Multimodal Communication in the Human-Cat Relationship: A Pilot Study.

Across all species, communication implies that an emitter sends signals to a receiver, through one or more channels. Cats can integrate visual and auditory signals sent by humans and modulate their behaviour according to the valence of the emotion perceived. However, the specific patterns and channels governing cat-to-human communication are poorly understood. This study addresses whether, in an extraspecific interaction, cats are sensitive to the communication channel used by their human interlocutor. We examined three types of interactions-vocal, visual, and bimodal-by coding video clips of 12 cats living in cat cafés. In a fourth (control) condition, the human interlocutor refrained from emitting any communication signal. We found that the modality of communication had a significant effect on the latency in the time taken for cats to approach the human experimenter. Cats interacted significantly faster to visual and bimodal communication compared to the "no communication" pattern, as well as to vocal communication. In addition, communication modality had a significant effect on tail-wagging behaviour. Cats displayed significantly more tail wagging when the experimenter engaged in no communication (control condition) compared to visual and bimodal communication modes, indicating that they were less comfortable in this control condition. Cats also displayed more tail wagging in response to vocal communication compared to the bimodal communication. Overall, our data suggest that cats display a marked preference for both visual and bimodal cues addressed by non-familiar humans compared to vocal cues only. Results arising from the present study may serve as a basis for practical recommendations to navigate the codes of human-cat interactions.

Discrimination of cat-directed speech from human-directed speech in a population of indoor companion cats (Felis catus).

In contemporary western cultures, most humans talk to their pet companions. Speech register addressed to companion animals shares common features with speech addressed to young children, which are distinct from the typical adult-directed speech (ADS). The way dogs respond to dog-directed speech (DDS) has raised scientists' interest. In contrast, much less is known about how cats perceive and respond to cat-directed speech (CDS). The primary aim of this study was to evaluate whether cats are more responsive to CDS than ADS. Secondarily, we seek to examine if the cats' responses to human vocal stimuli would differ when it was elicited by their owner or by a stranger. We performed playback experiments and tested a cohort of 16 companion cats in a habituation-dishabituation paradigm, which allows for the measurement of subjects' reactions without extensive training. Here, we report new findings that cats can discriminate speech specifically addressed to them from speech addressed to adult humans, when sentences are uttered by their owners. When hearing sentences uttered by strangers, cats did not appear to discriminate between ADS and CDS. These findings bring a new dimension to the consideration of human-cat relationship, as they imply the development of a particular communication into human-cat dyads, that relies upon experience. We discuss these new findings in the light of recent literature investigating cats' sociocognitive abilities and human-cat attachment. Our results highlight the importance of one-to-one relationships for cats, reinforcing recent literature regarding the ability for cats and humans to form strong bonds.

How's my kitty? Acoustic parameters of cat-directed speech in human-cat interactions.

In Western cultures, humans tend to use a specific kind of speech when talking to their pets, characterised, from an acoustical point of view, by elevated pitch and greater pitch modulation. Pet-directed speech (PDS), which has been mainly studied in dogs, shares some acoustic features with infant-directed speech (IDS), used when talking to young children. The purpose of this study was to test the hypothesis that adult humans also modify characteristics of their voice when talking to a cat. We compared acoustic parameters of speech directed to cats (CDS) and speech directed to adult humans (ADS). In a first experiment, we compared ADS and CDS utterances of male and female participants, addressing cats through video recordings, under controlled laboratory conditions. Both men and women used a higher pitch (mean fundamental frequency, or mean F0) in CDS vs. ADS. The second experiment was conducted under conditions allowing direct cat-human interactions, in a cohort of women. Once again, mean F0 was significantly higher in CDS vs. ADS. Overall, these data confirm our hypothesis that humans change the way they speak when addressing a cat, mainly by increasing the pitch of their voice. Further research is needed to fully investigate specificities of this speech.

Handlers' Representations on Therapy Dogs' Welfare.

