Group size, partner choice and collaborative actions in male Indo-Pacific bottlenose dolphins (Tursiops aduncus).

Due to the diversity of the phenomenon, dolphin cooperation has attracted considerable research interest in both wild populations and those under human care. Dolphins cooperate in various contexts, including group hunting, alloparental care, social learning, social play and alliance formation for securing mates. This investigation focused on the effect of group size and partner choice in a cooperative task using systematic group testing. A cooperative enrichment device was made of a PVC tube containing fish and ice that was temporarily sealed with two PVC caps with rope handles attached. The device was designed to be operated by pairs of dolphins, opened by simultaneous pull of its two handles. The analysis focused on two behaviours, cooperative opening and cooperative play with the device. Testing focused on an adult male dolphin group including four to six individuals and using a single or two devices. Altogether five group testing arrangements and a pairwise testing phase were conducted. Out of the six dolphins, five showed active involvement. All ten possible pairs of the five active dolphins were successful in opening and playing with the device cooperatively. Cooperation increased with group size, but the social networks showed no significant differences among group arrangements. However, the cooperative pairs showed a significant difference in success rate during pairwise vs group testing, while demonstrating a strong partner preference. This study provides the first evidence for partner choice with regards to cooperation in male dolphins.

An exploratory analysis of head-tilting in dogs.

Little is known about head-tilts in dogs. Based on previous investigations on the head turning and the lateralised brain pattern of human speech processing in dogs, we hypothesised that head-tilts may be related to increased attention and could be explained by lateralised mental functions. We observed 40 dogs during object-label knowledge tests and analysed head-tilts occurring while listening to humans requesting verbally to fetch a familiar toy. Our results indicate that only dogs that had learned the name of the objects tilted their heads frequently. Besides, the side of the tilt was stable across several months and tests. Thus, we suggest a relationship between head-tilting and processing relevant, meaningful stimuli.

Testing use of the first multi-partner cognitive enrichment devices by a group of male bottlenose dolphins.

Alliance formation plays a crucial part in male dolphins' lives. These partnerships may last for decades or even for a lifetime; thus, partner choice and the maintenance of these relationships are both considered key components of alliance formation. In our previous investigations, pairs of adult male dolphins showed a high success rate in cooperative manipulation of a cognitive enrichment device. Here, we introduced two novel cognitive enrichment devices to the group of five dolphins, facilitating simultaneous actions for not only pairs, but for three or even four dolphins. The devices were made of PVC tubes, fittings and caps equipped with rope handles, creating a three-way (T-shape) and a four-way (TT-shape) device. The devices were filled with fish and ice and were designed to be opened by simultaneous pull of the handles. Both devices were tested in 12 trials (each lasted for 15 min), separately. Only one of the caps could be opened, the others were affixed with the position of the openable cap counter-balanced over the trials. Although the dolphins received no training regarding the manipulation of the devices, they were successful in cooperatively opening the three-way devices in 10/12 of trials (70% by two and 30% by three dolphins) and the four-way devices also in 10/12 trials (50% by two, 40% by three and 10% by four dolphins). The dolphins interacted with the devices during the entire testing time, and this was mostly spent in cooperative play (77% and 56% of the test duration with the three-way and four-way device, respectively). The majority of the cooperative play was observed between one particular pair of dolphins that was temporarily associated with a third or sometimes even with a fourth dolphin. These findings demonstrate the first successful use of multi-partner cooperative enrichment devices, providing information on the social organization of a male dolphin group.

Conserved transcriptomic profiles underpin monogamy across vertebrates.

Social monogamy, typically characterized by the formation of a pair bond, increased territorial defense, and often biparental care, has independently evolved multiple times in animals. Despite the independent evolutionary origins of monogamous mating systems, several homologous brain regions and neuropeptides and their receptors have been shown to play a conserved role in regulating social affiliation and parental care, but little is known about the neuromolecular mechanisms underlying monogamy on a genomic scale. Here, we compare neural transcriptomes of reproductive males in monogamous and nonmonogamous species pairs of Peromyscus mice, Microtus voles, parid songbirds, dendrobatid frogs, and Xenotilapia species of cichlid fishes. We find that, while evolutionary divergence time between species or clades did not explain gene expression similarity, characteristics of the mating system correlated with neural gene expression patterns, and neural gene expression varied concordantly across vertebrates when species transition to monogamy. Our study provides evidence of a universal transcriptomic mechanism underlying the evolution of monogamy in vertebrates.

