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.     

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.     

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.     

Effect of taste preexposure on taste and odor aversions

Thirsty Sprague-Dawley rats drank flavored water in a wind tunnel prior to lithium-induced toxicosis. Flavors were presented for 5 min; 30 min later a toxin, lithium chloride, was injected. After the rats had recovered, subsequent aversions to the taste and the odor were assessed separately. In Experiment 1, extensive preexposure to the taste component of the flavor attenuated neophobia to the flavor and the subsequent taste aversion. However, the subsequent odor aversion was unaffected. Experiment 2 partially replicated the results of Experiment 1 and showed that, in a situation in which only taste-potentiated odor aversions are usually found, nonpotentiated aversions were evident. Experiment 3 found that, in addition to attenuating taste aversions, taste preexposure enhances the capacity of rats to learn nonpotentiated odor aversions. The results are interpreted with a neural-based model of conditioned flavor aversions.     

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.     

The role of oral and postingestional cues in the conditioning of taste preferences based on differing caloric density and caloric outcome in weanling and mature rats

Experiments 1, 2, and 3 demonstrated that 23-day-old rat pups can acquire a conditioned taste preference for a flavor paired with either a more calorically dense diet or a diet that provides more calories than another diet of equal caloric density. Experiment 4 showed that 42–49-day-old rats also can acquire conditioned taste preferences based on differing caloric density and differing caloric outcome. Experiments 5 and 6 demonstrated that adult rats form conditioned taste preferences based only on caloric density. Although these experiments indicate that young rats are apparently capable of more diverse learning than adult rats, the question of what US underlies conditioned taste preferences based on diets of differing caloric density remains unresolved.     

Pentobarbital-induced place aversion learning

Pentobarbital is self-administered by rats but has also been reported to produce a conditioned place aversion. Since the self-administration and place preference paradigms both are considered to assess drug reward, we further examined the hedonic properties of pentobarbital, using place conditioning. In Experiment 1, a dose of 15 mg/kg (intraperitoneal) of pentobarbital produced a conditioned place aversion after 4 conditioning trials of various durations (5, 15, 30, or 60 min). Since rats are typically drug experienced in the self-administration paradigm, in Experiments 2 and 3, we examined the effect of drug history on pentobarbital-induced place conditioning. Although preexposure to pentobarbital attenuated the place aversion, it never resulted in a place preference. As has been reported with alcohol, pentobarbital is hedonically aversive in rats, when novel.     

Context specificity of conditioning in flavor-aversion learning: Extinction and blocking tests

In three experiments using rats as subjects, we investigated the degree to which a conditioned flavor aversion transfers from one context to another. Experiment 1, using a one-trial conditioning procedure, found no effect of a change of context on a conditioned aversion. Experiment 2 employed a multitrial procedure and demonstrated that a conditioned aversion was extinguished more rapidly after a change of context. Experiment 3 showed that context change decreased the effectiveness with which a conditioned flavor could block acquisition of an aversion by a second flavor. It is argued that these data cannot be explained in associative terms, and that they constitute evidence of conditionality in a simple aversive conditioning procedure.     

Location preference and flavor aversion reinforced by amphetamine in rats

To associate the identical drug state with both a location and a flavor, rats were given intraperitoneal amphetamine injections and then confined for 20 min in one side of a shuttlebox with access to a flavored solution; on control trials without injections, they were confined for 20 min in the opposite side with a different flavor. In the first experiment, the rats were placed in the shuttlebox immediately after injections; in the second experiment, they were placed in the shuttlebox 20 min after injections. Subsequent free-choice tests in both experiments revealed an increased choice of the side of the shuttlebox associated with amphetamine but also an aversion to the flavored solution associated with the drug.     

Competition between ethanol-induced reward and aversion in place conditioning

Previous place conditioning studies in mice have shown that injection of ethanol immediately before a conditioned stimulus (CS+) produces conditioned preference, whereas injection of ethanol immediately after CS+ produces conditioned aversion. In the present experiments, we examined the learning that occurs when ethanol is injected in “ambiguous“ procedures that provide the opportunity for both types of conditioning. When ethanol was given midway through the CS (Experiments 1 and 2) or both before and after the CS (Experiment 3), the direction of place conditioning was the same as when mice were exposed only to whichever contingency occurred first (a primacy effect). That is, injection of ethanol in the middle of the CS conditioned aversion, whereas injection both before and after the CS conditioned preference. Because these results support the idea that ethanol elicits both aversive and rewarding effects, they are most consistent with conditioning theories that conceptualize unconditioned stimuli (USs) as events that can activate multiple representational components.     

