Duration of cats' (Felis catus) working memory for disappearing objects

This study explored the duration of cats' working memory for hidden objects. Twenty-four cats were equally divided into four groups, which differed according to the type of visual cues displayed on and/or around the hiding boxes. During eight sessions, the four groups of cats were trained to locate a desirable object hidden behind one of the four boxes placed in front of them. Then, the cats were tested with retention intervals of 0, 10, 30 and 60 s. Results revealed no significant differences between the groups during training or testing. In testing, the cats' accuracy to locate the hidden object rapidly declined between 0 and 30 s but remained higher than chance with delays of up to 60 s. The analysis of errors also indicated that the cats searched as a function of the proximity of the target box and were not subjected to intertrial proactive interference. This experiment reveals that the duration of cats' working memory for disappearing objects is limited and the visual cues displayed on and/or around the boxes do not help the cats to memorize a hiding position. In discussion, we explore why the duration of cats' working memory for disappearing objects rapidly declined and compare these finding with those from domestic dogs. The irrelevance of visual cues displayed on and around the hiding boxes on cats' retention capacity is also discussed.     

Invisible displacement understanding in domestic dogs (<Emphasis Type="Italic">Canis familiaris</Emphasis>): the role of visual cues in search behavior

Recently, (Collier-Baker E, Davis JM, Suddendorf T (2004) J Comp Psychol 118:421–433) suggested that domestic dogs do not understand invisible displacements. In the present study, we further investigated the hypothesis that the search behavior of domestic dogs in invisible displacements is guided by various visual cues inherent to the task rather than by mental representation of an object’s past trajectory. Specifically, we examined the role of the experimenter as a function of the final position of the displacement device in the search behavior of domestic dogs. Visible and invisible displacement problems were administered to dogs (N = 11) under two conditions. In the Visible-experimenter condition, the experimenter was visible whereas in the Concealed-experimenter condition, the experimenter was visibly occluded behind a large rigid barrier. Our data supported the conclusion that dogs do not understand invisible displacements but primarily search as a function of the final position of the displacement device and, to a lesser extent, the position of the experimenter.     

Deceptive-like behaviour in dogs (<Emphasis Type="Italic">Canis familiaris</Emphasis>)

Deception, the use of false signals to modify the behaviour of the receiver, occurs in low frequencies even in stable signalling systems. For example, it can be advantageous for subordinate individuals to deceive in competitive situations. We investigated in a three-way choice task whether dogs are able to mislead a human competitor, i.e. if they are capable of tactical deception. During training, dogs experienced the role of their owner, as always being cooperative, and two unfamiliar humans, one acting ‘cooperatively’ by giving food and the other being ‘competitive’ and keeping the food for themselves. During the test, the dog had the options to lead one of these partners to one of the three potential food locations: one contained a favoured food item, the other a non-preferred food item and the third remained empty. After having led one of the partners, the dog always had the possibility of leading its cooperative owner to one of the food locations. Therefore, a dog would have a direct benefit from misleading the competitive partner since it would then get another chance to receive the preferred food from the owner. On the first test day, the dogs led the cooperative partner to the preferred food box more often than expected by chance and more often than the competitive partner. On the second day, they even led the competitive partner less often to the preferred food than expected by chance and more often to the empty box than the cooperative partner. These results show that dogs distinguished between the cooperative and the competitive partner, and indicate the flexibility of dogs to adjust their behaviour and that they are able to use tactical deception.     

