Trace and Delay Eyeblink Conditioning: Contrasting Phenomena of Declarative and Nondeclarative Memory

Clark, Robert E.; Manns, Joseph R.; Squire, Larry R.

doi:10.1111/1467-9280.00356

Cited by 94 publications

(122 citation statements)

References 21 publications

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“…In this respect, our results are clear cut. They clearly generalize the eye-blink results previously described by Perruchet (1985) and Clark et al (2001) to the domain of voluntary responses. Our results strengthen the conclusions of studies showing that RTs may follow the same general laws as classically conditioned reactions (Los & Heslenfeld, 2005;Los, Knol, & Boers, 2001;Los & Van Den Heuvel, 2001;Razran, 1936;Rexroad, 1936;Stephens, 1937).…”

Section: Discussionsupporting

confidence: 89%

“…A promising strategy concerns the search for the conditions in which the expression of these processes is favored or hampered. In eyeblink conditioning, Clark et al (2001) reported that the dissociation between subjective expectancy and eyeblink responses disappeared and even turned into an association when the tone terminated before the onset of the air puff (a preparation known as "trace conditioning") instead of coterminating with E2 (a preparation known as "delay conditioning"). The reliability of this pattern of data has been questioned.…”

Section: Discussionmentioning

confidence: 99%

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Dissociating the effects of automatic activation and explicit expectancy on reaction times in a simple associative learning task.

Perruchet

Cleeremans²,

Destrebecqz³

2006

Journal of Experimental Psychology: Learning, Memory, and Cogni

169

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After repeated associations between two events, E1 and E2, responses to E2 can be facilitated either because participants consciously expect E2 to occur after E1 or because E1 automatically activates the response to E2, or because of both. In this article, the authors report on 4 experiments designed to pit the influence of these 2 factors against each other. The authors found that the fastest responses to a target in a reaction time paradigm occurred when automatic activation was highest and conscious expectancy lowest. These results, when considered together with previous findings indicating that, under most conditions, the relation between expectancy and reaction times is in the opposite direction, are indicative of a reversed association-an interaction pattern that J. C. Dunn and K. Kirsner (1988) demonstrated to be the only one that unambiguously points to the involvement of independent processes.Keywords: learning, conditioning, expectancy, automatism, dissociations When two normatively unrelated events, E1 and E2, are repeatedly displayed in close temporal succession, the presentation of E1 modifies (and generally improves) the behavioral response to E2. This phenomenon has been investigated in several independent areas of research, such as associative priming (where E1 is the prime, and E2 is the target), classical conditioning (where E1 is the conditioned stimulus [CS], and E2 is the unconditioned stimulus [US]), and studies of motor behavior (where E1 is the warning stimulus, and E2 is the imperative stimulus in a reaction time paradigm). In each case, and even though different terminology has often been used, the same distinction between two general classes of interpretations has been proposed. The first focuses on the conscious expectancy for E2 that is initiated by the occurrence of E1. The second class of interpretations posits some form of automatic activation through which the occurrence of E1 facilitates the response to E2 as a mandatory consequence of their having been repeatedly associated in the past. In this context, automatic activation is therefore assumed to reflect previous experience with the association, independent of the agent's conscious expectancy for E2.

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Section: Discussionsupporting

confidence: 89%

Section: Discussionmentioning

confidence: 99%

Dissociating the effects of automatic activation and explicit expectancy on reaction times in a simple associative learning task.

Perruchet

Cleeremans²,

Destrebecqz³

2006

Journal of Experimental Psychology: Learning, Memory, and Cogni

169

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“…Such a pattern was reported during eye-blink conditioning (37). We failed to find any significant trend in response slope.…”

Section: Discussionsupporting

confidence: 62%

Working memory and fear conditioning

Carter

Hofstötter

Tsuchiya

et al. 2003

Proc. Natl. Acad. Sci. U.S.A.

