The Espinet and the perceptual learning effects in flavour aversion conditioning: do they depend on a common inhibitory mechanism?

In three experiments rats were given short or long preexposure (4 or 10 sessions) to two compound flavours, AX and BX, according to an intermixed or a blocked schedule. Following preexposure, aversion conditioning trials were given with AX as the conditioned stimulus (CS). In Experiments 1 and 2, retardation and summation tests were then carried out to assess the inhibitory properties of B (an Espinet procedure). In Experiment 3, test trials evaluated generalization from AX to BX (the standard perceptual learning procedure). The results showed that B performed as an inhibitor of the unconditioned stimulus (US; an Espinet effect) only after long intermixed preexposure, whereas a reliable perceptual learning effect was observed both after short and after long preeexposure. The observation that B had no detectable inhibitory properties after short preexposure casts doubt on the suggestion that inhibitory learning is responsible for perceptual learning after brief exposure to AX and BX.     

Salience modulation in serial preexposure: implications for perceptual learning

In three experiments rats were given serial preexposure to two flavor stimuli. In Experiment 1, some animals were given exposure to AX followed by the presentation of BX, a forward schedule; the others were given backward preexposure (BX→AX). Conditioning and test trials with the A element showed that salience or effectiveness of A was better protected in the forward than in the backward condition. Experiments 2 and 3 assessed the relevance of this salience modulation mechanism for perceptual learning. In these experiments, generalization of a conditioned aversion from AX to BX was reduced in the forward (but not in the backward) condition only after prolonged exposure, indicating that the establishment of an inhibitory link from B to A is required for successful discrimination. However, generalization to a novel compound stimulus, NX, was reduced in the forward group both after short and long preexposure, suggesting the existence of salience modulation processes that work in parallel with associative inhibition. These results seem to support the existence of a salience modulation mechanism that seems to be beyond the scope of current theories of perceptual learning.     

Enhanced discrimination between flavor stimuli: roles of salience modulation and inhibition

Rats were given intermixed preexposure to the compound flavors AX and BX and to the compound CX in a separate block of trials (4 presentations of each compound). In Experiment 1, rats showed less generalization of conditioned aversion from AX to BX than from CX to BX, a perceptual learning effect. Experiment 2 showed that the formation of an excitatory association proceeded more readily between A and B than between C and B, suggesting that intermixed preexposure maintains the effective salience of A and B and does not establish inhibition between them, a process that would require prolonged preexposure. According to this analysis, salience modulation and associative inhibition may contribute to perceptual learning at different stages of preexposure.     

Representation-mediated Inhibitory Learning in the Conditioned-suppression Procedure

In three experiments, rats in group I (for intermixed) were given non-reinforced exposure to two compound stimuli, AX and BX, where A and B represent different auditory cues, and X represents a visual cue. AX and BX were presented in alternation. Group B (blocked) received similar exposure except that subjects experienced a block of AX trials and then a block of BX trials. Subsequent shock reinforcement of A was found to endow B with inhibitory strength in group I, as assessed by retardation (Experiments 1 and 2) and summation tests (Experiment 3). This outcome confirms and extends the results reported by Espinet, Iraola, Bennett, and Mackintosh (1995) and constitutes a further example of mediated learning in which the associative strength of a stimulus is found to be modified as a consequence of training given to some other event with which that stimulus is associated.     

Analysis of the role of associative inhibition in perceptual learning by means of the same-different task

In Experiment 1a, participants were exposed, over a series of trials, to separate presentations of 2 similar checkerboard stimuli, AX and BX (where X represents a common background). In one group, AX and BX were presented on alternating trials (intermixed), in another, they were presented in separate blocks of trials (blocked). The intermixed group performed to a higher standard than the blocked group on a same-different test. A superiority of intermixed over blocked exposure was also evident in a within-subject design (Experiment 1b) and when the test required discrimination between a preexposed stimulus and the background (e.g., AX vs. X), even if the background changed between preexposure and test (AY vs. Y) (Experiment 2). In Experiment 3, the intermixed/blocked effect was observed when, in preexposure, stimulus presentations were alternated with the background alone (e.g., AX/X). This suggests that the perceptual learning effect is not the consequence of inhibitory associations between unique features but to increased salience of those features. Experiment 4 confirmed this finding and also ruled out an account of the effect in terms of trial spacing.     

