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Brain mechanisms of extinction of the classically conditioned eyeblink response.

It is well established that the cerebellum and its associated circuitry are essential for classical conditioning of the eyeblink response and other discrete motor responses (e.g., limb flexion, head turn, etc.) learned with an aversive unconditioned stimulus (US). However, brain mechanisms underlying extinction of these responses are still relatively unclear. Behavioral studies have demonstrated extinction as an active learning process distinct from acquisition. Experimental data in eyeblink conditioning suggest that plastic changes specific to extinction may play an important role in this process. Both cerebellar and hippocampal systems may be involved in extinction of these memories. The nature of this phenomenon and identification of the neural substrates necessary for extinction of originally learned responses is the topic of this review.

Animals↗

PubMed2005

Opposing effects of amygdala and orbital prefrontal cortex lesions on the extinction of instrumental responding in macaque monkeys.

Extinction is a well-known behavioural phenomenon that allows organisms to respond flexibly to a changing environment. Although recent work implicates the amygdala and orbital prefrontal cortex (PFo) in extinction of Pavlovian conditioned fear and aversion, much less is known about the neural bases of instrumental extinction. To explore the contribution of the macaque amygdala to flexible responding in the face of changing reward contingency, we tested the effects of selective, excitotoxic lesions of the amygdala on extinction of an instrumental response. For comparison, we evaluated the effects of ablation of PFo on the same task. Amygdala lesions facilitated the extinction of instrumental responses, whereas lesions of PFo had the opposite effect.

Amygdala↗

PubMed1991

Tactile extinction in childhood hemiplegia.

Tactile extinction was investigated by the Quality Extinction Test (QET) of Schwartz in 39 patients, 34 of whom had congenital hemiplegia and five early-acquired hemiplegia. Extinction values were significantly higher in hemiplegia patients than in controls and usually contralateral to the side of brain damage, except for four cases with right hemiplegia. Values were also higher in males than in females. There were no significant differences between patients with left and right hemiplegia, and no correlations between extinction and IQ or the presence of epilepsy. There was a correlation between QET scores and neuroradiologically proven cerebral atrophy. The meaning of extinction in hemiplegic patients and the principal theories regarding the pathogenesis of neglect are discussed.

Adolescent↗

PubMed2006

Evidence for the role of infectious disease in species extinction and endangerment.

Infectious disease is listed among the top five causes of global species extinctions. However, the majority of available data supporting this contention is largely anecdotal. We used the IUCN Red List of Threatened and Endangered Species and literature indexed in the ISI Web of Science to assess the role of infectious disease in global species loss. Infectious disease was listed as a contributing factor in <4% of species extinctions known to have occurred since 1500 (833 plants and animals) and as contributing to a species' status as critically endangered in <8% of cases (2,852 critically endangered plants and animals). Although infectious diseases appear to play a minor role in global species loss, our findings underscore two important limitations in the available evidence: uncertainty surrounding the threats to species survival and a temporal bias in the data. Several initiatives could help overcome these obstacles, including rigorous scientific tests to determine which infectious diseases present a significant threat at the species level, recognition of the limitations associated with the lack of baseline data for the role of infectious disease in species extinctions, combining data with theory to discern the circumstances under which infectious disease is most likely to serve as an agent of extinction, and improving surveillance programs for the detection of infectious disease. An evidence-based understanding of the role of infectious disease in species extinction and endangerment will help prioritize conservation initiatives and protect global biodiversity.

Animals↗

PubMed1996

Reversible inactivations of the cerebellum prevent the extinction of conditioned nictitating membrane responses in rabbits.

1. Studies show that reversible inactivation of the anterior interpositus nucleus (AIP) of the cerebellum with muscimol (a GABAA agonist) prevents acquisition of the classically conditioned nictitating membrane response (NMR) in the rabbit. Here, we have used reversible inactivations of the AIP with muscimol to investigate the role of the cerebellum in the extinction of this response. 2. Experimental subjects were implanted with cannulae targeted to the AIP, through which muscimol could be infused via an injector cannula. This experiment was divided into three phases lasting 4 days, separated by 3 day intervals. Experimental and unoperated control subjects received acquisition training in phase 1; in phases 2 and 3 they received extinction training. 3. Presentation of the conditioned stimulus (CS) alone in phase 2 produced normal extinction in control subjects. Muscimol inactivation of the AIP in experimental subjects during phase 2 prevented extinction of conditioned responses (CRs), shown by initial high CR frequency in the first post-drug session of phase 3, which then extinguished in a manner indistinguishable from controls in phase 2. 4. Our findings support the suggestion that similar cerebellar circuitry is engaged in acquisition and extinction of NMR conditioning.

