Neophobias and conditioned taste aversions in rats following exposure to novel flavors

Rats were successively exposed to three solutions with distinctively different flavors and then tested for both neophobia and propensity to form conditioned taste aversions to a fourth distinctively flavored solution. All permutations between the four solutions (salty, bitter, sweet, and sour) were examined. The prior exposures resulted in attenuation of neophobia to novel salty and sour solutions, but not to equally novel bitter or sweet solutions. These effects were found to depend upon thediversity of the prior ingestive events rather than upon either a single specific flavor experience or a summation of the reductions in generalized neophobia accrued by each substance separately; both of the latter findings are inconsistent with stimulus generalization being responsible for the observed attenuation of neophobia to salty and sour solutions following exposure to diverse different solutions. A further test of generalization between the salty and bitter solutions, consisting of associating one flavor with poison and extinguishing the avoidance response in half the animals prior to testing for generalization of conditioned taste aversion to the other flavor, also proved negative. Although these effects of exposure to flavors distinctly different from the test solution may be dependent upon solution concentrations, further research found that the same pretest exposures and same test concentrations failed to inhibit formation of conditioned taste aversions. A demonstration of “latent inhibition” attested to the sensitivity of our procedure to potential interference with acquisition of conditioned taste aversions. The results are considered in light of the relationship between neophobia and conditioned tasted aversions, the differential biological relevancy of specific tastes, and abstraction as a cognitive capability of rats. The possibility is raised that the defense against toxins is not the primary function of neophobia.     

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.     

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.     

Effects of pretraining on conditioning-enhanced neophobia: Evidence for separable mechanisms of neophobia and aversion conditioning

In two experiments, rats (n = 228) received pretraining access to a distinctive novel flavor (saline) followed by aversion conditioning to a different novel conditioned stimulus (CS) (saccharin). Then the rats were tested for aversion to the CS (saccharin) or for conditioning-enhanced neophobia to a third novel flavor (casein hydrolysate). Pretraining access to a distinctive novel flavor that differed from the CS reliably reduced the magnitude of conditioning-enhanced neophobia to casein, but did not reliably affect conditioned aversion effects to the CS. Pretraining access to the CS reduced aversion effects to the CS and reduced postconditioning neophobia to casein to the performance level shown by ingestion-toxin controls. Results were consistent with the view (Braveman & Jarvis, 1978) that conditioned aversion and neophobia may be independent phenomena with separable underlying mechanisms.     

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.     

Stimulus generalization of an illness-induced aversion to different intensities of colored water in Japanese quail

After quail (Coturnix coturnix japonica) experienced a toxicosis following ingestion of water colored dark blue or light blue, the stimulus generalization of the aversion to light, medium, and dark blue water was investigated. Birds made ill following ingestion of dark blue water demonstrated a reduced aversion when tested with intensities of blue water weaker than the conditioning intensity. Birds made ill following ingestion of light blue water showed stronger aversions as a function of increasing test intensity, suggesting a stimulus intensity dynamism effect. The stimulus generalization and stimulus-intensity dynamism effects observed in this study where visual cues were used is in agreement with other results within the illness-induced aversions paradigm where rats were tested with varying concentrations of a gustatory stimulus.     

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.     

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.     

Conditioned taste aversion in Philippine rice rats (R. r. mindanensis): Comparisons among drugs,dosages, modes of administration,and sexes

The effects of select drugs, dosages, and modes of administration upon learned taste aversions were compared among groups of wild-caught male and female Philippine rice rats (R. r. mindanensis). Experiment 1 compared two-choice saccharin aversions for 28 days among groups intubated with copper sulfate, cyclophosphamide, lithium chloride, red squill, sodium chloride (control), or deionized water (control). Main results were that 375 mg/kg lithium chloride produced the greatest sustained aversions, whereas 198 mg/kg cyclophosphamide and 210 mg/kg red squill produced moderate aversions, with males showing more resistance to extinction than females. Experiment 2 compared saccharin aversion among matched groups of male and female rats that received low (36 mg/kg), moderate (105 mg/kg), or high (368 mg/kg) dosages of lithium by gavage, ip injection, or ingestion. Sex differences in rates of extinction were found for the ingestion and injection-dosed rats, but no sex difference was again found for rats dosed by gavage. A significant mode × day interaction indicated that extinction progressed more rapidly for rats dosed by gavage. For all modes of administration, high dosages yielded intense 28-day aversion, moderate dosages produced intermediate 3–5 day aversion, and low dosages caused no aversion.     

