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Motivation, development and object play: Comparative perspectives with lessons from dogs
Abstract
Object play occurs in diverse animals in addition to birds and mammals. Although many carnivores engage in object play in a predatory context, many non-predators do so also. Conjectures over the years on the motivation to play are reviewed dealing with intrinsic, developmental, and stimulus factors. We then report on quantitative studies of the play of puppies from 6 litters (3 breeds) when given 5 different toys with different sensory and functional properties at half week intervals from 3 to 7 weeks of age. The propensity to engage with objects begins early, play complexity increases rapidly, the structure of the play is similar to adult object play, and breed differences were found. Object play with predatory characteristics appears before weaning, suggesting that hunger is not the primary motivation. Studying the development of object play in different dog breeds may be useful in addressing questions of domestication and play evolution. © 2016
... (accessed on 23 February 2023)" with a long-term goal of studying the influence of phylogeny and function (herding, retrieving, guarding, etc.) on the development of behavior. The subjects are identical to those in the earlier study [27], other than the addition of Welsh Terrier L3, which was added to create more equivalent breed sample sizes. To minimize kennel effects, different breeders with similar types of within-home housing, medical care, environmental enrichment, and husbandry practices were chosen for this study. ...
... Researchers' classification of pup weeks varies [45,48]. Our meaning of week usage here is the same week scheme used in [27,28] and is clarified here by denoting the equivalent day age of each week timepoint as follows: 3.0 weeks (21 days); 4.0 weeks (28 days); 5.0 weeks (35 days); 6.0 weeks (42 days); and 7.0 weeks (49 days). Mid-week sessions were made on day 3 or 4 of each week. ...
... Each litter was assigned one set of toys based on breed for the duration of filming. Details regarding object sizes and provision to puppies based on breed are provided in the earlier study [27]. ...
Play behavior is a prominent aspect of juvenile behavior for many animals, yet early development, especially play with objects, has received little attention. Our previous study on object play introduced our general methods, focusing on litter differences in the developmental trajectory of object play and toy preferences. Here, we present a detailed ethogram of more than 30 observed object play behaviors. We focus on breed differences in the development of play in the three following breeds: Welsh Terriers, Vizslas, and standard Poodles. Puppies were video recorded from 3 to 7 weeks of age at half-week intervals upon the introduction of a standard set of five toys into their home environments. Ten minutes of video from each session for each puppy were analyzed using the Noldus Observer XT program. Aside from analyzing individual behaviors, they were also grouped into three behavioral categories. These were behaviors that occurred only in a solitary context, only in a social context, or in both contexts. Solitary object play developed first, and social object play developed later across breeds. There was a significant three-way interaction between breed, developmental age, and the context in which play occurred. Pairwise comparisons within each breed, age, and context are discussed, but a prominent result is that the onset of many behaviors occurred later in Welsh Terriers compared to the other breeds.
... Hence the primary aim of this specific ethological study on wolf pups was to document the ontogenetic developments of object exploration and play over 2-9 weeks of age (14-63 days). The subjects were handreared wolf pups from two litters tested with diverse commercial dog toys and focuses on some of the same topics as in the Burghardt et al. (2016) paper. We had the following predictions: (1) Given that Lord (2013) found wolves start to investigate environmental stimuli at 2.0 weeks of age, while dogs do not until 4.0 weeks of age, we predict the wolf pups will show the onset of object related play significantly before it emerges in dogs. ...
... Researcher meaning of pup weeks varies (see Lord, 2013vs. Pal, 2010, hence our meaning of week usage for the rest of this paper (same week scheme used in Burghardt et al., 2016) is clarified here by denoting the equivalent day age of each week timepoint: 2.0 weeks (14 days), 3.0 weeks (21 days), 4.0 weeks (28 days), 5.0 weeks (35 days), 6.0 weeks (42 days), 7.0 weeks (49 days), 8.0 weeks (56 days), 9.0 weeks (63 days). Mid-week sessions were made on day 3 or 4 of each week since there are 7 days in a week. ...
... Each litter was tested with its own standardized set of 5 different PetSafe® toys (hereafter referred to as 'objects'): plush squirrel (Wild Squirrel TM ), plush puff (Pogo Plush TM ), braided cloth rope attached to hard rubber ball (Roly Rope TM ), a blue hard rubber disk (Twist 'n Treat TM ) and a red rigid rubber bar-bell-shaped bone (Waggle TM ). We used the medium size of the rope, disk and bone (for more details on stimuli see Burghardt et al., 2016). ...
Studies exploring the development of object play of wolf pups are lacking. Comparisons of wolves vs. dogs can aid in teasing out the influences of domestication on any differences uncovered between wolves and various dog breeds. We investigated the development of object play from 2 to 9 weeks of age (14–63 days) in 2 litters of hand-reared wolf pups (Canis lupus). Both litters of pups were video-recorded in their home enclosures at week and mid-week timepoints. Each litter was given a separate set of the same five commercial dog (Canis lupus familiaris) toys with differing sensory and functional qualities. The first 10 min of activity from each session were analyzed using the Noldus Observer XT program and a previously developed ethogram of object related behaviors. The results were analyzed using repeated measures LMMs. Percent of time spent playing with toys, behavioral counts and behavioral diversity all increased significantly with age for wolf pups, and they were generally developmentally ahead of dog puppies. Wolves, like dogs, had preferences for some toys. These findings expand our grasp of infant wolf behavior, the comparative development of object play in canines, and provide a model for the study of domestication on play behavior.
... Domestic dogs also represent an ideal model for examining possible genetic influences on behavior, given the large variety of distinct dog breeds today (Burghardt, Albright, & Davis, 2016;. At present, however, the relative roles of breed-related and environmental factors in accounting for differences in play frequency and structure among domestic dogs have received little empirical attention until recently (Bradshaw et al., 2015). ...
... At present, however, the relative roles of breed-related and environmental factors in accounting for differences in play frequency and structure among domestic dogs have received little empirical attention until recently (Bradshaw et al., 2015). One recent exception, however, was shown by Burghardt et al. (2016) that found breed differences in object play in Vizslas. Standard Poodles, and Welsh Terriers when exposed to five standardized toys. ...
... Furthermore, breed differences in the motivation to engage in different forms of play (due to different artificial selection pressures; Rooney, Bradshaw, & Robinson, 2000) has been recently suggested as a topic worthy of study and relevant to our understanding of both the evolutionary and lifetime factors that govern play behavior (Bradshaw et al., 2015); specifically, breed-specific predatory motor patterns may represent an underlying biological mechanism central to our theoretical understanding of play in domestic dogs. Although breed differences in object play has been shown previously (Burghardt et al., 2016), no study to date has demonstrated an effect of breed on play behavior in domestic dogs with a direct comparison to social and solitary play (Bauer & Smuts, 2007;Bradshaw et al., 2015). In addition, it is currently unknown whether breeds that possess different predatory motor sequences differ in their propensities to engage in different forms of play (i.e., social and solitary play) at different rates, or in the impact Learn Behav of environmental manipulations on rates and forms of play behavior, and whether there are interactions between breed and the impact of environmental manipulations. ...
The domestic dog is an ideal model species in which to study the genetic and environmental factors that influence play behavior. Dogs exist in a wide variety of breeds and frequently engage in multiple forms of play. In the present study, we investigated whether the levels of solitary and social play differed between dogs of three breed types with distinct predatory motor pattern sequences (herding dogs, retrievers, and livestock guarding dogs [LGDs]). Furthermore, we investigated how environmental factors (social and nonsocial contexts) influenced play in dogs of these breed types. Groups of breed-matched dyads with working experience and of equivalent age, sex, and neuter status ratios were exposed to four experimental test conditions and two control conditions in randomized orders. With respect to solitary play, environmental context did have a significant effect, with toys reliably producing the highest levels of solitary play across all breed types. Retrievers engaged in significantly higher levels of solitary play overall than LGDs, and there was a trend in comparison to herding dogs. In contrast, neither environmental context nor breed had a significant effect on social play levels; however, neuter status of the dyads did have a significant effect on social play, with mixed-status dyads engaging in significantly higher levels of social play than same-status dyads. Our findings provide experimental evidence for identifying proximate, environmental stimuli that reliably facilitate social and solitary play and discuss possible genetic (i.e., breed type) and lifetime influences on the form of play in domestic dogs.
