Extinction of a saccharin—lithium association: Assessment by consumption and taste reactivity

Extinction of a conditioned palatability shift preceded extinction of conditioned taste avoidance whether rats were tested using a within-subjects design or a between-subjects design. In each of two experiments, consumption of 0.1% saccharin was paired with either 20 ml/kg of 0.15 M LiCl or equivolume physiological saline on a single trial. In Experiment 1, on each of 10 extinction trials, rats were given a taste reactivity test immediately prior to a consumption test. In Experiment 2, half of the rats were extinguished by taste reactivity testing and half of the rats were extinguished by a consumption test on each of 10 extinction trials. In both experiments, the aversive reactions of gaping and passive dripping were extinguished in a single trial and the suppression of ingestive reactions was extinguished in 2 trials; however, extinction of taste avoidance required 4–5 trials. These results suggest that rats continue to avoid a lithium-paired flavor even when they do not have an aversion to the taste.     

Reinstatement-induced recovery of a taste-LiCl association following extinction

Water-deprived rats were given a single exposure to saccharin and LiCl, either paired or unpaired. Half the subjects then received three saccharin-only exposures (extinction) in the training enclosure, followed by a single LiCl-only presentation (unconditioned stimulus reinstatement) 8 days after conditioning. The remaining subjects received six saccharin-only exposures, followed by LiCl reinstatement 13 days after conditioning. In both cases LiCl reinstatement occurred outside the training/test context. Appreciable recovery from extinction was observed after the partial loss of taste aversion obtained with three extinction sessions and the 8-day conditioning-reinstatement interval, but not after the asymptotic loss of taste aversion obtained with six extinction sessions and the 13-day conditioning-reinstatement interval. Conditioned taste aversions appear to be similar to more traditional associations with respect to both extinction and reinstatement-induced recovery from extinction. The results are discussed with reference to the event-memory, contextual-conditioning, and facilitated-retrieval hypotheses of postextinction reinstatement effects.     

Impact of brief or extended extinction of a taste aversion on inhibitory associations: Evidence from summation,retardation, and preference tests

In five conditioned taste aversion experiments with rats, summation, retardation, and preference tests were used to assess the effects of extinguishing a conditioned saccharin aversion for three or nine trials. In Experiment 1, a summation test showed that saccharin aversion extinguished over nine trials reduced the aversion to a merely conditioned flavor (vinegar), whereas three saccharin extinction trials did not subsequently influence the vinegar aversion. Experiment 2 clarified that result, with unpaired controls equated on flavor exposure prior to testing; the results with those controls suggested that the flavor extinguished for nine trials produced generalization decrement during testing. In Experiment 3, the saccharin aversion reconditioned slowly after nine extinction trials, but not after three. Those results suggested the development of latent inhibition after more than three extinction trials. Preference tests comparing saccharin consumption with a concurrently available fluid (water in Experiment 4, saline in Experiment 5) showed that the preference for saccharin was greater after nine extinction trials than after three. However, saccharin preference after nine extinction trials was not greater, as compared with that for either latent inhibition controls (Experiments 4 and 5) or a control given equated exposures to saccharin and trained to drink saline at a high rate prior to testing (Experiment 5). Concerns about whether conditioned inhibition has been demonstrated in any flavor aversion procedure are discussed. Our findings help explain both successes and failures in demonstrating postextinction conditioned response recovery effects reported in the conditioned taste aversion literature, and they can be explained using a memory interference account.     

