Subscribe: Journal of Comparative Psychology - Vol 123, Iss 4
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Journal of Comparative Psychology - Vol 130, Iss 3

The Journal of Comparative Psychology publishes original empirical and theoretical research from a comparative perspective on the behavior, cognition, perception, and social relationships of diverse species.

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Copyright: Copyright 2016 American Psychological Association

Reasoning versus association in animal cognition: Current controversies and possible ways forward.


The study of animal cognition is rife with controversy, and among the most long-standing and most intensely debated controversies in the field is the question of the extent to which the behavior of nonhuman animals can be fully understood on the basis of purely associative principles, or whether some behaviors exhibited by animals necessitate the assumption of inferential capacities in animals that defy an associative explanation. Remarkably, the continuing debate on the topic seems to be spawning little genuine progress in terms of substantial accumulation of new, generally accepted insights. As an introduction to a special section of the Journal of Comparative Psychology on the topic, the present article outlines a number of reasons for the stalemate and suggests ways to refertilize the debate. In particular, we claim that progress will not come from the adoption of general principles like Morgan’s canon or the primacy of prediction over postdiction. Instead, emphasis should be placed on a careful analysis of what it is that different sides in the debate do and do not agree on and an increased willingness to engage in adversarial collaboration, in the spirit of a shared interest in furthering our understanding of animal behavior. (PsycINFO Database Record (c) 2016 APA, all rights reserved)(image)

Beyond the information (not) given: Representations of stimulus absence in rats (Rattus norvegicus).


Questions regarding the nature of nonhuman cognition continue to be of great interest within cognitive science and biology. However, progress in characterizing the relative contribution of “simple” associative and more “complex” reasoning mechanisms has been painfully slow—something that the tendency for researchers from different intellectual traditions to work separately has only exacerbated. This article reexamines evidence that rats respond differently to the nonpresentation of an event than they do if the physical location of that event is covered. One class of explanation for the sensitivity to different types of event absence is that rats’ representations go beyond their immediate sensory experience and that covering creates uncertainty regarding the status of an event (thus impacting on the underlying causal model of the relationship between events). A second class of explanation, which includes associative mechanisms, assumes that rats represent only their direct sensory experience and that particular features of the covering procedures provide incidental cues that elicit the observed behaviors. We outline a set of consensus predictions from these two classes of explanation focusing on the potential importance of uncertainty about the presentation of an outcome. The example of covering the food-magazine during the extinction of appetitive conditioning is used as a test case for the derivation of diagnostic tests that are not biased by preconceived assumptions about the nature of animal cognition. (PsycINFO Database Record (c) 2016 APA, all rights reserved)(image)

All the way up or all the way down?: Some historical reflections on theories of psychological continuity.


In this paper I chart the history of the development of theories of psychological continuity in the modern period, beginning with the contrasting positions of René Descartes and Julian Offrey de la Mettrie during the period of the scientific revolution in Europe. In providing the logical geography of competing positions, I distinguish between two forms of strong psychological continuity and discontinuity, between theories of strong continuity and discontinuity between cognitive and associative processes and between theories of strong continuity and discontinuity between human and animal psychology and behavior. I note that both forms of strong continuity and discontinuity have tended to be affirmed or denied together, and have only rarely and recently been decoupled, opening up a new theoretical position in the debate, which affirms strong discontinuity between cognitive and associative processes but strong continuity between human psychology and some forms of animal psychology. Although the historical trend in the late 19th and early 20th century was to extend explanations in terms of association “all the way up” to the highest human cognitive processes, some contemporary theorists have tried to extend cognitive explanations “all the way down” to associative processes in both humans and nonhuman animals. I draw some tentative conclusions about the theoretical options in contemporary research on psychological continuity. (PsycINFO Database Record (c) 2016 APA, all rights reserved)(image)

Associative learning as higher order cognition: Learning in human and nonhuman animals from the perspective of propositional theories and relational frame theory.


We aim to provide a new perspective on the old debate about whether evidence for higher order cognition in nonhuman animals can be reinterpreted in terms of associative learning. Our starting point is the idea that associative learning is best thought of as an effect (i.e., the impact of paired events on behavior) rather than a specific mental process (e.g., the formation of associations). This idea allows us to consider (a) propositional theories according to which associative learning is mediated by higher order mental processes akin to problem solving and (b) relational frame theory that allows one to think of seemingly simple associative learning effects as instances of a complex phenomenon known as arbitrarily applicable relational responding. Based on these 2 theories, we argue that (a) higher order cognition and associative learning are not necessarily mutually exclusive and (b) a more sophisticated conceptualization of higher order cognition is warranted. (PsycINFO Database Record (c) 2016 APA, all rights reserved)(image)

Overcoming associative learning.


