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 learning: Alternating and blocked exposure to a compound of flavors and to an element of that compound

Rats received different schedules of pre-exposure to a compound flavor (AX) and to one element of that compound (X). In Group ALT, exposure consisted of alternating trials with AX and X; Group BLK received a block with all AX trials before a separate block with all X trials (or vice versa). Discrimination between AX and X was assessed then by establishing an aversion to X and measuring the generalization of this aversion to AX. In Experiments 1A and 1B, generalization was less in Group ALT than in Group BLK. In Experiment 2, this latter result was confirmed and furthermore only Group ALT, and not Group BLK, showed less generalization than a group that received exposure to X alone. These results are discussed in terms of their implications for theories of perceptual learning.     

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.     

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).     

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.     

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.     

Potentiation and overshadowing in preweanling and adult rats

Four experiments compared the aversion acquired by 18-day-old and 60-day-old rats to a flavor that was either tasted alone or in combination, simultaneously or successively, with another flavor when paired with illness. The purpose was to study temporal variables and theoretical issues pertinent to potentiation and overshadowing while investigating ontogenetic differences in these phenomena. When either the preweanlings or the adults were presented a simultaneous compound flavor (sucrose/coffee) followed by lithium chloride-induced illness, greater sucrose aversions were found than for animals conditioned on sucrose alone--that is, potentiation. Preweanlings demonstrated greater potentiation than did adults, whether the conditioned stimulus-unconditioned stimulus (CS-US) delay was 0 or 1 hr. This potentiation was eliminated by nonreinforced presentations of the alternative CS element. Potentiation was not seen when the two flavors were presented successively as the CS; instead, overshadowing occurred. Tests of configuring by extinction procedures indicated a tendency for these animals to form a configured representation of the simultaneous compound solution. This disposition for configuring tended to be more pronounced for preweanlings than for adults. Ontogenetic differences in response to configuration, CS saliency, and generalization decrement seem consistent with at least one model of potentiation and the ontogenetic differences in potentiation seen in the present experiments.     

Overshadowing of a context aversion by a novel incentive in operant conditioning

Two experiments examined the processes underlying the suppression of instrumental behaviours by lithium in rats, as reported by Meachum (1988 and this issue). Experiment 1 examined whether presenting a novel sucrose solution prior to lithium chloride administration would overshadow aversion learning to either the stimuli of the operant chamber or to familiar food pellets. After lever pressing had been established, and in the absence of responding, animals received free deliveries of a novel sucrose solution, familiar food pellets, or both, or they were exposed to only the cues of the operant chamber, prior to lithium injections. Lever pressing for food pellets was then assessed. It was found that the animals receiving the novel sucrose, either alone or with the familiar food pellets, pressed more for pellets than either the group receiving only food pellets or the group exposed to only the context. In addition, there was no appreciable difference in the response rates between the context-only group and the group that received the familiar food pellets. These outcomes were interpreted in terms of the novel sucrose overshadowing aversion learning to the context. Experiment 2 investigated whether in fact aversive contextual conditioning could be obtained using the present parameters. This was accomplished by directly manipulating the contexts. In this experiment animals were trained to lever press in two distinctive contexts. Subsequently, one context was paired with the novel sucrose, and the second was experienced in the absence of reinforcement prior to toxicosis. During a subsequent non-reinforced test it was found that responding in the context paired with the novel sucrose was considerably higher than responding in the context that was experienced alone. These findings stand in contrast to the taste-mediated contextual potentiation observed when a consumatory response is used to assess aversive contextual conditioning.     

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 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.     