Effects of preexposure on stimulus discrimination: an investigation of the mechanisms responsible for human perceptual learning

The effect of preexposure on human perceptual learning was investigated in four experiments. In Experiments 1a and 1b, participants were preexposed to one pair of visual stimuli on an intermixed schedule (AX/BX) and one on a blocked schedule (CX_DX). The ability to discriminate between AX and BX and between CX and DX was then assessed by examining the extent to which key presses assigned to each member of a stimulus pair generalized to the other member (Experiment 1a) and by looking at the accuracy of same-different responses (Experiment 1b). Stimuli were more easily discriminated following intermixed than following blocked preexposure on both the generalization and same-different tasks. This suggests that two stimuli are more perceptually distinct after intermixed preexposure. Experiments 2a and 2b investigated the mechanisms responsible for perceptual learning using same-different tasks. The results support the suggestion that the enhanced discrimination observed after intermixed preexposure is due to increases in the salience of the unique elements.     

Memory for, and salience of, the unique features of similar stimuli in perceptual learning

In two experiments, participants received exposure to complex checkerboards (e.g., AX and BX) that consisted of small distinctive features (A and B) superimposed on a larger common background (X). Subsequent discrimination between AX and BX, assessed by a same-different task, was facilitated when the stimuli were presented on alternate trials in preexposure--a perceptual learning effect (Experiment 1). The hypothesis that this form of exposure results in more accurate representations of the unique features was supported in Experiment 1, which showed that participants were well able to match the color of the feature with its shape. Experiment 2 showed that exposure to A and B in isolation, intermixed with presentations of AX and BX, enhanced the perceptual learning effect, which confirmed that the better encoding of the unique features during intermixed preexposure is a direct cause of the enhanced discrimination observed following preexposure on this schedule.     

Effects of preexposure on stimulus discrimination: An investigation of the mechanisms responsible for human perceptual learning

The effect of preexposure on human perceptual learning was investigated in four experiments. In Experiments 1a and 1b, participants were preexposed to one pair of visual stimuli on an intermixed schedule (AX/BX) and one on a blocked schedule (CX_DX). The ability to discriminate between AX and BX and between CX and DX was then assessed by examining the extent to which key presses assigned to each member of a stimulus pair generalized to the other member (Experiment 1a) and by looking at the accuracy of same–different responses (Experiment 1b). Stimuli were more easily discriminated following intermixed than following blocked preexposure on both the generalization and same–different tasks. This suggests that two stimuli are more perceptually distinct after intermixed preexposure. Experiments 2a and 2b investigated the mechanisms responsible for perceptual learning using same–different tasks. The results support the suggestion that the enhanced discrimination observed after intermixed preexposure is due to increases in the salience of the unique elements.     

Attention and relative novelty in human perceptual learning

Four experiments examined the role of attention in human perceptual learning. In Experiment 1, participants were preexposed to a pair of visual (checkerboard) stimuli AX and BX, with common elements X and unique features A and B. A same-different task was then used to assess discrimination of AX and BX and a pair of control stimuli, CY and DY. In addition, participants' eye movements were recorded to assess the role of attentional processes. The results showed that preexposure enhanced discrimination between AX and BX. Furthermore, participants showed greater attention to the preexposed unique features A and B than to the novel unique features C and D, as measured by the eye gaze monitor. Experiments 2 and 3 examined the prediction that perceptual learning is due to the relative familiarity of the common and unique stimulus features. Experiment 4 replicated the intermixed-blocked effect and showed that the way in which AX and BX are presented is also important for perceptual learning. The results generally support the idea that intermixed preexposure to AX and BX increases attention to the unique stimulus features A and B. Some aspects of the results are consistent with a relative novelty account, whereas others implicate a high-level attentional process that is not driven by stimulus novelty.     

Perceptual learning in flavor aversion: evidence for learned changes in stimulus effectiveness

Rats were exposed to the compound flavors AX and BX, presented in alternation, and to CX on a separate block of trials. Generalization to BX after aversion conditioning with AX was less than to CX. An equivalent effect was found when the nature of the common element was changed after preexposure but not when the common element was omitted during preexposure, during conditioning and test, or both. Rats conditioned with X alone again showed less aversion to BX than to CX; similarly, rats conditioned with a novel flavor (Y) showed less aversion to BY than to CY. These effects support the proposal that intermixed preexposure to AX and BX enhances the perceptual effectiveness of their unique features, A and B.     

