Aversion conditioning and enhanced neophobia: role of test stimuli

In Experiment 1, 100 rats (Rattus norvegicus) received 10% sucrose or 5% casein hydrolysate followed, after 10 min, by a LiCl or saline injection or, after 12 h, by a LiCl injection. Subsequently, rats received aversion testing to the CS or neophobia testing to the opposite novel flavor. Aversion effects were reliably greater to casein than to sucrose. However, conditioning with sucrose yielded a reliably greater increase in neophobia to casein (relative to controls) than conditioning with casein yielded to sucrose. In Experiment 2, 60 rats received distilled water followed, after 10 min, by LiCl or saline injection or, after 12 h, by LiCl injection. Aversion effects occurred to distilled water. Neophobia testing to casein and sucrose showed that, relative to controls, neophobia increased reliably more to casein. The results of Experiments 1 and 2 were not attributable to differences in baseline intakes between casein and sucrose flavors. Together, these experiments indicated that the demonstration of conditioning-enhanced neophobia may depend more on the characteristics of the neophobia test flavor than on the strength of aversion established because of CS characteristics.     

Potentiation of odor by taste in rats: tests of some nonassociative factors

The contribution of nonassociative neophobia and sensitization to the potentiation of odor by taste in rats was tested in three experiments. In Experiment 1, neophobia for almond odor (O), saccharin taste (T), and odor-taste compound (OT) cues was tested before and after noncontingent lithium chloride poisoning and compared with conditioned aversions produced by OT-LiCl temporal pairing. The OT compound potentiated unconditioned neophobia, but there was no evidence of poison-enhanced neophobia, disinhibition of neophobia, or sensitization by noncontingent LiCl; temporal pairing produced aversions for the compound and its elements. In Experiment 2, generalization to a novel odor was tested after O-LiCl or compound OT-LiCl pairing. The potentiated odor aversion did not generalize to the novel odor; it was specific to the odor paired with taste and LiCl. In Experiment 3, potentiation of the odor component by a discriminant or nondiscriminant taste component was tested. Potentiation was evident only when a novel discriminant taste was in compound with odor prior to LiCl poisoning. These studies support an associative "indexing" hypothesis of the potentiation effect in rats.     

Stimulus generalization of conditioned taste aversion in rats

Relatively little information is available regarding the intradimensional stimulus generalization of conditioned taste aversion (CTA). Experiment 1 employed a between-groups generalization test to examine the extent to which conditioned flavor aversion to one sucrose solution generalized to other concentrations of sucrose in adult rats. Evidence of a gradient of aversion was obtained. Because generalization gradients in other tasks have been found to flatten over a retention interval, Experiment 2 investigated the effects of delayed testing (2, 7, or 21 days) upon the slope of the generalization gradient. The generalization gradient flattened at the intervals, suggesting that subjects forgot the specific attributes of the conditioning concentration and avoided generalized stimuli as if they were the original CS. Experiment 3 used a long delay between taste and toxicosis to degrade the associative contingency and found no evidence that the generalization gradients found in the first two experiments could be explained in terms of enhanced neophobia due to poisoning. These findings provide further evidence (cf. A. W. Logue, 1979, Psychological Bulletin, 86, 276-296; M. Domjan, 1980, in J. S. Rosenblatt, R. A. Hinde, C. Beer, & M. Busnel (Eds.), Advances in the study of behavior, Vol. 11, New York: Academic Press) that CTA shares a number of similarities with other learning processes. Further, they illustrate that stimulus forgetting can be detected in a paradigm considered relatively immune to retention loss.     

Effects of flavor salience on aversion performance following relatively long-delay backward conditioning procedures

Male albino rats (n = 144) received a 0.15 M injection of lithium chloride (at 2.0% body wt), followed 10, 30, or 75 min later by a 5.0% casein CS or a 10.0% sucrose CS, casein being the more salient CS. For each CS one-third of the rats received no fluid during the toxin-CS interval. The remaining two-thirds of the rats received 2-min access to distilled water or to a novel flavor 5 min after onset of the toxin-CS interval. For sucrose CS groups, the novel flavor was casein; for casein CS groups, the novel flavor was sucrose. Groups which received no fluid during the toxin-CS interval showed reliably greater aversion effects to the casein CS than to the sucrose CS. Results of Test Trial 1 showed that aversion to casein was relatively unchanged across toxin-CS intervals, while aversion to sucrose decreased reliably from the 10-min to the 30- and 75-min intervals. However, for each toxin-CS interval, aversion to the sucrose CS was reliably enhanced when casein access occurred in the interval, relative to that for distilled water or no fluid access. For the casein CS, access to sucrose or distilled water in the toxin-CS interval altered aversion effects, relative to the no fluid condition, depending on the interval.     