... By tradition, play in non-human animals has been divided into three categories: (a) play with others ("social play"), (b) play with objects ("object play"), and (c) solitary play involving the animal moving its body in peculiar ways (e.g., rapid runs, jumps, body twists and rotations) ("locomotor-rotational play") (Fagen, 1981;Burghardt, 1998;Burghardt, 2005). Sometimes, components of two or more of the three can be combined (e.g., Biben, 1982;Pellis, 1991;Donaldson et al., 2002;Shimada, 2012;Burghardt et al., 2016;Manitzas Hill et al., 2023). Even in their pure forms, the content of each category can be diverse, for example, social play can involve behavior typically associated with sex, maternal activities, predation, or conspecific aggression (Pellis, 1988;Pellis and Pellis, 2009). ...
... As the name implies, play fighting resembles serious fighting, although given its differences with serious fighting, some authors prefer to call this behavior rough-and-tumble play as it emphasizes the cooperative, pleasurable side of the behavior (e.g., Palagi, Pellegrini, 2002;Scott and Panksepp, 2003;Burghardt et al., 2016;Smith and StGeorge, 2022), especially when presenting the phenomenon to audiences that may be hostile to any hint of aggression or violence (Pellis et al., 2022a). Nonetheless, a central feature of play fighting is competition, which is what makes it resemble serious fighting, but competition for what? ...
Play fighting has been one of the most intensely studied forms of play and so has provided some of our deepest insights into the understanding of play in general. As the label implies, this behavior resembles serious fighting, in that the animals compete for an advantage over one another, but unlike true aggression, for play fighting to remain playful, it also incorporates a degree of cooperation and reciprocity-restrained competition seems to be its hallmark. Despite these common features, it should be noted that both the advantage competed over and the mechanisms by which restraint is achieved varies across species. Such variation mitigates simple generalities. For example, how empirical support for a proposed adaptive function in one species not being replicated in another, is to be interpreted. What has emerged over the past few decades is that play fighting is diverse, varying across several dimensions, some superficial, some fundamental, making choosing species to compare a challenge. In this paper, we explore various design features that constitute play fighting and the ways these can be modified across different species and lineages of species. Given that a major pillar of ethology is that description precedes explanation, having a good grasp of the behavioral diversity of play fighting is an essential starting point for detailed analyses of the mechanisms and functions of play. We show that commonalities across species likely involve different mechanisms than do species idiosyncrasies, and that different styles of play fighting likely afford different adaptive opportunities.
... Given that some species lineages have evolved one type of play to the exclusion of another (e.g., social play but not object play) and for those that engage in multiple forms of play, some are more dominant than others (e.g., locomotor is more frequent than social), it is likely that the general system has been tapped into separately by neural circuits that are specifically engaged when playing in a particular mode. However, some species seem to be able to fluidly combine different types of play (e.g., Shimada, 2012;Burghardt et al., 2016;Palagi, 2018;Palagi and Bergman, 2021), suggesting that the specific neural circuits have become connected in a network. Indeed, the evolution of play proceeding from single forms to multiple and then multiple forms that can merge together may require a series of changes in the neural systems involved . ...
... The behavioural structure of object play in puppies is similar to that in adult dogs 29,32 . There are some notable breed differences in solitary play from as young as 7 weeks of age, for example retrievers are more likely to engage in solitary play than livestock-guarding dogs, but social play with other dogs does not appear to differ according to breed 29,33 . ...
Domesticated animals are famous for the ease with which they can accommodate to diverse human environments and roles, but less well-studied is the ease with which domestic animals can manipulate their human caregivers to their own ends. For example, domestic animals may start and end their play behaviour with humans at times of their choice. Here we present the results of a survey of 924 cat owners who report fetching behaviour in 1154 cats. The overwhelming majority (94.4%) of these owners report that fetching emerged in the absence of explicit training. Fetching was primarily first noticed when the cats were less than one year old (n = 701) or 1–7 years old (n = 415). Cats initiated and terminated fetching bouts more often than did their owners. Thus, cats who fetch demonstrate independent and co-ordinated agency in the onset and maintenance of fetching behaviour with their human partners. Additional findings highlight the diversity of objects fetched and the diversity in household demographics. Our thematic analysis reveals owners’ perspectives on (a) the process of a fetching session, (b) the initial acquisition of fetching, and (c) the circumstantial factors that influence fetching patterns. In summary, cats who fetch largely determine when they engage in fetching sessions and actively influence the play behaviour of their owners.
... Third, some forms of object play appear to have reached a level of structural, causal, and functional coherence in their organization that is characteristic of behavior systems (Pelletier et al., 2017;Pellis et al., 2019). Fourth, the numerous parallels drawn between the evolution and ontogeny of object manipulation in general (Heldstab et al., 2020) and object play in particular (Bjorklund and Gardiner, 2011;Pellegrini and Bjorklund, 2004;Pellegrini and Pellegrini, 2012;Pellegrini et al., 2007), and between the ontogeny and mechanisms of object play (Baldwin et al., 1993;Bigelow et al., 2004;Burghardt et al., 2016;Caro, 1980) are in line with Bergman and Beehner's (2022) proposition to group evolution, ontogeny, and mechanisms under an umbrella category of causes. Fifth, regarding long-term consequences, (object) play behavior can alter local environmental conditions via cultural or niche construction processes (Pellegrini and Pellegrini, 2012). ...
My main goal in this paper is to propose a reformulation of foundational models in behavioral research, including Tinbergen's (1963) well-known four levels of analysis (namely, ontological, mechanistic, functional, and evolutionary questions) and Mayr's (1961) dichotomy between proximate and ultimate causations. After critically evaluating these influential but problematic models, I present a three-level neo-Tinbergian approach to behavior that considers the triadic integration of behavioral causes, structure, and consequences along a single temporal continuum. I then argue that object-directed play is a good candidate behavior to apply this new paradigm by presenting significant examples of the combined analysis of at least two of these three levels. Finally, I show how stone handling, a form of culturally-transmitted object play in macaques, is perfectly amenable to this unified three-level explanatory framework. My proposed approach fits recent theoretical and empirical advances in behavioral biology, has a heuristic value, and may provide numerous benefits to a range of behavioral scientists.
... While social play in rat breeds is the most studied, dogs also are prime targets for study of breed differences, since they have been selected for many features involving hunting, guarding, herding, and so on (e.g., Coppinger & Coppinger, 1998), and a comprehensive molecular phylogeny of breeds is available (Parker et al., 2017). Differences among breeds in the ontogeny of object play have been documented (Burghardt et al., 2016). ...
... Play is critical for development [151][152][153][154][155][156][157][158][159][160]. In dogs, play is thought to have three primary functions: locomotory development, training for the unexpected, and social cohesion. ...
There are over 10 million pet dogs in the UK alone, and they have become a member of modern human families. If not properly socialised as puppies, dogs have a higher risk of problematic behaviours during adulthood, yet socialisation studies are lacking. Much of the experimental research was carried out at least 50 years ago, and the importance of socialisation was demonstrated so clearly that further studies with unsocialised controls would be deemed unethical. In this review, the aim was to evaluate all literature relevant to canine socialisation. This review used PRISMA-P guidelines to identify 29 studies: 14 were questionnaire-based studies (two of which also had a testing element), 15 included some form of experimental manipulation relating to socialisation, and one was a purely observational study. Based on this literature review, we recommend future research into minimum necessary socialisation levels, as well as breed differences in the timing of effective socialisation. Such studies will help owners and breeders produce well-adjusted adult dogs.
... We used food and toys as both are considered to function as rewards in dogs (Gerencsér et al., 2018). However, they can be associated with different appetitive behavioral actions (i.e., ingestion of a palatable item vs. object manipulation), motivational states (e.g., Burghardt et al., 2016), and individual responsiveness (Gerencsér et al., 2018). Based on the previous evidence that the inner brow raiser serves a communicative function (Kaminski et al., 2017), but does not reflect an emotional state (Caeiro et al., 2017;Kaminski et al., 2017;Bremhorst et al., 2019), we predicted a higher incidence of the inner brow raiser in the social context (when facing a human) than in the non-social context, but no effect of trial valence. ...
Whilst humans undisputedly shape and transform most of earth's habitats, the number of animals (domestic and wild) living on this planet far outnumbers that of humans. Inevitably, humans have to interact with animals under a variety of circumstances, such as during conservation efforts, wildlife and zoo management, livestock husbandry, and pet keeping. Next to the question of how humans deal with these interactions and conflicts, it is crucial to understand the animal's point of view:
How do animals perceive and differentiate between humans?