Transcriptomics of Parental Care in the Hypothalamic-Septal Region of Female Zebra Finch Brain.

(1) Background: The objective of this study was to uncover genomic causes of parental care. Since birds do not lactate and, therefore, do not show the gene expressional changes required for lactation, we investigate gene expression associated with parenting in caring and non-caring females in an avian species, the small passerine bird zebra finch (Taeniopygia guttata). Here, we compare expression patterns in the hypothalamic-septal region since, previously, we showed that this area is activated in parenting females. (2) Methods: Transcriptome sequencing was first applied in a dissected part of the zebra finch brain related to taking care of the nestlings as compared to a control group of social pairs without nestlings. (3) Results: We found genes differentially expressed between caring and non-caring females. When introducing a log2fold change threshold of 1.5, 13 annotated genes were significantly upregulated in breeding pairs, while 39 annotated genes were downregulated. Significant enrichments of dopamine and acetylcholine biosynthetic processes were identified among upregulated pathways, while pro-opiomelanocortin and thyroid hormone pathways were downregulated, suggesting the importance of these systems in parental care. Network analysis further suggested neuro-immunological changes in mothers. (4) Conclusions: The results confirm the roles of several hypothesized major pathways in parental care, whereas novel pathways are also proposed.

Did we find a copycat? Do as I Do in a domestic cat (Felis catus).

This study shows evidence of a domestic cat (Felis catus) being able to successfully learn to reproduce human-demonstrated actions based on the Do as I Do paradigm. The subject was trained to reproduce a small set of familiar actions on command "Do it!" before the study began. To test feature-contingent behavioural similarity and control for stimulus enhancement, our test consisted of a modified version of the two-action procedure, combined with the Do as I Do paradigm. Instead of showing two different actions on an object to different subjects, we applied a within-subject design and showed the two actions to the same subject in separate trials. We show evidence that a well-socialized companion cat was able to reproduce actions demonstrated by a human model by reproducing two different actions that were demonstrated on the same object. Our experiment provides the first evidence that the Do as I Do paradigm can be applied to cats, suggesting that the ability to recognize behavioural similarity may fall within the range of the socio-cognitive skills of this species. The ability of reproducing the actions of a heterospecific human model in well-socialized cats may pave the way for future studies addressing cats' imitative skills.

Cognitive enrichment device provides evidence for intersexual differences in collaborative actions in Indo-Pacific bottlenose dolphins (Tursiops aduncus).

Social living, long lifespan and advanced cognitive skills provided favourable conditions for the development of pro-social behaviours and cooperative activities in cetacean. Dolphins have been observed to collaborate for various purposes, finding food, finding mates or raising and teaching younger individuals. This study investigated the potential impact of demographic factors (sex and age), social factors (relatedness and group size), and individual experience in a cooperative problem solving task. A cognitive enrichment device was tested with 22 dolphins in 11 group settings. The device consisted of a tube, containing ice and fish, sealed by two caps with rope handles and designed to be operated by pairs of dolphins. The investigation focused on the differences in trial outcome (success rate of cooperative opening of the device) and on cooperative play (dolphin pairs engaging in synchronous swim with the device). From the five potential factors, sex showed the highest impact. Cooperative openings were more than four times more frequent in males than in females (75% vs 17%, respectively), and cooperative play was exclusively displayed by adult males. Given the strong correlation between cooperative opening and cooperative play, we argue the two behaviours can be regarded as parts of a cooperative action chain. This study provides the first evidence for intersexual differences in collaborative actions in dolphins under systematic testing conditions.

Perception of animacy in dogs and humans.

