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).     

Effects of distractor familiarity on habituation of neophobia

When the first presentation of a neophobic flavor is immediately followed by a distractor flavor, habituation of the neophobic response is typically attenuated. This is manifested by the fact that neophobia is still shown to the target flavor on its second presentation. Three experiments investigated the prediction that this effect will occur only for novel, but not familiar, distractor solutions. Experiments 1 and 2 found that, contrary to this prediction, both novel and familiar distractors can attenuate the habituation of a neophobic response. In Experiment 3, however, when the distractor was made very much more familiar, it lost its ability to interfere with the habituation of neophobia to the target solution. These results are discussed in terms of Wagner’s (1981) theory of habituation.     

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.     

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.     

Learning and nonlearned neophobia enhancement both contribute to the formation of illness-induced taste aversions by deer mice (Peromyscus maniculatus bairdi)

An investigation was made of the occurrence of learned and nonlearned aversions in the acquisition of illness-induced taste aversions in mice of the genusPeromyscus. It was determined: (1) that illness following the ingestion of a novel flavor both produced aversions specific to that flavor and also enhanced neophobia directed toward novel flavors in general; (2) that the specific aversion and the enhanced neophobia appeared to be mediated by independent processes, with no indication that the enhanced neophobia was dependent upon the integrity of the specific aversion; and (3) that illness following the ingestion of familiar water produced enhanced neophobia, which did not appear to be mediated by an aversion to water. It was noted that the results were fundamentally in agreement with those previously obtained with laboratory rats, except that a demonstration of the independence between the two types of aversions has not yet been reported in those animals.     

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.     

Extinction and retraining of simultaneous and successive flavor conditioning

In three experiments, rats received pairings of flavor conditioned stimuli with polycose unconditioned stimuli, in either a simultaneous or a sequential relation. Both temporal relations produced excellent conditioned increases in consumption of the flavors. Separate presentation of the flavors resulted in extinction in both cases. However, restoring the pairing of the flavor with polycose resulted in reconditioning only with the sequential, not with the simultaneous, relation.     

Taste preferences as a function of food deprivation during original taste exposure

It has been reported previously that rats prefer a flavor they consumed under high deprivation to a flavor they consumed under low deprivation (Revusky, 1967). Here it was found that this preference occurs only if nutritive solutions are used and the flavors are given preceding and following eating. If flavors are given separately from the daily feeding, rats prefer the flavor given under low deprivation, whether or not a nutritive solution is used (Experiment 3). If flavors are given before and after the daily feeding, rats prefer the flavor they had under high deprivation (before feeding) more if sucrose solutions are used than if saccharin solutions are used and more on a high-deprivation test than on a low-deprivation test (Experiments 1 and 2). It was concluded that the “incentive value” of consumption is not necessarily higher under high deprivation than under low deprivation. The preference for the low-deprivation flavor obtained here may reflect a greater proportional rewarding effect of consumption under low deprivation or may reflect an aversion to the flavor consumed under high deprivation. Perhaps a small taste of flavor under high deprivation initiates responses of digestion that are unsatisfied and thus aversive, and the more so the higher the deprivation level.     

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.     

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.     

Interval between preexposure and test determines the magnitude of latent inhibition: Implications for an interference account

The effect of a retention interval on latent inhibition was studied in three experiments by using rats and the conditioned taste-aversion procedure. In Experiment 1, we demonstrated an apparent loss of latent inhibition (i.e., a strengthening of the aversion) in preexposed subjects that experienced a retention interval of 12 days between conditioning and the test. In Experiment 2, we found no effect of this retention interval on the habituation of neophobia produced by the phase of exposure to the flavor. In Experiment 3, we showed that interposing a retention interval between preexposure and conditioning produced effects exactly comparable to those seen in Experiment 1. The implications of these results for rival theories of latent inhibition, as an acquisition deficit or as a case of interference at retrieval, are discussed.     

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.     

State-dependent retention induced by postacquisition exposure to pentobarbital or shock stress in rats

Although state-dependent retention (SDR) has been the focus of considerable research in recent years, there has been little work demonstrating this phenomenon when the treatment is administeredafter learning. In the present study, the effectiveness of two different treatments, pentobarbital and shock-induced stress, in producing postacquisition SDR in rats was examined. In all experiments, water-deprived rats were exposed to a novel flavor, apple juice, on each of 2 days. Subjects that remember their initial exposure to this substance should show an increase in consumption on their second exposure (i.e., an attenuation of neophobia). It was found that subjects exposed to pentobarbital (Experiment 1) or shock-induced stress (Experiment 2) immediately after their initial exposure did not increase their intake on the subsequent exposure unless they were reexposed to pentobarbital (Experiment 1) or shock-induced stress (Experiment 2b) shortly prior to that exposure. These results clearly show that this “attenuation of neophobia” paradigm can be used to investigate postacquisition SDR.     

Conditioned flavor preferences as a function of deprivation level: Preferences or aversions?

