Familiarity and relatedness: Effects on social learning about foods by Norway rats and Mongolian gerbils

In recent experiments in which the social influences on feeding in Mongolian gerbils were investigated, observer gerbils acquired food preferences from conspecific demonstrators only if the demonstrators and observers were either related or familiar. Even then, the effects of demonstrator gerbils on observers’ food choices lasted less than 24 h. In similar experiments with Norway rats, the familiarity/relatedness of demonstrators and observers had little effect on social learning, and the demonstrators’ influence on observers’ food choices lasted many days. We examined the causes of these differences and found that, after observer gerbils interacted with either unfamiliar or familiar conspecific demonstrators that had been fed using procedures typically used to feed demonstrator rats, they showed long-lasting social learning about foods, whereas observer rats interacting with conspecific demonstrators that had been fed as demonstrator gerbils normally are fed showed effects of familiarity/relatedness to demonstrators on their social learning about foods. Procedural differences, rather than species differences, seem to be responsible for reported inconsistencies in social learning about foods by rats and gerbils.     

Socially mediated attenuation of taste-aversion learning in Norway rats: Preventing development of “food phobias”

After interacting with rat demonstrators that had eaten a novel, palatable diet, many observer rats exhibited either attenuation or total blockade of their subsequent acquisition of a lithium-chloride-induced aversion to that diet. In natural circumstances, such social attenuation of aversion learning could prevent new recruits to a population (weanlings or recent immigrants) from learning maladaptive aversions (“food phobias”) to tainted or spoiled samples of normally safe foods that others of their social group were eating.     

What observer rats don’t learn about foods from demonstrator rats

Naive “observer” rats that interact with conspecific “demonstrators” fed a distinctive food increase intake of the food their demonstrators have eaten. Here we found that observer rats that had interacted simultaneously with 2 demonstrator rats, 1 fed a distinctively flavored, protein-poor food, the other a distinctively flavored, protein-rich food, did not prefer the former. Similarly, observer rats ate equal amounts of two distinctively flavored foods after interacting simultaneously with 2 demonstrator rats, 1 that had consumed all food available to it, the other fed from a surplus of the second food. Last, observer rats that had interacted with both a “trustworthy” demonstrator (1 an observer had learned ate only nutritious foods) and an “untrustworthy” demonstrator (1 an observer had learned ate noxious substances) did not prefer unfamiliar foods eaten by trustworthy demonstrators to those eaten by untrustworthy demonstrators. These findings suggest limits on social information observers use in selecting foods.     

Delays after eating: Effects on transmission of diet preferences and aversions

Previous studies have demonstrated that a naive rat (an observer), after interacting with a previously fed conspecific (a demonstrator), will exhibit an enhanced preference for the diet its demonstrator ate. Furthermore, observers poisoned after interacting with demonstrators exhibit an aversion to their respective demonstrators’ diets. In the present paper, we examined the effects, on transmission of information from demonstrator to observer, of introducing delays between the end of demonstrator feeding and initiation of demonstrator-observer interaction. We found that (1) for at least 4 h after ingestion, demonstrator rats emitted diet-related cues sufficient to alter observers’ subsequent diet preferences, and (2) diet-related cues emitted by demonstrators for 1 to 2 h after a meal were adequate conditional stimuli for aversion learning by their observers.     

Further evidence that Norway rats do not socially transmit learned aversions to toxic baits

Three experiments were undertaken to examine the effects of interactions with demonstrator rats made ill by injection of lithium chloride (LiCl) on the later food choices of their observers. We found that (1) observer rats that had been taught an aversion to an unfamiliar diet exhibited a substantial reduction of that aversion after interacting with poisoned demonstrators that had eaten the diet to which the observers had learned an aversion, (2) exposure of an observer rat to poisoned demonstrator rats that had eaten a diet interfered with later acquisition by the observer of an aversion to the diet that the poisoned demonstrators had eaten, and (3) after interacting with poisoned demonstrators that had eaten one of two diets, observers that ate both diets and were then made ill formed an aversion to whichever diet their respective, poisoned demonstrators had not eaten. The present experiments, like previous studies both in our laboratory and elsewhere, failed to provide any evidence that naive observer rats will learn to avoid a food as a result of interacting with demonstrator rats that had eaten the food and exhibit symptoms of toxicosis. To the contrary, observer rats in the present experiments exhibited an enhanced preference for foods eaten by sick demonstrators.     