While research on the benefits of animal-assisted interventions is beginning to build a significant body of work, studies on the well-being of therapy dogs are still in their infancy. Since handlers are the people responsible for their therapy dog's welfare, we interviewed 111 French handlers through an online questionnaire. Our results underlined that (i) therapy dogs' welfare is multidimensional when physical and psychological welfare, a balance between work and dog life and the settings and interactions of sessions are all taken into consideration. (ii) The response of our handlers emphasized that considering therapy dog welfare is important for the quality and safety of AAI. (iii) Three categories of risks factors were highlighted: the spatio-temporal framework (planning and environment), the interactions with beneficiaries and the handler themselves. It is particularly important that handlers talk about the negative impact of interactions with beneficiaries since they are at the heart of AAI, however there are few studies focusing on interactions as a stressor for dogs in this practice. Moreover, since there is a potential for positive bias in the handlers' representations, it is important that they be trained to identify and manage the stress in their dogs. Future research is particularly needed on the impact of interactions during sessions on therapy dog welfare.

French handlers' perspectives on Animal-Assisted Interventions.

BACKGROUND: Animal-Assisted Interventions (AAI) are well implemented in human healthcare, in France as elsewhere; yet there are still difficulties in characterizing these practices and misconceptions about their mechanisms - little is known about the French practice of AAI and about the human-animal team. OBJECTIVES: This study aims to characterize AAI by exploring their specificities through French handlers' perspectives. MATERIAL AND METHOD: An online survey addressed to French handlers working in AAI with mainly one dog was carried out. This research included questions about their practice in AAI (registration status, beneficiaries, and animals) and their background (training in AAI, training in the medico-social field, training in animal behavior). We then examined a phenomenological understanding of handlers' definitions of their practice in AAI, their motivations to work with these approaches, and the expectations of the human-animal team. We used an open coding strategy and created major themes from their answers. RESULTS: 111 handlers participated in this study. The quantitative data highlighted a heterogeneity of handlers' profiles and professional backgrounds, although most profiles had previous training in healthcare. Five themes characterizing AAI emerged from the qualitative analysis: (1) AAI as additional approaches to care settings, (2) AAI as person-centered approaches, (3) the complementarity between handlers and their animal(s), (4) the shared role of mediator, and (5) handlers' beliefs about the human-animal relationship related to their personal experiences. This survey allowed us to understand how the French use AAI and its role in the care system. CONCLUSION: The benefits of AAI are numerous both for care settings and for the caregivers mainly by making the care more humane. AAI seem to put the wellbeing of beneficiaries and the relationship with the caregiver at the center of the care. The complementarity of the human-animal team is the common feature of these practices and is critical to their success. Future interdisciplinary studies are required to explore the particularities of these interspecific approaches and the differences between countries.

Consistency of female preference for male song in the domestic canary using two measures: Operant conditioning and vocal response.

Variation of female preferences is often reported in the literature and could be related to an artefact derived from multiple different methods used. Thus, there is a need to evaluate the influence of different methods when assessing female preferences. The present study aims to compare female preferences obtained from an operant conditioning test and from female vocal responses to male song in the domestic canary (Serinus canaria). In an operant conditioning test, females had the possibility to choose between two keys; a peck on one key elicited a supposed very attractive canary song while a peck on the other key elicited a less attractive song. Meanwhile, female vocal responses were recorded. Our results revealed that female canaries preferred to peck on the key eliciting the attractive song and that they emitted more copulation calls in response to the attractive song compared to the less attractive song. This study shows the congruence of these two methods and further suggests that they are reliable to study female preferences in laboratory conditions.

Pet-directed speech draws adult dogs' attention more efficiently than Adult-directed speech.

Humans speak to dogs using a special speech register called Pet-Directed Speech (PDS) which is very similar to Infant-Directed Speech (IDS) used by parents when talking to young infants. These two type of speech share prosodic features that are distinct from the typical Adult-Directed Speech (ADS): a high pitched voice and an increased pitch variation. So far, only one study has investigated the effect of PDS on dogs' attention. We video recorded 44 adult pet dogs and 19 puppies when listening to the same phrase enounced either in ADS or in PDS or in IDS. The phrases were previously recorded and were broadcasted via a loudspeaker placed in front of the dog. The total gaze duration of the dogs toward the loudspeaker, was used as a proxy of attention. Results show that adult dogs are significantly more attentive to PDS than to ADS and that their attention significantly increases along with the rise of the fundamental frequency of human' speech. It is likely that the exaggerated prosody of PDS is used by owners as an ostensive cue for dogs that facilitates the effectiveness of their communication, and should represent an evolutionarily determined adaptation that benefits the regulation and maintenance of their relationships.