Humans have a tendency to perceive inanimate objects as animate based on simple motion cues. Although animacy is considered as a complex cognitive property, this recognition seems to be spontaneous. Researchers have found that young human infants discriminate between dependent and independent movement patterns. However, quick visual perception of animate entities may be crucial to non-human species as well. Based on general mammalian homology, dogs may possess similar skills to humans. Here, we investigated whether dogs and humans discriminate similarly between dependent and independent motion patterns performed by geometric shapes. We projected a side-by-side video display of the two patterns and measured looking times towards each side, in two trials. We found that in Trial 1, both dogs and humans were equally interested in the two patterns, but in Trial 2 of both species, looking times towards the dependent pattern decreased, whereas they increased towards the independent pattern. We argue that dogs and humans spontaneously recognized the specific pattern and habituated to it rapidly, but continued to show interest in the 'puzzling' pattern. This suggests that both species tend to recognize inanimate agents as animate relying solely on their motions.

Do as I Did! Long-term memory of imitative actions in dogs (Canis familiaris).

This study demonstrates long-term declarative memory of imitative actions in a non-human animal species. We tested 12 pet dogs for their ability to imitate human actions after retention intervals ranging from 1 to 24 h. For comparison, another 12 dogs were tested for the same actions without delay between demonstration and recall. Our test consisted of a modified version of the Do as I Do paradigm, combined with the two-action procedure to control for non-imitative processes. Imitative performance of dogs remained consistently high independent of increasing retention intervals, supporting the idea that dogs are able to retain mental representations of human actions for an extended period of time. The ability to imitate after such delays supports the use of long-term declarative memory.

Spontaneous action matching in dog puppies, kittens and wolf pups.

We investigated the spontaneous tendency of dog puppies, kittens and wolf pups to match their behaviour to actions demonstrated by a human, in the absence of food reward. Based on dogs' inherent sociality and domestication history, we predicted that the tendency to match human actions is more pronounced in this species than in the other two. To test this, we exposed N = 42 dog puppies, N = 39 kittens and N = 8 wolf pups to ostensive human demonstrations of an object-related action. We found that dog puppies paid more attention to the demonstration than kittens and wolf pups. Dog puppies and wolf pups matched the demonstrated actions in more trials than kittens. Dog puppies also tended to reproduce the human demonstration that differed from the action they typically preformed in the absence of demonstration. These results support that dog puppies show a tendency to attend to humans and conform their behaviour to human demonstrations in the absence of extrinsic food rewards. This spontaneous tendency is also relevant for practical applications, by providing the basis to devise puppy-training methods that rely less on food rewards, and instead exploit puppies' natural predisposition for social learning.

Individual variation and the resolution of conflict over parental care in penduline tits.

Eurasian penduline tits (Remiz pendulinus) have an unusually diverse breeding system consisting of frequent male and female polygamy, and uniparental care by the male or the female. Intriguingly, 30 to 40 per cent of all nests are deserted by both parents. To understand the evolution of this diverse breeding system and frequent clutch desertion, we use 6 years of field data to derive fitness expectations for males and females depending on whether or not they care for their offspring. The resulting payoff matrix corresponds to an asymmetric Snowdrift Game with two alternative evolutionarily stable strategies (ESSs): female-only and male-only care. This, however, does not explain the polymorphism in care strategies and frequent biparental desertion, because theory predicts that one of the two ESSs should have spread to fixation. Using a bootstrapping approach, we demonstrate that taking account of individual variation in payoffs explains the patterns of care better than a model based on the average population payoff matrix. In particular, a model incorporating differences in male attractiveness closely predicts the observed frequencies of male and female desertion. Our work highlights the need for a new generation of individual-based evolutionary game-theoretic models.

Cognitive Group Testing Promotes Affiliative Behaviors in Dolphins.