An extinction cue reduces spontaneous recovery of a conditioned taste aversion

The effect of an auditory cue presented during extinction on spontaneous recovery of a conditioned taste aversion was investigated in three experiments. Experiment 1 demonstrated that the presence of the cue during extinction did not influence saccharin consumption during that phase, and that an aversion to saccharin in the absence of the cue was stronger at 18 days than at 1 day after extinction, representing spontaneous recovery rather than a renewal effect. Experiment 2 showed that a cue presented during extinction and testing reduced spontaneous recovery. Experiment 3 replicated that effect and showed that it depended on the cue’s correlation with extinction and not on an unconditioned effect; cues that had been presented during or prior to conditioning did not reduce spontaneous recovery when presented during testing. The cue’s potential to reduce spontaneous recovery through conditioned inhibition or configural cue learning is discussed, as is the possibility that the cue retrieves a saccharin extinction memory in a manner consistent with Bouton’s (1993) account of spontaneous recovery.     

Flavor-flavor associations induce hedonic shifts in taste preference

Two experiments allowed rats to drink freely two neutral flavors (almond and vanilla) in simultaneous compound with two hedonically valued flavors (quinine and saccharin). The neutral flavor previously paired with saccharin was subsequently preferred. The neutral flavor that had been paired with quinine was subsequently avoided. Experiment 3 found similar results when the animals were hand-fed a preset amount of the solution. Preference shifts were not obtained when differential amounts of the neutral flavors were consumed in isolation. The data indicate that flavor-flavor associations can shift taste preferences.     

A mixed design reveals that glucose moieties facilitate extinction of a conditioned taste aversion in rats

Separate groups of water-deprived rats had four trials with 15-min access to 0.0073 M saccharin, 0.3 M alanine, 0.3 M glucose, 0.1 M maltose, 0.3 M fructose, 0.06 M sucrose, or 0.03 M Polycose. Trials 1–3 were followed by injections of either 0.15 M LiCl (1.33 ml/100 g b.w., i.p.) or saline; Trial 4 (Test) was CS only. Extinction included either 48-h access to water alone or to the appropriate CS, both followed by a 24-h, two-bottle choice of CS and water. This 3-day cycle was repeated five to six times. All rats acquired comparable conditioned taste aversions (CTAs), but extinction rates varied with the test and the taste CS. No CTA extinguished during the two-bottle choices following 2 water days. During one-bottle CS exposure, all CTAs extinguished, but the aversion continued longer in the probe two-bottle tests. Intake of glucose moieties recovered rapidly, often in two cycles; the other CSs took four to six cycles. Thus, CTA extinction varies with the nature of the taste CS.     

Injected flavor as a CS in the conditioned aversion preparation

Three experiments were conducted to determine the effectiveness of intravenous (IV) flavor injections in the formation of conditioned taste aversions and in the attenuation of neophobia. In Experiment 1, two groups of rats were permitted to drink either a .1% saccharin solution or tap water followed immediately by IV injections of lithium chloride (LiCl), and two more groups were given IV injections of a 2% saccharin solution followed immediately by IV injections of either LiCl or distilled water. Injected flavor did not serve as an effective CS for the conditioning of an aversion to .1% saccharin. The second experiment employed a two-bottle procedure to detect attenuation of neophobia using the injected-flavor technique. It was found that, whether saccharin had been injected intravenously (2%), injected intraperitoneally (2% IP), or orally consumed (.1%), neophobia for .5% saccharin was attenuated equally relative to controls. CS-US intervals were manipulated in the final experiment such that IP injections of 2% saccharin solution were followed 0–480 min later by IP injections of LiCl. In this case, it was shown that injected flavor (2% saccharin) could act as an effective CS if the US was delayed (optimally about 120 min) and when the test solution was .1% saccharin. The delay gradient found in Experiment 3 was interpreted as a generalization gradient where optimum conditioning was displayed at the point where the concentration of saccharin circulating in the animal at the time of illness onset most closely matched the concentration of the test solution.     

