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Audit Form - Welfare of Cattle in Feedlots (Updated Septebmer 2018)

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2. Cattle Exit Speed Score - Percentage of cattle that walk or trot out of the squeeze chute. 75% acceptable and 90% excellent. Fail if more than 2% fall while exiting the squeeze chute. Cattle that run or jump when they exit the squeeze chute will have lower weight gains.

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Cattle ingest a grain based feed and they are grazers. They like to have open access to land to freely graze. They are ruminants therefore they chew their cud. So, they will regurgitate their food and chew it for a second time. https://climateprotection.org/gotcha-happy-april-fools-day/10144-black-angus-cows-grazing-in-a-pasture-pv/

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Young will nurse from their mothers soon after birth.

Young will stay near their mother.

Young will try to stand and walk for the first time, may have difficulty at first.

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Mother will lick their young to groom them.

Mother will ingest her placenta to clean up the area.

Mother will be still to allow their young to nurse.

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It is common for females to mount other cows while in heat.

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It is common for males to sniff females to know if they are ready to be bred.
https://www.slideshare.net/rhfayed/reproductive-behaviour-1sexual-behaviour-in-animals

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The type of dominance hierarchy mostly seen in cattle is head to head fighting. Head butting.
I learned this in class.

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Pictured are two bulls exhibiting agonistic behavior by butting heads. They may be showing their dominance. One is showing that he  has power over the other and that this is his territory. https://fineartamerica.com/shop/canvas+prints/black+angus+bull?page=5

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Visual- Cattle use their sight to see where other cattle are relevant to where they need or want to be.

Auditory- "mooing" is used to call out certain names for other cattle and for when they need help.

Tactile- Cattle will head butt to assert dominance over others.

Chemical/smell- Cattle use their
sense of smell to detect an undesirable situation and to know when to mate and whom with.

http://www.publish.csiro.au/ebook/chapter/9781486301614_Chapter4
https://www.telegraph.co.uk/news/newstopics/howaboutthat/11297269/Cows-communicate-using-individual-sounds-like-human-names.html

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https://www.grandin.com/inc/cattlecorrals.html

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Below is a simple typical cattle handling facility. This system utilizes an alley, a sweep, chute and a head gate. The system is meant to gather and hold the cattle in one place. This system is using negative punishment by removing the animals freedom. A positive punishment may be portrayed when head gate pressure is applied. A positive reinforcement may be when the animal is let go after cooperating and the head gate is loosened.

http://tarterusa.com/product-lines/cattle-handling/

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Cattle are color blind to red. Therefore, a trainer would not want to pick this color to signal cattle to do something it is asked.
https://www.livescience.com/33700-bulls-charge-red.html

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An example of a sensitive period for a Black Angus calf would be when they are being weaned, they are transitioning from their mothers milk to solid foods. https://www.beefmagazine.com/mag/beef_weaning_twostep

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I chose Black Angus as my animal, because I would like to know more about them. They are often bred and raised for their meat. They are very commonly raised in Illinois. From what I know, they have good temperament. They are huge cows, when they get full size. I find them to be friendly. I also think that they are a very cool animal, because of how large and strong they are. Growing up I would always see hundreds of them out in fields, on livestock farms. I never really knew much about them. I am excited to learn more about them and their behaviors.

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In many species behavior can be linked to genetics. In this article, temperament is discussed as something that is linked to genetics. There are many different tests that are done to see how different breeds react in different situations. Breeders will choose to breed certain animals for desired behaviors.

