Potentiation and blocking of conditioned flavour and context aversions

Experiments 1, 2, and 3 demonstrated that the place in which rats had been made sick subsequently blocked the development of an aversion to a novel flavour that was presented in that place and paired with illness. These experiments also showed that a poisoned flavour subsequently potentiated the acquisition of an aversion to a novel place in which it was presented and paired with illness. Experiments 4 and 5 examined the hypothesis that the potentiation effect was mediated by the association between the place and the averted flavour rather than by the association between the place and the illness. However, there was no evidence that the place acquired aversive control over ingestive behaviours simply as a consequence of it having contained the averted flavour. Further, reinforcement of the aversive flavour outside the place-flavour compound increased flavour aversions but decreased, rather than increased, place aversions. These failures to detect evidence for an association between the place and the poisoned flavour led to a consideration of whether the animals were sensitive to the relation between the place and the poisoned flavour. Experiment 6 demonstrated that the potentiation effect was in fact contingent upon the relation between the place and the poisoned flavour, as was required by the view that the poisoned flavour had facilitated the association between the place and the illness.     

Enhanced preference for a flavour following reversed flavour—glucose pairing

In two experiments hungry rats showed enhanced preference for a flavour as a result of a procedure whereby the flavour followed presentation of 10% glucose—i.e. the reverse of the usual sequence. This provides the first example of excitatory conditioning resulting from reversed pairing involving an appetitive reinforcer. Experiment 1 compared a Cinnamon group, for which ingestion of glucose was followed after 3 min by cinnamon and on other days water was followed by wintergreen, with a Wintergreen group in which wintergreen followed glucose and cinnamon followed water. In a final 2-bottle test relative preference for cinnamon was greater in the Cinnamon than in the Wintergreen group. This effect was replicated in Experiment 2, which compared an Immediate group, for which, as above, drinking glucose was followed 3 min later by access to cinnamon, with a Delay group, for which there was a delay of 1 hr between glucose and subsequent access to cinnamon. Enhanced preference for cinnamon was found only in the Immediate group and appeared after only four glucose-cinnamon pairings. An important finding came from a choice test between glucose and saccharin in which the Immediate group showed a weaker preference for glucose than the Delay group. This indicated that a novel flavour may overshadow the association between the flavour of glucose and its consequences, which normally serves to enhance preference for glucose.     

Interaction between preexposure and overshadowing: Further analysis of the extended comparator hypothesis

Three experiments with rats used conditioned suppression of barpress to test predictions of the extended comparator hypothesis, which assumes that the effectiveness of (first-order) comparator stimuli in modulating responding to a target conditioned stimulus (CS) is itself modulated by other (second-order) comparator stimuli. Experiment 1 demonstrated that both pretraining exposure to the target CS alone (i.e., CS-preexposure effect, also known as latent inhibition) and pretraining exposure to a compound of the target CS and nontarget CS (i.e., compound-CS-preexposure effect) counteract overshadowing, and that posttraining deflation (i.e., extinction) of the overshadowing stimulus attenuates responding to the target CS when overshadowing is preceded by a CS-preexposure treatment (i.e., yields a CS-preexposure effect), but not when overshadowing is preceded by a compound-CS-preexposure treatment. Experiment 2 examined the consequences of posttraining associative inflation of the overshadowing stimulus or the preexposure companion stimulus following conjoint compound-CS-preexposure and overshadowing treatment. Experiment 3 examined the consequences of posttraining inflation of the overshadowing stimulus or the context following conjoint CS-alone preexposure and overshadowing treatment. The results support the expression-focused comparator view in contrast to recent acquisition-focused models of retrospective revaluation.     

Short-term flavour memory in the rat

In a series of experiments, rats were exposed twice to a flavour at times T1 and T2, and the second of these exposures was followed by toxicosis. The level of the subsequent aversion was viewed as an index of whether the flavour had been recognised as familiar at T2, with a familiar flavour accruing less aversiveness than an unfamiliar one. A flavour was recognised as familiar at time T2 after a long flavour-exposure at time T1 (Experiment Ia) when moderate (3·5 h) and long (27·5 h) T1-T2 intervals were employed but was so recognised after a brief exposure at T1 (Experiment Ib) only when a moderate T1-T2 interval was employed. The memorial processes underlying flavour recognition after a brief flavour exposure were assumed, therefore, to be transient. The remaining experiments employed a brief flavour exposure at T1 and moderate T1-T2 intervals in various attempts to disrupt flavour recognition. Recognition at T2, however, was not disrupted when one (Experiment II), or three (Experiment IV) distractor flavours were interpolated between the target flavour's presentations at T1 and T2. This failure was not due to the distractor having proactively interfered with the associability of the target flavour with illness at T2 (Experiment III). Further, recognition was not disrupted when the position of the target flavour's presentation at T1 was varied across a list of distractor flavours (Experiment V), nor when the similarity of the distractor and the target flavour was varied (Experiment VI). The results indicate that the processes subserving recognition after a brief presentation of that flavour, although transient, are resistant to interference and were discussed in terms of current theories of short-term memory in animals.     

