Evidence that appetitive responses for dehydration and food-deprivation are learned.

Rats do not seek water when cellularly dehydrated until they are about 4 weeks of age. This lack of appetitive 'seeking' behavior in young rats differs from their precocious ingestive responses such as an increased intake of solutions infused into their mouths when they are dehydrated as young as 2 days of age. Using video analysis of appetitive behavior in a structured environment, we document this early absence of appetitive responding and the subsequent acquisition of dehydration-elicited appetitive behavior. Weaning age pups were separated into four conditions: (i) experienced, dehydrated; (ii) experienced, nondehydrated; (iii) inexperienced, dehydrated; and (iv) inexperienced, nondehydrated. 'Experienced' rats received a dehydration and drinking experience prior to the test, and 'dehydrated' rats were dehydrated (by injection of a salt load) at the time of test. At the test, all water and food was removed from the test cages, eliminating the confounding of appetitive and consummatory measures. Despite the fact that pups in all conditions had experience with water and had previously drunk, only the 'experienced' pups differentially sought water when dehydrated. Parallel experiments with food deprivation produced similar results. Pups did not exhibit food-seeking behavior when food-deprived unless they had previous experience with food deprivation and eating. The appetitive 'seeking' behavior for feeding also appears to be learned. Directed appetitive behavior in general may thus be acquired.

Effects of prior experience with dehydration and water on the time course of dehydration-induced drinking in weanling rats.

Although cellular dehydration increases oral responding and swallowing of orally infused water in rats as young as 2 days old, it is not until well after the time of weaning that dehydration stimulates immediate water-seeking and initiation of drinking in situations where the water source must be approached voluntarily. Recent work has shown that the goal-directed appetitive sequence for drinking-orienting, approaching, and initiating contact with water-matures much later than the more precocial oral licking and swallowing behaviors, and normally comes to be elicited by dehydration only after post-weaning experience with dry food. In the current experiments we evaluate some critical features of post-weaning experience with dehydration and drinking, and find that prior experience with initiating drinking while dehydrated, but not experience with dehydration nor water per se, alters the time course of water intake during a subsequent hydrational challenge. The effects of experience are manifested as an increased proportion of water consumed in the early portion of the test, rather than a general increase in total consumption. These findings are consistent with the interpretation that prior experience is necessary for the coordination of water-oriented appetitive behaviors that lead to the initiation and maintenance of drinking bouts, and provide further evidence for an associative learning account of the acquisition of dehydration-induced drinking.

Thirst modulates a perception.

Does thirst make you more likely to think you see water? Tales of thirsty desert travelers and oasis mirages are consistent with our intuitions that appetitive state can influence what we see in the world. Yet there has been surprisingly little scrutiny of this appetitive modulation of perception. We tested whether dehydrated subjects would be biased towards perceptions of transparency, a common property of water. We found that thirsty subjects have a greater tendency to perceive transparency in ambiguous stimuli, revealing an ecologically appropriate modulation of the visual system by a basic appetitive motive.

Olfactory conditioning facilitates diet transition in human infants.

We evaluated whether Pavlovian conditioning methods could be used to increase the ingestion of non-preferred solutions by formula-fed human infants. In baseline measures, 5-7 month old infants sucked less frequently and consumed less water than regular formula. During a 3-day olfactory conditioning period, parents placed a small scented disk, the conditioned stimulus, on the rim of their infants' formula bottle at every feeding. Following this training, infants' responses to water were tested when their water bottles had a disk scented with the training odor, a novel odor, or no odor. Infants tested with the training odor sucked more frequently and consumed significantly more water than they had at baseline. Infants tested with no odor or a novel odor consumed water at or below baseline levels. These data demonstrate that olfactory conditioning can be used to enhance ingestion in infants and suggest that such methods may be useful for infants experiencing difficulty when making transitions from one diet to another.

Conditioned changes in appetitive and consummatory responses to flavors paired with oral or nutrient reinforcement among adult rats.

Recent studies of the behavioral organization of conditioned flavor preferences have suggested the involvement of at least two separate learning systems-an appetitive response system sensitive to the oral hedonic properties of the reinforcer, and a consummatory response system sensitive to its nutrient properties. However, these prior studies were conducted with weanling rats, that differ from adults in terms of their prior experience with food, their learning competencies, and the peculiar ontogenetic constraints on their behavior. It is, therefore, unknown whether flavor preference behaviors are similarly organized in adult rats. In this experiment, adult rats were trained to associate a specific CS flavor with either the sweet taste or the postingestive nutrient effects of sucrose. Conditioned appetitive orienting and consummatory oral responding to the CS flavors were then measured. Unlike weanling rats, adult rats exhibited both conditioned appetitive behavior and conditioned consummatory behavior in response a CS that was previously paired with either oral hedonic or nutrient reinforcement. These results suggest a set of important developmental changes in the neurobehavioral mechanisms of flavor preference learning in the postweaning period.

The acquisition of an appetite.

