Sedentary antlion larvae (Neuroptera: Myrmeleontidae) use vibrational cues to modify their foraging strategies

Learning abilities are exhibited by many animals, including insects. However, sedentary species are typically believed to have low capacities and requirements for learning. Despite this view, recent studies show that even such inconspicuous organisms as larval antlions, which employ an ambush predation strategy, are capable of learning, although their learning abilities are rather simple, i.e., limited to the association of vibrational cues with the arrival of prey. This study demonstrates, for the first time, that antlion larvae can use vibrational cues for complex modifications of their foraging strategies. Specifically, antlion larvae rapidly learn to differentiate between the vibrational cues associated with prey of different sizes, and they save resources by ignoring smaller prey in favour of larger, more energetically profitable prey. Moreover, antlion larvae can learn to associate vibrational cues with the loss of their prey, and they respond by burying their victims under the sand more often and more rapidly than do individuals with no opportunities to form such associations. These findings provide not only new insights into the cognitive abilities of animals but also support for the optimal foraging strategy concept, suggesting the importance of maximizing fitness output by balancing the costs and benefits of alternative foraging strategies.     

Friend or foe? The role of latent inhibition in predator and non-predator labelling by coral reef fishes

In communities of high biodiversity, the ability to distinguish predators from non-predators is crucial for prey success. Learning often plays a vital role in the ability to distinguish species that are threatening from those that are not. Many prey animals learn to recognise predators based on a single conditioning event whereby they are exposed to the unknown predator at the same time as alarm cues released from injured conspecifics. The remarkable efficiency of such learning means that recognition mistakes may occur if prey inadvertently learn that a species is a predator when it is not. Latent inhibition is a means by which prey that are pre-exposed to an unknown species in the absence of negative reinforcement can learn that the unknown animal is likely not a threat. Learning through latent inhibition should be conservative because mistakenly identifying predators as non-predators can have fatal consequences. In this study, we demonstrated that a common coral reef fish, lemon damselfish, Pomacentrus moluccensis can learn to recognise a predator as non-threatening through latent inhibition. Furthermore, we showed that we could override the latent inhibition effect by conditioning the prey to recognise the predator numerous times. Our results highlight the ability of prey fish to continually update the information regarding the threat posed by other fishes in their vicinity.     

Successive olfactory reversal learning in honeybees

Honeybees Apis mellifera can associate an originally neutral odor with a reinforcement of sucrose solution. Forward pairings of odor and reinforcement enable the odor to release the proboscis extension reflex in consecutive tests. Bees can also be conditioned differentially: They learn to respond to a reinforced odor and not to a nonreinforced one. They can also learn to reverse their choice. Here we ask whether honeybees can learn successive olfactory differential conditioning tasks involving different overlapping pairs of odors. The conditioning schedules were established in order to train the animals with 3, 2, 1, or 0 reversals previous to a last differential conditioning phase in which two additional reversals were present. We studied whether or not successive reversal learning is possible and whether or not learning olfactory discrimination reversals affects the solving of subsequent discrimination reversals. Therefore we compared the responses of bees that had experienced reversals with those of bees that had not experienced such reversals when both are confronted with a new reversal situation. In experiment 1 we showed that bees that had experienced three previous reversals were better in solving the final reversal task than bees with no previous reversal experience. In experiment 2, we showed that one reversal learning is enough for bees to perform better in the final reversal task. The successive different reversals trained in our experiments resemble the natural foraging situation in which a honeybee forager has to switch successively from an initial floral species to different ones. The fact that experiencing such changes seems to improve a bee's performance in dealing with further new exploited food sources has therefore an adaptive impact for the individual and for the colony as a whole.     