How do they generalize their behavior towards humans?
And how does knowledge about humans spread socially?
In this Research Topic, we aim to collect original empirical work and review articles to get a more comprehensive and diverse picture on how humans are part of the sensory and cognitive world of non-human animals. We strongly invite contributions that pinpoint shortcomings and limitations in interpreting the available research findings, that provide new cross-disciplinary frameworks (e.g. links between conservation biology and comparative psychology, or human-animal interactions at zoos and animal welfare) and that discuss the applied implementation of these findings (e.g. for conservation attempts or livestock husbandry management).
... Some wild animals have been shown to use humanmade objects for nesting or shelter; few examples exist of their utilization for play (Paulos et al., 2010). Domestic dog pups use objects for play at a very early age prior to weaning, suggesting such interactions are not associated with hunger and are indeed object play (Burghardt et al., 2016). We have a strong foundation for understanding the various effects of group composition, breeder condition, and even environmental fluctuations on reproduction in social carnivores (Ausband et al., 2017;Bateman et al., 2011;Brainerd et al., 2008;Stahler et al., 2013). ...
Some animals use humanmade objects for building and constructing nests or shelter and even for play. Gray wolves (Canis lupus) gather and use humanmade objects discovered in their natural environment. Gathering humanmade objects is a peculiar behavior particularly when there is no immediately apparent benefit to survival or reproduction. I opportunistically documented 46 different types of humanmade objects with plastic bottles and aluminum cans being the most common items found at wolf pup-rearing sites. Many objects were made of materials that appeared suitable to alleviate pain in teething pups. For some objects, however, it was not immediately obvious that they would alleviate teething pain due to their unpliable material. Additionally, such objects were quite rare in wolves' natural environment although it was not uncommon to find them at pup-rearing sites. Rare humanmade objects may provide a novelty that stimulates pups more than common objects. I hypothesize that objects used by wolf pups 1) alleviate pain from teething, and 2) provide adults respite from energetic pups. The latter is an important distinction because it implies the benefit of object play is to the adults and not the pups per se. Gathering novel objects that occupy energetic and hungry pups may influence the overall ability of social carnivores to leave young unattended while they hunt, to rest upon their return, and ultimately rear young successfully.
... This may have contributed to the lack of a size effect by reducing the likelihood the stimuli were perceived as potential prey. However, evidence suggests that use of domestic dogs as predators in a "chase" game with nonedible toys is likely to replicate predator behavior in an ecologically relevant way (Burghardt et al., 2016). Dogs' play has been characterized as the expression of "natural" behaviors such as predation, but with less intensity and where the end goal is not reached, as in this experiment where the toy "snake" is not killed and consumed (Bradshaw et al., 2015). ...
Caudal autotomy is a dramatic antipredator adaptation where prey shed their tail in order to escape capture by a predator. The mechanism underlying the effectiveness of caudal autotomy as a pre‐capture defense has not been thoroughly investigated. We tested two nonexclusive hypotheses, that caudal autotomy works by providing the predator with a “consolation prize” that makes it break off the hunt to consume the shed tail, and the deflection hypothesis, where the autotomy event directs predator attacks to the autotomized tail enabling prey escape. Our experiment utilized domestic dogs Canis familiaris as model predator engaged to chase a snake‐like stimulus with a detachable tail. The tail was manipulated to vary in length (long versus short) and conspicuousness (green versus blue), with the prediction that dog attacks on the tail should increase with length under the consolation‐prize hypothesis and conspicuous color under the deflection hypothesis. The tail was attacked on 35% of trials, supporting the potential for pre‐capture autotomy to offer antipredator benefits. Dogs were attracted to the tail when it was conspicuously colored, but not when it was longer. This supports the idea that deflection of predator attacks through visual effects is the prime antipredator mechanism underlying the effectiveness of caudal autotomy as opposed to provision of a consolation prize meal.
... We used food and toys as both are considered to function as rewards in dogs (Gerencsér et al., 2018). However, they can be associated with different appetitive behavioral actions (i.e., ingestion of a palatable item vs. object manipulation), motivational states (e.g., Burghardt et al., 2016), and individual responsiveness (Gerencsér et al., 2018). Based on the previous evidence that the inner brow raiser serves a communicative function (Kaminski et al., 2017), but does not reflect an emotional state (Caeiro et al., 2017;Kaminski et al., 2017;Bremhorst et al., 2019), we predicted a higher incidence of the inner brow raiser in the social context (when facing a human) than in the non-social context, but no effect of trial valence. ...
The inner brow raiser is a muscle movement that increases the size of the orbital cavity, leading to the appearance of so-called 'puppy dog eyes'. In domestic dogs, this expression was suggested to be enhanced by artificial selection and to play an important role in the dog-human relationship. Production of the inner brow raiser has been shown to be sensitive to the attentive stance of a human, suggesting a possible communicative function. However, it has not yet been examined whether it is sensitive to human presence. In the current study, we aimed to test whether the inner brow raiser differs depending on the presence or absence of an observer. We used two versions of a paradigm in an equivalent experimental setting in which dogs were trained to expect a reward; however, the presence/absence of a person in the test apparatus was varied. In the social context, a human facing the dog delivered the reward; in the non-social context, reward delivery was automatized. If the inner brow raiser has a communicative function and dogs adjust its expression to an audience, we expect it to be shown more frequently in the social context (when facing a person in the apparatus) than in the non-social context (when facing the apparatus without a person inside). The frequency of the inner brow raiser differed between the two contexts, but contrary to the prediction, it was shown more frequently in the non-social context. We further demonstrate that the inner brow raiser is strongly associated with eye movements and occurs independently in only 6% of cases. This result challenges the hypothesis that the inner brow raiser has a communicative function in dog-human interactions and suggests a lower-level explanation for its production, namely an association with eye movements.
... One such example is the domestic dog, a species that spends substantial amounts of energy and resources on humandirected play behaviour (e.g. [9][10][11]). The domestic dog also differs from its ancestral species, the wolf, in that play behaviour and in particular humandirected play behaviour is much more frequent and maintained throughout adulthood. ...
Human-directed play behaviour is a distinct behavioural feature of domestic dogs. But the role that artificial selection for contemporary dog breeds has played for human-directed play behaviour remains elusive. Here, we investigate how human-directed play behaviour has evolved in relation to the selection for different functions, considering processes of shared ancestry and gene flow among the different breeds. We use the American Kennel Club (AKC) breed group categorization to reflect the major functional differences and combine this with observational data on human-directed play behaviour for over 132 breeds across 89 352 individuals from the Swedish Dog Mentality Assessment project. Our analyses demonstrate that ancestor dogs already showed intermediate levels of human-directed play behaviour, levels that are shared with several modern breed types. Herding and Sporting breeds display higher levels of human-directed play behaviour, statistically distinguishable from Non-sporting and Toy breeds. Our results suggest that human-directed play behaviour played a role in the early domestication of dogs and that subsequent artificial selection for function has been important for contemporary variation in a behavioural phenotype mediating the social bond with humans.
... For some breeds, such selection has led to engaging in these actions toward non-edible objects being reinforcing in itself, manifest as play behavior and an obsession-like desire for object-play (3). Moreover, the act of performing the behavior appears to be intrinsically rewarding and is unrelated to satisfying nutritional needs (23). The desire for object-play is then harnessed as a potent reinforcer allowing for the repetition of hundreds of trials without the risk of satiety (3), and functions as a powerful motivator to work over extended periods of time. ...
Detection dogs are widely considered the most effective and adaptive method for explosives detection. Increases in emerging sophisticated threats are accelerating the demand for highly capable explosives detection, causing a strain on available supplies of quality canines worldwide. These strains are further compounded by rigorous behavioral standards required to meet mission-specific capabilities, leading to high rates of dogs disqualified from training or deployment. Ample research has explored the behavioral characteristics important for assistance, guide, and other traditional working roles, while those corresponding to more specialized tasks such as detection of explosives are not as well-understood. In this review we aim to identify the behavioral characteristics important for operational tasks of explosives detection dogs, contrasting with that of other working roles and highlighting key differences between explosives and other types of detection dogs. Further, we review the available research on methods for assessing and selecting candidate detection dogs and make recommendations for future directions and applications to the industry. Improvements and standardization in assessment technology allowing for the identification and enhancement of behavioral characteristics will be key to advancing canine detection technology in general.
