The effect of food deprivation on self-control

Two experiments examined the effect of food deprivation on choice in a discrete-trials self-control paradigm, choice between a larger, more-delayed reinforcer and a smaller, less-delayed reinforcer. In Experiment 1, four pigeons were each deprived to 65%, 80%, and 90% of their free-feeding weights, and the delay to the smaller reinforcer was varied. Deprivation level did not affect choice, but the rate of ineffective key pecks made during the reinforcer delays increased as deprivation increased. In Experiment 2, four pigeons were exposed to conditions in which they were fed up to their 80% free-feeding weights following experimental sessions, and in which they were given no postsession feedings. Both the pigeons' weights and their latencies to insert their heads into the food hopper when food was available were lower when the pigeons were not fed following experimental sessions. Choice showed no change. Deprivation level affects response rate and eating behavior in these procedures, but not choice.     

The “lunching” effect: Pigeons track motion towards food more than motion away from it

Four experiments measured pigeons’ pecking at a small touch-screen image (the CS) that moved towards or away from a source of food (the US). The image's effectiveness as a CS was dependent on its motion, direction, and distance relative to the US. Pecking to the CS increased with proximity to the US when the CS moved towards the US (Experiment 1). This held true even when a departing CS signalled a US of greater magnitude (Experiment 2). Response rates to stationary stimuli were greater the closer they were to the hopper; but rate was less than when the same spot was part of a motion towards food, and greater than when it was part of a motion away (Experiment 3). The rate of responding in all three cases (motion towards, stationary, motion away) decreased exponentially with distance from the hopper. The distance and motion effects observed under these Pavlovian contingencies were different when pecking to the spot was required for reinforcement (Experiment 4).     

Interactions among unit-price, fixed-ratio value, and dosing regimen in determining effects of repeated cocaine administration

Previous research has shown tolerance to cocaine that was dependent on fixed-ratio (FR) parameter size in the context of a multiple FR schedule of food reinforcement. Completion of the FR requirement in these studies resulted in the same magnitude of reinforcement, regardless of ratio size. The cost-to-benefits ratio (unit-price) was therefore not equated across the different FR components. The current study examined the effects of repeated administration of cocaine to pigeons when unit-price under FR schedules was either the same or different. Additionally, the role of a chronic variable-dosing versus chronic fixed-dosing procedure was examined when unit-price was equated across different ratio values. Pigeons were trained to key peck in daily sessions under a three-component multiple FR schedule of food presentation, according to which either 10, 30, or 100 pecks were required for each delivery of food. In Experiment 1, completion of the FR 10, FR 30, and FR 100 resulted in 1.5, 4.5, and 15.0 s access to food, respectively. That is, the response requirement was correlated with access to food time so that unit-price (pecks per second of access to food) was equated across components. After assessing acute effects of a range of doses of cocaine, drug administration occurred daily before each session, with the dose varying from day to day. Tolerance, the magnitude of which was unrelated to the peck requirement, developed under the repeated-dosing regimen. In order to assess whether having equal unit-price was responsible for producing similar levels of tolerance across components, daily drug administration continued in Experiment 2 using the variable-dose regimen, but the amount of food presented each time was fixed at 4.5 s access to food, yielding different unit-prices under the three pecking requirements. Subsequently, the conditions of Experiment 1 were reinstated, i.e., unit-price was equated. Making unit-price different or the same had little influence on effects of cocaine. In Experiment 3, a fixed dose of cocaine was administered before each session while programmed unit-price remained the same across components. Under these conditions, tolerance became peck-requirement related. Specifically, tolerance was most prevalent under the smaller requirements and less robust or absent when the largest requirement was in effect. Differences in unit-price, therefore, were not related to degree of tolerance, but work requirement was. Differences in effects of cocaine across responses requirements, however, were observed only when each session was preceded by the same dose, not when dose varied from session to session.     