Taste potentiates color-sickness associations in pigeons and quail

The present experiments assessed cue utilization in pigeons and quail on similar tests of poison-based aversion learning. In Experiments 1 and 2, three groups of pigeons were given colored water, flavored water, or colored flavored water prior to induction of sickness; these experiments differed only as to the specific colors and flavors used as stimuli. In both experiments, the birds trained with flavored water exhibited reliable taste aversions when tested with uncolored flavored water. Similar degrees of aversion were observed whether the flavored water had been colored or uncolored during training, suggesting that the color cue had little or no effect on the conditioning of the flavor cue. In contrast, the flavor cue had a pronounced effect on the conditioning of the color cue. When tested with unflavored colored water, the birds trained with colored flavored water exhibited significantly stronger color aversions than those trained with unflavored colored water. That is, the flavor cue enhanced or potentiated the conditioning of the color cue. In a third experiment, quail were trained in the same way as the pigeons with virtually the same result. The pattern of cue utilization observed in the present experiments with pigeons and quail differs markedly from that proposed by Wilcoxon, Dragoin, and Kral (1971) for quail. However, a reexamination of the results obtained by Wilcoxon et al. suggested that they are susceptible to an alternative interpretation consistent with the present results.     

Attenuation of the CS-preexposure effect after a retention interval in preweanling rats

The effects of flavor preexposure and retention interval were assessed in 6- and 12-day-old rats. Conditioned aversions to a flavor appeared at both ages. The conditioning of the younger pups was unaffected by conditioned stimulus (CS) preexposure and was not evident after a 10-day retention interval. For the 12-day-old rats, preexposure to either the flavor CS or a different flavor attenuated aversion strength when the rats were tested soon after conditioning. Other 12-day-old rats that were tested 10 days after conditioning also expressed substantial aversions, but with a retention interval of this length, the aversions were equivalent for animals preexposed to the CS and those not preexposed before conditioning. This loss of the CS-preexposure effect over a long interval, which has also been observed in adult rats, identifies the locus of this effect as postacquisition and perhaps at the stage of memory retrieval.     

Effects of context exposure during conditioning on conditioned taste aversions

Rats were used in a conditioned taste aversion procedure in order to examine the effects of context exposure duration during the conditioning sessions on conditioned responding. One flavor was paired with lithium chloride during a long session in one context, whereas another flavor was conditioned during a short session in another context. Testing occurred in the home cage. The results showed that conditioning during short sessions produced strong conditioned taste aversions. Conditioning during long sessions produced strong conditioned taste aversions when the conditioned-stimulus-unconditionedstimulus (CS-US) pairing occurred at the end of the lengthy session. Other results showed that context-US associations were formed during the short duration sessions and that these associations supported conditioned responding to the CS trained in that context. The results are discussed with respect to the different influences that contextual cues can exert on conditioned responding.     

Increase in retention of a taste aversion by weanling rats after a long interval

Weanling rats were tested for retention of an aversion to a novel flavor (chocolate milk) that had been conditioned as a single-element conditioned stimulus (CS) or in compound with a novel ambient odor (banana). The presence of the ambient odor during conditioning had no effect on flavor aversion shortly thereafter, confirming previous results. The flavor aversion observed 21 days after conditioning, however, was significantly stronger for pups conditioned with the single-element CS than for those given the flavor-odor compound as the CS. This retention effect was due to a surprisingincrease in the conditioned aversion observed 21 days after conditioning with the single-element CS. A second experiment confirmed this paradoxical increase in retention of the aversion to chocolate milk. This experiment also verified that no such increase occurred in retention of the conditioned aversion to a different flavor (saccharin), whether the initial aversion was strong or weak. The results may be explained in terms of generalized latent inhibition from consumption of mother’s milk.     

Variable processing of flavors in rat STM

Hooded Lister rats exhibited less neophobia towards (i.e., drank more of) a novel fluid (3% lemon or 5% sucrose) on a 10-min test if given a 6-min exposure to that fluid 6 h earlier. Presentation of a distractor (1.26% coffee) immediately after preexposure to the test solution enhanced neophobia habituation to lemon (Experiment 1), but disrupted habituation to sucrose (Experiment 3). This bidirectional distractor effect was not due to distractor-induced change in the hedonic value of the preexposed test flavor (Experiment 4). Evidence was obtained (Experiment 5) indicating that the rat perceives lemon to be more similar to coffee than is sucrose. It is suggested that when test flavor and distractor are dissimilar, processing of the distractor denies the preexposed test flavor sufficient processing in STM to allow encoding of information about that flavor in LTM. Consequently, the rat responds to a subsequent presentation of the test flavor as it would to a novel stimulus. When test flavor and distractor are similar, however, the distractor elicits less processing in STM (cf. Wagner, 1976) and is therefore less able to disrupt STM processing of the preexposed test flavor. The resultant loss of neophobia to the test flavor resulting from encoding of information about that flavor in LTM may then be augmented by generalization of attenuated neophobia to the distractor. Consistent with this analysis, coffee was shown to suffer more proactive interference when preceded by lemon than when preceded by sucrose (Experiment 6).     