Effect of interaction type on the characteristics of pet-directed speech in female dog owners.

Recent studies focusing on the interspecific communicative interactions between humans and dogs show that owners use a special speech register when addressing their dog. This register, called pet-directed speech (PDS), has prosodic and syntactic features similar to that of infant-directed speech (IDS). While IDS prosody is known to vary according to the context of the communication with babies, we still know little about the way owners adjust acoustic and verbal PDS features according to the type of interaction with their dog. The aim of the study was therefore to explore whether the characteristics of women's speech depend on the nature of interaction with their dog. We recorded 34 adult women interacting with their dog in four conditions: before a brief separation, after reuniting, during play and while giving commands. Our results show that before separation women used a low pitch, few modulations, high intensity variations and very few affective sentences. In contrast, the reunion interactions were characterized by a very high pitch, few imperatives and a high frequency of affectionate nicknames. During play, women used mainly questions and attention-getting devices. Finally when commanding, women mainly used imperatives as well as attention-getting devices. Thus, like mothers using IDS, female owners adapt the verbal as well as the non-verbal characteristics of their PDS to the nature of the interaction with their dog, suggesting that the intended function of these vocal utterances remains to provide dogs with information about their intentions and emotions.

On Public Influence on People's Interactions with Ordinary Biodiversity.

Besides direct impacts of urban biodiversity on local ecosystem services, the contact of city dwellers with urban nature in their everyday life could increase their awareness on conservation issues. In this paper, we focused on a particularly common animal urban species, the feral pigeon Columba livia. Through an observational approach, we examined behavioral interactions between city dwellers and this species in the Paris metropolis, France. We found that most people (mean: 81%) do not interact with pigeons. Further, interactions (either positive or negative) are context and age-dependent: children interact more than adults and the elderly, while people in tourist spots interact more than people in urban parks or in railway stations, a result that suggests that people interacting with pigeons are mostly tourists. We discuss these results in terms of public normative pressures on city dwellers' access to and reconnection with urban nature. We call for caution in how urban species are publically portrayed and managed, given the importance of interactions with ordinary biodiversity for the fate of nature conservation.

Female signalling to male song in the domestic canary, Serinus canaria.

Most studies on sexual selection focus on male characteristics such as male song in songbirds. Yet female vocalizations in songbirds are growing in interest among behavioural and evolutionary biologists because these vocalizations can reveal the female's preferences for male traits and may affect male display. This study was designed to test whether male song performance influences the different female signals in the domestic canary (Serinus canaria). Female canaries were exposed to three types of song performance, differing in the repetition rate of sexy syllables. This experiment demonstrates that female birds are engaged in multimodal communication during sexual interaction. The results support the copulation solicitation hypothesis for female-specific trills: these trills were positively correlated and had a similar pattern to the copulation solicitation displays; responses were higher to the songs with higher performance and responses decreased with the repetition of the stimulation. Also, we observed a sensitization effect with the repetition of the song of the highest performance for the simple calls. Simple trills and other calls were more frequent during the broadcast of canary songs compared with the heterospecific control songs. The differential use of female signals in response to different song performance reveals a highly differentiated female signalling system which is discussed in light of the role of female traits to understand sexual selection in a broader perspective.

Male song quality modulates c-Fos expression in the auditory forebrain of the female canary.

In canaries, specific phrases of male song (sexy songs, SS) that are difficult to produce are especially attractive for females. Females exposed to SS produce more copulation displays and deposit more testosterone into their eggs than females exposed to non-sexy songs (NS). Increased expression of the immediate early genes c-Fos or zenk (a.k.a. egr-1) has been observed in the auditory forebrain of female songbirds hearing attractive songs. C-Fos immunoreactive (Fos-ir) cell numbers were quantified here in the brain of female canaries that had been collected 30min after they had been exposed for 60min to the playback of SS or NS or control white noise. Fos-ir cell numbers increased in the caudomedial mesopallium (CMM) and caudomedial nidopallium (NCM) of SS birds as compared to controls. Song playback (pooled SS and NS) also tended to increase average Fos-ir cell numbers in the mediobasal hypothalamus (MBH) but this effect did not reach full statistical significance. At the individual level, Fos expression in CMM was correlated with its expression in NCM and in MBH but also with the frequency of calls that females produced in response to the playbacks. These data thus indicate that male songs of different qualities induce a differential metabolic activation of NCM and CMM. The correlation between activation of auditory regions and of the MBH might reflect the link between auditory stimulation and changes in behavior and reproductive physiology.

