The role of amount consumed in flavor preexposure effects and neophobia

Rats received either a small or a large amount of a novel saccharin solution prior to a conditioning episode in which the saccharin flavor was paired with lithium-induced toxicosis. When the time between the preexposure and the conditioning episode was 3.5 h, both the small and large amounts of saccharin decremented conditioning. However, when the time between the preexposure and the conditioning episode was 23.6 h, only the consumption of a large amount of saccharin decremented conditioning. In a second experiment, rats received either a short or a long exposure to the novel saccharin solution and were then given a choice test between the saccharin and water. Rats given the choice test immediately following their preexposure to saccharin avoided consuming it. If they received the choice test 4 h later, they consumed more of the saccharin irrespective of the length of the preexposure. In contrast, when they were given a choice test 24 h after the preexposure, only rats that had received a long preexposure displayed a significant preference for the saccharin. The present results demonstrate that the flavor preexposure effect and the attenuation of neophobia are both determined by the time between preexposure to the novel flavor and the conditioning episode or choice test and the amount of preexposure to the novel flavor. These findings are discussed in relation to the information-processing model developed by Wagner (1976, 1978) and are used to provide an account, in terms of this model, of the delay-gradient seen in flavor-aversion learning.     

The effect of nonnutritive satiation on the learning of a flavor preference by rats

On each day of training in Experiment 1, hungry rats were given one flavored saccharin solution followed by a differently flavored saccharin solution. The rats drank more of the first flavor during training, but preferred the second flavor in a subsequent choice test. In Experiment 2, the two flavored saccharin solutions were provided on alternate days, with one flavor being preceded by nothing and the other flavor by plain saccharin. The rats drank more of the flavor preceded by nothing during training, but preferred the other flavor in a subsequent choice test. These results suggest that a state of nonnutritive satiation can reinforce a flavor preference.     

Effects of expected vs. unexpected proximal US preexposure on taste-aversion learning

Animals were first conditioned to expect lithium treatment following exposure to one taste solution (the CS+) and to expect no drug treatment following exposure to another flavor (the CS?). All subjects then received a saccharin taste-aversion conditioning trial. In Experiment 1, this conditioning trial was preceded 0, 1, 2, 4, or 6 h earlier by exposure to the CS+ flavor for independent groups. The CS+ exposure attenuated saccharin aversion learning if it occurred immediately before the saccharin conditioning trial but not if it occurred 1 h or more before conditioning. In Experiment 2, the saccharin conditioning trial was preceded 3 or 4.5 h earlier by a lithium injection. This proximal US preexposure injection was either unannounced (Li) or preceded by exposure to the CS+ (CS+Li) or the CS? (CS?Li) stimuli. The US preexposure attenuated saccharin aversion learning in all cases. However, the interference effect was less when the preexposure injection was expected (CS+Li) than when it was unexpected (CS?Li). This outcome could not be explained in terms of direct effects of the CS+ and CS? stimuli on the saccharin conditioning trial, and shows that the proximal US preexposure effect is a function of not only the drug dosage and preexposure interval, but also the anticipation of the drug pretreatment.     

The effects of taste extinction on ingestional potentiation in weanling rats

Four experiments investigated taste potentiation in weanling rats. In Experiment 1, the animals that drank a conditioning compound of denatonium and saccharin consumed significantly less on the test than controls that drank only saccharin during conditioning. This enhanced saccharin aversion was decremented by postconditioning extinction to denatonium in Experiment 2, and no generalization of saccharin aversions to the denatonium was observed in Experiment 3. Extinction of either saccharin or denatonium aversions after compound conditioning was shown in Experiment 4 to result in substantial decrements in aversions to the compound. The relationship of these outcomes to a multiple-association account of potentiation and to the role of discrimination processes in ingestional learning is discussed.     