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Abstract

Animal temperament can be defined as a response to environmental or social stimuli. There are a number of temperament traits in cattle that contribute to their welfare, including their response to handling or milking, response to challenge such as human approach or intervention at calving, and response to conspecifics. In a number of these areas, the genetic basis of the trait has been studied. Heritabilities have been estimated and in some cases quantitative trait loci (QTL) have been identified. The variation is sometimes considerable and moderate heritabilities have been found for the major handling temperament traits, making them amenable to selection. Studies have also investigated the correlations between temperament and other traits, such as productivity and meat quality. Despite this, there are relatively few examples of temperament traits being used in selection programmes. Most often, animals are screened for aggression or excessive fear during handling or milking, with extreme animals being culled, or EBVs for temperament are estimated, but these traits are not commonly included routinely in selection indices, despite there being economic, welfare and human safety drivers for their. There may be a number of constraints and barriers. For some traits and breeds, there may be difficulties in collecting behavioral data on sufficiently large populations of animals to estimate genetic parameters. Most selection indices require estimates of economic values, and it is often difficult to assign an economic value to a temperament trait. The effects of selection primarily for productivity traits on temperament and welfare are discussed. Future opportunities include automated data collection methods and the wider use of genomic information in selection.

Keywords: temperament, animal welfare, genetic variation, animal personality, genetic correlation

Introduction

Genetic improvement, including selection between breeds, crossing and within-breed selection, is widely used in farm livestock and has led to dramatic changes in performance in dairy and beef cattle over the last 50 years or so (e.g., Simm, 1998). Historically, most emphasis has been on traits that are most directly associated with profitability, and most easily measured, such as milk yield or body weight. However, selection between or within breeds for a broader set of traits, including health and "fitness" traits, is becoming more widespread as producers realize that productivity can only be maintained or improved with a more holistic view of animal performance. Reproduction, longevity and health traits are used in a number of breeding programmes for dairy and beef cattle, and there is growing interest in behavioral traits associated with animal welfare and ease of management. Temperament traits such as fearfulness or aggressiveness are important to consider as they affect how the animal responds to the husbandry and handling conditions on the farm and during procedures like transport. The aim of this review is to determine what progress has been made in the steps in the chain from trait definition through to the use of these traits in selection, including the recent opportunity for genomic selection. We also review the research that has investigated associations between temperament traits and productivity, health and reproductive traits to determine whether selection for these traits may be altering temperament indirectly.

What is temperament?

Farmers and others involved with the keeping of cattle and other livestock are well aware that there are differences between individual animals in their behavioral response to alarming or challenging situations. Furthermore, individuals are often consistent in the way they respond when the challenge is repeated. In cattle, the magnitude of response, and the difference between animals are of most importance to humans in situations that involve human interaction, such as where animals are handled, moved or milked. Some animals are calm and docile, while others are distressed and struggle to escape. Animals may also show consistency in their response in other situations, such as response to a new-born calf, and aggression or affiliation toward herd-mates.

This observed consistency of response within the animal, and the variation shown between individual animals or groups of animals, has historically been given a number of different labels, depending on whether the user is from a psychological, farm livestock or behavioral ecology background. In human psychology, it is known as personality, while in behavioral ecology the term "behavioral syndrome" is used to describe differences in clustering of traits between animal populations. In animal husbandry settings, the term "temperament" is largely used. In cattle, temperament is often described as an animal's response to handling or forced movement by humans (Tulloh, 1961; Burrow, 1997). This definition appears to have come from the terminology that farmers use to describe the way their cattle behave during handling (e.g., Hassall, 1974, a paper from a beef producer). It is also similar to the term "disposition" used in North America (Beef Improvement Federation Guidelines, 2010). This human-focussed definition of temperament has been used broadly across the cattle sector, particularly in beef cattle. A number of authors have used the term "temperament" with a situation "specifier" to describe the context (e.g., Brown, 1974 uses the term "maternal protective temperament"). Thus the term "handling temperament" can be used to differentiate the response from other contexts. The use of terms such as "maternal temperament" and "aggressive temperament" or simply a descriptor term such as "aggressiveness" and "sociability" are found in studies that consider consistency in the animal's response in contexts other than handling (Brown, 1974; Kilgour and Dalton, 1984; Reale et al., 2007; Gutierrez-Gil et al., 2008; Gibbons et al., 2009a, 2010).

Why is temperament important?

