Finkemeier & Puppe 2018personality Concepts

REVIEW
published: 28 June 2018

doi: 10.3389/fvets.2018.00131
Personality Research in Mammalian

Farm Animals: Concepts, Measures,
and Relationship to Welfare
Marie-Antonine Finkemeier 1,2 , Jan Langbein 1* and Birger Puppe 1,2*
1
Institute of Behavioural Physiology, Leibniz Institute for Farm Animal Biology (FBN), Dummerstorf, Germany, 2 Behavioural
Sciences, Faculty of Agricultural and Environmental Sciences, University of Rostock, Rostock, Germany
Measuring and understanding personality in animals is a rising scientific field. Much

research has been conducted to assess distinctive individual differences in behavior in
a large number of species in the past few decades, and increasing numbers of studies
include farm animals. Nevertheless, the terminology and definitions used in this broad
scientific field are often confusing because different concepts and methods are used
to explain often synonymously applied terms, such as personality, temperament and
Edited by: coping style. In the present review we give a comprehensive overview of the concepts
Edna Hillmann,
and terms currently used in animal personality research and critically reveal how they are
Humboldt-Universität zu Berlin,
Germany defined and what they measure. First, we shortly introduce concepts describing human
Reviewed by: personality and how these concepts are used to explain animal personality. Second, we
T. Bas Rodenburg, present which concepts, methods and measures are applied in farm animal personality
Wageningen University and Research,
Netherlands
research and show that the terminology used seems to be somehow species-related.
Céline Tallet, Finally, we discuss some findings on the possible impact of personality on the welfare of
INRA Centre Bretagne-Normandie,
farm animals. The assessment of personality in farm animals is of growing scientific and
France
practical interest. Differences in theoretical frameworks and methodological approaches
*Correspondence:
Jan Langbein may also entail the diverse use of the different concepts between basic and applied
langbein@fbn-dummerstorf.de research approaches. We conclude that more consistency is needed in using different
Birger Puppe
puppe@fbn-dummerstorf.de
theoretical concepts, terms and measures, especially in farm animal personality research.
The terms coping style and temperament, which are used in different ways, should not
Specialty section: be examined as independent concepts, but rather should be considered as different
This article was submitted to
Animal Behavior and Welfare,
aspects of the whole personality concept. Farm animal personality should be increasingly
a section of the journal considered for the improvement of animal housing, management, breeding and welfare.
Frontiers in Veterinary Science
Keywords: personality, temperament, coping style, welfare, farm animals
Received: 06 March 2018
Accepted: 29 May 2018
Published: 28 June 2018
INTRODUCTION
Citation:
Finkemeier M-A, Langbein J and Measuring animal personality has become quite important in the past few decades, but many
Puppe B (2018) Personality Research
studies use different concepts and terminology to explain individual differences in behavior and
in Mammalian Farm Animals:
Concepts, Measures, and
physiology. Nevertheless, studies on farm animals do measure personality traits with methods that
Relationship to Welfare. have been used to assess animal behavior. It must be born in mind that the concepts and terms such
Front. Vet. Sci. 5:131. as personality, temperament, coping style used in animal personality research have their origin in
doi: 10.3389/fvets.2018.00131 human personality research.
Frontiers in Veterinary Science | www.frontiersin.org 1 June 2018 | Volume 5 | Article 131

Finkemeier et al. Personality in Mammalian Farm Animals
In human personality research, the term personality is used research. This category classifies behavior and cognitive abilities
to describe a distinctive and relatively stable set of mental that are used by the individual to face the problem (approach)
traits that explain the organism’s behavior, and the concept or to divert the individual’s attention away (avoidance) from the
of personality is explained with the “Five-Factor Model of problem (13).
Personality.” The five factors describing human personality Aim of the present review was to give a comprehensive
are extraversion, agreeableness, conscientiousness, neuroticism, overview how these concepts and terms, which have their
and openness (1). Goldberg (2) describes which traits define origin in human psychology, are currently used in animal and
each factor—extraversion (or surgency) comprises traits like especially farm animal personality research and to critically
talkativeness, assertiveness and activity levels on one end of the reveal how they are defined and what they measure. To
spectrum and silence, passivity and reserve on the other end; obtain an overview of published studies in selected mammalian
agreeableness (or pleasantness) includes traits like kindness, trust farm animal species we carried out a systematic literature
and warmth and the opposite traits of hostility, selfishness and search using the Web of Science (state December 2017)
distrust; conscientiousness (or dependability) comprises traits with keywords including the respective animal species (cattle,
like organization, thoroughness and reliability on one end of the goat, sheep, horse, pig) and different synonyms applied in
spectrum and carelessness, negligence and unreliability on the personality research (temperament, personality, coping). Then
other end; neuroticism (or emotional stability) includes traits like we exemplarily assessed the found studies within the context
nervousness, moodiness and temperamentality; and openness of terms and methodological approaches in general personality
to experience (or intellect) describes traits like imagination, research. Finally, we discuss some findings on the possible impact
curiosity and creativity on one end of the spectrum and of personality on the welfare of farm animals to especially
shallowness and imperceptiveness on the other end. highlight a future direction of personality research.
The term temperament in human psychology research is
closely related to personality. Temperament is defined as the
inherited, early appearing tendencies that continue throughout GENERAL CONCEPTS AND TERMS IN
life and serve as the foundation for personality (3–5) because ANIMAL PERSONALITY RESEARCH
temperament is observable in infants and is tied to basic
psychological processes (6, 7). According to Buss (8), the term Personality
temperament is used to reflect genetic behavioral differences, In animal personality research, many terms are used to explain
while the term personality, which is based on temperament, individual differences in behavior, besides personality itself [e.g.,
seems to be used to reflect non-genetic differences. Rothbart behavioral syndromes (14) and coping style (15, 16)]. Definitions
et al. (7) suggested that understanding temperament is central of the different terms used in animal personality research are
to understanding personality because it influences and is shown in Table 1: Personality is defined as a correlated set of
influenced by the experiences of each individual and because individual behavioral and physiological traits that are consistent
one of its outcomes is the adult personality. However, most over time and contexts (15, 29).
theorists assume that temperament provides the starting place Personality may explain individual differences in dominance
for personality development [reviewed by McCrae et al. (6)]. rank, coping, cognitive abilities and physiology. At least
Temperament provides process-oriented models by establishing one of the five factors which are used to describe human
links between individual differences in behavior and their personality (i.e., extraversion, neuroticism, agreeableness,
psychological and biological substrates in humans as well as in openness, and conscientiousness) has been investigated in many
non-human animals, while personality provides subject-oriented different species. The most commonly measured personality
models (7). factors in animal personality research are exploration, activity,
Coping is a very broad and a multidimensional concept with a aggressiveness, sociability, and boldness (29–31). Exploration
long and complex history. In human psychology research, coping seems to resemble the human factor openness, aggressiveness
strategies refer to intentional cognitive or behavioral attempts seems to resemble agreeableness, and sociability seems to
by the individual to manage a stressor (9). Lazarus (10) has resemble extraversion, while boldness and activity seem to be
emphasized coping as a key concept for research on adaptation combined in the human factor neuroticism [reviewed by Gosling
and health. Because coping is a behavioral reaction to aversive and John (32)]. According to Gosling and John (32), the human
situations that induce several physiological stress reactions (11), factor conscientiousness can only be found in chimpanzees
individual ways of dealing with stress have an enormous impact and gorillas. Figure 1 shows the five personality factors in
on human health (12). Lazarus (10) suggests to differentiating humans and how they describe the personality type of two
between two approaches to coping, one that treats coping as a imaginary individuals [adapted from (33)] and the equivalent
personality characteristic (style) and another that refers to the personality factors found in animals. While individual A scores
efforts to manage stress (process). Whereas the process approach highly in exploration (openness), sociability (extraversion) and
primarily addresses the behavioral and physiological mechanisms aggressiveness (agreeableness), individual B scores highly on
involved in the adaptational response to stress, coping style boldness and activity (neuroticism) and, if this individual is a
rather seems to describe a personal or individual consistency in human, in conscientiousness (Figure 1). A discussed factor in
this response. The classification of coping into an “approach- animals is the dominance factor that seems to be defined by
avoidance” category can also be found in animal personality boldness, physical aggression and low levels of fearfulness (32).

