Genetics of animal temperament: aggressive behaviour at mixing is genetically associated with the response to handling in pigs

Aggression when pigs are mixed into new social groups has negative impacts on welfare and production. Aggressive behaviour is moderately heritable and could be reduced by genetic selection. The possible wider impacts of selection for reduced aggressiveness on handling traits and activity in the home pen were investigated using 1663 male and female pedigree pigs (898 purebred Yorkshire and 765 Yorkshire × Landrace). Aggressive behaviour was observed over 24 h after pigs were mixed at 10 weeks of age into groups balanced for unfamiliarity and weight. Aggression was highly heritable (duration of involvement in reciprocal fighting h2 = 0.47 ± 0.03, and duration of delivering one-sided aggression h2 = 0.34 ± 0.03). Three weeks after mixing, home pen inactivity (indicated by the frequency of lying) was observed over 24 h. Inactivity was weakly heritable (h2 = 0.05 ± 0.01) but showed no significant genetic association with aggression. Pigs' behaviour during handling by humans was assessed on entry to, whilst inside and on exit from a weigh crate at both mixing and end of test at 22 weeks. Pigs were generally easy to handle, moving easily into and out of the crate. Scores indicating 'very difficult to move' were rare. Handling scores at weighing were weakly heritable (h2 = 0.03 to 0.17), and moderately correlated across the two weighings (rg = 0.28 to 0.76). Aggressive behaviour at mixing was genetically associated with handling at the end of test weighing: pigs that fought and delivered one-sided aggression had handling scores indicating more active behaviour at weighing (e.g. moving quickly into the crate v. fighting rg = 0.41 ± 0.05 and v. bullying rg = 0.60 ± 0.04). Also, there was a genetic association between receiving one-side aggression at mixing and producing high-pitched vocalisations in the weigh crate (rg = 0.78 ± 0.08). Correlated behavioural responses occurring across different challenging situations (e.g. social mixing and human handling) have been described by the concept of animal temperament (also known as coping styles, personality or behavioural syndromes), but this has rarely been demonstrated at the genetic level in farm animals. These findings may have practical implications for the development of breeding programmes aimed at altering animal temperament. Breeding to reduce aggression could result in some reduction in activity at weighing. This would have consequences for animal production, because pigs which are inactive at weighing take longer to move into and out of the weigh crate, and perhaps also for animal welfare.     

Promoting positive states: the effect of early human handling on play and exploratory behaviour in pigs

It is known that tactile stimulation (TS) during ontogeny modifies brain plasticity and enhances the motor and cognitive skills. Our hypothesis was that early handling including TS would increase play and exploratory behaviour in commercial pigs under standardized test conditions. Piglets from 13 litters were subjected to three handling treatments from 5 to 35 days of age: all the piglets were handled (H), none of the piglets were handled (NH) or half of the piglets in the litter were handled (50/50). At 42 days of age, the pigs’ behaviour was observed in pairs in a novel pen with a ‘toy’ (tug rope). The main results were that more locomotor play was performed by pigs from litters where all or half of them had been handled, whereas social exploratory behaviour was more pronounced in pigs from litters where half of them had been handled. Although behaviour was affected by the interaction of treatment with sex or with weight category, we propose that the handling procedure does seem to have acted to increase locomotor skills and that handling half of the piglets in the litter may have triggered a series of socio-emotional interactions that were beneficial for the whole group.     

Aggression and Affiliation during Social Conflict in Pigs

Social conflict is mostly studied in relation to aggression. A more integral approach, including aggressive and affiliative behaviour as well as physiology, may however give a better understanding of the animals'' experience during social conflict. The experience of social conflict may also be reflected in the spatial distribution between conspecifics. The objective was to assess the relationship between behaviour, physiology, and spatial integration in pigs (Sus scrofa) during social conflict. Hereto, 64 groups of pigs (9 wk of age) were studied in a 24 h regrouping test whereby pairs of familiar pigs were grouped with 2 unfamiliar pairs, in either barren or straw-enriched housing. Data on aggressive and affiliative behaviour, skin lesions, body weight, and haptoglobin could be summarized into three principal component analysis factors. These three factors were analysed in relation to spatial integration, i.e. inter-individual distances and lying in body contact. Pigs stayed up to 24 h after encounter in closer proximity to the familiar pig than to unfamiliar pigs. Pigs with a high factor 1 score were more inactive, gave little social nosing, had many skin lesions and a high body weight. They tended to space further away from the familiar pig (b = 1.9 cm; P = 0.08) and unfamiliar ones (b = 0.7 cm; P = 0.05). Pigs that were involved in much aggression (factor 2), and that had a strong increase in haptoglobin (factor 3), tended to be relatively most far away from unfamiliar pigs (b = 0.03 times further; P = 0.08). Results on lying in body contact were coherent with results on distances. Pigs in enriched housing spaced further apart than pigs in barren housing (P<0.001). The combined analysis of measures revealed animals that may either promote or slow down group cohesion, which may not have become clear from single parameters. This emphasizes the importance of an integral approach to social conflict.     