Animals↗

PubMed1987

Hemispace-visual field interactions in visual extinction.

Visual extinction was studied in a patient with neglect from a right hemispheric lesion. Extinction occurred during double simultaneous stimulation within the same visual hemi-field (VHF). This finding suggests that interhemispheric rivalry is not a critical factor. During double simultaneous stimulation within the same VHF as well as with stimuli in different VHFs, the severity of extinction was determined by both the retinotopic and the hemi-spatial position of the extinguished stimulus. The location of the other stimulus, however, did not seem relevant. It is proposed that damage to a corticolimbic-reticular system reduces attentional capacity and that extinction during double simultaneous stimulation in the visual modality reflects an inability to distribute the limited attentional resources equally to two different locations. The severity of extinction is a measure of an unequal distribution of attention along a gradient within visual space.

Attention↗

PubMed2003

Temporal modulation of spatial tactile extinction in right-brain-damaged patients.

Unilateral and bilateral electrotactile stimuli were delivered to both hands of 11 right-brain-damaged (RBD) patients with left tactile extinction and 20 healthy subjects. Bimanual stimuli could be presented simultaneously or with varying stimulus onset asynchronies (SOAs). Subjects indicated their detection of unilateral or bilateral stimuli, their judgements of whether stimuli were simultaneous or successive, and, in the latter case, which side came first. In RBD patients, extinction was maximal with simultaneous presentations and decreased as SOA increased. With short SOAs, omissions of left-sided stimuli occurred with both right-side and left-side stimulus precedence, suggesting a forward and backward interference of the right stimulus on the processing of the left stimulus within a time window of at least 100 msec. In contrast, there was no interference of the left stimulus on the detection of the right stimulus. Unlike controls, extinction patients rarely expressed simultaneity judgements, but those that were produced tended to be veridical or nearly so, like in normal controls. Whereas controls expressed generally accurate judgements of right or left precedence, patients showed a bias toward a right precedence and a maximal uncertainty between left-first and right-first choices when the left stimulus had a lead between 100 and 200 msec. The results are consistent with the hypothesis that, as in other sensory modalities, tactile extinction is associated with an abnormal persistent bias of attention toward the ipsilesional side that delays the processing of contralesional stimuli. However, the finding that both extinction and explicit judgements of simultaneity tended to occur with simultaneous bilateral stimuli suggest the presence of some residual neural capacity to detect precise temporal coincidence.

Adult↗

PubMed1987

Asymmetry of lateralised hemispheric functions in schizophrenia. Influence of clinical and epidemiological characteristics on quality extinction test performance.

The Quality Extinction Test was used to detect lateralised abnormalities of hemispheric functions in schizophrenic patients. Course of illness significantly affected the distribution of tactile extinctions, chronic patients showing more left-side extinctions than sub-chronic ones. Ages significantly affected the number of left extinctions, and sex the number of right extinctions. The importance of clinical and epidemiological characteristics of the patients in determining the quality and degree of hemispheric dysfunction in schizophrenia was confirmed.

Extinction, Psychological↗

PubMed1990

Effects of changes in stimulus contingency on visual extinction.

Left visual field (LVF) extinction during double simultaneous stimulation (DSS) is common in patients with right cerebral lesions. A postulate is that during DSS, the intact left hemisphere's limited attentional capacity is directed to right hemispace, resulting in LVF extinction. External cueing may help direct attention to the LVF and improve performance. In the present study, we varied patterns of unilateral stimulation preceding DSS in an attempt to redirect attention through expectancy. Nine patients (7 stroke, 2 tumor) with right hemisphere lesions each had 40 DSS trials with the distribution of unilateral stimulation trials systematically varied. Mean extinction percentages on DSS trials were 17% following 5 right unilateral trials, 30% following 1 right unilateral trial, 52% following 1 left unilateral trial, and 63% following 5 left unilateral trials. These findings indicate that the probability of extinction decreases following unilateral stimulation to the right visual field and increases following unilateral stimulation to the LVF. These results suggest that expectancy alters the attentional bias which mediates extinction.

Adult↗

PubMed2004

Distinct roles of hippocampal de novo protein synthesis and actin rearrangement in extinction of contextual fear.

