Domestication and breeding objective did not shape the interpretation of physical and social cues in goats (Capra hircus).

Artificial selection by humans, either through domestication or subsequent selection for specific breeding objectives, drives changes in animal cognition and behaviour. However, most previous cognitive research comparing domestic and wild animals has focused on companion animals such as canids, limiting any general claims about the effects of artificial selection by humans. Using a cognitive test battery, we investigated the ability of wild goats (non-domestic, seven subjects), dwarf goats (domestic, not selected for milk production, 15 subjects) and dairy goats (domestic, selected for high milk yield, 18 subjects) to utilise physical and social cues in an object choice task. To increase the heterogeneity of our test samples, data for domestic goats were collected by two experimenters at two research stations (Agroscope; Research Institute for Farm Animal Biology). We did not find performance differences between the three groups in the cognitive test battery for either physical or social cues. This indicates that for a domestic non-companion animal species, domestication and selection for certain breeding objectives did not measurably shape the physical and cognitive skills of goats.

How do goats "read" 2D-images of familiar and unfamiliar conspecifics?

To study individual recognition in animals, discrimination tasks are often conducted by presenting 2D images of real conspecifics. However, animals may discriminate the images merely as visual stimulus combinations without establishing referential relationships to the individuals depicted. In the current study, we investigated whether goats are able to discriminate photos of familiar and unfamiliar conspecifics, whether they not only process the photos as visual stimuli, but also understand them as virtual copies of real conspecifics and whether they grasp the concept of familiarity. Using a computer-controlled learning device, in three tests, goats of two experimental groups (A and B) had to discriminate portrait (Te1), profile (Te2) or headless body photos (Te3) of conspecifics. Tests were presented as 4-choice tasks, with one photo from Group A (rewarded) plus three photos from Group B (distractors). That is, the rewarded photo was familiar to Group A, but unfamiliar to Group B. Finally, in a reversal test (Te4) we reversed this principle. The goats learned the discriminations in Te1 to Te3 within two (Te1 and Te2) and three training days (Te3), respectively, and they needed between 91 [CL (66, 126)] and 174 [CL (126, 241)] trials to reach the learning criterion, with no statistically significant differences between the groups. In Te4, in contrast, the animals took 403 [Group A; CL (291, 557)] and 385 [Group B; CL (286, 519)] trials, respectively, to learn the task. The lack of spontaneous preferences for the photo of the familiar conspecific in the pretests of Te1 to Te3 in Group A, as well as the lack of differences in the number of trials to learn the discriminations between both groups, do not at first glance suggest that the goats established a correspondence between real conspecifics and their 2D representations. However, the higher number of trials in Te4 suggests that both groups formed the learning rule of choosing either the known (Group A) or the unknown goat (Group B) over the course of Te1 to Te3 and then failed after the rule was reversed, providing evidence that goats can associate 2D photos of conspecifics with real animals.

Setting the stage to tag "n" track: a guideline for implementing, validating and reporting a radio frequency identification system for monitoring resource visit behavior in poultry.

Passive radio frequency identification (RFID) can advance poultry behavior research by enabling automated, individualized, longitudinal, in situ, and noninvasive monitoring; these features can usefully extend traditional approaches to animal behavior monitoring. Furthermore, since the technology can provide insight into the visiting patterns of tagged animals at functional resources (e.g., feeders), it can be used to investigate individuals' welfare, social position, and decision-making. However, the lack of guidelines that would facilitate implementing an RFID system for such investigations, describing it, and establishing its validity undermines this technology's potential for advancing poultry science. This paper aims to fill this gap by 1) providing a nontechnical overview of how RFID functions; 2) providing an overview of the practical applications of RFID technology in poultry sciences; 3) suggesting a roadmap for implementing an RFID system in poultry behavior research; 4) reviewing how validation studies of RFID systems have been done in farm animal behavior research, with a focus on terminologies and procedures for quantifying reliability and validity; and 5) suggesting a way to report on an RFID system deployed for animal behavior monitoring. This guideline is aimed mainly at animal scientists, RFID component manufacturers, and system integrators who wish to deploy RFID system as an automated tool for monitoring poultry behavior for research purposes. For such a particular application, it can complement indications in classic general standards (e.g., ISO/IEC 18000-63) and provide ideas for setting up, testing, and validating an RFID system and a standard for reporting on its adequacy and technical aspects.