Saccharin aversions induced by lithium chloride toxicosis in a backward conditioning paradigm

Seven experimental groups of seven rats each were allowed to consume saccharin solution at different times relative to intubation of lithium chloride solution. Six backward conditioning (BWD) groups were intubed 0.5, 1, 2, 3, 4, and 8 h before saccharin consumption, and a forward conditioning (FWD) group was intubed 0.5 h after saccharin consumption. A no-lithium control group of 14 rats received no intubation. Only the 0.5-h FWD and the 0.5-h BWD groups showed an aversion to saccharin relative to the no-lithium controls. The aversion to saccharin in the 0.5-h FWD group was more pronounced than that in the 0.5-h BWD group. This shows that the aversive effects of lithium toxicosis dissipate far sooner than the aversive effects of X-irradiation.     

Taste quality and extinction of a conditioned taste aversion in rats

Rats (Rattus norvegicus) that received a taste cue (saccharin, saline, quinine, or sucrose) paired with a lithium chloride (LiCl) injection displayed a robust decrease in consumption of that taste, relative to controls that had the taste unpaired with LiCl. Consumption of the paired taste increased with each nonreinforced presentation (i.e., extinction). After asymptotic extinction, rats that had had a 0.1% saccharin cue paired with LiCl consumed less of the saccharin solution than did controls. A similar data pattern was observed with a 10% sucrose solution. These results are consistent with the view that some aspect of the excitatory CS-US association remains after extinction. On the other hand, rats that had a bitter (0.005% or 0.001% quinine) or salty (1% or 0.5% saline) solution paired with LiCl drank similar amounts of the fluid as controls after asymptotic extinction treatment. Together, these experiments suggest that a taste that is either sweet or preferred is required in order to demonstrate the chronic decrease in fluid consumption after extinction treatment. The data suggest that the conditioning experience prevents the later development of a preference for the sweet taste, rather than there being a retained aversion that suppresses fluid consumption.     

Independence of neophobia and taste aversion learning

Animals were presented with (1) one solution which differed from that of the test solution, (2) a series of distinctly flavored solutions whose flavors differed from that of the test solution, or (3) with a flavored solution whose flavor was the same as that of the test solution. When animals received the solution whose flavor was the same as that of the test solution prior to a test for neophobia and prior to a conditioning trial, neophobia was reduced and aversions were weakened. However, when animals received a solution or a series of solutions whose flavors differed from that of the test solution, neophobia was reduced but conditioned aversions were unaffected. Presentations of solutions that differed from the test solution following aversion formation left the association between the taste of the test solution and the effects of the aversion-inducing treatment intact. In a final experiment it was discovered that neophobia was reduced as much when animals drank solutions whose flavors changed every third day as when they drank the same solution throughout testing.     

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.     

Effects of food deprivation on learning and expression of flavor preferences conditioned by saccharin or sucrose

In four experiments, food deprivation was varied during conditioning and testing of conditioning of flavor preferences by sweeteners. Conditioned preferences for a flavor associated with a more concentrated solution were enhanced by increased deprivation in training whether sucrose or saccharin was used when rats consumed solutions freely during training. When consumption of solutions was controlled and higher deprivation levels were used, preference for the higher concentration of sucrose was still enhanced by increased deprivation in training, but this did not occur with saccharin. We suggest that deprivation may enhance the reinforcing value of sweetness only when calories increase along with sweetness. We also suggest that deprivation can enhance flavor preference learning by increasing consumption and thereby increasing exposure to the flavored solutions.     

Higher order conditioning of a taste aversion

Two groups of rats each received 5 drops of NaCl solution from a dropper placed directly inside the mouth. The experimental group was then injected with lithium chloride to establish a conditioned taste aversion. The control group was injected 24 h later. After a recovery day the above procedures were repeated. On the next day both groups received 5 drops of a saccharin solution followed immediately by 5 drops of the NaCl solution. Subsequent preference tests established that the experimental group had learned an aversion to the saccharin solution as a result of its pairing with the NaCl solution which had previously been associated with poisoning. These results demonstrate that higher order conditioning of a taste aversion can be established using tastes as both the first-order stimulus and the second-order stimulus.     