Conditioned flavor preferences based on hunger level during original flavor exposure

In five experiments, rats’ preference for a flavor was greater if the flavor had previously been consumed under low rather than high deprivation. This preference was conditioned in as few as three flavor-deprivation pairings (Experiment 1), and persisted through 28 test days, half under each deprivation level (Experiment 2). Rats never preferred the flavor associated with high deprivation even when calories were increased by giving 40 ml of 8% sucrose or when caloric density was increased to the equivalent of 20% sucrose. The preference for the low-deprivation flavor was greater when saccharin solutions were used rather than sucrose solutions, but the preference did emerge when sucrose solutions were used as testing proceeded and when a lower concentration of sucrose was used. We suggest that these preferences may be a result of flavor-taste associations rather than associations between flavors and postingestive consequences, and that the taste of the solutions under low deprivation is preferred to the taste under high deprivation.     

An exposure effect opposes flavor-nutrient learning

When a caloric substance is followed by a flavored substance, preference for the flavor that followed the calories can increase because of a flavor-nutrient association. We showed here that this effect is opposed by a fullness effect: consuming the caloric substance itselfreduces-consuniption of the flavor that follows. Because consumption of this flavor was less than consumption of a flavor that was given alone, there was a reduced preference for the flavor that followed the calories—an effect opposing flavor-nutrient learning. The preference for the flavor that was given alone and consumed in greater amounts seems to be due to exposure per se and is not a result of contrast. When the amount of the flavor given alone was directly controlled so that it was one half the amount of the flavor that followed the calories, preference for the flavor following the calories was no higher than when consumption of the two flavors was equal. Thus, exposure can interfere with flavor-nutrient learning, but does not enhance it.     

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.     

Interstimulus interval determines whether ethanol produces conditioned place preference or aversion in mice

DBA/2J mice were exposed to a distinctive floor stimulus (CS+) and ethanol (2 g/kg) in a place conditioning paradigm. A different floor stimulus (CS?) was presented with saline. Mice injected just before or 30 min before CS exposure (Groups 0, ?30) showed conditioned place preference, whereas mice injected right after exposure to the CS (Group 5) displayed place aversion (Experiment 1). None of the other groups (?120, ?60, 15, 60) showed place conditioning. Handling and saline injection given just before or after CS exposure were unable to produce place conditioning (Experiment 2). However, there was a positive relationship between ethanol concentration (10% vs. 20%) and test performance, suggesting that peritoneal irritation influences place conditioning (Experiment 3). Overall, these findings support the suggestion that intraperitoneal injection of ethanol produces an initial short-duration aversive effect that is followed by a longer lasting positive motivational effect.     

Expression of a taste aversion conditioned with an odor-taste compound: Overshadowing is relatively weak in weanlings and decreases over a retention interval in adults

Adult rats were injected with lithium chloride (LiCl) after consumption of a novel flavor (chocolate milk) that either was or was not presented together with a novel ambient odor (banana) as a compound conditioned stimulus (CS). In Experiment 1, the adults’ consumption of the flavor 24 h after conditioning was compared with that of weanling rats given the same conditioning treatment on Postnatal Day 21. The results confirmed previous indications that the reduction in aversion observed for adults conditioned with the compound CS (overshadowing) was weak or nonexistent in weanlings. After a longer retention interval (21 days), there was no evidence of overshadowing in adults despite maintained retention of the basic conditioned aversion. In Experiment 2 this decrease in overshadowing after a long retention interval was replicated with adult animals and extended to a different method of testing. The form of the effect was the same as in Experiment 1: The decrease in overshadowing occurred over the retention interval without loss in retention of the basic taste aversion; the decrease in overshadowing was a consequence of anincrease in the flavor aversion displayed by animals conditioned with the compound CS. The impaired flavor aversion (i.e., the overshadowing) observed shortly after conditioning apparently was due to factors associated with memory retrieval, rather than to reduced attentional or associative strength.     

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.     