Personality traits affect learning performance in dwarf goats (Capra hircus).

A wide range of species exhibit time- and context-consistent interindividual variation in a number of specific behaviors related to an individual's personality. Several studies have shown that individual differences in personality-associated behavioral traits have an impact on cognitive abilities. The aim of this study was to investigate the relationship between personality traits and learning abilities in dwarf goats. The behavior of 95 goats during a repeated open field (OF) and novel object test (NO) was analyzed, and two main components were identified using principal component analysis: boldness and activity. In parallel, the goats learned a 4-choice visual initial discrimination task (ID) and three subsequent reversal learning (RL) tasks. The number of animals that reached the learning criterion and the number of trials needed (TTC) in each task were calculated. Our results show that goats with the lowest learning performance in ID needed more TTC in RL1 and reached the learning criterion less frequently in RL2 and RL3 compared to animals with better learning performance in ID. This suggests a close relationship between initial learning and flexibility in learning behavior. To study the link between personality and learning, we conducted two analyses, one using only data from the first OF- and NO-test (momentary personality traits), while the other included both tests integrating only animals that were stable for their specific trait (stable personality traits). No relationship between personality and learning was found using data from only the first OF- and NO-test. However, stability in the trait boldness was found to have an effect on learning. Unbold goats outperformed bold goats in RL1. This finding supports the general hypothesis that bold animals tend to develop routines and show less flexibility in the context of learning than unbold individuals. Understanding how individual personality traits can affect cognitive abilities will help us gain insight into mechanisms that can constrain cognitive processing and adaptive behavioral responses.

Responsiveness of domesticated goats towards various stressors following long-term cognitive test exposure.

Current evidence suggests that frequent exposure to situations in which captive animals can solve cognitive tasks may have positive effects on stress responsiveness and thus on welfare. However, confounding factors often hamper the interpretation of study results. In this study, we used human-presented object-choice tests (in form of visual discrimination and reversal learning tests and a cognitive test battery), to assess the effect of long-term cognitive stimulation (44 sessions over 4-5 months) on behavioural and cardiac responses of female domestic goats in subsequent stress tests. To disentangle whether cognitive stimulation per se or the reward associated with the human-animal interaction required for testing was affecting the stress responsiveness, we conditioned three treatment groups: goats that were isolated for participation in human-presented cognitive tests and rewarded with food ('Cognitive', COG treatment), goats that were isolated as for the test exposure and rewarded with food by the experimenter without being administered the object-choice tests ('Positive', POS treatment), and goats that were isolated in the same test room but neither received a reward nor were administered the tests ('Isolation', ISO treatment). All treatment groups were subsequently tested in four stress tests: a novel arena test, a novel object test, a novel human test, and a weighing test in which goats had to enter and exit a scale cage. All treatment groups weretested at the same two research sites, each using two selection lines, namely dwarf goats, not selected for production traits, and dairy goats, selected for high productivity. Analysing the data with principal component analysis and linear mixed-effects models, we did not find evidence that cognitive testing per se (COG-POS contrast) reduces stress responsiveness of goats in subsequent stress tests. However, for dwarf goats but not for dairy goats, we found support for an effect of reward-associated human-animal interactions (POS-ISO contrast) at least for some stress test measures. Our results highlight the need to consider ontogenetic and genetic variation when assessing stress responsiveness and when interacting with goats.

Learned control of urinary reflexes in cattle to help reduce greenhouse gas emissions.

Indiscriminate voiding of excreta by cattle contributes to greenhouse gas (GHG) emissions and soil and water contamination1,2. Emissions are higher in animal-friendly husbandry offering cattle more space2 - a trade-off we call the 'climate killer conundrum'. Voiding in a specific location (latrine) would help resolve this dilemma by allowing ready capture and treatment of excreta under more spacious farming conditions. For urination, toileting requires self-control and coordination of a complex chain of behaviors including awareness of bladder fullness, overriding of excretory reflexes, selection of a latrine and intentional relaxation of the external urethral sphincter3. Attempts to train toileting in cattle have so far been only partly successful4-6, even though their excretion and associated neurophysiological control are similar to those in species capable of toileting3. Similarly, very young infants have been considered incapable of self-initiated voiding, but they can be taught with extensive training7. Using a backward chaining, reward-based training procedure, we here show that cattle can control their micturition reflex and use a latrine for urination. Such self-control provides evidence that animals can learn to respond to and reveal internal experiences via appropriately trained operant behaviors, thereby providing another way to explore their subjective states.