... For some breeds, such selection has led to engaging in these actions toward non-edible objects being reinforcing in itself, manifest as play behavior and an obsession-like desire for object-play (3). Moreover, the act of performing the behavior appears to be intrinsically rewarding and is unrelated to satisfying nutritional needs (23). The desire for object-play is then harnessed as a potent reinforcer allowing for the repetition of hundreds of trials without the risk of satiety (3), and functions as a powerful motivator to work over extended periods of time. ...
Detection dogs are widely considered the most effective and adaptive method for explosives detection. Increases in emerging sophisticated threats are accelerating the demand for highly capable explosives detection, causing a strain on available supplies of quality canines worldwide. These strains are further compounded by rigorous behavioral standards required to meet mission-specific capabilities, leading to high rates of dogs disqualified from training or deployment. Ample research has explored the behavioral characteristics important for assistance, guide, and other traditional working roles, while those corresponding to more specialized tasks such as detection of explosives are not as well-understood. In this review we aim to identify the behavioral characteristics important for operational tasks of explosives detection dogs, contrasting with that of other working roles and highlighting key differences between explosives and other types of detection dogs. Further, we review the available research on methods for assessing and selecting candidate detection dogs and make recommendations for future directions and applications to the industry. Improvements and standardization in assessment technology allowing for the identification and enhancement of behavioral characteristics will be key to advancing canine detection technology in general
... Furthermore, behavioral development in these highly altricial animals can be theoretically and comparatively important. Dogs, wolves, and related canids are the focus of much developmental research [19][20][21][22][23][24], while bears, although less social as adults than many other carnivores, have a far longer period of maternal care, and can provide tests of hypotheses deriving from research on canids. ...
Denning behavior has long remained the least observed aspect of bear behavior. During 2010–2013, we used webcams, microphones, the internet, and 14,602 h of archived video to document the denning behaviors of two adult wild black bears (Ursus americanus) as they gave birth and cared for four litters through six winters in northeastern Minnesota. Observations included types of dens, labor, pre-parturient genital swelling, birthing positions, post-partum vocalizations, mothers removing amniotic tissues and warming newborn cubs in sub-freezing temperatures, frequency of nursing, cubs establishing nipple order, yearlings suckling, the ingestion of snow and icicles, the ingestion of foot pads, urination and defecation in latrine areas, toilet-licking, eye opening, reciprocal tongue-licking, play, rapid eye movement (REM) sleep and possible dreaming, and reactions to wildlife intruders. The use of this new method for observing natural bear dens allowed the identification of many behaviors undescribed for any species of wild bear in dens. We also discuss the need for future studies and how the depth and duration of black bear hibernation varies with body condition and geographic region.
... These phylogenetic considerations suggest that play evolved multiple times in the animal kingdom, with different types of play evolving independently of one another, although a few species that have more than one type of play in their repertoire show evidence that they can mix those types in novel ways (e.g., Burghardt et al., 2016;Shimada, 2012). It is in the context of this patchy distribution of play and its various forms, that the questions related as to whether play constitutes a behavior system needs to be considered. ...
Given that many behavior patterns cluster together in sequences that are organized to solve specific problems (e.g., foraging), a fruitful perspective within which to study behaviors is as distinct ‘behavior systems’. Unlike many behavior systems that are widespread (e.g., anti-predator behavior, foraging, reproduction), behavior that can be relegated as playful is diverse, involving behavior patterns that are typically present in other behavior systems, sporadic in its phylogenetic distribution and relatively rare, suggesting that play is not a distinct behavior system. Yet the most striking and complex forms of play have the organizational integrity that suggests that it is a behavior system. One model that we develop in this paper, involves three stages of evolutionary transition to account for how the former can evolve into the latter. First, play-like behavior emerges from the incomplete development of other, functional behavior systems in some lineages. Second, in some of those lineages, the behavior patterns typical of particular behavior systems (e.g., foraging) are reorganized, leading to the evolution of specific ‘play behavior systems’. Third, some lineages that have independently evolved more than one such play behavior system, coalesce these into a ‘super system’ allowing some animals to combine behavior patterns from different behavior systems during play. Alternative models are considered, but irrespective of the model, the overall message from this paper is that the conceptual framework of the behavior system approach can provide some new insights into the organization and diversity of play present in the animal kingdom.
... When measuring humandirected playfulness, tug-of-war is among the three most commonly performed types of heterospecific social play between dogs and owners (Horowitz & Hecht, 2016). However, because behavioral play repertories vary with context (Rooney et al., 2000) and juvenile dogs show object preferences in playful situations (Burghardt, Albright, & Davis, 2016), future research should investigate both context-and age-specific play responses in dogs and This document is copyrighted by the American Psychological Association or one of its allied publishers. ...
The domestication of animals and plants offers an exceptional opportunity to study evolutionary adaptations. In particular, domesticated animals display several behavioral alterations, including increased sociability and decreased fearfulness and aggression, when compared with their wild ancestors. However, studies quantifying simultaneous changes in multiple behaviors during domestication are lacking. Moreover, the role of human-directed play behavior has been largely neglected when studying the domestication process. Here we address these issues by examining behavioral changes during the domestication of the dog ( Canis familiaris ) from the gray wolf ( Canis lupus ) using a standardized behavioral test applied to wolf hybrids and several dog breeds. Contrary to expectations, our study provides little support for collective behavioral alterations. Specifically, although we found that wolf hybrids were less playful and overall more fearful than dogs, we did not detect any differences in sociability or aggression between wolf hybrids and dog breeds. Instead, our results suggest that behavioral alterations during domestication do not necessarily occur in concert and point to an important, but previously overlooked, role of selection on play behavior directed at humans during the domestication of dogs.
... A commonly held view of play is that it involves intermixing behavior patterns from multiple functional behaviors (Heymer, 1977;Meyer-Holzapfel, 1956;Millar, 1981) and there are empirical studies that support this view (e.g., Bekoff, 1974;Burghardt, Albright, & Davis, 2016;Donaldson, Newberry, Špinka & Cloutier, 2002;Petrů, Špinka, Charvátová, & Lhota, 2009;Shimada, 2012). In contrast, other studies of play fighting in species in which more than one type of behavior is simulated has shown that sequences of one type of play fighting are not interrupted with shifts to targeting the body parts typical of another type of behavior (Pasztor, Smith, MacDonald, Michener, & Pellis, 2001;Pellis et al., 2000). ...
Play fighting is a commonly reported form of play that involves competitive interactions that generally do not escalate to serious fighting. Although in many species what are competed over are the body targets that are bitten or struck in serious fighting, for many others, the competition can be over other forms of contact, such as sex, social grooming, and predation. In primates, the most detailed studies have been of species such as Old World monkeys, that engage in play fighting that simulates serious fighting, but reports of a number of others, especially among nocturnal prosimians, have noted that play fighting can also involve simulation of sex and grooming. The present study on captive born gray mouse lemurs (Microcebus murinus) provides a quantitative assessment of the relative engagement by juveniles in play fighting involving agonistic and amicable targets. About 80% of play fighting involves competing to groom or mount one another, with a minority involving competing to bite. That these forms of play fighting may be distinct from one another is suggested by the finding that attack on one target does not lead to counterattack on another. The findings are discussed in terms of the evolution and mechanisms underlying play fighting in primates and more widely among animals. (PsycINFO Database Record
With the improvement of people's consumption levels and the increasing demand for emotional companionship, the status of pets in families has risen significantly. Some pet - owners affectionately call their pets "fur - kids", which reflects the care and attention they give to their pets. According to the China Pet Industry White Paper 2023 - 2024, the number of pet dogs and pet cats in China reached 51.75 million and 69.8 million respectively in 2023. Behind these huge numbers lies a pet consumption market with great potential, and pet toys, as an important part of the pet supplies market, have seen a growing consumption market year by year. This study aims to explore and analyze consumers' consumption intentions for pet toys. It mainly collects and analyzes multi - dimensional data such as consumers' basic information, performance expectations, effort expectancy, facilitating conditions, consumption habits, and security through the method of distributing questionnaires. The study will also explore the impact of different age, gender, education, occupation and other social demographic characteristics on consumption intentions, so as to reveal the consumption trends and potential opportunities in the pet toy consumption market. This paper deeply analyzes various factors affecting consumers' consumption intentions for pet toys and puts forward many targeted improvement suggestions, hoping to provide some new inspiration for related enterprises, enabling them to better meet consumers' needs and jointly promote the healthy development of the pet consumption market.