Social influences on the identification of toxic foods by Norway rats

In the present experiments, a naive “observer” rat first interacted with a “demonstrator” rat previously fed a diet unfamiliar to the observer. The observer then sampled two unfamiliar diets, one of which was the diet its demonstrator had eaten. The observer was then injected with LiCl and, following recovery from toxicosis, was offered a choice between the two diets it sampled prior to toxicosis induction. It was found that: (1) each observer rat formed an aversion to whichever diet its demonstrator had not eaten, (2) effects of demonstrators on aversion learning by observers were present even if there was a 7- or 8-day delay between interaction of a demonstrator and observer and diet sampling by the observer, and (3) observers interacting with 3 demonstrators, each fed a different diet, subsequently exhibited a reduced tendency to form an aversion to each of the diets eaten by their demonstrators. Taken together, the results indicate that information acquired from conspecifics as to the diets they have eaten can play an important role in determining the foods to which otherwise naive rats will learn aversions.     

How New Caledonian crows solve novel foraging problems and what it means for cumulative culture

New Caledonian crows make and use tools, and tool types vary over geographic landscapes. Social learning may explain the variation in tool design, but it is unknown to what degree social learning accounts for the maintenance of these designs. Indeed, little is known about the mechanisms these crows use to obtain information from others, despite the question’s importance in understanding whether tool behavior is transmitted via social, genetic, or environmental means. For social transmission to account for tool-type variation, copying must utilize a mechanism that is action specific (e.g., pushing left vs. right) as well as context specific (e.g., pushing a particular object vs. any object). To determine whether crows can copy a demonstrator’s actions as well as the contexts in which they occur, we conducted a diffusion experiment using a novel foraging task. We used a nontool task to eliminate any confounds introduced by individual differences in their prior tool experience. Two groups had demonstrators (trained in isolation on different options of a four-option task, including a two-action option) and one group did not. We found that crows socially learn about context: After observers see a demonstrator interact with the task, they are more likely to interact with the same parts of the task. In contrast, observers did not copy the demonstrator’s specific actions. Our results suggest it is unlikely that observing tool-making behavior transmits tool types. We suggest it is possible that tool types are transmitted when crows copy the physical form of the tools they encounter.     

Experimental identification of social learning in wild animals

Field experiments can provide compelling demonstrations of social learning in wild populations. Social learning has been experimentally demonstrated in at least 23 field experiments, in 20 species, covering a range of contexts, such as foraging preferences and techniques, habitat choice, and predator avoidance. We review experimental approaches taken in the field and with wild animals brought into captivity and note how these approaches can be extended. Relocating individuals, introducing trained individual demonstrators or novel behaviors into a population, or providing demonstrator-manipulated artifacts can establish whether and how a particular act can be socially transmitted in the wild and can help elucidate the benefits of social learning. The type, strength, and consistency of presented social information can be varied, and the provision of conditions favoring the performance of an act can both establish individual discovery rates and help determine whether social information is needed for acquisition. By blocking particular avenues of social transmission or removing key individuals, routes of transmission in wild populations can be investigated. Manipulation of conditions proposed to favor social learning can test mathematical models of the evolution of social learning. We illustrate how field experiments are a viable, vital, and informative approach to the study of social learning.     

Defensive burying in rats and hamsters

Hooded rats and golden hamsters were shocked by one of two prods in a chamber with a sawdust-covered floor. Rats buried the prod through which they had been shocked, but hamsters displayed no burying behavior. Hamsters may not have buried the prod because they could not perform the required motor pattern. However, hamsters can carry and pile food pellets. Therefore, in a second experiment, rats and hamsters were shocked in a chamber with wooden blocks on the floor. Rats piled blocks around the prod through which they had been shocked, but hamsters did not. The third experiment established that, like rats, hamsters can associate a prod with shock in one trial, since they showed differential avoidance of a prod through which they had been shocked. Since hamsters are nonsocial and rats are social, these results are consistent with suggestions that burying sources of aversive stimulation evolved as an altruistic behavior.     

Social influence on avoidance of dangerous stimuli by rats

Three experiments were conducted to determine whether a naive observer rat would avoid contact with a shock prod after watching a demonstrator rat contact, be shocked by, and defensively bury the prod. We found that observer rats took longer to contact prods that had delivered a shock to and been buried by a demonstrator rat than to contact prods that had not delivered shock and had not been buried. However, observer rats contacted prods buried by an unseen demonstrator rat or by an unseen experimenter with the same latencies as those for prods they had seen deliver shock to and be buried by a demonstrator rat. In large enclosures, subjects took 1–2 h longer to contact buried prods than to contact unburied prods. We conclude that alteration of the physical environment by individuals receiving noxious stimulation can significantly reduce the probability that conspecifics will contact the noxious stimulus. Observational learning per se, however, need not be involved.     

