Encoding specific associative memory: Evidence from behavioral and neural manipulations.

Within-subjects procedures with rats assessed the associative structures acquired during conditioning trials in which the interval between the stimuli and food was either short or long (i.e., A–10 s→food and B–40 s→food). In Experiments 1 and 2, after these conditioning trials, A and B served as second-order reinforcers for 2 further stimuli (i.e., X→A and Y→B); whereas Experiment 3 used a sensory preconditioning procedure in which X→A and Y→B trials occurred before the conditioning trials, and rats were finally tested with X and Y. In each experiment, Y elicited greater responding at test than did X. This finding supports the contention that the long-lived trace of B (associated with food on B–40 s→food trials) is more similar to the memory of B that was associatively provoked by Y, than is the short-lived trace of A (associated with food on A–10 s→food trials) to the memory of A that was associatively provoked by X. These conclusions were reinforced by the effects of a neural manipulation that disrupted discrimination learning involving the short traces of stimuli but not the long traces of the same stimuli. (PsycINFO Database Record (c) 2011 APA, all rights reserved)     

Temporal Coding in Conditioned Inhibition: Analysis of Associative Structure of Inhibition.

Two experiments with rats as subjects were conducted to investigate the associative structure of temporal control of conditioned inhibition through posttraining manipulation of the training excitor-unconditioned stimulus (US) temporal relationship. Experiment 1 found that following simultaneous Pavlovian inhibition training (i.e., A → US/XA-no US) in which a conditioned stimulus (CS A) was established as a delay excitor, maximal inhibition was observed on a summation test when CS X was compounded with a delay transfer CS. Furthermore, posttraining shifts in the A-US temporal relationship from delay to trace resulted in maximal inhibition of a trace transfer CS. Experiment 2 found complementary results to Experiment 1 with an A-US posttraining shift from serial to simultaneous. These results suggest that temporal control of inhibition is mediated by the training excitor-US temporal relationship. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Contrasting reduced overshadowing and blocking.

Preexposure of a cue without an outcome (X-) prior to compound pairings with the outcome (XZ→O) can reduce overshadowing of a target cue (Z). Moreover, pairing a cue with an outcome (X→O) before compound training can enhance its ability to compete with another cue (i.e., blocking). Four experiments were conducted in a conditioned bar-press suppression preparation with rats to determine whether spacing of the X- or X→O trials would differentially affect reduced overshadowing and blocking. Experiment 1a showed that reduced overshadowing was larger with massed trials than with spaced trials. Experiment 1b found that blocking was larger with spaced trials than with massed trials. Experiments 2a and 2b indicated that these effects of trial spacing were both mediated by the associative status of the context at test. The results are interpreted in the framework of contemporary learning theories. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Integrative priming occurs rapidly and uncontrollably during lexical processing.

Lexical priming, whereby a prime word facilitates recognition of a related target word (e.g., nurse → doctor), is typically attributed to association strength, semantic similarity, or compound familiarity. Here, the authors demonstrate a novel type of lexical priming that occurs among unassociated, dissimilar, and unfamiliar concepts (e.g., horse → doctor). Specifically, integrative priming occurs when a prime word can be easily integrated with a target word to create a unitary representation. Across several manipulations of timing (stimulus onset asynchrony) and list context (relatedness proportion), lexical decisions for the target word were facilitated when it could be integrated with the prime word. Moreover, integrative priming was dissociated from both associative priming and semantic priming but was comparable in terms of both prevalence (across participants) and magnitude (within participants). This observation of integrative priming challenges present models of lexical priming, such as spreading activation, distributed representation, expectancy, episodic retrieval, and compound cue models. The authors suggest that integrative priming may be explained by a role activation model of relational integration. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Sequential and simultaneous feature positive discriminations: Occasion setting and configural learning in human Pavlovian conditioning.