Unlike older animals, weanling-age rats do not seek water to drink when they are dehydrated, despite the fact that a physiological sensitivity to dehydration is present very soon after birth. We demonstrate here that the appetitive behaviors needed to approach and obtain water become linked to dehydration only as a result of specific postnatal learning experience. Preventing early experience with dehydration retards the developmental emergence of dehydration-induced, water-oriented behavior in young rats. But a single pairing of water with dehydration can establish an appetitive response. These findings reveal a critical role of early learning in the development of goal-oriented behavior. Such a learning process is potentially characteristic of other behavioral systems, from the most basic appetites to complex motives.

Evidence that oral and nutrient reinforcers differentially condition appetitive and consummatory responses to flavors.

Rats tend to increase their intake of a flavor that has previously been paired with either sweet taste or with caloric repletion. However, it is unclear whether such a change in intake is caused by changes in appetitive behaviors such as orienting and approach, or changes in consummatory behaviors and oral responsiveness. Also, it is unclear whether oral reinforcers (sweetness) and postingestive reinforcers (nutrients) lead to the same kinds of behavioral change. In the current experiments, weanling rats with oral and gastric cannulas repeatedly experienced a flavor paired with either sweetness, high caloric density, or neither. Rats were then tested for differences in appetitive olfactory orienting and consummatory oral responsiveness elicited by the flavor. Results suggest that oral reinforcement (sweetness) produces conditioning of appetitive responding to the flavor, while postingestive reinforcement produces conditioning of consummatory responding. A second experiment indicates that these behavioral changes are specific increases in responsiveness conditioned by flavor + unconditioned stimulus (US) pairing, and are unlikely to be nonspecific effects of daily unconditioned stimulus exposure.

Electromyographic analysis of oral habituation in rat pups.

Rat pups show decreases in mouthing activity in response to a series of repeated oral infusions of a diet. This decrease in mouthing activity has been termed "oral habituation" and these changes have been readily recorded with simple behavioral observations. Oral habituation appears to be a component of satiety in young rats. In the present study, to more specifically characterize changes in motor response topography during habituation in muscle groups used for mastication, mouthing activity was recorded by implanting fine wire electromyographic electrodes in the superficial masseter, anterior digastric, sternohyoideus, and genioglossus muscles of 12-day-old rat pups. During testing, pups received a series of brief oral infusions of a 10% sucrose diet delivered through an oral cannula. The results demonstrated that mouthing activity as observed and scored behaviorally was highly correlated with mouthing behavior recorded by EMG, with oral habituation distinctly emerging in both measures. In addition, the pattern of motor activity in the four masticatory muscles changed during the course of oral habituation. Within the minute following a single infusion, the cycle frequency, duration of activity, and relative onset time of activity in the four muscles changed. In addition, across the course of habituation, both cycle frequency and relative onset times of muscle activity changed. These results demonstrate the general reliability of behavioral observations of masticatory motor activity in young rats and provide further information on how the pattern of activity of muscles involved in the mouthing motor pattern is altered during the course of oral habituation.

Sensitization of ingestive responding in preweanling rats: time course and specificity.

Two experiments measured the response-enhancing effects of oral stimulation in young rats with no feeding experience. In Experiment 1, pups were implanted with oral cannulas and deprived for 0, 6, or 24 hr. In nondeprived rats, sucrose exposure transiently sensitized oral responding to subsequent stimuli for several minutes. Among 24-hr-deprived rats, baseline oral responding had increased to where sucrose exposure was no longer sensitizing. Unlike oral responding, sensitization of activity by sucrose exposure increased with deprivation. Sensitization was also evident in the microstructure of responses to infusions. In Experiment 2, the specificity of sensitization was examined by varying stimulus palatability. Sucrose exposure sensitized oral responding to both sucrose and quinine-adulterated sucrose, but quinine-adulterated sucrose was not sensitizing. The results suggest that in inexperienced rats, oral stimulation initiates changes in responding that may mimic the endogenous sequelae of deprivation, perhaps representing early substrates for learned controls of intake.

Unilateral conditioning of an odor aversion in 6-day-old rat pups.

In young rats, several forms of olfactory memory can be functionally localized to one side of the brain by restricting training stimuli to one naris and corresponding olfactory bulb. In the experiment, the analysis of lateralized olfactory learning in 6-day-old rat pups was extended by an evaluation of the consequences of aversive reinforcers in a unilateral olfactory conditioning paradigm. An olfactory aversion was conditioned by delivering a mild footshock to animals in the presence of a novel odor. Olfactory stimulation was confined to one naris and corresponding olfactory bulb by inserting a soft rubber plug into the opposite naris. This lateralization in processing resulted in unilateral memory for the odor aversion that was only expressed when the trained naris was open during an odor preference test. The hypothesis that the hedonic value (or valence) component of conditioning is represented in structures that are unilaterally accessed during training and testing is discussed.

Evaluation of a distance consulting service based on interactive video and integrated computerized technology.