Odor-aversion learning and retention span in neonatal mouse pups

One hundred and sixty-four litters of Swiss CD-1 random-bred mice were used to assess learning and retention capacities during the first postnatal week. In Experiment 1, whole 7-day litters were exposed for 65 min to commercial extracts of either mint or lemon sprinkled over wood shavings. Five minutes after the beginning of the exposure, half of the litters were injected ip with the illness-inducing agent lithium chloride (LiCl; 0.20 M, 2% of body weight); the other half was treated with saline solution (8% NaCl). On Postnatal Day 10, the animals were singly introduced in a warmed arena for a 180-s preference test, and the time spent in the mint- and lemon-scented areas of the apparatus was recorded. When compared with saline-injected pups, mice that experienced lemon-LiCl pairings showed a significant aversion for the lemon-scented area, while the mint aversion in the mint-LiCl group just missed statistical significance. Three additional control groups (unhandled on Day 7, or only LiCl- or saline-injected) did not show significant preferences for either the mint or the lemon odor. In Experiment 2, litters of 3, 5, or 7 days were similarly exposed to lemon-scented shavings for either 5 or 20 min, injected with LiCl or saline, and then exposed for an additional 60 min to the shavings. On Postnatal Day 10, tests like those of Experiment 1 showed a significant odor-aversion in animals conditioned on Day 7, but not in those conditioned on Day 3 or 5. In Experiment 3, 3- and 5-day old pups were exposed to lemon odor-LiCl or -NaCl pairings, and tested for aversion after 3 or 7 days (CS duration 5 min before injection and either 30 or 60 min after injection). Only when the conditioning-testing interval was limited to 3 days did LiCl-injected groups show a significant aversion, which did not depend on duration of CS exposure.     

Taste aversion after ingestion of lithium chloride: An associative analysis

In five experiments with rats we examined the aversion established by consumption of a solution of lithium chloride (LiCl). Experiment 1 showed that consumption of LiCl established an aversion to saline (NaCl). Experiment 2 showed that the size of the aversion was reduced in rats given pre-exposure to saline (a latent inhibition effect). Experiment 3 showed that experience of a sucrose-saline compound prior to consumption of LiCl generated an aversion to sucrose (a sensory preconditioning effect). Experiments 4 and 5 examined the effects produced by consumption of a sucrose-LiCl compound and demonstrated reciprocal overshadowing between the two tastes. These results confirm that consumption of LiCl establishes an aversion to the taste of this substance. Their implications for the use of orally consumed LiCl as a technique for the control of predatory behaviour are discussed.     

Effects of prior exposure on conditioned taste aversion in the rat: androgen- and estrogen-dependent events

The effects of preexposure and gonadal hormone manipulation on the extinction of a conditioned taste aversion were investigated. In Experiment 1, male rats were given one prior exposure to sucrose at some selected time (Days 4, 2, or 1) before a second exposure (Day 0) to sucrose and an LiCl injection. Controls received no prior exposure to sucrose but experienced only the sucrose + LiCl condition (Day 0). under the single exposure condition (Day 0) castrated animals extinguished the aversion faster than either testosterone-treated castrated rats or sham-operated rats. Sham-operated and castrated rats that had received preexposure treatment (Days 4, 2, or 1) were not distinguishable on a within-groups comparison of behavior but differed from their respective controls by exhibiting faster rates of extinction. However, in the testosterone-treated group only the Day 4 preexposure group exhibited a faster extinction rate. In Experiment 2, estradiol, dihydrotestosterone, and testosterone were studied by using only a Day 1 preexposure condition. The testosterone-treated group maintained the aversion for the longest period of time, followed by dihydrotestosterone-treated, sham, castrated, and estradiol-treated groups. It appears that estradiol augments the castration effect whereas dihydrotestosterone attenuates the effect. In Experiment 3, estradiol was administered alone or in combination with two different doses of dihydrotestosterone, and a Day 1 preexposure condition was used. The findings indicate that the outcome of behavior is dependent on the ratio of estradiol to dihydrotestosterone, with variations in this ratio resulting in fast (estrogen effect) to slow (androgen effect) rates of extinction.     

Instrumental Outcome Devaluation Is Attenuated by the Anti-emetic Ondansetron

In three experiments we assessed the effect of an anti-emetic, the selective S-HT antagonist ondansetron, on (1) the conditioning of a taste aversion using lithium chloride (LiCl); (2) the expression of that aversion; and (3) instrumental outcome-devaluation effects. In Experiment 1 it was found that ondansetron reduced the aversion induced by LiCl when administered prior to the LiCl injection and also attenuated the expression of that aversion when administered prior to test sessions. In Experiments 2 and 3, thirsty rats were trained, in a single session, to lever press and chain pull for sucrose and saline solutions concurrently before being injected with LiCl. They were then re-exposed to both solutions, one after injection of vehicle and the other after injection of ondansetron. In a choice extinction test on the levers and chains, animals performed more of the action whose training outcome was re-exposed under ondansetron than the other action, whether the test was conducted after an injection of vehicle or after one of ondansetron.     