Using a conditioned suppression preparation, the authors investigated sequential (X→A+/A–) versus simultaneous (XA+/A–) feature positive (FP) discrimination learning in humans. The sequential discrimination was expected to be resolved by means of a Feature X Modulated Target A–US association and the simultaneous discrimination by a feature X–US association. After sequential FP training, extinction of Feature X did not affect discriminative X→A/A responding (Experiment 1), and X transferred its modulatory ability only to new targets, B, that had also been modulated (Experiment 2). This suggests that the sequential FP discrimination indeed resulted in occasion setting. Unlike expected, Feature X Extinction did not affect discriminative XA/A responding after simultaneous FP training (Experiment 3), while at the same time Feature X did show the predicted nonselective transfer to new targets, B (Experiment 4). J. M. Pearce's (1987) configural learning theory can account for most but not all findings of Experiments 3 and 4. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Associative History Affects the Associative Change Undergone by Both Presented and Absent Cues in Human Causal Learning.

R. A. Rescorla (2000) noted that a number of influential theories of associative learning do not take the associative history of cues (i.e., the prior training that they have received) into account when calculating the associative change undergone by those cues. The authors tested this assumption in a human causal learning paradigm and found associative history to be an important determinant of the learning undergone by cues that are presented on a trial. Moreover, associative history was also found to influence the amount of retrospective revaluation undergone by absent cues. These findings conflict with models of causal learning in which the associative change undergone by an element of a cue compound is governed by a summed error term (e.g., R. A. Rescorla & A. R. Wagner, 1972). (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Transfer between contextual conditional discriminations: An examination of how stimulus conjunctions are represented.

In 2 experiments, when rats were placed in 1 pair of contexts, A and B, 2 relationships were in force (X→food and Y→no food), and when they were placed in another pair of contexts, C and D, the complementary relationships were operative (Y→food and X→no food). In Experiment 1, rats then received a 2nd discrimination that was either contextually congruent (in A and B, Y→food and X→no food; in C and D, X→food and Y→no food) or contextually incongruent (in A and D, Y→food and X→no food; in C and B, X→food and Y→no food) with the 1st discrimination. In Experiment 2, the 1st discrimination, involving X and Y, was interleaved with a 2nd discrimination, involving V and W, that was again either congruent (in A and B, V→food and W→no food) or incongruent (in A and D, V→food and W→no food) with the 1st discrimination. The congruent discriminations were acquired more readily than were the incongruent discriminations. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Differential age effects for implicit and explicit conceptual associative memory.

Older adults show disproportionate declines in explicit memory for associative relative to item information. However, the source of these declines is still uncertain. One explanation is a generalized impairment in the processing of associative information. A second explanation is a more specialized impairment in the strategic, effortful recollection of associative information, leaving less effortful forms of associative retrieval preserved. Assessing implicit memory of new associations is a way to distinguish between these viewpoints. To date, mixed findings have emerged from studies of associative priming in aging. One factor that may account for the variability is whether the manipulations inadvertently involve strategic, explicit processes. In two experiments we present a novel paradigm of conceptual associative priming in which subjects make speeded associative judgments about unrelated objects. Using a size classification task, Experiment 1 showed equivalent associative priming between young and older adults. Experiment 2 generalized the results of Experiment 1 to an inside/outside classification task, while replicating the typical age-related impairment in associative but not item recognition. Taken together, the findings support the viewpoint that older adults can incidentally encode and retrieve new meaningful associations despite difficulty with the intentional recollection of the same information. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Learning about associations: Evidence for a hierarchical account of occasion setting.

In 2 experiments rats were trained on a switching discrimination, with 4 occasion setters, A, B, C, and D and 2 target stimuli, x and y. When signaled either by A or by B, x was reinforced with food and y was not, whereas when signaled either by C or by D these reinforcement relations were reversed (i.e., A: → x+, A: y → ?, B: x → +, B: y → ?, C: x → ?, C: y → +, D: x → ?, D: y → +). In a subsequent Stage A was paired with shock, and then the degree to which food–reinforced (Experiment 1a) and nonreinforced (Experiment 1b) presentations of x and y were capable of eliciting fear was assessed. Those conditioned stimulus (CS)/unconditioned stimulus (US) relations that had been operative in the presence of the fear-eliciting occasion setter A (i.e., x → +, y → ?) elicited more fear than the alternative CS/US combinations (i.e., x → ?, y → +). The implications of these findings are discussed with reference to theories of occasion setting and of configural learning. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Effects of concurrent load on feature- and rule-based generalization in human contingency learning.