Ex-situ research in aquariums and zoological settings not only support scientific advancement, they also provide opportunities for education, facilitating both mental and physical stimulation, consequently improving welfare. This study aimed to investigate the impact of cognitive testing on the well-being of a group of male Indo-Pacific bottlenose dolphins. The occurrence of affiliative, aggressive and potentially stereotypical behaviors was assessed based on seven monitored behavior types and compared between "Session days" and "Non-session days." The consistency of the impact was assessed over a three-year period. The analyses revealed that "Play with enrichment," "Affiliative tactile," "Social play" and "Synchronous swim" were significantly higher, while "Aggression" was significantly lower on Session days than on Non-session days. Individual analysis showed significant increase in the positive welfare indicators in all dolphins during Session days. The social network analysis of aggressive interactions between group members also supported an overall decrease of aggression during Session days. These results indicate that dolphin groups that voluntarily participate in cognitive tests under human care benefit from the testing and show an improvement in animal welfare while contributing to scientific advancement.

Sexual conflict predicts morphology and behavior in two species of penduline tits.

BACKGROUND: The evolutionary interests of males and females rarely coincide (sexual conflict), and these conflicting interests influence morphology, behavior and speciation in various organisms. We examined consequences of variation in sexual conflict in two closely-related passerine birds with contrasting breeding systems: the Eurasian penduline tit Remiz pendulinus (EPT) exhibiting a highly polygamous breeding system with sexually antagonistic interests over parental care, and the socially monogamous Cape penduline tit Anthoscopus minutus (CPT). We derived four a priori predictions from sexual conflict theory and tested these using data collected in Central Europe (EPT) and South Africa (CPT). Firstly, we predicted that EPTs exhibit more sexually dimorphic plumage than CPTs due to more intense sexual selection. Secondly, we expected brighter EPT males to provide less care than duller males. Thirdly, since song is a sexually selected trait in many birds, male EPTs were expected to exhibit more complex songs than CPT males. Finally, intense sexual conflict in EPT was expected to lead to low nest attendance as an indication of sexually antagonistic interests, whereas we expected more cooperation between parents in CPT consistent with their socially monogamous breeding system. RESULTS: Consistent with our predictions EPTs exhibited greater sexual dimorphism in plumage and more complex song than CPTs, and brighter EPT males provided less care than duller ones. EPT parents attended the nest less frequently and less simultaneously than CPT parents. CONCLUSIONS: These results are consistent with sexual conflict theory: species in which sexual conflict is more manifested (EPT) exhibited a stronger sexual dimorphism and more elaborated sexually selected traits than species with less intense sexual conflict (CPT). Our results are also consistent with the notion that EPTs attempt to force their partner to work harder as expected under sexual conflict: each member of the breeding pair attempts to shift the costs of care to the other parent. More brightly colored males benefit more from desertion than dull ones, because they are more likely to remate with a new female. Taken together, the comparison between two closely related species with contrasting breeding systems suggest that sexual conflict over care has influenced the evolution of behavior and morphology in penduline tits.

Mental representation and episodic-like memory of own actions in dogs.

We investigated whether dogs remember their spontaneous past actions relying on episodic-like memory. Dogs were trained to repeat a small set of actions upon request. Then we tested them on their ability to repeat other actions produced by themselves, including actions performed spontaneously in everyday situations. Dogs repeated their own actions after delays ranging from a few seconds to 1 hour, with their performance showing a decay typical of episodic memory. The combined evidence of representing own actions and using episodic-like memory to recall them suggests a far more complex representation of a key feature of the self than previously attributed to dogs. Our method is applicable to various species, paving the way for comparative investigations on the evolution and complexity of self-representation.

Comparing the tractability of young hand-raised wolves (Canis lupus) and dogs (Canis familiaris).

Dogs live in 45% of households, integrated into various human groups in various societies. This is certainly not true for wolves. We suggest that dogs' increased tractability (meant as individual dogs being easier to control, handle and direct by humans, in contrast to trainability defined as performance increase due to training) makes a crucial contribution to this fundamental difference. In this study, we assessed the development of tractability in hand-raised wolves and similarly raised dogs. We combined a variety of behavioural tests: fetching, calling, obeying a sit signal, hair brushing and walking in a muzzle. Wolf (N = 16) and dog (N = 11) pups were tested repeatedly, between the ages of 3-24 weeks. In addition to hand-raised wolves and dogs, we also tested mother-raised family dogs (N = 12) for fetching and calling. Our results show that despite intensive socialization, wolves remained less tractable than dogs, especially in contexts involving access to a resource. Dogs' tractability appeared to be less context dependent, as they followed human initiation of action in more contexts than wolves. We found no evidence that different rearing conditions (i.e. intensive socialization vs. mother rearing) would affect tractability in dogs. This suggests that during domestication dogs might have been selected for increased tractability, although based on the current data we cannot exclude that the differential speed of development of dogs and wolves or the earlier relocation of wolves to live as a group explains some of the differences we found.

