Caloric balancing and sucrose intake of hamsters and gerbils

In Experiment 1, hamsters and gerbils were randomly assigned to groups that were given a choice between pairs of sucrose solutions differing in concentration. The groups consisted of a control along with the following pairs: 2% and 6%; 2% and 8%; 2% and 12%; 2% and 18%; 6% and 8%; 6% and 12%; 6% and 18%; 12% and 18%. They were tested for 15 days, and the results indicated that the animals drank more from the cylinder containing the higher concentration. The principle of "least consummatory effort" appeared to be substantiated by these results. In Experiment 2, other hamsters and gerbils were randomly assigned to one of six groups receiving either 0.5%, 2%, 6%, 10%, 14%, or 18% sucrose solution. Analysis of their intake of sucrose and Purina Lab Chow indicated that their total caloric intake was relatively constant, reflecting the fact that an increase of consumption of the solutes in the sucrose solution reduced their intake of the stock diet. The results from both experiments indicated that although the caloric intake of hamsters relative to body weight was greater than that of gerbils, the proportion of total calories taken from sucrose was greater for the latter than for the former.     

Flavour neophobia in gerbils (Meriones unguiculatus) and hamsters (Mesocricetus auratus)

Flavour neophobia in gerbils and hamsters was studied by comparing the ingestive/pouching behaviour of these animals when presented with either a familiar unflavoured peanut or a novel one that imparted basic flavours such as sweet, salty, sour, or bitter. Neophobia was expressed by an aversion towards the novel nut manifested by greater latency either to ingest it (gerbils) or to deposit it in the cheek pouch (hamsters) relative to that of the unflavoured one. This method, involving reactions to the taste of novel solid food, provides a more sensitive measure than the traditional one measuring intake of a novel flavoured fluid. The results indicated that gerbils and hamsters encountering either a sweet, salty, or sour nut for the first time showed a neophobic reaction. However, there was no apparent sign of neophobia when the animals were tested with a nut flavoured with quinine. The theoretical implications of these findings are discussed.     

Negative anticipatory contrast and preference conditioning: flavor cues support preference conditioning, and environmental cues support contrast.

In 2 experiments, access to a 0.15% saccharin solution was followed on alternating days by access to a 32% sucrose solution and the same saccharin solution. In Experiment 1, rats increased both intake of and preference for a flavored saccharin solution that predicted sucrose, but neither effect was found using a predictive odor cue alone. Experiment 2 replicated the predictive flavor results but showed suppression of saccharin intake when environmental cues predicted sucrose. When both flavor and environment predicted sucrose, saccharin intake did not change, but preference for the predictive flavor increased. Discriminative taste cues appear to facilitate the development of preference conditioning, but environmental cues favor negative anticipatory contrast effects. Also, preference conditioning and contrast may develop concurrently and compete for expression.     

Substitution and caloric regulation in a closed economy.

Three experiments were conducted to study the effect of an imperfect substitute for food on demand for food in a closed economy. In Experiments 1 and 2, rats pressed a lever for their entire daily food ration, and a fixed ratio of presses was required for each food pellet. In both experiments, the fixed ratio was held constant during a daily session but was increased between sessions. The fixed ratio was increased over a series of daily sessions once in the absence of concurrently available sucrose and again when sucrose pellets were freely available. For both series, increases in the fixed ratio reduced food intake, but body weight was reduced only in the no-sucrose condition. In the sucrose condition, body weight and total caloric intake (sucrose plus food) were relatively unaffected by increases in the fixed ratio. At all fixed ratios, food intake was proportionally reduced by the intake of sucrose. In Experiment 3, monkeys obtained food or saccharin by pressing keys; the fixed ratio of presses per food pellet was increased once when tap water was each monkey's only source of fluid, again when each monkey's water was sweetened with saccharin, and a third time when each monkey had concurrent access to the saccharin solution and plain water. Increases in the fixed ratio, but not the intake of the saccharin solution, reduced each monkey's food intake. Because neither rats' sucrose nor monkeys' saccharin intakes affected the slope of the respective demand curves for food, monkeys and rats increased their daily output of presses and thereby defended their daily intake of those complementary elements of food. However, sucrose reduced rats' food intake. The relative constancy of body weight and total caloric intake in the sucrose condition is consistent with the possibility that rats tended to regulate caloric intake.     