Traditionally pigs are perceived as farm animals, but in the last 70–80 years they are serving some other purposes, particularly as companion animals. It is necessary to have structured and comprehensive data on the specifics of their perception, communication and social behavior to manage their behavior and welfare effectively. This article gathers and analyzes information from currently available publications and studies conducted on wild boars, domestic pigs of productive breeds and miniature pigs, characterizing the functioning of their sense organs, specifics of social behavior and communication with humans.
Domesticated animals are famous for the ease with which they can accommodate to diverse human environments and roles, but less well-studied is the ease with which domestic animals can manipulate their human caregivers to their own ends. Here we present the results of a survey of 924 cat owners who report fetching behaviour in 1,154 cats. The overwhelming majority (94.4%) of these owners report that fetching emerged in the absence of explicit training. Fetching was primarily first noticed when the cats were less than one year old ( n = 701) or 1 to 7 years old ( n = 415). Cats initiated and terminated fetching bouts more often than did their owners. Thus, cats who fetch demonstrate independent and co-ordinated agency in the onset and maintenance of fetching behaviour with their human partners. Additional findings highlight the diversity of objects fetched and the diversity in household demographics. Our thematic analysis reveals owners’ perspectives on (a) the process of a fetching session, (b) the initial acquisition of fetching, and (c) the circumstantial factors that influence fetching patterns. In summary, cats who fetch largely determine when they engage in fetching sessions and actively influence the play behaviour of their owners.
Although many species of Cetacea play, few detailed studies of their play exist. The present thesis provides detailed descriptions of two types of social play (sexual play and mouth-to-mouth interaction play) and one type of non-social play (object play) in belugas (Delphinapterus leucas), an Artic dwelling species of toothed whale. The group studied contained animals of many ages of both sexes and was maintained under human care. Given the unique features of different types of play, a major conclusion of this thesis is that play is multi-functional. Indeed, even a single form of play (e.g., mouth-to-mouth interactions) may have more than one function. For example, while mouth-to-mouth interactions may help train motor coordination skills in immature animals, such play may be used to form and maintain social relationships in adults. For belugas, play is serving many functions, all depending on the type of play and the age of the animals.
Play fighting, the most iconic form of social play, has been hypothesized to serve multiple evolutionary roles as a function of the age of the players. Although widely practised by youngsters, in some mammal species this form of play can also be present in adults. Here, we aimed to test these hypotheses by looking at the play-fighting behaviour of spotted hyaenas, Crocuta crocuta, by analysing the behaviour across age classes. Spotted hyaenas live in fission–fusion societies characterized by a rigid, nepotistic system of dominance hierarchy. Yet, the species is also characterized by social flexibility, which is evident from the high levels of support, cooperative behaviours and alliances. All these social features make spotted hyaenas a valuable model to explore play fighting at every age. We found that both immature individuals and adults invested a comparable amount of time in playful activities and showed similar motivation in initiating and maintaining their playful interactions. By play fighting, immatures can improve their motor and physical skills as predicted by the motor training hypothesis and, in agreement with the social assessment hypothesis, adults can gain information on social partners with whom they will have to interact in the future. Finally, contrary to our expectations, we found that those playful interactions characterized by strong competition (measured via play asymmetry index) also had the longest durations independently from the age of the players involved. Owing to the absence of escalation into real fighting, both immatures and adults appear to be able to manage the playful sessions despite their unbalanced nature. All these findings suggest that play fighting in spotted hyaenas can function as a ‘safe social bridge’ navigating both immatures and adults into the real future competitive challenges of the clan.
Although play fighting has been studied for over a century in both human and non-human animals, quantitative data on marine mammals are still scarce. Here, we investigated play fighting in South American sea lions (Otaria flavescens), one of the most sexually dimorphic species with an extreme polygynous mating system, high levels of both intra- and intersexual competition. All these features make South American sea lions a
good model species to test some predictions on play fighting. Our results indicate play is restricted to juveniles, being inhibited among adults, and as to be expected in a species that shows a high degree of sexual dimorphism, it is mainly expressed in males. Even though playful interactions were punctuated by competitive behaviours, animals played in a highly symmetric way and were able to adjust their competitive playful interactions in a flexible manner and so reduce the risk of escalation to a minimum level. They were highly selective in their choice of playmates by limiting the number of players per session and playing more with agematched companions and friends. All these factors taken together are probably at the basis of the low risk of escalation recorded during the study. This result is predictive of a high ability and motivation of these
animals to engage in play behaviour which can have a possible role not only in the acquisition of dominance status, but also in establishing and maintaining social relationships, an unexpected role in a so highly competitive species.
Play behavior in crocodilians is not uncommon, but it remains virtually undescribed in scientific literature. I present the first overview of play behavior of three types (locomotor play, object play and social play) in crocodilians based on original observations, published reports and anecdotal evidence. Object play is the type most often reported; social play can include interactions with conspecifics and mammals. Apparently, play behavior is not particularly rare in crocodilians, but is underreported due to the difficulties of observing it and interpreting the observations.
Object play in primates is viewed as generally having no immediate functional purpose, limited for the most part to immature individuals. At the proximate level, the occurrence of object play in immatures is regarded as being intrinsically self-rewarding, with the ultimate function of supporting motoneuronal development and the acquisition of skills necessary to prepare them for survival as adults. Stone handling (SH), a solitary object play behaviour occurs, and has been studied, in multiple free-ranging and captive troops of provisioned Japanese macaques, as well as rhesus and long-tailed macaques for over 35 years now. A review of our combined findings from these observations reveal that infants acquire SH in the first 3-4 months of life and exhibit increasingly more complex and varied behavioural patterns with age. The longitudinal data shows that many individuals maintain this activity throughout life, practicing it under relaxed ecological and social conditions. The ultimate function may be bimodal, promoting motor development in young and neural maintenance and regeneration in adult and aging individuals.
In a recent paper,(1) we examined whether oxytocin in the domestic dog modulates the maintenance of close social bonds in non-reproductive contexts. We found that exogenous oxytocin promotes positive social behaviors not only toward conspecifics, but also toward human partners. Here we examined in further detail the effect that oxytocin manipulation has on social play. When sprayed with oxytocin, subjects initiated play sessions more often and played for longer periods of time than when sprayed with saline. Furthermore, after oxytocin nasal intake dogs displayed play signals more often than after saline administration, suggesting that oxytocin enhances dogs' play motivation. To our knowledge, this study provides the first evidence that oxytocin promotes social play in the domestic dog. We use these results to hypothesize on the potential therapeutic use of oxytocin for promoting social behaviors and treating social deficits in the domestic dog.
Social play behavior, abundant in the young of most mammalian species, is thought to be important for social and cognitive development. Social play is highly rewarding, and as such, the expression of social play depends on its pleasurable and motivational properties. Since the motivational properties of social play have only sporadically been investigated, we developed a setup in which rats responded for social play under a progressive ratio schedule of reinforcement. Dopaminergic neurotransmission plays a key role in incentive motivational processes, and both dopamine and noradrenaline have been implicated in the modulation of social play behavior. Therefore, we investigated the role of dopamine and noradrenaline in the motivation for social play. Treatment with the psychostimulant drugs methylphenidate and cocaine increased responding for social play, but suppressed its expression during reinforced play periods. The dopamine reuptake inhibitor GBR-12909 increased responding for social play, but did not affect its expression, whereas the noradrenaline reuptake inhibitor atomoxetine decreased responding for social play as well as its expression. The effects of methylphenidate and cocaine on responding for social play, but not their play-suppressant effects, were blocked by pretreatment with the dopamine receptor antagonist α-flupenthixol. In contrast, pretreatment with the α2-adrenoceptor antagonist RX821002 prevented the play-suppressant effect of methylphenidate, but left its effect on responding for social play unaltered. In sum, the present study introduces a novel method to study the incentive motivational properties of social play behavior in rats. Using this paradigm, we demonstrate dissociable roles for dopamine and noradrenaline in social play behavior: dopamine stimulates the motivation for social play, whereas noradrenaline negatively modulates the motivation for social play behavior and its expression.
In this quick guide, Gordon Burghardt considers the criteria for ascribing a particular animal behavior as "play", and in particular the evidence for play in fishes, frogs and reptiles.
Copyright © 2015 Elsevier Ltd. All rights reserved.