Group lactation from 7 or 14 days of age reduces piglet aggression at weaning compared to farrowing crate housing

Early life experiences can affect social behaviour in later life, but opportunities for socio-behavioural development are often overlooked in current husbandry practices. This experiment investigated the effects of rearing piglets in two-stage group lactation (GL) system from 7 or 14 days of age on piglet aggression at weaning. Three lactation housing treatments were applied to a total of 198 piglets from 30 litters of multiparous sows. All dams farrowed in standard farrowing crates (FCs). Group lactation litters were transferred with their dam at 7 (GL₇) or 14 days (GL₁₄) postpartum to GL pens (one pen of five sows at 8.4 m²/sow and one pen of seven sows at 8.1 m²/sow, per GL treatment). Farrowing crate litters remained with their dam in a single litter until weaning. At weaning, 10 to 14 piglets from two unfamiliar litters from the same housing treatment were mixed into pens (n=5 pens/treatment) and their behaviour was continuously recorded for 3.5 h. For each pen, the frequency of aggressive bouts (reciprocal and non-reciprocal aggression lasting <5 s), the frequency and duration of fights (reciprocal aggression lasting ⩾5 s) and bullying events (non-reciprocal aggression lasting ⩾5 s) were recorded, along with whether interactions involved familiar or unfamiliar piglets. Aggressive bouts delivered by FC piglets were approximately 1.5 and 3.0 times more frequent than that delivered by GL₇ and GL₁₄ piglets, respectively (40.5, 16.7 and 9.9 bouts/pig, respectively; P<0.05). Fighting was more frequent (1.6, 0.3 and 0.4 fights/pig, respectively; P<0.001) and fights were longer (83, 15 and 32 s fight/pig, respectively; P<0.001) between FC piglets than between GL₇ or GL₁₄ piglets. Bullying did not differ between housing treatments (P>0.05). GL₇ and GL₁₄ piglets engaged in a similar number of fights with unfamiliar as familiar piglets, but FC piglets had almost three times as many fights with unfamiliar than with familiar piglets (P<0.05). This experiment confirms the benefits of GL housing for pig social development. Further investigation is required to determine whether mixing before 14 days postpartum has implications for other indicators of animal welfare and productivity in a two-stage GL housing system.     

Effect of rearing system and mixing at loading on transport and lairage behaviour and meat quality: comparison of outdoor and conventionally raised pigs

The effect of raising pigs outdoors or conventionally in a barren environment was investigated with respect to behaviour during a 3 to 4 h journey and a 2 h lairage, blood chemistry at slaughter and meat quality characteristics. Pigs were either kept in farm pen groups or were mixed at loading and kept in the groups, so formed, until slaughter. Non-mixed outdoor pigs settled more quickly during transport and lay down to a greater extent at the end of the journey and lairage period compared with conventionally raised pigs. Mixing led to fewer pigs sitting and lying during transport for conventionally raised pigs, where nearly 80% were still standing at the end of the journey. Mixing had no effect on pig posture in the lairage. Outdoor pigs were less aggressive than conventionally raised pigs especially during lairage and had a lower frequency of unacceptable skin damage in the rear and shoulder area. Aggressive interactions were almost exclusively confined to mixed groups and occurred mainly between pigs from different farm pens, i.e. between unfamiliar animals. Mixing at loading led therefore to higher levels of unacceptable skin damage. Cortisol concentrations in slaughter blood were not affected by rearing system or mixing, but mixed, conventionally raised pigs had higher plasma creatine kinase (CK) activities than non-mixed ones (1132 v. 761 U/l, respectively, P < 0.05). Outdoor pigs had similar CK activities, irrespective of mixing (682 and 771 U/l for mixed and non-mixed, respectively). Muscle pH early post mortem was highest in outdoor pigs and muscle temperature lower, but no pig showed pH values below 6.0. Ultimate pH values were both higher (Semimembranosus (S)) and lower (Semispinalis capitis (SC)) than in conventionally raised pigs, and outdoor pigs tended to have a lower frequency of higher than normal pH values. Internal reflectance (MQM) values in Biceps femoris (BF) were highest in outdoor pigs but the incidence of pale, soft and exudative (PSE) meat was low, varying between 0% and 1% for experimental groups. In general, the effects of rearing system and mixing on meat quality measurements taken early post mortem or the day after slaughter were slight, but the trends seen support the CK results, and show that conventionally raised pigs may have found mixing pre-slaughter to be more physically stressful than outdoor pigs did.     

