Review: Early life predisposing factors for biting in pigs.

The pig industry faces many animal welfare issues. Among these, biting behaviour has a high incidence. It is indicative of an existing problem in biters and is a source of physical damage and psychological stress for the victims. We categorize this behaviour into aggressive and non-aggressive biting, the latter often being directed towards the tail. This review focusses specifically on predisposing factors in early life, comprising the prenatal and postnatal periods up to weaning, for the expression of aggressive and non-aggressive biting later in life. The influence of personality and coping style has been examined in a few studies. It varies according to these studies and, thus, further evaluation is needed. Regarding the effect of environmental factors, the number of scientific papers is low (less than five papers for most factors). No clear influence of prenatal factors has been identified to date. Aggressive biting is reduced by undernutrition, cross-fostering and socialization before weaning. Non-aggressive biting is increased by undernutrition, social stress due to competition and cross-fostering. These latter three factors are highly dependent on litter size at birth. The use of familiar odours may contribute to reducing biting when pigs are moved from one environment to another by alleviating the level of stress associated with novelty. Even though the current environment in which pigs are expressing biting behaviours is of major importance, the pre-weaning environment should be optimized to reduce the likelihood of this problem.

Temperament and dominance relate to feeding behaviour and activity in beef cattle: implications for performance and methane emissions.

In beef cattle, feeding behaviour and activity are associated with feed efficiency and methane (CH4) emissions. This study aimed to understand the underlying traits responsible for the contribution of cattle behaviour to individual differences in feed efficiency, performance and CH4 emissions. A total of 84 steers (530±114 kg BW) of two different breeds (crossbreed Charolais and Luing) were used. The experiment was a 2×2×3 factorial design with breed, basal diets (concentrate v. mixed) and dietary treatments (no additive, calcium nitrate or rapeseed cake) as the main factors. The individual dry matter intake (DMI; kg) was recorded daily and the BW was measured weekly over a 56-day period. Ultrasound fat depth was measured on day 56. Based on the previous data, the indexes average daily gain, food conversion and residual feed intake (RFI) were calculated. The frequency of meals, the duration per visit and the time spent feeding per day were taken as feeding behaviour measures. Daily activity was measured using the number of steps, the number of standing bouts and the time standing per day. Agonistic interactions (including the number of contacts, aggressive interactions, and displacements per day) between steers at the feeders were assessed as indicators of dominance. Temperament was assessed using the crush score test (which measures restlessness when restrained) and the flight speed on release from restraint. Statistical analysis was performed using multivariate regression models. Steers that spent more time eating showed better feed efficiency (P=0.039), which can be due to greater secretion of saliva. Feeding time was longer with the mixed diet (P<0.001), Luings (P=0.009) and dominant steers (P=0.032). Higher activity (more steps) in the pen was associated with poorer RFI, possibly because of higher energy expenditure for muscle activity. Frequent meals contributed to a reduction in CH4 emissions per kg DMI. The meal frequency was higher with a mixed diet (P<0.001) and increased in more temperamental (P=0.003) and dominant (P=0.017) steers. In addition, feed intake was lower (P=0.032) in more temperamental steers. This study reveals that efficiency increases with a longer feeding time and CH4 emissions decrease with more frequent meals. As dominant steers eat more frequently and for longer, a reduction in competition at the feeder would improve both feed efficiency and CH4 emissions. Feed efficiency can also be improved through a reduction in activity. Selection for calmer cattle would reduce activity and increase feed intake, which may improve feed efficiency and promote growth, respectively.

Current available strategies to mitigate greenhouse gas emissions in livestock systems: an animal welfare perspective.