The effect of concurrent load on generalization performance in human contingency learning was examined in 2 experiments that employed the combined positive and negative patterning procedure of Shanks and Darby (1998). In Experiment 1, we tested 32 undergraduates and found that participants who were trained and tested under full attention showed generalization consistent with the application of an opposites rule (i.e., single cues signal the opposite outcome to their compound), whereas participants trained and tested under a concurrent cognitive load showed generalization consistent with surface similarity. In Experiment 2, we replicated the effect with 148 undergraduates and provided evidence that it was the presence of concurrent load during training, rather than during testing, that was critical. Implications for associative, inferential, and dual-process accounts of human learning are discussed. (PsycINFO Database Record (c) 2011 APA, all rights reserved)     

The status of rapid response learning in aging.

Strong evidence exists for an age-related impairment in associative processing under intentional encoding and retrieval conditions, but the status of incidental associative processing has been less clear. In 2 experiments, we examined the effects of age on rapid response learning—the incidentally learned stimulus–response association that results in a reduction in priming when a learned response becomes inappropriate for a new task. Specifically, we tested whether priming was equivalently sensitive in both age groups to reversal of the task-specific decision cue. Experiment 1 showed that cue inversion reduced priming in both age groups with a speeded inside/outside classification task, and in Experiment 2, cue inversion eliminated priming on an associative version of this task. Thus, the ability to encode an association between a stimulus and its initial task-specific response appears to be preserved in aging. These findings provide an important example of a form of associative processing that is unimpaired in older adults. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Mechanisms of inferential order judgments in humans (Homo sapiens) and rhesus monkeys (Macaca mulatta).

If A > B, and B > C, it follows logically that A > C. The process of reaching that conclusion is called transitive inference (TI). Several mechanisms have been offered to explain transitive performance. Scanning models claim that the list is scanned from the ends of the list inward until a match is found. Positional discrimination models claim that positional uncertainty accounts for accuracy and reaction time patterns. In Experiment 1, we trained rhesus monkeys (Macaca mulatta) and humans (Homo sapiens) on adjacent pairs (e.g., AB, BC, CD, DE, EF) and tested them with previously untrained nonadjacent pairs (e.g., BD). In Experiment 2, we trained a second list and tested with nonadjacent pairs selected between lists (e.g., B from List 1, D from List 2). We then introduced associative competition between adjacent items in Experiment 3 by training 2 items per position (e.g., B?C?, B?C?) before testing with untrained nonadjacent items. In all 3 experiments, humans and monkeys showed distance effects in which accuracy increased, and reaction time decreased, as the distance between items in each pair increased (e.g., BD vs. BE). In Experiment 4, we trained adjacent pairs with separate 9- and 5-item lists. We then tested with nonadjacent pairs selected between lists to determine whether list items were chosen according to their absolute position (e.g., D, 5-item list > E, 9-item list), or their relative position (e.g., D, 5-item list     

Extracting functional equivalence from reversing contingencies.

In 2 experiments assessing acquired equivalence, human participants were initially presented with 4 cues, 2 of which were paired with 1 outcome and 2 of which were paired with a 2nd outcome. These contingencies were then reversed across several training blocks such that, although each cue was paired equally often with each of the two outcomes across blocks, cues A and B always signaled the same outcome within blocks (as did cues C and D). In both experiments, performance on a subsequent transfer discrimination was enhanced when participants were required to generalize between stimuli that had been paired with the same outcome within each block of training. Additional tests did not yield evidence of a bias toward a specific set of cue–outcome contingencies in either experiment. Moreover, interviews conducted at the end of Experiment 2 revealed that performance on the transfer discrimination was enhanced only in participants who discovered the equivalence relationships during initial training. The results challenge simple associative, and attentional, accounts of acquired equivalence and favor the view that this effect is mediated by comparisons of the similarity of adjacent cue–outcome structures. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Acquisition of automatic imitation is sensitive to sensorimotor contingency.