Perceptual learning in humans: Roles of preexposure schedule, feedback, and discrimination assay

In three experiments, humans received preexposure to two compound flavours (AX and BX: saline-lemon and sucrose-lemon) that were presented either in an intermixed (e.g., AX, BX, . . . BX, AX, . . .) or a blocked (e.g., AX, AX, . . . BX, BX. . .) fashion. Subsequently, AXwas paired with an unpleasant bitter taste, and the discriminability of AX and BX was assessed using the accuracy of same/different judgements and by the extent to which any learned dislike of AX generalized to BX. When participants received feedback about the accuracy of their same/different judgements during preexposure those given intermixed preexposure were more accurate in making these judgements during the test than those given blocked preexposure (Experiments 1 and 2A), however, there was no evidence of any learned dislike in these experiments. In Experiment 2B, in which participants did not receive feedback about the accuracy of their judgements, there was no effect of the preexposure regime on same/different judgements, but there was a learned dislike of AX, and this generalized less to BX in participants given intermixed than in those given blocked preexposure. The beneficial effects of intermixed preexposure are consistent with results from other species (chicks and rats), but the differences created by the presence or absence of feedback place constraints on the analysis of these effects.     

Stimulus Preexposure, Comparison, and Changes in the Associability of Common Stimulus Features

In four experiments, rats received preexposure either to both of two compound flavours (AX and BX), or to just one (BX). Experiment 1 demonstrated a perceptual learning effect, showing that, for animals given preexposure to both flavours, an aversion conditioned to AX generalized only poorly to BX. Subsequent experiments assessed the properties of the common feature, X. Experiment 3 showed that the two preexposure treatments did not differ in the extent to which they produced habituation of the neophobia evoked by X. Experiment 2 showed that conditioning to X proceeded more rapidly in subjects given preexposure to both AX and BX than in subjects preexposed to BX alone. In Experiment 4, a similar effect was found when the elements of the compounds were presented serially. It is concluded that the perceptual learning effect of Experiment 1 occurs in spite of the fact that preexposure to two stimuli tends to maintain the associability of their common elements.     

Analysis of the perceptual learning effect in flavour aversion learning: Evidence for stimulus differentiation

Rats received exposure to two compound flavours, AX and BX, where A and B were sucrose and saline and X was acid. For group intermixed (I), exposure consisted of alternating trials with AX and BX; group blocked (B) received a block of AX trials and a separate block of BX trials. Experiment 1 showed that generalization to BX after conditioning with AX was less profound in group I than in group B. Separate examination of the elements of the compound showed that the source of this difference lay in the strength acquired by the X element. X acquired less strength in group I than in group B (Experiments 1 and 2), whereas for the A element (Experiments 3 and 4) the reverse pattern was obtained. These results support the proposal that the perceptual learning effect (restricted generalization from AX to BX in group I) depends on a process that enhances the effectiveness of unique stimulus elements (A and B) and reduces that of common elements (such as X).     

Memory encoding of stimulus features in human perceptual learning

Two experiments analysed memory encoding in human perceptual learning. Both experiments started with preexposure without feedback to four checkerboards composed by a unique feature each and sharing a common feature (AX, BX, CX, and DX). Elements of one pair were presented intermixed and elements of the other pair were presented in separate blocks. Immediately after preexposure participants completed a memory recognition task in which the characteristics of the distractors were manipulated. Experiment 1 showed that only intermixed presentation results in good encoding of the unique features of the stimuli. Experiment 2 demonstrated that intermixed preexposure results in different encoding of unique versus common features of the stimuli: Participants are able to remember A and B better than they remember X, whereas for the blocked condition memory for C, D, and X does not differ. Overall, the results presented here support the proposal that intermixing stimuli results in differential memory traces for unique versus common features and that this contributes to the intermixed/blocked effect.     

Inhibitory associations contribute to perceptual learning in humans

Perceptual learning in humans was examined in 2 experiments. In Experiment 1, participants received intermixed exposure to 2 similar compounds (AX, BX, AX, BX, . . .) and blocked exposure to a 2nd pair of similar compounds (CY, CY, . . . , DY, DY, . . .). Aversions established to AX and CY generalized less to BX than to DY. In Experiment 2, 1 pair of compounds was presented in a forward order (i.e., AX3BX), whereas the 2nd pair of compounds was presented in a backward order (i.e., DY-->CY). Aversions established to AX and CY generalized less to BX than to DY. These results indicate that inhibitory associations contribute to perceptual learning in humans and thereby establish a fundamental similarity between the mechanisms that underlie perceptual learning in humans and rats.     