Global bias reliability in dogs (<Emphasis Type="Italic">Canis familiaris</Emphasis>)

Dogs enrolled in a previous study were assessed two years later for reliability of their local/global preference in a discrimination test with the same hierarchical stimuli used in the previous study (Experiment 1) and with a novel stimulus (Experiment 2). In Experiment 1, dogs easily re-learned to discriminate the positive stimulus; their individual global/local choices were stable compared to the previous study; and an overall clear global bias was found. In Experiment 2, dogs were slower in acquiring the initial discrimination task; the overall global bias disappeared; and, individually, dogs tended to make inverse choices compared to the original study. Spontaneous attention toward the test stimulus resembling the global features of the probe stimulus was the main factor affecting the likeliness of a global choice of our dogs, regardless of the type of experiment. However, attention to task-irrelevant elements increased at the expense of attention to the stimuli in the test phase of Experiment 2. Overall, the results suggest that the stability of global bias in dogs depends on the characteristics of the assessment contingencies, likely including the learning requirements of the tasks. Our results also clearly indicate that attention processes have a prominent role on dogs’ global bias, in agreement with previous findings in humans and other species.     

Reputation-like inference in domestic dogs (<Emphasis Type="Italic">Canis familiaris</Emphasis>)

Humans frequently interact with strangers absent prior direct experience with their behavior. Some conjecture that this may have favored evolution of a cognitive system within the hominoid clade or perhaps the primate order to assign reputations based on third-party exchanges. However, non-primate species’ acquisition of skills from experienced individuals, attention to communicative cues, and propensity to infer social rules suggests reputation inference may be more widespread. We utilized dogs’ sensitivity to humans’ social and communicative cues to explore whether dogs evidenced reputation-like inference for strangers through third-party interactions. Results indicated dogs spontaneously show reputation-like inference for strangers from indirect exchanges. Further manipulations revealed that dogs continued to evidence this ability despite reduction of specific components of the observed interactions, including reduction of visual social cues (i.e., face-to-face contact between the participants in the interaction) and the nature of the recipient (i.e., living, animate agent versus living, inanimate self-propelled agent). Dogs also continued to demonstrate reputation-like inference when local enhancement was controlled and in a begging paradigm. However, dogs did not evidence reputation-like inference when the observed interaction was inadvertent.     

Comprehension of human pointing gestures in young human-reared wolves (<Emphasis Type="Italic">Canis lupus</Emphasis>) and dogs (<Emphasis Type="Italic">Canis familiaris</Emphasis>)

Dogs have a remarkable skill to use human-given cues in object-choice tasks, but little is known to what extent their closest wild-living relative, the wolf can achieve this performance. In Study 1, we compared wolf and dog pups hand-reared individually and pet dogs of the same age in their readiness to form eye-contact with a human experimenter in an object-choice task and to follow her pointing gesture. The results showed that dogs already at 4 months of age use momentary distal pointing to find hidden food even without intensive early socialization. Wolf pups, on the contrary, do not attend to this subtle pointing. Accordingly in Studies 2 and 3, these wolves were tested longitudinally with this and four other (easier) human-given cues. This revealed that wolves socialized at a comparable level to dogs are able to use simple human-given cues spontaneously if the human’s hand is close to the baited container (e.g. touching, proximal pointing). Study 4 showed that wolves can follow also momentary distal pointing similarly to dogs if they have received extensive formal training. Comparing the wolves to naïve pet dogs of the same age revealed that during several months of formal training wolves can reach the level of dogs in their success of following momentary distal pointing in parallel with improving their readiness to form eye-contact with a human experimenter. We assume that the high variability in the wolves’ communicative behaviour might have provided a basis for selection during the course of domestication of the dog.     

Quantity-based judgments in the domestic dog (<Emphasis Type="Italic">Canis lupus familiaris</Emphasis>)