Heaven it's my wife! Male canaries conceal extra-pair courtships but increase aggressions when their mate watches.

Many animals live in a communication network, an environment where individuals can obtain information about competitors or potential mates by observing interactions between conspecifics. In such an environment, interactants might benefit by changing their signalling behaviour in the presence of an audience. This audience effect seems widespread among species, has been observed during various types of interaction (e.g. intra-sexual vs. inter-sexual interaction) and varies according to the social context (e.g. gender, hierarchical or mating status of the audience). However, the way individuals might adapt their signalling behaviour to a combination of these factors remains poorly understood. To address this question, we studied how the presence of an audience affects the behaviour of male domestic canaries Serinus canaria during two types of interactions: (i) an extra-pair interaction and (ii) a male-male competition for food. Males were observed under three conditions: (a) in the absence of audience, (b) in the presence of their mate or (c) of a familiar female. Our results show that male domestic canaries minutely adapt their courting and agonistic behaviours to a combination of: (i) the type of interaction (extra-pair interaction/male-male competition), (ii) the social context (mate, familiar female or nobody in audience) and (iii) the behaviours of both the audience and the interactant. These results highlight the ability of animals to subtly adapt their behaviour to the social environment. This also raises questions about the cognitive foundations and evolution of these processes especially considering that canaries are known neither for having high cognitive abilities nor for being a typical example for the social intelligence hypothesis.

Pigeons discriminate between human feeders.

Considered as plague in many cities, pigeons in urban areas live close to human activities and exploit this proximity to find food which is often directly delivered by people. In this study, we explored the capacity of feral pigeons to take advantage of this human-based food resource and discriminate between friendly and hostile people. Our study was conducted in an urban park. Pigeons were fed by two experimenters of approximately the same age and skin colour but wearing coats of different colours. During the training sessions, the two human feeders displayed different attitudes: one of the feeders was neutral and the second was hostile and chased away the pigeons. During the two test phases subsequent to the training phase, both feeders became neutral. Two experiments were conducted, one with one male and one female feeder and the second with two female feeders. In both experiments, the pigeons learned to quickly (six to nine sessions) discriminate between the feeders and maintained this discrimination during the test phases. The pigeons avoided the hostile feeder even when the two feeders exchanged their coats, suggesting that they used stable individual characteristics to differentiate between the experimenter feeders. Thus, pigeons are able to learn quickly from their interactions with human feeders and use this knowledge to maximize the profitability of the urban environment. This study provides the first experimental evidence in feral pigeons for this level of human discrimination.

Maternal effects and beta-carotene assimilation in Canary chicks.

Carotenoids are pigments responsible for the red, orange and yellow coloration of plants and animals. They may be beneficial in two ways; they have a powerful antioxidant activity, and they can behave as an immunostimulant. Animals however cannot synthesize carotenoids de novo, they must obtain them through their diet. In our experiments on Canaries, we investigated how mothers transfer their dietary carotenoid-related benefits to their offspring; either through the egg, or through the diet (during chicks' feeding). Female Canaries were allowed to access beta-carotene enriched food during egg formation and/or chicks' feeding. We sorted the chicks into four groups using the period when they assimilated the beta-carotene as a variable. The four groups were: (i) before hatching (from yolk), (ii) after hatching (from maternal feeding), (iii) before and after hatching, or (iv) never. Colorimetry and HPLC analysis from sub-samples of yolks confirmed the maternal transfer of dietary carotenoids to the yolk. Our results show that benefits from maternal dietary carotenoids are transferred to the chicks, but according to the period when they are assimilated by the chicks, the physiological effects are different. It was found that the chicks' growth was enhanced when carotenoids were assimilated both before and after hatching. However an increase in cellular immunity efficiency only occurs when the assimilation takes place after hatching.