TABLE 1 | Different personality-related terms in alphabetical order used to

describe individual differences in animal behavior and their definitions.
Personality- Definition References

related
term
Behavioral Behavioral response to handling (17)

response
Behavioral A suite of correlated behaviors (14)
syndrome reflecting an individual’s consistency
in behavior across multiple situations;
a population or species can exhibit a
behavioral syndrome; within the
syndrome, individuals have a
behavioral type
Coping personality Coping strategies that may reflect (18)
type different personality types
Coping style Based on the animal’s reaction to its (11, 15, 16, FIGURE 1 | The five personality factors in humans and how they describe the
environment with respect to reducing 19) personality type of two imaginary individuals [adapted from (33)]. The
effects of aversive stimuli: equivalent personality factors described in animals are written in bold and
- fight or flight response italics. Scores are given from zero to 100 in percentages. For example,
- approach or avoidance Individual A (black dashed lines and points) scored high in openness,
- boldness or shyness extraversion and agreeableness, while Individual B (gray dashed lines and
Emotional Social reactivity (i.e., active vs. (20) points) scored high on neuroticism and conscientiousness.
reactivity passive strategy); exploratory activity;
reactivity to humans
Identity profile Describes individuality, personality (21)
and their relationship with certain influenced by the social organization. In hyenas, the dominance
morphological traits of the animals; rank is transmitted through a matrilineal system, and females
four groups of similar animals:
are larger than males. Sex differences in personality seem to
aggressive, affiliative, passive,
avoiders be related to the ecological niches occupied by the two sexes
Individual Individual variation; intra-animal (22–25) [reviewed by Gosling and John (32)].
differences in repeatability; the relationships David and Dall (37) state in their review that most of the
behavior between different test situations and confusion about terminology in animal personality studies is due
the frequency distributions of various to two main concepts, which can also be used complementarily:
measures of behavior; consistency of
individual variability
the intra-individual variability and the life-history approach. The
first concept tries to explain the occurrence of between-individual
Temperament Inherited, early appearing tendencies (3–6)
that continue throughout life and differences in the consistency of any behavior (34), which means
serve as the foundation for that studies employing this concept focus on the consistency and
personality; observable in infants and flexibility of behavioral expression. The second concept focuses
animals and tied to basic only on boldness-like behaviors, which means that studies using
psychological processes
this concept concentrate on the individual average behavior and
Personality A correlated set of individual (15, 26–28)
behavioral and physiological traits that
the between-individual relationships between life history and
are consistent over time and contexts behavior (37). One approach using both concepts is the pace-of-
life syndrome (POLS) [reviewed by Réale et al. (26)]. This concept
includes the idea that between-individual differences in behavior
are associated with individual variations in life-history traits [e.g.,
In humans, these traits seem to be related to extraversion because habitat use, predation avoidance, dispersal and/or social behavior
humans participate in multiple dominance hierarchies that are (14, 38–40)] because both may contribute to evolutionary trade-
less clearly defined and involve widely divergent skills (32). This offs and may have co-evolved. And physiological traits that differ
means that species not only differ in the type of relevant factors, between individuals, such as growth, metabolic rate, reproductive
but also in the number of such factors (34): While chimpanzees value and hormone levels have also become important for animal
share a common six-factor structure, including dominance, personality research (28, 40–42) because they can be influenced
extraversion, agreeableness, conscientiousness, neuroticism, by genetic predispositions and early environmental effects (27,
and openness (35), mountain gorillas seem to have only four 43).
personality factors: dominance, openness, sociability and Another important aspect of personality research in animals
proto-agreeableness (36). As in human personality research, sex is that despite the consistency of some factors, others seem to be
differences can be found in the expression of animal personality plastic and help the organism adapt to changing environments.
traits. For example, hyena (Crocuta crocuta) males are more This aspect is termed behavioral and/or physiological plasticity
fearful and nervous than females. This effect seems to be (43). Responsiveness to environmental conditions may span from

short-term (phenotypic flexibility) to long-term effects (44), and factor boldness is related to other investigated behavioral traits.
the early environment an individual experiences contributes to Boldness was found to be related to reactivity to humans in horses
between-individual differences in life histories and personality (77), cattle (50), and sheep (69): Individuals that interacted more
factors (41, 45). For example, in coral reef fish, individual scores with a novel stimulus were more interested in novel humans as
on activity, boldness and aggressiveness increased from 2.5- well. In sheep, bold individuals proved to be more explorative and
to 6-fold when induced by an increase in water temperature, the distance between individuals while grazing was greater than
indicating individual coping abilities and/or behavioral plasticity in shy individuals (69). Moreover, the personality factor activity
(46). Moreover, photoperiodic changes influenced fearlessness, proved to be related to boldness and reactivity to humans in
the stress response, the timing of maturation and resting sheep: More active sheep seem to be bolder und more reactive
metabolic rate (RMR) in wild cavies (Cavia aperea) and showed to humans than shy sheep (65, 101). Calves can be categorized
differences between sexes (47, 48). Behavioral and physiological in four different personality types based on a combination of
plasticity is an important factor for the survival of an organism. behavioral and heart rate variability (HRV) data (50).
Even if most of the personality factors proved to be consistent
over time and contexts, factors like RMR and fearlessness were Temperament
not consistent in cavies, whereas exploration and boldness In animal personality research, the term temperament is closely
were consistent regardless of the photoperiodic treatment (47). related to personality. Temperament often matches the definition
One might argue that factors such as RMR and fearlessness found in human personality research and is defined as inherited,
should not count as personality factors. However, an example early-appearing tendencies that continue throughout life and
in the European mink (Mustela lutreola) showed that traits that serve as the foundation for personality (6) (Table 1). Since
that describe factors like boldness and exploration can change the distinction of both terms by definition is often vague,
between seasons but that these changes remain consistent over some animal researchers simply treat them as synonyms (29),
time (49), which matches the definition of a personality factor. while others argue that the term temperament is used simply
These examples indicate that the plasticity and consistency to avoid using the term personality with regard to animals,
of personality factors are not exact opposites and indicate which might be associated with anthropomorphism by some
interesting asymmetries in the adjustments of personality factors ethologists (102). While temperament is assumed to be based
that seem somehow species-related. on early, stable predispositions in the emotional and behavioral
In farm animal studies, the term personality is scarcely used, responses of an individual like boldness, aggressiveness and
even if the methods employed (e.g., repeated measurements) pleasantness, personality is often reserved for patterns of
and measured variables (e.g., curiosity) can be described as reflection and reaction to environmental circumstances acquired
personality factors. In those studies, the term temperament during a lifetime in organisms with sophisticated cognitive
is used as an equivalent to the term personality. This could capacities. The behavioral factors that are the focus of
be due to the fact that farm animals are used to get meat, research on animal temperament in many different species are
milk and other products and that it is easier to consume farm reactivity, fearfulness, sociability, responsiveness, and aggression.
animal products when scientists do not attribute human-like Temperament in non-human animals is often described on a
characteristics to explain individual differences in farm animal one-dimensional scale using expressions such as “proactive vs.
behavior. Table 2 lists selected studies assessing personality reactive,” “aggressive vs. non-aggressive,” “bold vs. shy,” etc.
factors using different experimental designs in mammalian farm (103, 104). The actual theoretical frameworks on temperament
animals (Table 2A: cattle; Table 2B: goats; Table 2C: sheep; in non-human animals argue for scoring reactions in more
Table 2D: horses; Table 2E: pigs) and the behavioral tests used. than one dimension to reflect the entire nature of temperament
Some of these studies only measured and/or analyzed a small (29, 105). A multivariate approach has been used by Meager
aspect of personality, measured just one or two personality et al. (106) to investigate whether distinct temperaments are
traits or are examples of the very beginning of farm animal present in Atlantic cod (Gadus morhua). Their results refer to
personality research, while other studies, shown in this table, the multidimensionality of animal temperament and provide
show consistency and/or cross-context correlations between a clear indication that two distinct behavioral phenotypes are
traits. One can see that the use of a certain personality-related evident in fish. A study in non-human primates has shown
term is dependent on the species studied. In cattle, the term that temperament, based on a factor analysis including behavior
temperament is mostly used to explain individual differences in different dimensions in yearling rhesus monkeys (Macaca
in behavior, while in pigs, the term coping style is mainly mulatta), seems to be an important factor in an individual’s ability
used (Tables 2A,E). In horses and more recent farm animal to select friends and is related to later variations in the animals’
personality studies, one can find the term personality as well social networks (107). The subjects preferred peers that had
[e.g., (18, 76)] (Table 2D). In most of the farm animal studies similar temperament scores to their own even after accounting
investigating temperament, different personality factors can be for sex, rank and kinship. This example shows that the definition
found [e.g., curiosity shown in the general response to unfamiliar of temperament from the human domain can be used for non-
humans or the exit from a restraint device (50, 100)]. These can human animals as well. Foyer et al. (108) investigated the impact
be considered as personality factors when they are measured in at of the level of maternal care in dogs on the offspring’s behavior
least two different time periods of an animals’ life. Like in other at approximately 18 months of age. The authors conducted a
taxa, some farm animal studies can show that the personality set of temperament tests to score the behavior of the dogs in

TABLE 2 | Selected studies assessing personality factors using different experimental designs in mammalian farm animals [2 (A): cattle; 2 (B): goats; 2 (C): sheep; 2 (D):
horses; 2 (E): pigs].
Personality-related Test Measures Study

term used
(A): CATTLE (Bos taurus)