The effect of drug habituation before and after taste aversion learning in rats

In Experiment I, rats received eight habituation injections of either lithium chloride (LiCl) or sodium chloride (NaCl), then two aversion training trials in which access to saccharin solution was followed by LiCl injections, and finally eight extinction trials with saccharin but no injections. The rats habituated to LiCl showed less aversion to saccharin during training and extinction. In Experiment II, rats received two aversion training trials, then eight habituation trials to either LiCl or NaCl, then eight extinction trials, four more aversion training trials, and eight more extinction trials. The rats habituated to LiCl did not differ during the first extinction period from those habituated to NaCl, but showed less aversion to saccharin during the second training and extinction periods. Consequently, habituation to LiCl reduces the learning of an aversion to saccharin but does not reduce the performance of a previously learned aversion.     

Second-order conditioning detects unexpressed morphine-induced salt aversion

Morphine failed to condition a salt taste aversion at a dose (15 mg/kg) sufficient to produce a robust aversion to a saccharin taste. Indeed, three different concentrations of salt (1%, 1.5%, and 2%) paired with the same morphine dose yielded no direct evidence for conditioned aversion. Yet, when a novel saccharin taste was paired in compound with the previously conditioned salt conditioned stimulus, we found evidence for a conditioning to the saccharin cue alone in three separate experiments. Control groups eliminated alternative accounts such as neophobia and differential exposure to morphine. Combined, these findings indicate that morphine conditioned a salt aversion. Although this aversion was not directly expressed, a second-order conditioning procedure was able to provide a more sensitive index of conditioning.     

Second-order conditioned taste aversion in rats: Shared modality is not sufficient to promote an association between S2 and SI

Second-order conditioning (SOC) in pigeons, but not rats, appears to involve an association between the second-order stimulus (S2) and the first-order stimulus (SI). Nairne and Rescorla (1981) suggested it was the use of stimuli from the same modality that promoted an association between S2 and SI in pigeon SOC studies. In support of their hypothesis, they demonstrated that pigeons, like rats, did not form an association between S2 and SI when these stimuli were from different modalities. In this study, we sought to determine whether rats, like pigeons, would associate S2 with SI when these stimuli shared the same modality. Female Lister rats injected with LiCl after consuming .12M saline solution (SI) showed an aversion to a 15% sucrose solution (S2) that was subsequently paired with the saline. This was so regardless of whether S2 and SI had been presented sequentially (Experiment 1) or simultaneously (Experiment 2). Only in Experiment 2, however, did extinction of the aversion to saline diminish the aversion to sucrose; that is, employing stimuli from the same modality was not a sufficient condition, of itself, to allow rats to associate S2 with SI.     

Taste and emotionality in rats selectively bred for high versus low saccharin intake

Rats were selectively bred for high versus low saccharin ingestion, a putative measure of enhanced stress and emotionality (Dess, 1991). In Experiment 1, third-generation Occidental high-saccharin (HiS) and low-saccharin (LoS) rats were tested for saccharin ingestion and emotionality. The saccharin test confirmed that the lines differed on the selection phenotype. In addition, LoS rats were more emotional, as evidenced by longer emergence latencies and more defecation in a modified open-field test. In Experiment 2, LoS rats had lower quinine preference scores and drank saccharin-adulterated glucose less avidly. These outcomes are reminiscent of the behavior shown by inescapably shocked rats. Unlike helpless rats, however, LoS rats drank less avidly during a dilute sucrose test, an effect more reminiscent of chronic mild stress. The lines did not differ reliably on intake of concentrated glucose or Polycose, even when the latter was mixed with saccharin. In Experiment 3, LoS rats preferred saccharin less strongly than did HiS rats at concentrations of 0.05% to 0.7% and had an aversion to a 1.0% solution. In Experiment 4, LoS rats were affected more by shock, as assessed by stress-induced anorexia. These and other recent findings support the notion of shared mechanisms for taste, emotionality, and stress vulnerability.     