We examined the ability of domestic dogs to choose the larger versus smaller quantity of food in two experiments. In experiment 1, we investigated the ability of 29 dogs (results from 18 dogs were used in the data analysis) to discriminate between two quantities of food presented in eight different combinations. Choices were simultaneously presented and visually available at the time of choice. Overall, subjects chose the larger quantity more often than the smaller quantity, but they found numerically close comparisons more difficult. In experiment 2, we tested two dogs from experiment 1 under three conditions. In condition 1, we used similar methods from experiment 1 and tested the dogs multiple times on the eight combinations from experiment 1 plus one additional combination. In conditions 2 and 3, the food was visually unavailable to the subjects at the time of choice, but in condition 2, food choices were viewed simultaneously before being made visually unavailable, and in condition 3, they were viewed successively. In these last two conditions, and especially in condition 3, the dogs had to keep track of quantities mentally in order to choose optimally. Subjects still chose the larger quantity more often than the smaller quantity when the food was not simultaneously visible at the time of choice. Olfactory cues and inadvertent cuing by the experimenter were excluded as mechanisms for choosing larger quantities. The results suggest that, like apes tested on similar tasks, some dogs can form internal representations and make mental comparisons of quantity.     

A test of object permanence in a new-world monkey species,cotton top tamarins (<Emphasis Type="Italic">Saguinus oedipus</Emphasis>)

Cotton top tamarins were tested in visible and invisible displacement tasks in a method similar to that used elsewhere to test squirrel monkeys and orangutans. All subjects performed at levels significantly above chance on visible (n=8) and invisible (n=7) displacements, wherein the tasks included tests of the perseverance error, tests of memory in double and triple displacements, and "catch" trials that tested for the use of the experimenter's hand as a cue for the correct cup. Performance on all nine tasks was significantly higher than chance level selection of cups, and tasks using visible displacements generated more accurate performance than tasks using invisible displacements. Performance was not accounted for by a practice effect based on exposure to successive tasks. Results suggest that tamarins possess stage 6 object permanence capabilities, and that in a situation involving brief exposure to tasks and foraging opportunities, tracking objects' movements and responding more flexibly are abilities expressed readily by the tamarins. Electronic Publication     

Cross-species referential signalling events in domestic dogs (<Emphasis Type="Italic">Canis familiaris</Emphasis>)

Referential gestures are used by a signaller to draw a recipient’s attention to a specific object, individual or event in the environment. These gestures have received much research attention in relation to human and non-human primates with great apes being shown to possess impressive gestural repertoires. Domestic dogs (Canis familiaris) meanwhile provide an ideal non-primate candidate for investigating referential signalling due to their unique relationship with humans that centres on non-verbal communication with frequent interaction. Here we observed 37 pet dogs in their own homes. Owners recorded 242 videos containing 47 potential referential gesture events. We analysed those recordings to reveal evidence of 19 referential gestures performed by domestic dogs during everyday communicative bouts with humans, showing that the gestures conform to the five features of referential signalling. Our study exposes impressive gesturing abilities in a non-primate mammal; especially when viewed in the context of the cross-species rather than intraspecific communication.     

Piagetian object permanence and its development in Eurasian jays (<Emphasis Type="Italic">Garrulus glandarius</Emphasis>)

Object permanence in Eurasian jays (Garrulus glandarius) was investigated using a complete version of the Uzgiris and Hunt scale 1. Nine hand-raised jays were studied, divided into two groups according to their different developmental stages (experiment 1, older jays: 2–3 months old, n = 4; experiment 2, younger jays: 15 days old, n = 5). In the first experiment, we investigated whether older jays could achieve piagetian stage 6 of object permanence. Tasks were administered in a fixed sequence (1–15) according to the protocols used in other avian species. The aim of the second experiment was to check whether testing very young jays before their development of “neophobia” could influence the achievement times of piagetian stages. Furthermore, in this experiment tasks were administered randomly to investigate whether the jays’ achievement of stage 6 follows a fixed sequence related to the development of specific cognitive abilities. All jays tested in experiments 1 and 2 fully achieved piagetian stage 6 and no “A not B” errors were observed. Performance on visible displacement tasks was better than performance on invisible ones. The results of experiment 2 show that “neophobia” affected the response of jays in terms of achievement times; the older jays in experiment 1 took longer to pass all the tasks when compared with the younger, less neophobic, jays in experiment 2. With regard to the achieving order, jays followed a fixed sequence of acquisition in experiment 2, even if tasks were administered randomly, with the exception of one subject. The results of these experiments support the idea that piagetian stages of cognitive development exist in avian species and that they progress through relatively fixed sequences. Electronic Supplementary Material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Short-term observational spatial memory in Jackdaws (<Emphasis Type="Italic">Corvus monedula</Emphasis>) and Ravens (<Emphasis Type="Italic">Corvus corax</Emphasis>)