Behavioral response Docility test Aggressiveness against the handler, running time and number of escapes (17)
per minute of test period in presence or absence of the handler
Individual differences in Human approach test Behavioral response when a human approaches the cow (23, 24)
behavior Lateralization test The side the animal uses to avoid an obstacle
Novel human test Behavioral response toward an unknown human
Novel object test Behavioral response toward an unknown object
Open field test Activity and exploration behavior in an unknown arena
Surprise test Latency to start eating again after encountering a blast of air
Temperament score Chute score, velocity exit, pen score, flight speed
Temperament Combined social isolation Activity and exploration behavior in an unknown arena and activity during (50–61)
and open field test social isolation
Docility test Aggressiveness against the handler, running time and number of escapes
per minute of test period in presence or absence of the handler
Handling test Touching the cow from head to tail
Human approach test Behavioral response when a human approaches the cow
Social isolation test Activity and exploration behavior during social isolation
Temperament score Chute score, velocity exit, pen score, flight speed
(B): GOATS (Capra hircus)
Identity profiles Behavioral observation Direct observation during milking and/or over a certain period of time (21)
Temperament Behavioral observation Direct observation during milking and/or over a certain period of time (62, 63)
Personality Novel object test Behavioral response toward an unknown object (64)
Social isolation test Call, activity, and exploration behavior during social isolation
(C): SHEEP (Ovis aries)
Emotional reactivity Human approach test Behavioral response when a human approaches the sheep (20)
Temperament Novel object test Behavioral response toward an unknown object (65–68)
Personality Open field test Activity and exploration behavior in an unknown arena (69)
(D): HORSES (Equus caballus)
Coping type Behavior after reintroduction Expression of submissive or dominant behavior toward conspecifics (70)
in a group
Owner ratings Questionnaire about the expression of certain behaviors
Water spray test Scoring of the horse’s reaction to a spray of water
Temperament Behavior after reintroduction Expression of submissive or dominant behavior toward conspecifics (70–75)
in a group
Behavioral observation Expression of behaviors in the home pen
Handling test Latency to allow the human to touch the leg about three times
Horse personality Questionnaire about 43 behaviorally defined adjectives
questionnaire (HPQ)
Surprise test Latency to start eating again after opening an umbrella
Water spray test Scoring of the horse’s reaction to a spray of water
(Continued)

TABLE 2 | Continued
Personality-related Test Measures Study

term used
Personality Handling test Latency to allow the human to touch the leg about three times of the horse (76–87)
Horse personality Questionnaire about 43 behaviorally defined adjectives
questionnaire (HPQ)
(E): PIGS (Sus scrofa)
Coping Personality Backtest number of escape attempts, latency to first escape attempt (18)
Type
Coping Type Backtest Number of escape attempts, latency to first escape attempt (88–96)
Behavioral observation Expression of behaviors in the home pen
Combined open field and Activity and exploration behavior in an unknown arena with an unknown
novel object test object
Human approach test Behavioral response when a human approaches the pig
Resident-intruder test Latency to attack a conspecific, number and duration of fights, lesion score
Social support test Socio-positive behavior toward a conspecific
Individual differences in Backtest Number of escape attempts, latency to first escape attempt (25, 97)
behavior Combined open field and Activity and exploration behavior in an unknown arena with an unknown
novel object test object
Human approach test Behavioral response when a human approaches the pig
Open door test Latency to leave the home pen
Temperament Human approach test Behavioral response when a human approaches the pig (98)
Open door test Latency to leave the home pen
Personality Backtest Number of escape attempts, latency to first escape attempt (99)
The column “Personality-related term used” refers to the term the studies used to explain their results. The column “Test” refers to the behavioral tests used, while the column “Measures”
refers to the measured variables.
different factors, such as sociability and fearfulness. The study included in the breeding objectives of some countries [reviewed
showed that the level of maternal care differed consistently and by Gibbons (111)]. Furthermore, high reactive bulls (reactivity
that it has a significant impact on the adult temperament of was measured with a temperament score) had a greater rectal
the offspring, mainly in terms of sociability and dominance. temperature and a higher cortisol and epinephrine concentration
The responses of cats across different behavioral factors were prior to and after transportation than low reactive bulls (53).
assessed to calculate a feline temperament profile (109). Later, As these examples measure personality, because measured in
the reactions of the cats in a 3-min stress test were measured. adult animals, these examples show how complex it is to describe
The authors did not find correlations between temperament individual personality in animals and that temperament reflects a
scores and measures of behavioral and adrenocortical responses sub-aspect of the whole concept of animal personality.
in the stress test. However, many studies concerning individual
differences in animal behavior do not make the distinction Coping Style
between temperament and personality consistently, which makes The concept of coping styles has also been used in the past few
it more difficult to understand the results with regard to decades to better understand animal personality and is based
temperament and/or personality. on the animal’s reaction to its environment with respect to
In many farm animal studies, the term temperament is used reducing the effect of aversive stimuli (16, 19) (Table 1). Like
as an equivalent to the term personality, which also can be in humans, animals show individual differences in coping with
seen in Table 2. General responses to unfamiliar humans or different environmental changes, confirming that personality
situations are quite important for handling, management and and coping style are closely linked [(11, 15, 16, 19, 112);
selective breeding (65). And traits such as fearfulness, happiness, but see Zidar et al. (113) for an example of a missing
alertness and docility [(51, 52, 110) can be considered to describe relationship in the red junglefowl]. According to the boldness-
temperament when measured during the juvenile period of shyness continuum, animals can be categorized into three sub-
an animal and/or once in an animal’s life. Animals that are groups based on their risk-taking behavior: bold, intermediate
more likely to cope with handling procedures are considered and shy (112) or proactive, intermediate and reactive based
to have a “good” temperament (51) and dairy temperament on the animal’s reaction to its environment with regard to
(generally defined as the animal’s response to milking) has been altering the effect of aversive stimuli (fight/flight response) (11,

15, 16, 19). Proactive individuals are considered to be more are relatively independent from one another, which means that
aggressive toward conspecifics, show more dominant behavior, they must not co-occur [reviewed by Uher (118)]. Each factor
and are considered more explorative, bold and active. Moreover, summarizes the shared variation of diverse inter-correlating
they respond with a strong sympathetic activation and an traits. The rank-orders of the same individuals on the same
increase in noradrenergic stimulation when confronted with factor can vary considerably across different situations. An
a challenging situation, while reactive animals are considered individual can have different scores on each factor [reviewed by
the opposite behavioral phenotype, with behavior that is more Uher (118)]. The individual-oriented perspective (like in humans
submissive, less explorative and less active and responding with and shown in Figure 1) can also be applied to the analyses
a strong hypothalamic-pituitary-adrenocortical reactivity toward of multiple individuals such that individuals sharing a similar
challenging situations (31, 114–117). profile can be grouped statistically into configurational types.
Figure 2 shows a possible multidimensional approach to Such groups represent distinct and discontinuous categories of
describing the five basic factors in animal personality research prototypes (e.g., proactive, intermediate and reactive) [reviewed
applied to the concept of animal coping style using an example by Uher (118)]. According to Koolhaas (119), proactive
of two imaginary animals. In addition, there is a potential sixth individuals perform better under highly predictable conditions,
factor, dominance. There is an ongoing discussion as to whether while reactive individuals perform better under variable and
dominance is a factor by itself or just a social outcome, and it also unpredictable environmental conditions. Proactive individuals
represents other species-related factors that are often missed in also grow faster and have a higher RMR, which they can afford
measurements (34, 102). Considering the description of proactive due to their higher rates of food acquisition. When food is
and reactive individuals above, the animal that scores low on the abundant, proactive individuals may perform better than reactive
aggressiveness, activity and sociability axes would be described individuals [reviewed by Careau et al. (120)]. Many studies have
as being more peaceful, inactive and highly social and would shown that different coping styles correlate with dominance rank,
represent a reactive coping style. The animal that scores high cognitive abilities and physiological measures [e.g., immunology
on the boldness, exploration and aggressiveness axes would be (119)]. In fish, bold female guppies (Poecilia reticulate) learned
described as being more aggressive, explorative and bold and a task more quickly than shy females (121), and in zebra finches
would be considered as a proactive individual. Intermediate (Taeniopygia guttata), more active birds were faster at solving a
individuals would express behavior described by a mixture of task, while less active individuals needed longer or did not solve
the scores on the factors resulting in individual differences on a the task at all, which indicates that higher activity may lead to
continuum. Some factors show only low inter-correlations and routine-forming behavior instead of being attentive to external
cues (122). Proactive Senegalese sole (Solea senegalensis) juveniles
exhibited a shorter feeding latency, a higher duration of escape
attempts and a lower cortisol level than reactive individuals (123).
An example in mammals revealed that different coping styles in
wild alpine marmots (Marmota marmota) are accompanied by
different baseline and stress-induced plasma oxidative statuses
(28). In male rainbowfish (Melanotaenia duboulayi), personality
factors like aggression, activity and boldness covaried with
the male’s position in the hierarchy, directly influencing their
reproductive success (124). Moreover, an example of spiders
living in colonies showed that individuals with different coping
styles do different tasks. While aggressive spiders perform prey
capture and colony defense, less aggressive spiders perform brood
care (22).
Farm animal research on coping styles became more and
more important because the coping style covers the aspect
of an animal’s skill in coping with environmental challenges.
Farm animals like pigs (Sus scrofa), cattle (Bos taurus), and
horses (Equus caballus) have to cope with numerous challenging
situations (e.g., regrouping, housing, management) and technical
FIGURE 2 | The five basic factors in animal personality research (bold lines) material during their lives. Therefore, investigating their coping
that can also be used to describe coping style. The dominance factor is strategies and cognitive abilities appears to be a very important
presented in dashed lines because there are insufficient data to consider it as research topic, because like temperament, it reflects sub-aspects
a fully accepted sixth factor in animal personality. The dots represent two
imaginary animals with different coping styles. The animal behaving as
of the whole concept of animal personality. Aggression, boldness
described by the black-gray dots would represent a reactive coper, while the and exploration seem to be related to results in backtests that
animal behaving as described by the black dots would be considered as a determine the specific coping-type in pigs assessed by the early
proactive individual. Intermediate individuals would express behavior described escape behavior of piglets: Proactive pigs proved to be more
by a mixture of scores on the factors resulting in individual differences along a
aggressive, explorative and bolder than reactive pigs (88, 97).
continuum.
Recently, a number of studies were conducted to investigate