Neuronal activation in zebra finch parents associated with reintroduction of nestlings.

Recent studies of the brain mechanisms of parental behaviors have mainly focused on rodents. Using other vertebrate taxa, such as birds, can contribute to a more comprehensive, evolutionary view. In the present study, we investigated a passerine songbird, the zebra finch (Taeniopygia guttata), with a biparental caring system. Parenting-related neuronal activation was induced by first temporarily removing the nestlings, and then, either reuniting the focal male or female parent with the nestlings (parental group) or not (control group). To identify activated neurons, the immediate early gene product, Fos protein, was labeled. Both parents showed an increased level of parental behavior following reunion with the nestlings, and no sexual dimorphism occurred in the neuronal activation pattern. Offspring-induced parental behavior-related neuronal activation was found in the preoptic, ventromedial (VMH), paraventricular hypothalamic nuclei, and in the bed nucleus of the stria terminalis. In addition, the number of Fos-immunoreactive (Fos-ir) neurons in the nucleus accumbens predicted the frequency of the feeding of the nestlings. No difference was found in Fos expression when the effect of isolation or the presence of the mate was examined. Thus, our study identified a number of nuclei involved in parental care in birds and suggests similar regulatory mechanisms in caring females and males. The activated brain regions show similarities to rodents, while a generally lower number of brain regions were activated in the zebra finch. Furthermore, future studies are necessary to establish the role of the apparently avian-specific neuronal activation in the VMH of zebra finch parents.

Social behavior in dog puppies: Breed differences and the effect of rearing conditions.

INTRODUCTION: Most of the studies investigating the effect of early rearing environment in dogs used laboratory dogs and reported that early experiences markedly affect the puppies' behavior. However, the subjects of these experiments cannot be considered as representatives of family dogs. METHODS: In this study, we investigated whether different raising conditions shape social behavior toward humans in 8-week-old family dog puppies of two breeds, Labrador and Czechoslovakian wolf dog. The puppies were tested in a series of tests that represented typical situations of family dogs. RESULTS: We found that Czechoslovakian wolf dog puppies were more active than Labrador puppies in general, as they were more likely to explore the environment and the objects and spent more time doing so. Tendency to gaze at humans also varied between breeds, but in a context-specific way. Additionally, puppies housed separately from their mother interacted more with toys, puppies housed in a kennel tended to stay closer to the experimenter than puppies raised in the house, and puppies housed in a kennel tended to stay in the proximity of the experimenter more than puppies raised in the house. CONCLUSIONS: Our results provide evidence for early keeping conditions influencing social behavior and also highlight breed differences in puppies' behavior. Whether these differences are due to different developmental patterns and/or behavioral predispositions remains to be explored.

Social learning of goal-directed actions in dogs (Canis familiaris): Imitation or emulation?

A goal-directed action is composed of two main elements on which the observer may focus its attention: the movement performed (i.e., the action) and the outcome (i.e., the goal). In a social learning situation, consequently, the observer may imitate the action of the model or emulate the result of its action. In humans and primates, the tendency to selectively engage in any of these two processes is considered to be dependent upon the availability and saliency of information about the goal, implying the capacity to recognize the goals of others' actions. Dogs are skillful in learning socially from humans, and, when trained with the Do as I Do method, they imitate human actions. Here, we tested trained dogs for engaging in imitation or emulation based on information about the goal. We found that dogs observing the demonstration of an object-related action in the absence of a clear goal tended to solve the task by matching the body movement of the human demonstrator. In contrast, when they could observe the exact same movement, but the goal was apparent, they attempted to solve the task by their own means, that is, by emulation, instead of imitating the demonstrated action. These results provide experimental evidence for dogs being able to recognize the goals of others and adjust their behavior accordingly, relying flexibly on imitation or emulation. (PsycINFO Database Record (c) 2019 APA, all rights reserved).

