Contingency learning and causal detection in Hermissenda: II. Cellular mechanisms.

Neural correlates of contingency-sensitive behavioral changes in the nudibranch Hermissenda were studied by in vitro conditioning of the isolated nervous system. The additions of unpaired light-alone or hair cell-alone stimulation presentations to sequences of light-hair cell stimulation pairings were found to attenuate the cumulative depolarization of Type B photoreceptors that normally result from pairings. Unpaired light presentations produced a transitory depression of the light-induced depolarizing generator potential of the B cell. Unpaired hair cell stimulation synaptically hyperpolarized the B photoreceptor and thus acted in a retroactive manner to partially reverse the cumulative depolarization owing to prior pairings. Behavioral experiments revealed a striking temporal specificity for the decremental effects of added light and rotational stimulation. The present experiments indicate that contiguity and contingency relations are both encoded and stored in the Type B photoreceptors, and to a first approximation reflect similar biophysical mechanisms. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Interactive contributions of intracellular calcium and protein phosphatases to massed-trials learning deficits in Hermissenda.

Using Hermissenda as subjects, massed-trials training deficits were examined. Associative pairings of light and rotation induced a progressively greater conditioned foot contraction in response to light as the intertrial interval (ITI) was extended (up to 8 min). In contrast, a short ITI (30 s) produced no evidence of learning. In a corresponding in vitro conditioning experiment that mimicked training of the intact animal, facilitation of neuronal excitability in the animal's B photoreceptors paralleled the results obtained in vivo. Imaging of intracellular Ca2+ using Fura-2 indicated that Ca2+ levels remained elevated during short ITIs. This Ca2+ accumulation appears to induce activation of protein phosphatases because normal facilitation of the B photoreceptors was induced with a short ITI if training occurred in the presence of a phosphatase inhibitor. These results suggest that intracellular Ca2+ and protein phosphatases contribute interactively to the kinetics of memory formation and provide evidence that an accumulation of intracellular Ca2+ across training trials may impede memory formation. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Contingency learning and causal detection in Hermissenda: I. Behavior.

The addition of extra light-alone or rotation-alone presentations to sequences of light-rotation pairings reduced the associative suppression of phototaxic behavior for the nudibranch Hermissenda. Training-induced changes in Type B photoreceptor light responses were found to parallel the training-induced behavioral changes in the intact animal. The decremental effects of the degraded contingency treatments upon neural and behavioral changes normally occasioned by light-rotation pairings reflected two processes. One factor was the increased stimulation frequency entailed by degraded contingency training. The second factor reflected the specifically unpaired character of the added light-alone or rotation-alone presentations, independent of frequency changes. The attenuation of phototaxic suppression was not because of a general habituation process or adaptation to the effects of either visual or vestibular stimulation. Instead, attenuation seemed to reflect a local interference effect of interspersed unpaired stimuli. The present experiments demonstrate a sensitivity to stimulus contingencies for Hermissenda similar to that of many vertebrates and indicate that contiguity and contingency relations are both encoded and stored in the Type B photoreceptors. The results indicate that similar neurophysiological mechanisms are involved. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

A mutation in the AMPA-type glutamate receptor, glr-1, blocks olfactory associative and nonassociative learning in Caenorhabditis elegans.

The α-amino-3-hydroxy-5-methyl-4-isoxazole propionate (AMPA)-type ionotropic glutamate receptor mediates fast excitatory neurotransmission in the vertebrate brain and is important for synaptic plasticity and the initial induction of long-term potentiation (LTP). This study found that the putative Caenorhabditis elegans AMPA receptor gene, glr-1, plays a significant role in experience-dependent behavior in C. elegans. glr-1 mutants are deficient in an olfactory associative learning task, in which diacetyl (DA) is paired with acetic acid solution. glr-1 mutant nematodes are also impaired in nonassociative learning (habituation) with the same DA stimulus. The C. elegans learning mutants, lrn-1 and lrn-2, are impaired in chemosensory associative learning yet have no deficits in habituation. The results suggest that although associative and nonassociative learning can be genetically dissociated ( lrn-1 and lrn-2), they also share some common molecular processes, including glr-1 -mediated neurotransmission. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Acquisition of conditioned associations in Hermissenda: Additive effects of contiguity and the forward interstimulus interval.