Overshadowing between landmarks on the touchscreen and in arena with pigeons

The role of generalization decrement in spatial overshadowing was evaluated using a landmark-based spatial search task in both a touchscreen preparation (Experiment 1a) and in an Automated Remote Environmental Navigation Apparatus (ARENA, Experiment 1b). A landmark appeared as a colored circle among a row of eight (touchscreen) or six (ARENA) potential locations. On overshadowing trials, Landmark X was located two positions away from a hidden goal, while another landmark, A, was in the position between X and the goal. On control trials, Landmark Y was positioned two locations away from the goal but without a closer landmark. All subjects were then tested with separate trials of A, X, Y, and BY. Testing revealed poor spatial control by X relative to A and Y, thereby replicating the spatial overshadowing effect. Spatial control by Y was similar when tested in compound with novel landmark (BY) and on trials of Y alone. Thus, overshadowing in a small-scale environment does not appear to be due to a process of generalization decrement between training and testing.     

Perceptual and conceptual training mediate the verbal overshadowing effect in an unfamiliar domain

Describing memory for a nonverbal perceptual stimulus can impair recognition. This verbal overshadowing effect may occur when perceptual expertise exceeds verbal expertise, but previous evidence for this hypothesis has been strictly correlational. We conducted two experiments, exploring the relation between expertise and verbal overshadowing by manipulating perceptual and conceptual knowledge. Participants received perceptual training (categorizing mushrooms), conceptual training (lecture), or no training, then viewed a mushroom and described it or not. A recognition test followed. In Experiment 1, perceptual training increased susceptibility to verbalization relative to the no training condition. However, conceptual training did not insulate individuals from the verbal overshadowing effect. Experiment 2 was modified to enhance both forms of training. It replicated the finding that perceptual training increases vulnerability to verbal overshadowing, and indicated that, when sufficiently emphasized, verbalization may be useful following conceptual training. The results demonstrate the role that different forms of learning and expertise may play in mediating language influences on recognition memory.     

A Study of Blocking and Overshadowing in Filial Imprinting

The occurrence of blocking and overshadowing in filial imprinting was investigated in junglefowl chicks (Gallus gallus spadiceus). When subjects were exposed to a novel object in the presence of a familiar one, imprinting on the novel object was impaired in one of two experimental groups (Experiment 1). When subjects were exposed to two objects from the beginning, imprinting on each object was impaired (Experiment 2). These results suggest that phenomena resembling blocking and overshadowing in conditioning may occur in imprinting. The fact that overshadowing was much more prevalent and convincing than blocking is discussed by referring to processes involved in the formation of internal representations. It is suggested that processes connecting these representations to the executive system of filial behaviour may follow the rules of associative learning.     

Overshadowing and stimulus duration

In 3 experiments, the authors investigated the effect of stimulus duration on overshadowing. Experiments 1 and 2 examined responding to a target conditioned stimulus (CS1) when it was conditioned in compound with a coterminating overshadowing stimulus (CS2) that was longer, shorter, or of the same duration (the long, short, and matched groups, respectively). Equal overshadowing of conditioning to CS1 was obtained in all 3 conditions relative to a control group conditioned to the light alone. There were, however, differences in responding to CS2 as a function of its absolute duration. Experiment 3 examined the contribution of the food-food interval/CS onset-food interval ratio to these findings. In Experiments 1 and 2, the ratio differed for the overshadowing CS but not for the target CS. In Experiment 3, this arrangement was reversed, but the pattern of results remained the same. The implications of these findings for trial-based and real-time models of conditioning are discussed.     

Lithium-mediated disruptions of latent inhibition: Overshadowing by the unconditioned stimulus in flavor conditioning

In a series of experiments, the ability of a single lithium preexposure to disrupt CS effectiveness was assessed using a latent inhibition procedure. Lithium preexposure administered proximal (90 min) to a saccharin familiarization trial reduced latent inhibition whereas a similar administration more distal (360 min) to flavor familiarization failed to do so. Additional experiments demonstrated that this socalled “US overshadowing” effect was not attributable to sensitization (Experiment 2), excitatory backward conditioning (Experiment 3), or state dependency (Experiment 4). The implications of US overshadowing for proximal US-preexposure effects are discussed.     