Goats (Capra hircus) From Different Selection Lines Differ in Their Behavioural Flexibility.

Given that domestication provided animals with more stable environmental conditions, artificial selection by humans has likely affected animals' ability to learn novel contingencies and their ability to adapt to changing environments. In addition, the selection for specific traits in domestic animals might have an additional impact on subjects' behavioural flexibility, but also their general learning performance, due to a re-allocation of resources towards parameters of productivity. To test whether animals bred for high productivity would experience a shift towards lower learning performance, we compared the performance of dwarf goats (not selected for production, 15 subjects) and dairy goats (selected for high milk yield, 18 subjects) in a visual discrimination learning and reversal learning task. Goats were tested individually in a test compartment and were rewarded by choosing either a white or a black cup presented by the experimenter on a sliding board behind a crate. Once they reached a designated learning criterion in the initial learning task, they were transferred to the reversal learning task. To increase the heterogeneity of our test sample, data was collected by two experimenters at two research stations following a similar protocol. Goats of both selection lines did not differ in the initial discrimination learning task in contrast to the subsequent reversal learning task. Dairy goats reached the learning criterion slower compared to dwarf goats (dairy goats = 9.18 sessions; dwarf goats = 7.74 sessions; P = 0.016). Our results may indicate that the selection for milk production might have affected behavioural flexibility in goats. These differences in adapting to changing environmental stimuli might have an impact on animal welfare e.g., when subjects have to adapt to new environments or changes in housing and management routines.

How can Cattle be Toilet Trained? Incorporating Reflexive Behaviours into a Behavioural Chain.

Untrained cattle do not defecate or urinate in defined locations. The toilet training of cattle would allow urine and faeces to be separated and stored, reducing climate-damaging emissions and improving animal health. In a proof-of-concept study, we evaluated a novel protocol for toilet training in cattle. Five heifer calves (and yoked controls) were trained in the voluntary (operant) behaviours of a toileting chain. Then, reflexive urinating responses were incorporated into the chain, with toileting signalled by a tactile (vibratory) stimulus. On 95% of occasions, the calves inhibited/interrupted urination when receiving the stimulus, and on 65% of these occasions, reinitiated urination in the latrine. Furthermore, during 63% of urinations in the latrine, the calves oriented to the reward location before any food was delivered, providing additional evidence that calves can be successfully toilet trained with food rewards. Yoked controls failed to learn most of the operant elements and all the reflexive responses of toilet training. The results show that reflexive behaviours can be incorporated into voluntary toileting sequences with cattle and extend the range of species that can be toilet trained. Future refinement of the protocol to allow training under practical farm conditions offers the potential to mitigate climate damage and improve animal health.

Conditionability of 'voluntary' and 'reflexive-like' behaviors, with special reference to elimination behavior in cattle.

Typically, cattle urinate and defecate with little or no control over time and place. The resulting excreta contributes to a range of adverse effects on the environment and the animals themselves. These adverse effects could be substantially ameliorated if livestock could be toilet trained. Toilet training requires an animal to suppress impending voiding (a reflexive-like behavior), move to a latrine (voluntary behavior) and reinitiate voiding. Here, we review the neurophysiological processes and learning mechanisms regulating toileting. The suppression and initiation of voiding occur primarily via the coordinated activity of smooth and striated anal and urinary sphincter muscles. The autonomic and somatic nervous systems, along with central processes, regulate these muscles. In several mammalian species, voluntary control of the sphincters has been demonstrated using classical and/or operant conditioning. In this review, we demonstrate that the neurophysiological and behavioral regulation of voiding in cattle is likely to be similarly conditionable. The management of excreta deposition in cattle could have major benefits for reducing livestock greenhouse gas emissions and improving animal health/welfare.