The effect of long or chopped straw on pig behaviour

In the EU, pigs must have permanent access to manipulable materials such as straw, rope, wood, etc. Long straw can fulfil this function, but can increase labour requirements for cleaning pens, and result in problems with blocked slatted floors and slurry systems. Chopped straw might be more practical, but what is the effect on pigs’ behaviour of using chopped straw instead of long straw? Commercial pigs in 1/3 slatted, 2/3 solid pens of 15 pigs were provided with either 100 g/pig per day of long straw (20 pens) or of chopped straw (19 pens). Behavioural observations were made of three focal pigs per pen (one from each of small, medium and large weight tertiles) for one full day between 0600 and 2300 h at each of ~40 and ~80 kg. The time spent rooting/investigating overall (709 s/pig per hour at 40 kg to 533 s/pig per hour at 80 kg), or directed to the straw/solid floor (497 s/pig per hour at 40 kg to 343 s/pig per hour at 80 kg), was not affected by straw length but reduced with age. Time spent investigating other pigs (83 s/pig per hour at 40 kg), the slatted floor (57 s/pig per hour) or pen fixtures (21 s/pig per hour) was not affected by age or straw length. Aggressive behaviour was infrequent, but lasted about twice as long in pens with chopped straw (2.3 s/pig per hour at 40 kg) compared with pens with long straw (1.0 s/pig per hour at 40 kg, P=0.060). There were no significant effects of straw length on tail or ear lesions, but shoulders were significantly more likely to have minor scratches with chopped straw (P=0.031), which may reflect the higher levels of aggression. Smaller pigs showed more rooting/investigatory behaviour, and in particular directed towards the straw/solid floor and the slatted floor than their larger pen-mates. Females exhibited more straw and pen fixture-directed behaviour than males. There were no effects of pig size or sex on behaviour directed towards other pigs. In summary, pigs spent similar amounts of time interacting with straw/solid floor when long and chopped straw were provided, and most aspects of pig-directed behaviour and injuries were not affected by straw length. There was an increase in pigs with minor shoulder lesions with chopped straw, perhaps because of increased aggression. The use of chopped straw as an enrichment material for pigs warrants further investigation in larger and more detailed studies.     

Effect of rearing system and mixing at loading on transport and lairage behaviour and meat quality: comparison of free range and conventionally raised pigs

Free range pigs, born outdoors and reared after weaning in enriched indoor accommodation, were compared with conventionally raised pigs from a farm, matched for effects on meat quality, with respect to behaviour during transport and lairage, blood chemistry at slaughter and meat quality characteristics. Pigs were either kept in farm pen groups or were mixed at loading and kept in the groups, so formed, until slaughter. Free range pigs tended to settle faster during the 2½ h transport and 2 h lairage than conventionally raised pigs and were more likely to lie as resting posture during transport. Mixing at loading had no effect on posture during transport or during lairage for free range pigs but mixed conventionally raised pigs showed a greater variability in posture during lairage compared to non-mixed conventionally raised pigs, presumably as a result of disturbance from fighting conspecifics. Conventionally raised and free range pigs showed similar levels of aggression during transport but conventionally raised pigs were more aggressive during the lairage (average for mixed groups 12 v. 2 fights, P < 0.001). Aggressive interactions, such as one-way bites, were almost exclusively confined to mixed groups and all fights with mutual biting in mixed groups occurred solely between unfamiliar animals. The frequency of unacceptable skin damage in the middle and shoulder was highest in conventionally raised pigs and in mixed groups. Cortisol concentration and creatine kinase (CK) activity in slaughter blood were not affected by the rearing system. Mixing did not affect cortisol concentrations but led to higher CK activities compared to non-mixing (957 v. 588 U/l, respectively, P < 0.05). The rate of pH fall after slaughter was not affected by the rearing system but muscle temperatures early post mortem were highest in free range pigs. Mixing did not affect pH or temperature early post mortem. Neither rearing system nor mixing at loading affected ultimate pH or internal reflectance (meat quality marbling values).     

