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.     

Undernutrition during suckling and latent learning ability of rehabilitated adult male rats

The present report investigates the effects of early undernutrition on the latent learning ability of rehabilitated adult male rats in a simple maze task. Rats were undernourished during suckling by feeding their dams an 8% casein diet. Well-nourished dams received a 25% casein diet during the same period. Rats were weaned at 21 days of age and nutritionally rehabilitated until they became adults, when behavioral task was conducted. Under a nonappetitive condition, rats were exposed either to an open field or to a maze apparatus. They were thereafter deprived of water and tested in the maze apparatus. Both well-nourished and undernourished rats that had been previously exposed to the maze performed better than those exposed to the open field. Nutritional treatment had no effect on performance of either the latent learning or of the open-field groups. These results suggest that rehabilitated adult rats are able to learn about the environment when no immediate reinforcement is involved. The discrepancy between our findings and results reported by others may be due to differences in task complexity and/or perhaps to the fact that nutritional rehabilitation also plays a role in reversing some of the deleterious effects of early undernutrition on learning ability of rats.     

Risky choice as a function of amount and variance in food supply.

In Experiment 1, 4 rats earned their daily food ration by choosing on a trials basis between a "risky" and a "riskless" lever. The risky lever produced either 15 45-mg food pellets or no pellets, and on average provided five pellets per choice. The riskless lever always produced three pellets. Across conditions, the number of trials per session was varied. Body weight and choice of the risky lever decreased as the number of trials per session decreased, even though body weight could only be defended by increased choice of the risky lever. In Experiment 2, trials per session were fixed, but the number of pellets delivered by the risky and riskless levers was either at the same level as in Experiment 1 or tripled from those levels. Now choice of the risky lever was inversely related to the size of reinforcement and to body weight. The results of these experiments show that risk aversion covaries with the amount of food available in a session and the daily variance in the amount of food earned.     

Adult blood pressure reduction in spontaneously hypertensive rats reared by normotensive Sprague-Dawley mothers.

Spontaneously hypertensive (SHR), Wistar-Kyoto (WKY) normotensive, and Sprague-Dawley (SD) normotensive rats were either reared by their natural mothers (controls) or fostered to a mother of one of the other strains. The effects of rearing environment were assessed by measures of body weight at weaning and adult resting mean arterial blood pressure (MAP) and heart rate (HR). At weaning, SHR control rats weighed less than WKY controls which in turn weighed less than SD controls. SHR and WKY rats reared by SD mothers weighed significantly more than their respective controls, whereas SD rats reared by either SHR or WKY mothers weighed less than SD control rats. Manipulation of the maternal environment was attended by a change in adult resting MAP in only one group. SHR rats reared by SD mothers had MAPs which were significantly lower than SHRs reared by their natural mothers. No within-strain adult HR differences were observed between control and cross-fostered groups. These data indicate that the effects of cross-fostering on adult resting MAP previously reported for SHR and WKY animals are generalizable to another normotensive strain. That is, genetically hypertensive (SHR) animals exposed to a normotensive (WKY or SD) maternal environment display a reduction in adult resting MAP, whereas hypertension or an increase in adult MAP is not induced in normotensive rats (WKY or SD) as a consequence of exposure to a hypertensive maternal environment.     

Visual Reinforcement Signals Interfere with the Effects of Reinforcer Magnitude Manipulations

Three experiments examined the effect of reinforcement magnitude on free-operant response rates. In Experiment 1, rats that received four food pellets responded faster than rats that received one pellet on a variable ratio 30 schedule. However, when the food hopper was illuminated during reinforcer delivery, there was no difference between the rates of response produced by the two magnitudes of reward. In Experiment 2, there was no difference in response rates emitted by rats receiving either one or four pellets of food as reward on a random interval (RI) 60-s schedule. In Experiment 3, rats responding on an RI 30-s schedule did so at a lower rate with four pellets as reinforcement than with one pellet. This effect was abolished by the illumination of the food hopper during reinforcement delivery. These results indicate that the influence of magnitude is obscured by manipulations which signal the delivery of reinforcement.     

Growth and social behavior of previously undernourished mice

Developing mice were growth-retarded by underfeeding of their mothers during lactation. Control mothers were well nourished throughout. All offspring were fed ad libitum from weaning till their behaviour was tested in adulthood. The undernourished mice had weight deficits at weaning of 49% compared with controls. Nevertheless, they showed partial catch-up growth thereafter, in that their absolute weight deficits decreased between weaning and adulthood. Social behaviour observations were made on pairs of mice of the same treatment group. Previously undernourished mice were slower to start attacking one another, spent more time near or in contact with one another and did less tail-rattling than their controls.     