Opportunities and constraints when studying social learning: Developmental approaches and social factors

Identifying social learning in wild populations is complicated by the relative lack of ability to conduct controlled experiments in natural habitats. Even in more controlled captive settings, tracking the innovation and spread of behavior among known individuals can be challenging, and these studies often suffer from a lack of ecological validity. In recent years, a host of new approaches have been undertaken to attempt to provide more quantitative control and empirical demonstration of social learning, both in the wild and in captive settings that more closely mimic natural contexts. Developmental approaches are being undertaken more regularly that allow us to study the ontogenetic trajectory of complex skills in a variety of taxa. Likewise, a spirited focus on the social context of social learning has emerged, and researchers have begun to meticulously analyze the influences of social systems and the characteristics of demonstrators and observers. Here, we provide a review of these studies and summarize the opportunities and constraints that exist when one attempts to study learning in social species. We suggest that although the study of social learning in nonhuman animals is becoming much more complex, addressing this complexity provides a fruitful model for understanding the evolution of human cultural behavior.     

Human cumulative culture in the laboratory: Effects of (micro) population size

Traditionally, experiments on social learning (in both humans and nonhumans) involve dyads, with an experimenter or experimenter-trained conspecific serving as the demonstrator and the participant as the observer. But social learning in nature often involves multiple potential models, and the models themselves were once learners. We discuss our studies of social learning by adult humans in interactive group settings in the absence of formal demonstrations by experimenters, which tracked transmission over multiple learner generations. In these experiments, we found evidence for cumulative learning over generations. This has allowed us to manipulate learning conditions in order to test hypotheses regarding the necessary conditions for cumulative culture. We also report results from a further experiment using similar methods, which compared conditions of varying cohort size. Participants were given the task to build a paper airplane to fly as far as possible. Contrary to expectations, there was no advantage for larger cohort sizes, in terms of the cumulative effects observed.     

Social transmission of food preferences among norway rats by marking of food sites and by gustatory contact

Three experiments were conducted to investigate the social learning and transmission of food preferences by excretory marking among adult male Norway rats. The experiments extend our earlier findings that rats prefer to eat from a food bowl marked by the excretory deposits of conspecifics and that this mechanism can result in the communication and social learning of food preferences (Laland & Plotkin, 1991). Here we investigate whether a tradition of food and food site preferences can become established by these means. Experiment 1 established that the residual cues deposited by rats lose their powers of communication as “markers” of food sites over a 72-h period. Experiment 2 showed that while a socially enhanced preference for one flavored diet could be transmitted from one animal to the next along a chain, it was unstable for an alternative diet. This suggests that social transmission may be more stable when it reinforces a prior preference than when it conflicts with one. In Experiment 3, the stability of socially transmitted food preferences was bolstered by the addition of a second process for the communication of diet preferences-namely, gustatory cues on the demonstrator’s breath. This finding suggests that when a socially transmitted trait is mediated by more than one process, the processes may interact, and the diffusion is likely to be more stable.     

Overshadowing and blocking between landmark learning and shape learning: the importance of sex differences

Rats were trained in a triangular-shaped pool to find a hidden platform that maintained a constant relationship with two sources of information, an individual landmark and one part of the pool with a distinctive shape. In Experiment 1, shape learning overshadowed landmark learning but landmark learning did not overshadow shape learning in males, while landmark learning overshadowed shape learning but shape learning did not overshadow landmark learning in females. In Experiment 2, rats were pretrained either with the single landmark relevant or with the shape relevant, in the absence of the alternative cue. Final test trials, without the platform, revealed reciprocal blocking only in females; in males, shape learning blocked landmark learning, but not viceversa (Experiment 2a). In Experiment 2b, male rats received a longer pretraining with the single landmark relevant, and now landmark learning blocked shape learning. The results thus confirm the claim that males and females partially use different types of spatial information when solving spatial tasks. These results also agree with the suggestion that shape learning interacts with landmark learning in much the same way as does learning about any pair of stimuli in a Pavlovian conditioning experiment.     