Pigs' aggressive temperament affects pre-slaughter mixing aggression, stress and meat quality

Pre-slaughter stress has a negative impact on animal welfare and on meat quality. Aggressive behaviour when pigs are mixed together for transportation to, or on arrival at, the abattoir is an important factor in pre-slaughter stress. Aggressiveness of pigs varies between individuals in the population, and this study investigated its effects on stress and meat quality at slaughter. We mixed pigs at a young age to identify individuals of high (H) or low (L) aggressive temperament using the previously validated approach of lesion scoring. To contrast extremes of social stress single-sex groups of eight pigs were mixed according to their aggressiveness in HH, HL or LL combinations or left unmixed (U) prior to transport and slaughter (n = 271). Each treatment was replicated in at least two groups in each of four slaughter batches. Mixing per se had little effect, but mixed groups composed of aggressive pigs (HH) had more carcass skin lesions and higher levels of plasma cortisol at slaughter and had loin muscle samples with higher pH at 24 h, and lower redness (a*) and yellowness (b*) compared to the other treatments. Females had higher levels of plasma cortisol at slaughter, a more rapid decline in pH post-slaughter and greater lean content of meat. Lactate and creatine kinase (CK) levels and meat pH were affected by the interaction of sex and treatment. Genetic factors, dam and sire line composition, and halothane locus (ryanodine receptor 1, RYR1) genotype, also affected a number of production and meat quality parameters as expected. Additionally, 'commercially normal' levels of social stress were studied in four further slaughter batches with no manipulation of group composition (n = 313). In these pigs, the proportion of unfamiliar pigs and group size of lairage groups explained limited variation in lesion scores at slaughter, but earlier aggressiveness did not. High numbers of skin lesions on the carcass were associated with high levels of cortisol and lactate and low glucose at slaughter, but not with meat quality measures. When stress and meat quality measures were compared for all pigs, high lactate was associated with low early pH and high drip loss, while high cortisol and CK were associated with high pH at 24 h and changes in meat colour. In conclusion, mixing pigs of above average aggressiveness resulted in greater aggression and stress, and changes in meat quality parameters, consistent with the effects of pre-slaughter stress on muscle chemistry.     

The use of olfactory and other cues for social recognition by juvenile pigs

Social recognition is essential for the maintenance of a stable group structure. Failure to recognise familiar conspecifics in social groups of juvenile pigs may initiate agonistic encounters that can compromise welfare and productivity. Current housing systems may allow build up of atmospheric ammonia that might, in turn, interfere with the olfactory system and compromise olfactory perception. In the present study, 16 juvenile pigs were housed in fresh air while another 16 pigs were kept in an ammoniated atmosphere (approximately 36ppm) for 1 week prior to test and another week during testing. We then assessed the role of olfaction in social recognition and determined whether chronic exposure to ammonia compromised discrimination based on olfactory perception by comparing the pigs' responses to selected cues from a familiar and an unfamiliar pig presented simultaneously in separate chambers of a modified Y-maze in each of two test situations (near, remote). Visual, auditory, olfactory, and tactile cues were all provided in the "near" test situation; here, the stimulus pigs were presented in two separate chambers behind clear perspex walls containing an aperture that allowed nose-to-nose contact between the test and stimulus pigs. On the other hand, the "remote" test provided only olfactory cues via air passed from the chambers containing the stimulus pigs into the test chamber. Each test lasted 5min and the pigs' behaviour was recorded via overhead video cameras; we then measured the accumulated times spent near and the numbers of visits made to the familiar and the unfamiliar stimulus pigs as well as the transitions between them. Overall, pigs made more visits to and spent significantly longer near both the stimulus pigs in the near test than in the remote one (ANOVA, P<0.001). They also made more transitions between the stimuli in the former test than the latter (P<0.001). Pigs from both the ammonia and the fresh-air treatment groups showed social discrimination. However, pigs that had received chronic exposure to ammonia visited the familiar pig more often and spent longer near it than the unfamiliar one regardless of the test situation (P<0.05) whereas those reared in fresh air spent longer near the unfamiliar animal (P<0.05). The present results suggest that pigs from both treatment groups employed olfactory cues in social recognition, but that chronic exposure to ammonia did not interfere with this ability. However, ammonia treatment seemingly affected social preferences, thus indicating an unknown and more fundamental effect of living in ammoniated atmospheres.     