Response latency of gerbils and hamsters to nuts flavoured with bitter-tasting substances

The unconditioned reactions of Mongolian gerbils to peanuts rolled in sucrose octa-acetate (SOA) or denatomium benzoate (Bitrex) were studied in Experiment 1. Although SOA evokes as high a bitterness rating among humans as quinine salts, the gerbils' latency to ingest a nut flavoured with the former was similar to that to an unflavoured nut. Their oral latency to a nut that was flavoured with Bitrex was longer than to either an unflavoured one or one flavoured with SOA. However, the fact that they ingested a nut flavoured with the “most bitter substance known to man” within 70 sec after contact is surprising. Experiment 2 dealt with the effects of a peanut that was either unflavoured or flavoured with SOA or quinine and then paired with a LiCl injection. The oral latency of these gerbils to a nut conveying such flavours was assessed for six days after this conditioning. A nut flavoured with quinine acquired aversive properties but not ones that were flavoured with SOA or were unflavoured. Similar results were observed in Experiment 3, when hamsters were tested under comparable conditions. The theoretical implications of these results are discussed.     

Apparent Incentive Contrast Effects in Autoshaping with Rats

The effects of shifts in reward quality and quantity on Pavlovian acquisition were studied in rats. In Experiment 1, animals preexposed to unsignaled food pellets, 10% sucrose solution, or home cage controls subsequently received autoshaping training (response-independent lever-pellet or lever-solution pairings, in three groups each). Unsignaled preexposure to sucrose solution facilitated autoshaping for pellets (relative to unshifted controls), whereas unsignaled preexposure to pellets retarded autoshaping for sucrose solution. In Experiment 2, unsignaled preexposure to 30% sucrose solution impaired acquisition reinforced by food pellets, relative to 2% solution. Using a choice procedure, Experiment 3 demonstrated that rats prefer pellets to either 2 or 10% sucrose solutions, but they prefer the 30% solution to the pellets. Experiment 4 demonstrated the facilitatory effect after an upward shift in reward magnitude rather than quality (from 1 to 12 pellets), but provided weaker evidence for retardation following a downward magnitude shift. Experiment 5 was similar to Experiment 4, except that animals received autoshaping training from the outset. No evidence of successive positive contrast was obtained, but there was a significant successive negative contrast effect. Moreover, extinction was faster after acquisition with 12 pellets rather than 1. These results suggest the presence of incentive contrast effects under Pavlovian training conditions.     

Learned preference for a hedonically negative flavor is observed after pairings with positive post-ingestion consequences rather than with a palatable flavor

In two experiments, thirsty rats consumed a compound of sucrose and a non-preferred flavor. In Experiment 1, a conditioned preference was observed in the experimental group when animals were tested both thirsty and hungry, but not when they were tested just thirsty. Animals in the control group, which experienced the flavor and the sucrose unpaired, never showed a preference. Experiment 2 replicated the absence of a preference in the experimental group when rats were tested thirsty, but provided evidence that a flavor-taste association had been formed during training. After conditioning, sucrose was paired with LiCl in group Dev whereas it was unpaired in group NonDev. The sucrose devaluation produced a decrease in CS preference in group Dev, and an increment in group NonDev. Taken together, these results show that preference for a non-preferred flavor can be readily observed after pairings with the positive consequences of the US (calories or absence of an expected illness) rather than with a palatable flavor.     

Incentive learning and the motivational control of instrumental performance

Hungry rats were trained to press a lever and pull a chain concurrently, with one action being reinforced with a sucrose solution and the other with food pellets. In addition, in the first two experiments all animals experienced non-contingent presentations of the two incentives in the absence of the operant manipulanda while either thirsty or hungry and either before (Experiment 1A) or after (Experiment 1B) the instrumental training. When lever pressing was assessed subsequently in extinction under thirst, the animals pressed at a relatively high rate only if (1) this action had been reinforced with the sucrose solution rather than the food pellets during training and (2) they had received the non-contingent presentations of the sucrose solution and food pellets on days on which they were thirsty rather than hungry. A third experiment demonstrated that non-contingent exposure to the sucrose solution alone, but not to water under thirst was sufficient to bring about this type of motivational control of instrumental performance.     