Incentive contrast produced by deprivation shifts

Deprivation shifts, using conditions analogous to those which produce strong incentive contrast when reward quantity is changed (extended preshift training, short interval from the last preshift trial to the first postshift trial, large preshift differences in deprivation) resulted in contrast of runway speed and choice behavior. In the first experiment, a downshift of hunger during rats’ runway training produced a slow lessening of speed below that of a group trained continually at low hunger. In the second experiment, rats were trained to traverse a runway to one food-containing goalbox when very hungry and to another, distinctively different food-containing goalbox when not very hungry. The rats were next given a series of choice trials between the two goalboxes. There was a brief preference for the high-hunger goalbox, followed by a preference for the low-hunger goalbox. The results of the second experiment suggest that deprivation affects the strength of conditioning of a cue-reinforcer expectancy, while the slow development of contrast in the first experiment indicates that deprivation also affects the development of either habit strength or a response-reinforcer expectancy.     

Attenuation by contextual cues of retroactive interference of a conditional discrimination in rats

We investigated the effect of associating unique contextual cues with an interpolated learning task on retroactive interference in long-term memory. Rats were originally trained in a two-bar operant chamber with an auditory conditional discrimination stimulus. During interpolated learning, which occurred in either the original or a new context, some rats were trained on a probability learning task that did not include the auditory stimuli present during original learning. Subsequent retraining on the original conditional discrimination task in the original context showed that (1) significant retroactive interference occurs in rats, and (2) the presence of unique contextual cues during interpolated learning significantly reduces this interference. These results extend the conditions under which the susceptibility to retroactive interference can be altered by contextual cues.     

Postshock learning and conditioned defensive burying

Natural sources of aversive stimuli are frequently well-defined material objects that are present both before and after the aversive event. In the present experiment, rats acquired information about such a source after the aversive event and used this information to guide their subsequent defensive reactions to it. The rats were shocked by one of two possible sources, either a black or a striped prod, mounted on opposite end walls of the test chamber. Immediately following the shock, the houselights were momentarily extinguished and the patterns on the two prods were automatically switched for subjects in the experimental condition or left unchanged for subjects in the control condition. The rats were left in the chamber for another 5 min with the patterns in their new positions before being removed for 2 min while the two prods were mounted on the side walls. During the ensuing test of conditioned defensive burying, the rats in the control condition directed the majority of their burying behavior at the prod exhibiting the pattern displayed by the shock source prior to and during the shock administration. In contrast, the rats in the experimental group buried the prod exhibiting the pattern displayed by the shock source during the postshock period more than they did the prod displaying the pattern present on the shock source prior to and during the shock administration.     

Irrelevant cues and shape discrimination learning by cats

Experimentally naive eats learned to discriminate a pair of objects (circle and triangle) that differed in external contour, with no irrelevant cues (N = 30), with irrelevant brightness cues (N = 26), and with irrelevant size and brightness cues (N = 26). In a second experiment, naive cats were trained to discriminate bidimensional patterns (circle and triangle) with no irrelevant visual cues (N = 24) or with irrelevant size and brightness cues (N = 26). Irrelevant cues did not significantly affect the rate of shape discrimination learning in either experiment. The findings disagree with the results of several similar experiments with rats.     

Demonstrator influence on observer diet preference: Effects of simple exposure and the presence of a demonstrator

Previous studies in our laboratory have demonstrated that a naive rat (an observer), after interacting briefly with a previously fed conspecific (a demonstrator), will exhibit an enhanced preference for the diet its demonstrator had been fed. The present studies were undertaken to determine whether demonstrator-induced alterations in observer diet preference were the result of simple exposure of observers to diet-identifying cues emitted by demonstrators during the period of demonstrator-observer interaction. Our results indicated that observer experience of diet-related cues in the stimulus context provided by the presence of a demonstrator was sufficient to enhance observer preference for a diet, whereas simple exposure to that diet was not. We concluded that demonstrator influence on observer diet preferences was not the consequence of simple exposure of observers to demonstrator-emitted cues reflecting demonstrators’ diet.     

Dissociation of patterned alternation learning and the partial reinforcement extinction effect in preweanling rats

Sprague-Dawley rat pups aged 14 or 18 days were trained on a patterned (single) alternation schedule with either an 8- or a 105-sec intertriai interval (ITI). At the 8-sec ITI, alternation learning was obtained at both ages, but the older age group learned more rapidly. There was no evidence of response alternation at the 105-sec ITI at either age. Continuously reinforced (CRF) and partially reinforced (PRF) groups trained and extinguished along with the patterned alternation (PA) group at the 105-sec ITI showed a robust partial reinforcement extinction effect (PREE) at both ages. Moreover, there was no difference in the rate of extinction of the PRF and PA groups at either age (i.e., no effect of N-length). A PREE can therefore be obtained in infant rats under conditions that apparently preclude the formation of sequential associations. The implications of this finding for the ontogeny of instrumental learning and extinction are discussed.