Influence of social mixing and group size on skin lesions and mounting in organic entire male pigs

Alternatives to surgical castration are needed, due to stress and pain caused by castration of male pigs. One alternative is production of entire male pigs. However, changed behaviour of entire males compared with castrated males might adversely affect the welfare of entire males and changes in management procedures and production system might be needed. Elements from the organic pig production system might be beneficial in this aspect. The aim of this article is to investigate the effect of grouping strategy including social mixing and group size on levels of mounting behaviour and skin lesions, hypothesising that procedures that disrupt the social stability (e.g. regrouping) will have a larger negative effect in small groups compared with large groups. Approximately 1600 organic entire male pigs of the breed (Landrace×Yorkshire)×Duroc were reared in parallel in five organic herds, distributed across four batches in a 2×2 factorial design in order to test the influence of social mixing (presence or absence of social mixing at relocation) and group size (15 and 30 animals). Animals were able to socialise with piglets from other litters during the lactation period, and were all mixed across litters at weaning. A second mixing occurred at insertion to fattening pens for pigs being regrouped. Counting of skin lesions (1348 or 1124 pigs) and registration of mounting behaviour (1434 or 1258 pigs) were done on two occasions during the experimental period. No interactive effects were found between social mixing and group size on either skin lesions or mounting frequency. Herd differences were found for both mounting frequency and number of skin lesions. No association between skin lesions and mounting were revealed. Social mixing and group size were shown as interacting effects with herds on mounting frequency (P<0.0001), but with no consistent pattern across all herds. In addition, no effect of social mixing was found on mean number of skin lesions, but more lesions were observed in large groups (P<0.036). This could indicate that keeping entire male pigs in groups of 30 animals as compared with smaller groups of 15 may marginally decrease the welfare of these animals.     

Effects of previous experience on the agonistic behaviour of male crickets, Gryllus bimaculatus

Male solitary animals frequently enter aggressive interactions with conspecific individuals to protect their territory or to gain access to females. After an agonistic encounter, the loser (subordinate individual) changes its behaviour from aggression to avoidance. We investigated agonistic interactions between pairs of male crickets to understand how dominance is established and maintained. Two na?ve males readily entered into agonistic interactions. Fights escalated in a stereotyped manner and were concluded with the establishment of dominance. If individuals were isolated after the first encounter and placed together 15 minutes later, subordinate crickets tended to avoid any further contact with the former dominant opponent. Moreover, subordinate males also avoided unfamiliar dominant and na?ve opponents. They displayed aggressive behaviour only towards unfamiliar subordinate opponents. This suggests that the subordinate male change their behaviour depending on the dominance status of the opponent. Dominant crickets, in contrast, displayed aggressive behaviour towards familiar as well as unfamiliar opponents. If the interval between the first and second encounter was longer than 30 minutes, the former subordinate male showed aggressive behaviour again. However, if the subordinate cricket was paired with the same opponent three consecutive times within 45 minutes, it avoided the former dominant opponent for up to 6 hours following the third encounter. Our results suggest that the maintenance of dominance in male crickets depends largely on the behavioural change of subordinate individuals. Possible mechanisms to maintain dominance are discussed.     

Reducing Mouse Anxiety during Handling: Effect of Experience with Handling Tunnels

Handling stress is a well-recognised source of variation in animal studies that can also compromise the welfare of research animals. To reduce background variation and maximise welfare, methods that minimise handling stress should be developed and used wherever possible. Recent evidence has shown that handling mice by a familiar tunnel that is present in their home cage can minimise anxiety compared with standard tail handling. As yet, it is unclear whether a tunnel is required in each home cage to improve response to handling. We investigated the influence of prior experience with home tunnels among two common strains of laboratory mice: ICR(CD-1) and C57BL/6. We compared willingness to approach the handler and anxiety in an elevated plus maze test among mice picked up by the tail, by a home cage tunnel or by an external tunnel shared between cages. Willingness to interact with the handler was much greater for mice handled by a tunnel, even when this was unfamiliar, compared to mice picked up by the tail. Once habituated to handling, C57BL/6 mice were most interactive towards a familiar home tunnel, whereas the ICR strain showed strong interaction with all tunnel handling regardless of any experience of a home cage tunnel. Mice handled by a home cage or external tunnel showed less anxiety in an elevated plus maze than those picked up by the tail. This study shows that using a tunnel for routine handling reduces anxiety among mice compared to tail handling regardless of prior familiarity with tunnels. However, as home cage tunnels can further improve response to handling in some mice, we recommend that mice are handled with a tunnel provided in their home cage where possible as a simple practical method to minimise handling stress.     