Farm Animal Cognition-Linking Behavior, Welfare and Ethics.

Farm animal welfare is a major concern for society and food production. To more accurately evaluate animal farming in general and to avoid exposing farm animals to poor welfare situations, it is necessary to understand not only their behavioral but also their cognitive needs and capacities. Thus, general knowledge of how farm animals perceive and interact with their environment is of major importance for a range of stakeholders, from citizens to politicians to cognitive ethologists to philosophers. This review aims to outline the current state of farm animal cognition research and focuses on ungulate livestock species, such as cattle, horses, pigs and small ruminants, and reflects upon a defined set of cognitive capacities (physical cognition: categorization, numerical ability, object permanence, reasoning, tool use; social cognition: individual discrimination and recognition, communication with humans, social learning, attribution of attention, prosociality, fairness). We identify a lack of information on certain aspects of physico-cognitive capacities in most farm animal species, such as numerosity discrimination and object permanence. This leads to further questions on how livestock comprehend their physical environment and understand causal relationships. Increasing our knowledge in this area will facilitate efforts to adjust husbandry systems and enrichment items to meet the needs and preferences of farm animals. Research in the socio-cognitive domain indicates that ungulate livestock possess sophisticated mental capacities, such as the discrimination between, and recognition of, conspecifics as well as human handlers using multiple modalities. Livestock also react to very subtle behavioral cues of conspecifics and humans. These socio-cognitive capacities can impact human-animal interactions during management practices and introduce ethical considerations on how to treat livestock in general. We emphasize the importance of gaining a better understanding of how livestock species interact with their physical and social environments, as this information can improve housing and management conditions and can be used to evaluate the use and treatment of animals during production.

Better, Not Just More-Contrast in Qualitative Aspects of Reward Facilitates Impulse Control in Pigs.

Delay-of-gratification paradigms, such as the famous "Marshmallow Test," are designed to investigate the complex cognitive concepts of self-control and impulse control in humans and animals. Such tests determine whether a subject will demonstrate impulse control by choosing a large, delayed reward over an immediate, but smaller reward. Documented relationships between impulsive behavior and aggression in humans and animals suggest important implications for farm animal husbandry and welfare, especially in terms of inadequate social behavior, tail biting and maternal behavior. In a preliminary study, we investigated whether the extent of impulse control would differ between quantitatively and qualitatively different aspects of reward in pigs. Twenty female piglets were randomly divided into two groups, with 10 piglets each. After a preference test to determine individual reward preference among six different food items, a discrimination test was conducted to train for successful discrimination between different amounts of reward (one piece vs. four pieces) and different qualitative aspects of reward (highly preferred vs. least preferred food item). Then, an increasing delay (2, 4, 8, 16, 24, 32 s) was introduced for the larger/highly preferred reward. Each piglet could choose to get the smaller/least preferred reward immediately or to wait for the larger/highly preferred reward. Piglets showed clear differences in their preference for food items. Moreover, the "quality group" displayed faster learning in the discrimination test (number of sessions until 90% of the animals completed the discrimination test: "quality group"-3 days vs. "quantity group"-5 days) and reached a higher level of impulse control in the delay-of-gratification test compared to the "quantity group" (maximum delay that was mastered: "quality group"-24 s vs. "quantity group"-8 s). These results demonstrate that impulse control is present in piglets but that the opportunity to get a highly preferred reward is more valued than the opportunity to get more of a given reward. This outcome also underlines the crucial role of motivation in cognitive test paradigms. Further investigations will examine whether impulse control is related to traits that are relevant to animal husbandry and welfare.

Evaluating the temporal and situational consistency of personality traits in adult dairy cattle.