The associative sequence learning model proposes that the development of the mirror system depends on the same mechanisms of associative learning that mediate Pavlovian and instrumental conditioning. To test this model, two experiments used the reduction of automatic imitation through incompatible sensorimotor training to assess whether mirror system plasticity is sensitive to contingency (i.e., the extent to which activation of one representation predicts activation of another). In Experiment 1, residual automatic imitation was measured following incompatible training in which the action stimulus was a perfect predictor of the response (contingent) or not at all predictive of the response (noncontingent). A contingency effect was observed: There was less automatic imitation indicative of more learning in the contingent group. Experiment 2 replicated this contingency effect and showed that, as predicted by associative learning theory, it can be abolished by signaling trials in which the response occurs in the absence of an action stimulus. These findings support the view that mirror system development depends on associative learning and indicate that this learning is not purely Hebbian. If this is correct, associative learning theory could be used to explain, predict, and intervene in mirror system development. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Analysis of the content of configural representations: The role of associatively evoked and trace memories.

Two experiments examined the content of configural learning in rats. In Experiment 1, after simple pre-exposure to two hybrid contexts (AB and CD), rats acquired a configural discrimination involving two of the contexts (A and C) and two auditory stimuli (X and Y; AX→food, AY→no food, CX→no food, and CY→food). When rats were then placed in context B, they were more likely to respond to X than Y, and when they were placed in context D the reverse was the case. Experiment 2 demonstrated that rats can acquire a configural discrimination involving the presence of context (A) and its memory trace (a; AX→food, AY→no food, aX→no food, and aY→food). These results show that associatively provoked memories (Experiment 1) and memory traces (Experiment 2) can participate in configural discriminations. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Using context to resolve temporal ambiguity.

Three conditioned lick suppression experiments with rats examined the role of the context in the selection and integration of independently acquired interval relationships. In Experiment 1, rats were exposed to separate conditioned stimuli 1 and 2 (CS1–CS2) pairings with 2 different interval relationships, each in its own distinctive context, X or Y. The resultant integration was determined by the training context (X or Y) in which unconditioned stimulus (US)–CS2 backward pairings occurred, as assessed in a third neutral context (Z). In Experiment 2, rats experienced CS1–CS2 pairings with 2 different interval relationships as in Experiment 1, and then received US–CS2 pairings in both contexts X and Y. The testing context (i.e., X or Y) determined the resultant integration. In Experiment 3, rats were exposed to CS1–CS2 pairings in 2 different interval relationships each in different phases (i.e., Phases 1 and 2), and then in Phase 3 received US–CS2 pairings. The temporal context of testing (i.e., short or long retention interval) determined the resultant integration. Thus, both physical and temporal context can be used to disambiguate conflicting temporal information. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Effects of emotion on associative recognition: Valence and retention interval matter.

In two experiments, we examined the effects of emotional valence and arousal on associative binding. Participants studied negative, positive, and neutral word pairs, followed by an associative recognition test. In Experiment 1, with a short-delayed test, accuracy for intact pairs was equivalent across valences, whereas accuracy for rearranged pairs was lower for negative than for positive and neutral pairs. In Experiment 2, we tested participants after a one-week delay and found that accuracy was greater for intact negative than for intact neutral pairs, whereas rearranged pair accuracy was equivalent across valences. These results suggest that, although negative emotional valence impairs associative binding after a short delay, it may improve binding after a longer delay. The results also suggest that valence, as well as arousal, needs to be considered when examining the effects of emotion on associative memory. (PsycINFO Database Record (c) 2011 APA, all rights reserved)     

Hydrogen bonding, solvent exchange, and coupled proton and electron transfer in the oxidation and reduction of redox-active tyrosine Y(Z) in Mn-depleted core complexes of photosystem II