It is believed that de novo protein synthesis is fundamentally linked to synaptic changes in neuronal circuits involved in acquisition and extinction of conditioned responses. Recent studies show that neuronal plasticity may be also altered by cytoskeletal rearrangement independently of protein synthesis. We investigated the role of these processes in the hippocampus during acquisition and extinction of context-dependent conditioned fear in mice. Intrahippocampal injections of the protein synthesis inhibitors anisomycin and puromycin, or of the actin rearrangement inhibitors cytochalasin D and latrunculin A, prevented the acquisition of context-dependent fear. Unexpectedly, anisomycin and puromycin enhanced extinction without erasing the fear memory. In contrast, cytochalasin D and latrunculin A prevented extinction of context-dependent freezing. On the basis of these findings, it is suggested that certain hippocampal mechanisms mediating extinction of conditioned contextual fear are inhibited by protein synthesis and involve actin rearrangement. Such mechanisms might predominantly elicit modifications of hippocampal circuits that store the conditioning memory.

Acoustic Stimulation↗

PubMed2004

The transition from memory retrieval to extinction.

Memory is measured by measuring retrieval. Retrieval is often triggered by the conditioned stimulus (CS); however, as known since Pavlov, presentation of the CS alone generates extinction. One-trial avoidance (IA) is a much used conditioned fear paradigm in which the CS is the safe part of a training apparatus, the unconditioned stimulus (US) is a footshock and the conditioned response is to stay in the safe area. In IA, retrieval is measured without the US, as latency to step-down from the safe area (i.e., a platform). Extinction is installed at the moment of the first unreinforced test session, as clearly shown by the fact that many drugs, including PKA, ERK and protein synthesis inhibitors as well as NMDA receptor antagonists, hinder extinction when infused into the hippocampus or the basolateral amygdala at the moment of the first test session but not later. Some, but not all the molecular systems required for extinction are also activated by retrieval, further endorsing the hypothesis that although retrieval is behaviorally and biochemically necessary for the generation of extinction, this last process constitutes a new learning secondary to the unreinforced expression of the original trace.

Animals↗

PubMed1993

Analysis of the reinforcement and extinction components in DRO contingencies with self-injury.

Previous research has shown that self-injurious behavior (SIB) maintained by positive reinforcement may be reduced under differential-reinforcement-of-other-behavior (DRO) contingencies. In this study, we conducted an analysis of the reinforcement and extinction components of DRO while treating the self-injury of 3 women with developmental disabilities. A functional analysis revealed that each subject's SIB was maintained by positive reinforcement in the form of attention. Subsequent reinforcer assessments identified preferred and nonpreferred stimuli for later use in conjunction with DRO. Results showed high rates of SIB for all 3 subjects during baseline, which persisted when DRO was implemented without the relevant extinction component (withholding of attention for SIB) for 2 of the subjects. Low rates of SIB were observed for all subjects when DRO plus extinction was implemented or when extinction was implemented alone, suggesting that extinction may be a critical component of DRO schedules.

Adult↗

PubMed1994

Reemergence and extinction of self-injurious escape behavior during stimulus (instructional) fading.

Based on results of a functional analysis indicating that the self-injurious behavior (SIB) of 3 individuals was maintained by negative reinforcement (escape from instructional situations), the effects of stimulus (instructional) fading were evaluated in a multiple baseline design across subjects. The rate of instructions was reduced to zero at the beginning of treatment and was gradually increased (faded in) across sessions as long as SIB remained low. However, if SIB remained high for 10 consecutive sessions, extinction was implemented until SIB decreased, at which point extinction was withdrawn and fading was resumed. Treatment was completed when the rate of instructions was the same as in baseline (two per minute), and SIB remained below 0.5 responses per minute for two consecutive sessions. Results showed that instructional fading (without extinction) virtually eliminated SIB initially, but these effects were not maintained. All 3 subjects required multiple exposures to extinction and over 150 treatment sessions in order to meet the end-of-treatment criteria. Advantages and limitations of fading procedures without an extinction component, as well as extensions of both interventions to other clinical problems, are discussed.

Adult↗

PubMed1996

Responding maintained by intermittent reinforcement: implications for the use of extinction with problem behavior in clinical settings.