An inhibitory within-compound association attenuates overshadowing

According to the comparator hypothesis (Miller & Matzel, 1988), cue competition depends on the association between a target stimulus (X) and a competing cue (e.g., an overshadowing cue [A]). Thus, it was expected that overshadowing would be reduced by establishing an inhibitory-like relationship between X and A before compound conditioning. In three lever press suppression experiments with rats, this expectation was supported. Experiment 1 showed that establishing an inhibitory X-A relationship reduced overshadowing. In Experiment 2, degrading the inhibitory-like relationship before conditioning allowed reinforced AX compound trials to result in overshadowing. Experiment 3 replicated the results of Experiment 2 when the inhibitory relationship was degraded after compound conditioning. The results support the view that within-compound associations are necessary not only for retrospective revaluation, but also for conventional cue competition.     

Contrasting reduced overshadowing and blocking

Preexposure of a cue without an outcome (X-) prior to compound pairings with the outcome (XZ-->O) can reduce overshadowing of a target cue (Z). Moreover, pairing a cue with an outcome (X-->O) before compound training can enhance its ability to compete with another cue (i.e., blocking). Four experiments were conducted in a conditioned bar-press suppression preparation with rats to determine whether spacing of the X- or X-->O trials would differentially affect reduced overshadowing and blocking. Experiment 1a showed that reduced overshadowing was larger with massed trials than with spaced trials. Experiment 1b found that blocking was larger with spaced trials than with massed trials. Experiments 2a and 2b indicated that these effects of trial spacing were both mediated by the associative status of the context at test. The results are interpreted in the framework of contemporary learning theories.     

Flavour aversion produced by running and attenuated by prior exposure to wheels

In two experiments hungry rats were given access to running wheels. When given the novel flavour, almond, prior to novel access to the wheels, a conditioned aversion to almond was revealed by a subsequent two-bottle test. No such aversion was found in rats with previous experience of wheel running, whether this prior running occurred in the absence of any novel flavour, as in Experiment 1, or following access to saccharin, as in Experiment 2. These results suggest that the failure of rats with prior experience of the running wheels to develop a flavour aversion (unconditioned stimulus, US, preexposure effect) is unlikely to be due to associative blocking. Instead it seems that increasing exposure to a wheel produces habituation of its nausea-inducing properties.     

Contrast effects in flavour preference learning

In four experiments the role of contrast effects in producing learned flavour preferences was examined. The experiments showed that contrast effects are pervasive in flavour preference learning, producing results that are often paradoxical from a traditional reinforcement point of view. In Experiment 1, rats preferred a reinforced flavour over a nonreinforced flavour more if the reinforcer was 1% sucrose than if the reinforcer was 8% sucrose. Because the cue flavour was dissolved in 8% sucrose, this represents an anticipatory positive contrast effect. In Experiment 2, the relationship between cue and consequence was shown to be important in flavour preference learning, as expected if contrast effects are involved. In Experiment 3, rats preferred a flavour that was reinforced 4 times, and they preferred this flavour more than one reinforced 8 times, presumably because the greater the expectancy of the consequence, the greater the anticipatory negative contrast. In Experiment 4, rats consumed less of 0.15% saccharin if they received 32% sucrose randomly 90 min either before or after the 0.15% saccharin than if they received only 0.15% saccharin, a simultaneous negative contrast effect.     

The Influence of Retention Interval on the US Preexposure Effect: Changes in Contextual Blocking over Time

A number of studies manipulating the length of the interval between conditioning and testing indicate spontaneous recovery from overshadowing, suggesting that certain instances of overshadowing represent a deficit in memory retrieval rather than a failure of animals to form an association between the overshadowed stimulus and the US. The present series of experiments examined the influence of lengthening the retention interval on blocking, another stimulus selection phenomenon that is typically interpreted as an acquisition deficit. The results indicated that when subjects were tested shortly (3 days) after training conditioning to a taste blocked subsequent conditioning to an odor conditioned in compound with that taste (Experiment 1), whereas prior conditioning to an odor did not block subsequent conditioning to a taste conditioned in compound with that odor (Experiment 2). This pattern of results was essentially unchanged when testing occurred at a longer (21-day) retention interval. However, there was evidence of a US preexposure effect in Experiment 2 when subjects in the US ONLY control condition were tested at the 3-day retention interval, but not when testing occurred 21 days after conditioning. Experiments 3 and 4 examined whether this loss of the US preexposure effect over time might actually represent a change in the degree of contextual blocking as the retention interval is lengthened. Exposure to the conditioning context either during the interval between Phase 1 and Phase 2 of conditioning (Experiment 3) or prior to Phase 1 of conditioning (Experiment 4) alleviated this US preexposure effect suggesting that the loss of the US preexposure effect as the retention interval is lengthened observed in Experiment 2 is due to changes in the degree of blocking by contextual stimuli over time. The results are discussed in terms of differential susceptibility of forgetting of two functional roles played by a contextual stimuli in the current situation-context as a CS and context as a retrieval cue for other CS-US associations.     