An Experimental Analysis of Spatial Position Effects on Foraging and Vigilance in Brown-Headed Cowbird flocks

Several observational studies have found that the costs and benefits of social foraging vary as a function of spatial position in the group. However, it is difficult to make mechanistic inferences because several confounding factors, such as food deprivation levels, food availability, neighbor distance, and group size can mask or amplify spatial position effects. We attempted to address experimentally the effect of spatial position on foraging and vigilance in a group, controlling for many confounding factors. We used enclosures that restricted physical but not visual interactions between brown-headed cowbirds and manipulated spatial position, flock size, and neighbor distance. Pecking rate (number of pecks per trial duration) was not related with position, but instantaneous pecking rate (number of pecks per foraging bout duration) was higher at the edge. The proportion of time spent head-up (scanning and food-handling) was also higher at the edge. For pecking rate and proportion of time spent scanning, changes in neighbor distance influenced the behavior of edge birds to a lesser extent than central birds. These results suggest that cowbirds at the edge perceived greater predation risk and that during the limited foraging time available, edge birds tried to compensate by foraging at a faster rate.     

Courtship as a reinforcement for key pecking in the pigeon, Columba livia

This report describes a within-subjects comparison of the reinforcing effects of food and display on a single response, key pecking. Three male pigeons socially isolated and food deprived for 22 hr pecked a key-light when pecks resulted in a brief sighting of their respective mates. Pecks to a second key-light concurrently available resulted in brief food rewards. Responses to the female key had a greater latency than responses to the food key, irrespective of the nature of the preceding reinforcement. The two responses also differed in their form and in the extent to which they were accompanied by other behaviours performed around the key-lights. This is discussed in terms of both response-reinforcer interaction and the intrusion of Pavlovian conditioning effects.     

How does food distance influence foraging in the ant Lasius niger: the importance of home-range marking

We study the influence of food distance on the individual foraging behaviour of Lasius niger scouts and we investigate which cue they use to assess their distance from the nest and accordingly tune their recruiting behaviour. Globally, the number of U-turns made by scouts increases with distance resulting in longer travel times and duration of the foraging cycle. However, over familiar areas, home-range marking reduces the frequency and thereby the impact of U-turns on foraging times leading to a quicker exploitation of food sources than over unmarked set-ups. Regarding information transfer, the intensity of the recruitment trail reaching the nest decreases with increasing food distance for all set-ups and is even more reduced in the absence of home-range marking. Hence, the probability of a scout continuing to lay a trail changes along the homeward journey but in a different way according to home-range marking. Over unexplored setups, at a given distance from the food source, the percentage of returning trail-laying ants remains unchanged for all tested nest-feeder distances. Hence, the tuning of the trail recruiting signal by scouts was not influenced by an odometric estimate of the distance already travelled by the ants during their outward journey to the food. By contrast, over previously explored set-ups, a distance-related factor – that is the intensity of home-range marking – strongly influences their recruiting behaviour. In fact, over a home-range marked bridge, the probability of returning ants maintaining their trail-laying behaviour increases with decreasing food distance while the gradient of home-range marks even induces ants which have stopped laying a trail to resume this behaviour in the nest vicinity. We suggest that home-range marking laid passively by walking ants is a relevant cue for scouts to indirectly assess distance from the nest but also local activity level or foraging risks in order to adaptively tune trail recruitment and colony foraging dynamics. Received 13 July 2004; revised 26 January and 20 May 2005; accepted 2 July 2005.     

How do pigeons represent numbers?: Studies of number scale bisection

Pigeons' bisection of two number scales, 1-16 and 2-32, was examined in two experiments. In Experiment 1, pigeons learned to choose different comparison keys following 1 or 16 key pecks and 2 or 32 key pecks and then were tested with intermediate numbers of key pecks on nonreinforced probe trials. Psychophysical curves that plotted choice of the large number key against number of pecks showed superposition of the two number scales and indifference between the two keys at the geometric means of each scale. The same procedures were used in Experiment 2, but pigeons were trained to bisect each scale at its arithmetic mean. Two asymmetries appeared in curves that plotted discrimination performance against number: near the midpoints of the scales, accuracy was higher for high numbers than for low numbers, but this relationship reversed at the ends of the scales, with low numbers discriminated more accurately than high numbers. An associative model that assumed stimulus generalization between numbers was used to predict the findings of these two experiments. The model showed a good qualitative fit to the obtained data when it was assumed that number was represented on a log scale and generalization was constant at all numbers but not when it was assumed that number was represented on a linear scale and generalization was scalar.     