Whether play occurs in fishes has long been a contentious issue, but recent observations document that social, object, and locomotor play can all be found in some species of teleosts. However, quantitative studies and those documenting individual differences are rare. We recorded hundreds of occurrences of an unusual behavior in three male Tropheus duboisi. The target behavior of attacking and deflecting an object that rapidly returned to its upright position not only fit the criteria for play behavior, but differed quantitatively and qualitatively among the individuals. This behavior has not been observed in other species of cichlids and other kinds of fishes. The presence or absence of food or other fish either within the aquarium or visible in an adjacent aquarium had no marked or consistent effect on the occurrence of the behavior. Various explanations for the origin and function of the behavior are discussed.
Social play behavior Social play behavior is the most vigorous and characteristic form of social interaction displayed by developing mammals. The laboratory rat is an ideal species to study this behavior, since it shows ample social play that can be easily recognized and quantified. In this chapter, we will first briefly describe the structure of social play behavior in rats. Next, we will discuss studies that used social isolation rearing during the period in life when social play is most abundant to investigate the developmental functions of social play behavior in rats, focusing on the consequences of play deprivation on social, cognitive, emotional, and sensorimotor development. Last, we will discuss the neural substrates of social play behavior in rats, with emphasis on the limbic corticostriatal circuits that underlie emotions and their influence on behavior.
Laboratory rats have been widely used to study the development and neural underpinnings of play behavior. However, it is not known whether domestic rats play in the same way and at the same frequency as their wild counterparts. In this study, the play of juvenile rats from a colony of wild rats maintained in captivity was compared to that of a strain of domesticated rats (e.g., Long Evans hooded). Three predictions were tested. First, it was predicted that wild rats would incorporate more agonistic behavior in their play. This was not found, as in all cases, both the wild and the laboratory rats attacked and defended the nape during play, a nonagonistic body target. Second, because play is typically more frequent in domesticated animals than their wild progenitors, it was predicted that the wild rats should play less than the laboratory rats. This was found to be the case. Third, because wild animals tend to be less tolerant of proximity by conspecifics and tend to be more agile in their movements, it was predicted that there would be less contact between wild pair mates. This was found to be the case; data show that the play of laboratory rats involves the same target (i.e., the nape of the neck) and tactics of defense as those used by wild rats. However, the laboratory rats initiated playful attacks more frequently, and were more likely to use tactics that promoted bodily contact. These similarities and differences need to be considered when using laboratory animals as models for play in general. (PsycINFO Database Record (c) 2013 APA, all rights reserved).
The brain endocannabinoid system plays a crucial role in emotional processes. We have previously identified an important role for endocannabinoids in social play behavior, a highly rewarding form of social interaction in adolescent rats. Here, we tested the hypothesis that endocannabinoid modulation of social play behavior occurs in brain regions implicated in emotion and motivation. Social play increased levels of the endocannabinoid anandamide in the amygdala and nucleus accumbens (NAc), but not in prefrontal cortex or hippocampus of 4- to 5-week-old male Wistar rats. Furthermore, social play increased phosphorylation of CB1 cannabinoid receptors in the amygdala. Systemic administration of the anandamide hydrolysis inhibitor URB597 increased social play behavior, and augmented the associated elevation in anandamide levels in the amygdala, but not the NAc. Infusion of URB597 into the basolateral amygdala (BLA) increased social play behavior, and blockade of BLA CB1 cannabinoid receptors with the antagonist/inverse agonist SR141716A prevented the play-enhancing effects of systemic administration of URB597. Infusion of URB597 into the NAc also increased social play, but blockade of NAc CB1 cannabinoid receptors did not antagonize the play-enhancing effects of systemic URB597 treatment. Last, SR141716A did not affect social play after infusion into the core and shell subregions of the NAc, while it reduced social play when infused into the BLA. These data show that increased anandamide signaling in the amygdala and NAc augments social play, and identify the BLA as a prominent site of action for endocannabinoids to modulate the rewarding properties of social interactions in adolescent rats.
The combination of exploration, habituation, and exploratory play was investigated in octopuses (
Octopus dofleini). Eight octopuses were given 10 trials to investigate a floating pill bottle. Exploration consisted of palpation of the object with the arms. Habituation was noticeable in the 1st trial but was more complex across trials. Two octopuses appeared to show exploratory play. This play consisted of aiming water jets through the flexible funnel, which caused regular transport of the object to and return by the aquarium intake current. In this situation, the amount of the 3 activities appeared to be minimally correlated. The results raise questions about the course of habituation, the definition and the extent of play, and the relation of exploratory play to exploration in complex animals. (PsycINFO Database Record (c) 2012 APA, all rights reserved)
Traces the historical background of the field of behavioral studies from the 17th-century works of Rene Descartes to the present. An overview of the book is included, stressing problems in experimental methods and interpretation of empirical data. (29 ref.) (PsycINFO Database Record (c) 2012 APA, all rights reserved)
The dog was the first domesticated animal but it remains uncertain when the domestication process began and whether it occurred just once or multiple times across the Northern Hemisphere. To ascertain the value of modern genetic data to elucidate the origins of dog domestication, we analyzed 49,024 autosomal SNPs in 1,375 dogs (representing 35 breeds) and 19 wolves. After combining our data with previously published data, we contrasted the genetic signatures of 121 breeds with a worldwide archeological assessment of the earliest dog remains. Correlating the earliest archeological dogs with the geographic locations of 14 so-called "ancient" breeds (defined by their genetic differentiation) resulted in a counterintuitive pattern. First, none of the ancient breeds derive from regions where the oldest archeological remains have been found. Second, three of the ancient breeds (Basenjis, Dingoes, and New Guinea Singing Dogs) come from regions outside the natural range of Canis lupus (the dog's wild ancestor) and where dogs were introduced more than 10,000 y after domestication. These results demonstrate that the unifying characteristic among all genetically distinct so-called ancient breeds is a lack of recent admixture with other breeds likely facilitated by geographic and cultural isolation. Furthermore, these genetically distinct ancient breeds only appear so because of their relative isolation, suggesting that studies of modern breeds have yet to shed light on dog origins. We conclude by assessing the limitations of past studies and how next-generation sequencing of modern and ancient individuals may unravel the history of dog domestication.
The effects of the selective 5HT(1A) agonist 8-OH-DPAT were assessed on the play behavior of juvenile rats. When both rats of the test pair were comparably motivated to play, the only significant effect of 8-OH-DPAT was for play to be reduced at higher doses. When there was a baseline asymmetry in playful solicitation due to a differential motivation to play and only one rat of the pair was treated, low doses of 8-OH-DPAT resulted in a collapse of asymmetry in playful solicitations. It did not matter whether the rat that was treated initially accounted for more nape contacts or fewer nape contacts, the net effect of 8-OH-DPAT in this model was for low doses of 8-OH-DPAT to decrease a pre-established asymmetry in play solicitation. It is concluded that selective stimulation of 5HT(1A) receptors changes the dynamic of a playful interaction between two participants that are differentially motivated to play. These results are discussed within a broader framework of serotonergic involvement in mammalian playfulness.
Studying play behavior in octopuses is an important step toward understanding the phylogenetic origins and function of play as well as the cognitive abilities of invertebrates. Fourteen Octopus vulgaris (7 subadults and 7 adults) were presented 2 Lego objects and 2 different food items on 7 consecutive days under 2 different levels of food deprivation. Nine subjects showed play-like behavior with the Lego objects. There was no significant difference in play-like behavior corresponding to food deprivation, age, and sex of the octopuses. The sequence of behaviors, from exploration to play-like behavior, had a significant influence on the establishment of play-like behavior, as it occurred mostly on Days 3-6 of the 7-day experiment. The pattern of development of play-like activities after a period of exploration and habituation in this study agrees with the hypothesis that object play follows object exploration. A homologous origin of this behavioral trait in vertebrates and invertebrates is highly unlikely, as the last common ancestor might not have had the cognitive capacity to possess this trait.
Reptile species are rarely the subjects of environmental enrichment programs, and there is little data available in the literature from enrichment studies involving these species. The authors share their studies of enriching the environments of Nile soft-shelled turtles.
Why do animals play? Play has been described in animals as diverse as reptiles, birds and mammals, so what benefits does it provide and how did it evolve? Careful, quantitative studies of social, locomotor and object play behaviour are now beginning to answer these questions and to shed light on many other aspects of both animal and human behaviour. This interdisciplinary volume, first published in 1998, brings together the major findings about play in a wide range of species including humans. Topics about play include the evolutionary history of play, play structure, function and development, and sex and individual differences. Animal Play is destined to become the benchmark volume in this subject, and will provide a source of inspiration and understanding for students and researchers in behavioural biology, neurobiology, psychology and anthropology.