From contrast to reinforcement: role of response contingency in anticipatory contrast

Intake of a 0.15% saccharin solution is suppressed if access to the saccharin is followed by access to 32% sucrose in brief daily pairings. The present series of four experiments was concerned with factors that lead to this anticipatory contrast effect (suppressed saccharin intake) rather than a reinforcement effect. In Experiment 1, anticipatory contrast was obtained with an autoshaping procedure (no lick requirement on the initial tube), and degree of contrast did not vary as a function of intersolution interval in the range of 0-15 s. Experiments 2 and 3 showed that requirements of 10, 100, 200, or 400 licks on the first tube available led to a reinforcement effect in latency, but a requirement of 0 licks (autoshaping procedure) led to a contrast effect in licks and latency. In Experiment 4, a group with a 200-contingent-lick requirement showed a reinforcement effect in latency, but a group yoked to this contingent group showed a contrast effect in both latency and licks. Overall, the results suggest that anticipatory contrast occurs under conditions of a "relaxed" instrumental contingency. The data are discussed in terms of control of behavior by stimulus-stimulus, response-stimulus, and stimulus-response associations, and the results are related to behavioral contrast, to flavor-outcome associations, and to "misbehavior" produced by Pavlovian-instrumental interactions.     

Ethanol-mediated taste aversions and state-dependency in preweanling (16-day-old) rats

Requirements for conditioning of an ethanol-mediated taste aversion in 16-day-old rat pups were examined. Experiment 1 demonstrated that preweanling rats are capable of acquiring, in two trials, an aversion to a 15% sucrose solution when followed by intragastric intubation of a 1.2 g/kg dose of 17% v/v ethanol, but not when followed by a 0.4 g/kg dose. Comparison was with control animals given sucrose followed by an equivalent volume isocaloric Half and Half. When the 0.4 g/kg dose of ethanol preceded sucrose presentation by 30 min (Experiment 2), the aversion was learned, suggesting that the effective delay between the sucrose and the critical consequences of the ethanol had been too long with the former procedure. Expression of the sucrose aversion required, however, the reinstatement of the context of intoxication--state-dependent retention. Finally, the results of Experiment 3B indicated that, in addition to the association between the sucrose and the aversive consequences of alcohol intoxication, the orosensory cues resulting from alcohol's direct elimination, via such processes as respiration and salivation, became associated with the appetitive properties of the sucrose. This was evidenced by a conditioned increase in preference for ethanol odor. Possible age-related differences in the ability to associate stimuli with alcohol's unconditioned consequences, and in state dependency are discussed.     

Genetic differences in avoidance learning by Rattus norvegicus: escape/avoidance responding, sensitivity to electric shock, discrimination learning, and open-field behavior

The behaviors of rats selectively bred for either good or poor shuttle box avoidance learning were studied. The results of Experiment 1 indicated that the phenotypic difference in avoidance learning is not associated with differences in speed of escape or avoidance responding. Differences between the lines in frequency of intertrial responses (ITRs), which appear during training but not during pretest, suggest that ITRs in animals of the low-avoidance (SLA) line are more suppressed by electric shock than in animals of the high-avoidance (SHA) line. This result suggests that SLA animals may be more emotionally responsive than SHA animals. Experiment 2 demonstrated that the animals of the two lines do not differ in absolute sensitivity to electric shock, and Experiment 3 showed that the poor performance of the SLA line is not due to an inability to learn. Experiment 3 also provided evidence which suggests that the poor avoidance learning by SLA animals is due to their emotional reactivity. Observations of open-field behavior in Experiment 4 are consistent with this hypothesis. The major consistent correlate of the phenotypic difference in avoidance learning is greater emotionality or emotional reactivity in SLA than in SHA animals.     

