backward" procedures be modified to include a more molecular analysis of the UCS event. (21 ref.) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Systematic manipulation of individual events in a truly random control in rats.

Truly random sequences of tone (conditioned stimuli, CSs) and shock (unconditioned stimuli, UCSs) were given to a total of 78 male Holtzman albino rats. Conditioning to the CS was measured using a conditioned suppression procedure. In Exp I eliminating chance CS-UCS pairings by systematically removing CSs that overlapped UCSs weakened conditioning monotonically as a function of the number of paired CSs removed. In Exp II systematically delaying early chance pairings, while holding constant the number of CSs, UCSs, and pairings, produced a nonsignificant weakening of conditioning. In Exp III delaying pairings again produced a nonsignificant weakening of conditioning when the early nonpaired events were CSs but significantly weakened conditioning when the early nonpairings were UCSs. Data suggest that each chance pairing in the truly random control produces an increment in conditioning unless blocked by prior UCSs alone. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Blocking, Unblocking, and Overexpectation of Fear: A Role for Opioid Receptors in the Regulation of Pavlovian Association Formation.

Injection of the opioid receptor antagonist naloxone facilitated acquisition of fear to contextual and auditory conditioned stimuli (CSs) in Experiments 1A and 1B. Experiment 2 showed that prior conditioning to a distinctive context blocked conditioning to an auditory CS. Blocking of CS fear was prevented by administrations of naloxone or increases in footshock intensity. Blocking of CS fear was facilitated by decreases in footshock intensity in a naloxone-reversible manner. Experiment 3 showed that compound conditioning of two CSs, each previously and separately paired with shock, produced overexpectation of fear that was reversed by naloxone. These results are consistent with a role for opioid receptors controlling Pavlovian association formation by regulating the discrepancy (A--ΣV) described by R. A. Rescorla and A. R. Wagner (1972). (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Control of fluid palatability by exteroceptive Pavlovian signals.

Two experiments, with 15 male albino rats, investigated whether discrete auditory conditioned stimulus/stimuli (CS) that signal the availability or onset of taste unconditioned stimulus/stimuli (UCS) (sucrose, quinine) can control orofacial responses in the absence of those UCSs. In Exp I, one auditory stimulus (CS+) was paired with the delivery of a sucrose solution to the magazine floor, and another auditory stimulus (CS–) was never followed by sucrose. Following conditioning, oral infusions of water that were preceded by the CS+ were found to elicit more ingestive (sucrose-typical) orofacial responses than did water alone or water preceded by the CS–. In Exp II, the conditioned ingestive reactions to a signal for sucrose observed in Exp I again occurred, and conditioned aversive (quinine-typical) orofacial responses occurred in response to water infusions preceded by a former signal for quinine. Data suggest that perceived palatability may be influenced by Pavlovian associations involving exteroceptive CSs and illustrate the importance of supporting stimuli in modulating the effects of Pavlovian associations on behavior. (41 ref) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Associations between Pavlovian CSs and context.

Three experiments, with 48 female Carneaux pigeons, used autoshaping procedures to assess associations between Pavlovian CSs and the contexts in which they occur. Three contexts were identified by different visual wall patterns; conditioning of those contexts was assessed by the amount of general activity in their presence. Exp I found that key-light CSs given discriminative autoshaping with food in 1 context could differentially 2nd-order conditioned activity to 2 new contexts. Exp II used a sensory preconditioning procedure to detect associations between 2 contexts and neutral key lights given in their presence. When those key lights were differentially paired with food in a 3rd context, activity changed differentially in the original 2 contexts. Exp III found that if 2 key lights were paired with food in 2 different contexts, differential extinction of those key lights induced differential activity in their contexts. Consequently, CSs that are either previously conditioned, neutral, or currently undergoing conditioning all become associated with the context in which they appear. Exps II and III found that the changes in context value induced by manipulation of the key lights influenced learning and performance to other CSs. A context that was made valuable through manipulation of its CS was better able to block conditioning to another CS but also better able to promote performance to previously trained CSs. Results agree with outcomes of direct conditioning of a context by a UCS and suggest that CS-induced changes in context are quite powerful. (20 ref) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Differential control by context in the inflation and reinstatement paradigms.

Examined, in 5 conditioned suppression experiments, the influence of summation between fear of the CS and the context in experimental paradigms in which the rat is exposed to UCSs following conditioning or extinction. Context-preference tests assessed contextual fear. In Exps I–III with 88 female Wistar rats, the inflation paradigm, in which fear of a CS paired with a weak UCS is enhanced by exposure to intense UCS alone, was investigated. Results show that the contextual fear that was present when the target CS was tested was reduced by presenting the intense UCSs in a different context, by exposing Ss to the context following their presentation, and by signaling the intense UCSs with a 2nd CS. In Exp IV with 32 female Wistar rats, UCS exposures following conditioning or extinction both produced contextual fear, but only fear of the extinguished CS was reinstated by that fear. In Exp V with 32 female Wistar rats, identical amounts of contextual fear reinstated fear of an extinguished CS, but not a nonextinguished CS, when the 2 types of CSs were arranged to evoke comparable amounts of fear prior to testing. It is suggested that contextual fear plays a role in the reinstatement paradigm but not in the inflation paradigm. (33 ref) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Single neurons in the medial prefrontal cortex of the rat exhibit tonic and phasic coding during trace fear conditioning.