119

113

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In trace conditioning, a short interval is interposed between the termination of the conditioned stimulus (CS) and the onset of the unconditioned stimulus (US). In delay conditioning, the CS and US overlap. We here investigate the extent to which human classical fear conditioning depends on working memory. Subjects had to carry out an n-back task, requiring tracking an item 1 or 2 back in a sequentially presented list of numbers, while simultaneously being tested for their ability to associate auditory cues with shocks under a variety of conditions (single-cue versus differential; delay versus trace; no task versus 0-, 1-, and 2-back). Differential delay conditioning proved to be more resilient than differential trace conditioning but does show a reduction due to task interference similar in slope to that found in trace conditioning. Explicit knowledge of the stimulus contingency facilitates but does not guarantee trace conditioning. Only the single-cue delay protocol shows conditioning during the more difficult working memory task. Our findings suggest that the larger the cognitive demands on the system, the less likely conditioning occurs. A postexperimental questionnaire showed a positive correlation between conditioning and awareness for differential trace conditioning extinction. P avlovian conditioning is widely used to study associative learning in species ranging from mollusks to flies, rodents, monkeys, and humans (1-10). This form of learning involves the association of an initially neutral stimulus, the conditioned stimulus (CS), with a correlated meaningful stimulus, the unconditioned stimulus (US). An unresolved question concerns the extent to which certain forms of classical conditioning depend on higher-level cognitive processes, including selective attention, working memory, and awareness (11-17). Eye-blink conditioning is an associative learning paradigm where the role of explicit knowledge͞awareness is being investigated. The paradigm involves the association of an eye blink (a somatic motor response) with a previously meaningless stimuli (CS).Recent data showed that, in humans, associative trace conditioning of eye-blink responses requires awareness of the contingency between the CS (a tone) and the US (a puff of air to the eye), whereas this is not the case for delay conditioning (11,(18)(19)(20). In delay conditioning, the start of the US is temporally contiguous with the CS, whereas in trace conditioning, an interval is interposed between the end of the CS and the start of the US. Distracting subjects by having them perform a secondary task (for example, a verbal shadowing task) during a trace procedure prevents conditioning. Furthermore, subjects' ability to report the exact nature of the CS͞US relationship (e.g., ''I believe the tone came before the airpuff'') is greatly impaired with concurrent distraction during trace conditioning. Conversely, associative delay conditioning appears to be insensitive to distractors. Other experiments find that both trace and delay associative differentia...

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“…Early evidence comes from lesion studies with experimental animals showing that acquisition of trace conditioning requires intact hippocampal formation (4,5) and medial prefrontal cortex (6), whereas delay conditioning can occur even with the entire forebrain removed (7,8). Later studies involving human subjects further validate the involvement of different brain circuits in these two conditioning variants and even suggest, more surprisingly, that conscious awareness might be a prerequisite for trace but not delay conditioning (9,10). It is then hypothesized that the participation of hippocampus and neocortex, as well as the associated higher cognitive function, is necessary in trace conditioning to maintain a representation of the CS or CS/US contingency so as to bridge the temporal gap (11,12).…”

mentioning

confidence: 99%

Distinct molecular underpinnings of Drosophila olfactory trace conditioning

Shuai

Qin

et al. 2011

Proc. Natl. Acad. Sci. U.S.A.

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Trace conditioning is valued as a simple experimental model to assess how the brain associates events that are discrete in time. Here, we adapted an olfactory trace conditioning procedure in Drosophila melanogaster by training fruit flies to avoid an odor that is followed by foot shock many seconds later. The molecular underpinnings of the learning are distinct from the well-characterized simultaneous conditioning, where odor and punishment temporally overlap. First, Rutabaga adenylyl cyclase (Rut-AC), a putative molecular coincidence detector vital for simultaneous conditioning, is dispensable in trace conditioning. Second, dominant-negative Rac expression, thought to sustain early labile memory, significantly enhances learning of trace conditioning, but leaves simultaneous conditioning unaffected. We further show that targeting Rac inhibition to the mushroom body (MB) but not the antennal lobe (AL) suffices to achieve the enhancement effect. Moreover, the absence of trace conditioning learning in D1 dopamine receptor mutants is rescued by restoration of expression specifically in the adult MB. These results suggest the MB as a crucial neuroanatomical locus for trace conditioning, which may harbor a Rac activity-sensitive olfactory "sensory buffer" that later converges with the punishment signal carried by dopamine signaling. The distinct molecular signature of trace conditioning revealed here shall contribute to the understanding of how the brain overcomes a temporal gap in potentially related events.learning and memory | olfaction | cAMP | Rho GTPase I n trace conditioning, the conditional stimulus (CS) and the unconditional stimulus (US) are separated in time by a stimulus-free interval (1). This so-called "trace interval" can last for a fraction of a second in eyeblink conditioning but many seconds in fear conditioning, which poses a challenging question: how does the brain overcome this temporal gap to form the association between the CS and US (2)? Intriguingly, trace conditioning in mammals engages neural substrates fundamentally different from delay conditioning, where the CS precedes but also temporally overlaps with the US (3). Early evidence comes from lesion studies with experimental animals showing that acquisition of trace conditioning requires intact hippocampal formation (4, 5) and medial prefrontal cortex (6), whereas delay conditioning can occur even with the entire forebrain removed (7,8). Later studies involving human subjects further validate the involvement of different brain circuits in these two conditioning variants and even suggest, more surprisingly, that conscious awareness might be a prerequisite for trace but not delay conditioning (9, 10). It is then hypothesized that the participation of hippocampus and neocortex, as well as the associated higher cognitive function, is necessary in trace conditioning to maintain a representation of the CS or CS/US contingency so as to bridge the temporal gap (11, 12). However, little is known about what form this representation takes and how it e...

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Trace and Delay Eyeblink Conditioning: Contrasting Phenomena of Declarative and Nondeclarative Memory

Cited by 94 publications

References 21 publications

Dissociating the effects of automatic activation and explicit expectancy on reaction times in a simple associative learning task.

Dissociating the effects of automatic activation and explicit expectancy on reaction times in a simple associative learning task.

Working memory and fear conditioning

Distinct molecular underpinnings of Drosophila olfactory trace conditioning

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