the impact of different coping styles on psychophysiological factors for each species. Different species have different
measures and molecular correlates later in life. Reactive cows environmental requirements because they have different habitats
can be distinguished by their baseline HRV (54) and pigs and different life-histories. Koski (34) indicated that most studies
characterized as belonging to different coping styles differed about personality focus on boldness, aggressiveness, activity,
in their general autonomic reaction (e.g., HR, HRV) and in exploration and sociability as the most prominent personality
their affective appraisal in relation to common husbandry-related factors. However, she writes that the focus on these personality
situations like feeding or human handling (125), indicating factors derived from human personality research may ignore
personality-depended emotional assessment of environmental the possibility of other factors that are more important for
situations. There is also evidence that individual coping styles the investigated species and that such a focus limits our
of pigs are reflected in their immune responses indicating understanding of the full repertoire of personality factors [also
that proactive pigs might favor molecular pathways enabling reviewed by Gosling and John (32)]. Therefore, it is possible that
a more effective strategy for defense and recovery than in species not only differ in the type of relevant factors, but also in
reactive pigs (89, 90). Moreover, a recently conducted genome- the number of such factors (34): A study done on 1223 horses
wide association study revealed that hypothalamic genes known of eight different breeds with a Horse Personality Questionnaire
to be involved in pathways regarding the immune system, also identified approximately six personality factors (78). Carter
telomere function signaling, neurotransmitter receptors as well et al. (131) found boldness to be one of the most commonly
as circadian rhythms were differently expressed in pigs with measured personality factors in animal personality research,
different coping styles (126). and it has been interpreted as the propensity to take risks,
especially in novel situations, and as an individual’s response
to a risky situation faced alone. Boldness is often tested by
MEASURES AND METHODS IN FARM quantifying behavioral responses to novel objects, responses
ANIMAL PERSONALITY RESEARCH to a novel environment, and/or responses to predation risk.
However, these three types of tests are not necessarily comparable
Personality, temperament and/or coping style and how they and demonstrate a lack of standardization for quantifying this
are interrelated with a diversity of other external or internal behavior. For example, measuring boldness in cavies is slightly
stimuli are investigated using different experimental setups. The different than measuring boldness in pigs and cattle. For the
main problem is to know exactly which personality factor or three species, one can use the novel object test. While in cavies,
variable is being investigated using a certain experimental design the novel object test is conducted in the home enclosure (30,
(e.g., tests of exploration in an open field, test of boldness 47, 48), with pigs and cattle, the test is normally conducted in
in investigating a novel object). Murphy et al. (127) identify an arena outside the home pen after a habituation period or
in their review the most important aspects of experiments after an open field test [reviewed by Forkman et al. (132)]. Tests
used to investigate farm animal personality. A test should be like the novel human test or the open-door test are conducted
ecologically valid and the animal should be able to display its with farm animal species and measure the reactivity of an
natural emotion-related behaviors. Since emotions have recently individual toward unknown persons or places (55, 71, 98). This
been discussed to be a personality factor by itself [see below type of test is only feasible with domesticated species that are
reviewed by Koolhaas et al. (128)], tests should be sensitive used to human manipulation such as farm animals. In most
enough to capture slight differences in levels of emotional cases, the measured trait is suggested to mirror fearfulness when
arousal (bodily activation or excitation, the first dimension of describing an animal having a higher latency in approaching
emotion) or valence (negative or positive, the second dimension an unknown person [reviewed by Murphy et al. (127)]. In
of emotion) (129, 130), which is important for longitudinal rodents, tests like the elevated plus maze [e.g., in mice and rats
studies. Furthermore, the test itself and the variables measured (133, 134)] or leaving a plastic shelter in an open field [e.g.,
should be standardized to allow for comparisons between studies in cavies (47)] are used to measure fearfulness or its opposite,
(127). In his review, Gosling (3) describes the variables that can fearlessness. Exploration is measured in most mammal studies
be used to analyse personality factors: (i) reactivity, emotionality, and in most farm animal species with an open field test [reviewed
fearfulness: e.g., measurement of the defecation rate in an open by Forkman et al. (132), but see recent critiques by Perals et al.
field; (ii) exploration/boldness: e.g., interactions with a novel (135)].
object; (iii) sociability: e.g., frequency of social encounters; (iv) To measure different coping strategies in pigs, Hessing et al.
aggression: e.g., latency to attack another individual; (v) activity: (136) suggested conducting the backtest at an early age to
e.g., amount of enclosures (e.g., open field or arena test) covered; measure defensive reactions like struggling while the piglet
(vi) dominance/assertiveness: e.g., the individual’s rank in the is turned over on its back. Zebunke et al. (91) used more
dominance hierarchy. Of course, these variables can be used than 3,000 individuals and repeated the test four times with
to analyse other personality factors as well, and the mentioned each individual. The results showed a moderate consistency
personality factors can be measured by other variables or even a of behavioral reactions across repeated testing, which cannot
mixture thereof. be attributed to a randomly occurring pattern. The authors
The most important aspect is to find the most appropriate concluded that the backtest does not provide phenotypic
test with the most appropriate variables to record the evidence for definitive coping styles that are clearly separable.
behaviors that are desired to analyse the intended personality Instead, they found pronounced individual dispositions along

a continuum from proactive to reactive coping behaviors. farm animals is a possible means of measuring states of welfare
The results of an equivalent to the backtest conducted with and can also help to increase welfare. Individuals that are less well
juvenile cavies before weaning (the struggle test) show a adapted to their environment may have reduced welfare, which
similar pattern: the time spent struggling was not significantly in turn can lead to reduced productivity (145, 146). Figure 4
repeatable but showed a trend (137). Therefore, the backtest represents the influence of personality on individual welfare.
may indicate a certain coping style in pigs, which is discussed Personality directly influences behavior and physiology and
to be a personality factor by itself (113, 128). The backtest therefore influences individual welfare, while as in a feedback-
should not be used as a reference on its own but rather loop, welfare can directly influence behavior and physiology.
should be used as an addition to other tests measuring Behaviors can influence physiology and vice versa in a sort of
behavior and a certain personality type. Moreover, these positive feedback system as well. Especially in farm animals,
examples confirm that one test can simultaneously be influenced domestication has an impact on behavior and physiology and
by and therefore measure two or more personality factors directly influences breeding. During domestication of most farm
(29). animal species, behavior changed to lower levels of aggression
Besides the fear of anthropomorphism in using the term and activity (147). Artificial selection for the improvement
personality, to know what is measured is one of the biggest of production traits may have resulted in the selection of
lacking points in the field of farm animal personality research (see animals that would count as reactive copers (147). Genetic
Table 2). Inspired by the definitions in Table 1, Figure 3 shows studies of captive animals often rely on the selection of specific
how researchers could proceed to know which personality related production traits (148–150), because specific production traits
term they should use for their interpretation. Temperament are emphasized in the individual pedigree of an animal used
and coping style overlap with personality, because they are for breeding [(151, 152); reviewed by Laine and van Oers
sub-aspects of the whole personality concept. With all the (153)]. Recently, personality traits have been considered to a
behavioral tests used in farm animal personality research (e.g., greater extent in the calculation of breeding indices because
open field test, novel object test, open door test, backtest etc.) some of these traits show moderate to high heritability (150,
applied ethologist can measure different aspects of personality: 154–156). Aggression is highly heritable in pigs [ranging from
It depends of the approach they use. For instance, (i) using h2 = 0.32 to h2 = 0.48 (157–159)]. In pigs as well as in
a behavioral test to measure a response to an aversive or cattle, aggressive behavior toward stockpersons and group-
stressful stimulus or situation (e.g., approach or avoidance) members is related to increased maternal behavior, which can
means that primarily a specific coping style is assessed, be problematic [reviewed by Haskell et al. (155, 158)]. In beef
(ii) conducting various behavioral tests in a combination and dairy cattle, handling shows moderate to high heritability
and/or conducted at least twice during lifetime, measures scores: e.g., chute score (h2 = 0.24), flight speed (h2 = 0.36),
cross-context correlations and/or consistency and therefor and docility (h2 = 0.26) [reviewed by Haskell et al. (56, 155,
assesses personality, (iii) conducting behavioral tests during 160)]. Especially in dairy cattle, milking temperament shows a
the juvenile stage and only once during lifetime, would assess moderate heritability (e.g., on average h2 = 0.19), but is also
temperament. related to production traits such as milk yield [reviewed by
Haskell et al. (56, 155)]. In France the docility test has been
used to select for improved temperament in Limousin cattle since
PERSONALITY AND WELFARE IN FARM 1992, and dairy temperament (generally defined as the animal’s
ANIMALS response to milking) and milking speed have been included in the
breeding objectives of some countries [e.g., United Kingdom and
In human psychology, studies show that subjective well-being Norway reviewed by Gibbons (111)]. Horses, especially stallions
is associated with health (138) and is linked to personality considered for breeding purposes, are judged for performance
[reviewed by Weiss et al. (139)]. Well-being correlates with the (e.g., the gaits under the rider, jumping ability, rideability,
five-factor model of personality, especially with neuroticism, fitness, health, stamina) and personality (e.g., behavior during
extraversion and conscientiousness (140). In zoo animals, studies handling (labeled as character), attention and reactivity (labeled
indicate that zoo keepers are able to reliably rate animal as temperament) and braveness, willingness and ability to learn
personality traits and that those ratings are implemented into [labeled as willingness to work (72)].
zoo management practices to improve the welfare of captive Another approach to explain the relationship between
animals (141). Personality research in farm animals has become personality and welfare has been reviewed by Koolhaas and
important as well because welfare not only comprises the actual van Reenen (128), and it describes a three-dimensional model
health status of an animal but also is affected by individual with coping style, emotionality and sociability as independent
differences in behavior and physiology (142). Dawkins (143) factors. These factors are defined as being stable over time
already stated that individual behavior has a big advantage in and across contexts within the individual. Emotionality seems
welfare studies because it might become an “early warning to be one important aspect to increasing welfare because it
system” of trouble yet to come. Changes in behavior (e.g., makes the distinction between fearful animals that are highly
aggressiveness toward the caretaker) can be a hint of pain or other emotionally aroused by a challenging situation and non-fearful
problems (144). Therefore, investigating and understanding animals that do not perceive the same situation as stressful or
individual differences in behavior, respective to personality, in alarming (128). A highly emotionally aroused animal would