Examined conditioned suppression of photokinesis (CSPK) by the marine mollusc in 3 experiments. In each experiment, groups of Ss received light (conditioned stimulus, CS) paired with high-speed orbital rotation (unconditioned stimulus, UCS), light and rotation explicitly unpaired, or no exposure to these stimuli. 24 hrs after training, all Ss were tested for CSPK in the presence of the light. 50 CS–UCS pairings resulted in a marginal CSPK, whereas 100 and 150 pairings produced strong CSPK. In Exp 2, delay between CS onset and UCS onset was varied between 1 and 10 s. The 10-s interstimulus interval (ISI) did not support conditioning, whereas 1-s and 2-s ISIs were effective. In Exp 3, CS–UCS pairings in which the CS preceded the onset of the UCS and ended with the offset of the UCS evoked stronger CSPK than either a CS that preceded the UCS and ended with its onset or a CS that was paired in simultaneous compound with the UCS. CS–UCS contiguity and the forward ISI act additively to establish the CS–UCS association. No differences were observed between groups that were untreated and that received the CS and UCS unpaired. Similarities are noted in the temporal characteristics of associative learning in these Ss and vertebrate species. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Spatial EEG correlates of nonassociative and associative olfactory learning in rabbits.

Recent studies have shown that spatially distributed olfactory bulbar activity correlates with odor-specific behavioral responding (e.g., W. J. Freeman and K. A. Grajski [see PA, Vol 75:28775]; Freeman and W. Schneider [see PA, Vol 68:5257]). The present studies established olfactory bulbar spatial EEG correlates of nonassociative and associative learning in odorant stimulation in rabbits. Behavior was quantified by measuring magnitude and probability of the sniff response. It was shown that (a) olfactory bulbar spatial EEG amplitude patterns do not simply reflect odor (peripheral) stimulation, (b) repeated presentations of a nonreinforced odor initially reveal a transient EEG pattern change but the pattern change does not recur after the subject has habituated to the odor, and (c) repeated presentations of a reinforced odor (mild cutaneous shock), with a second nonreinforced odor serving as a control, reveal that coexisting, odorspecific spatial EEG amplitude patterns emerge with the acquisition of differential behavioral responding. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Associative learning changes intrinsic to Hermissenda Type A photoreceptors.

The eyes of the nudibranch mollusc Hermissenda contain 2 classes of photoreceptors. Type B photoreceptors exhibit increased light responses (LRs) and membrane excitability after repeated pairings of light and rotation and play an important role in the mediation of associatively produced reductions in phototactic behavior. Type A photoreceptors (TAPs) have also been shown to change with associative training. In previous research it was not possible to determine whether the effects of associative training reflected changes in synaptic input to TAPs or intrinsic changes in somatic conductances. In the present study, intracellular recordings from synaptically isolated TAPs were obtained on retention days after training, and pairing-specific decreases in light-induced generator potentials and decreases in resting input resistance were observed. Current- and voltage-clamp analysis of TAPs from untrained animals revealed that an important determinant of the steady-state LR was a calcium-activated K+ current (IK–Ca). Thus, TAPs also appear to be a primary site for associative information storage in Hermissenda. It is suggested that enhancement of IK–Ca by associative training may contribute to the diminished LR of TAPs. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Exposure to inescapable stress persistently facilitates associative and nonassociative learning in rats.

The present study examined the proactive effects of inescapable stress on aversive Pavlovian conditioning. Stressed rats were restrained and exposed to 90 1-mA tailshocks. Twenty-four hours later, all rats were exposed to 10 conditioned stimuli (CS; 350 ms of white noise at 85 dB). Rats then received either paired training in which the CS coterminated with a 100-ms, 0.7-mA periorbital shock or the same stimuli presented in an explicitly unpaired fashion. After the unpaired exposures, these rats were also exposed to paired training. Previously stressed rats exhibited persistent sensitization to the white-noise stimulus. Stressed rats exposed to unpaired stimuli, and no longer exhibiting a sensitized response, acquired the eyeblink conditioned response at a facilitated rate when these stimuli were presented in a paired fashion. These results also demonstrate that the effect of stress on classical conditioning is long-lasting, in excess of 48 hr. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Stimulus intensity modulates associative and nonassociative responding in preweanling rats

The purpose of the present study was to determine whether preweanling rats respond differentially to the intensity and energy source of a stimulus. Previous studies have suggested that infants process compound stimuli based on net stimulus intensity regardless of the energy source of the compound's elements, but more direct tests have been needed of the infant's response to the stimulus attributes of intensity and energy source. This response was tested for auditory and visual stimuli that had been equated (Experiment 1) in terms of perceived intensity (low and high). Intensity level and energy source of the stimuli were varied independently within nonassociative (Experiment 2) and associative (Experiment 3) procedures. The overall results indicate that stimuli of a low-perceived intensity were processed in terms of their intensity, whereas high-intensity stimuli were processed on the basis of energy source. These results are relevant to contemporary issues of cognitive development in humans and animals.     