Livestock production is a major contributor to greenhouse gas (GHG) emissions, so will play a significant role in the mitigation effort. Recent literature highlights different strategies to mitigate GHG emissions in the livestock sector. Animal welfare is a criterion of sustainability and any strategy designed to reduce the carbon footprint of livestock production should consider animal welfare amongst other sustainability metrics. We discuss and tabulate the likely relationships and trade-offs between the GHG mitigation potential of mitigation strategies and their welfare consequences, focusing on ruminant species and on cattle in particular. The major livestock GHG mitigation strategies were classified according to their mitigation approach as reducing total emissions (inhibiting methane production in the rumen), or reducing emissions intensity (Ei; reducing CH4 per output unit without directly targeting methanogenesis). Strategies classified as antimethanogenic included chemical inhibitors, electron acceptors (i.e. nitrates), ionophores (i.e. Monensin) and dietary lipids. Increasing diet digestibility, intensive housing, improving health and welfare, increasing reproductive efficiency and breeding for higher productivity were categorized as strategies that reduce Ei. Strategies that increase productivity are very promising ways to reduce the livestock carbon footprint, though in intensive systems this is likely to be achieved at the cost of welfare. Other strategies can effectively reduce GHG emissions whilst simultaneously improving animal welfare (e.g. feed supplementation or improving health). These win-win strategies should be strongly supported as they address both environmental and ethical sustainability. In order to identify the most cost-effective measures for improving environmental sustainability of livestock production, the consequences of current and future strategies for animal welfare must be scrutinized and contrasted against their effectiveness in mitigating climate change.

Prediction of reduction in aggressive behaviour of growing pigs using skin lesion traits as selection criteria.

Aggression at regrouping is a common issue in pig farming. Skin lesions are genetically and phenotypically correlated with aggression and have been shown to have a significant heritable component. This study predicts the magnitude of reduction in complex aggressive behavioural traits when using lesion numbers on different body regions at two different time points as selection criteria, to identify the optimum skin lesion trait for selection purposes. In total, 1146 pigs were mixed into new social groups, and skin lesions were counted 24 h (SL24h) and 3 weeks (SL3wk) post-mixing, on the anterior, centre and posterior regions of the body. An animal model was used to estimate genetic parameters for skin lesion traits and 14 aggressive behavioural traits. Estimated breeding values (EBVs) and phenotypic values were scaled and standardised to allow direct comparison across multiple traits. Individuals with SL24h and SL3wk EBVs in the least aggressive 10% of the population were compared with the population mean to predict the expected genetic and phenotypic response in aggressive behaviour to selection. At mixing, selection for low anterior lesions was predicted to affect substantially more behavioural traits of aggressiveness than lesions obtained on other body parts, with EBVs between -0.21 and -1.17 SD below the population mean. Individuals with low central SL24h EBVs also had low EBVs for aggressive traits (-0.33 to -0.55). Individuals with high SL3wk EBVs had low EBVs for aggression at mixing (between -0.24 and -0.53 SD below the population mean), although this was predicted to affect fewer traits than selection against SL24h. These results suggest that selection against anterior SL24h would result in the greatest genetic and phenotypic reduction in aggressive behaviour recorded at mixing. Selection for increased SL3wk was predicted to reduce aggression at mixing; however, current understanding about aggressive behaviour under stable social conditions is insufficient to recommend using this trait for selection purposes.

Why are most EU pigs tail docked? Economic and ethical analysis of four pig housing and management scenarios in the light of EU legislation and animal welfare outcomes.

To limit tail biting incidence, most pig producers in Europe tail dock their piglets. This is despite EU Council Directive 2008/120/EC banning routine tail docking and allowing it only as a last resort. The paper aims to understand what it takes to fulfil the intentions of the Directive by examining economic results of four management and housing scenarios, and by discussing their consequences for animal welfare in the light of legal and ethical considerations. The four scenarios compared are: 'Standard Docked', a conventional housing scenario with tail docking meeting the recommendations for Danish production (0.7 m2/pig); 'Standard Undocked', which is the same as 'Standard Docked' but with no tail docking, 'Efficient Undocked' and 'Enhanced Undocked', which have increased solid floor area (0.9 and 1.0 m2/pig, respectively) provision of loose manipulable materials (100 and 200 g/straw per pig per day) and no tail docking. A decision tree model based on data from Danish and Finnish pig production suggests that Standard Docked provides the highest economic gross margin with the least tail biting. Given our assumptions, Enhanced Undocked is the least economic, although Efficient Undocked is better economically and both result in a lower incidence of tail biting than Standard Undocked but higher than Standard Docked. For a pig, being bitten is worse for welfare (repeated pain, risk of infections) than being docked, but to compare welfare consequences at a farm level means considering the number of affected pigs. Because of the high levels of biting in Standard Undocked, it has on average inferior welfare to Standard Docked, whereas the comparison of Standard Docked and Enhanced (or Efficient) Undocked is more difficult. In Enhanced (or Efficient) Undocked, more pigs than in Standard Docked suffer from being tail bitten, whereas all the pigs avoid the acute pain of docking endured by the pigs in Standard Docked. We illustrate and discuss this ethical balance using numbers derived from the above-mentioned data. We discuss our results in the light of the EU Directive and its adoption and enforcement by Member States. Widespread use of tail docking seems to be accepted, mainly because the alternative steps that producers are required to take before resorting to it are not specified in detail. By tail docking, producers are acting in their own best interests. We suggest that for the practice of tail docking to be terminated in a way that benefits animal welfare, changes in the way pigs are housed and managed may first be required.