Thorndike (1898, 1911) rejected the idea that animal behavior was the consequence of reasoning, and suggested instead that the gradual acquisition of associations formed the basis of behavior—a contention that has had a significant impact on the development of animal learning theory. Despite this, comparative psychology provides a number of examples of behaviors that have been considered to be above and beyond the explanation of associative-, or reinforcement-learning mechanisms. These behaviors have motivated some researchers to propose higher-order cognitive abilities in animals, including (but not limited to) reasoning, sensitivity to ambiguity, and metacognition. However, other authors have questioned this claim, and provided alternative explanations for these behaviors from an associative perspective. With relevant examples, the steps that must be taken in order to overcome an associative explanation of behavior are described. (PsycINFO Database Record (c) 2016 APA, all rights reserved)(image)

Causal reasoning versus associative learning: A useful dichotomy or a strawman battle in comparative psychology?


The debate about whether or not one could/should ascribe reasoning abilities to animals has deep historical roots and seems very up-to-date in the light of the immense body of new empirical data originating from various species and research paradigms. Associative learning (AL) seems to be a ubiquitous low-level contender for any cognitive interpretation of animal behavior, mostly because of the assumed mechanistic simplicity and phylogenetic prevalence. However, the implicit assumption that AL is simple and therefore the most parsimonious mechanism to describe seemingly complex behavior can and must be questioned on various grounds. Using recent empirical findings with chimpanzees as an example, I argue that at times inferential reasoning might be the most likely candidate to account for performance differences between experimental and control conditions. Finally, a general conclusion drawn from the current debate(s) in the field of comparative psychology could be that a dichotomist battle of 2 conceptual camps—each of which is lacking a clear and homogeneous theoretical framework—is a scientific deadlock. (PsycINFO Database Record (c) 2016 APA, all rights reserved)(image)

Spatial generalization of imitation in dogs (Canis familiaris).


Dogs, like human infants, are able to imitate human actions after a delay (deferred imitation). This study demonstrates that in deferred imitation tasks, dogs can generalize imitation across context modification to a certain extent. Specifically, they can imitate an object-related action if the object used by the demonstrator is displaced to a different location. However, if the object is interchanged with a different one, their imitative performance drops while they show a spatial bias toward the location of demonstration. We used a combination of the 2-action procedure and the “Do as I do” paradigm and displaced the target objects manipulated by the demonstrator, so that, at the time of recall, dogs could only match either the original location of demonstration or the displaced object, but not both. In conditions with a single object present and displaced after the demonstration, dogs matched the action and the object on which it was shown. In conditions with the location of 2 objects interchanged, dogs more likely matched the location. However, when humans provided ostensive cues and pointing gestures to draw the subjects’ attention toward the displaced target object, dogs’ predisposition to follow human communication outweighed their spatial bias and, as a consequence, their object matching and imitative performance increased. We conclude that object’s physical features function as retrieval cues that facilitate recalling the action. In addition to figurative information, dogs rely strongly on spatial information in Do as I do tasks. The results are discussed concerning dogs’ representational system of imitative actions. (PsycINFO Database Record (c) 2016 APA, all rights reserved)(image)

Three actions, two groups: Looking for the origin of primate manual lateralization.


Handedness is the most evident behavioral asymmetry in humans: to study nonhuman primate hand preference could be optimal to investigate the evolutionary origin of handedness in our species, even though behavioral asymmetries are widespread among vertebrates. This study investigated hand preferences in 32 Old World monkeys and 26 great apes during 3 spontaneous actions, assessing the effect of taxonomic group, sex and age on primate handedness. Data about foraging, locomotion, and manipulation were collected and 50 bouts per behavioral category per subject were recorded. A bias toward right-hand use for starting locomotion was reported in both Old World monkeys and great apes. Furthermore, in the great apes, a group-level right-hand preference for manipulation was found. Results suggest an important role of factors such as posture and task complexity in the evolution of primate manual lateralization. The effect of taxonomic group, sex, and age on the hand preference are also discussed. (PsycINFO Database Record (c) 2016 APA, all rights reserved)(image)

Pattern of nipple use by puppies: A comparison of the dingo (Canis dingo) and the domestic dog (Canis familiaris).