Duration discrimination is better with food access as the signal than with light as the signal

In Experiment 1 a go/no-go discrimination procedure was used to compare control of five pigeons' keypecking by food-access duration with control by light duration. Pecks to an illuminated key were reinforced with grain following 10-sec presentations of food access or houselight, but not after 5-sec presentations of either stimulus. Each subject discriminated food-access duration faster and to a greater degree than light duration. In four between-subject replications, pigeons discriminated food-access duration better than the duration of a localized light, the feeder light and a keylight, and with either water or food as reinforcement. In Experiment 2 control by durations of food access and light was compared using a conditional right-left choice procedure (two pigeons), and a delayed symbolic matching-to-sample procedure (six pigeons). Under both, choice accuracy again was higher on food-access trials. The results of Experiment 3, in which two pigeons received generalization trials with durations of food access and light that were intermediate to the training values, confirmed that responding was controlled by the duration dimension of both food access and light. The superior control by food access is consistent with previous evidence that food is an effective and memorable stimulus, possibly because of its biological importance. These results also provided empirical support for the commonly made assumption that stimuli differ in effectiveness. As well, the results show that the stimulus to be discriminated can play an important role in the accuracy of duration disciminations, a fact which has implications for the study of temporal discriminations in animals.     

Motivational control of instrumental performance following a shift from thirst to hunger

Thirsty rats were trained to press a lever for either a sucrose solution or saline before performance was tested in extinction while the animals were either hungry alone or experiencing both hunger and a sodium appetite. Reinforcer-specific motivational control was observed in that the animals trained with the sucrose solution pressed more than those trained with the saline when they were tested hungry, but not when they were tested under combined hunger and sodium appetite. In order to assess the role of a Pavlovian incentive process in this effect, thirsty animals received non-contingent pairings of one stimulus with the sucrose solution and another with saline in the second experiment. In an extinction test the sucrose stimulus augmented lever pressing relative to the saline stimulus when the animals were hungry, but not when they were thirsty. In the subsequent experiments the contribution of the Pavlovian process was equated by giving concurrent training with both incentives. Lever pressing and chain pulling were reinforced concurrently, one with the sucrose solution and the other with saline, while the animals were thirsty. Once again, the animals pressed more in extinction if this action had been trained with the sucrose solution rather than the saline, but only if they were hungry rather than thirsty. Thus, instrumental performance across a thirst-to-hunger shift can also be controlled by an instrumental incentive process. The direct engagement of the instrumental process by this motivational shift contrasts to the absence of such control following a hunger-to-thirst transition (Dickinson & Dawson, 1987a), a fact attributed to the asymmetrical motivational interactions produced by water and food deprivation.     

Signalling and incentive processes in instrumental reinforcer devaluation

We have previously reported that conditioning an aversion to the reinforcer using an isotonic lithium chloride (LiCl) solution following instrumental training reduces performance in a subsequent extinction test only if animals are re-exposed to the reinforcer prior to the test. Rescorla (1992), in contrast, reported an immediate devaluation effect using a hypertonic LiCl solution that did not depend upon re-exposure. In two experiments we examined the effect of using a hypertonic LiCl solution to condition the aversion to the reinforcer on subsequent instrumental performance in extinction, with and without re-exposure. In Experiment 1 thirsty rats were trained to press a lever for a sucrose solution before being injected with 0.6 M LiCl either immediately or after a delay. Half of the immediate and delay groups were then re-exposed to the sucrose in the absence of the lever, with the remainder being exposed to water. Contrary to the previously reported effects of isotonic LiCl, a hypertonic solution induced a reinforcer devaluation effect in all the immediately poisoned animals, which did not depend upon re-exposure to the reinforcer. In Experiment 2 the possibility that this devaluation effect was induced by the discomfort associated with the hypertonicity of the solution was assessed by replicating Experiment 1 but, in addition, using two immediately poisoned groups given the LiCl injection under anaesthesia. In the absence of anaesthesia, the devaluation effect observed without re-exposure to the reinforcer in Experiment 1 was replicated. When the injection was given under anaesthesia, however, a reinforcer devaluation effect was observed only in animals that were re-exposed to the reinforcer prior to the extinction test. These results were interpreted as evidence that a reinforcer devaluation effect induced by pairing the reinforcer with illness depends upon a process of incentive learning, whereas a devaluation effect mediated by learning a signalling relationship between the reinforcer and somatic discomfort does not.     

Determinants of the enhancement of flavored-water intake by prior exposure.