Children's acquisition and reversal behavior in a probability learning situation as a function of programed instruction,internal-external control,and schedules of reinforcement

A three-choice, contingent task was used with 192 fifth and sixth graders in a 2 × 2 × 3 factorial design combining instruction (programed instruction on selected probability concepts vs no programed instruction), locus of control (internal vs external), and schedules of reinforcement (33, 66, and 100%). The dependent measures were the percentage of correct acquisition responses, of correct reversal responses, and of pattern responses, as well as posttests on probability concepts. The major findings of the study were associated with schedule of reinforcement. In acquisition and reversal, Ss under 100% reinforcement during acquisition tended to maximize the greatest, followed by the 66 and 33% conditions, in that order. The ordinal relationship among schedules was the exact reverse of the maximizing approach for the pattern responses. A partial reinforcement effect was obtained in reversal. Evidence indicated that programed instruction and locus of control affected maximizing behavior, patterning behavior, and resistance to extinction (though these variables did not interact with reinforcement schedule in the predicted direction). Finally the posttest data showed that instructed Ss did learn more relative to noninstructed Ss.     

Elasmobranch cognitive ability: using electroreceptive foraging behaviour to demonstrate learning, habituation and memory in a benthic shark

Top predators inhabiting a dynamic environment, such as coastal waters, should theoretically possess sufficient cognitive ability to allow successful foraging despite unpredictable sensory stimuli. The cognition-related hunting abilities of marine mammals have been widely demonstrated. Having been historically underestimated, teleost cognitive abilities have also now been significantly demonstrated. Conversely, the abilities of elasmobranchs have received little attention, despite many species possessing relatively large brains comparable to some mammals. The need to determine what, if any, cognitive ability these globally distributed, apex predators are endowed with has been highlighted recently by questions arising from environmental assessments, specifically whether they are able to learn to distinguish between anthropogenic electric fields and prey bioelectric fields. We therefore used electroreceptive foraging behaviour in a model species, Scyliorhinus canicula (small-spotted catshark), to determine cognitive ability by analysing whether elasmobranchs are able to learn to improve foraging efficiency and remember learned behavioural adaptations. Positive reinforcement, operant conditioning was used to study catshark foraging behaviour towards artificial, prey-type electric fields (Efields). Catsharks rewarded with food for responding to Efields throughout experimental weeks were compared with catsharks that were not rewarded for responding in order to assess behavioural adaptation via learning ability. Experiments were repeated after a 3-week interval with previously rewarded catsharks this time receiving no reward and vice versa to assess memory ability. Positive reinforcement markedly and rapidly altered catshark foraging behaviour. Rewarded catsharks exhibited significantly more interest in the electrical stimulus than unrewarded catsharks. Furthermore, they improved their foraging efficiency over time by learning to locate and bite the electrodes to gain food more quickly. In contrast, unrewarded catsharks showed some habituation, whereby their responses to the electrodes abated and eventually entirely ceased, though they generally showed no changes in most foraging parameters. Behavioural adaptations were not retained after the interval suggesting learned behaviour was not memorised beyond the interval. Sequences of individual catshark search paths clearly illustrated learning and habituation behavioural adaptation. This study demonstrated learning and habituation occurring after few foraging events and a memory window of between 12 h and 3 weeks. These cognitive abilities are discussed in relation to diet, habitat, ecology and anthropogenic Efield sources.     

An eight-alternative concurrent schedule: foraging in a radial maze.

In two experiments conducted in an eight-arm radial maze, food pellets were delivered when a photocell beam was broken at the end of each arm via a nose poke, according to either fixed-interval or random-interval schedules of reinforcement, with each arm providing a different frequency of reinforcement. The behavior of rats exposed to these procedures was well described by the generalized matching law; that is, the relationships between log behavior ratios and log pellet ratios were approximated by linear functions. The slopes of these log-log functions, an index of sensitivity to reinforcement frequency, were greatest for nose pokes, intermediate for time spent in an arm, and least for arm entries. Similar results were obtained with both fixed-interval and random-interval schedules. Addition of a 10-s changeover delay in both experiments eliminated the slope differentials between nose pokes and time spent in an arm by reducing the slopes of the nose-poke functions. These results suggest that different aspects of foraging may be differentially sensitive to reinforcement frequency. With concurrent fixed-interval schedules, the degree of temporal control exerted by individual fixed-interval schedules was directly related to reinforcement frequency.     