The effect of group housing on feeding patterns and social behaviour of previously individually housed growing pigs

Group housed pigs make less frequent feeder visits of longer duration, and eat at a faster rate than pigs housed individually. They also have lower growth rates which may be due to elevated stress levels resulting from changes in the concentrations of hormones, such as cortisol and adrenaline associated with aggression and social stress. The aim of this experiment was to investigate the effects of grouping on feeding pattern, time budgeting and the social behaviour of pigs kept as individuals from weaning until grouping. In total, 12 Large WhitexLandrace male pigs (four pigs per block) mean (+/-S.E.) start weight 22.5+/-0.7kg were housed individually for 3 weeks (Period 1) after which in two replicates (Blocks 1 and 3), pigs were combined into a group of four (Period 2) before being returned to individual housing for a further 3 weeks (Period 3). In Block 2, the four pigs remained as individuals across periods but were moved between pens at the end of Periods 1 and 2 to account for any pen effects. Feeding pattern and food intake were recorded throughout and pigs were weighed three times a week. Video recordings and live behavioural observations were made to record time budgets and social behaviour. Grouped pigs made less visits to the feeder in Period 2 than when they were housed individually in Periods 1 (P<0.001) and 3 (P<0.01). Visit duration was longer in Period 2 than in Periods 1 (P<0.01) and 3 (P<0.05). Food intake and weight gain were greater in Period 3 than in Periods 1 and 2 (P<0.001 and P<0.01, respectively). There were no significant effects of moving pigs between pens in Block 2 on feeding behaviour and timebudgets. In Period 2, grouped pigs slept more (P<0.01) and spent less time feeding (P<0.01) and rooting (P<0.01) than in Periods 1 and 3. The frequency of aggression decreased over time from mixing (P<0.001). Possible explanations for the changes in feeding behaviour when pigs are moved from individual to group housing are competition, group cohesion, or that the high frequency of feeder visits when the pigs are housed individually is a consequence of a lack of social stimulation. Of these different possibilities, the results suggest that group cohesion is most likely to have been causal in the observed changes in feeding behaviour.     

Shoaling Decisions in Angelfish: The Roles of Social Status and Familiarity

Abstract Past research has shown that angelfish, Pterophyllum scalare , are capable of discriminating between shoals composed of familiar dominant and subordinate companions, whereas they show no preference for shoals of unfamiliar conspecifics. In this study, the relative importance of familiarity and social status (shoal factors) on the shoaling decision of juvenile angelfish, which also differed in social status (individual factor), was investigated as very little is known about such tradeoffs in fishes. Dominant and subordinate individuals were given the choice to shoal with a group of conspecifics composed of familiar dominants vs. unfamiliar dominants and composed of familiar subordinates vs. unfamiliar subordinates. The findings demonstrate that fish with different social status differed in their shoaling preference. Subordinate test fish showed a preferential association with familiar subordinates over unfamiliar subordinates, but preferred the unfamiliar shoal over the familiar one when both shoals were constituted by dominant individuals. The shoaling behaviour shown by dominant test fish, on the other hand, indicated no significant preference for any of the shoals regardless of their composition. A replicate preference test carried out 2 h 30 min after the first one indicated that the association pattern was relatively consistent. Results suggest that angelfish are able to differentiate between the stimulus shoals and demonstrate that the pervasive influence of familiarity on the shoaling decision may be restrained or overridden by the composition of the familiar shoals and the social status of the test fish.     

Social familiarity relaxes the constraints of limited attention and enhances reproduction of group‐living predatory mites