Observational spatial memory (OSM) refers to the ability of remembering food caches made by other individuals, enabling observers to find and pilfer the others' caches. Within birds, OSM has only been demonstrated in corvids, with more social species such as Mexican jays (Aphelocoma ultramarine) showing a higher accuracy of finding conspecific' caches than less social species such as Clark's nutcrackers (Nucifraga columbiana). However, socially dynamic corvids such as ravens (Corvus corax) are capable of sophisticated pilfering manoeuvres based on OSM. We here compared the performance of ravens and jackdaws (Corvus monedula) in a short-term OSM task. In contrast to ravens, jackdaws are socially cohesive but hardly cache and compete over food caches. Birds had to recover food pieces after watching a human experimenter hiding them in 2, 4 or 6 out of 10 possible locations. Results showed that for tests with two, four and six caches, ravens performed more accurately than expected by chance whereas jackdaws did not. Moreover, ravens made fewer re-visits to already inspected cache sites than jackdaws. These findings suggest that the development of observational spatial memory skills is linked with the species' reliance on food caches rather than with a social life style per se.     

The effect of oxytocin on biological motion perception in dogs (<Emphasis Type="Italic">Canis familiaris</Emphasis>)

Recent studies have shown that the neuropeptide oxytocin is involved in the regulation of several complex human social behaviours. There is, however, little research on the effect of oxytocin on basic mechanisms underlying human sociality, such as the perception of biological motion. In the present study, we investigated the effect of oxytocin on biological motion perception in dogs (Canis familiaris), a species adapted to the human social environment and thus widely used to model many aspects of human social behaviour. In a within-subjects design, dogs (N = 39), after having received either oxytocin or placebo treatment, were presented with 2D projection of a moving point-light human figure and the inverted and scrambled version of the same movie. Heart rate (HR) and heart rate variability (HRV) were measured as physiological responses, and behavioural response was evaluated by observing dogs’ looking time. Subjects were also rated on the personality traits of Neuroticism and Agreeableness by their owners. As expected, placebo-pretreated (control) dogs showed a spontaneous preference for the biological motion pattern; however, there was no such preference after oxytocin pretreatment. Furthermore, following the oxytocin pretreatment female subjects looked more at the moving point-light figure than males. The individual variations along the dimensions of Agreeableness and Neuroticism also modulated dogs’ behaviour. Furthermore, HR and HRV measures were affected by oxytocin treatment and in turn played a role in subjects? looking behaviour. We discuss how these findings contribute to our understanding of the neurohormonal regulatory mechanisms of human (and non-human) social skills.     

Enhanced recognition of emotional stimuli in the chimpanzee (<Emphasis Type="Italic">Pan troglodytes</Emphasis>)

Humans often have a better memory of emotional events than neutral events. From the comparative-cognitive perspective, we explored the enhancement of recognition memory by emotion in chimpanzees (Pan troglodytes) using a serial probe recognition task. In this task, we sequentially presented a list of pictures to subjects and then tested their recognition of specific pictures from within the list. We selected pictures of aggressive chimpanzees as emotional stimuli and less tensed, relaxed chimpanzees as neutral stimuli. In Experiment 1, we gave four-item lists to two young chimpanzees; one showed significantly greater recognition of pictures depicting aggressive chimpanzees than neutral ones. In Experiment 2, this chimpanzee was further tested using a recognition task with eight-item lists. The subject again showed better recognition of emotional stimuli than neutral. Furthermore, the presence of an emotional stimulus in the list also facilitated recognition of the neutral item immediately following it. Overall, although only one of the two chimpanzees showed enhanced recognition memory by emotional stimuli, this is the first demonstration of such a response in the chimpanzee. The findings are discussed in comparison with those of human studies.     