Learning about non-predators and safe places: the forgotten elements of risk assessment

A fundamental prerequisite for prey to avoid being captured is the ability to distinguish dangerous stimuli such as predators and risky habitats from non-dangerous stimuli such as non-predators and safe locations. Most research to date has focused on mechanisms allowing prey to learn to recognize risky stimuli. The paradox of learned predator recognition is that its remarkable efficiency leaves room for potentially costly mistakes if prey inadvertently learn to recognize non-predatory species as dangerous. Here, we pre-exposed embryonic woodfrogs, Rana sylvatica, to the odour of a tiger salamander, Ambystoma tigrinum, without risk reinforcement, and later try to teach the tadpoles to recognize the salamander, a red-bellied newt Cynops pyrrhogaster—a closely related amphibian, or a goldfish, Carassius auratus, as a predator. Tadpoles were then tested for their responses to salamander, newt or fish odour. Pre-exposure to salamander did not affect the ability of tadpoles to learn to recognize goldfish as a predator. However, the embryonic pre-exposure to salamanders inhibited the subsequent learning of salamanders as a potential predator, through a mechanism known as latent inhibition. The embryonic pre-exposure also prevented the learned recognition of novel newts, indicating complete generalization of non-predator recognition. This pattern does not match that of generalization of predator recognition, whereby species learning to recognize a novel predator do respond, but not as strongly, to novel species closely related to the known predator. The current paper discusses the costs of making recognition mistakes within the context of generalization of predators and dangerous habitats versus generalization of non-predators and safe habitats and highlights the asymmetry in which amphibians incorporate information related to safe versus risky cues in their decision-making. Mechanisms such as latent inhibition allow a variety of prey species to collect information about non-threatening stimuli, as early as during their embryonic development, and to use this information later in life to infer the danger level associated with the stimuli.     

Behavioral ephemera,difficult discriminations,and behavioral stability

Every species possesses abilities for successfully interacting with its environment. These result from phylogeny. In the laboratory, one may arrange artificial conditions that thwart an organism's abilities. The result may be a “phenomenon.” With sufficient training, however, the phenomenon may prove to be ephemeral, as the organism's basic abilities reassert themselves. Pigeons respond extremely well to differences and nondifferences in rate of obtaining food. This ability may be thwarted in a variety of ways, but the results tend to be ephemeral. An example is an experiment that pitted pigeons' preference for unimpeded responding against their ability to respond to food rate. In a concurrent-chains procedure, the terminal links were identical variable-interval schedules, but in one terminal link, every response produced a timeout. The duration of the timeout varied, and preference varied with it, but the relation vanished with training, in keeping with the equality of food rate across the 2 terminal links. Some other examples of “phenomena” that tend to disappear with sufficient training and sufficient variation in experimental parameters are behavioral contrast, conditioned reinforcement, resistance to extinction, and suboptimal choice. These “phenomena” depend on pigeons' failing to make difficult discriminations. They appear to be behavioral ephemera.     

Learning, memorizing and apparent forgetting of chemical cues from new predators by Iberian green frog tadpoles

Many antipredator adaptations are induced by the prey’s ability to recognize chemical cues from predators. However, predator recognition often requires learning by prey individuals. Iberian green frog tadpoles (Pelophylax perezi) have the ability to learn new potential predators. Here, we tested the memory capabilities of Iberian green frog tadpoles. We conditioned tadpoles with chemicals cues from a non-predatory fish in conjunction with conspecific alarm cues, and examined whether tadpoles retained their conditioned response (reduction of activity level). We found that conditioned tadpoles reduced their activity levels in subsequent exposures to the non-predatory fish cues alone. Tadpoles were able to remember this association and reduced movement rate at least for 9 days after. The ability to learn and memorize potential predators may be especially important for the survivorship of prey species that are likely to find a high variety of predators. However, after those 9 days, there was a lack of response to the non-predatory fish cues alone in the absence of reinforcement. This could be explained if tadpoles behave according to the threat-sensitive predator avoidance hypothesis, and the perceived risk to the learning cue diminished over time, or it could be due to an apparent forgetting process to avoid non-adaptative responses to chemical cues of non-dangerous species that were randomly paired with alarm cues. Thus, this study demonstrates that green frog tadpoles in the absence of reinforcement remember the chemical cues of a learned predator only for a limited time that may be adaptative in a threat-sensitive context.     