Salience and the effects of CS preexposure on aversion conditioning

Rats (n = 84) received preexposure to distilled water or to one of two differently salient flavors, 5.0% casein or 10.0% sucrose, casein being the more salient. Each preexposure group then received aversion conditioning to a 5.0% casein or a 10.0% sucrose CS. Aversion effects were reliably more enduring to casein than to sucrose. Relative to water-preexposed groups, preexposure to casein attenuated aversion effects to the casein CS reliably less than preexposure to sucrose attenuated aversion effects to the sucrose CS. During preexposure, neophobia was reliably greater to casein than to sucrose, suggesting that the demonstration of salience in taste aversion learning may be based on the inherent aversive properties of novelty.     

ACTH and ACTH4-10 modification of neophobia and taste aversion responses in the rat.

A series of experiments assessed the effects of ACTH and the ACTH analogue ACTH4-10 on drinking in conditioned taste aversion and neophobic situations. Both substances delayed the extinction of a conditioned taste aversion established by a single pairing of lithium chloride with milk (Experiment 1). However, in this situation, the ACTH parent peptide was more potent behaviorally. Administration of ACTH suppressed milk consumption in animals with no toxicosis experience (Experiment 2). This effect was apparently not due to the conditioning of a taste aversion (Experiment 3) with ACTH serving as a weak aversive unconditioned stimulus. Administration of exogenous ACTH (Experiment 4) or ACTH4-10 (Experiment 5) did not enhance neophobia; however, repeated injections of ACTH suppressed drinking. This ACTH suppression was related to the familiarity/novelty of the subtance being consumed. The neophobic response to milk eas no accompanied by pituitary-adrenal activation (Experiment 6). Both neophobic and conditioned taste aversion situation appear to be useful for assessing peptide effects on consummatory behavior.     

Effects of preexposure flavor concentration on conditioned aversion and neophobia

In Experiment 1, 128 experimentally naive, water-deprived rats (Rattus norvegicus) received pretraining access to either 0.25 or 1.5% saccharin, distilled water, or 2.0% saline, followed either by a pairing of 0.25 or 1.5% saccharin with an intraperitoneal injection of 0.15 M lithium chloride (LiCl) or by a pairing of distilled water with LiCl. Preexposure to either saccharin concentration reliably reduced conditioned aversion effects to 0.25% saccharin, relative to that for preexposure to distilled water or saline. But only preexposure to 1.5% saccharin reduced aversion effects to that concentration. In Experiment 2, 48 naive, water-deprived rats received preexposure procedures as in Experiment 1. Afterwards, the rats were tested for neophobia to 0.25 or 1.5% saccharin. Neophobia was reliably greater to the 1.5% concentration. However, preexposure to either saccharin concentration obliterated evidence for neophobia to saccharin, relative to that following preexposure to distilled water or saline.     

Developmental flavor experience affects utilization of odor, not taste in toxiphobic conditioning

Feeding experiences were varied in developing rats and the effects upon flavor neophobia and lithium chloride-induced flavor aversions were observed. In Experiment 1, nursing experience of neonate rats was reduced by artificial feeding via intragastric cannula; the rats then were tested with apple juice paired with lithium chloride injection at weaning or maturity. Conditioned aversions were not affected, but neophobia to novel apple juice was attenuated in artificially-reared rats tested at maturity. In Experiment 2, rats received enriched feeding experience after weaning, which consisted of (a) obtaining many complex flavors, a few of which were paired with poisoning, effortlessly in the home cage, or (b) foraging for various foods on an elevated maze. No dramatic effects on neophobia or conditioned taste aversion for saccharin water were apparent. In Experiment 3, rats were given experience after weaning with vanilla-scented water either paired or unpaired with quinine water, and then tested with the odor of almond or that odor compounded with saccharin water for neophobia and lithium-induced aversions. Flavor-experienced rats exhibited more pronounced odor conditioning and more resistance to extinction of the odor aversion after both simple and compound conditioning. In contrast, saccharin taste aversions were relatively unchanged. Apparently, enriched feeding and drinking experience facilitates the utilization of odor more than taste cues.     