The temperament traits that have received most attention are generally those that have adverse production, welfare or human safety consequences. The foremost of these is handling temperament, and the impacts of poor temperament on farm management efficiency and animals has been a key driver for many studies (e.g., Burrow, 1997; Barrozo et al., 2012). A beef animal that responds to confinement in a chute, weigh crush or handling race by struggling violently and trying to escape is at a higher risk of injury to itself, human handlers and other animals than an animal that responds calmly (Voisinet et al., 1997a). This type of animal is also more likely to make the process of handling a group of animals for weighing or drafting much slower and less efficient. A number of studies have shown that handling temperament is also linked to growth, feeding efficiency and meat quality in beef cattle. Understanding the extent of this association has driven a great deal of research that will be discussed below. For dairy cattle, a calm response to the milking procedure is important both to maximize the efficiency of the milking process and to minimize the residual milk volume. Docility in dairy cattle at milking and during handling is a trait that has been under selection informally and formally for generations, so extreme responses are rare. However, the problems created by an animal that is not easy to handle and milk mean that "dairy temperament" (which is measured as strength of response to the milking procedure) has been investigated and is still part of many dairy breeding programmes worldwide (Interbull: www.interbull.org/ib/geforms).

There are other temperament traits that have received less attention in the literature, but are important from an animal welfare or human safety standpoint. Maternal aggressiveness, where a dam shows defensive aggressiveness toward any human or animal attempting to interfere with her calf, is a trait that clearly had evolutionary advantages for wild animals, and still does in some extensive production environments. However, when this aggression is directed at stockworkers or members of the public entering grazing fields, it clearly becomes much more problematic (Turner et al., 2013). Other traits that are important for welfare include resource-based aggression, where an animal shows aggression toward another when in competition for a resource such as feed or water, and social motivation or sociability, which is the willingness to be in close proximity to group-mates.

Trait definition and measurement

Given the potential adverse effects of excitable temperament on human safety and handling efficiency, the use of selective breeding to improve temperament is important. A number of steps must be taken to enable selection to take place (Figure ). Firstly, the trait (in this case a behavior or response) must be defined, which typically includes a definition of the context in which it is important. The next step is to devise a measurement system so that the trait can be assessed in a rapid, quantifiable and reliable way by non-scientists, and then to validate it against other measures of the trait if possible, so that the chosen measure accurately characterizes the response. This measure can then be used in a number of ways. It can be used as a "screening" tool, such as when individual animals with poor scores for a temperament trait are culled or not considered for breeding, or the measure can be used as part of a genetic improvement programme. This section will investigate the progress with regards to trait definition and measurement.

Handling-beef cattle

Fearful or excitable responses may be expressed by animals in many novel or challenging contexts, such as during interactions with other animals or when entering a new field or pen, but it is largely during handling that this characteristic becomes a problem. A fearful response to handling manifests itself in a variety of ways. Animals may struggle, show agitated movements, attempt to escape, vocalize, show increased respiration rates, defaecate, show changes in their ear, head and tail positions and facial expressions and be more or less motivated to move away from the handling area or handler. The challenge is to find a scale or measure that adequately represents these varied responses. In beef cattle, there are some very well established assessments: flight speed or flight time, chute (known as a crush in Australasia/Europe) score and the docility score. These have sometimes been grouped into restrained and non-restrained categories (Burrow, 1997). Restrained tests are primarily those assessing the response to restraint in a handling chute, confinement in a pen or raceway, or alternatively measuring the response to that confinement by assessing the flight time or speed to move away from the place of confinement. Unrestrained tests are those in which the animal is not confined, but the animal's response to being approached, moved or handled is scored. These unrestrained tests are also characterized by more directly measuring the response to proximity to a human, whereas the restrained tests may measure the response to physical restraint as well as the proximity to humans. The main tests are described below.

Handling and milking-dairy cattle

Typically, milking temperament is seen as the response to the whole milking procedure, and is mostly scored by the farmer or milking staff. A categorical scale based on descriptive definitions of different levels of response to the milking and handling procedures are often used, with scores from 1-5 or 1-9 typically representing poor to good milking temperament. Temperament scores are often combined with other assessments such as milking speed to derive a "workability" trait. Milking temperament data are collated by herd improvement or milk recording organizations in many countries ( www.interbull.org). A number of researchers have used more objective assessments such as an assessment of the number of steps, kicks or flinches the cow makes in response to the milking procedure (e.g., Willis, 1983; Breuer et al., 2000). In experimental situations, human approach or flight distance tests have been used with dairy cattle (e.g., Waiblinger et al., 2003; Gibbons et al., 2009b), and shown to have good within-animal repeatability. The tests involve scoring the response of the animal as the experimenter moves toward her. Gibbons et al. ( 2011) found that approach distance was related to flight speed, but not chute score, in dairy heifers.