Trace fear conditioning is a learning task that requires the association of an auditory conditioned stimulus (CS) and a shock unconditioned stimulus (US) that are separated by a 20-s trace interval. Single neuron activity was recorded from the prelimbic and infralimbic areas of the medial prefrontal cortex in rats during trace fear conditioning or nonassociative unpaired training. Prelimbic neurons showed learning-related increases in activity to the CS and US, whereas infralimbic neurons showed learning-related decreases in activity to these stimuli. A subset of prelimbic neurons exhibited sustained increases in activity during the trace interval. These sustained prelimbic responses may provide a bridging code that allows for overlapping representations of CS and US information within the trace fear conditioning circuit. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Conditioned diminution of the unconditioned response in rabbit eyeblink conditioning: Identifying neural substrates in the cerebellum and brainstem.

Several models of Pavlovian conditioning assume that processing of an unconditioned stimulus/stimuli (UCS) is diminished by a CS with which it had been paired. Two experiments evaluated the hypothesis that UCS processing may be diminished by CS-dependent activation of the cerebellum. Experiment 1 showed that electrical brain stimulation (EBS) of the cerebellar interpositus nucleus diminished the peak amplitude of the rabbit's unconditioned eyeblink response. This effect was bilateral, was systematically related to the intensity of EBS, maximal 50 ms after the offset of EBS, and substantially reversed by naloxone. Experiment 2 showed that inactivating the contralateral red nucleus with γ-D-glutamylglycine blocked the decremental effect of interpositus stimulation. Implications for neural systems mediating the inhibitory effects of cerebellar activation and the antinociceptive role of noradrenergic and opioid systems in Pavlovian conditioning phenomena are discussed. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Classical conditioning of autonomic fear responses is independent of contingency awareness.

The role of contingency awareness in classical conditioning experiments using human subjects is currently under debate. This study took a novel approach to manipulating contingency awareness in a differential Pavlovian conditioning paradigm. Complex sine wave gratings were used as visual conditional stimuli (CS). By manipulating the fundamental spatial frequency of the displays, we were able to construct pairs of stimuli that varied in discriminability. One group of subjects was given an “easy” discrimination, and another was exposed to a “difficult” CS+ and CS–. A 3rd group was exposed to a stimulus that was paired with the unconditional stimulus (UCS) 50% of the time and served as a control. Skin conductance response (SCR) and continuous UCS expectancy data were measured concurrently throughout the experiment. Differential UCS expectancy was found only in the easy discrimination group. Differential SCRs were found in the easy discrimination group as well as in the difficult discrimination group, but not in the 50% contingency control. The difficult discrimination group did not exhibit differential UCS expectancy but did show clear differential SCR. These observations support a dual process interpretation of classical conditioning whereby conditioning on an implicit level can occur without explicit knowledge about the contingencies. The role of contingency awareness in classical conditioning experiments using human subjects is currently under debate. This study took a novel approach to manipulating contingency awareness in a differential Pavlovian conditioning paradigm. Complex sine wave gratings were used as visual conditional stimuli (CS). By manipulating the fundamental spatial frequency of the displays, we were able to construct pairs of stimuli that varied in discriminability. One group of subjects was given an “easy” discrimination, and another was exposed to a “difficult” CS+ and CS–. A 3rd group was exposed to a stimulus that was paired with the unconditional stimulus (UCS) 50% of the time and served as a control. Skin conductance response (SCR) and continuous UCS expectancy data were measured concurrently throughout the experiment. Differential UCS expectancy was found only in the easy discrimination group. Differential SCRs were found in the easy discrimination group as well as in the difficult discrimination group, but not in the 50% contingency control. The difficult discrimination group did not exhibit differential UCS expectancy but did show clear differential SCR. These observations support a dual process interpretation of classical conditioning whereby conditioning on an implicit level can occur without explicit knowledge about the contingencies. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Functional MRI of human amygdala activity during Pavlovian fear conditioning: Stimulus processing versus response expression.

Although laboratory animal studies have shown that the amygdala plays multiple roles in conditional fear, less is known about the human amygdala. Human subjects were trained in a Pavlovian fear conditioning paradigm during functional magnetic resonance imaging (fMRI). Brain activity maps correlated with reference waveforms representing the temporal pattern of visual conditional stimuli (CSs) and subject-derived autonomic responses were compared. Subjects receiving paired CS-shock presentations showed greater amygdala activity than subjects receiving unpaired CS-shock presentations when their brain activity was correlated with a waveform generated from their behavioral responses. Stimulus-based waveforms revealed learning differences in the visual cortex, but not in the amygdala. These data support the view that the amygdala is important for the expression of learned behavioral responses during Pavlovian fear conditioning. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Neurotoxic or electrolytic lesions of the ventral subiculum produce deficits in the acquisition and expression of Pavlovian fear conditioning in rats.

The effects of neurotoxic or electrolytic ventral subicular (vSUB) lesions on the acquisition and expression of Pavlovian fear conditioning in rats were examined. Conditioning consisted of the delivery of tone–footshock trials in a novel observation chamber, and freezing served as the measure of conditional fear. Pretraining vSUB lesions produced a severe tone freezing deficit and a modest context freezing deficit, whereas posttraining lesions produced severe deficits in freezing to both a tone -and a context conditional stimulus (CS). Similar impairments were produced by neurotoxic and electrolytic lesions. Increases in motor activity associated with the lesions could not account for freezing deficits. These results reveal that neurons in the vSUB have an important role in both the acquisition and expression of Pavlovian fear conditioning to contextual and acoustic CSs. (PsycINFO Database Record (c) 2010 APA, all rights reserved)