Altered grooming responses to stress in rats exposed prenatally to ethanol

The effects of prenatal alcohol exposure on grooming, locomotion, and rearing in response to stress were examined in adult rats whose mothers consumed a liquid diet containing 35% ethanol-derived calories (EDC). Offspring of both pair-fed 0% EDC mothers and ad libitum chow-fed mothers were included as controls. In Experiment 1, females groomed more than males following placement into a novel test chamber, but no differences due to prenatal treatment were observed. Ethanol-exposed animals groomed more than controls following the stress of a forced 1-min swim (Experiment 2), but when rats tested in Experiment 1 were observed again after forced swim stress (Experiment 3), no differences due to prenatal treatment or sex were observed. Experiment 4 examined the effects of pretreatment with 1 mg/kg naloxone on novelty-induced grooming and as in Experiment 1 prenatal treatment did not affect grooming responses. Females again groomed significantly more than males and naloxone reduced grooming equally for all groups. The results suggest that novelty-induced grooming is a sex-influenced behavior, with females grooming more than males, and that animals exposed prenatally to alcohol and tested as adults may have altered responses to certain stressors (i.e., forced swim) under specific conditions. The altered grooming response of alcohol-exposed rats to swim stress can be eliminated by preexposing them to novelty stress.     

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.     

Choice between constant and variable alternatives by rats: effects of different reinforcer amounts and energy budgets

Two experiments, using rats as subjects, investigated the effect of different reinforcer amounts and energy budgets on choice between constant and variable alternatives under a closed economy. Rats were housed in the chamber and were exposed to a modified concurrent-chains schedule in which the choice phase was separated from a rest phase during which the rats could engage in other activities. In the choice phase, a single variable-interval schedule arranged entry into one of two equal terminal links (fixed-interval schedules). The constant terminal link ended with the delivery of a fixed number of food pellets (two or three, depending on the condition), whereas the variable terminal link ended with a variable number of food pellets (means of two or three, depending on the condition). Energy budget was defined as positive when body weights were over 90% of free-feeding weights, and as negative when they were under 80% of free-feeding weights. The different body weights were produced by varying the duration of the equal terminal-link schedules within daily 3-hr sessions. In Experiment 1, rats chose between a constant and a variable three pellets under both energy budgets. Rats preferred the constant three pellets more under the positive energy budget, whereas they were indifferent under the negative energy budget. In Experiment 2, rats chose between a constant three pellets and a variable two pellets, and chose between a constant two pellets and a variable three pellets under both energy budgets. The rats strongly preferred the constant three pellets over the variable two pellets under both energy budgets. In contrast, rats preferred the variable three pellets over the constant two pellets only under the negative energy budget, whereas they were indifferent under the positive energy budget. These results indicate that rats choices are sensitive to the difference in reinforcer amounts and to the energy budgets defined by the level of body weight. The present results are consistent with those obtained with small granivorous birds as well as with the predictions of a recent risk-sensitive foraging theory.     

Operant hoarding: a new paradigm for the study of self-control.

In the first of four experiments, rats were exposed to a modified multiple continuous reinforcement-extinction schedule during 15-min daily sessions. In one condition (saves condition) with the cuelight on, a single lever press produced a food pellet, briefly extinguished the cuelight, and started a clock. Saves (additional lever presses with interresponse times less than 1 s) produced an additional food pellet, briefly extinguished the cuelight, and restarted the interresponse time clock. The cuelight was extinguished 1 s after the last lever press and remained off during a 10-s period of extinction, during which no food pellets were delivered. In the other condition (savings account condition), the contingencies were the same except that the cuelight was extinguished and was not reilluminated after the initial lever press, and the delivery of all food pellets in the reinforcement component was delayed until the onset of extinction. In both conditions, rats made saves, but mean saves (total saves divided by the number of reinforcement components) were slightly reduced in the savings account condition. In Experiment 2, using six equally spaced 15-min sessions per day on alternate days, saves were either followed immediately with food and brief cuelight offset (saves condition) or were not reinforced at all. Mean saves were much greater when saves were reinforced. In Experiment 3, during 5-min daily sessions, saves earned a single pellet (savings account condition) or a number of pellets equal to the ordinal number of the lever press (interest condition). Rats made fewer mean saves, with little change in the food rate, when saves earned interest. In Experiment 4, the rats earned all their food in the operant situation during 24 daily 5-min sessions, these separated by 55-min intersession intervals during which no food was available; otherwise, the conditions were the same as in Experiment 3. In Experiment 4, the shift to interest for saves led to an increase in mean daily mean saves (total daily mean saves divided by the number of daily sessions) as well as to an increase in the number of food pellets delivered in each session. The results are discussed in terms of self-control and behavioral economics.     