Genetic associations of short- and long-term aggressiveness identified by skin lesion with growth, feed efficiency, and carcass characteristics in growing pigs.

The objective of this study was to investigate the genetic relationships between skin lesion traits in group housed growing pigs as a measure of short- (in a newly mixed group) and long- (in a socially stable group) term aggression and commonly used commercial performance measures: growth, feed intake, feed efficiency, and carcass traits. Data on 2,413 growing pigs (138 groups) were available. Pigs were mixed into new social groups of 18 animals, and skin lesions were counted 24 h (SL24h) and 5 wk (SL5wk) postmixing. The animal model was used to estimate genetic parameters for skin lesion traits, test daily gain, lifetime daily gain, daily feed intake, feed efficiency (calculated as test daily gain divided by daily feed intake), loin depth, back fat, and HCW. Skin lesions had a heritable component, ranging from 0.08 for anterior SL24h to 0.22 for central SL5wk and would, therefore, be suitable as a method of phenotyping aggression for selection purposes. Significant positive genetic correlations were found between SL24h and SL5wk (0.46 to 0.81). Positive genetic correlations were also found between SL24h (central and posterior body regions) or SL5wk (all body regions) and the production traits lifetime daily gain, test daily gain, and HCW (0.29 to 0.54). Central SL24h, anterior SL5wk, and posterior SL5wk were found to correlate positively with feed efficiency (0.39 to 0.50), suggesting that pigs with more lesions convert feed more efficiently. Where significant, the magnitude of phenotypic correlations was low but positive (0.07 to 0.10). These results suggest that, genetically, animals that receive many lesions show improved performance compared to those with few lesions, except for anterior SL24h, which had previously been shown to be genetically positively correlated with the initiation of nonreciprocal attacks. It may, therefore, be possible, via selection against anterior skin lesions at mixing, to reduce this form of 1-sided aggression without adversely affecting production traits.

Injurious tail biting in pigs: how can it be controlled in existing systems without tail docking?

Tail biting is a serious animal welfare and economic problem in pig production. Tail docking, which reduces but does not eliminate tail biting, remains widespread. However, in the EU tail docking may not be used routinely, and some 'alternative' forms of pig production and certain countries do not allow tail docking at all. Against this background, using a novel approach focusing on research where tail injuries were quantified, we review the measures that can be used to control tail biting in pigs without tail docking. Using this strict criterion, there was good evidence that manipulable substrates and feeder space affect damaging tail biting. Only epidemiological evidence was available for effects of temperature and season, and the effect of stocking density was unclear. Studies suggest that group size has little effect, and the effects of nutrition, disease and breed require further investigation. The review identifies a number of knowledge gaps and promising avenues for future research into prevention and mitigation. We illustrate the diversity of hypotheses concerning how different proposed risk factors might increase tail biting through their effect on each other or on the proposed underlying processes of tail biting. A quantitative comparison of the efficacy of different methods of provision of manipulable materials, and a review of current practices in countries and assurance schemes where tail docking is banned, both suggest that daily provision of small quantities of destructible, manipulable natural materials can be of considerable benefit. Further comparative research is needed into materials, such as ropes, which are compatible with slatted floors. Also, materials which double as fuel for anaerobic digesters could be utilised. As well as optimising housing and management to reduce risk, it is important to detect and treat tail biting as soon as it occurs. Early warning signs before the first bloody tails appear, such as pigs holding their tails tucked under, could in future be automatically detected using precision livestock farming methods enabling earlier reaction and prevention of tail damage. However, there is a lack of scientific studies on how best to respond to outbreaks: the effectiveness of, for example, removing biters and/or bitten pigs, increasing enrichment, or applying substances to tails should be investigated. Finally, some breeding companies are exploring options for reducing the genetic propensity to tail bite. If these various approaches to reduce tail biting are implemented we propose that the need for tail docking will be reduced.

