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).     

Comparison and Contrast as a Mechanism of Perceptual Learning?

In a series of flavour aversion experiments, rats received different schedules of pre-exposure to two compound flavours (AX and BX). Discrimination between them was assessed by establishing an aversion to AX and measuring generalization of this aversion to BX. Experiment 1 demonstrated that alternating pre-exposure to AX and BX resulted in less generalization than did blocked exposure, where animals received all exposure to AX before exposure to BX (or vice versa). This difference was not accompanied by any difference in the strength of the aversion conditioned to the common X element. Varying the interval between exposure to AX and BX in the alternating condition from a minute or two to several hours had no effect on generalization. However, Experiments 2 and 3 showed that when the interval between exposure to AX and that to BX was reduced to zero sec, the alternating schedule increased generalization between AX and BX. In this case, the increase in generalization was accompanied by an increase in the strength of the aversion conditioned to X.     

Alternating exposure to two compound flavors creates inhibitory associations between their unique features

Rats were exposed to two compound solutions, saline-lemon and sucrose-lemon. In Group ALT, trials with one solution alternated with trials with the other. Group BLK received all trials with one solution before any trials with the other. Previous retardation tests had implied that only alternating exposure would establish sucrose as an inhibitor of saline. To provide a complementary summation test for this inhibition, in Experiment 1, all the animals received pairings of peppermint and saline and were tested for consumption of peppermint-sucrose under sodium depletion. Consumption was increased by sodium depletion only in Group BLK. In Experiment 2, a retardation test was used to show that presentation of saline-lemon before sucrose-lemon on each exposure day would establish sucrose as an inhibitor of saline. Neither exposure to sucrose-lemon before saline-lemon nor alternating exposure to sucrose and saline alone had the same effect. These results provide support for an associative theory of perceptual learning that suggests that exposure to complex stimuli aids later discrimination partially as a result of establishing inhibitory associations between their unique elements.     

Evidence for inhibitory associations between the unique elements of two compound flavours

In each of two experiments, rats were pre-exposed to two flavoured solutions, saline-lemon and sucrose-lemon. For group ALT, trials with one solution alternated with trials with the other. Group BLK received all trials with one solution in a block, before any trials with the other. An associative theory suggests that the alternating, but not the blocked, schedule would establish an inhibitory association between sucrose and saline. To provide a retardation test of this inhibition, some animals in each group were then given a single pairing of saline and sucrose, experienced sodium depletion, and were finally tested for their consumption of sucrose. Sodium depletion increased consumption of sucrose more in group BLK than in group ALT. In groups given no saline-sucrose pairing, sodium depletion had only a small effect on sucrose consumption, which was the same in both groups. After multiple pairings of saline and sucrose, sodium depletion had an equally large effect on sucrose consumption in both ALT and BLK groups. These results imply that alternating pre-exposure to two compound solutions does establish an inhibitory association between their unique elements, and thus provide support for an associative theory of perceptual learning.     

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.     

Mediated configural learning in rats

Three experiments investigated mediated configural learning in male rats. In Experiment 1, after exposure to audio-visual compounds AX and BY, rats received trials where X was paired with shock, and Y was not. The idea that conditioning with X enables the evoked configural representation of AX to be linked to shock received support from the facts that while AX provoked more fear than BX, there was no difference between BY and AY. Similarly, Experiment 2 showed that after exposure to AX and BY, separate pairings of X and Y with shock resulted in more fear to AX and BY than AY and BX. In Experiment 3, rats in group consistent received separate exposures to A and X in Context C, and B and Y in D, while those in group inconsistent received A and X (and B and Y) in both C and D. After rats had received shocks in both C and D, rats in group consistent showed more fear to AX and BY than to BX and AY, but this was not the case in group inconsistent. These results indicate that configural representations, formed either by presenting auditory and visual stimuli as parts of a compound or in a shared context, are subject to a process of mediated learning.     

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.     

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.     

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.     

Taste preconditioning augments odor-aversion learning

On the basis of previous work that has shown a taste can potentiate odor-aversion conditioning in AX+ conditioning, 6 experiments used rats to examine the effects of pairing a preconditioned taste (A) with a novel odor cue (X) in an A+/AX+ aversion conditioning design. Experiments 1A and 1B demonstrated that a preconditioned taste produced a robust odor aversion that was significantly stronger than a potentiated odor aversion. The results of Experiment 2 showed that the robust odor aversion produced by A+/AX+ conditioning was not the result of the potentiated odor aversion summating with generalization from the taste aversion. The augmented odor aversion was produced only when the taste and odor stimuli were presented simultaneously (Experiment 3) and the preconditioned taste aversion was intact at compound conditioning (Experiment 4). Pairing a novel odor with a preconditioned taste was not sufficient to condition an aversion to odor (Experiment 5), although other results implicated a role for an association between odor and taste in the odor augmentation effect (Experiment 6). The present results have implications for current models of taste + odor interactions in flavor-aversion conditioning.     

Augmentation, not blocking, in an A+/AX+ flavor-conditioning procedure

An A+/AX+ Pavlovian conditioning design typically produces weakened or blocked conditioning to stimulus X. Two experiments were conducted in which rats first received an odor (A+) paired with an emetic US, and then received odor and taste (AX+) paired with the US. In both experiments, the preconditioned odorfacilitated conditioning to the taste. In Experiment 1, a group that received two odor-illness pairings in A+ conditioning had a stronger taste aversion than a group that only had a single odor-illness pairing. Experiment 2 demonstrated that the strengthened taste aversion in the A+/AX+ condition was not due to stimulus generalization. The results represent a unique outcome in the flavor-aversion literature that is similar to potentiation. We propose that this facilitated conditioning to X in the A+/AX+ design be termedaugmentation.     