Weight Loss in Rats Produced by Running: Effects of Prior Experience and Individual Housing

To investigate factors affecting activity-based anorexia (ABA) or activity-stress (AS), rats were given 2-hr access to a running wheel immediately prior to their daily 1.5-hr food access during the light cycle. This produced a reduction in food intake, a steady increase in running, and a large drop in body weight with a prolonged delay before weight recovery began. Experiment 1 found that these effects were reduced in rats with prior experience of eating at this time of day. In contrast, prior experience of running in the wheel when on ad lib food enhanced these effects in Experiment 2, where a subsequent change for half the subjects to individual housing produced a further decrease in body weight. The latter factor was investigated from the outset of Experiment 3 and again individually housed rats showed greater weight loss than did group-housed rats. This experiment also found that in rats of the same age a low initial body weight predicts greater vulnerability to ABA. It was concluded that ABA results from activity-induced reduction of feeding, which prolongs adaptation to a new feeding schedule and is accentuated by social isolation.     

The effect of deprivation level during noncontingent pairings and instrumental learning on subsequent instrumental performance

In Experiment 1, rats received either response-noncontingent pairings of a tone stimulus with food or response-contingent instrumental training during which only responses in the presence of the tone were reinforced. During this training rats were maintained at either 70 or 90% of their growth-adjusted predeprivation body weights. In a subsequent test phase, one-half of the subjects in each training condition were tested under either 70 or 90% body weight. Subjects which had received response-contingent instrumental training under 70% body weight responded significantly faster to presentations of the tone than did 90% body weight-trained subjects, regardless of test deprivation condition. There was no effect of the deprivation level in effect during response-noncontingent pairings. In Experiment 2, rats received noncontingent tone-reward pairings or nonpairings under either 70 or 90% body weight. In a later test phase under 90% body weight, instrumental responding to the tone was significantly faster for subjects which had received the tone-reward pairings. Deprivation level during the pairings again produced no effect. These results support two conclusions. First, the expectancy learning process appears to be relatively independent of deprivation conditions in effect when such learning takes place; and secondly, deprivation conditions in effect during instrumental learning affect the strength of S-R associative bonds formed.     

Context modulation of US signal value

Context modulation of unconditioned stimulus (US) signal value was examined in two experiments. In Experiment 1, rats showed significantly more magazine entries after a single food pellet US in Context A (in which three additional pellets shortly followed) than in Context B (in which the delivery of three additional pellets was delayed). Experiment 2 replicated this effect and showed that any differences in contextual associative strength at the time the single food pellet was delivered did not importantly contribute to context modulation. Implications for theories of occasion setting and context modulation are considered.     

Determinants of reinforcer accumulation during an operant task.

Responses by rats on an earn lever made available food pellets that were delivered to a food cup by responses on a second, collect, lever. The rats could either collect and immediately consume or accumulate (defined as the percentage of multiple earn responses and as the number of pellets earned before a collect response) earned pellets. In Experiment 1, accumulation varied as a function of variations in the earn or collect response requirements and whether the earn and collect levers were proximal (31 cm) or distal (248 cm) to one another. Some accumulation occurred under all but one of the conditions, but generally was higher when the earn and collect levers were distal to one another, particularly when the earn response requirement was fixed-ratio (FR) 1. In Experiment 2, the contributions of responses and time to accumulation were assessed by comparing an FR 20 earn response requirement to a condition in which only a single earn response was required at the end of a time interval nominally yoked to the FR interval. When 248 cm separated the earn and collect levers, accumulation was always greater in the FR condition, and it was not systematically related to reinforcement rate. In Experiment 3, increasing the earn response requirement with a progressive-ratio schedule that reset only with a collect response increased the likelihood of accumulation when the collect and earn levers were 248 cm apart, even though such accumulation increased the next earn response requirement. Reinforcer accumulation is an understudied dimension of operant behavior that relates to the analysis of such phenomena as hoarding and self-control, in that they too involve accumulating versus immediately collecting or consuming reinforcers.     