The effect of nonreinforced stimulus exposure on the strength of a conditioned taste aversion as a function of retention interval: Do latent inhibition and extinction involve a shared process?

Two experiments examined the effects of nonreinforced flavor exposure on the strength of a conditioned taste aversion. Rats were conditioned by pairing maple flavor with LiC1. Prior to or subsequent to this pairing, some animals received nonreinforced exposure to either maple or saccharin. In separate subjects, preference for maple was tested 1 or 21 days after the last training episode. In the first experiment, the nonreinforced stimulus exposure occurred before conditioning (latent inhibition, or LI, procedure); in the second experiment, the nonreinforced exposure occurred after conditioning (extinction, or EXT, training). In both experiments, nonreinforced exposure to maple or saccharin reduced the magnitude of a conditioned maple aversion when testing occurred soon after conditioning. When testing was delayed, however, the attenuation due to nonreinforced saccharin exposure dissipated, both with the LI procedure and with EXT. In contrast, the nonreinforced exposure to maple was found to attenuate conditioned reactions at both short and long retention intervals. The release from generalized LI and spontaneous recovery following generalized EXT training are discussed in terms of retrieval processing. The possibility that the same mechanism may underlie LI and EXT is considered.     

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 effects of stimulus preexposure on taste-mediated environmental conditioning: Potentiation and overshadowing

On four occasions, Holtzman rats drank saccharin in a distinctive environment prior to lithium-induced toxicosis. Preconditioning exposure to saccharin either in the home cage or in the distinctive environment interfered significantly with the establishment of an environmental aversion. Animals preexposed to the experimental environment, however, showed environmental aversions substantially stronger than those in animals preexposed to saccharin and only slightly higher than those with no preexposure to either the taste or the environment. Subsequent saccharin tests revealed significantly stronger aversions in the group that received environmental preexposure than in any of the other groups. This pattern of outcomes demonstrates taste-mediated potentiation of novel and familiar environmental stimuli as well as overshadowing of the taste by novel environmental stimuli. Furthermore, it indicates that previous demonstrations of taste-mediated environmental potentiation involve facilitated conditioning of the environmental stimuli and decremented conditioning of the taste stimuli.     

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.     

Posttraining flavor exposure in hungry rats after simultaneous conditioning with a nutrient converts the CS into a conditioned inhibitor

The present study investigated the decrement in nutrient-based conditioned flavor preference found in hungry rats exposed to a flavor following simultaneous flavor–sucrose conditioning while thirsty. Although a significant decrease in preference was found in the experimental group in each experiment, there was no evidence of either spontaneous recovery (Experiment 1) or reinstatement (Experiment 2). In addition, posttraining flavor exposure weakened the original flavor–sucrose association (Experiment 3). These results suggested that the flavor–US association might have been impaired after posttraining flavor exposure. Two further experiments assessed whether the flavor acquired the properties of a net inhibitor, using the retardation and summation tests for conditioned inhibition. Experiment 4 revealed that the flavor suffered retardation when retraining was conducted after the exposure phase. In Experiment 5, the target flavor decreased the preference shown for a different flavor previously paired simultaneously with sucrose when both were presented forming an unreinforced compound in the summation tests. None of these effects was found in a control group, which had received serial flavor → nutrient presentations during training. Together, these results suggest that a flavor simultaneously paired with sucrose acquires the properties of a net inhibitor when it is subsequently presented outside the compound to hungry animals.     