The influence of word shading and word length on eye movements during reading

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.     

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.     

The effects of habituation training on compound conditioning are not reversed by an associative activation treatment

Rats received habituation to either 2 compound flavors (AX and BY; the activation group) or a compound and an element alone (AX and Y; the habituation group). They also received additional presentations of Y alone either after (Experiment 1) or intermixed (Experiment 2) with habituation. In the habituation group, A had undergone habituation whereas B had not; in the activation group, both A and B had undergone habituation, but presenting Y alone should result in associative activation of B and that, according to G. Hall (2003), should increase B's efficacy. A supplementary experiment demonstrated that the presentation of Y does activate a representation of B. In both experiments, an aversion was established to AB, and subsequently the habituation group showed a greater aversion to B than to A. However, in neither experiment was there any indication that the activation group showed a greater aversion to B than to A. These results are inconsistent with the suggestion that the associative activation of a stimulus representation in the absence of the stimulus reverses the effects of habituation training.     

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.     

Perceptual learning in flavour aversion conditioning

In each of four experiments, rats drank a solution of saline or of lemon and saline shortly before receiving an injection of lithium chloride, and the generalization of the resulting aversion to sucrose or to lemon and sucrose was measured. There was little generalization from saline alone to sucrose alone, and prior exposure to the two solutions had no effect on their discriminability. An aversion conditioned to lemon-saline, however, did generalize to lemon-sucrose, and the extent of this generalization was substantially reduced by prior exposure to the two compound solutions. This perceptual learning effect was partly, but not entirely, attributable to the latent inhibition of the common element, lemon, produced by exposure to the two compounds: animals pre-exposed to lemon alone discriminated between lemon-saline and lemon-sucrose better than animals pre-exposed to saline and sucrose alone; but exposure to the three elements in isolation was not as effective as exposure to the two compound solutions in enhancing their discriminability. The final experiment established that one critical feature of compound pre-exposure is that it involves experience of saline and sucrose in the presence of the same common element. According to an associative theory of perceptual learning, this would result in the establishment of inhibitory associations between saline and sucrose, thus reducing generalization between the two compound solutions.     

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.     

Inhibitory sensory preconditioning detected with a sodium depletion procedure

In each of two experiments, two groups of rats were exposed to three flavoured solutions: A (citric acid), B (salt), and AX (a compound of citric acid and saccharin). Both experiments used a between-subjects design in which a paired group received presentations of A followed by B, alternating with presentations of AX (i.e., A → B/AX), and an unpaired group received alternating presentations of A, B, and AX (i.e., A/B/AX). This arrangement was expected to establish X as an inhibitor of B in group paired but not in group unpaired. In Experiment 1, after preexposure all subjects received a single presentation of an XB compound, then experienced sodium depletion, and were tested for their consumption of X, which was greater in group unpaired than in group paired. In Experiment 2, after preexposure, all subjects received four presentations of a new flavour, C, in compound with B and subsequently, under sodium depletion, were tested for consumption of XC. Intake of the XC compound was less in group paired than in group unpaired. These results suggest that, in group paired, X acquired an inhibitory relationship with B both retarding the acquisition of an excitatory association with B (retardation test, Experiment 1) and reducing the response to a new stimulus, C, strongly associated to B (summation test, Experiment 2). These results provide direct evidence of inhibition between two neutral stimuli and, therefore, of inhibitory sensory preconditioning.     

Inhibitory sensory preconditioning detected with a sodium depletion procedure

In each of two experiments, two groups of rats were exposed to three flavoured solutions: A (citric acid), B (salt), and AX (a compound of citric acid and saccharin). Both experiments used a between-subjects design in which a paired group received presentations of A followed by B, alternating with presentations of AX (i.e., A → B/AX), and an unpaired group received alternating presentations of A, B, and AX (i.e., A/B/AX). This arrangement was expected to establish X as an inhibitor of B in group paired but not in group unpaired. In Experiment 1, after preexposure all subjects received a single presentation of an XB compound, then experienced sodium depletion, and were tested for their consumption of X, which was greater in group unpaired than in group paired. In Experiment 2, after preexposure, all subjects received four presentations of a new flavour, C, in compound with B and subsequently, under sodium depletion, were tested for consumption of XC. Intake of the XC compound was less in group paired than in group unpaired. These results suggest that, in group paired, X acquired an inhibitory relationship with B both retarding the acquisition of an excitatory association with B (retardation test, Experiment 1) and reducing the response to a new stimulus, C, strongly associated to B (summation test, Experiment 2). These results provide direct evidence of inhibition between two neutral stimuli and, therefore, of inhibitory sensory preconditioning.     

Assessments of changes in the effective salience of stimulus elements as a result of stimulus preexposure

Rats received exposure to 3 flavor compounds, AX and BX, presented in alternation, and CX, presented on a separate block of trials. The hypothesis that this treatment would leave B effectively more salient than C was tested in 3 ways. Experiment 1 showed that the unconditioned response evoked by B was stronger than that evoked by C. Experiment 2 showed that B was more effective than C when used as a reinforcer in a sensory preconditioning procedure. Experiment 3 showed that B was learned about more readily than C as a conditioned stimulus in flavor aversion conditioning. Alternating preexposure to 2 similar stimuli may protect their distinctive features from the loss of salience normally produced by nonreinforced exposure to a stimulus.