Taste aversion learning: simulation of interference with the gustatory cue during conditioning

Rats were taught an aversion to a sucrose taste cue of varying strengths. The concentration of the sucrose solution was either, 10, 7.5, 5, 2.5, 1, 0.5, or 0.25% which the animals drank for 5 min. Thirty minutes later they were poisoned with lithium chloride. On the test day all animals had access to a 10% sucrose solution regardless of the concentration they had drunk on the conditioning day. Animals conditioned with a 10, 7.5, or 5% sucrose cue subsequently displayed an identically strong aversion to the 10% cue. Only those animals conditioned with a sucrose cue which was 1% or less displayed a significantly weaker aversion to the 10% cue. The results are discussed in terms of the theory that interference with taste aversion learning by such agents as pentylenetetrazol and electroconvulsive shock may have their effect by disrupting the gustatory engram. If this assumption is correct then it suggests that the memory of the gustatory cue may be stored, at least prior to poisoning, in a quite labile state and an apparently limited disruption of taste aversion learning may in fact represent a substantial amnesic effect.     

Restoring the associability of a pre-exposed CS by a surprising event

Three experiments are reported which use rats and the conditioned suppression technique. The first two confirmed a previous finding that prior exposure to a stimulus predicting a weak shock retards further learning when this same stimulus is subsequently used to signal a stronger shock. They further showed that this loss of stimulus associability could be attenuated by inserting trials on which the stimulus was presented alone in the absence of shock before the phase of training with the stronger shock. Experiment III demonstrated that, for animals given prior exposure to two stimuli, the insertion of nonreinforced trials with one of the stimuli will restore the associability only of that stimulus. These results are taken to show that a surprising event (the omission of an expected shock) can restore the associability of a pre-exposed conditioned stimulus.     

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.     

The relative contributions of predictiveness and salience in flavor aversion learning

Two experiments investigated the relative influence of cue predictiveness and stimulus salience on flavor aversion learning. In the first study experimental subjects experienced a nonsalient strawberry flavor and a different, more salient flavor prior to illness on each of the first three conditioning trials; the nonsalient strawberry and salient lemon were paired with illness on the fourth trial. This treatment resulted in a strong aversion to the strawberry flavor and on aversion to the lemon flavor. In contrast, animals which had experienced only a single pairing of strawberry, lemon, and illness acquired a strong aversion to lemon and no aversion to strawberry, while animals which had experienced only the pairing of different flavors and illness on each of the first three trials developed a strong aversion to both strawberry and lemon. Experiment 2 replicated the results of the experimental condition of the first study but also included a “blocking” condition in which subjects received three strawberry and illness pairings prior to a strawberry-lemon-illness pairing. Subjects in the blocking condition developed only a weak aversion to strawberry, and this aversion did not block the development of a strong lemon aversion. Apparently, in order for a nonsalient flavor cue to block the acquisition of an aversion to a more salient cue, the nonsalient flavor must not only be paired with illness but also be more predictive of illness than the more salient flavors also paired with illness. The data were discussed in terms of the R. A. Rescorla and A. R. Wagner's associative model (1972, in Classical conditioning II: Current research and theory, New York: Appleton-Century-Crofts), R. A. Rescorla's catalytic model (1982, Journal of Experimental Psychology: Animal Behavior Processes,8, 131–141), and N. J. Mackintosh's attentional model (1975, Psychological Review,82, 276–298) of conditioning.     

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.     

Failure of serial taste-taste compound presentations to produce overshadowing of extinction of conditioned taste aversion

Two experiments were conducted to study overshadowing of extinction in a conditioned taste aversion preparation. In both experiments, aversive conditioning with sucrose was followed by extinction treatment with either sucrose alone or in compound with another taste, citric acid. Experiment 1 employed a simultaneous compound extinction treatment and found results indicative of overshadowing of extinction. By contrast, Experiment 2, in which extinction treatment involved serial compound presentations, failed to obtain evidence of overshadowing of extinction. The results of Experiment 2 indicate that the serial presentation of two tastes was processed as equivalent to the separate presentation of the tastes. The results are discussed in relation to: (1) Convergent evidence from research on latent inhibition, (2) competing theories of learning and, (3) their possible adaptive value in food-selection learning.