Results of basic research have demonstrated that behavior maintained on an intermittent schedule of reinforcement (INT) will be extinguished more slowly than behavior maintained on a continuous schedule (CRF). Although these findings suggest that problem behaviors may be difficult to treat with extinction if they have been maintained on INT rather than on CRF schedules, few applied studies have examined this phenomenon with human behavior in clinical settings. The purpose of this study was to determine whether problem behavior maintained on CRF schedules would be extinguished more rapidly than behavior maintained on INT schedules. Three individuals diagnosed with profound mental retardation participated after results of pretreatment functional analyses had identified the sources of reinforcement that were maintaining their self-injury, aggression, or disruption. Subjects were exposed to extinction following baseline conditions with CRF or INT schedules alternated within reversal or multielement designs. Results suggested that problem behavior may not be more difficult to treat with extinction if they have been maintained on INT rather than CRF schedules. However, switching from an INT to a CRF schedule prior to extinction may lower the baseline response rate as well as the total number of responses exhibited during extinction.

Adult↗

PubMed1996

Developing a technology for the use of operant extinction in clinical settings: an examination of basic and applied research.

Extinction of operant behavior, which involves terminating the reinforcement contingency that maintains a response, is important to the development, generalization, and reduction of behavior in clinical settings. We review basic and applied research findings on variables that influence the direct and indirect effects of extinction and discuss the potential value of a general technology for the use of extinction. We suggest that current research findings are not sufficient for the development of a comprehensive, applied technology of extinction and provide extensive guidelines for further studies on factors that may affect the course of extinction in clinical settings.

Adult↗

PubMed2000

Effects of noncontingent reinforcement on problem behavior and stimulus engagement: the role of satiation, extinction, and alternative reinforcement.

This study examined the effects of noncontingent reinforcement (NCR) with and without extinction on problem behavior and stimulus engagement (consumption of reinforcement) of 4 participants. Reductions in problem behavior using NCR have frequently been attributed to both satiation of the reinforcer and extinction. In the current study, aspects of the NCR treatment effects were difficult to explain based solely on either a satiation or an extinction account. Specifically, it was found that stimulus engagement remained high throughout the NCR treatment analysis, and that problem behavior was reduced to near-zero levels during NCR without extinction. The implications of these findings are discussed with respect to the satiation and extinction hypotheses frequently described in the applied literature. Findings from basic studies examining the effects of response-independent schedules are presented, and are used as the basis for a matching theory account of NCR-related effects. It is proposed that reductions in problem behavior observed during NCR interventions may be a function of the availability of alternative sources of reinforcement.

Child↗

PubMed2002

Response rate viewed as engagement bouts: resistance to extinction.

Rats obtained food pellets by nose poking a lighted key, the illumination of which alternated every 50 s during a session between blinking and steady, signaling either a relatively rich (60 per hour) or relatively lean (15 per hour) rate of reinforcement. During one training condition, all the reinforcers in the presence of the rich-reinforcement signal were response dependent (i.e., a variable-interval schedule); during another condition only 25% were response dependent (i.e., a variable-time schedule operated concurrently with a variable-interval schedule). An extinction session followed each training block. For both kinds of training schedule, and consistent with prior results, response rate was more resistant to extinction in the presence of the rich-reinforcement signal than in the presence of the lean-reinforcement signal. Analysis of interresponse-time distributions from baseline showed that differential resistance to extinction was not related to baseline differences in the rate of initiating response bouts or in the length of bouts. Also, bout-initiation rate (like response rate) was most resistant to extinction in the presence of the rich-reinforcement signal. These results support the proposal of behavioral momentum theory (e.g., Nevin & Grace, 2000) that resistance to extinction in the presence of a discriminative stimulus is determined more by the stimulus-reinforcer (Pavlovian) than by the stimulus-response-reinforcer (operant) contingency.

Animals↗

PubMed2006

Extinction of a saccharin-lithium association: assessment by consumption and taste reactivity.

Extinction of a conditioned palatability shift preceded extinction of conditioned taste avoidance whether rats were tested using a within-subjects design or a between-subjects design. In each of two experiments, consumption of 0.1% saccharin was paired with either 20 ml/kg of 0.15 M LiCl or equi-volume physiological saline on a single trial. In Experiment 1, on each of 10 extinction trials, rats were given a taste reactivity test immediately prior to a consumption test. In Experiment 2, half of the rats were extinguished by taste reactivity testing and half of the rats were extinguished by a consumption test on each of 10 extinction trials. In both experiments, the aversive reactions of gaping and passive dripping were extinguished in a single trial and the suppression of ingestive reactions was extinguished in 2 trials; however, extinction of taste avoidance required 4-5 trials. These results suggest that rats continue to avoid a lithium-paired flavor even when they do not have an aversion to the taste.

Animals↗