Women use voice parameters to assess men's characteristics.

The purpose of this study was: (i) to provide additional evidence regarding the existence of human voice parameters, which could be reliable indicators of a speaker's physical characteristics and (ii) to examine the ability of listeners to judge voice pleasantness and a speaker's characteristics from speech samples. We recorded 26 men enunciating five vowels. Voices were played to 102 female judges who were asked to assess vocal attractiveness and speakers' age, height and weight. Statistical analyses were used to determine: (i) which physical component predicted which vocal component and (ii) which vocal component predicted which judgment. We found that men with low-frequency formants and small formant dispersion tended to be older, taller and tended to have a high level of testosterone. Female listeners were consistent in their pleasantness judgment and in their height, weight and age estimates. Pleasantness judgments were based mainly on intonation. Female listeners were able to correctly estimate age by using formant components. They were able to estimate weight but we could not explain which acoustic parameters they used. However, female listeners were not able to estimate height, possibly because they used intonation incorrectly. Our study confirms that in all mammal species examined thus far, including humans, formant components can provide a relatively accurate indication of a vocalizing individual's characteristics. Human listeners have the necessary information at their disposal; however, they do not necessarily use it.

Social competition and plasma testosterone profile in domesticated canaries: an experimental test of the challenge hypothesis.

The challenge hypothesis predicts that plasma testosterone (T) concentration is high when male-male competitions are high and decreases when males are engaged in paternal care. In monogamous species, T concentration increases at the beginning of the breeding period and decreases after egg laying. According to the challenge hypothesis, increasing competition should also lead to T increase. The aim of our study was to test this hypothesis. In a first experiment, we measured the T profile of domesticated canaries housed with their mate in separated cages without competition. In a second one, we created a competition by housing male and female domestic canaries together (in an aviary) and emphasized this competition by limiting food access. We also studied social status effect. Our results showed no effect of social status in both sexes and no differences in female's T concentration. Concerning males, we obtained a clear monogamous T profile from the ones housed in a low competition situation and a polygamous profile from the others housed in high competition situation. Thus, our results support the hypothesis of the plasticity of the mechanisms controlling T concentration according to environmental conditions.

Sexually attractive phrases increase yolk androgens deposition in Canaries (Serinus canaria).

The androgen concentration in birds' eggs varies with laying order, breeding conditions, and mate attractiveness. In passerine birds, mate attractiveness depends upon song quality. The aim of our study was to evaluate the effect of one criterion used by females to assess male song quality that is to say the presence of sexually attractive phrases on yolk androgen deposition. Twenty-five female Canaries were assigned to three experimental groups; in the first group, the females were allowed to hear songs made up with attractive phrases; in the second group, they were allowed to hear songs made up with non-attractive phrases; and in the control group, the females could not hear any song. Our results show that females allowed to hear songs with attractive phrases deposit significantly higher amounts of androgens (mostly testosterone) in their eggs than females without acoustical stimulation. The females exposed to songs with non-attractive phrases had androgen amounts halfway between the two other groups. This suggests that when females are paired with mates able to sing attractive phrases they can allocate more androgens in their eggs during the pre-laying period.

Female canaries produce eggs with greater amounts of testosterone when exposed to preferred male song.

Male birdsong has a great influence in the stimulation of female reproduction. However, female physiological responsiveness to song may depend on the degree of complexity of male song. This is expected because females of iteroparous organisms may increase their fitness by matching their reproductive investment to the predicted value of each reproductive attempt. To the extent that the expression of male ornaments is a signal of male quality, we expect females to increase their investment when paired to highly ornamented males. However, female investment may be cryptic and difficult to detect, such as androgen content in the eggs. In this study, we exposed female canaries (Serinus canaria) to attractive and unattractive song repertoires using a crossover design. As predicted, females invested greater concentrations of testosterone in their eggs when exposed to attractive repertoires than when exposed to unattractive repertoires. This implies that song repertoires convey important information about the reproductive value of a given male and suggests that testosterone deposition in egg yolk may be costly.