Response strategies during acquisition of serial learning in pigeons (Columba livia)

Pigeons were exposed on fixed duration sessions to four chromatic stimuli and trained to perform a 2-item sequence. A detailed analysis of incorrect responses performed on each trial prior to completion of the sequence led to distinguish three learning stages. Already on initial sessions animals responded significantly more frequently to relevant stimuli than to irrelevant ones indicating stimulus control by salience. During an intermediary stage, responding to item 2 increased significantly and animals started an increasing number of trials with item 2, suggesting stimulus control by recency. On the final session the correct sequence outnumbered significantly the reverse sequence, suggesting control by order, while recency and salience effects were present to a lesser extent. Repeated pecks, i.e. consecutive pecks on the same cue, occurred on the four cues on the first session and decreased significantly on the last session for irrelevant colours. In addition, when key pecks were followed by a brief sound signal, repeats on relevant stimuli decreased while efficient colour pecks (i.e. when repeats were ignored) on these stimuli increased. These findings are discussed in the light of those obtained in traditional chaining paradigms.     

Trophallaxis in honeybees, Apis mellifera (L.), as related to their past experience at the food source

Forager honeybees returning to the hive after a successful foraging trip unload the collected liquid to recipient hivemates through mouth-to-mouth food exchange contacts (trophallaxis). The speed at which the liquid is transferred (unloading rate) from donor to recipient is related to the profitability offered by the recently visited food source. However, because a forager's evaluation of the profitability of a food source, as measured by dancing behaviour, is influenced by previous foraging experience, we investigated whether trophallaxis might also be influenced by previous foraging experience. We measured unloading rate for a given profitability condition at the food source (in terms of solution flow rate) in three groups of foragers that differed in their previous experience at the source. One group experienced the same flow rate of solution in five successive visits (control group), another group experienced a lower flow rate in the first four visits and the third group experienced a higher flow rate in the first four visits. The results of the present study show that animals trained to a lower flow rate increased their unloading rate compared with the control group, indicating an influence of past foraging experience on their evaluation of food source profitability. This influence was not observed in the group trained to a higher flow rate, which responded as the control group. Additional experiments indicated that foragers appear to evaluate the profitability of the source by integrating an overall flow rate throughout the entire visit, instead of measuring only the current flow rate delivered by the feeder. Copyright 2003 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.      

Foragers of the stingless bee Plebeia droryana inform nestmates about the direction,but not the distance to food sources

1. The tropical stingless bees have evolved intricate communication systems to recruit nestmates to food locations. Some species are able to accurately communicate the location of food, whereas others simply announce the presence of food in the environment.
2. Plebeia droryana is a tiny Neotropical stingless bee that, until recently, was thought to use a solitary foraging strategy, that is without the use of a recruitment communication system. However, recent research has indicated that P. droryana might be able to recruit nestmates to specific food source locations.
3. We tested this by studying whether foragers can guide nestmates in the direction and the distance of artificial feeders placed in the vicinity of the colony. We trained bees to a scented sucrose solution feeder at 10 m and placed different feeders either in different directions (experiment 1) or in different distances (experiment 2). We found that P. droryana directs newcomers in the right direction, but distance information does not seem to be communicated.
4. Moreover, we then tested whether newcomers use chemical and visual cues originating from nestmates foraging at the food source, but found no evidence for the use of these social cues provided by conspecifics.
5. The potential mechanism that P. droryana may use to orient recruits toward the food source, however, remains unknown and requires further study.