FIGURE 3 | Various behavioral tests that can be used to assess different aspects of personality. It becomes evident that coping style and temperament are
sub-aspects of the concept of personality.
Some studies have shown evidence that emotionality, and

therefore personality, seem to have an impact on production
traits, such as meat quality and milk production. Studies on steers
show that individual differences in stress responses (e.g., increase
of cortisol in a stressful situation) or in blood lactate levels have
an impact on meat tenderness (57, 58). In dairy cows, personality
has an impact on behavioral and physiological responses to
milking and on the stress associated with being milked in a
novel environment (50, 161). A study on horses shows that
personality has an influence on pain expression. Horses that were
highly affected by vertebral problems showed more aggressive
behavior toward humans than horses with no vertebral problems
(144). Lameness was more expressed by highly extraverted
horses even if the severity level of the injury threshold was
lower when compared to more neurotic individuals (76). A
study on female sheep (Ovis aries) showed that animals typed
as “nervous” [measurement of the behavioral reactivity to the
psychosocial stress of social isolation and selected for “calm”
or “nervous” temperament for 17 generations (66)] seem to
show a decrease in agitation score, the frequency of vocalizations
and the plasma concentrations of cortisol when treated with
FIGURE 4 | Relationship between personality and welfare. Personality directly lavender oil as an alternative treatment to alleviate anxiety
influences behavior and physiology and therefore influences individual welfare, compared to the response of “calm” sheep (66). Moreover,
while as in a feedback-loop, welfare can directly influence behavior and these “nervous” sheep produced a lower volume of higher
physiology. Behavior can influence physiology and vice versa in a sort of
viscosity colostrum than “calm” sheep, and this disparity could
positive feedback system. Especially in farm animals, domestication has an
impact on behavior and physiology and directly influences breeding.
be corrected by nutritional supplementation (with barley), which
only had an effect on “nervous” sheep (162). These examples
indicate that personality seems to be an important factor in the
therefore exhibit activation of neuroendocrine systems while efficacy of certain substances and nutritional supplementations.
a non-fearful animal do not show any enhanced biological Personalized medicine in animals and humans already indicates
responses (128). that personality is a strong indicator for pathology development,

medical treatment and substance efficacy (163). Understanding terms for consistent behavioral responses in animals, but as
the individual personalities of farm animals is not only important different aspects of the whole personality concept. Research on
for their welfare but also has an impact on economic factors for personality of farm animals is currently far from covering all
farmers [reviewed by Clark et al. (164)]. Therefore, it is important possible aspects, but focuses in particular on the phenotyping
to think about considering to draw an individual personality of personality traits and potential relationships with cognition,
profile like in Figure 1 [adapted from Costa and McCrae (33)] to emotion and welfare. We conclude that the assessment of
picture farm animal personality and in concordance to improve personality in farm animals is of growing scientific, practical
management, handling, breeding, medical treatment and the and economic interest, because it has an obvious verifiable
design of housing systems that allow the animal to perform impact on the individual behavioral reaction to different housing
effective coping behavior [reviewed by Wechsler (11)]. systems, management practices and veterinary interventions
and is therefore important for the improvement of animal
CONCLUSION welfare.
This review gives an impression of how diverse farm animal AUTHOR CONTRIBUTIONS
personality research is and which aspects have to be considered
in investigating personality in different farm animal species. M-AF, JL, and BP contributed to the conception and structure of
Terminology in farm animal personality research is somewhat the manuscript. Main parts of the manuscript were written by M-
confusing and in some cases difficult to compare because AF. The final version of the manuscript was prepared and edited
the terms used seem somehow to be species-related. While by M-AF, JL, and BP.
studies in different mammals, birds, and other taxa widely
use the term personality to describe between-individual ACKNOWLEDGMENTS
consistency in behavioral variation, in farm animal research
the terms temperament and coping style are predominantly We thank Christian Nawroth, Katrin Siebert, Evelin Normann
used, probably because personality might be associated with and the reviewers for valuable comments and corrections on
anthropomorphism. The broad field of personality research earlier versions of the manuscript. The publication of this article
generally needs more consistency in using theoretical concepts, was funded by the Open Access Fund of the Leibniz Association
terms and measures and we recommend that the terms and the Open Access Fund of the Leibniz Institute for Farm
should neither be regarded as synonyms nor as independent Animal Biology (FBN).
REFERENCES 11. Wechsler B. Coping and coping strategies: a behavioural view. Appl Anim
Behav Sci. (1995) 43:123–34. doi: 10.1016/0168-1591(95)00557-9
1. John OP, Srivastava S. The Big Five trait taxonomy: history, measurement 12. Henry JP, Stephens PM. Stress, Health, and the Social Environment. New
and theoretical perspectives. In: Lawrence AP, Oliver PJ, editors. York, NY: Springer (1977)
Handbook of Personality: Theory and Research. New York: Guilford (1999). 13. Moos RH. Coping Response Inventory Youth Form: Professional Manual.
p. 102–38. Odessa, FL: Psychological Assessment Resources, Inc. (1993).
2. Goldberg LR. The structure of phenotypic personality traits. Am Psychol. 14. Sih A, Bell A, Chadwick Johnson J. Behavioural syndromes: an
(1993) 48:26–34. doi: 10.1037/0003-066X.48.1.26 ecological and evolutionary overview. Trends Ecol Evol. (2004) 19:372–8.
3. Gosling SD. From mice to men: what we can learn about doi: 10.1016/j.tree.2004.04.009
personality from animal research? Psychol Bull. (2001) 127:45–86. 15. Koolhaas JM, Korte SM, De Boer SF, Van Der Vegt BJ, Van Reenen
doi: 10.1037/0033-2909.127.1.45 CG, Hopster H, et al. Coping styles in animals: current status in
4. Buss AH. Personality: Temperament, Social Behavior and the Self. Needham behaviour and stress-physiology. Neurosci Biobehav Rev. (1999) 23:925–35.
Heights, MA: Allyn & Bacon (1995). doi: 10.1016/S0149-7634(99)00026-3
5. Goldsmith H, Buss A, Plomin R, Rothbart M, Thomas A, Chess S, et al. 16. Korte SM, Koolhaas JM, Wingfield JC, McEwen BS. The Darwinian
Roundtable: what is temperament? Four approaches. Child Dev. (1987) concept of stress: benefits of allostasis and costs of allostatic load and the
58:505–29. doi: 10.2307/1130527 trade-offs in health and disease. Neurosci Biobehav Rev. (2005) 29:3–38.
6. McCrae RR, Costa JR, Paul T, Ostendorf F, Angleitner A, Hrebíčkovà M, et al. doi: 10.1016/j.neubiorev.2004.08.009
Nature over nurture: temperament, personality and life span development. J 17. Grignard L, Boissy A, Boivin X, Garel JP, Le Neindre P. The social
Pers Soc Psychol. (2000) 78:173–86. doi: 10.1037/0022-3514.78.1.173 environment influences the behavioural responses of beef cattle to handling.
7. Rothbart MK, Ahadi SA, Evans DE. Temperament and personality: Appl Anim Behav Sci. (2000) 68:1–11. doi: 10.1016/S0168-1591(00)00085-X
origins and outcomes. J Pers Soc Psychol. (2000) 78:122–35. 18. Melotti L, Oostindjer M, Bolhuis JE, Held S, Mendl M. Coping
doi: 10.1037/0022-3514.78.1.122 personality type and environmental enrichment affect aggression
8. Buss AH. Personality: primate heritage and human distinctiveness. In: at weaning in pigs. Appl Anim Behav Sci. (2011) 133:144–53.
Aronoff J, Rabin A, editors. Emergence of Personality. New York, NY: doi: 10.1016/j.applanim.2011.05.018
Springer Publishing Co. (1987). p. 57–101. 19. Cannon WB. Bodily Changes in Pain, Hunger, Fear and Rage. New York, NY:
9. Affleck G, Tennen H. Construing benefits from adversity: adaptional Appleton (1929).
significance and dispositional underpinnings. J Pers. (1996) 64:899–922. 20. Boissy A, Bouix J, Orgeur P, Poindron P, Bibe B, Le Neindre P. Genetic
doi: 10.1111/j.1467-6494.1996.tb00948.x analysis of emotional reactivity in sheep: effects of the genotypes of
10. Lazarus RS. Coping theory and research: past, present, and future. Psychosom the lambs and of their dams. Genet Select Evol. (2005) 37:381–401.
Med. (1993) 55:234–47. doi: 10.1097/00006842-199305000-00002 doi: 10.1186/1297-9686-37-5-381