Hepatic expression patterns in psychosocially high-stressed pigs suggest mechanisms following allostatic principles.

Psychosocial challenges are known to introduce cellular and humoral adaptations in various tissues and organs, including parts of the sympatho-adrenal-medullary system and hypothalamic-pituitary-adrenal axis as well as other peripheral tissue being responsive to cortisol and catecholamines. The liver is of particular interest given its vital roles in maintaining homeostasis and health as well as regulating nutrient utilization and overall metabolism. We aimed to evaluate whether and how response to psychosocial stress is reflected by physiological molecular pathways in liver tissue. A pig mixing experiment was conducted to induce psychosocial stress culminating in skin lesions which reflect the involvement in aggressive behavior and fighting. At 27 weeks of age, animals prone to psychosocially low- and high-stress were assigned to mixing groups. Skin lesions were counted before mixing and after slaughter on the carcass. Individual liver samples (n=12) were taken. The isolated RNA was hybridized on Affymetrix GeneChip porcine Genome Arrays. Relative changes of mRNA abundances were estimated via variance analyses. Molecular routes related to tRNA charging, urea cycle, acute phase response, galactose utilization, and steroid receptor signaling were found to be increased in psychosocially high-stressed animals, whereas catecholamine degradation and cholesterol biosynthesis were found to be decreased. In particular, psychosocially high-stressed animals show decreased expression of catechol-O-methyltransferase (COMT) which has been linked to molecular mechanisms regulating aggressiveness and stress response. The expression patterns of high-stressed animals revealed metabolic alterations of key genes related to energy-mobilizing processes at the expense of energy consuming processes. Thus, the coping following psychosocial challenges involves transcriptional alterations in liver tissue which may be summarized with reference to the concept of allostasis, a strategy which is critical for survival.

Short-term temperament tests in beef cattle relate to long-term measures of behavior recorded in the home pen.

Handling temperament tests for beef cattle have been related to production traits, with calmer temperaments having greater growth rates. In most tests of temperament or personality, observation of the animal takes place over a short period of time, sometimes completed in a matter of minutes. This study investigated whether behavior observed in a temperament test was reflective of the steer's behavior in the home pen. Indoor-housed, crossbred, Bos taurus beef steers (n = 67) were fitted with triaxial activity monitors (IceTags; IceRobotics Ltd., South Queensferry, Edinburgh, Scotland) and activity was recorded for 2 periods of 14 d each. Also, each steer was scored on 4 measures of temperament: 2 handling tests (flight speed and chute score) and 2 feeding behavior scores (aggression at feeders and ability to displace at feeders). Each temperament observation was repeated 4 times, with repeatability of the traits ranging from 0.23 (aggression) to 0.48 (flight speed). Activity measures derived from the accelerometer data, such as bout lengths, were found to be highly repeatable between the 2 periods of activity monitoring (repeatabilities of 0.67 and 0.70 for average lying bout duration and average standing bout duration, respectively). Steers with high flight speeds were also more active in the home pen (MotionIndex: rs = 0.35, P = 0.004; average step count: rs = 0.34, P = 0.005) than steers with low flight speeds. Steers that were more capable of displacing other steers at feeders had longer average standing bout durations (rs = 0.26, P = 0.036), which were more variable (standing time SD: rs = 0.27, P = 0.030), and lay down for less time (rs = -0.35, P = 0.004). No correlations were found between aggression at feeders or chute score and home pen behavior. Results of this study are the first to demonstrate that short-term temperament tests are related to longer-term behavior data in beef steers and these results should be taken into consideration for future research.

Precalving temperament and maternal defensiveness are independent traits but precalving fear may impact calf growth.