Other traits

Other temperament traits such as sociability, intra-specific aggression and response to novelty and social separation have been assessed in beef and dairy cattle, and maternal behavior in beef cattle. Maternal behavior, or maternal aggressiveness is a human safety as well as a calf survival issue, particularly in farming systems in which humans come in close contact with cows and calves (Turner et al., 2013). Improvement of animal welfare is the main driver for assessing many of the other traits, as well as the desire to understand the relationship between the specific handling tests and the wider personality of the animal (Kilgour et al., 2006). It is thought that animals that are not excessively fearful of novel objects or isolation from other animals will cope better with modern intensive or semi-intensive farming systems than more reactive animals (Kilgour et al., 2006; Gibbons et al., 2009b). Similarly, it has been hypothesized that an animal with high social motivation will integrate and cope better with group housing than low sociability animals, and that animals showing low aggression will suffer less stress and have less negative impact on other animals (Gibbons et al., 2009a, 2010). Methods to quantify these characteristics have involved assessing the response of animals to novel objects, social isolation or to a competitive situation. A number of studies have shown moderate to high repeatabilities of scores for individual animals, indicating that they can be classed as temperament traits (e.g., novelty: Kilgour et al., 2006; Gibbons et al., 2009b, 2010; aggression: Gibbons et al., 2009a; MacKay et al., 2013).

In terms of trait definition, it would appear that there are some very good definitions for a number of temperament traits, particularly for beef handling and dairy cow milking temperament. These traits have established measurement protocols and measurement scales. There are other traits that have received less attention, but which show good repeatability.

Genetic variation between and within breeds

Once a trait has been defined and a reliable measurement system created, the degree of genetic variation within and between breeds must be determined, if genetic improvement is to be made. Genetic variation can be exploited in one of three ways currently-selection between breeds (or breed substitution-replacing one breed with another, superior breed), crossbreeding (crossing different breeds to create animals with intermediate performance to the parent breeds, or to produce animals with attributes of both parental breeds, or to exploit heterosis or "hybrid vigor"-the boost in performance often seen in crosses, over and above that expected from the mean performance of the parent breeds), or selection within breeds. Whether crossbreeding leads to significant heterosis effects on temperament traits such as handling ease has not been studied, but warrants investigation (Burrow, 1997). A fourth option, direct genetic modification, is also available, but this is largely confined to experimental use rather than commercial practice at the moment. This may change as techniques such as "gene editing" used in human gene therapy begin to be applied to allow targeted changes in livestock (Lilico et al., 2013).

Breed differences

The choice of breed or strain by producers is influenced by temperament, but choice is often based on subjective information. Differences in performance of breeds managed in the same environment provide more objective evidence that a trait is under genetic control. Substitution of one breed by another is a rapid way to effect genetic change. Information on the differences between breeds and their crosses is also a prerequisite for the design of optimal crossbreeding schemes. Stark differences in handling ease between the relatively docile Bos taurus and relatively flighty Bos indicus cattle are well known (Hearnshaw et al., 1979; Becker and Lobato, 1997; Voisinet et al., 1997a; Buchenauer, 1999; Burrow, 2001). Large differences between individual breeds of Bos taurus cattle have also been demonstrated, although individual reports often conflict (e.g., Hearnshaw and Morris, 1984; Gauly et al., 2001; Boissy et al., 2005; Hoppe et al., 2010). There are also reported differences between the dairy breeds in their milking temperament (Sewalem et al., 2010). In many cases, these reported differences are most likely to be due to differences in the way in which cattle from the different breeds were raised and their level of exposure to humans. However, in those studies in which the rearing environment was standardized, breed differences have still been found, indicating that the response of cattle to handling by humans is, at least at the level of the breed, under some genetic control. Other than the distinction between Bos indicus and Bos taurus, studies in which different breeds have been reared and handled together in a standardized manner are not numerous enough to allow a "league table" of breed temperament to be created at present.