Effects of proximal unconditioned stimulus preexposure on ingestional aversions learned as a result of taste presentation following drug treatment

Taste aversions were conditioned by exposing subjects to a 1.0% saccharin solution 30 min after an injection of lithium chloride. The aversion learning was disrupted if subjects had also received an additional lithium injection some time earlier (Experiments 1–3). This interference effect of US preexposure was a decreasing function of the preexposure interval, beyond the optimal interval (105 min) for observing the phenomenon (Experiment 1), and was directly related to the dose of the preexposure injection (Experiment 2). No interference with conditioning occurred at short (e.g., 30-min) preexposure intervals (Experiment 1), probably because under these circumstances the preexposure injection itself conditioned a strong aversion (Experiment 4). At moderate (105-min) but not at short (30-min) preexposure intervals, the interference with aversions learned as a result of taste exposure following drug injection was comparable to the interference with learning in a more conventional forward conditioning procedure (Experiments 3 and 4). These findings are similar to previously documented effects of proximal CS- and US-preexposure and are consistent with recent stimulus rehearsal and opponent-process theories.     

Extinction of the overshadowing CS after overshadowing in conditioned taste aversion

Using a conditioned taste aversion preparation overshadowing of flavor-illness association was produced through the presentation of a second flavor during the interval between the first flavor and illness. The modulatory effects of extinguishing the association between the second (over-shadowing) flavor and illness on conditioned responding to the target flavor was investigated. In Experiment 1, we found that, following one-trial overshadowing, extinction of the overshadowing flavor had no effect on conditioned responding to the target flavor. In Experiment 2, we found a similar absence of an effect of extinction of the overshadowing stimulus in a multitrial over-shadowing paradigm. Experiment 3 confirmed the results of Experiments 1 and 2 using conditioning parameters that were designed to weaken the association between the overshadowed flavor and illness. In Experiments 4 and 5, we used simultaneous presentation of the flavors during conditioning and obtained a weakened aversion to the overshadowed flavor when the overshadowing CS was extinguished. These findings are inconsistent with previous observations in conditioned fear preparations that suggest that extinction of the association between the overshadowing stimulus and the unconditioned stimulus attenuates overshadowing. Possible reasons for the discrepant results are discussed.     

Toxiphobia conditioning with exteroceptive cues

The types of conditioned properties acquired by novel (i.e., nonpreexposed) or familiar (i.e., preexposed) exteroceptive cues that were paired with toxicosis, in the absence of a flavor CS, were evaluated in four experiments. In Experiment 1, the conditioned properties of novel exteroceptive cues served to block the acquisition of an aversion to a flavor CS during flavor conditioning and to suppress the ingestion response during flavor testing; animals failed to suppress their ingestion of either the flavor CS or a neutral flavor when tested in the absence of the exteroceptive CS, but suppressed their ingestion of both the flavor CS and the neutral flavor when tested in the presence of the exteroceptive CS. In Experiment 2, preexposed exteroceptive cues that had been paired with toxicosis failed to provide evidence of such conditioned properties. Experiment 3 demonstrated that preexposed contextual cues that were reinforced in compound with novel exteroceptive cues failed to acquire the conditioned properties acquired by nonpreexposed contextual cues under the same conditions of reinforcement. Finally, in Experiment 4, the conditioned properties of the novel exteroceptive cues served to evoke a conditioned gastrointestinal response that gradually extinguished as a function of repeated nonreinforced exposures to the exteroceptive cues, and, in the absence of such extinction, the conditioned properties served to block the acquisition of a flavor aversion.     

Inhibitory associations between neutral stimuli in flavor-aversion conditioning

In Experiments 1 and 2, rats were exposed to two compound flavors, AX and BX, containing one flavor in common (X). Following this exposure phase, an aversion was conditioned to A in the experimental group by pairing its consumption with an injection of lithium, while a control group drank A without being poisoned. The effect of this treatment was to establish B as a conditioned inhibitor. In Experiment 1, experimental animals were slower than controls to condition an aversion to B when its consumption was paired with lithium (a retardation test of conditioned inhibition). In Experiment 2, B alleviated the suppression of intake of another flavor previously paired with lithium (a summation test). Experiments 3 and 4 established that these effects depended upon prolonged prior exposure to AX and BX.