Recognition of human faces by dogs (<Emphasis Type="Italic">Canis familiaris</Emphasis>) requires visibility of head contour

Researchers have suggested that dogs are able to recognise human faces, but conclusive evidence has yet to be found. Experiment 1 of this study investigated whether dogs can recognise humans using visual information from the face/head region, and whether this also occurs in conditions of suboptimal visibility of the face. Dogs were presented with their owner’s and a stranger’s heads, protruding through openings of an apparatus in opposite parts of the experimental setting. Presentations occurred in conditions of either optimal or suboptimal visibility; the latter featured non-frontal orientation, uneven illumination and invisibility of outer contours of the heads. Instances where dogs approached their owners with a higher frequency than predicted by chance were considered evidence of recognition. This occurred only in the optimal condition. With a similar paradigm, Experiment 2 investigated which of the alterations in visibility that characterised the suboptimal condition accounted for dogs’ inability to recognise owners. Dogs approached their owners more frequently than predicted by chance if outer head contours were visible, but not if heads were either frontally oriented or evenly illuminated. Moreover, male dogs were slightly better at recognition than females. These findings represent the first clear demonstration that dogs can recognise human faces and that outer face elements are crucial for such a task, complementing previous research on human face processing in dogs. Parallels with face recognition abilities observed in other animal species, as well as with human infants, point to the relevance of these results from a comparative standpoint.     

Long-term memory for concepts in a California sea lion (<Emphasis Type="Italic">Zalophus californianus</Emphasis>)

An adult California sea lion (Zalophus californianus) with extensive experience in performing discrimination learning tasks was tested to evaluate her long-term memory for two previously learned concepts. An associative concept, that of equivalence classification, was retested after a retention interval of approximately 1 year. The sea lion had originally shown emergent equivalence classification with nonsimilarity-based classes of stimuli in a simple discrimination repeated-reversal procedure as well as in a matching-to-sample procedure. The 1-year memory test revealed no decrement in classification performance in either procedure. A relational concept, that of generalized identity matching, was retested after approximately 10 years. The sea lion had originally received trial-and-error exemplar training with identity matching-to-sample problems prior to transferring the concept to novel stimulus configurations. In the 10-year memory test, the sea lion immediately and reliably applied the previously established identity concept to familiar and novel sets of matching problems. These are the first reports of long-term conceptual memory in a nonprimate species. The experimental findings are consistent with a variety of observations of sea lions in natural settings, which indicate that natal sites, feeding areas, and individuals may be remembered over long periods of time. Electronic Supplementary Material is available if you access this article at http://dx.doi.org/10.1007/s10071-002-0153-8. On that page (frame on the left side), a link takes you directly to the supplementary material. Electronic Publication     

Investigating emotional contagion in dogs (<Emphasis Type="Italic">Canis familiaris</Emphasis>) to emotional sounds of humans and conspecifics

Emotional contagion, a basic component of empathy defined as emotional state-matching between individuals, has previously been shown in dogs even upon solely hearing negative emotional sounds of humans or conspecifics. The current investigation further sheds light on this phenomenon by directly contrasting emotional sounds of both species (humans and dogs) as well as opposed valences (positive and negative) to gain insights into intra- and interspecies empathy as well as differences between positively and negatively valenced sounds. Different types of sounds were played back to measure the influence of three dimensions on the dogs’ behavioural response. We found that dogs behaved differently after hearing non-emotional sounds of their environment compared to emotional sounds of humans and conspecifics (“Emotionality” dimension), but the subjects responded similarly to human and conspecific sounds (“Species” dimension). However, dogs expressed more freezing behaviour after conspecific sounds, independent of the valence. Comparing positively with negatively valenced sounds of both species (“Valence” dimension), we found that, independent of the species from which the sound originated, dogs expressed more behavioural indicators for arousal and negatively valenced states after hearing negative emotional sounds. This response pattern indicates emotional state-matching or emotional contagion for negative sounds of humans and conspecifics. It furthermore indicates that dogs recognized the different valences of the emotional sounds, which is a promising finding for future studies on empathy for positive emotional states in dogs.     