Recent research suggests that personality, defined as consistent individual behavioral variation, in farm animals could be an important factor when considering their health, welfare, and productivity. However, behavioral tests are often performed individually and they might not reflect the behavioral differences manifested in every-day social environments. Furthermore, the contextual and longer-term temporal stability of personality traits have rarely been investigated in adult dairy cattle. In this study, we tested three groups of lactating Holstein cows (40 cows) using an individual arena test and a novel object test in groups to measure the contextual stability of behavior. Among the recorded individual test parameters, we used seven in the final analysis, which were determined by a systematic parameter reduction procedure. We found positive correlations between novel object contact duration in the group test and individual test parameters object contact duration (Rs = 0.361, P = 0.026) and movement duration (Rs = 0.336, P = 0.039). Both tests were repeated 6 months later to investigate their temporal stability whereby four individual test parameters were repeatable. There was no consistency in the group test results for 25 cows tested twice, possibly due to group composition changes. Furthermore, based on the seven individual test parameters, two personality traits (activity/exploration and boldness) were identified by principal component analysis. We found a positive association between the first and second tests for activity/exploration (Rs = 0.334, P = 0.058) and for boldness (Rs = 0.491, P = 0.004). Our results support the multidimensional nature of personality in adult dairy cattle and they indicate a link between behavior in individual and within-group situations. The lack of stability according to the group test results implies that group companions might have a stronger influence on individual behavior than expected. We suggest repeating the within-group behavioral measurements to study the relationship between the social environment and the manifestation of personality traits in every-day situations.

Human-directed behaviour in goats is not affected by short-term positive handling.

In addition to domestication, interactions with humans or task-specific training during ontogeny have been proposed to play a key role in explaining differences in human-animal communication across species. In livestock, even short-term positive interactions with caretakers or other reference persons can influence human-animal interaction at different levels and over different periods of time. In this study, we investigated human-directed behaviour in the 'unsolvable task' paradigm in two groups of domestic goats (Capra aegagrus hircus). One group was positively handled and habituated to a plastic box by the experimenter to retrieve a food reward, while the other group only received standard husbandry care and was habituated to the box without human assistance. In the unsolvable task, the lid was fixed to the box, with the reward inaccessible to the subjects. The goats were confronted with the unsolvable task three times. We observed no difference between the two groups regarding gaze and contact alternations with the experimenter when confronted with the task they cannot solve by themselves. The goats did not differ in their expression rates of both gaze and contact alternations over three repetitions of the unsolvable task; however, they showed earlier gaze and contact alternations in later trials. The results do not support the hypothesis that short-term positive handling or task-specific training by humans facilitates human-directed behaviour in goats. In contrast, standard husbandry care might be sufficient to establish humans as reference persons for farm animals in challenging situations.

Motor self-regulation in goats (Capra aegagrus hircus) in a detour-reaching task.

Motor self-regulation is the ability to inhibit a prepotent response to a salient cue in favour of a more appropriate response. Motor self-regulation is an important component of the processes that interact to generate effective inhibitory control of behaviour, and is theorized to be a prerequisite of complex cognitive abilities in humans and other animals. In a large comparative study using the cylinder task, motor self-regulation was studied in 36 different species, mostly birds and primates. To broaden the range of species to comprehensively evaluate this phenomenon, motor self-regulation was studied in the domestic goat, which is a social ungulate species and moderate food specialist. Using the cylinder task, goats were first trained to perform a detour-reaching response to retrieve a reward from an opaque cylinder. Subsequently, an otherwise identical transparent cylinder was substituted for the opaque cylinder over 10 test trials. The goats' ability to resist approaching the visible reward directly by touching the cylinder and to retain the trained detour-reaching response was measured. The results indicated that goats showed motor self-regulation at a level comparable to or better than that of many of the bird and mammal species tested to date. However, the individual reaction patterns revealed large intra- and inter-individual variability regarding motor self-regulation. An improvement across trials was observed only in latency to make contact with the reward; no improvement in the proportion of accurate trials was observed. A short, distinct pointing gesture by the experimenter during baiting did not have any impact on the side of the cylinder to which the goats detoured. In half of goats, individual side biases were observed when detouring to the side of the cylinder, but there was no bias at the population level for either the left or right side. The results underline the need for a detailed examination of individual performance and additional measures to achieve a complete understanding of animal performance in motor self-regulation tasks.

Personality Research in Mammalian Farm Animals: Concepts, Measures, and Relationship to Welfare.