Slow reacquisition of a conditioned taste aversion

Rats were used to examine the extent to which extinction of an acquired conditioned taste aversion retards subsequent reacquisition. A saccharin-flavored solution (sac) was paired with LiCl and then followed by CS-alone extinction trials with this flavor. A control group received a different flavor, decaffeinated-coffee (coff), during initial conditioning and extinction. Sac was then paired with LiCl for all rats during a second conditioning phase. Reacquisition of the aversion to sac was retarded relative to the acquisition of an aversion to sac by the control group. A similar experiment with fewer extinction trials, but still with complete loss of the initial aversion, did not obtain slow reacquisition. The results are discussed with respect to an interference view of extinction and the slow-reacquisition effect.     

Habituation to illness: Effects of prior experience with the US on the formation of learned taste aversions in rats

Experiment 1 investigated the effects of US habituation on the acquisition and extinction of learned taste aversions in rats. Subjects receiving five noncontingent LiCl intubations prior to conditioning failed to develop a conditioned taste aversion, while control subjects experiencing a single saccharin/LiCl pairing displayed a pronounced taste aversion which weakened during subsequent poisonings. Experiment 2 examined whether habituation, defined as a waning of responses to repeated presentation of an illness stimulus, was a possible mechanism for explaining the results of Experiment 1. Subjects showed a decrease in motor activity following an initial LiCl intubation, but less attenuation of activity with successive intubations.     

Taste aversion learning and schedule-induced polydipsia in rats

Experiment 1 sought to determine whether schedule-induced drinking could be abolished by means of a taste aversion. Polydipsic rats were given access to a .4% saccharin solution while they were exposed to an intermittent food schedule. Immediately after the session, they received an intraperitoneal injection of either lithium chloride or sodium chloride. Following a recovery day with water in the experimental chamber, the animals were again exposed to the saccharin solution. The poisoned animals (lithium chloride) drank very little saccharin compared to the control animals (sodium chloride), indicating that they had learned a taste aversion in only one conditioning trial. Experiment 2 established that polydipsic rats can learn a taste aversion despite a long delay between schedule-induced saccharin consumption and poisoning, and that the delay gradient displayed by polydipsic rats is similar to that observed in thirst-motivated rats.     

Superlatent inhibition and spontaneous recovery: Differential effects of pre- and postconditioning CS-alone presentations after long delays in different contexts

In two pairs of three-stage conditioned taste aversion experiments, we examined the effects of delay interval (1 or 21 days) between the second and third stages, and of context in which the animals spent the delay (same as or different from the context of the other stages) on latent inhibition (LI) and spontaneous recovery following extinction. In the LI experiments (Experiments 1A and 1B), the first stage comprised nonreinforced presentations to saccharin or to water. In the second stage, rats were conditioned by saccharin paired with LiCl. In the extinction experiments (Experiments 2A and 2B), the order of the stages was reversed. For all experiments, Stage 3, the test stage, consisted of three presentations of saccharin alone. There was a super-LI effect in the saccharin-preexposed group that spent the 21- day delay in the different context (Experiment 1A). When the delay was spent in the same context, there was no difference in the amount of LI between the short- and long-delay groups (Experiment 1B). Conversely, there was a spontaneous recovery effect in the long-delay/same-context group (Experiment 2B), but not in the long-delay/different-context group (Experiment 2A). The pattern of results, incompatible with current explanations of delay-induced changes in memory performance, was interpreted in terms of an interaction between the delay conditions (same or different delay context), which modulate the extinction of previously acquired context-CS-nothing associations (during CS-alone presentations), and primacy effects.     

Conditioning with taste as the CS in conditioned flavor preference learning

There appear to be unconditioned affective reactions to the four basic tastes: liking for sweet and salt and disliking for sour and bitter. We attempted to modify these reactions by pairing the tastes with calories and with sweeter tastes. Differing sucrose concentrations were ineffective in reversing the preference for salt over citric acid or for saccharin over quinine. We could, however, reverse the preference for salt over citric acid, producing an actual preference for sour over salt, by using sucrose and saccharin as the reinforcers. The initial reactions to tastes could also be modified by reducing the initial difference in affect produced by the tastes through mixing both tastes with sucrose.     