Poison-induced neophobia in rats: Role of stimulus generalization of conditioned taste aversions

Rats repeatedly injected with lithium chloride were subsequently tested drinking novel and familiar solutions of both casein hydrolysate and vinegar. Injections in the absence of edibles result in only a small, and sometimes not reliable, increased avoidance of the novel casein and vinegar solutions. In contrast, if subjects acquired an aversion to saccharin as a result of the lithium injections, this learned aversion generalized to casein hydrolysate, with the generalization greatly enhanced by novelty of the casein flavor. However, the saccharin aversions did not generalize to the novel vinegar solution nearly as much as to the novel casein flavor. These results suggest that previous observations of poison-induced neophobia were probably in part a result of the stimulus generalization of conditioned taste aversions and that in addition to test stimulus novelty some other factor, such as stimulus salience or similarity to the conditioned aversive flavor, is also involved in the generalization of learned taste aversions.     

Specificity of cue to consequence in aversion learning in the rat: Control for US-induced differential orientations

In four experiments, each using a single conditioning trial, rats avoided a light more than a saccharin solution after these stimuli had been paired with footshock, whereas saccharin was avoided more than the light after these stimuli had been paired with lithium injection. The use of a single conditioning trial precludes possible US-induced differential orientations from influencing which stimuli will be associated on the conditioning trial. This cue-consequence specificity effect was obtained even when subjects conditioned with lithium received a non-contingent footshock prior to the test session, and when subjects conditioned with footshock received a noncontingent lithium injection before testing (Experiments 2–4). Weak aversions to the light in rats given a light-lithium pairing and noncontingent footshock and to the saccharin in subjects that received a saccharin-footshock pairing and noncontingent lithium administration were obtained in Experiment 2. However, these weak aversions were not obtained when subjects were given three nonreinforced exposures to the test chamber before the test session (Experiments 3 and 4). These results indicate that US-induced differential orientations do not mediate the cue-consequence effect in aversion learning.     

Learned flavor preferences based on calories are independent of initial hedonic value

In three experiments, the learning of flavor preferences due to pairing with calories was examined. In Experiment 1, the relative hedonic values of four isocaloric solutions and saccharin were assessed by offering these substances simultaneously to naive rats. The caloric solutions were then used to condition a flavor preference in separate groups of rats. Although the solutions were reliably different in unconditioned hedonic value, the conditioned flavor preferences were identical. In Experiment 2, we compared solutions of sucrose and saccharin that were equal in unconditioned hedonic value. Only the sucrose conditioned a preference. Finally, in Experiment 3, preferences were found to be sensitive to the number of calories available during conditioning. These results are discussed in terms of a peripheral cholecystokinin (CCK) reflex and the integration of that information along with taste information at the area postrema (AP) and surrounding nuclei. It is proposed that CCK acts centrally to adjust the incentive motivation or hedonic value of flavors.     

Recovery from conditioned suppression to a partially overlapping compound stimulus

In a conditioned suppression paradigm, a partially overlapping compound stimulus signaled occurrences of electric shock. That compound CS consisted of 3 min of continuous illumination of the houselight with three discrete 5-see presentations of a tone superimposed. A .5-sec electric shock was coterminous with each tone presentation. Rats that received this treatment in early conditioning sessions showed considerable recovery from conditioned suppression to the houselight as the experiment progressed. However, the effect was not reversible, and it could not be demonstrated in rats that experienced extensive prior conditioning to the houselight alone. These results are discussed in relation to a hypothesis concerning the modulation of behavioral control exerted by elements of compound stimuli and as they relate to a recent theoretical model for Pavlovian fear conditioning.     

Potentiation of taste and extract stimuli in conditioned flavor preference learning

In these experiments, we investigated the nature of potentiation in the conditioned flavor preference paradigm. Almond and banana extracts, which have strong odor components, were combined with salt and saccharin (liked tastes; Experiment 1) or quinine and citric acid (disliked tastes; Experiment 2) in a flavor preference procedure that mixed these solutions with a caloric reinforcer (polycose). The results showed that liked tastes potentiated preference conditioning to extracts (Experiment 1), whereas extracts potentiated preference conditioning to disliked tastes (Experiment 2). In both experiments, the presumably less liked stimulus (i.e., the extract in Experiment 1 and the disliked taste in Experiment 2) was the potentiated cue.