     

The influence of the food size,distance and food site on food carrying behavior in rats (Rattus norvegicus)

The behavior to foods in rats (Rattus norvegicus) was investigated in a seminatural enclosure and in a straight alley in the laboratory. In both situations, the size of food (0.045-, 1-, and 3-g pellets were used in the enclosure and 0.045-, 0.2-, 1-, and 3-g pellets in the alley) and distance between food sites and nest site (1.8, 3.6, 5.4, and 7.2 m in the enclosure and 2, 4, and 6 m in the alley) were varied. In both situations, when food was presented in only 1 place, the food carrying behavior increased as the size of food increased from 0.045 to 1-g. One and three g pellets were always carried, regardless of the distance to nest. On the other hand, when multiple food sites were presented in the alley, rats were more likely to carry small pellet from the further food site, whereas large pellet was always carried and were not influenced by the distance. The behavior to foods in the enclosure, however, was not influenced in either food site condition. This might be due to the difference in familiarity to the environment and anxiety from perceived predation risk.     

Leptin inhibits food-deprivation-induced increases in food intake and food hoarding

Food deprivation stimulates foraging and hoarding and to a much lesser extent, food intake in Siberian hamsters. Leptin, the anorexigenic hormone secreted primarily from adipocytes, may act in the periphery, the brain, or both to inhibit these ingestive behaviors. Therefore, we tested whether leptin given either intracerebroventricularly or intraperitoneally, would block food deprivation-induced increases in food hoarding, foraging, and intake in animals with differing foraging requirements. Hamsters were trained in a running wheel-based food delivery foraging system coupled with simulated burrow housing. We determined the effects of food deprivation and several peripheral doses of leptin on plasma leptin concentrations. Hamsters were then food deprived for 48 h and given leptin (0, 10, 40, or 80 microg ip), and additional hamsters were food deprived for 48 h and given leptin (0, 1.25, 2.5, or 5.0 microg icv). Foraging, food intake, and hoarding were measured postinjection. Food deprivation stimulated food hoarding to a greater degree and duration than food intake. In animals with a foraging requirement, intracerebroventricular leptin almost completely blocked food deprivation-induced increased food hoarding and intake, but increased foraging. Peripheral leptin treatment was most effective in a sedentary control group, completely inhibiting food deprivation-induced increased food hoarding and intake at the two highest doses, and did not affect foraging at any dose. Thus, the ability of leptin to inhibit food deprivation-induced increases in ingestive behaviors differs based on foraging effort (energy expenditure) and the route of administration of leptin administration.     

A further study of choice and percentage reinforcement

In Experiment I four pigeons were trained in a concurrent chains procedure with fixed-ratio schedules (FR1) in the initial components and fixed-time schedules in the terminal components. Pecking one of the keys when both keys were white initiated a fixed time schedule on that key. A peck to the left key produced three stripes on the key. At the termination of the fixed-time component food always occurred. Pecking the other key produced either a circle or a triangle. If a circle appeared, reinforcement occurred. If a triangle appeared a brief timeout was given. Initially the stripes appeared on the left key and the circle and triangle on the left. This was reversed during the course of the experiment. In addition, sessions were conducted in which both circle and triangle sometimes preceded reinforcement and sometimes timeout. For most birds under most conditions there was a preference for the key that produced the circle and triangle. When these were uncorrelated with reinforcement and time out three of the birds preferred the key producing 100% reinforcement.

In Experiment II three factors were varied and VI 20 sec schedules were used in the initial links instead of FR1. The results showed that pigeons preferred the 50% condition more 1) the greater the duration of the terminal links, 2) the smaller the value on the initial link VI schedules and 3) the less the probability of food in the terminal link with stripes on the key.     

