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.     

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.     

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.     

Generalized conditioned flavor aversions: Effects of toxicosis training with one flavor on the preference for different novel flavors

In two experiments, experimental rats were trained to have strong aversions to one of 10 novel-flavored solutions through contingent lithium injections; control rats were injected with lithium in the absence of prior consumption. Each animal was then tested with each of the 10 solutions, and the preferences of the experimental rats and the control rats were compared. A generalization gradient of the aversion to each conditioned aversive flavor was obtained for each test flavor.     

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.     

Cross-modality contrast: Exteroceptive context habituation enhances taste neophobia and conditioned taste aversions

The relationship between absolute and relative stimulus novelty was examined within the context of the conditioned taste aversion paradigm in which the relative novelty of the conditioned interoceptive stimulus was manipulated by differential exteroceptive context habituation. Rats received similar isolation histories but either 5 or 30 days of habituation to the test environment prior to treatment. One group was administered lithium chloride following saccharin consumption, a second group was administered isotonic saline following saccharin consumption, and a third group was administered saline after water consumption. The animals habituated for 30 days exhibited greater conditioned avoidance and greater neophobic avoidance of saccharin than did animals habituated for only 5 days. The results are interpreted in terms of a cross-modality stimulus contrast effect which implicates context habituation as an important parameter of both taste neophobia and taste aversion learning.     

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.     

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.     

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.     

Changes in the retrievability of associations to elements of the compound CS determine the expression of overshadowing

In two experiments, rats were presented with a taste conditioned stimulus (CS) alone, an odor CS alone, or an odor-taste compound followed by lithium chloride injection. When tested 1 day following conditioning, there was evidence that the odor cue overshadowed conditioning to the taste; however, there was no indication of overshadowing following a longer (21-day) retention interval, despite undiminished strength of the aversion in animals conditioned with only the single element (taste). The overshadowing observed at the 1-day retention interval was not reciprocal. Rats conditioned with the odor CS alone or with the compound CS expressed odor aversions of comparable strength—that is, no overshadowing. However, in contrast to the taste aversion, overshadowing of conditioning to the odor by taste was evident following a 21-day retention interval. Rather than reflecting a failure of the overshadowed stimulus to acquire associative strength, these data suggest that overshadowing may be expressed, or not expressed, as a result of changes in the relative retrievability of learned associations over time.     

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.     

Transfer of a taste aversion from food to water under various states of deprivation

The importance of ingestive contexts (feeding and drinking) and deprivation states to rats’ transfer of a taste aversion were examined, In Experiments 1 and 2, rats were trained with novel saccharin-treated foods while either food deprived or food and water deprived, They were then tested with a 1,0% saccharin solution while either water deprived or food and water deprived, Comparable aversions to the solution were displayed regardless of deprivation states, Two further experiments examined transfer to a .1% saccharin solution in conjunction with deprivation state change, When both stimulus properties and deprivation were widely discrepant from training to test, reduced transfer was noted. The results suggest that stimulus similarity was a stronger controlling variable than deprivation state similarity in facilitating the transfer of an aversion from a feeding context to a drinking context, The results were viewed as being consistent with the known parameters affecting generalization gradients.     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

Lack of reinstatement of an extinguished taste aversion

Three experiments with rat subjects were designed to investigate the possibility that an extinguished saccharin aversion might be reinstated if the animals are made ill with lithium chloride (in the absence of saccharin) following extinction. Although reinstatement can be obtained when the unconditioned stimulus is presented following the extinction of other kinds of conditioned behaviors, the present experiments provided no evidence that an extinguished taste aversion can be reinstated. No reinstatement was observed, even when the aversion had been only partially extinguished and when multiple injections of lithium chloride were administered in an attempt to reinstate the aversion.