The main aim of this book is to provide a basis for a complete dog behavioural biology based on concepts derived from contemporary ethology. Thus, dog behaviour is viewed from both functional (evolution and ecology) and mechanistic and developmental points of view. The study of dogs is placed in a comparative context which involves comparison with their ancestors (wolves), as well as with humans with which dogs share their present environment. Instead of advocating a single theory which would explain the emergence of dogs during the last 20,000 years of human evolution, this book gives an overview of present knowledge which has been collected by scientists from various fields. It aims to find novel ways to increase our understanding of this complex evolutionary process by combining different methods originating from different scientific disciplines. This is facilitated by describing complementing knowledge provided by various field of science, including zooarchaeology, cognitive and comparative ethology, human-animal interaction, behaviour genetics, behavioural physiology and development, and behavioural ecology. This interdisciplinary approach to the study of dogs deepens our biological understanding of dog behaviour, but also utilizes this knowledge to reveal secrets to behavioural evolution in general, even with special reference to the human species.
Cephalopods are generally regarded as the most intelligent group among the invertebrates. Despite their popularity, relatively little is known about the range and function of their cognitive abilities. This book fills that gap, accentuating the varied and fascinating aspects of cognition across the group. Starting with the brain, learning and memory, Part I looks at early learning, memory acquisition and cognitive development in modern cephalopods. An analysis of the chambered nautilus, a living fossil, is included, providing insight into the evolution of behavioural complexity. Part II surveys environmental responses, especially within the active and learning-dependent coleoids. The ever-intriguing camouflage abilities of octopus and cuttlefish are highlighted, alongside bioluminescence, navigation and other aspects of visual and cognitive competence. Covering the range of cognitive function, this text underscores the importance of the cephalopods within the field of comparative cognition generally. It will be highly valuable for researchers, graduates and senior undergraduate students.
Four infant coyotes (Canis latrans) were studied in order to describe quantitatively the development of predatory behaviour. Our results indicated that prior play and agonistic experience had virtually no effect on later predatory success. Also, there was no relationship between an individual's social rank and its prey-killing ability. Latency to kill was shortened when animals were tested in, pairs and hunger level was not related to latency to kill. The results are discussed with respect to current ‘functionalist’ theories of play behaviour and Leyhausen's concept of the relative hierarchy of moods. The practice theory of play should be reconsidered in light of the results of this and other recent studies.
Twenty-four free-ranging dog puppies belonging to six litters were observed from birth to 13 weeks of age to study the play behaviour in early ontogeny. Only a single annual breeding cycle with synchronized breeding season was recorded in this study. Mean litter size was 6.67 ranging from 5 to 8 with a male-biased sex ratio of 1.22:1. Social investigation was first observed with 3 weeks of age, and then subsequently developed other play behaviours (play-fighting, play-mounting, aggressive play, objects play and pseudo-sexual play). The litters were significantly different from each other in relation to the number of total play bouts (χ2=475.42, d.f.=5, P
Toys are often provided for adult dogs housed in kennels, but their effectiveness as environmental enrichment is not well documented. At a minimum, toys need to elicit interest in the animal for which they are intended, before any “enrichment” can be claimed. In this study we have examined short-term preferences for toys with a range of characteristics, using two methods of presentation, in both long-stay dogs in complex kennels, and short-stay dogs in rehoming kennels. The dogs, one sample in residential kennels (LSE, N=30) and the other in rehoming kennels (RH, N=66), were tested individually with four robust toys, presented both hanging and on the floor, over two 15min trials. The trial was also repeated with a second RH sample (N=34) comparing the four robust toys with less robust toys, all presented on the floor. Latency to and duration of interaction with each toy were recorded remotely. In the first trial, 34% of RH dogs and 43% of LSE dogs interacted with the toys; of the dogs that interacted, the average duration of interaction was higher among RH dogs (120s) than among LSE dogs (28s). Toys on the floor were interacted with for significantly longer than hanging toys by both LSE and RH dogs. RH dogs were faster to interact with the floor toys than the hanging toys, but the LSE dogs did not appear to discriminate between hanging and floor toys in latencies to interact. In the second trial, 76% of the RH dogs interacted with one or more of the toys, interacting for significantly longer with the four less robust toys, but their latencies to interact were similar between the robust and less robust toys. Average duration of interaction (227s) was higher than in the first trial. Our findings support previous proposals that robust toys are little used by kennel housed dogs. However, with less robust toys, interaction was relatively prolonged, indicating that interest to the dog may be enhanced if the toy can be chewed easily and/or makes a noise. Hanging toys were not favoured, although these have been reported to stimulate high levels of interaction in juvenile laboratory beagles.
The purpose of this chapter is to present a general framework for studying the development of behavior. The thesis to be defended
here is that the building blocks of behavior are various kinds of perceptual, central, and motor components, all of which
can exist independently. The study of development is primarily the study of changes in these components themselves and in
the connections among them
Animal social play represents an important tool for self- and social-assessment purposes during the juvenile phase. Nevertheless, this activity may continue into adulthood as well providing immediate benefits to the playmates. In this study, I investigated the dynamics of adult play in a wolf colony hosted at the Pistoia Zoo (Italy). The study wolves performed social play to a greater extent compared to solitary play. Play distribution was not affected by relationship quality (measured by body contact and agonistic support frequencies) and aggression levels. Probably, in wolves other behavioural strategies are employed for strengthening inter-individual relationships and reducing conflicts among fellows. Play was distributed throughout the entire group independently of the sex of playmates. The absence of sexual-dimorphism in play may be linked to the fact that in the wolf pack males and females share the same roles and behavioural repertoire. Rank distance between conspecifics negatively correlated with play distribution: by playing wolves with closest ranking positions tested each other for acquiring information on skills of possible competitor and gaining hierarchical advantage over it. Finally, in agreement with previous studies, my findings showed that wolves significantly reduced their playful activity during contests of high conflict of interests such as mating period and feeding time.
In this study we systematically investigate the mode of acquisition and the developmental process of stone handling, a form of solitary object play, in a captive troop of Japanese macaques (Macaca fuscata) housed in an outdoor enclosure at the Kyoto University Primate Research Institute, Japan. This study was conducted to evaluate two alternative hypotheses regarding the mode of acquisition of stone handling in infants: (1) environmental stimuli (availability of and exposure to stones) and (2) social stimuli (exposure to stone handling individuals). Early exposure to stones in the environment had no significant effect on when infants acquired the behavior. No significant correlations were recognized between the age of stone handling acquisition and number of stones encountered per hour from birth to acquisition, or the time spent in a specific area of the enclosure as a function of the number of available stones therein. However, being exposed to a stone handling model(s) was a social stimulus that had an effect on the age of acquisition, with a significant negative correlation between a mother's stone handling frequency and the age of acquisition by her infant. Infants of non-stone handling mothers acquired the behavior much later than others. Infant peers who acquired stone handling earlier played no significant role as stone handling models. Of the factors tested here, the timing of acquisition depended mainly on the level of proximity to a demonstrator and the frequency at which those available demonstrators performed the behavior.