The redox-active tyrosines, Y(Z) and Y(D), of Photosystem II are oxidized by P680+ to the neutral tyrosyl radical. This oxidation thus involves the transfer of the phenolic proton as well as an electron. It has recently been proposed that tyrosine Y(Z) might replace the lost proton by abstraction of a hydrogen atom or a proton from a water molecule bound to the manganese cluster, thereby increasing the driving force for water oxidation. To compare and contrast with the intact system, we examine here, in a simplified Mn-depleted PSII core complex, isolated from a site-directed mutant of Synechocystis PCC 6803 lacking Y(D), the role of proton transfer in the oxidation and reduction of Y(Z). We show how the oxidation and reduction rates for Y(Z), the deuterium isotope effect on these rates, and the Y(Z)* - Y(Z) difference spectra all depend on pH (from 5.5 to 9.5). This simplified system allows examination of electron-transfer processes over a broader range of pH than is possible with the intact system and with more tractable rates. The kinetic isotope effect for the oxidation of P680+ by Y(Z) is maximal at pH 7.0 (3.64). It decreases to lower pH as charge recombination, which shows no deuterium isotope, starts to become competitive with Y(Z) oxidation. To higher pH, Y(Z) becomes increasingly deprotonated to form the tyrosinate, the oxidation of which at pH 9.5 becomes extremely rapid (1260 ms(-1)) and no longer limited by proton transfer. These observations point to a mechanism for the oxidation of Y(Z) in which the tyrosinate is the species from which the electron occurs even at lower pH. The kinetics of oxidation of Y(Z) show elements of rate limitation by both proton and electron transfer, with the former dominating at low pH and the latter at high pH. The proton-transfer limitation of Y(Z) oxidation at low pH is best explained by a gated mechanism in which Y(Z) and the acceptor of the phenolic proton need to form an electron/proton-transfer competent complex in competition with other hydrogen-bonding interactions that each have with neighboring residues. In contrast, the reduction of Y(Z)* appears not to be limited by proton transfer between pH 5.5 and 9.5. We also compare, in Mn-depleted Synechocystis PSII core complexes, Y(Z) and Y(D) with respect to solvent accessibility by detection of the deuterium isotope effect for Y(Z) oxidation and by 2H ESEEM measurement of hydrogen-bond exchange. Upon incubation of H2O-prepared PSII core complexes in D2O, the phenolic proton of Y(Z) is exchanged for a deuterium in less than 2 min as opposed to a t(1/2) of about 9 h for Y(D). In addition, we show that Y(D)* is coordinated by two hydrogen bonds. Y(Z)* shows more disordered hydrogen bonding, reflecting inhomogeneity at the site. With 2H ESEEM modulation comparable to that of Y(D)*, Y(Z)* would appear to be coordinated by two hydrogen bonds in a significant fraction of the centers.     

Blocking and Unblocking in Human Causal Learning.

Three experiments sought to develop the suggestion that, under some circumstances, common associative learning mechanisms might underlie animal conditioning and human causal learning, by demonstrating, in humans, an effect analogous to the unblocking by reinforcer omission observed in animal conditioning. Experiment 1 found no such effect. Experiment 2, designed to prevent inhibitory influences that might have masked excitatory unblocking in Experiment 1, demonstrated unblocking, indicating common human-animal associative learning mechanisms in which the associability of a stimulus varies as a function of its predictive history. Experiment 3, using a similar design but with a procedure promoting application of rational inference processes, failed to detect the same unblocking effect, indicating that associative and cognitive mechanisms may influence human causal learning. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Blickets and babies: The development of causal reasoning in toddlers and infants.

Previous research has suggested that preschoolers possess a cognitive system that allows them to construct an abstract, coherent representation of causal relations among events. Such a system lets children reason retrospectively when they observe ambiguous data in a rational manner (e.g., D. M. Sobel, J. B. Tenenbaum, & A. Gopnik, 2004). However, there is little evidence that demonstrates whether younger children possess similar inferential abilities. In Experiment 1, the authors extended previous findings with older children to examine 19- and 24-month-olds' causal inferences. Twenty-four-month-olds' inferences were similar to those of preschoolers, but younger children lacked the ability to make retrospective causal inferences, perhaps because of performance limitations. In Experiment 2, the authors designed an eye-tracking paradigm to test younger participants that eliminated various manual search demands. Eight-month-olds' anticipatory eye movements, in response to retrospective data, revealed inferences similar to those of 24-month-olds in Experiment 1 and preschoolers in previous research. These data are discussed in terms of associative reasoning and causal inference. (PsycINFO Database Record (c) 2010 APA, all rights reserved)