Potentiation of phototactic suppression in Hermissenda by a chemosensory stimulus during compound conditioning.

Modifications of Hermissenda's phototactic behavior by compound pairings of light, scallop extract, and rotation were assessed. In general, the scallop extract potentiated phototactic suppression. Potentiation was dependent on (a) conjunctive presentations of scallop and light, (b) number of conditioning trials, and (c) scallop extract concentration. In related experiments, no second-order conditioning or sensory preconditioning of phototactic suppression was observed, indicating that within-compound associations did not contribute appreciably to potentiation. These results represent the first detailed analysis of compound conditioning in a mollusk using discrete presentations of well-characterized conditioned stimuli from distinct sensory modalities. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Dissociation of associative and nonassociative concomitants of classical fear conditioning in the freely behaving rat.

In this article we describe three experiments aimed at determining why the conditioned stimulus/stimuli (CS) elicits similar increases in heart rate in groups given paired and random training. The first study demonstrates that regardless of the pseudoconditioning control procedure used (random, backwards, shock-alone, or naive), the same pattern of results is obtained: the increases in arterial pressure are greater in the paired than in each control group, but the heart rate rises to the same extent in all groups. The second study determined that the context in which the responses are tested (conditioning apparatus vs novel test chamber) does not affect the general pattern of results obtained. The third study demonstrates that the superficially similar increases in heart rate in conditioned and pseudoconditioned rats are achieved by different physiological mechanisms: coactivation of the sympathethic and parasympathetic nervous systems in conditioned rats and sympathetic excitation alone in pseudoconditioned rats. Thus, the heart is influenced by associative emotional processes, but heart rate is not, under these conditions, a particularly useful index of those influences. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Mutations that prevent associative learning in C. elegans.

The nematode Caenorhabditis elegans offers a promising system for the reductionist study of learning and memory. In this article, classical conditioning in C. elegans is demonstrated with a variety of associative learning assays. These assays allowed for the isolation and behavioral characterization of 2 mutant C. elegans lines impaired in associative learning. Both lines show no short-term or long-term associative conditioning; however, they appear relatively normal in tests of nonassociative learning and sensorimotor function. In combination with the well-described genetics and neuroanatomy of C. elegans, the isolation of mutants selectively, yet completely, blocked in associative learning provides the basis for an effective characterization of the cellular and molecular aspects of associative learning. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Associative training results in persistent reductions in a calcium-activated potassium current in Hermissenda Type B photoreceptors.

Type B photoreceptors from the eyes of associatively trained Hermissenda had significantly greater light responses and net input resistances than Type B cells from control Ss on retention days following conditioning. Differences were still apparent when the contribution of a rapidly inactivating K+ current (IA) to the photoresponse was minimized by either depolarization-induced inactivation or block by 4-aminopyridine ions. Training-produced differences in Type B cell light responses were abolished by treatments that reduced the contribution of a calcium-activated K+ current (IK-Ca) to the light response. Under voltage-clamp conditions in which IK-Ca was isolated from other components of outward current, it was selectively reduced by associative training. The associative reduction of IK-Ca could not be attributed to training-produced reductions in the voltage-dependent calcium current of Type B cells. ICa was enhanced by associative training. A slower component of light-induced inward current was greater for Type B cells from associatively trained Ss. Previous results suggest that this slower component may be due to light-induced reduction of a calcium-activated K+ current. Training-produced reductions in a calcium-activated K+ may be responsible for enhanced photoresponses of Type B cells from conditioned Ss. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Olfactory associative learning in Caenorhabditis elegans is impaired in lrn-1 and lrn-2 mutants.

The C. elegans mutants, lrn-1 and lrn-2, are impaired in associative learning using conditioned taste cues. Both mutants are defective in associative learning about appetitive and aversive events, indicating that lrn-1 and lrn-2 exert effects across motivational boundaries. In a new olfactory associative learning paradigm, in which wild type worms learn to avoid a previously attractive diacetyl odor after it has been paired with an aversive acetic acid solution, lrn-1 and lrn-2 are impaired. Although defective in associative learning using a conditioned olfactory cue, nonassociative learning (habituation and dishabituation) using this same olfactory cue is unaffected. The discovery that lrn-1 and lrn-2 are defective in associative learning with both taste and olfactory cues may suggest that associative learning in different sensory modalities converges on a common genetic pathway in C. elegans that is subserved by lrn-1 and lrn-2. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Attentional and error-correcting associative mechanisms in classical conditioning.