Extinction of conditioned taste aversions: Effects of concentration and overshadowing

Extinction of conditioned taste aversions was examined as a function of taste concentration and of the presence of an additional taste. The results of Experiment 1 were consistent with previous evidence in that a conditioned aversion to high concentration saline was more persistent in extinction than an aversion to a low concentration. However, when floor effects were avoided the rate of extinction was faster for the higher (1%) concentration than for 0.2% saline (Experiment 2), a result consistent with accounts of extinction in other preparations. Three further experiments examined extinction of a conditioned sucrose aversion. The addition of 1% saline, but not of 0.2% saline, to sucrose during extinction produced overshadowing ("protection from extinction"; Experiment 3). Such overshadowing by saline was detected after two, but not after a single extinction trial (Experiment 4). This last finding suggests that under the conditions of the present experiments sweet and salty tastes function as elemental stimuli competing for loss of associative strength. No overshadowing was found when almond (an aqueous odour) was used in place of saline as the added stimulus, even when high concentrations of almond were used that produced observable neophobia (Experiments 5A and 5B).     

Trace decay and the priming of short-term memory in long-delay taste aversion learning: Disruptive effects of novel exteroceptive stimulation

A variation of Kalat and Rozin's two-presentation paradigm was used to test the hypothesis that the first, as opposed to the second, presentation of a flavor conditioned stimulus (CS) constitutes the functional CS in two-presentation experiments involving moderate interflavor intervals (IFIs), and results in flavor aversions that are a function of the primary, as opposed to the secondary, conditioned stimulus-unconditioned stimulus (CS-US) interval. Contrary to the hypothesis, it was shown in Experiment 1 that holding the primary CS-US interval constant at 4 hr for each of three groups, while decreasing the secondary CS-US interval (i.e., the interval between the second flavor presentation and the illness) from 3.75 hr to 2.5 hr to .5 hr, resulted in the flavor aversion increasing as the secondary CS-US interval decreased. However, the aversion acquired by the group with a 0.5 hr secondary CS-US interval was also found to be significantly weaker than that acquired by a single-presentation 0.5 hr control group. In Experiment 2 it was demonstrated that animals exposed to novel exteroceptive stimulation (NES) immediately prior to a second flavor presentation that preceded the US by 0.5 hr acquired an aversion as strong as that acquired by a 0.5-hr control group. In Experiment 3 it was demonstrated that, in the absence of a second flavor presentation, animals exposed to novel exteroceptive stimulation 0.5 hr prior to the US acquired a weaker flavor aversion than did animals not exposed to novel exteroceptive stimulation during the 4-hr flavor CS-illness US interval. The contrasting effects of novel exteroceptive stimulation observed in Experiments 2 and 3 were replicated in Experiment 4. The results suggest, consistent with the trace-decay hypothesis and Wagner's (1976) general model of stimulus processing, that exposure to novel exteroceptive stimulation disrupts continued processing of the short-term memory (STM) trace of the initial presentation of a flavor CS, and hence minimizes stimulus preexposure effects attributable to the priming of STM.     

Effects of prior exposure on conditioned taste aversion in the rat: androgen- and estrogen-dependent events

The effects of preexposure and gonadal hormone manipulation on the extinction of a conditioned taste aversion were investigated. In Experiment 1, male rats were given one prior exposure to sucrose at some selected time (Days 4, 2, or 1) before a second exposure (Day 0) to sucrose and an LiCl injection. Controls received no prior exposure to sucrose but experienced only the sucrose + LiCl condition (Day 0). under the single exposure condition (Day 0) castrated animals extinguished the aversion faster than either testosterone-treated castrated rats or sham-operated rats. Sham-operated and castrated rats that had received preexposure treatment (Days 4, 2, or 1) were not distinguishable on a within-groups comparison of behavior but differed from their respective controls by exhibiting faster rates of extinction. However, in the testosterone-treated group only the Day 4 preexposure group exhibited a faster extinction rate. In Experiment 2, estradiol, dihydrotestosterone, and testosterone were studied by using only a Day 1 preexposure condition. The testosterone-treated group maintained the aversion for the longest period of time, followed by dihydrotestosterone-treated, sham, castrated, and estradiol-treated groups. It appears that estradiol augments the castration effect whereas dihydrotestosterone attenuates the effect. In Experiment 3, estradiol was administered alone or in combination with two different doses of dihydrotestosterone, and a Day 1 preexposure condition was used. The findings indicate that the outcome of behavior is dependent on the ratio of estradiol to dihydrotestosterone, with variations in this ratio resulting in fast (estrogen effect) to slow (androgen effect) rates of extinction.     