The intermixed-blocked effect in human perceptual learning is not the consequence of trial spacing

A robust finding in humans and animals is that intermixed exposure to 2 similar stimuli (AX/BX) results in better discriminability of those stimuli on test than does exposure to 2 equally similar stimuli in 2 separate blocks (CX_DX)--the intermixed-blocked effect. This intermixed-blocked effect may be an example of the superiority of spaced over massed practice; in the intermixed, but not the blocked exposure regime, each presentation of a given stimulus (e.g., AX) is separated from the next by the presentation of its partner (BX). Two experiments with human participants replicated the intermixed-blocked effect and showed that the effect was not due to the spacing of exposure trials. A mechanism for the intermixed-blocked effect is proposed, which combines theories from associative learning and memory.     

Superior discrimination between similar stimuli after simultaneous exposure

Human participants received unsupervised exposure to difficult-to-discriminate chequerboard stimuli (e.g., AX and BX), before learning a discrimination between them. Experiment 1 demonstrated that prior exposure enhanced later discrimination and that intermixed exposure (AX, BX, AX, BX …) resulted in better subsequent discrimination than did blocked exposure (CY, CY … DY, DY …). Experiment 2 showed that simultaneous exposure to two similar stimuli (AX–BX, BX–AX …) facilitated the later acquisition of a successive discrimination, more than successive exposure (AX–AX, BX–BX …). These results parallel those observed by Mundy, Honey, and Dwyer (2007) who used pictures of human faces as stimuli and establish the generality of the fact that simultaneous exposure produces a particularly marked perceptual learning effect.     

Retrospective Revaluation and Inhibitory Associations: Does Perceptual Learning Modulate our Perception of the Contingencies Between Events?

The reported experiments seek to reconcile the conflict between proposed explanations of retrospective revaluation of contingency judgements in experiments with humans (e.g. Dickinson & Burke, 1996) and those of a perceptual learning phenomenon discovered in rats (Espinet, Iraola, Bennett, &Mackintosh, 1995). The experiments employ a medical diagnosis scenario, in which subjects are asked to judge whether particular symptoms signal the presence or absence of a fictitious disease. In Experiment 1, following alternating exposure to pairs of symptoms, AX and BX, subjects were informed that symptom A was predictive of illness. As a result of this training, subjects rated symptom B as predictive of the absence of the illness. In Experiment 2, a possible mechanism behind this effect was considered. In particular, it was established that pre-exposure in which the AX pair reliably preceded the BX pair produced the same pattern of results as those shown in Experiment 1. In contrast, when the order of pairs was reversed, so that BX reliably preceded AX, the reduction in ratings for symptom B was not obtained. Several extant associative theories are discussed, but none are found to account adequately for this pattern of results. To overcome these inadequacies, a parsimonious theory is suggested that explains all the effects observed in these experiments.     

Effects of varying the amount of preexposure to spatial cues on a subsequent navigation task

In each of two experiments rats were preexposed to four compound landmarks (AX, BX, CX, and DX) one at a time; they were then trained to find a submerged platform located in a fixed position in a swimming pool using these same landmarks. When the preexposure was SHORT (4 sessions) it facilitated subsequent learning (a perceptual learning effect), whereas when rats were given a LONG preexposure phase (8 sessions) this facilitatory effect disappeared. EXTRA-LONG preexposure (16 sessions) reversed the facilitatory effect-that is to say, we observed retarded learning. The results show that rats' ability to navigate towards an invisible goal is affected by the length of their preexposure to the spatial cues that signal the location of the goal. These data are consistent with an associative analysis of the swimming pool navigation task.     

Superior discrimination between similar stimuli after simultaneous exposure

Human participants received unsupervised exposure to difficult-to-discriminate chequerboard stimuli (e.g., AX and BX), before learning a discrimination between them. Experiment 1 demonstrated that prior exposure enhanced later discrimination and that intermixed exposure (AX, BX, AX, BX ...) resulted in better subsequent discrimination than did blocked exposure (CY, CY ... DY, DY ...). Experiment 2 showed that simultaneous exposure to two similar stimuli (AX-BX, BX-AX ...) facilitated the later acquisition of a successive discrimination, more than successive exposure (AX-AX, BX-BX ...). These results parallel those observed by Mundy, Honey, and Dwyer (2007) who used pictures of human faces as stimuli and establish the generality of the fact that simultaneous exposure produces a particularly marked perceptual learning effect.