Lorenz describes the nature and program of comparative ethology, which may be defined as the comparative study of innate behavior. After criticizing the reciprocal errors made by workers in other fields with respect to patterns of behavior, directedness, spontaneity, and innate behavior, he emphasized the necessity of a through inventory of –iall–n behavior patterns at the disposal of a given species as a starting point for meaningful research. Next is application of the true comparative method, a study of "the similarities and dissimilarities of homologous characters of allied forms… ." 46 references. (PsycINFO Database Record (c) 2012 APA, all rights reserved)
Most studies on animal motivation have been confined to the biological or homeostatic drives that were discussed in the previous chapter. This emphasis on drives such as hunger and thirst has been a source of criticism leveled at animal psychologists by those interested in human motivation. The argument is that most human behavior does not seem to be directed toward the satisfaction of basic biological needs. In our culture, severe hunger or thirst, for example, is an atypical condition. Society is organized to provide for the fundamental necessities of life. Rather than basing a theory of human motivation on the drives investigated in the animal laboratory, some psychologists have proposed a different level of conceptualization. Many motivational theorists, for instance, maintain that the need for self-actualization, the need to know and understand, the need for belongingness and love are vastly more important in modern society than are the biological needs. At first glance it may appear that animal research can make little or no contribution to a comprehensive theory of motivation, since, according to these theoreticians, the motives of man are entirely different from those of lower animals. It should be made perfectly clear, however, that animal behavior is not based exclusively on the so-called "primary biological drives." These particular drives are most frequently utilized in the laboratory because they can be controlled while the investigator focuses his attention on some other phenomenon, and for decades it usually has been the phenomenon of learning. Indeed, nearly every important problem in learning can be studied by observing how a hungry rat, cat, or monkey obtains food under different experimental conditions. It is almost as if the motivation of animals has been exploited rather than investigated. But recently some animal psychologists have become interested in motivation as an area of investigation in its own right, and their research shows decidedly that motivational mechanisms other than the primary biological drives are important determinants of behavior. Others, while working on the problem of motivation, are essentially extending the previous research on learning. This chapter will describe some experiments resulting from these two different approaches. (PsycINFO Database Record (c) 2012 APA, all rights reserved)
Domestic dogs are reported to show intense but transient neophilia towards novel objects. Here, we examine habituation and dishabituation to manipulable objects by kennel-housed dogs. Labrador retrievers (N = 16) were repeatedly presented with one toy for successive 30-s periods until interaction ceased. At this point (habituation), a different toy was presented that contrasted with the first in both colour and odour (since the dog's saliva would have accumulated on the first), colour alone, or odour alone. No effect of the type of contrast was detected in the number of presentations to habituation, the difference in duration of interaction between the first presentation of the first toy and the presentation of the second toy (recovery), or the duration of interaction with the second toy (dishabituation). Varying the time interval between successive presentations of the first toy up to habituation between 10 s and 10 min had no effect on the number of presentations to habituation, nor did it alter the extent of dishabituation. Varying the delay from habituation to presentation of the second toy, between 10 s and 15 min, affected neither the recovery nor the dishabituation. Overall, the study indicates that loss of interest in the object during object-orientated play in this species is due to habituation to the overall stimulus properties of the toy rather than to any single sensory modality and is also atypical in its insensitivity to the interval between presentations.
Behaviours that always appear playful (play markers) are distinguished from behaviours that appear playful in some contexts, but not others (context-dependent play components). Age changes in the frequency of performance of both kinds of playful behaviours are described, as are age changes in the frequency with which context-dependent play components accompany play markers in baboon social interactions. Some quantitative properties of social interactions containing and lacking play markers are compared. Interactions with play markers last longer and have a higher rate of change of constituent behaviours than interactions without. Animals are also more persistent in the face of changes in their partner's behaviour in interactions with play markers. It is suggested that an attempt to produce a definition of play is not a useful enterprise, but that it is important to investigate causal mechanisms underlying behaviours that appear playful to human observers, and to clarify the relationships among those mechanisms. Data are presented suggesting that at least two sets of mechanisms, not totally separate, underly the performance of baboon social play. Functions of the behaviours controlled by these mechanisms are discussed.
Play behavior is a fundamental and intrinsic neurobehavioral process in the mammalian brain. Using rough-and-tumble play in the juvenile rat as a model system to study mammalian playfulness, some of the relevant neurobiological substrates for this behavior have been identified, and in this review this progress. A primary-process executive circuit for play in the rat that includes thalamic intralaminar nuclei, frontal cortex and striatum can be gleaned from these data. Other neural areas that may interact with this putative circuit include amygdala, ventral hypothalamus, periaqueductal gray (PAG), and deep tectum, as well as ascending dopamine systems which participate in all types of seeking urges. At the neurochemical level, considerable evidence points to specific cholinergic and dopaminergic controls, but also endogenous opioids and cannabinoids as having a positive modulatory influence over playfulness, with all neuropeptides known to have aversive effects to reduce play. Monoamines such as norepinephrine and serotonin certainly modulate play, but they influence all psychobehavioral systems, suggesting non-specific effects. We proceed to discuss how increased insights into the neurobiological mechanisms of play can inform our understanding of normal and abnormal childhood development.
Juvenile Fischer 344 rats are known to be less playful than other inbred strains, although the neurobiological substrate(s) responsible for this phenotype is uncertain. In the present study, Fischer 344 rats were compared to the commonly used outbred Sprague-Dawley strain on several behavioral and physiological parameters in order to ascertain whether the lack of play may be related to compromised activity of brain dopamine (DA) systems. As expected, Fischer 344 rats were far less playful than Sprague-Dawley rats, with Fischer 344 rats less likely to initiate playful contacts with a playful partner and less likely to respond playfully to these contacts. We also found that Fischer 344 rats showed less of a startle response and greater pre-pulse inhibition (PPI), especially at higher pre-pulse intensities. The increase in PPI seen in the Fischer 344 rat could be due to reduced DA modulation of sensorimotor gating and neurochemical measures were consistent with Fischer 344 rats releasing less DA than Sprague-Dawley rats. Fast scan cyclic voltammetry (FSCV) revealed Fischer 344 rats had less evoked DA release in dorsal and ventral striatal brain slices and high-performance liquid chromatography revealed Fischer 344 rats to have less DA turnover in the striatum and prefrontal cortex. We also found DA-dependent forms of cortical plasticity were deficient in the striatum and prefrontal cortex of the Fischer 344 rat. Taken together, these data indicate that deficits in play and enhanced PPI of Fischer 344 rats may be due to reduced DA modulation of corticostriatal and mesolimbic/mesocortical circuits critical to the execution of these behaviors.
Biologists, breeders and trainers, and champion sled dog racers, Raymond and Lorna Coppinger have more than four decades of experience with literally thousands of dogs. Offering a scientifically informed perspective on canines and their relations with humans, the Coppingers take a close look at eight different types of dogsâhousehold, village, livestock guarding, herding, sled-pulling, pointing, retrieving, and hound. They argue that dogs did not evolve directly from wolves, nor were they trained by early humans; instead they domesticated themselves to exploit a new ecological niche: Mesolithic village dumps. Tracing the evolution of today's breeds from these village dogs, the Coppingers show how characteristic shapes and behaviorsâfrom pointing and baying to the sleek shapes of running dogsâarise from both genetic heritage and the environments in which pups are raised. For both dogs and humans to get the most out of each other, we need to understand and adapt to the biological needs and dispositions of our canine companions, just as they have to ours.
A drive-reduction theory of learning, emphasizing the internal physiological state, is untenable. Learning efficiency is far better related to "tensions in the brain than in the belly." The key to human learning is motivation aroused by external stimulation. 35 references.
Two rhesus monkeys, given 60 two-hour sessions with a six-device mechanical puzzle showed clear evidence of learning, the curve showing ratio of incorrect to correct responses appearing quite comparable to similar curves obtained during externally rewarded situations. When, on the thirteenth day of tests, the subjects were presented with the puzzle 100 times at 6-minute intervals, the number of devices manipulated decreased regularly throughout the day, although there was no significant change in the number of times the problem assembly was attacked.
Stone handling (SH) in Japanese macaques, a form of solitary-object play, is newly acquired only by young individuals, and is the first example of a directly nonadaptive behavior that is maintained as a behavioral tradition within free-ranging provisioned social troops. We report here the first systematic investigation of this behavior in a stable captive social troop, the Takahama troop, which is housed in an outdoor enclosure of the Primate Research Institute (PRI), Kyoto University, Japan. This study was conducted to evaluate relevant competing hypotheses regarding the function of object play (e.g., misdirected foraging behavior and motor training) to explain the proximal causes and ultimate function(s) of SH. The "misdirected foraging behavior" hypothesis can be ruled out because of the lack of a clear temporal relationship between feeding and the occurrence of SH in any age class. Age-related differences in SH performance and behavioral patterns were observed, suggesting possible differences in the immediate cause and ultimate function between young and adults. Young individuals engaged in frequent bouts of short duration, involving locomotion and vigorous body actions throughout the day, which is typical for play by young in general. This pattern of behavior is consistent with the motor training hypothesis, which states that play occurs during the development of motor and perceptual skills and is thus potentially critical for neural and cognitive development. This practice is continued by those who acquire it at an early age, with adults engaging in significantly fewer but longer bouts that involve more stationary, complex manipulative patterns, almost exclusively in the late afternoon. We propose that for adults, at the proximate level SH is psychologically relaxing, but ultimately functions to maintain and regenerate neural pathways, and potentially helps to slow down the deterioration of cognitive function associated with advanced age in long-lived provisioned and captive macaques.