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.     

Differential Acquisition of Aversion by Two Distinctive Contexts Paired with Lithium-Induced Illness

In three experiments using rats as subjects, we investigated the role of the hedonic properties of the CS in context aversion conditioning. In Experiment 1, rats were allowed to drink water in two different contexts, L-S (Light room and Small cage) and D-B (Dark room and Big cage). Rats' consumption of water was higher in the D-B than in the L-S context. This result was taken as evidence for the L-S context eliciting a state of uneasiness that interferes with drinking plain water. In Experiments 2 and 3, rats were injected with lithium after placement in the L-S or the D-B context. In a blocking test (Symonds & Hall, 1997), the L-S context showed conditioning since it blocked the acquisition of aversion by a novel taste, quinine (Experiment 2) or salt (Experiment 3). By contrast, the D-B context failed to show conditioning, that is, no blocking was observed with salt or quinine. These results suggest that the hedonic properties of the context used as the CS may affect its acquisition of associative strength when paired with an internal aversive stimulus.     

CER suppression, passive-avoidance learning, and stress-induced suppression of drinking in the Syracuse high- and low-avoidance strains of rats (Rattus norvegicus)

The Syracuse strains of Long-Evans rats were selectively bred for good (SHA) or poor (SLA) avoidance learning in a two-way shuttle box, which resulted in a phenotypic difference that is correlated with behavior patterns indicative of emotional reactivity, SLA animals showing evidence of greater emotional reactivity than SHA animals. The first three experiments examined conditioned suppression of bar pressing and compared paired and unpaired conditioned- and unconditioned-stimulus presentations to evaluate the influence of conditioning versus primary aversive stimulation on baseline responding. SLA animals acquired conditioned suppression faster than SHA animals and also showed greater suppression of baseline responding than SHA animals. In Experiment 4, SLA animals learned a passive-avoidance task faster than SHA animals. In Experiment 5, SLA animals showed greater stress-induced suppression of drinking a weak quinine solution than SHA animals. These data are consistent with the hypothesis that SLA animals are more emotionally reactive than SHA animals.     

Mere exposure to telereceptive cues facilitates intake of a novel flavor in chickens (Gallus domesticus)

This experiment investigated whether mere exposure to visual cues of a distinctively colored solution (water) would facilitate intake of a novel flavor (vinegar) of the same distinctive color. Eighty-four chicks received access to clear water, clear vinegar, red water, or red vinegar under conditions which permitted drinking or they received access to clear water or red water under conditions (mere exposure) which prevented drinking. All chicks were then tested on intake of red vinegar. Chicks which had received mere exposure or ingestional experience with red-colored water drank reliably more red vinegar than did chicks which had received prior ingestional experience with the novel taste, vinegar, but not with the visual cues of red. However, intake of red vinegar was reliably greater following ingestional experience with red water than following visual exposure without ingestion. Results supported the mere exposure hypothesis.     

Aversiveness of denatonium saccharide and quinine in grasshopper mice Onychomys leukogaster

The responses toward denatonium saccharide and quinine by three rodent species differ from those exhibited by humans. This study measured the relative aversiveness of these two compounds in grasshopper mice (Onychomys leucogaster) to determine if this species responded similarly to or differently from these other rodents. In a choice test, the mice clearly preferred quinine over denatonium saccharide at concentrations of 1/10,000 and 1/50,000, but only marginally preferred quinine at a concentration of 1/1000. When denatonium or quinine was paired with toxicosis, both suppressed drinking at concentrations from 1/10,000 to 1/50,000, but only denatonium suppressed drinking at 1/100,000. Neither compound suppressed drinking at 1 part in million. Grasshopper mice perceived denatonium saccharide as more aversive than quinine in both experiments. Such a response resembled the behavior of humans more than that of the other three rodents previously tested.     

Olfactory aversive conditioning in the newborn (3-hr-old) rat impairs later suckling for water and milk

An olfactory conditioning paradigm tested whether newborn rats can acquire a conditioned aversion to olfactory events associated with their first postnatal meal 3-5 hr after birth. Exposure to lemon odor (conditioned stimulus [CS]) paired with intraoral infusions of 0.1% quinine (unconditioned stimulus) resulted in explicit conditioning. Responsiveness to a surrogate nipple providing water in the presence of the CS was significantly lower than the 3 control conditions. The conditioning dramatically suppressed responsiveness to a surrogate nipple providing milk, which normally is expressed voraciously in terms of sustained nipple attachment and milk intake. These findings suggest that as early as 3-5 hr after birth newborn rats are capable of aversive conditioning to odors in the context of suckling behavior.     