Role of exteroceptive background context in taste-aversion conditioning and extinction

In two experiments, saccharin (CS) and lithium chloride (US) were paired in a context consisting of specific visual, auditory, tactual, and olfactory cues. The saccharin aversion was then extinguished in a context free from conditioning-context cues. Later, saccharin preference tests were given in the presence and absence of these cues. The results indicated that the background cues of the conditioning trial controlled the amount of saccharin drunk on extinction trials, and, furthermore, that extinction of the taste aversion was context specific; i.e., groups given extinction trials in a different (from conditioning) context retained their saccharin aversion in the conditioning context only. The results indicate an important role played by the exteroceptive context in taste-aversion conditioning.     

Nonreinforced flavor exposure attenuates the effects of conditioned taste aversion on both flavor consumption and cue palatability

Nonreinforced exposure to a cue tends to attenuate subsequent conditioning with that cue—an effect referred to as latent inhibition (LI). In the two experiments reported here, we examined LI effects in the context of conditioned taste aversion by examining both the amount of consumption and the microstructure of the consummatory behavior (in terms of the mean size of lick clusters). The latter measure can be taken to reflect affective responses to, or the palatability of, the solution being consumed. In both experiments, exposure to a to-be-conditioned flavor prior to pairing the flavor with nausea produced by lithium chloride attenuated both the reduction in consumption and the reduction in lick cluster sizes typically produced by taste aversion learning. In addition, we observed a tendency (especially in the lick cluster measure) for nonreinforced exposure to reduce neophobic responses to the test flavors. Taken together, these results reinforce the suggestion from previous experiments using taste reactivity methods that LI attenuates the effects of taste aversion on both consumption and cue palatability. The present results also support the suggestion that the failure in previous studies to see concurrent LI effects on consumption and palatability was due to a context specificity produced by the oral taste infusion methods required for taste reactivity analyses. Finally, the fact that the pattern of extinction of conditioned changes in consumption and in lick cluster sizes was not affected by preexposure to the cue flavors suggests that LI influenced the quantity but not the quality of conditioned taste aversion.     

Sexually dimorphic extinction of a conditioned taste aversion in rats

The existence of a sexual dimorphism in the extinction of conditioned taste aversions in both fluid-deprived and nondeprived rats was investigated. When nondeprived rats were poisoned with LiCl after consuming a sucrose solution, males extinguished the subsequent conditioned taste aversion more slowly than females. There was no difference in the rate of extinction of a LiCl-induced conditioned taste aversion in fluid-deprived rats. Using marihuana extract distillate as the toxin, it was shown that the dimorphism found in the nondeprived rats is not a toxin-specific effect, and does not appear to be due to the males being made more ill. Possible explanations for the appearance of the dimorphism and its relation to the deprivational state of the organism are discussed.     

Cross-modality transfer effects in conditioning-enhanced neophobia in chicks (Gallus domesticus): Evidence for the separability of novelty from specific stimulus characteristics

In this study, we investigated the separability of novelty from specific stimulus characteristics (e.g., color or taste quality) in the transfer of aversion effects. Ninety-six chicks (Gallus domesticus) received a novel visual (red water) or taste (3.0% vinegar) CS paired with an injection of lithium chloride or saline. The chicks were then tested for aversion to the CS or for conditioning-enhanced neophobia in response to a different novel visual cue (green water) or taste cue (1.0% saline). Aversions to the CSs were reliable and similar to each other. Reliable evidence of conditioning-enhanced neophobia occurred with respect to each test stimulus, irrespective of the type of CS, but conditioning with the vinegar CS produced reliably greater enhancement of neophobia than did conditioning with red water. For each CS, postconditioning neophobia was more persistent in testing with saline than with green water. The results for postconditioning neophobia suggested that novelty is a general stimulus property that is separable from specific stimulus characteristics.