Combinations of <Emphasis Type="Italic">Stit</Emphasis> with <Emphasis Type="Italic">Ought</Emphasis> and <Emphasis Type="Italic">Know</Emphasis>

This paper presents a short survey of recent developments in stit theories, with an emphasis on combinations of stit and deontic logic, and those of stit and epistemic logic.     

Precis of <Emphasis Type="Italic">Philosophy and Neuroscience</Emphasis>: <Emphasis Type="Italic">A Ruthlessly Reductive Account</Emphasis>

This book precis describes the motives behind my recent attempt to bring to bear “ruthlessly reductive” results from cellular and molecular neuroscience onto issues in the philosophy of mind. Since readers of this journal will probably be most interested in results addressing features of conscious experience, I highlight these most prominently. My main challenge is that philosophers (even scientifically-inspired ones) are missing the nature and scope of reductionism in contemporary neuroscience by focusing exclusively on higher-level cognitive neuroscience, and ignoring the discipline's cell-physiological and molecular-biological core.     

Interspecies transmission of emotional information via chemosignals: from humans to dogs (<Emphasis Type="Italic">Canis lupus familiaris</Emphasis>)

We report a study examining interspecies emotion transfer via body odors (chemosignals). Do human body odors (chemosignals) produced under emotional conditions of happiness and fear provide information that is detectable by pet dogs (Labrador and Golden retrievers)? The odor samples were collected from the axilla of male donors not involved in the main experiment. The experimental setup involved the co-presence of the dog’s owner, a stranger and the odor dispenser in a space where the dogs could move freely. There were three odor conditions [fear, happiness, and control (no sweat)] to which the dogs were assigned randomly. The dependent variables were the relevant behaviors of the dogs (e.g., approaching, interacting and gazing) directed to the three targets (owner, stranger, sweat dispenser) aside from the dogs’ stress and heart rate indicators. The results indicated with high accuracy that the dogs manifested the predicted behaviors in the three conditions. There were fewer and shorter owner directed behaviors and more stranger directed behaviors when they were in the “happy odor condition” compared to the fear odor and control conditions. In the fear odor condition, they displayed more stressful behaviors. The heart rate data in the control and happy conditions were significantly lower than in the fear condition. Our findings suggest that interspecies emotional communication is facilitated by chemosignals.     

Dogs (<Emphasis Type="Italic">Canis familiaris</Emphasis>) adjust their social behaviour to the differential role of inanimate interactive agents

Dogs are able to flexibly adjust their social behaviour to situation-specific characteristics of their human partner’s behaviour in problem situations. However, dogs do not necessarily detect the specific role played by the human in a particular situation: they may form expectations about their partners’ behaviour based on previous experiences with them. Utilising inanimate objects (UMO—unidentified moving object) as interacting agents offers new possibilities for investigating social behaviour, because in this way we can remove or control the influence of previous experience with the partner. The aim of the present study was to investigate whether dogs are able to recognise the different roles of two UMOs and are able to adjust their communicative behaviour towards them. In the learning phase of the experiment, dogs were presented with a two-way food-retrieval problem in which two UMOs, which differed in their physical appearance and abilities, helped the dog obtain a piece of food in their own particular manner. After a short experience with both UMOs, dogs in the test phase faced one of the problems in the presence of both inanimate agents. Overall, dogs displayed similar levels of gazing behaviour towards the UMOs, but in the first test they looked, approached and touched the relevant partner first. This rapid adjustment of social behaviour towards UMOs suggests that dogs may generalise their experiences with humans to unfamiliar agents and are able to select the appropriate partner when facing a problem situation.