Taste aversions conditioned by the aversiveness of insulin and formalin: role of CS specificity

Experimenters in the past have reported that when insulin is used as the unconditioned stimulus (US), rats will learn an aversion to a sodium chloride but not a sucrose solution, whereas with formalin as the US, they will learn an aversion to a sucrose but not a saline solution. The present experiments failed to confirm these findings. Aversions to sucrose were conditioned with insulin and aversions to sodium chloride were conditioned with formalin. The use of a more concentrated sucrose solution in the present study may have been responsible for the successful sucrose-aversion conditioning with insulin. Although the source of the discrepancy in findings concerning aversion conditioning with formalin remains unclear, experiments ruled out numerous possibilities. These experiments also showed that sodium chloride aversion conditioning with formalin is a highly robust phenomenon that occurs with a variety of conditioned stimulus durations and formalin doses, with distributed and massed training, in male and female rats, and even if saline is not the only novel solution presented during conditioning. Furthermore, the aversion can be detected with both single-stimulus and choice test procedures.     

Acquisition and extended retention of a conditioned taste aversion in preweanling rats

The time course of memory decay for infant rats may shed light on the processes responsible for infantile amnesia. A taste aversion conditioning procedure appropriate for both neonatal and adult rats was employed in four experiments to investigate the ontogeny of extended retention. In Experiment 1, rats trained at 1, 10, 20, or 60 days of age were tested for retention of the taste aversion 25 days later. At testing, only those rats conditioned when 20 or 60 days old demonstrated significant taste aversions. Experiments 2 and 3 established that rats 14-15 days old and older were able to retain significant taste aversions following a 25-day retention interval. Younger rats did, however, acquire and retain the aversion for several days and showed a gradual retention loss over progressively longer retention intervals (Experiment 4). These findings suggest that preweanling rats demonstrate initial consolidation, storage, and retrieval of conditioned taste aversions. It is only after this initial period that retention deficits become evident.     

Short- and long-term decrements in toxicosis-induced odor-aversion learning: the role of duration of exposure to an odor

Six experiments employed an odor-aversion paradigm to investigate the role of the duration of exposure to an odor in determining that odor's subsequent associability with illness. Rats were exposed to an odor at times T1 and T2, and the second of these exposures was followed by toxicosis. When the initial odor exposure was brief, the odor aversion was attenuated with a moderate T1-T2 interval of 3 hr (Experiment 1) but not with long intervals of 28 hr and 76 hr (Experiment 2). In contrast, when the initial odor exposure was long, the odor aversion was attenuated at a long T1-T2 interval (Experiment 3). With a T1-T2 interval of 24 hr, a brief initial exposure did not attenuate odor aversions when the context either remained the same or was changed from T1 to T2, whereas a long initial exposure attenuated such aversions when the context remained the same but not when the context was changed (Experiment 4). With a T1-T2 interval of 3 hr, a brief initial exposure attenuated odor aversions when the context remained the same or was changed from T1 to T2, whereas a long initial exposure attenuated such aversions when the context remained the same but not when the context was changed (Experiment 5). A brief exposure at T1, either with or without a subsequent context "extinction," attenuated odor aversions when the T1-T2 interval was 3 hr but not when this interval was 24 hr; a long initial exposure at T1, without a subsequent context "extinction," attenuated odor aversions when the T1-T2 interval was 4 hr and 24 hr but with a subsequent context "extinction" did not attenuate such aversions at either 4-hr or 24-hr T1-T2 intervals (Experiment 6). The results demonstrate that the duration of exposure to an odor determined whether that odor presentation caused short- or long-term decrements in odor conditionability and are discussed in terms of the relation between self- and retrieval-generated processes.     

Stimulus Preexposure, Comparison, and Changes in the Associability of Common Stimulus Features

In four experiments, rats received preexposure either to both of two compound flavours (AX and BX), or to just one (BX). Experiment 1 demonstrated a perceptual learning effect, showing that, for animals given preexposure to both flavours, an aversion conditioned to AX generalized only poorly to BX. Subsequent experiments assessed the properties of the common feature, X. Experiment 3 showed that the two preexposure treatments did not differ in the extent to which they produced habituation of the neophobia evoked by X. Experiment 2 showed that conditioning to X proceeded more rapidly in subjects given preexposure to both AX and BX than in subjects preexposed to BX alone. In Experiment 4, a similar effect was found when the elements of the compounds were presented serially. It is concluded that the perceptual learning effect of Experiment 1 occurs in spite of the fact that preexposure to two stimuli tends to maintain the associability of their common elements.     