Noncontingent reinforcement, alternative reinforcement, and the matching law: a laboratory demonstration

Basic researchers, but not most applied researchers, have assumed that the behavior-decelerating effects of noncontingent reinforcement result at least partly from adventitious reinforcement of competing behaviors. The literature contains only sketchy evidence of these effects because few noncontingent reinforcement studies measure alternative behaviors. A laboratory model is presented in which concurrent schedules of contingent reinforcement were used to establish a "target" and an "alternative" behavior. Imposing noncontingent reinforcement decreased target behavior rates and increased alternative behavior rates, outcomes that were well described by the standard quantitative account of alternative reinforcement, the generalized matching law. These results suggest that adventitious reinforcement of alternative behaviors can occur during noncontingent reinforcement interventions, although the range of conditions under which this occurs remains to be determined in future studies. As an adjunct to applied studies, laboratory models permit easy measurement of alternative behaviors and parametric manipulations needed to answer many research questions.     

Variable-time reinforcement schedules in the treatment of socially maintained problem behavior

Noncontingent reinforcement (NCR) consists of delivering a reinforcer on a time-based schedule, independent of responding. Studies evaluating the effectiveness of NCR as treatment for problem behavior have used fixed-time (FT) schedules of reinforcement. In this study, the efficacy of NCR with variable-time (VT) schedules was evaluated by comparing the effects of VT and FT reinforcement schedules with 2 individuals who engaged in problem behavior maintained by positive reinforcement. Both FT and VT schedules were effective in reducing problem behavior. These findings suggest that VT schedules can be used to treat problem behavior maintained by social consequences.     

Choice, changeover, and travel

Since foraging in nature can be viewed as instrumental behavior, choice between sources of food, known as “patches,” can be viewed as choice between instrumental response alternatives. Whereas the travel required to change alternatives deters changeover in nature, the changeover delay (COD) usually deters changeover in the laboratory. In this experiment, pigeons were exposed to laboratory choice situations, concurrent variable-interval schedules, that were standard except for the introduction of a travel requirement for changeover. As the travel requirement increased, rate of changeover decreased and preference for a favored alternative strengthened. When the travel requirement was small, the relations between choice and relative reinforcement revealed the usual tendencies toward matching and undermatching. When the travel requirement was large, strong overmatching occurred. These results, together with those from experiments in which changeover was deterred by punishment or a fixed-ratio requirement, deviate from the matching law, even when a correction is made for cost of changeover. If one accepted an argument that the COD is analogous to travel, the results suggest that the norm in choice relations would be overmatching. This overmatching, however, might only be the sign of an underlying strategy approximating optimization.     

Win-shift and win-stay learning in the short-beaked echidna (<Emphasis Type="Italic">Tachyglossus aculeatus</Emphasis>)

Numerous previous investigators have explained species differences in spatial memory performance in terms of differences in foraging ecology. In three experiments we attempted to extend these findings by examining the extent to which the spatial memory performance of echidnas (or "spiny anteaters") can be understood in terms of the spatio-temporal distribution of their prey (ants and termites). This is a species and a foraging situation that have not been examined in this way before. Echidnas were better able to learn to avoid a previously rewarding location (to "win-shift") than to learn to return to a previously rewarding location (to "win-stay"), at short retention intervals, but were unable to learn either of these strategies at retention intervals of 90 min. The short retention interval results support the ecological hypothesis, but the long retention interval results do not. Electronic Publication     

Delay reduction and optimal foraging: variable-ratio search in a foraging analogue.

The present study investigated conditions under which the conditioned reinforcement principles of delay-reduction theory and views based on simple maximization of reinforcement rate make ordinally opposing predictions with respect to foraging-related choice behavior. The use of variable-ratio schedules in the choice phase also represents an extension of delay-reduction theory to schedules that may better mimic the effort involved in searching. Pigeons responded on modified concurrent-chains schedules in which equal variable-ratio schedules led to unequal variable-interval outcomes and unequal reinforcer amounts. All 4 subjects completed a minimum of two replications of conditions for which the predictions of delay-reduction theory and a simple rate-maximizing theory were opposed. Results were consistent with delay reduction's ordinal predictions in 11 of 11 replications of the divergent predictions favoring the smaller, more immediate alternative. The predictions of rate maximization were upheld only when they were consistent with those of delay reduction. Results are discussed in terms of conditioned reinforcement, sensitivity to reductions in delay to food, and possible rules of thumb that may be useful in characterizing foraging.     