21. Miranda-de la Lama GC, Sepúlveda WS, Montaldo HH, María GA, 42. Bouwhuis S, Quinn JL, Sheldon BC, Verhulst S. Personality and basal
Galindo F. Social strategies associated with identity profiles in dairy goats. metabolic rate in a wild bird population. Oikos (2013) 123:56–62.
Appl Anim Behav Sci. (2011) 134:48–55. doi: 10.1016/j.applanim.2011. doi: 10.1111/j.1600-0706.2013.00654.x
06.004 43. West-Eberhard MJ. Phenotypic accommodation: adaptive innovation
22. Holbrook CT, Wright CM, Pruitt JN. Individual differences in personality due to developmental plasticity. J Exp Zool. (2005) 304B:610–8.
and behavioural plasticity facilitate division of labour in social spider doi: 10.1002/jez.b.21071
colonies. Anim Behav. (2014) 97:177–83. doi: 10.1016/j.anbehav.2014.09.015 44. Piersma T, Drent J. Phenotipic flexibility and the evolution
23. Kilgour RJ, Melville GJ, Greenwood PL. Individual differences in the reaction of organismal design. Trends Ecol Evol. (2003) 18:228–33.
of beef cattle to situations involving social isolation, close proximity of doi: 10.1016/S0169-5347(03)00036-3
humans, restraint and novelty. Appl Anim Behav Sci. (2006) 99:21–40. 45. Groothuis TGG, Maestripieri D. Parental influences on offspring personality.
doi: 10.1016/j.applanim.2005.09.012 In: Carere, C, Maestripieri D, editors. Animal Personalities, Chicago. IL:
24. Boissy A, Bouissou MF. Assessment of individual differences in University of Chicago Press (2013). p. 317–52.
behavioural reactions of heifers exposed to various fear-eliciting situations. 46. Biro PA, Beckmann C, Stamps JA. Small within-day increases in temperature
Appl Anim Behav Sci. (1995) 46:17–31. doi: 10.1016/0168-1591(95) affects boldness and alters personality in coral reef fish. Proc R Soc Lond Biol.
00633-8 (2009) 277:71–7. doi: 10.1098/rspb.2009.1346
25. Reimert I, Rodenburg TB, Ursinus WW, Kemp B, Bolhuis JE. Responses 47. Finkemeier MA, Trillmich F, Guenther A. Match-mismatch experiments
to novel situations of female and castrated male pigs with divergent using photoperiod expose developmental plasticity of personality traits.
social breeding values and different backtest classifications in barren Ethology (2016) 122:80–93. doi: 10.1111/eth.12448
and straw-enriched housing. Appl Anim Behav Sci. (2014) 151:24–35. 48. Guenther A, Trillmich F. Photoperiod influences the behavioral and
doi: 10.1016/j.applanim.2013.11.015 physiological phenotype during ontogeny. Behav Ecol. (2013) 24:402–11.
26. Réale D, Garant D, Humphries MM, Bergeron P, Careau V, Montiglio doi: 10.1093/beheco/ars177
PO. Personality and the emergence of the pace-of-life syndrome concept 49. Haage M, Bergvall UA, Maran T, Kiik K, Angerbjörn A. Situation
at the population level. Philos Trans R Soc B. (2010) 365:4051–63. and context impacts the expression of personality: the influence of
doi: 10.1098/rstb.2010.0208 breeding season and test context. Behav Process. (2013) 100:103–9.
27. Réale D, Dingemanse NJ (eds.). Animal personality. In eLS (2012). doi: 10.1016/j.beproc.2013.08.009
doi: 10.1002/9780470015902.a0023570 50. Sutherland MA, Rogers AR, Verkerk GA. The effect of temperament
28. Constantini D, Ferrari C, Pasquaretta C, Cavallone E, Carere C, Von and responsiveness towards humans on the behavior, physiology
Hardenberg A, et al. Interplay between plasma oxidative status, cortisol and and milk production of multi-parous dairy cows in a familiar
coping styles in wild alpine marmots, Marmota marmota. J Exp Biol. (2012) and novel milking environment. Physiol Behav. (2012) 107:329–37.
215:374–83. doi: 10.1242/jeb.062034 doi: 10.1016/j.physbeh.2012.07.013
29. Réale D, Reader SM, Sol D, McDougall PT, Dingemanse NJ. Integrating 51. Sant’Anna AC, Paranhos da Costa MJR. Validity and feasibility of qualitative
animal temperament within ecology and evolution. Biol Rev Camb Philos Soc. behavior assessment for the evaluation of Nellor cattle temperament. Livest
(2007) 82:291–318. doi: 10.1111/j.1469-185X.2007.00010.x Sci. (2013) 157:254–62. doi: 10.1016/j.livsci.2013.08.004
30. Guenther A, Finkemeier MA, Trillmich F. The ontogeny of 52. Webb LE, van Reenen CG, Jensen MB, Schmitt O, Bokkers EAM. Does
personality in the wild guinea pig. Anim Behav. (2014) 90:131–9. temperament affect learning in calves? Appl Anim Behav Sci. (2015) 165:33–9.
doi: 10.1016/j.anbehav.2014.01.032 doi: 10.1016/j.applanim.2015.01.013
31. Verbeek MEM, Boon A, Drent PJ. Exploration, aggressive behavior and 53. Burdick NC, Carroll JA, Hulbert LE, Dailey JW, Willard ST, Vann RC,
dominance in pair-wise confrontations of juvenile male great tits. Behaviour et al. Relationships between temperament and transportation with rectal
(1996) 113:945–63. doi: 10.1163/156853996X00314 temperature and serum concentrations of cortisol and epinephrine in bulls.
32. Gosling SD, John OP. Personality dimensions in nonhuman animals: Livest Sci. (2010) 129:166–72. doi: 10.1016/j.livsci.2010.01.020
a cross-species review. Curr Dir Psychol Sci. (1999) 8:69–75. 54. Frondelius L, Järvenranta K, Koponen T, Mononen J. The effects of body
doi: 10.1111/1467-8721.00017 posture and temperament on heart rate variability in dairy cows. Physiol
33. Costa PT, McCrae RR. Revised NEO Personality Inventory (NEO PI-R) Behav. (2015) 139:437–41. doi: 10.1016/j.physbeh.2014.12.002
and NEO Five-Factor Inventory (NEO-FFI): Professional Manual. Oxford: 55. Gibbons J, Lawrence A, Haskell M. (2009). Responsiveness of dairy cows to
Psychological Assessment Resources, Incorporated (1992). human approach and novel stimuli. Appl Anim Behav Sci. (2009) 116:163–73.
34. Koski SE. Broader horizons for animal personality research. Front Ecol Evol. doi: 10.1016/j.applanim.2008.08.009
(2014) 2:70. doi: 10.3389/fevo.2014.00070 56. Graunke KL, Nürnberg G, Repsilber D, Puppe B, Langbein J. Describing
35. Altschul DM, King JE, Inoue-Murayama M, Ross SR, Weiss A. Longevity and temperament in an ungulate: a multidimensional approach. PLoS ONE
personality in captive chimpanzees (Pan troglodytes). Peer J. Preprints (2016). (2013) 8:1–12. doi: 10.1371/journal.pone.0074579
4:e1916w2. doi: 10.7287/peerj.preprints.1916v2 57. Boles JA, Kohlbeck KS, Meyers MC, Perz KA, Davis KC, Thomson JM. The
36. Eckardt W, Steklis HD, Steklis NG, Fletcher AW, Stoinski TA, Weiss A. use of blood lactate concentration as an indicator of temperament and its
Personality dimensions and their behavioral correlates in wild virunga impact on growth rate and tenderness of steaks from Simmental × Angus
mountain gorillas (Gorilla beringei beringei). J Comp Psychol. (2015) 129:26– steers. Meat Sci. (2015) 103:68–74. doi: 10.1016/j.meatsci.2015.01.003
41. doi: 10.1037/a0038370 58. Cafe LM, Robinson DL, Ferguson DM, Geesink GH, Greenwood PL.
37. David M, Dall SRX. Unravelling the philosophies underlying ‘animal Temperament and hypothalamic-pituitary-adrenal axis function are related
personality’ studies: a brief re-appraisal of the field. Ethology (2016) 122:1–9. and combine to affect growth, efficiency, carcass, and meat quality
doi: 10.1111/eth.12445 traits in Brahman steers. Domest Anim Endocrinol. (2011) 40:230–40.
38. Dall SRX, Houston AI, McNamara JM. The behavioural ecology of doi: 10.1016/j.domaniend.2011.01.005
personality: consistent individual differences from an adaptive perspective. 59. Phocas F, Boivin X, Sapa J, Trillat G, Boissy A, Le Neindre P. Genetic
Ecol Lett. (2004) 7:734–9. doi: 10.1111/j.1461-0248.2004.00618.x correlations between temperament and breeding traits in Limousin heifers.
39. Dingemanse NJ, Both C, Van Noordwijk AJ. Rutten AL, Drent PJ. Natal Anim Sci. (2006) 82:805–11. doi: 10.1017/ASC200696
dispersal and personalities in great tits (Parus major). Proc R Soc Lond Biol. 60. Brand B, Hadlich F, Brandt B, Schauer N, Graunke KL, Langbein
(2003) 270:741–7. doi: 10.1098/rspb.2002.2300 J, et al. (2015). Temperament type specific metabolite profiles of the
40. Dingemanse NJ, Réale D. Natural selection and animal personality. prefrontal cortex and serum in cattle. PLoS ONE (2015) 10:e0125044.
Behaviour (2005) 142:1159–84. doi: 10.1163/156853905774539445 doi: 10.1371/journal.pone.0125044
41. Biro PA, Stamps JA. Are animal personality traits linked to 61. del Campo M, Brito G, Soares de Lima J, Hernández P, Montossi F. Finishing
life-history productivity? Trends Ecol Evol. (2008) 23:361–8. diet, temperament and lairage time effects on carcass and meat quality traits
doi: 10.1016/j.tree.2008.04.003 in steers. Meat Sci. (2010) 86:908–14. doi: 10.1016/j.meatsci.2010.07.014