The effects of preexposure-test and conditioning-test intervals on the magnitude of latent inhibition

Using the conditioned taste aversion paradigm, two experiments were conducted to examine the effects of the interval between preexposure and test and that between conditioning and test on the magnitude of latent inhibition. Experiment 1 revealed that the degree of latent inhibition was attenuated when rats were given a 21‐day interval between preexposure and test. It was also found that this attenuation was more marked in subjects which were given conditioning immediately after preexposure than those which were conditioned shortly before the test. Retention interval between preexposure and test was reduced to 12 days in Experiment 2, and exactly the same pattern of results as those found in Experiment 1 was obtained. These findings suggest that the memory of conditioning as well as that of preexposure decreases its retrievability after a long retention interval, although the former is more retainable than the latter.     

Relationship between vomiting and taste aversion learning in the ferret: studies with ionizing radiation, lithium chloride, and amphetamine.

The relationship between emesis and taste aversion learning was studied in ferrets (Mustela putorius furo) following exposure to ionizing radiation (50-200 cGy) or injection of lithium chloride (1.5-3.0 mEq/kg, ip). When 10% sucrose or 0.1% saccharin was used as the conditioned stimulus, neither unconditioned stimulus produced a taste aversion, even when vomiting was produced by the stimulus (Experiments 1 and 2). When a canned cat food was used as the conditioned stimulus, lithium chloride, but not ionizing radiation, produced a taste aversion (Experiment 3). Lithium chloride was effective in producing a conditioned taste aversion when administration of the toxin was delayed by up to 90 min following the ingestion of the canned cat food, indicating that the ferrets are capable of showing long-delay learning (Experiment 4). Experiment 5 examined the capacity of amphetamine, which is a qualitatively different stimulus than lithium chloride or ionizing radiation, to produce taste aversion learning in rats and cats as well as in ferrets. Injection of amphetamine (3 mg/kg, ip) produced a taste aversion in rats and cats but not in ferrets which required a higher dose (> 5 mg/kg). The results of these experiments are interpreted as indicating that, at least for the ferret, there is no necessary relationship between toxin-induced illness and the acquisition of a CTA and that gastrointestinal distress is not a sufficient condition for CTA learning.     

Food choice in rats (Rattus norvegicus): the effect of exposure to a poisoned conspecific

Three experiments were conducted to examine the effects of exposure to a poisoned conspecific on subsequent food aversion in rats. In Experiment 1A, rats that had been aversively conditioned to a cocoa-flavored food were exposed to a poisoned conspecific that had eaten the same food. On the subsequent choice test, the animals increased their aversion to that food. These results were reconfirmed in Experiment 1B, in which a cinnamon-flavored food was used as the stimulus. In Experiment 2, subjects were first exposed to a poisoned conspecific and then conditioned to the food which the conspecific had eaten. On the test, they showed no sign of increased aversion to that food.     

Effects of neophobia and habituation on the poison-induced avoidance of exteroceptive stimuli in the rat.

Two experiments on the role of neophobia in poison-induced aversions to exteroceptive stimuli are reported. In Experiment 1, rats were given either 10 or 25 days of habituation to the test situation prior to conditioning. Those animals with the longer habituation period avoided a complex of novel exteroceptive stimuli while those with the shorter habituation period did not. In Experiment 2 rats initially avoided the more novel of two containers, but gradually came to eat equal amounts from both. A single pairing of toxicosis with consumption from either the novel or the familiar container reinstated the avoidance of the novel container in both cases. The results were discussed in terms of an interaction between habituation and conditioning procedures. It was suggested that previously reported differences between interoceptive and exteroceptive conditioning effects may have been influenced by the differential novelty of the two classes of stimuli in the test situation. It was further suggested that non-contingently poisoned control groups should routinely be included in poison avoidance conditioning studies.     

Disruption of conditioned taste aversion: evidence that ECS weakens the gustatory engram

Two experiments are reported concerning the neural substrate of conditioned taste aversion (CTA) and the amnesic mechanisms involved when such learning is disrupted by electroconvulsive shock (ECS). Subjects were adult male rats. In the first experiment it is demonstrated that an aversion established to a 2.5% sucrose solution can simulate the learning loss reported in earlier studies which was caused by interpolating ECS between tasting and illness when the aversion was established using a 10% sucrose cue. It is concluded that if ECS acts to disrupt the memory of the taste cue, it possibly reduces it to only about one-quarter of its original strength, a substantial deficit not readily apparent simply from the degree of aversion displayed. In the second experiment, CTAs were established using either a 2.5% sucrose cue or a 10% cue with ECS interpolated during the taste-illness interval. Animals in the two groups were subsequently confronted with a range of sucrose stimuli and their respective aversions were compared. Near identical responses were observed under all conditions tested. These findings are consistent with the theory that ECS disrupts CTA by weakening the gustatory engram.