Brain Distribution and Sexually Dimorphic Expression of Amylin in Different Reproductive Stages of the Zebra Finch (Taeniopygia guttata) Suggest Roles of the Neuropeptide in Song Learning and Social Behaviour.

The expression of the recently identified neuropeptide, amylin, is restricted in rodents to the postpartum preoptic area and may play a role in the control of parental behaviours and food intake. These processes are substantially different between bird and rodent parents as birds do not lactate but often show biparental care of the offspring. To establish the presence and role of amylin in the bird brain, in the present study, we investigated the distribution of amylin in brains of adult male and female zebra finches in three different reproductive stages (i.e. paired without young, incubating eggs or provisioning nestlings) and in unpaired control birds living in same sex flocks. Amylin mRNA was identified in the hypothalamus of zebra finch by RT-PCR, which was also used to produce probes for in situ hybridisation. Subsequently, in situ hybridisation histochemistry was performed in brain sections, and the labelling signal was quantified and compared between the groups. Amylin showed a much wider brain distribution than that of rodents. A strong and, in some regions, sexually dimorphic label was found in the striatum and several brain regions of the social behavioural network in both males and females. Many regions responsible for the learning of birdsong also contained amylin-positive neurons, and some regions showed sex differences reflecting the fact that vocalisation is sexually dimorphic in the zebra finch: only males sing. Area X (Ar.X), a striatal song centre present only in males, was labelled in paired but not unpaired male. Ar.X, another song centre, the lateral part of the magnocellular nucleus of the anterior nidopallium (lMAN) also contained amylin and had higher amylin label in paired, as opposed to unpaired birds. The wider distribution of amylin in birds as compared to rodents suggests a more general role of amylin in social or other behaviours in avian species than in mammals. Alternatively, parental care in birds may be a more complex behavioural trait involving a wider set of brain regions. The sex differences in song centres, and the changes with reproductive status suggest a participation of amylin in social behaviours and related changes in the singing of males.

Sexual conflict and consistency of offspring desertion in Eurasian penduline tit Remiz pendulinus.

BACKGROUND: The trade-off between current and future parental investment is often different between males and females. This difference may lead to sexual conflict between parents over care provisioning in animals that breed with multiple mates. One of the most obvious manifestations of sexual conflict over care is offspring desertion whereby one parent deserts the young to increase its reproductive success at the expense of its mate. Offspring desertion is a wide-spread behavior, and its frequency often varies within populations. We studied the consistency of offspring desertion in a small passerine bird, the Eurasian penduline tit Remiz pendulinus, that has an extremely variable breeding system. Both males and females are sequentially polygamous, and a single parent (either the male or the female) incubates the eggs and rears the young. About 28-40% of offspring are abandoned by both parents, and these offspring perish. Here we investigate whether the variation in offspring desertion in a population emerges either by each individual behaving consistently between different broods, or it is driven by the environment. RESULTS: Using a three-year dataset from Southern Hungary we show that offspring desertion by females is consistent between nests. Male desertion, however, depends on ambient environment, because all males desert their nests early in the season and some of them care late in the season. Therefore, within-population variation in parental care emerges by sexually different mechanisms; between-individual variation was responsible for the observed pattern of offspring desertion in females, whereas within-individual variation was responsible for the observed pattern in males. CONCLUSION: To our knowledge, our study is the first that investigates repeatability of offspring desertion behavior in nature. The contrasting strategies of the sexes imply complex evolutionary trajectories in breeding behavior of penduline tits. Our results raise an intriguing question whether the sexual difference in caring/deserting decisions explain the extreme intensity of sexual conflict in penduline tits that produces a high frequency of biparentally deserted (and thus wasted) offspring.