Some characteristics of spatial associative memory in the pigeon, Columba livia

Willson and Wilkie (1993) developed a novel procedure to assess pigeons' memory for the spatial location of food. Only one of four locations provided food each daily session. Each location consisted of an illuminated pecking key and grain feeder. Over different days different locations, randomly selected, provided food during a 16-min session. The pigeons tended to revisit the location at which food was found on the previous day thereby demonstrating memory for food-spatial location associations over 24 h. Three experiments were conducted to further investigate this phenomenon. In Experiment 1 the session duration was varied between 4 and 32 min. Longer sessions had no detectable effect on their ability to remember the rewarded location 24 h later, a result that suggests that only brief encounters with food at a particular location are necessary for recall. In Experiment 2 the necessity of an active search for the day's rewarded location was removed; a 5-min period in which only the rewarded key was lit preceded the regular 16-min session. Pecks to the lit key in this 5-min period produced grain on the standard schedule. This manipulation facilitated the pigeons' discovery of food but did not affect their ability to remember the rewarded location, suggesting that the process of search and discovery is not essential to the associative memory process. In Experiment 3, food was available during the complete session (non-depleting condition) or was available only during the first half of the session (depleting condition). No detectable differences in the birds' memory of yesterday's profitable location were found. This suggests that non-depletion of food is not a necessary condition for day-to-day recall of food location. Taken together these findings enlarge our understanding of the spatial associative memory process.     

PYY(3-36) into the arcuate nucleus inhibits food deprivation-induced increases in food hoarding and intake

Central administration of neuropeptide Y (NPY) increases food intake in laboratory rats and mice, as well as food foraging and hoarding in Siberian hamsters. The NPY-Y1 and Y5 receptors (Rs) within the hypothalamus appear sufficient to account for these increases in ingestive behaviors. Stimulation of NPY-Y2Rs in the Arcuate nucleus (Arc) has an anorexigenic effect as shown by central or peripheral administration of its natural ligand peptide YY (3-36) and pharmacological NPY-Y2R antagonism by BIIE0246 increases food intake. Both effects on food intake by NPY-Y2R agonism and antagonism are relatively short-lived lasting ∼4 h. The role of NPY-Y2Rs in appetitive ingestive behaviors (food foraging/hoarding) is untested, however. Therefore, Siberians hamsters, a natural food hoarder, were housed in a semi-natural burrow/foraging system that had (a) foraging requirement (10 revolutions/pellet), no free food (true foraging group), (b) no running wheel access, free food (general malaise control) or (c) running wheel access, free food (exercise control). We microinjected BIIE0246 (antagonist) and PYY_(3-36) (agonist) into the Arc to test the role of NPY-Y2Rs there on ingestive behaviors. Food foraging, hoarding, and intake were not affected by Arc BIIE0246 microinjection in fed hamsters 1, 2, 4, and 24 h post injection. Stimulation of NPY-Y2Rs by PYY_(3-36) inhibited food intake at 0–1 and 1–2 h and food hoarding at 1–2 h without causing general malaise or affecting foraging. Collectively, these results implicate a sufficiency, but not necessity, of the Arc NPY-Y2R in the inhibition of food intake and food hoarding by Siberian hamsters.     

Attention, the presolution period, and choice accuracy in pigeons

Six pigeons were trained first on a color then on a form discrimination; four other pigeons were trained first on form and then on color. One of two colors or one of two forms (sample stimuli) appeared in the center of a touch sensitive monitor for 5 pigeons and in the center and in 16 other locations for 5 other pigeons. A peck anywhere within the region in which the sample stimuli appeared produced two white disks (comparison stimuli), one on the left and one on the lower right corners of the screen. Correct left-right choices provided food. Although of no consequence, the location of pecks in presence of the sample was predictive of the pigeon's subsequent choice. Accuracy, choice of the correct comparison stimulus, was greater when the sample stimuli appeared in the center as well as 16 other locations than when it appeared only in the center. The presolution period, the period of chance accuracy prior to evidence of discrimination learning, was decreased on each task following training on the other task. This evidence of facilitation following an extra-dimensional shift was attributed to continued relevance of the conditions under which the first task was learned. The duration of the presolution period was inversely related to asymptotic accuracy-data accounted for by Heinemann's (1983) theory of information processing during the presolution period.     