R. A. Rescorla (2000, Rescorla, 2001, Rescorla, 2002) reported that the associative changes undergone by 2 conditioned stimuli that are reinforced or not reinforced in compound depend on their initial associations. The results contradict the predictions of simple error-correction models but can be explained by models that incorporate a “constrained” error-correction rule. A model of classical conditioning presented by N. A. Schmajuk, Y. Lam, and J. A. Gray (1996) suggests that attentional mechanisms, acting during both compound training and testing, have an important role in producing those results. Moreover, the model suggests that those attentional mechanisms might obscure the evaluation of the associative changes undergone by the conditioned stimuli during compound training. Two experiments that differentiate our model from competing theories are proposed. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Temporal order sensitivity of associative neural and behavioral changes in Hermissenda..

Hermissenda's neural and behavioral changes produced by light-rotation pairings were assessed as a function of the temporal relations between visual and vestibular stimulation. The results of in vitro simulations of conditioning indicated that simultaneous pairings (synchronous onsets and offsets of light and caudal hair cell stimulation) resulted in significantly greater cumulative depolarization of Type B photoreceptors than did either forward (light preceded hair cell stimulation) or backward (hair cell stimulation preceded light) pairings. Further experiments revealed that the attenuation of cumulative depolarization produced by the forward and backward pairings reflected the asynchrony of stimulus offsets that characterize these conditioning sequences, rather than their onsets. Analogous behavioral experiments revealed that intact animals trained with forward or backward pairings exhibited significantly less conditioning than those trained with simultaneous pairings. Strong parallels between the magnitude of cumulative depolarization from in vitro conditioning studies and the behavioral results for intact animals were also observed in experiments in which stimulus onset synchrony was held constant but offsets were made asynchronous, and vice versa. Thus Hermissenda exhibits a sensitivity to the temporal arrangement of light and rotation, and the results of behavioral conditioning can be predicted accurately from the outcome of in vitro conditioning of the isolated nervous system. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Potentiation of phototactic suppression in Hermissenda by compound conditioning results in potentiated excitability changes in Type B and A photoreceptors.

Ocular photoreceptor responses were recorded from Hermissenda exposed to compound conditioning, light conditioning, or one of several control treatments. Type A photoreceptors from light-conditioned animals responded with smaller generator potentials than those from untrained animals; type B photoreceptors from light-conditioned animals responded with greater generator potentials and action potential frequencies than cells from controls. These photoresponse changes were accompanied by increases and decreases in the input resistances of type B and A photoreceptors, respectively. The same photoresponse changes occurred in exaggerated form in cells from compound-conditioned animals, but no additional input resistance changes were observed. The results suggest that potentiation of phototactic suppression by compound conditioning involves a facilitation of the same photoreceptor excitability changes produced by mere pairings of light and rotation. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Dissociations between expectancy and performance in simple and two-choice reaction-time tasks: A test of associative and nonassociative explanations.

Perruchet, Cleeremans, and Destrebecqz (2006) reported a striking dissociation between trends in the conscious expectancy of an event and the speed of a response that is cued by that event. They argued that this indicates the operation of independent processes in human associative learning. However, there remains a strong possibility that this dissociation is not a consequence of associative learning and is instead caused by changes in vigilance or sensitivity based on the recency of events on previous trials. Three experiments tested this possibility with versions of a cued reaction time task in which trends in performance could not be explained by these nonassociative factors. Experiment 1 introduced a dual-response version of the task, in which response-related vigilance should be held relatively constant, and Experiments 2 and 3 used a differential conditioning procedure to separate the influence of recent response cue presentation from the recent associative history of the trial events. In all experiments, similar trends in reaction time were evident, suggesting a genuine influence of associative learning on response performance. Experiment 3 demonstrated that the associative contribution to these trends was not caused by commensurate changes in expectancy of the response cue. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Dissociative affective and associative priming effects in the lexical decision task: Yes versus no responses to word targets reveal evaluative judgment tendencies.

The affective priming effect (AP; i.e., shorter evaluative or lexical decision latencies for affectively congruent prime–target pairs) has often been interpreted as evidence for spreading activation from the prime to affectively congruent targets. The present study emphasizes the view that in the lexical decision task, the prime-target configuration is implicitly evaluated as a question of the form "Is (prime) (target)?" (e.g., "Is death wise?") so that there is a tendency to affirm in cases of congruency and to negate in cases of incongruency. Therefore, after establishing the AP with the lexical decision task in Experiment 1, in Experiment 2 the assignment of yes responses to words and nonwords was varied. For the word?=?yes condition, the AP emerged, whereas the data pattern was reversed for the word?=?no condition. In Experiment 3, a comparable pattern of results was not found for symmetrical or backward associatively related prime–target pairs. (PsycINFO Database Record (c) 2010 APA, all rights reserved)