Human safety can be compromised by the response of beef cows to handling or when defending their calf. However, little is known about how precalving temperament, postcalving defensiveness, and maternal care are related. The impacts of cow temperament on calf neonatal vigor and ADG are also unknown. Data were collected on 2 farms (Farm 1, n = 143, 1 parity; Farm 2, n = 237, 2 parities). Temperament was recorded before calving when restrained in a crush (crush score), on exit from the crush (flight speed), and when isolated with a handler. Defensiveness was recorded within 4 d after calving during handling of the calf. Maternal interactions with the calf and calf vigor were recorded for 3 h after calving (Farm 1 only) and ADG was measured over 7 mo. Crush score and flight speed were repeatable within a parity (range in repeatability 0.33 to 0.49; P < 0.001). Crush score (0.50; P < 0.001) and defensiveness (up to 0.71; P < 0.001) were repeatable across parities. Temperament and defensiveness were unrelated on Farm 1; on Farm 2 a fearful crush score was associated with heightened defensiveness as measured by vigorous movement during calf handling (P < 0.001). Temperament and defensiveness were unrelated to calving ease or the amount of maternal behavior shown to the calf. At Farm 1, cows that exited the crush quickly had calves with a lighter birth weight (P = 0.023) and those that were agitated when isolated had calves with a decreased ADG (P = 0.017). Defensiveness was unrelated to ADG and neither temperament nor defensiveness affected calf vigor. Cow precalving temperament and postcalving defensiveness are repeatable but appear to be independent traits, neither of which is related to maternal interactions with the neonatal calf. Reducing precalving fearfulness should not affect postcalving behavior and changing postcalving defensiveness should not affect other maternal care traits. Fearful cows may produce calves with decreased birth weight and ADG, which, if confirmed, suggests that cow fearfulness may have wider economic implications than previously realized.

Board invited review: The importance of the gestation period for welfare of calves: maternal stressors and difficult births.

The prenatal period is of critical importance in defining how individuals respond to their environment throughout life. Stress experienced by pregnant females has been shown to have detrimental effects on offspring biology in humans and a variety of other species. It also is becoming increasingly apparent that prenatal events can have important consequences for the behavior, health, and productivity of offspring in farmed species. Pregnant cattle may experience many potentially important stressors, for instance, relating to their social environment, housing system and physical environment, interactions with humans and husbandry procedures, and their state of health. We examined the available literature to provide a review of the implications of prenatal stress for offspring welfare in cattle. The long-term effects of dystocia on cattle offspring also are reviewed. To ensure a transparent and repeatable selection process, a systematic review approach was adopted. The research literature clearly demonstrates that prenatal stress and difficult births in beef and dairy cattle both have implications for offspring welfare and performance. Common husbandry practices, such as transport, were shown to influence offspring biology and the importance of environmental variables, including thermal stress and drought, also were highlighted. Maternal disease during pregnancy was shown to negatively impact offspring welfare. Moreover, dystocia-affected calves suffer increased mortality and morbidity, decreased transfer of passive immunity, and important physiological and behavioral changes. This review also identified considerable gaps in our knowledge and understanding of the effects of prenatal stress in cattle.

Genetic associations between behavioral traits and direct-social effects of growth rate in pigs.

This study examined the behavioral consequences of selecting pigs using a social genetic model for growth. Calculations enable each member of a group of pigs to be given a direct breeding value (DBV) and a social breeding value (SBV), which can be summarized into a total breeding value (TBV) for growth. Selection for growth TBV could affect animal behavior because social effects account for within-group interactions. Data were recorded from 96 groups of Yorkshire and Yorkshire × Landrace pigs in a nucleus herd. Each group contained 15 pigs fed ad libitum from 2 feeders; the space allowance was 0.85 m2/pig. Average daily gain was quantified from 35 to 100 kg of BW. Fighting and bullying activity at mixing (period 1), lying frequency 3 wk after mixing (period 2), and counts of skin lesions in periods 1 and 2 were recorded. The DBV for these traits were estimated with a classic animal model. We simulated different correlations between the direct genetic effect and the social genetic effect on growth rate (r(DS)), 2 components that respectively determine a pig's genetic capacity to grow and its genetic influence on growth of group mates: r(DS) was successively assumed to be 0 and ±0.12, ±0.20, ±0.29, and ±0.58. Finally, the correlations between DBV, SBV, and TBV for ADG, as well as the DBV for behavior and skin lesions, were calculated and tested for a level of significance at P < 0.05. The gradient from negative to positive values of r(DS) refers to a progressive path running from genetic antagonism to genetic mutualism for growth. If rDS in the population truly ranged between -0.58 and -0.20, correlations for TBV for ADG with DBV for fighting and bullying progressively increased with rDS. Consequently, if rDS was low (between -0.12 and +0.12) or positive (>+0.12), pigs with high TBV for ADG had higher DBV for bullying other pigs in the group and for fighting than pigs with lower TBV for ADG. Pigs with high TBV for ADG did not differ from other pigs in their DBV for lesions to the anterior part of the body, but they had a lower DBV for posterior lesions, whereas in period 2, they had higher DBV for posterior lesions and lower DBV for lying. Under genetic mutualism for growth and in housing conditions similar to those in the present study, selection for growth TBV would promote the rapid establishment of the dominance relationships, with more aggressive contests among group mates at mixing. Pigs would subsequently be more active but, judging by skin lesions, less willing to fight in a more stable social situation.