Genetic variation within breeds

Having identified the optimal breeds or crosses for a given production system, there are opportunities for further genetic improvement via selection of the best parents within the chosen breed, or within each of the breeds making up the chosen crossbreed. Objective within-breed selection usually requires knowledge of the traits affecting profitability (breeding goal traits) and their relative economic values, potential proxy traits on which to base selection (selection criteria) if breeding goal traits can not be measured directly (e.g., if they are expressed late in life, can only be measured post-mortem or are time-consuming and costly to measure), and estimates of the genetic and phenotypic variances and covariances among these traits.

Typically, estimates of heritability (the ratio of additive genetic variation to total phenotypic variation) are required to establish the degree to which the traits of interest are under genetic control, and hence the scope for changing them by selection (the variation in the trait concerned is also important here). Accurate estimates of heritability require measures of the trait of interest, as well as pedigree information, on many animals. Heritability estimates alone are sufficient to produce simple (univariate) predicted or estimated breeding values (EBVs) which are predictions of the genetic merit of candidates for individual traits of interest. Most modern breeding programmes use more sophisticated statistical techniques (based on "best linear unbiased prediction") to produce multivariate EBVs (a suite of EBVs for traits of interest, that takes into account relationships among animals, and associations among traits). This requires estimates of phenotypic and genetic variances and covariances among all traits (these are also required to derive regressions or correlations usually used to quantify associations among traits). Often multivariate EBVs are weighted and combined in a selection index, producing a single score identifying animals with the highest predicted genetic merit for overall economic performance. This requires estimates of the economic value of all traits that contribute to the overall breeding goal. Figure illustrates the steps involved in prediction of conventional breeding values.

Relationship between temperament traits and other traits

In this section, the relationship between temperament traits, and productivity, health and fitness traits are reviewed. Some studies have investigated the mechanisms underlying these correlations. (See also Supplementary Material Tables A1-A7 for a list of papers and results).

Consequences of selection for production on temperament

It would appear the inclusion of temperament in selection indices for both beef and dairy would have benefits for productivity and also animal welfare although many of the phenotypic associations between temperament and economic traits require further investigation at the genetic level. In beef cattle, calmer animals grow faster and have better feed conversion rates. Meat quality is better in calmer animals, and there may be benefits in terms of health and reproduction. In dairy cattle, milk production and milking speed is higher in calmer animals. Survival is higher in calmer animals, perhaps because farmers are more liable to cull animals that are difficult to milk. The health and fertility benefits are less clear in dairy animals.

In beef cattle, the low genetic correlations between productivity and temperament traits suggest that while selection for efficiency and growth would improve temperament, the correlated response to selection will be low. However, this also implies that current selection goals focussed on productivity alone will result in only a slow improvement in temperament. This may justify placing selection pressure on temperament itself in order to achieve more significant genetic progress in behavior and welfare which may be especially desirable for Bos indicus animals (Sant'Anna et al., 2012). Inclusion of temperament into a selection index would result in a reduction in selection pressure on other economically important productivity traits, and the implications of this would need to be quantified and considered.

Molecular approaches: QTLs and GWAS

Over the last 30 years there has been a great deal of work worldwide to investigate the molecular genetic basis of a wide range of traits of interest in livestock production. This has included studies intended to detect quantitative trait loci (QTL), which are loci explaining a portion of the variation in traits of interest, as well as work to develop increasingly dense genome maps for farm livestock, and studies investigating associations between molecular genetic markers and traits of interest.