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.     

Generalization of the CS- preexposure effect transfers to taste- aversion learning

Rats repeatedly exposed to a maple, vanilla, banana, or water solution were subsequently poisoned with lithium chloride after drinking the maple solution. Subjects preexposed to water showed the strongest aversions to maple, followed, in descending order, by rats preexposed to banana, vanilla, and maple. These results suggest that the effects of prepoisoning experience with a novel flavor generalized to the maple CS and interfered with the development of the maple aversion. The technique reported here is potentially applicable to study of gustatory discrimination processes.     

Poison-avoidance learning to food-related tactile stimuli: Avoidance of texture cues by rats

Rats injected with lithium chloride after ingesting familiar food pellets presented in textured metal sleeves learned aversions to the sleeved food. In a choice between sleeved and unsleeved food, the aversions were evident following conditioning with toxicosis delayed as long as 120 min after exposure to the sleeved food (Experiment 1). Texture-specific aversions resulted from procedures in which rats were exposed to food in both rough- and smooth-textured sleeves but were injected with lithium only in conjunction with one of the textures (Experiments 2–4). This differential aversion learning occurred when lithium treatment was delayed 30 min after exposure to the sleeved food (Experiments 3 and 4) and was equally evident in rats conditioned and tested in total darkness or in normal room-level illumination (Experiment 4). However, differential texture aversion learning was not observed with 90- or 300-min delayed toxicosis (Experiment 3). The present experiments highlight the importance of tactile cues in the poison-avoidance learning of species that handle their food during the course of ingestion.     

CS–US interval determines the transition from overshadowing to potentiation with flavor compounds

The present series of five flavor aversion experiments with rat subjects examined compound conditioning at varying CS-US intervals. Using a taste-taste design, Experiments 1A and 1B demonstrated overshadowing at a 0-min CS-US interval and potentiation at a 120-min CS-US interval, and these effects occurred with both tastes of the compound. Experiment 2 showed that the aversion to a single element is reduced when the CS-US interval is increased to 120 min, but the aversion for a compound taste is not. Experiments 3A and 3B explored odor + taste compound conditioning; the results demonstrated odor potentiation across the trace interval and a transition from taste overshadowing to taste potentiation. Collectively, the data show that the change from overshadowing to potentiation was not due to changes in the aversions produced by compound conditioning but, instead, was due to a more rapid loss of conditionability across a trace interval prior to the US in single-element conditioning. These experiments suggest that following compound conditioning, the aversion to each element represents generalization decrement from the configured compound, but the designation of overshadowing or potentiation actually depends on the status of conditioning in the single-element control.     

Socially induced diet preference can partially reverse a LiCl-induced diet aversion

I examined the capacity of a socially induced enhanced diet preference to reverse the effects of a LiCl-induced diet aversion. I found that rats poisoned after eating a novel diet (Diet NPT) would consume substantial amounts of Diet NPT following interaction with a conspecific that had eaten Diet NPT. Neither rats interacting with a conspecific fed some other diet nor rats exposed to Diet NPT itself exhibited reduced aversion to Diet NPT. This surprising capacity of social interaction to ameliorate even profound toxicosis-induced aversions suggests that social influence may be a major experiential determinant of the diet preferences of free-living rats.     

Variations in the retention of taste aversions: Evidence for retrieval competition

In six experiments, we examined taste and compound taste/taste aversions at different retention intervals. In Experiment 1, saccharin aversions were significantly weaker 1 day after conditioning than 21 days after conditioning. This effect was determined not to be caused by the aftereffects of illness or differential hydration. With the use of a saccharin/denatonium compound, Experiment 2 demonstrated overshadowing of a denatonium aversion at 21- and 1-day retention intervals, Experiment 4 showed a potentiated saccharin aversion only at the 21-day retention interval, and both Experiments 2 and 4 revealed that the aversion of the taste-only controls was stronger at the later retention interval. Experiments 3 and 5 demonstrated that the differences at the two retention intervals were not caused by unconditioned changes in taste preference. Finally, Experiment 6 showed that extinction of the conditioning environment prior to testing results in stronger saccharin aversions than occur in nonextinguished controls. Collectively, these experiments suggest that testing within a 24-h period after conditioning will result in significantly weaker taste aversions. Also, these results support a retrieval-competition explanation that may account for the weakened aversions at the 1-day testing interval of both groups conditioned to single elements and those conditioned to compounds.