In many group‐living animals, within‐group associations are determined by familiarity, i.e. familiar individuals, independent of genetic relatedness, preferentially associate with each other. The ultimate causes of this behaviour are poorly understood and rigorous documentation of its adaptive significance is scarce. Limited attention theory states that focusing on a given task has interrelated cognitive, behavioural and physiological costs with respect to the attention paid to other tasks. In multiple signal environments attention has thus to be shared among signals. Assuming that familiar neighbours require less attention than unfamiliar ones, associating with familiar individuals should increase the efficiency in other tasks and ultimately increase fitness. We tested this prediction in adult females of the group‐living, plant‐inhabiting predatory mite Phytoseiulus persimilis. We evaluated the influence of social familiarity on within‐group association behaviour, activity, predation and reproduction. In mixed groups (familiar and unfamiliar), familiar predator females preferentially associated with each other. In pure groups (either familiar or unfamiliar), familiar predator females produced more eggs than unfamiliar females at similar predation rates. Higher egg production was correlated with lower activity levels, indicating decreased restlessness. In light of limited attention theory, we argue that the ability to discriminate between familiar and unfamiliar individuals and preferential association with familiar individuals confers a selective advantage because familiar social environments are cognitively and physiologically less taxing than unfamiliar social environments.     

Effects of Familiarity and Population Density on Competitive Interactions and Growth: An Experimental Study on a Territorial Salmonid Fish

The deleterious effect of competition for space and food in animals increases with increasing population density. In contrast, familiarity towards conspecifics can relax the intensity of interference competition. Here, we hypothesized that familiarity towards conspecifics mitigates the effect of density‐dependent growth and dispersal behaviour in territorial animals. To test this, wild‐captured juvenile brown trout were subjected to two consecutive laboratory experiments. First, growth and fin erosion were measured for 40 d in a 2 × 2 factorial design manipulating density and familiarity. The density was manipulated via size of experimental tanks, while per capita food abundance and fish number was constant. All fish were subsequently exposed to an emergence test, giving them the option to leave their group and disperse to a novel unoccupied environment. The results show that familiarity increases growth and decreases the level of fin erosion (i.e. proxy of intensity of aggressive interactions). We found no significant effect of population density on growth rate. However, there was a tendency towards higher fin erosion in fish kept under high density. The growth of individuals was also affected by their size rank within the group, with the largest individuals in each group growing disproportionally faster than the rest of the group, probably due to their high social rank. However, the second and third fish in the size rank did not grow significantly faster and tended to suffer higher mortality than the rest of the group. During the emergence test, the largest individuals in the familiar groups left the shelter either as the first (six of 12 groups) or last (five of 12 groups) individual in the group, while no such pattern was observed in unfamiliar groups. Our results suggest that individuals in familiar groups receive less aggression and stress (i.e. fin damage) and grow faster than fish in unfamiliar groups. The mechanisms indicated in this laboratory study may be especially important in highly fecund organisms like fish which undergo density‐dependent bottlenecks during early life.     

Piglet- and stockperson-directed sow aggression after farrowing and the relationship with a pre-farrowing,human approach test

The aims of the experiment were to determine the incidence and severity of piglet- and stockperson-directed aggression in two different farrowing systems and to determine whether either could be predicted using behavioural and physiological measures recorded previously during a standard human approach test. Sixty-two gilts, permanently housed in groups of 4–5, were approach tested at 6–8 weeks prior to parturition. Gilts were moved to a handling area, fitted with a heart rate monitor and moved to a square test arena. After 2 min familiarisation, an unfamiliar human entered the pen and stood for 3 min against one wall. Then, the human approached the gilt and touched her snout. Behaviour and heart rate were recorded continuously. Social rank was determined for each gilt using pair-wise food competition tests. At farrowing, gilts were randomly assigned to either open pens or conventional crates. At parturition, litter size and piglet weights were recorded and causes of any deaths ascertained, with emphasis on those piglets dying due to savaging. The aggression directed at the stockperson during piglet weighing at birth, 7 days and 14 days of age was scored on a scale of 1–5 (non-aggressive to extremely aggressive). Five gilts savaged their piglets and five other gilts were particularly aggressive towards the stockperson. Savaging of piglets and levels of stockperson-directed aggression were higher in the open pen system than in farrowing crates. Gilts that savaged piglets were more likely to show ‘shy’ behaviour during the human approach test, whereas gilts that were highly aggressive towards the stockperson were more likely to show ‘bold’ behaviour in the same test. Stockperson-directed aggression was displayed consistently, both within the same lactation and between subsequent lactations. Stockperson-directed aggression was unrelated to piglet survival but did confer advantages in piglet growth rate between birth and 7 days of age.     