Behavioral contrast produced by a signaled decrease in local rate of reinforcement

Pigeons' key pecking in the presence of one stimulus (S1) was reinforced according to a response-dependent variable-interval schedule. Pecking rate during S1 increased (behavioral contrast) when a second stimulus (S2) [associated with either a response-dependent fixed-interval schedule (Experiment I) or a response-independent reinforcement schedule in which reinforcement availability was signaled by visual (Experiment II) or temporal (Experiment III) stimuli] alternated with S1. These experiments suggest that a discriminable, signaled decrease in local reinforcement rate during S2 is an antecedent of the behavioral contrast response rate increases during S1.     

Failure of sodium- and calcium-deficient rats to acquire conditioned taste aversions to the object of their specific hunger.

Attempts were made to condition taste aversions to the objects of two mineral-specific hungers. Both the innate preference of adrenalectomized rats for sodium and the learned preference of parathyroidectomized rats for calcium were studied. None of the sodium-deficient rats poisoned after drinking sodium chloride (NaCl) reached a taste-avoidance criterion, even after nine pairings of salt ingestion with aversive lithium chloride injections. Six of 11 calcium-deficient rats did not meet the salt-avoidance criterion after 10 pairings. Nondeficient control subjects learned to avoid these salt solutions completely after an average of only three such pairings. Besides unmasking a surprising degree of similarity between the learned and innate specific hungers studied, the results clearly demonstrate a powerful influence of physiological need on aversion conditioning.     

The effects of a pre-time-out stimulus on matching-to-sample of humans,

In each of two experiments human subjects were intermittently reinforced with money on a fixed-ratio schedule for emitting correct matching responses. A pre-time-out stimulus which signaled removal of positive reinforcement was periodically superimposed. In the first experiment the superimposed pre-time-out stimulus was paired with a 1-min or 4-min response-independent time out. In the second experiment the pre-time-out stimulus was paired with a 1-min or 4-min time out contingent on the incorrect responses. The pre-time-out stimulus did not markedly influence performance when the time out was response independent. In contrast, the pre-time-out stimulus markedly suppressed incorrect responding when the time out was contingent on the incorrect responses. When duration of this time-out was increased from 1-min to 4-min, suppression of incorrect responding increased and correct responding was suppressed. Therefore, behavioral suppression by a pre-time-out stimulus was obtained only when the signaled aversive event-time out-was response produced. In this case, suppression was influenced by time-out duration.     

Potentiation and overshadowing in odor-aversion learning: Role of method of odor presentation,the distal-proximal cue distinction,and the conditionability of odor

Four experiments with rat subjects examined interactions between odors and tastes in compound aversion conditioning. In Experiments 1 and 2, a saccharin taste potentiated the conditioning of an almond odorant presented on a cup near the location of the drinking spout, but overshadowed the same odorant when it was mixed in drinking water. This difference depended on the type of odor administered during conditioning rather than testing (Experiment 2). Experiments 3 and 4 examined implications of two other differences between the cup and drink odors: (1) while the cup odor was a distal cue that suppressed approach to the spout, the drink odor was primarily a proximal cue that suppressed consumption, and (2) the cup odor was less conditionable than the drink odor when the stimuli were conditioned alone. In Experiment 3, unlike the proximal saccharin taste, the proximal drink odor failed to potentiate the conditioning of a different distal odor. In Experiment 4, substantial dilution of the proximal drink's concentration resulted in both weaker conditioning of the stimulus alone, and potentiation of its conditioning by saccharin. The results suggest that the difference in the conditionability of the cup and drink odors, and not their status as distal and proximal cues, may be critical in the production of potentiation. Weakly conditionable target stimuli are uniquely potentiated by saccharin.     

Loss of associability by a compound stimulus comprising excitatory and inhibitory elements

In each of two experiments rats learned a discrimination between a stimulus (A) that signaled shock and a compound stimulus (AB) that signaled no shock. In Experiment 1 it was found that the AB compound acquired excitatory strength only slowly when, in a second phase of training, it was made to signal the occurrence of shock. In Experiment 2 the acquisition of inhibitory strength by the compound was similarly found to be retarded. This second experiment also replicated the results of Experiment 1. The relevance of these results to current theories of latent inhibition and attention is discussed.     

The role of taste and oral somatosensation in olfactory localization

Although there is only one set of olfactory receptors, odours are experienced as smells when sniffing things (e.g., sniffing a wine's bouquet) and as flavours when the olfactory stimulus is present in the mouth (e.g., drinking wine). How this location binding—external versus internal environment—is achieved is poorly understood. Experiment 1 employed a new procedure to study localization, which was then used to explore whether localization is primarily dependent upon simultaneous oral somatosensation. Experiment 2, using solutions of varying viscosity, and Experiment 3, using oral movement of varying vigour, revealed that sniffed odours are not localized to the mouth by somatosensation alone. Instead, Experiment 4 demonstrated that a tastant needs to be present and that increasing tastant concentration generates increasing oral localization. Experiment 5 found that this reliance upon gustation reflects the previously observed “confusion” that people show for taste and smell stimuli in the mouth. We suggest that this “confusion” reflects the gustatory system's superior ability to suppress olfactory attention, thus assisting flavour binding.     