Surprisingly little information is available about the behavior of newborn mammals in the functionally vital context of suckling. We have previously reported notable differences in the pattern of nipple use by kittens of the domestic cat and puppies of the domestic dog. Whereas kittens rapidly develop a “teat order,” with each individual using principally 1 or 2 particular nipples, puppies show no such pattern. We asked whether the more “chaotic” behavior seen in puppies of the domestic dog (Canis familiaris) could be the result of relaxed selection due to domestication. In a first test of this hypothesis, we studied suckling behavior in 4 litters of wild-type captive dingoes (Canis dingo), a canid species that has inhabited the Australian mainland in substantial numbers for at least 5,000 years with minimal human influence. On all measures of individual puppies’ behavior—time spent attached to nipples, lack of individual use of particular nipples and consequent absence of a teat order, lack of synchronized suckling with other littermates, lack of agonistic behavior—we found no differences between the 2 species. In conclusion, we suggest that the difference between the pattern of suckling behavior of kittens of the domestic cat (and other felids) and the domestic dog is not an artifact of domestication, but rather reflects phylogenetic differences between felids and canids as a consequence of their different lifestyles and associated patterns of parental care. These findings emphasize the need for comparative studies to avoid simplistic generalizations from 1 or 2 species across broad taxonomic groups. (PsycINFO Database Record (c) 2016 APA, all rights reserved)(image)

Unveiling the “secret” of play in dogs (Canis lupus familiaris): Asymmetry and signals.


Due to their playful propensity, dogs are a good model to test some hypotheses about play dynamics (length, asymmetry, features of players) and communication (play bow [PBOW]; relaxed open-mouth [ROM] display). We video-recorded 203 play sessions between dogs in an off-leash dog park in Palermo, Italy. Contrary to the expectation, play asymmetry (particularly high in this species) did not differ between stranger and familiar dogs, thus suggesting the limited role of play in forming dominance relationships. Asymmetry negatively affected the duration of the session, whereas the increasing number of players was positively linked to the duration of playful interactions. The number of PBOWs exchanged by players may exert a certain influence on the session length as well. PBOWs were performed independently from the kind of play (locomotor vs. contact) the dogs were engaging in. Conversely, ROMs were preferentially emitted during contact play when “face-to-face” interactions were more likely. Body closeness is also required in case opening the mouth has not a signal function but only preludes a bite. However, in the 82% of cases play bites did not follow a ROM, thus suggesting that dogs place ROMs in the appropriate context to optimize signal detectability. In conclusion, 2 tactics may concur in coping with the asymmetry and unpredictability of play sessions in dogs. First, whenever the asymmetry increases dogs shorten the duration of their sessions thus limiting the risk of possible escalation. Second, dogs make use of a good communicative system based on the reciprocal exchange of playful signals. (PsycINFO Database Record (c) 2016 APA, all rights reserved)(image)

Wolves (Canis lupus) and dogs (Canis familiaris) differ in following human gaze into distant space but respond similar to their packmates’ gaze.


Gaze following into distant space is defined as visual co-orientation with another individual’s head direction allowing the gaze follower to gain information on its environment. Human and nonhuman animals share this basic gaze following behavior, suggested to rely on a simple reflexive mechanism and believed to be an important prerequisite for complex forms of social cognition. Pet dogs differ from other species in that they follow only communicative human gaze clearly addressed to them. However, in an earlier experiment we showed that wolves follow human gaze into distant space. Here we set out to investigate whether domestication has affected gaze following in dogs by comparing pack-living dogs and wolves raised and kept under the same conditions. In Study 1 we found that in contrast to the wolves, these dogs did not follow minimally communicative human gaze into distant space in the same test paradigm. In the observational Study 2 we found that pack-living dogs and wolves, similarly vigilant to environmental stimuli, follow the spontaneous gaze of their conspecifics similarly often. Our findings suggest that domestication did not affect the gaze following ability of dogs itself. The results raise hypotheses about which other dog skills might have been altered through domestication that may have influenced their performance in Study 1. Because following human gaze in dogs might be influenced by special evolutionary as well as developmental adaptations to interactions with humans, we suggest that comparing dogs to other animal species might be more informative when done in intraspecific social contexts. (PsycINFO Database Record (c) 2016 APA, all rights reserved)(image)

Japanese monkeys (Macaca fuscata) quickly detect snakes but not spiders: Evolutionary origins of fear-relevant animals.


Humans quickly detect the presence of evolutionary threats through visual perception. Many theorists have considered humans to be predisposed to respond to both snakes and spiders as evolutionarily fear-relevant stimuli. Evidence supports that human adults, children, and snake-naive monkeys all detect pictures of snakes among pictures of flowers more quickly than vice versa, but recent neurophysiological and behavioral studies suggest that spiders may, in fact, be processed similarly to nonthreat animals. The evidence of quick detection and rapid fear learning by primates is limited to snakes, and no such evidence exists for spiders, suggesting qualitative differences between fear of snakes and fear of spiders. Here, we show that snake-naive Japanese monkeys detect a single snake picture among 8 nonthreat animal pictures (koala) more quickly than vice versa; however, no such difference in detection was observed between spiders and pleasant animals. These robust differences between snakes and spiders are the most convincing evidence that the primate visual system is predisposed to pay attention to snakes but not spiders. These findings suggest that attentional bias toward snakes has an evolutionary basis but that bias toward spiders is more due to top-down, conceptually driven effects of emotion on attention capture. (PsycINFO Database Record (c) 2016 APA, all rights reserved)(image)