Reinforcement and the organisation of behaviour in golden hamsters: Differential overshadowing of a CS by different responses

Are different responses differentially associable with their consequences? An overshadowing design was used to try to answer this question for three responses previously studied in golden hamsters. In Experiment I, scrabbling was rapidly suppressed by electric shock punishment, and it overshadowed a tone which occurred between scrabbling and shock. In Experiment II, no evidence of response-shock association was obtained when open rearing was the punished response, and open rearing did not overshadow the tone. Punishment had some effect on face washing, but there was no statistically significant overshadowing with this response. These results are consistent with the idea that differences in punishment suppression among these three responses have an associative basis. They also demonstrate the usefulness of a novel paradigm for studying “preparedness” of response-reinforcer associations.     

Retrospective revaluation effects following serial compound training and target extinction

Using a conditioned suppression task, two experiments examined retrospective revaluation effects after serial compound training in a release from overshadowing design. In Experiment 1, serial X → A+ training produced suppression to target A, which was enhanced when preceded by feature X, whereas X by itself elicited no suppression. Subsequent A− presentations extinguished responding to A, but had no effect on either responding to X → A or X alone. However, the addition of A− trials did enhance the ability of feature X to elicit suppression to a novel target, B, suggesting retrospective revaluation of X’s properties. Experiment 2 showed that the enhanced transfer effect, observed in Experiment 1, was independent of the training history of the target (B− or Y → B+/B−). Together, these results suggest that feature X did not retrospectively acquire excitatory strength or occasion setting power, but rather a generalized ability to increase responding to any other cue.     

Proactive interference between cues trained with a common outcome in first-order Pavlovian conditioning

The nature of interference between cues (X, A) trained apart with a common outcome (O; an unconditioned stimulus) was explored by assessing proactive interference in first-order Pavlovian conditioning (i.e., A-O, X-O, resulting in attenuated responding to X). Three lick-suppression studies were conducted with water-deprived rats. Posttraining extinction of the interfering cue (A) attenuated proactive interference (Experiment 1), which mirrors the observation that extinction of the competing cue can reduce competition between cues trained together (e.g., recovery from overshadowing). Proactive interference was also attenuated with manipulations known to attenuate interference between outcomes trained apart (e.g., counterconditioning), namely reminder cues (Experiment 2) and renewal (Experiment 3). The findings suggest that similar processes underlie interference between cues trained apart, between cues trained together, and between outcomes trained apart.     

Blocking and overshadowing in two species of fish.

In Experiment 1, a mixed blocking-overshadowing effect of color on an auditory discrimination was demonstrated in goldfish. In Experiment 2 (with lines differing in color and angle), blocking of angle by color and of color by angle was demonstrated in goldfish, In Experiment 3 (again with lines differing in color and angle), overshadowing of angle by color was demonstrated in carp, but the same animals (like goldfish in a previous study) failed to show greater intradimensional than extradimensional transfer. These results are consistent with the hypothesis that blocking and overshadowing are general phenomena of vertebrate learning. They suggest also that the processes responsible for blocking and overshadowing are different from those which produce the dimensional transfer effect.     

Contextual factors in neophobia and its habituation: The role of absolute and relative novelty

In four experiments we investigated the role of contextual cues in the habituation of neophobia in rats. Experiment 1 showed that the consumption of a novel flavour increased across a series of presentations in one context (A) but fell when the flavour was subsequently presented in a second, novel, context (B). In Experiments 2 and 3, subjects again received exposure to a flavour in context A, but also were familiarized with the test context, B. These subjects consumed the flavour with equal readiness, whether it was presented in Context A or in Context B at test. Experiment 4 replicated the results of Experiment 1 and also showed that the consumption of a novel flavour was not influenced by whether it was presented in a novel or a familiar context. Several mechanisms by which the novelty or familiarity of the context might interact with the novelty or familiarity of the flavour were discussed.