How pigeons estimate rates of prey encounter.

Pigeons were trained on operant schedules simulating successive encounters with prey items. When items were encountered on variable-interval schedules, birds were more likely to accept a poor item (long delay to food) the longer they had just searched, as if they were averaging prey density over a short memory window (Experiment 1). Responding as if the immediate future would be like the immediate past was reversed when a short search predicted a long search next time (Experiment 2). Experience with different degrees of environmental predictability appeared to change the length of the memory window (Experiment 3). The results may reflect linear waiting (Higa, Wynne, & Staddon, 1991), but they differ in some respects. The findings have implications for possible mechanisms of adjusting behavior to current reinforcement conditions.     

DO CHILDREN PREFER CONTINGENCIES? AN EVALUATION OF THE EFFICACY OF AND PREFERENCE FOR CONTINGENT VERSUS NONCONTINGENT SOCIAL REINFORCEMENT DURING PLAY

Discovering whether children prefer reinforcement via a contingency or independent of their behavior is important considering the ubiquity of these programmed schedules of reinforcement. The current study evaluated the efficacy of and preference for social interaction within differential reinforcement of alternative behavior (DRA) and noncontingent reinforcement (NCR) schedules with typically developing children. Results showed that 7 of the 8 children preferred the DRA schedule; 1 child was indifferent. We also demonstrated a high degree of procedural fidelity, which suggested that preference is influenced by the presence of a contingency under which reinforcement can be obtained. These findings are discussed in terms of (a) the selection of reinforcement schedules in practice, (b) variables that influence children's preferences for contexts, and (c) the selection of experimental control procedures when evaluating the effects of reinforcement.     

Meal patterns of cats encountering variable food procurement cost.

The meal patterns of 2 cats in a laboratory habitat with variable foraging costs were examined in a foraging paradigm in which subjects could initiate meals at any time by completing a predetermined number of bar presses (the procurement price) and then could eat any amount. From meal to meal, the procurement price either was fixed or varied among a geometric series of five prices. As the fixed price or the mean of the variable prices increased, meal frequency decreased and meal size increased; daily intake was unaffected. Within variable-price schedules, meal size was not related to the just-paid procurement price. These results suggest that cats respond to the global rather than to the local cost structure of their habitat. They appear to respond to an average of the prices encountered, initiating meals of a frequency and size appropriate to that average. This was true even when the average price was high, meals were infrequent, and thus price encounters were widely separated in time. Therefore, the time window over which the consequences of behavior can affect behavior is longer than often conceived, at least in economies in which the animal controls its intake and the frequency, size, and distribution of its meals.     

Frequency versus magnitude of reinforcement: New data with a different procedure

Two pigeons, with previous exposure to concurrent schedules, were submitted to 29 sessions of 8 hours each with concurrent variable-interval variable-interval schedules in which reinforcement parameters changed from session to session. In the first nine sessions reinforcement durations were equal in both schedules while reinforcement frequencies varied; in Sessions 10 through 18, both frequency and duration of reinforcement were varied; in Sessions 19 through 29, only reinforcement duration was varied. Results with this different procedure confirm previous findings that behavior is more sensitive to changes in reinforcement frequency than to reinforcement magnitude.     

WITHIN‐SESSION TRANSITIONS IN CHOICE: A STRUCTURAL AND QUANTITATIVE ANALYSIS

The present study used within‐session transitions between two concurrent schedules to evaluate choice in transition. Eight female Long‐Evans rats were trained to respond under concurrent schedules of reinforcement during experimental sessions that lasted 22 hr. The generalized matching equation was used to model steady‐state behavior at the end of each session, while transitional behavior that emerged following the change in reinforcement schedules was modeled using a logistic equation. The generalized matching and logistic equations were appropriate models for behavior generated during single‐session transitions. A local analysis of behavior on the two response alternatives during acquisition was used to determine the source of preference as revealed in response ratios. The number of “low‐response” visits, those containing three to five responses, remained stable. Preference ratios largely reflected a sharp increase in the number of visits with long response bouts on the rich alternative and a decrease in the number of such visits to the leaner alternative.