Associations between response to handling and growth and meat quality in frequently handled Bos taurus beef cattle.

Fearful behavioral responses to handling (temperament) are undesirably associated with ADG and meat quality in infrequently handled Bos indicus cattle. It has never been assessed whether these relationships exist in calmer Bos taurus breeds in systems where handling is more frequent. Such systems predominate in some countries where beef production is a major agricultural activity. During fattening, 144 crossbred cattle from Limousin and Aberdeen Angus sires were assessed for temperament using 4 approaches: response to movement along a race (race score; 4 occasions), restraint in a crush (crush score; 4 occasions), flight speed from the crush (flight speed; 4 occasions), and isolation in a pen with a human (isolation score; 1 occasion in yr 1, 2 occasions in yr 2). Measurements of ADG were made between birth and slaughter and between 16 and 18 mo of age during fattening. Fattening occurred indoors on a complete mixed diet fed for ad libitum intake. Meat quality was measured by pH, color, and Volodkevitch shear force and by a sensory panel. The repeatability of temperament traits was 0.17 (race score), 0.35 (crush score), 0.51 (flight speed), and 0.36 (isolation score). The proportion of the total variance of temperament traits attributable to the sire and the social group was low (0.003 to 0.402). However, the sire did affect behavior in all tests apart from the crush score (ranging from P = 0.02 to P < 0.001). Correlations between behavior in the different tests (ranging from r = 0.21 to 0.54, and P = 0.02 to P < 0.001) apart from between-flight speed and isolation score indicate that fearful behavior was consistently shown across assessment methods. A calm response in the crush score test was associated with a greater ADG during fattening (P = 0.05), whereas a calm response during the isolation test was associated with a greater ADG in cold carcass weight (P = 0.02). Animals with a calm isolation score had less tender meat as judged by the sensory panel (P = 0.03), but no other effects were apparent between temperament and meat quality measures, although several tendencies (0.06 ≤ P ≤ 0.10) were found. Temperament did not appear to relate to meat quality in this study of frequently handled Bos taurus genotypes, which is in contrast to other studies using different beef production systems. Genetic correlations between temperament and meat quality under these conditions could, although not measured in this study, still exist in the absence of phenotypic correlations.

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.

Genetic validation of postmixing skin injuries in pigs as an indicator of aggressiveness and the relationship with injuries under more stable social conditions.

The objective of the study was to estimate genetic correlations between skin lesions and aggressive behavior postmixing and under more stable social conditions as a potential means of selecting against pig aggressiveness. Postmixing aggression in commercial pig production is common, compromises welfare and profitability, and cannot be significantly reduced by low-cost changes to the environment. A genetic component to individual aggressiveness has been described in pigs and other species. Selective breeding against aggressiveness ought to be possible if an easily measured indicator trait can be shown to be genetically associated with aggressive behavior. Aggressive behavior was recorded continuously for 24 h after mixing, and a count of skin lesions (lesion count, LC) was recorded at 24 h and 3 wk postmixing on 1,663 pigs. Two behavioral traits were found to have a moderate to high heritability similar to that of growth traits; duration of involvement in reciprocal fighting (0.43 +/- 0.04) and delivery of nonreciprocal aggression (NRA; 0.31 +/- 0.04), whereas receipt of NRA had a lower heritability (0.08 +/- 0.03). Genetic correlations (r(g)) suggested that lesions to the anterior region of the body 24 h after mixing were associated with reciprocal fighting (r(g) = 0.67 +/- 0.04), receipt of NRA (r(g) = 0.70 +/- 0.11), and to a lesser extent, delivery of NRA (r(g) = 0.31 +/- 0.06). Lesions to the center and rear were primarily genetically associated with receipt of NRA (r(g) = 0.80 +/- 0.05, 0.79 +/- 0.05). Genetic correlations indicated that pigs that engaged in reciprocal fighting delivered NRA to other animals (r(g) = 0.84 +/- 0.04) but were less likely to receive NRA themselves (r(g) = -0.41 +/- 0.14). A genetic merit index using lesions to the anterior region as one trait and those to the center or rear or both as a second trait should allow selection against animals involved in reciprocal fighting and the delivery of NRA. Positive correlations between LC 24 h and 3 wk after mixing were found, especially for lesions to the center and rear of the body, indicating that postmixing lesions are predictive of those received under more stable group conditions. As well as reducing immediate aggression at mixing, selection on postmixing LC is expected to have a long-term impact on injuries from aggression, even after dominance relationships are established.