Conditioned suppression, punishment, and aversion

Three experiments were conducted to assess the aversive properties of a visual stimulus in the presence of which one group of birds received response-contingent shock (discriminated punishment) while a yoked group of birds received non-contingent shocks (conditioned suppression). In Experiment 1, presentation of the visual stimulus contingent on key pecking reduced the response rate (conditioned punishment effect) for birds under the conditioned suppression procedure but did not reduce the response rate of birds under the discriminative punishment procedure. Non-contingent shocks also produced greater suppression of responding maintained by positive reinforcement in the presence of a visual stimulus than did response-contingent shocks. In Experiment 2, a greater shock intensity (2 mA) was used. All the differences between the two groups found in Experiment 1 were also found in Experiment 2. Experiment 3 demonstrated that response-contingent shock did not result in a conditioned punishment effect even when positive reinforcers were unavailable during the discriminative punishment schedule. The exteroceptive stimulus that was paired with shock in the conditioned suppression procedure acquired the ability to punish behavior. The exteroceptive stimulus in the discriminative punishment schedule did not acquire this ability.     

Signal modality and choice between signaled and unsignaled food

Choice between signaled and unsignaled response-independent food schedules was assessed in three experiments using a commitment procedure. In Experiment 1, subjects tested with a 5-s visual signal consistently changed from the signaled to the unsignaled schedule. Changing from the unsignaled to the signaled schedule was observed only occasionally and only at low levels. The same outcome was observed in Experiment 2 with different types of visual signals and with different stimulus combinations identifying the signal period, the signal-absent period, and the unsignaled schedule. In Experiment 3 the visual signal was replaced with an auditory signal for four of the subjects tested in Experiment 2. The subjects then changed from the unsignaled to the signaled schedule or showed a substantial reduction in choice for the unsignaled schedule. The data were assessed using a conditioned-reinforcement interpretation of choice.     

Effects of signaled and unsignaled shock on schedule-controlled lever pressing and schedule-induced licking: Shock intensity and body weight

Schedule-controlled lever pressing and schedule-induced licking were studied in rats under a multiple fixed-interval fixed-interval schedule of food reinforcement. Following acquisition of stable rates of pressing and licking, a multiple variable-time variable-time schedule of electric-shock delivery was superimposed upon the baseline schedule. In only one component of the multiple schedule, a 5-sec stimulus preceded each shock (signaled shock). In the other component shock was unsignaled. Several shock intensities (Experiment 1) and body weights (Experiment 2) were studied. Lever pressing and licking were affected similarly by experimental manipulations, although with parametric differences. Depending upon shock intensity and body weight, rates of lever pressing and licking were hardly suppressed, suppressed primarily in the unsignaled shock component (differential suppression), or markedly suppressed in both components. Differential suppression during components with signaled and unsignaled shock and conditioned suppression of responding during the preshock stimulus appeared not to be functionally related. Differential suppression depended more on the discriminability of shock-free time, and on shock intensity, body weight, and the type of response than on the “preparatory” behavior preceding shock.     

US habituation,like CS extinction,produces a decrement in conditioned fear responding that is NMDA dependent and subject to renewal and reinstatement

Just as fear can be learned, it can also be inhibited. The most common way of reducing learned fear is through extinction, where the conditioned stimulus (CS) previously paired with an aversive unconditioned stimulus (US) is repeatedly presented on its own. Another, much less commonly studied, way to inhibit learned fear is by habituating, or devaluing, the US. In this procedure, fear responding to a CS is reduced by repeatedly presenting the US in the absence of the CS following the conditioning phase. The purpose of the present study was to directly compare the effects of US habituation and CS extinction on a learned fear response (freezing). Experiment 1 demonstrated that US habituation given either after (Experiment 1A) or before (Experiment 1B) fear conditioning reduced freezing to the CS at test. We then showed that the reduction in freezing resulting from either US habituation or CS extinction was context-specific (i.e., a change in context led to a renewal of the learned fear response; Experiment 2) and, furthermore, was attenuated when a pre-test shock was given (i.e., reinstatement of fear was observed in both cases; Experiment 3). Finally, Experiment 4 demonstrated that an injection of the NMDA antagonist MK-801 prior to US habituation impaired long-term retention of the learning that takes place during this procedure. Together, these results suggest that the decrement in conditioned fear responses produced by US habituation and CS extinction could rely on overlapping processes.     