Measuring and understanding personality in animals is a rising scientific field. Much research has been conducted to assess distinctive individual differences in behavior in a large number of species in the past few decades, and increasing numbers of studies include farm animals. Nevertheless, the terminology and definitions used in this broad scientific field are often confusing because different concepts and methods are used to explain often synonymously applied terms, such as personality, temperament and coping style. In the present review we give a comprehensive overview of the concepts and terms currently used in animal personality research and critically reveal how they are defined and what they measure. First, we shortly introduce concepts describing human personality and how these concepts are used to explain animal personality. Second, we present which concepts, methods and measures are applied in farm animal personality research and show that the terminology used seems to be somehow species-related. Finally, we discuss some findings on the possible impact of personality on the welfare of farm animals. The assessment of personality in farm animals is of growing scientific and practical interest. Differences in theoretical frameworks and methodological approaches may also entail the diverse use of the different concepts between basic and applied research approaches. We conclude that more consistency is needed in using different theoretical concepts, terms and measures, especially in farm animal personality research. The terms coping style and temperament, which are used in different ways, should not be examined as independent concepts, but rather should be considered as different aspects of the whole personality concept. Farm animal personality should be increasingly considered for the improvement of animal housing, management, breeding and welfare.

Coping Style Modifies General and Affective Autonomic Reactions of Domestic Pigs in Different Behavioral Contexts.

Based on individual adaptive strategies (coping), animals may react differently to environmental challenges in terms of behavior and physiology according to their emotional perception. Emotional valence as well as arousal may be derived by measuring vagal and sympathetic tone of the autonomic nervous system (ANS). We investigated the situation-dependent autonomic response of 16 domestic pigs with either a reactive or a proactive coping style, previously selected according to the backtest which is accepted in piglets to assess escape behavior. At 11 weeks of age, the pigs were equipped with an implantable telemetric device, and heart rate (HR), blood pressure (BP) and their respective variabilities (HRV, BPV) were recorded for 1 h daily over a time period of 10 days and analyzed in four behavioral contexts (resting, feeding, idling, handling). Additionally, the first minute of feeding and handling was used for a short-term analysis of these parameters in 10-s intervals. Data from day 1-3 (period 1) and day 8-10 (period 2) were grouped into two separate periods. Our results revealed general differences between the coping styles during feeding, resting and handling, with proactive pigs showing higher HR compared to reactive pigs. This elevated HR was based on either lower vagal (resting) or elevated sympathetic activation (feeding, handling). The short-term analysis of the autonomic activation during feeding revealed a physiological anticipation reaction in proactive pigs in period 1, whereas reactive pigs showed this reaction only in period 2. Food intake was characterized by sympathetic arousal with concurrent vagal withdrawal, which was more pronounced in proactive pigs. In contrast, neither coping style resulted in an anticipation reaction to handling. Vagal activation increased in reactive pigs during handling, while proactive pigs showed an increase in sympathetically driven arousal in period 2. Our findings confirm significant context-related differences in the general autonomic reaction of pigs with different coping styles. Additionally, the two coping styles differ in their affective appraisal over the time course of the experiment, underlining the importance of taking individual differences into account when studying affect and emotion in humans and animals.

Comment on "Ducklings imprint on the relational concept of 'same or different'".

Martinho and Kacelnik (Reports, 15 July 2016, p. 286) reported that newly hatched ducklings imprinted on relational concepts. We argue that reanalyzing the data at the individual level shows that this conclusion cannot be applied for all sets of stimuli presented and that the ability to grasp relational concepts is limited to the stimulus category that is most beneficial for survival.

Adrenocortical Expression Profiling of Cattle with Distinct Juvenile Temperament Types.

Temperament affects ease of handling, animal welfare, and economically important production traits in cattle. The use of gene expression profiles as molecular traits provides a novel means of gaining insight into behavioural genetics. In this study, differences in adrenocortical expression profiles between 60 F2 cows (Charolais × German Holstein) of distinct temperament types were analysed. The cows were assessed in a novel-human test at an age of 90 days. Most of the adrenal cortex transcripts which were differentially expressed (FDR <0.05) were found between temperament types of 'fearful/neophobic-alert' and all other temperament types. These transcripts belong to several biological functions like NRF2-mediated oxidative stress response, Glucocorticoid Receptor Signalling and Complement System. Overall, the present study provides new insight into transcriptional differences in the adrenal cortex between cows of distinct temperament types. Genetic regulations of such molecular traits facilitate uncovering positional and functional gene candidates for temperament type in cattle.