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.

Bayesian analysis of genetic associations of skin lesions and behavioural traits to identify genetic components of individual aggressiveness in pigs.

There is increasing interest in genetic selection against behavioural traits that impact negatively on welfare and productivity in commercial livestock production. Post-mixing aggressiveness in pigs shows wide phenotypic variation, affects health, welfare and growth performance and is a routine feature of production. A Bayesian approach was used to estimate the heritability of three traits associated with aggressiveness in pigs during the 24 h post-mixing; duration in reciprocal aggression, and in receipt of, or delivery of non-reciprocal aggression (NRA). For the purposes of genetic selection, recording aggressive behaviour is excessively labour intensive. The genetic correlations were quantified between the behavioural traits and an easily measurable indicator trait; the number of skin lesions following mixing (lesion score, LS). The heritabilities for the three behavioural traits ranged from 0.17 to 0.46 (receipt of NRA and reciprocal aggression respectively). The duration in reciprocal aggression and in delivery of NRA showed a strong genetic correlation (r g = 0.79 with 95% Bayesian credibility interval of 0.62-0.94). The genetic correlation between LS and these two behaviours indicated that selection on breeding values of LS could be used to reduce aggressiveness. The duration in receipt of NRA appeared to be regulated by different genes or genomic effects compared with the other behavioural traits and LS. Although duration in receipt of NRA was not genetically associated with LS, it was lowly but significantly environmentally associated with the residuals of central and caudal LS (r e = 0.28-0.32), indicating that pigs that received NRA also received bites on the central and caudal third of the body. The pen that the animals were mixed into was found to be a very important factor for the analysed traits, in particular those representing behavioural characteristics. Based on the estimated genetic parameters, it is concluded that selection on breeding values for reduced LS (especially central LS) is expected to reduce reciprocal aggression and the delivery of NRA but will not change the receipt of NRA directly.

On the time-scales of magmatism at island-arc volcanoes.

Precise information on time-scales and rates of change is fundamental to an understanding of natural processes and the development of quantitative physical models in the Earth sciences. U-series isotope studies are revolutionizing this field by providing time information in the range 10(2)-10(4) years, which is similar to that of many modern Earth processes. I review how the application of U-series isotopes has been used to constrain the time-scales of magma formation, ascent and storage beneath island-arc volcanoes. Different elements are distilled-off the subducting plate at different times and in different places. Contributions from subducted sediments to island-arc lava sources appear to occur some 350 kyr to 4 Myr prior to eruption. Fluid release from the subducting oceanic crust into the mantle wedge may be a multi-stage process and occurs over a period ranging from a few hundred kyr to less than one kyr prior to eruption. This implies that dehydration commences prior to the initiation of partial melting within the mantle wedge, which is consistent with recent evidence that the onset of melting is controlled by an isotherm and thus the thermal structure within the wedge. U-Pa disequilibria appear to require a component of decompression melting, possibly due to the development of gravitational instabilities. The preservation of large (226)Ra disequilibria permits only a short period of time between fluid addition and eruption. This requires rapid melt segregation, magma ascent by channelled flow and minimal residence time within the lithosphere. The evolution from basalt to basaltic andesite probably occurs rapidly during ascent or in magma reservoirs inferred from some geophysical data to lie within the lithospheric mantle. The flux across the Moho is broadly andesitic, and some magmas subsequently stall in more shallow crustal-level magma chambers, where they evolve to more differentiated compositions on time-scales of a few thousand years or less.