Intertrial interval as a contextual stimulus: further analysis of a novel asymmetry in temporal discrimination learning

Four experiments investigated discrimination learning when the duration of the intertrial interval (ITI) signaled whether or not the next conditional stimulus (CS) would be paired with food pellets. Rats received presentations of a 10-s CS separated half the time by long ITIs and half the time by short ITIs. When the long ITI signaled that the CS would be reinforced and the short interval signaled that it would not be (Long+/Short-), rats learned the discrimination readily. However, when the short ITI signaled that the CS would be reinforced and the long interval signaled that it would not (Short+/Long-), discrimination learning was much slower. Experiment 1 compared Long+/Short- and Short+/Long- discrimination learning with 16-min/4-min or 4-min/1-min ITI combinations. Experiment 2 found no evidence that Short+/Long- learning is inferior because the temporal cue corresponding to the short interval is ambiguous. Experiment 3 found no evidence that Short+/Long- learning is poor because the end of a long ITI signals a substantial reduction in delay to the next reinforcer. Long+/Short- learning may be faster than Short+/Long-because elapsing time involves exposure to a sequence of hypothetical stimulus elements (e.g., A then B), and feature-positive discriminations (AB+/A-) are learned quicker than feature-negative discriminations (A+/AB-). Consistent with this view, Experiment 4 found a robust feature-positive effect when sequentially presented CSs played the role of elements A and B.     

Cholinergic dependence of taste memory formation: evidence of two distinct processes

Learning the aversive or positive consequences associated with novel taste solutions has a strong significance for an animal's survival. A lack of recognition of a taste's consequences could prevent ingestion of potential edibles or encounter death. We used conditioned taste aversion (CTA) and attenuation of neophobia (AN) to study aversive and safe taste memory formation. To determine if muscarinic receptors in the insular cortex participate differentially in both tasks, we infused the muscarinic antagonists scopolamine at distinct times before or after the presentation of a strong concentration of saccharin, followed by either an i.p. injection of a malaise-inducing agent or no injection. Our results showed that blockade of muscarinic receptors before taste presentation disrupts both learning tasks. However, the same treatment after the taste prevents AN but not CTA. These results clearly demonstrate that cortical cholinergic activity participates in the acquisition of both safe and aversive memory formation, and that cortical muscarinic receptors seem to be necessary for safe but not for aversive taste memory consolidation. These results suggest that the taste memory trace is processed in the insular cortex simultaneously by at least two independent mechanisms, and that their interaction would determine the degree of aversion or preference learned to a novel taste.     

An investigation into blocking of filial imprinting in the chick during exposure to a compound stimulus

The occurrence of "blocking" was investigated in jungle fowl chicks (Gallus gallus spadiceus B.) in an imprinting situation. In Experiment 1, chicks were simultaneously exposed to two stationary coloured cylinders, either two red cylinders (Group RR), a yellow and a red cylinder (YR), or two yellow cylinders (YY). After six days of exposure, the cylinders were removed from the cages and replaced by a yellow and a blue cylinder (i.e. YB) for each chick. This second phase of the experiment lasted for seven days. When the blue cylinder was presented alone during tests at different stages in Phase 2, the RR birds spent significantly more time with this cylinder and emitted fewer shrill calls than the chicks in the YR and YY groups. In Experiment 2, RR and YY birds were reared as in Experiment 1, except that in the second phase of the experiment they were exposed to a blue cylinder only. In this experiment the development of an attachment to the novel blue cylinder proceeded similarly in the RR and YY birds. In Experiment 3, it was found that chicks that were reared with a yellow and a red cylinder preferred the latter stimulus. Thus, although in the first phase of Experiment 1 the RR birds had been exposed to a more attractive stimulus, in tests during the second phase they spent more time with a novel stimulus (B) than the YY birds. These results are consistent with the suggestion that imprinting to a novel stimulus is "blocked" to some extent when that stimulus is presented in compound with another stimulus to which the animal has previously been exposed.     

Associative modulation of the orienting response: distinct effects revealed by hippocampal lesions

The ability of auditory stimuli to modulate rats' tendency to orient to visual targets was assessed. In Experiment 1, trials where an auditory stimulus (A) signaled one visual array (X) were intermixed with unsignaled presentations of a second array (Y). Comparison of the orienting responses (ORs) to X and Y revealed that A produced a transient (unconditioned) and an emerging (conditioned) disruptive influence on the OR to X. In Experiments 2 and 3, trials where A signaled X were intermixed with others where another auditory stimulus (B) signaled Y. Stimulus A's ability to modulate the OR to X was then assessed by presenting A prior to test arrays containing both X and Y. Control rats were more likely to orient to Y than X (Experiments 2 and 3) and rats with excitotoxic lesions of the hippocampus were more likely to orient to X than Y (Experiment 3). These results show that auditory stimuli exert distinct modulatory influences on the OR to visual stimuli with which they are associated.