62. Lyons DM, Price EO, Moberg GP. Individual differences in temperament of 82. Lansade L, Bouissou MF. Reactivity to humans: a temperament trait of horses
domestic dairy goats: constancy and change. Anim Behav. (1988) 36:1323–33. which is stable across time and situations. Appl Anim Behav Sci. (2008)
doi: 10.1016/S0003-3472(88)80201-X 114:492–508. doi: 10.1016/j.applanim.2008.04.012
63. Lyons DM. Individual differences in temperament of dairy goats and 83. Lansade L, Bouissou MF, Boivin X. Temperament in preweanling horses:
the inhibition of milk ejection. Appl Anim Behav Sci. (1989) 22:269–82. development of reactions to humans and novelty, and startle responses. Dev
doi: 10.1016/0168-1591(89)90022-1 Psychobiol. (2007) 49:501–13. doi: 10.1002/dev.20233
64. Nawroth C, Prentice PM, McElligott AG. Individual personality 84. Krüger K, Farmer K, Heinze J. The effects of age, rank and
differences in goats predict their performance in visual learning neophobia on social learning in horses. Anim Cogn. (2014) 17:645–55.
and non-associative cognitive tasks. Behav Process. (2017) 134:43–53. doi: 10.1007/s10071-013-0696-x
doi: 10.1016/j.beproc.2016.08.001 85. Lansade L, Philippon P, Herve L, Vidament M. Development of personality
65. Beausoleil NJ, Blache D, Stafford KJ, Mellor DJ, Noble ADL. Selection for tests to use in the field, stable over time and across situations, and linked to
temperament in sheep: domain-general and context-specific traits. Appl horses’ show jumping performance. Appl Anim Behav Sci. (2016) 176:43–51.
Anim Behav Sci. (2012) 139:74–85. doi: 10.1016/j.applanim.2012.02.020 doi: 10.1016/j.applanim.2016.01.005
66. Hawken PAR, Fiol C, Blache D. Genetic differences in temperament 86. Duberstein KJ, Gilkeson JA. Determination of sex differences in personality
determine whether lavender oil alleviates or exacerbates anxiety in sheep. and trainability of yearling horses utilizing a handler questionnaire. Appl
Physiol Behav (2012a) 105:1117–23. doi: 10.1016/j.physbeh.2011.12.005 Anim Behav Sci. (2010) 128:57–63. doi: 10.1016/j.applanim.2010.09.012
67. Hazard D, Moreno C, Foulquie D, Delval E, Francois D, Bouix J, et al. 87. Roberts K, Hemmings AJ, Moore-Colyer M, Parker MO, McBride SD.
Identification of QTLs for behavioral reactivity to social separation and Neural modulators of temperament: a multivariate approach to personality
humans in sheep using the OvineSNP50 BeadChip. BMC Genomics (2014) trait identification in the horse. Physiol Behav. (2016) 167:125–31.
15:778. doi: 10.1186/1471-2164-15-778 doi: 10.1016/j.physbeh.2016.08.029
68. McBride SD, Wolf B. Using multivariate statistical analysis to measure 88. Ruis MAW, te Brake JHA, Engel B, Buist WG, Blokhuis HJ, Koolhaas
ovine temperament; stability of factor construction over time and JM. Adaptation to social isolation acute and long-term stress responses of
between groups of animals. Appl Anim Behav Sci. (2007) 103:45–58. growing gilts with different coping characteristics. Physiol Behav. (2001)
doi: 10.1016/j.applanim.2006.04.030 73:541–51. doi: 10.1016/S0031-9384(01)00548-0
69. Sibbald AM, Erhard HW, McLeod JE, Hooper RJ. Individual personality 89. Oster M, Scheel M, Muráni E, Ponsuksili S, Zebunke M, Puppe B, et al. The
and the spatial distribution of groups of grazing animals: an example fight-or-flight response is associated with PBMC expression profiles related
with sheep. Behav Process. (2009) 82:319–26. doi: 10.1016/j.beproc.2009. to immune defence and recovery in swine. PLoS ONE (2015) 10:e0120153.
07.011 doi: 10.1371/journal.pone.0120153
70. Seaman SC, Davidson HPB, Waran NK. How reliable is temperament 90. Bolhuis JE, Parmentier HK, Schouten WGP, Schrama JW, Wiegant
assessment in the domestic horse (Equus caballus)? Appl Anim Behav Sci. VM. Effects of housing and individual coping characteristics
(2002) 78:175–91. doi: 10.1016/S0168-1591(02)00095-3 on immune responses of pigs. Physiol Behav. (2003) 79:289–96.
71. Lansade L, Simon F. Horses’ learning performances are under the influence doi: 10.1016/S0031-9384(03)00090-8
of several temperamental dimensions. Appl Anim Behav Sci. (2010) 125:30– 91. Zebunke M, Repsilber D, Nürnberg G, Wittenburg D, Puppe B. The backtest
37. doi: 10.1016/j.applanim.2010.02.010 in pigs revisited–an analysis of intra-situational behaviour. Appl Anim Behav
72. von Borstel UK, Pasing S, Gauly M. Towards a more objective assessment Sci. (2015) 169:17–25. doi: 10.1016/j.applanim.2015.05.002
of equine personality using behavioural and physiological observations 92. Bolhuis JE, Schouten WGP, de Leeuw JA, Schrama JW, Wiegant
from performance test training. Appl Anim Behav Sci. (2011) 135:277–85. VM. Individual coping characteristics, rearing conditions and
doi: 10.1016/j.applanim.2011.10.007 behavioural flexibility in pigs. Behav Brain Res. (2004) 152:351–60.
73. Lansade L, Bouissou MF, Erhard HW. Fearfulness in horses: a temperament doi: 10.1016/j.bbr.2003.10.024
trait stable across time and situations. Appl Anim Behav Sci. (2008) 115:182– 93. Reimert I, Bolhuis JE, Kemp B, Rodenburg TB. Social support in
200. doi: 10.1016/j.applanim.2008.06.011 pigs with different coping styles. Physiol Behav. (2014) 129:221–9.
74. Graf P, von Borstel UK, Gauly M. Practical considerations regarding doi: 10.1016/j.physbeh.2014.02.059
the implementation of a temperament test into horse performance 94. Spake JR, Gray KA, Cassady JP. Relationship between backtest and
tests: results of a large-scale test run. J Vet Behav. (2014) 9:329–40. coping styles in pigs. Appl Anim Behav Sci. (2012) 140:146–53.
doi: 10.1016/j.jveb.2014.08.004 doi: 10.1016/j.applanim.2012.06.007
75. Hausberger M, Muller C, Lunel C. Does work affect personality? A study in 95. Zebunke M, Nürnberg G, Melzer N, Puppe B. The backtest in pigs revisited
horses. PLoS ONE (2011) 6:e14659. doi: 10.1371/journal.pone.0014659 – inter-situational behaviour and animal classification. Appl Anim Behav Sci.
76. Ijichi C, Collins LM, Elwood RM. Pain expression is linked to (2017) 194:7–13. doi: 10.1016/j.applanim.2017.05.011
personality in horses. Appl Anim Behav Sci. (2014) 152:38–42. 96. Janczak AM, Pedersen LJ, Bakken M. Aggression, fearfulness and
doi: 10.1016/j.applanim.2013.12.007 coping styles in female pigs. Appl Anim Behav Sci. (2003) 81:13–28.
77. Górecka-Bruzda A, Jastrzebska E, Sosnowska Z, Jaworski Z, Jezierski doi: 10.1016/S0168-1591(02)00252-6
T, Chruszczewski MH. Reactivity to humans and fearfulness tests: field 97. Kooij EvErp-vd, Kuijpers AH, Schrama JW, van Eerdenburg FJCM,
validation in Polish Cold Blood Horses. Appl Anim Behav Sci. (2011) Schouten WGP, Tielen MJM. Can we predict behaviour in pigs? Searching for
133:207–15. doi: 10.1016/j.applanim.2011.05.011 consistency in behaviour over time and across situations. Appl Anim Behav
78. Lloyd AS, Martin JE, Bornett-Gauci HLI, Wilkinson RG. Horse personality: Sci. (2002) 75:293–305. doi: 10.1016/S0168-1591(01)00203-9
variation between breeds. Appl Anim Behav Sci. (2008) 112:369–83. 98. Brown JA, Dewey C, Delange CFM, Mandell IB, Purslow PP, Robinson JA,
doi: 10.1016/j.applanim.2007.08.010 et al. Reliability of temperament tests on finishing pigs in group-housing
79. Visser EK, van Reenen CG, Hopster H, Schilder MBH, Knaap JH, and comparison to social tests. Appl Anim Behav Sci. (2009) 118:28–35.
Barneveld A, et al. Quantifying aspects of young horses’ temperament: doi: 10.1016/j.applanim.2009.02.005
consistency of behavioural variables. Appl Anim Behav Sci. (2001) 74:241–58. 99. Ruis MAW, te Brake JHA, van de Burgwal JA, de Jong IC, Blokhuis
doi: 10.1016/S0168-1591(01)00177-0 HJ, Koolhaas JM. Personalities in female domesticated pigs: behavioural
80. Visser EK, van Reenen CG, Engel B, Schilder MBH, Barneveld A, and physiological indications. Appl Anim Behav Sci. (2000) 66:31–47.
Blokhuis HJ. The association between performance in show-jumping and doi: 10.1016/S0168-1591(99)00070-2
personality traits earlier in life. Appl Anim Behav Sci. (2003) 82:279–95. 100. Randle HD. Facial hair whorl position and temperament in cattle. Appl Anim
doi: 10.1016/S0168-1591(03)00083-2 Behav Sci. (1998) 56:139–47. doi: 10.1016/S0168-1591(97)00086-5
81. Olsen HF, Klemetsdal G. Temperament of the Norwegian horse breeds-a 101. Beausoleil NJ, Blach D, Stafford KJ, Mellor DJ, Noble ADL. Exploring the
questionnaire based study. Appl Anim Behav Sci. (2017) 193:60–6. basis of divergent selection for “temperament” in domestic sheep. Appl Anim
doi: 10.1016/j.applanim.2017.03.015 Behav Sci. (2008) 109:261–74. doi: 10.1016/j.applanim.2007.03.013