Rats tend to prefer flavors previously consumed under low deprivation to flavors previously consumed under high deprivation (Capaldi & Myers, 1982). We attempted to distinguish among possible associative explanations by determining whether this conditioning phenomenon was based upon conditioned preferences, conditioned aversions, or both. We compared preference for flavors presented exclusively under either high or low deprivation with preference for a neutral flavor. In Experiments 1A and 1B the neutral flavor was one that had been randomly paired with both high and low deprivation, whereas in Experiments 2 and 3 the neutral flavors had not been associated with either high or low deprivation. Our results strongly suggest that this conditioning phenomenon is based upon an actual increase in preference for the flavor consumed under low deprivation rather than on any form of aversion conditioning.     

The role of amount consumed in flavor preexposure effects and neophobia

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

Conditioned fear and neophobia following inescapable shock

Lick-suppression tests were used in seven experiments to assess the transsituational transfer of fear in the learned helplessness paradigm. Two sources of fear combined to suppress test drinking in inescapably shocked rats. A situational odor was strongly associated with shock pretreatments and mediated the transfer of conditioned fear during testing. Fear of the pretreatment odor was greater following inescapable shock than after escapable shock or restraint. This conditioned suppression was retained for at least 72 h after pretreatment. Neophobia was enhanced as a second, nonassociative reaction to inescapable shock. Unconditioned fear was augmented by a novel odor in the test context, but otherwise was weak and dissipated within 72 h. However, neophobia was necessary for differential conditioned suppression in inescapably shocked rats. The pretreatment odor elicited fear only when tested in a novel context. Initial habituation to the test apparatus reduced conditioned fear. These data provide additional evidence for odor-mediated transfer of helplessness. Conditioned fear and neophobia are discussed in relation to recent anxiety interpretations of the phenomenon.     

Rats’ memory for serially presented flavors: Effects of interstimulus interval and generalization decrement

In four experiments, the effect of sequential exposure to a series of five novel flavors on the subsequent neophobic response of water-deprived rats to those flavors when they were presented simultaneously was examined. After a list-test interval of 30 min and a list-interstimulus interval of 10 sec, the rats generally consumed more of the first and last flavors presented in the initial sequence. This finding was taken to reflect the existence of primacy and recency effects. Experiment 1 provided evidence that successive contamination can occur between flavors in the initial list, making subsequent recognition of later flavors in the list more difficult. However, this effect was overcome by presentation of water between each flavor during the list exposure. Experiments 2 and 4 showed that primacy was not a necessary result of successive contamination in this procedure, by demonstrating that increasing the interstimulus interval between list items decreased the size of the primacy effect. This result suggests that rats’ memory for serially presented items may be controlled by mechanisms different from those typically implicated in the human verbal memory literature. In Experiment 3, the question of whether the testing procedure adopted here could have introduced sources of artifactually produced serialposition effects was explored, but no such influence was found.     

Effect of taste context and ambient context changes on successive negative contrast

Rats shifted from 32% sucrose to 4% sucrose lick less than rats that experience oniy the 4% solution. Previous experiments have found this negative contrast effect to be reduced (“disinhibited”) by the addition of a novel tone in the postshift period. In Experiment 1 of this paper, the negative contrast effect was enhanced when a novel flavor was added to the sucrose solution in the postshift period. In Experiments 2–4, changes in the ambient context, even changes sufficient to produce disruptions in licking, did not alter the degree of negative contrast. Tiese results suggest that (1) rats compare rewards across substantially different contexts, (2) contrast may serve to enhance taste neophobia, and (3) a disinhibitory effect may be confined to the presentation of punctate, nontaste, novel stimuli within a familiar context.     

Serial habituation in two-, three-, and four-month-old infants.

Serial habituation of visual fixations in 2-, 3-, and 4-month-old infants was investigated through a design permitting cross-sectional, within-subject longitudinal, cohort longitudinal, and time-lag analyses. The components of the standard stimulus were displayed individually before and after successive exposures of the standard for familiarization. Relative saliencies of the components for each child were indexed by magnitude of responding to these parts in the initial presentation. Comparisons of the pre- and postexposure fixation times suggest that for all ages habituation was under way to the parts of the stimulus in orders of the relative saliencies. The nature of the data in general did not appear to be influenced significantly by any 1 methodology.     

Microstructural analysis of conditioned and unconditioned responses to maltodextrin

The microstructure of licking responses was analyzed to investigate the interaction between unconditioned responses to maltodextrin and the responses to flavor cues previously associated with maltodextrin. Experiment 1 demonstrated that although the consumption of maltodextrin peaked at intermediate concentrations, the mean lick cluster size showed a positive, monotonic increase with concentration. In Experiment 2, a (conditioned stimulus) CS+ flavor was paired with 16% maltodextrin, whereas a CS- flavor was paired with 2% maltodextrin. During test, consumption of the CS+ was higher than that of the CS- when the flavors were combined with 2% maltodextrin, but not when combined with 16% maltodextrin. In contrast, cluster size was larger with the CS+ than with the CS-, regardless of the concentration of maltodextrin present on test. Previous analyses of licking microstructure indicate that cluster size reflects the palatability of the ingested solution. Thus, the present results indicate that flavor conditioning can change the palatability of the cue flavors. Adding the CS+ flavor to maltodextrin produced results analogous to increasing the concentration of maltodextrin (in terms of both consumption and licking microstructure measures), which is consistent with the idea that after conditioning, responses to the CS+ flavor and to the unconditioned stimulus are mediated via the same representation.