Motivational properties of spaced food presentation in hungry rats: Long-lasting effects on drinking and eating

Hungry rats were given 60 food pellets either at spaced intervals or massed in one or two presentations, or no food, followed by a consumption test in the home cage. Drinking was higher after spaced food sessions than after massed food or no food sessions (Experiments 1, 2, 3, and 5) and higher after the presentation of standard food pellets than after quinine-flavored pellets (Experiment 2). An effect of spacing was found even with a delay of 1 hr between the end of the food session and the drinking test (Experiment 3). Spaced food presentation also produced an increase in the consumption of quinine-flavored food pellets after the end of the session (Experiment 4). These results suggest that the enhancement of behavior caused by intermittent reinforcement cannot be attributed entirely to the short-term eliciting effects of the reinforcer or to its thirst-inducing properties.     

Developmental expression of defensive responses during exposure to conspecific adults in preweanling rats (Rattus norvegicus).

I examined preweanling rats' (Rattus norvegicus) expression of ultrasounds and secretion of ACTH when exposed to unfamiliar adult male rats or to their mothers. Pups at 7 days of age produced similar levels of ultrasonic vocalization near both unfamiliar males and mothers. However, these pups could discriminate familiar from unfamiliar adults because ACTH was significantly higher in pups near adult males than in those near mothers. At 14 days of age, pups avoided adult males but not their mothers; therefore, adult males represented a significant threat. Importantly, 14-day-old rats significantly reduced ultrasound production only when near adult males. Pups at 21 days of age no longer emitted ultrasounds when socially isolated or when near conspecific adults. In addition, 14- and 21-day-old rats produced similar elevated ACTH levels across stimulus conditions. Results show significant changes in preweanling rats' responses to conspecific adults.     

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.     

Choice between delayed reinforcers in an adjusting-delay schedule: The effects of absolute reinforcer size and deprivation level

Choice between two reinforcers differing in magnitude and delay was investigated in rats using an adjusting-delay discrete-trials schedule in which the two reinforcers were associated with two levers (A and B). The delay to Reinforcer A (the smaller reinforcer) was always 2 sec, whereas the delay to Reinforcer B was varied in accordance with the distribution of choices in successive blocks of trials. In Experiment 1, the mean delay to the large reinforcer during the last 5 of 60 training sessions was greater when the rats were maintained at 80% than when they were maintained at 90% of their free-feeding body weights. In Experiment 2, the delay to the larger reinforcer was greater when the two reinforcers consisted of one and two 45-mg food pellets than when they consisted of three and six pellets. The results are consistent with a model of “self-control” which posits hyperbolic relations between reinforcer value and reinforcer magnitude, and between reinforcer value and delay of reinforcement.     

The effects of signalled and unsignalled lick-dependent delays on the development of schedule-induced drinking in rats

Food pellets were programmed to be delivered to rats every 60 sec (Fixed Time 60-sec schedule), and the development of schedule-induced drinking was measured in terms of the amount of water consumed and the number of licks per inter-pellet interval. For some rats (masters) 10-sec delays in food delivery were dependent on licks. Yoked-control rats received food at the same time as their masters and independently of their own behaviour. In Experiment 1, in which the delays were signalled by a blackout, the master rats began to drink, but this schedule-induced behaviour then decreased to levels lower than those shown by the yoked controls. When the signalled delays were discontinued, the drinking of the master rats recovered. In Experiment 2, in which the delays were not signalled, the master rats did not develop as much schedule-induced drinking as the yoked controls, and discontinuing the delays led to only small increases in drinking. These results support the view that schedule-induced drinking is subject to control by its consequences.     

Lean and mean rats: Some effects of acute changes in the food supply upon territorial aggression

In experiment 1, individually housed rats subjected to short-term food restriction displayed more territorial aggression toward conspecific intruders than controls maintained on a free-feeding diet. In experiment 2, small groups of three adult male rats had access to either a standard laboratory diet or the standard diet plus sucrose. Groups with the sucrose supplement were significantly less aggressive toward intruders than controls. Sucrose availability did not produce appreciable gains in body weight but it did reliably decrease within-colony weight variation. The results suggest the existence of an effective dietary mechanism that enables a social species such as Rattus norvegicus to tolerate each other in dense feeding aggregations when food is abundant. Conversely, when food is limited, social intolerance increases and serves to limit the development of large feeding groups.