Social inertia in white-throated sparrows results from recognition of opponents

Social inertia is a term for the stability of dominance relationships despite changes in the intrinsic dominating abilities of opponents. In a standard test for social inertia, low-ranking birds in an established hierarchy receive implants with testosterone (treated) and high-ranking birds receive empty implants (untreated). Social inertia occurs when the treated birds remain subordinate to untreated opponents in these groups, despite evidence that similarly treated birds dominate untreated strangers. In previous demonstrations of social inertia, however, treated and untreated birds were returned to their original aviaries and tested with familiar opponents, and thus the effects of familiarity with the location and those of familiarity with opponents were not separated. To address this issue, we investigated social inertia in 16 groups of white-throated sparrows Zonotrichia albicollis. When low-ranking treated birds were placed in new aviaries with familiar high-ranking, untreated opponents (treatment S, same opponents), dominance relationships showed social inertia. When such birds were placed in new aviaries with unfamiliar opponents (treatment N, new opponents), testosterone influenced dominance. When groups of high-ranking, untreated birds acquainted with each other were placed with unfamiliar treated opponents (treatment G, grouped dominants), 'coat-tail' effects (dominance by association with high-ranking individuals) sometimes outweighed the effects of testosterone. Social inertia in this species is thus a result of familiarity with opponents, rather than familiarity with locations of encounters. Measurements of aggressive tendencies confirmed a previous report that social inertia suppresses activation of aggression by testosterone. White-throated sparrows can thus recognize their opponents, and this ability affects the expression of both dominance and aggression. Copyright 1999 The Association for the Study of Animal Behaviour.     

The effects of weight asymmetry and resource distribution on aggression in groups of unacquainted pigs

The relationship between weight asymmetry and aggression when mixing groups of unacquainted pigs in two different environments was investigated. Ten groups of 4 female (LandracexYorkshire), unacquainted pigs with a weight asymmetry of 3.1+/-0.2 kg (mean weight: 16.6+/-0.6 kg) between each pig, and another 10 groups with a weight asymmetry of 1.2+/-0.1 kg (mean weight: 14.1+/-0.1 kg), were mixed at the age of 7 weeks. Ten of the groups (five of each) were mixed in an experimental pen with a heterogeneous distribution of straw. The pen consisted of two main compartments with straw in one of them, and a passage area with concrete floor in between. The other 10 groups were mixed in the same pen, but with a homogenous distribution of straw (straw spread all over the pen).The results showed that fighting duration was significantly shorter in groups with large weight asymmetry than in groups with small weight asymmetry irrespective of the environment. The number of bites delivered during the fights in the heterogeneous environment was lower in groups with large weight asymmetry than in groups with small weight asymmetry. In the homogenous environment, however, there was no significant difference between groups with large and small weight asymmetries regarding the number of bites. The combination of a limited straw area and a small weight asymmetry resulted in the greatest number of bites. In groups with large weight asymmetry, the largest pig won around 50% of the fights, and 25% of the variation in the percentage of fights won was explained by the weight asymmetry.In groups with small weight asymmetry, less than 10% of the variation in the percentage of fights won could be explained by weight asymmetry. The two largest pigs delivered significantly more bites and spent significantly more time fighting than the smallest pig. The second largest pig received significantly more bites and body lesions than the smallest pig in the groups. Fights between the two largest pigs had a significantly longer duration than when other pigs were involved. The results were discussed in relation to sequential assessment theory and resource defence theory.     

The Temporal Lobes Differentiate between the Voices of Famous and Unknown People: An Event-Related fMRI Study on Speaker Recognition

It is widely accepted that the perception of human voices is supported by neural structures located along the superior temporal sulci. However, there is an ongoing discussion to what extent the activations found in fMRI studies are evoked by the vocal features themselves or are the result of phonetic processing. To show that the temporal lobes are indeed engaged in voice processing, short utterances spoken by famous and unknown people were presented to healthy young participants whose task it was to identify the familiar speakers. In two event-related fMRI experiments, the temporal lobes were found to differentiate between familiar and unfamiliar voices such that named voices elicited higher BOLD signal intensities than unfamiliar voices. Yet, the temporal cortices did not only discriminate between familiar and unfamiliar voices. Experiment 2, which required overtly spoken responses and allowed to distinguish between four familiarity grades, revealed that there was a fine-grained differentiation between all of these familiarity levels with higher familiarity being associated with larger BOLD signal amplitudes. Finally, we observed a gradual response change such that the BOLD signal differences between unfamiliar and highly familiar voices increased with the distance of an area from the transverse temporal gyri, especially towards the anterior temporal cortex and the middle temporal gyri. Therefore, the results suggest that (the anterior and non-superior portions of) the temporal lobes participate in voice-specific processing independent from phonetic components also involved in spoken speech material.