The intake of a 2.0% sodium saccharin solution in rats was observed to increase as a function of both the number (Experiment 1) and the duration (Experiment 3) of prior periods of access to the saccharin flavor, but did not increase when subjects were maintained on a fluid deprivation procedure in the absence of saccharin exposure (Experiment 2). The enhancement of intake was further influenced by the schedule of saccharin preexposures in the absence of variations in the amount of solution tasted (Experiment 4). The effect was not a function of the opportunity for subjects to determine their own pattern of contact with the saccharin flavor, the opportunity for association of the flavor with hunger and thirst reduction, or the amount of saccharin swallowed during preexposure (Experiment 5). These results suggest that mere exposure to a flavored solution is sufficient to increase subsequent intakes. The phenomenon is discussed in terms of the attenuation of neophobia elicited by the novelty of flavored solutions.     

Alternating exposure to two compound flavors creates inhibitory associations between their unique features

Rats were exposed to two compound solutions, saline-lemon and sucrose-lemon. In Group ALT, trials with one solution alternated with trials with the other. Group BLK received all trials with one solution before any trials with the other. Previous retardation tests had implied that only alternating exposure would establish sucrose as an inhibitor of saline. To provide a complementary summation test for this inhibition, in Experiment 1, all the animals received pairings of peppermint and saline and were tested for consumption of peppermint-sucrose under sodium depletion. Consumption was increased by sodium depletion only in Group BLK. In Experiment 2, a retardation test was used to show that presentation of saline-lemon before sucrose-lemon on each exposure day would establish sucrose as an inhibitor of saline. Neither exposure to sucrose-lemon before saline-lemon nor alternating exposure to sucrose and saline alone had the same effect. These results provide support for an associative theory of perceptual learning that suggests that exposure to complex stimuli aids later discrimination partially as a result of establishing inhibitory associations between their unique elements.     

Within-subjects positive and negative contrast effects in rats.

Rats were given alternating 1-min. access periods to 2 tubes containing either 32% or 4% sucrose solutions for daily 6-min. test sessions. Lick rate for 32% was higher under comparison (32 vs. 4) than noncomparison (32 vs. 32) conditions; and lick rate for 4% was lower under comparison conditions (4 vs. 32) than under noncomparison conditions (4 vs. 4). All sucrose conditions were varied within subjects and both positive and negative contrast were obtained with a small n. In addition to lick rate, intake and latency measures also revealed contrast effects. Deprivation conditions altered latency but not lick rate measures of contrast. Reducing the test session to 3 min. (alternating 30-sec. access periods) did not greatly affect contrast. Additional experiments provided evidence for distinct within-days and between-days contrast effects, as well as a between-groups contrast effect.     

Food hoarding behaviour in the golden hamster (Mesocricetus auratus): Effects of body weight loss and hoard-size discrimination

Two experiments investigated some of the factors that influence food hoarding behaviour in the golden hamster (Mesocricetus auratus). In Experiment 1, hamsters given limited access to food failed to increase the amount of food they consumed at each meal. Instead, they exhibited a significant increase in the amount of food taken to hoard during the limited access period, and this permitted hamsters who were allowed to retain their hoard to maintain body weight at levels comparable to free-feeding controls. Hamsters that were not allowed to retain hoarded food rapidly lost body weight. There was also an inverse correlation between body weight and amount of food taken to hoard in all hamsters given limited access to food, suggesting that body weight loss was one factor directly related to increased hoarding during periods of limited food accessibility. Experiment 2 investigated the effect of hoard-size discrimination on food hoarding in hamsters by artificially manipulating hoard size. The results suggested that hoarding behaviour could be influenced by artificially depleting and repleting the hoard, and this factor appears to influence hoarding in the absence of any significant changes in body weight. These results are discussed in relation to the kinds of adaptive feeding strategies that hamsters may utilize in their natural environment.     

Effects of response–stimulus interval manipulation and articulatory suppression on task switching

We conducted three experiments using a list paradigm to examine how articulatory suppression and response–stimulus interval (RSI) manipulation affected task switching. Experiments 1 and 2 tested task-switching performance under a short and long RSI and three concurrent task conditions (control, articulatory suppression, and tapping) without external task cues. The results indicated that alternation had a greater effect under articulatory suppression than under the control and tapping conditions, and that articulatory suppression costs were unrelated to the RSI. In Experiment 3, an external task cue was provided with each stimulus, and the negative effect of articulatory suppression on alternation cost was eliminated. These results indicated that articulatory suppression effects did not differ between conditions of short and long RSI and that the availability of verbal representations of task information was independent of RSI length. This paper discussed the possible roles played by the phonological loop in task-switching paradigms.