QTLs which influence behavioral traits have been found in a number of breeds (Table ). Studies have shown significant or indicative QTL for a number of behavioral traits. Chromosomes 1, 8, 9, 16, and 29 are implicated across studies, although QTLs affecting behavior have been found on other chromosomes as well. Glenske et al. ( 2011) found an association between a candidate gene DRD4 on chromosome 29 and performance in the docility test. DRD4 is a dopamine receptor gene involved in curiosity and novelty seeking in mammals (Rubenstein et al., 1997). A database containing information on behavioral QTLs can be found at www.animalgenome.org/cgi-bin/QTLDB/index.

However, while there are a few traits of interest in livestock that are largely determined by genotype at a single locus or a few loci, there are many more traits of interest that appear to be polygenic in nature, and influenced by many, often hundreds, of loci (Hayes et al., 2009). Moreover, there are often rather few genes that have a large effect on these polygenic traits, and many more that individually have a small effect.

Increasingly dense genome maps are available for livestock with tens or hundreds of thousands of single nucleotide polymorphisms (SNPs) measured throughout the genome. These, coupled with automated platforms for genotyping on so-called SNP "chips," allow genome-wide association studies (GWAS) to be done relating markers to traits of interest, including temperament traits. In beef cattle, a study of temperament and meat quality in Nellore-Angus beef cattle found an association between response to social separation in a pen and a gene regulating sodium ion transport, indicating a difference in nervous system responsiveness (Hulsman Hanna et al., 2014). Additionally, a study in Brown Swiss cattle identified regions with high influence on temperament and aggression on chromosomes 4, 8, and 14 (Kramer et al., 2014). As mathematical techniques are developed that will allow evaluations across breeds and as costs of genotyping fall, more studies that include the assessment of temperament traits are likely.

The availability of dense genome maps and rapid, increasingly affordable genotyping has altered the paradigm for application of molecular genetics in livestock breeding, for many traits of interest. Rather than relying on genotypes at a few loci to predict genetic merit, predictions are increasingly based on information from tens or hundreds of thousands of SNPs throughout the genome. The prediction of genetic merit itself relies on GWAS in a "reference population" of animals-large populations of relevant animals that have both molecular genetic and phenotypic information available. GWAS followed by genomic selection is thought to be a particularly useful approach to improving traits that are difficult, expensive or time-consuming to measure, such as temperament traits. Once the trait has been measured in the reference population, candidates for selection from other similar populations need only be genotyped to predict their genetic merit for temperament (though associations need to be re-estimated periodically). Direct genomic breeding values (dGEBVs) can be predicted from molecular genetic information alone, but increasingly these are combined with EBVs derived from phenotypic records on candidates for selection and their relatives, to enhance accuracy. Figure illustrates the steps involved in prediction of genomic and conventional breeding values.

Both the dairy and beef industries are already using, or moving toward the use of, genomic estimated breeding values (GEBVs). Until recently, much of the genomic research has focussed on productivity traits, meat quality and reproductive traits (see Hayes et al., 2009; and Garrick, 2011 for reviews). This may be because the number of animals with phenotypes required is very large. The need to do the analysis on each breed individually, and the costs of phenotyping and genotyping relative to the perceived benefit of assessing temperament traits are likely to be (at least short term) constraints on the use of this technique. However, many phenotypes can be assessed in each study, allowing temperament to be assessed alongside traits seen to be more economically important.

The use of temperament traits in selection programmes

From the research reviewed above, it would appear that many of the building blocks for selection indexes that include temperament traits exist: the traits can be defined and measured, heritability estimates are available from studies on large numbers of animals in which the traits are carefully measured, and these are similar to heritabilities of many other traits currently under selection. Genetic correlations for a number of temperament traits with productivity measures have been estimated. There is not always consensus across the studies, but some of the larger studies provide strong evidence of favorable genetic correlations.

However, temperament is not often included in breeding indexes. In dairy cattle breeding, EBVs for milking temperament are available as stand-alone EBVs, or information on bulls is available in sub-index scores for "workability" that includes milking speed for some countries. The situation for beef cattle is similar. Although the correlation between handling temperament and growth and meat quality suggest that including temperament in a selection index would be beneficial from a profit and welfare point of view, it is not currently used. Animals may be excluded based on their raw score. Stand-alone EBVs are available (such as flight time for some Bos indicus breeds in Australia and North America, and docility scores for British and European breeds in some countries), but the trait is not currently included in a selection index (Johnston, personal communication).