Telemedicine equipment was installed at a rural site in Drumheller and at Calgary, 85 miles (136 km) away. It allowed consultation between health-care providers at Drumheller and specialists and subspecialists at the Faculty of Medicine in Calgary. The efficacy of the system in providing more equitable access to health care for persons living in geographically remote sites was evaluated during a 12-month pilot project. Seventy-five encounters were attempted during the study. A total of 55 encounters (42 clinical and 13 non-clinical) were completed. The completed clinical encounters were distributed across a wide spectrum of medical specialties. Users of the system reported favourably on the impact of the telemedicine system on access to health services for rural patients, on diagnostic, investigative and management decisions, on patient and physician travel times, on feelings of professional isolation and educational opportunities and on overall patient health status. The study provides new knowledge and demonstrates the success of the technology in this project. As expected, other problems were raised and addressed in a preliminary manner including: the potential for health-provider education; acquisition and retention of rural physicians; ethical, legal and patient confidentiality issues; minimum acceptable technology; and network management issues.

The use of advanced computer technology to enhance access to health care and to respond to community needs: the results of the evaluation of a technology-based clinical consultation service.

This paper describes a long distance, high tech, two-way interactive video pilot system that supports teleconsultations between primary health care givers at a rural site and specialists and subspecialists at the Faculty of Medicine, University of Calgary. Evaluation results from the 12-month trial are presented. The underlying philosophy behind the telemedicine outreach pilot program was to provide equal access to health services for those currently in under-served areas. The advanced technology system offered multiple clinical services in emergency, urgent, and elective situations. The impact of the telemedicine consultative service on health care delivery, patients, providers, and costs were evaluated. Findings from the 12-month trial demonstrate that quality care and access to care can be offered in spite of the disparate location of patients and health care providers. Some limitations and constraints to the implementation of the technology are noted and reviewed.

Does oral experience terminate ingestion?

Using data from studies of ingestive behavior in developing rat pups we demonstrate how oral experience can contribute to the termination of ingestion. In rat pups, repeated oral stimulation with sweet solutions causes a decline in oral responsiveness. The diminished responsiveness is specific to the flavor of the stimulus experienced orally and can persist for several hours. We suggest that this experience-based decrement in responsiveness is best considered "oral habituation" and that oral habituation largely accounts for the onset of satiety. Post-ingestive feedback signals may have their influence through the oral habituation process or act in the context of oral habituation. Oral habituation is also shown to depend on the pattern of stimulus presentation, a phenomenon that adds considerable complexity to assessing the contributions of oral experience to satiety. The concept of oral habituation may be useful in understanding the immediate control of ingestion and the moment-to-moment expression of ingestive behavior in adult animals.

Integration of oral habituation and gastric signals in decerebrate rat pups.

During a series of oral infusions of a sweet solution, the ingestive responses of young rat pups habituate; pups stop responding to the infusions even when their stomachs are empty and the infused diet is nonnutritive. The rate of this oral habituation is enhanced by the addition of gastric fill signals, even in decerebrate pups. In intact but not in decerebrate pups, prior deprivation gates out the influence of gastric fill on habituation. This oral habituation system, responsive to multiple ingestion-related signals, may serve as the elemental process that integrates physiological state with ongoing behavior to control ingestion.

Taste and smell sensations enhance the satiating effect of both a high-carbohydrate and a high-fat meal in humans.

The effects of meal sensory properties (tasty vs. bland) and nutrient composition [high-CHO (carbohydrate) vs. high-FAT] on hunger ratings, blood glucose and free fatty acids (FFA), taste perception, and subsequent food intake, were studied in human subjects. Aspartame and vanilla were used to augment meal palatability, yielding four isocaloric liquid meals: bland-FAT, tasty-FAT, bland-CHO, tasty-CHO. Normal-weight, nondieting young adults consumed each of the meals for breakfast on separate days. The main finding was that tasty versions of high-FAT and high-CHO meals were more satiating than nutritionally identical bland meals, as indicated by a greater decrease in hunger ratings following the tasty meals. Changes in blood glucose and FFA were related to meal nutrient composition, but not to meal sensory properties. High-CHO meals tended to be more satiating than high-FAT meals. Consumption of each of the meals produced a similar decrease in pleasantness ratings of food-related tastes. Intake of carbohydrates was significantly higher at a self-selected lunch 5.25 h following a tasty breakfast. These findings indicate that hunger is decreased to a greater extent by meals flavored with aspartame and vanilla relative to nutritionally identical, unflavored meals. The satiety-enhancing effect of oral stimulation was found for both high-FAT and high-CHO meals.

Control of ingestion by oral habituation in rat pups.

The role of habituation of mouthing activity in the control of ingestion was investigated in 6-, 12-, and 18-day-old rat pups. In pups at all ages, oral habituation to a flavored diet inhibited ingestion of a continuous oral infusion of that same diet. Twelve-day-old pups that had orally habituated to a diet continued to consume less of a continuous oral infusion of that diet both 30 min and 3 hr later, and the duration of suppressed ingestion was shown to be dependent on the rate of stimulus presentation during habituation experience. These data suggest that oral habituation may be a diet-specific influence on both intra- and intermeal patterning.