A direct effect of competition on food choice by pigeons

Previous research on food choice of pigeons foraging alone and in competition showed an indirect response to the competitor, mediated by the resource depletion associated with the competitor. This experiment showed, in addition, that pigeons can adjust their food choice in direct response to the competitor itself. Pigeons foraged for large, preferred grains of maize and smaller, unpreferred grains of wheat presented in bowls covered with sawdust. In a within-subject design, pigeons were tested alone or in competition with another pigeon. In competition, a higher choice proportion of the wheat grains was observed on the first two choices out of 48 items, and this effect increased with time in the experiment. This result underscores the role of learning in group foraging behaviour. It also suggests a possible ecological implication: that individuals that learn faster may enjoy a competitive advantage in terms of reduced delay in responding to the presence of a conspecific.     

Pigeons do not perceptually complete partly occluded photos of food: an ecological approach to the "pigeon problem"

Humans routinely complete partly occluded objects to recognize the whole objects. However, a number of studies using geometrical figures and even conspecific images have shown that pigeons fail to do so. In the present study, we tested whether pigeons complete partially occluded objects in a situation simulating a natural feeding context. In Experiment 1, we trained pigeons to peck at any photograph of food and not to peck at any containing a non-food object. At test, we presented both photos of food partly occluded by pigeon's feather and photos simply truncated at the same part. We predicted that if the pigeons perceptually completed the occluded portion, then they would discriminate the photos of occluded food better than the truncated photos. The result was that the pigeons pecked at the truncated photos earlier than the occluded photos. Placing the occluder next to all of the stimuli in Experiment 2 or substituting indented lozenge for the feather in Experiment 3 did not affect the results. Thus, even in a simulated ecologically significant situation, pigeons continued to not show evidence of perceptual completion.     

Characterizing the effects of d-amphetamine on temporal discrimination

Although the effects of dopamine agonists on temporal discrimination have been widely studied, conclusions as to their behavioral and neurophysiological effects are difficult due to a number of discrepant findings in the literature. This study examined whether a previously unexplored procedural difference could account for some of these discrepancies. In three experiments, pigeons categorized the duration of temporal samples during two variants of the interval-bisection procedure. In the position variant, responses to a side key that corresponded to the sample duration produced food. In the color-matching variant, responses to the key color that corresponded to the sample duration produced food. Across experiments, d-amphetamine produced a general disruption of temporal discrimination not accompanied by over or underestimation of time in both procedures. This effect occurred whether pigeons were exposed to both procedures within session (Experiment 1) or across conditions (Experiments 2 and 3) and whether the pigeons were drug experienced (Experiments 1 and 2) or naïve (Experiment 3). Analyses of the cumulative normal functions fitted to the proportion-long response data indicated that d-amphetamine produced its effects by selectively decreasing stimulus control, rather than by affecting timing.     

Reaction times identify a Pavlovian component in a two-choice discrimination

Six pigeons discriminated on discrete trials between two colors. In Experiment 1, two luminous spots were both either blue or green and the reinforced responses were “peck left” for blue and “peck right” for green. In Experiment 2, the hue of a center spot controlled subsequent choice pecks to left or right. In both experiments response bias was manipulated in two ways. During stimulus frequency (“SF”) sessions correct responses brought food on 40% of trials; in “imbalanced” blocks of sessions one hue appeared on 80% of trials and the other on 20%. During reinforcement probability (“RNF”) sessions the hues appeared equally often, but in imbalanced blocks the hues signaled different reinforcement probabilities, either 64% or 16%. In “balanced” control blocks the hues appeared equally often and were both reinforced at 40%. The experiments gave similar results. When bias was computed from choice percentages the imbalanced conditions yielded substantial response bias, and the amount of bias was about the same under RNF and SF treatments. However, reaction times (RTs) gave a different outcome. RNF imbalance slowed responses directed at the less reinforced stimulus, but SF imbalance had little RT effect (Experiment 1) or no effect (Experiment 2). These results suggest that choice was controlled by an instrumental stimulus-response-reinforcement association, whereas RTs were controlled by a Pavlovian stimulus-reinforcement association.