Constraints and barriers

There are a number of possible technical and producer motivational reasons why temperament traits are not incorporated into selection indexes. A major technical barrier to the use of temperament traits in selection indexes is the need for economic values to allow the trait to be weighted in a selection index. However, it should be possible to derive an economic weight for temperament from the effects that it has on meat quality and growth in beef cattle and the additional labor costs incurred from an animal that is difficult to handle or a cow that is slow or difficult to milk. This has indeed been done for Bos taurus cattle by a team in the US (Busby et al., 2006), but other estimates of economic values are lacking. Another issue is the lack of complete information on genetic and phenotypic correlations of temperament with all the parameters that could be used in selection indexes. For some breeds in some countries, correlations between productivity, meat quality, some fertility traits and temperament have been investigated, but by no means all. New traits are also being incorporated into selection indexes, such as calving ease, and the correlation of this trait with temperament must be determine before both traits can be included. Herd or industry scale and level of organization at the national level may also be important. The pattern of uptake suggests that in regions or countries where a breed is numerous and the breed society or governmental body is well-organized enough to provide support for the recording and evaluation of temperament traits, handling temperament traits may be evaluated. It may be that addition of temperament to an existing selection index has little impact on the overall response, but it should be examined.

Producer motivational factors are also involved. It is clear that temperament is generally poorer in Bos indicus breeds than Bos taurus animals, which may explain the greater motivation to assess temperament in Bos indicus animals. In some Bos taurus breeds, the perception of the breed as being flighty or difficult to handle appears to motivate the breed society to make genetic evaluations on temperament measures such as docility, to improve the trait and improve the popular image of the breed. Additionally, as the response to handling can be modified by repeated handling and habituation to the proximity of humans, the farming system used in any country or region will influence the necessity or motivation of producers to use genetic selection mechanisms to deal with temperament issues. On smaller farms, which are typical of much of Europe, animals arguably experience a higher level of human contact during pasture rotations or seasonal housing than the larger extensive rangeland or feedlot systems more typical of Australia and America. Extreme responses to handling may decline as repeated exposure allows the animal to habituate to human proximity and the handling process. Thus, husbandry conditions may reduce the necessity to use genetic selection to improve temperament. There is also the perception amongst some European producers that some degree of reactivity in animals is desirable, as it promotes survival and competitiveness. Clearly, farm extension and advisory work is needed to inform producers of the negative effects of poor temperament on productivity and profitability.

The future: moving forward and overcoming constraints

As it appears important for welfare and economic reasons to improve handling temperament, we need to facilitate the use of these traits in selection indexes. Research has shown that it is possible to clearly define and accurately record these traits. An increased understanding of the biological basis of these traits will also improve progress. Across the globe, several breed societies and countries have developed EBVs based on farmer-recorded assessments of temperament in dairy cattle and different types of temperament tests in beef cattle. In beef cattle, the lack of a single measure of temperament in beef cattle may impede progress. In the dairy sector in particular, standardization of recording-initially for milk-related traits, and latterly for a much wider set of traits-has helped to underpin improvement in these traits (along with the widespread use of artificial insemination, well-designed breeding programmes usually based on progeny testing, and development of statistical techniques to improve the prediction of genetic merit of animals within and across countries).

Genomic selection may provide an important opportunity for increased use of temperament traits, as increasingly, the prediction of the genetic merit of farm animals will include molecular genetic information. As the cost of genotyping falls, and the predictive power of the information increases, the rate-limiting step to application is likely to be the lack of high-quality records of traits of interest, or phenotypes, both to investigate associations between genotypes and traits of importance in the first place, or to allow ranking of candidates for selection. Temperament traits would therefore need to be measured in comprehensively recorded reference populations, and the correlations between these traits and all others estimated. Within the foreseeable future, GEBVs could also be based on complete DNA sequence information, at least for potentially influential animals.

We should also look to the inclusion of temperament traits other than the response to handling. The correlations between tests measuring response to handling and responses in other contexts can be low. Therefore, selection on the basis of chute test or flight speed may have little impact on traits which are contextually different, such as intra-specific aggressiveness, sociability and maternal defensiveness (e.g., Turner et al., 2013). This review suggests little work has been done on personality traits other than handling temperament, and yet selection for maternal ability and appropriate levels of aggression and sociability and flexibility may be important in terms of animal health and welfare, and also in terms of farm efficiency. Achieving improvements in the latter traits will require the development of automated methods for their measurement. This may come from technologies such as the use of automatic measurement of eye-white (Core et al., 2009), thermal imaging of body areas that show alteration due to stressful events or from other methods, or the use of data collected from activity monitors used to detect oestrus that can be used to characterize personality traits (MacKay et al., 2013). Automatic methods for assessment of meat quality in abattoirs will provide further incentives to improve temperament in beef cattle.

However, even when more efficient methods of phenotyping behavior and other correlated traits are developed, implementation of selection will continue to require that the industry recognizes the need for temperament traits to be used in breeding programs. The case for inclusion seems clearest for handling temperament in beef and dairy, but other traits require more research. Understanding the value of selection for temperament traits will be facilitated by continued effort to clarify and quantify the full range of economic and welfare implications of poor temperament. The primary focus of selection pressure primarily on "output" traits such as carcass weight or milk yield will most likely change in future, as new traits such as RFI are likely to be included in selection goals, particularly in beef, but possibly also in dairy cattle, responding to the need to reduce greenhouse gas emissions from agriculture or the economic necessity for improved feed use efficiency. Animal behavior, particularly behavioral responses to stress, has been hypothesized to be a determinant of feed use efficiency (Richardson and Herd, 2004). In reality, there is a paucity of information on how temperament correlates with feed use efficiency and the information available is contradictory (Petherick et al., 2002; Nkrumah et al., 2007; Elzo et al., 2009; Cafe et al., 2011b; Rolfe et al., 2011). Therefore, it has yet to be established whether improving feed use efficiency will bring with it improvements in temperament, but the low correlations shown in this review suggest that progress will be slow. Changes in some biological systems in response to improvements in feed use efficiency, such as a down-regulation of the rate of endogenous protein turnover, could compromise the animals' ability to respond to stress (Baldwin et al., 1980) leading to changes in behavior and implications for welfare. Given the global interest in improving feed use efficiency in cattle, there is therefore a need to understand the role of temperament as a driver of efficiency and, conversely, how changing feed use efficiency may impact on welfare through other routes.

As handling temperament and other behavioral traits clearly have economic value and animals that respond poorly to handling, and in other situations, suffer negative emotional and physical experiences, resulting in reduced welfare, it is clearly important to improve temperament. Genetic improvement will be important as well as investment in appropriate housing and handling systems. Genetic improvement may become more important against a background of increased herd size, intensification of beef and dairy enterprises and reduced availability of labor. Increased automation and advances in genomic techniques that allow identification of genetically superior animals once the markers have been located in training populations will contribute and quantitative methods will also continue to be important.

Funding

Neither SRUC nor any of the authors received any financial support from a third party in the preparation of this manuscript. SRUC receives funding from the Scottish Government.

Conflict of interest statement

The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

Acknowledgments

SRUC receives funding from RESAS, Scottish Government. We are grateful to Professor Georgios Banos, Dr. Raphael Mrode and Dr. Xavier Boivin for their advice, and to Dr. David Johnston and Paul Williams for information on Breedplan and the Australian Brahman Association.

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# data[].attributes.attachment_link.title
Genetic selection for temperament traits in dairy and beef cattle

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Animal temperament can be defined as a response to environmental or social stimuli. There are a number of temperament traits in cattle that contribute to their welfare, including their response to handling or milking, response to challenge such as human ...

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One of the main behavioral heritability  traits listed is docility. Economically, docility is very important because an animal that is more docile is easier to handle and breed. This also makes the animal easier to sell and more valuable.
https://www.angus.org/Nce/Heritabilities.aspx