102. Jones AC, Gosling SD. Temperament and personality in dogs (Canis 124. Colléter M, Brown C. Personality traits predict hierarchy rank in
familiaris): a review and evaluation of past research. Appl Anim Behav Sci. male rainbowfish social groups. Anim Behav. (2011) 81:1231–7.
(2005) 95:1–53. doi: 10.1016/j.applanim.2005.04.008 doi: 10.1016/j.anbehav.2011.03.011
103. Benus RF, Bohus B, Koolhaas JM, van Oortmerssen GA. Behavioral strategies 125. Krause A, Puppe B, Langbein J. Coping style modifies general and affective
of aggressive and nonaggressive male-mice in active shock avoidance. Behav autonomic reactions of domestic pigs in different behavioral contexts. Front
Processes (1989) 20:1–12. doi: 10.1016/0376-6357(89)90008-9 Behav Neurosci. (2017) 11:103. doi: 10.3389/fnbeh.2017.00103
104. Boersma GJ, Scheurink AJ, Wielinga PY, Steimer TJ, Benthem 126. Ponsuksili S, Zebunke M, Murani E, Trakooljul N, Krieter J, Puppe
L. The passive coping Roman Low Avoidance rat, a non-obese B, et al. Integrated genome-wide association and hypothalamus eQTL
rat model for insulin resistance. Physiol Behav. (2009) 97:353–8. studies indicate a link between the circadian rhythm-related gene
doi: 10.1016/j.physbeh.2009.03.005 PER1 and coping behaviour. Sci Rep. (2015) 5:16264. doi: 10.1038/srep
105. Posner J, Russell JA, Peterson BS. The circumplex model of 16264
affect: an integrative approach to affective neuroscience, cognitive 127. Murphy E, Nordquist RE, van der Staay FJ. A review of behavioural methods
development, and psychopathology. Dev Psychopathol. (2005) 17:715–34. to study emotion and mood in pigs, Sus scrofa. Appl Anim Behav Sci. (2014)
doi: 10.1017/S0954579405050340 159:9–28. doi: 10.1016/j.applanim.2014.08.002
106. Meager JJ, Fernö A, Skjæraasen JE, Järvi T, Rodewald P. Multidimensionality 128. Koolhaas JM, van Reenen CG. Animal Behavior and Well-Being
of behavioural phenotypes in Atlantic cod, Gadus morhua. Physiol Behav. Symposium: interaction between coping style/personality, stress, and
(2012) 106:462–70. doi: 10.1016/j.physbeh.2012.03.010 welfare: relevance for domestic farm animals. J Anim Sci. (2016) 94:2284–96.
107. Weinstein TAR, Capitanio JP. Individual differences in infant temperament doi: 10.2527/jas.2015-0125
predict social relationships of yearling rhesus monkeys, Macaca mulatta. 129. Mendl M, Burman OHP, Paul, ES. An integrative and functional framework
Anim Behav. (2008) 76:455–65. doi: 10.1016/j.anbehav.2008.01.024 for the study of animal emotion and mood. Proc R Soc Lond Biol. (2010)
108. Foyer P, Wilsson E, Jensen P. Levels of maternal care in dogs affect adult 277:2895–904. doi: 10.1098/rspb.2010.0303
offspring temperament. Sci Rep. (2016) 6:19253. doi: 10.1038/srep19253 130. Russell J. A circumplex model of affect. J Pers Soc Psychol. (1980) 39:1161–78.
109. Iki T, Ahrens F, Pasche KH, Bartels A, Erhard MH. Relationships between doi: 10.1037/h0077714
scores of the feline temperament profile and behavioural and adrenocortical 131. Carter AJ, Feeney WE, Marshall HH, Cowlishaw G, Heinsohn R. Animal
responses to a mild stressor in cats. Appl Anim Behav Sci. (2011) 132:71–80. personality: what are behavioural ecologists measuring? Biol Rev Camb Philos
doi: 10.1016/j.applanim.2011.03.008 Soc. (2013) 88:465–75. doi: 10.1111/brv.12007
110. Le Neindre P, Trillat G, Sapa J, Ménissier F, Bonnet JN, Chupin JM. Individual 132. Forkman B, Boissy A, Meunier-Salaün MC, Canali E, Jones RB. A critical
differences in docility in limousin cattle. J Anim Sci. (1995) 73:2249–53. review of fear tests used on cattle, pigs, sheep, poultry and horses. Physiol
doi: 10.2527/1995.7382249x Behav. (2007) 92:340–74. doi: 10.1016/j.physbeh.2007.03.016
111. Gibbons J. The Effect of Selecting for “Robustness” on Temperament in Dairy 133. Handley SL, Mithani S. Effects of alpha-adrenoceptor agonists and
Cows. Ph.D., thesis, The University of Edinburgh (2009). antagonists in a maze-exploration model of ‘fear’-motivated behaviour.
112. Coleman K, Wilson DS. Shyness and boldness in pumpkinseed sunfish: Naunyn Schmiedebergs Arch Pharmacol. (1984) 327:1–5.
individual differences are context-specific. Anim Behav. (1998) 56:927–36. 134. Montgomery KC. The relation between fear induced by novel stimulation
doi: 10.1006/anbe.1998.0852 and exploratory drive. J Comp Physiol Psychol. (1955) 48:254–60.
113. Zidar J, Balogh A, Favati A, Jensen P, Leimar O, Løvlie H. A comparison doi: 10.1037/h0043788
of animal personality and coping styles in the red junglefowl. Anim Behav. 135. Perals D, Griffin AS, Bartomeus I, Sol D. Revisiting the open-field test: what
(2017) 130:209–20. doi: 10.1016/j.anbehav.2017.06.024 does it really tell us about animal personality? Anim Behav. (2017) 123:69–79.
114. Carere C, Drent PJ, Privitera L, Koolhaas JM, Groothuis TGG. Personalities doi: 10.1016/j.anbehav.2016.10.006
in great tits, Parus mayor: stability and consistency. Anim Behav. (2005) 136. Hessing MJC, Hagelso AM, Vanbeek JAM., Wiepkema PR, Schouten WGP,
70:795–805. doi: 10.1016/j.anbehav.2005.01.003 Krukow R. Individual behavioral characteristics in pigs. Appl Anim Behav Sci.
115. Carere C, Caramaschi D, Fawcett T. Covariation between personalities (1993) 37:285–95. doi: 10.1016/0168-1591(93)90118-9
and individual differences in coping with stress: converging evidence and 137. Guenther A, Trillmich F. Within-litter differences in personality and
hypotheses. Curr Zool. (2010) 56:728–40. physiology relate to size differences among siblings in cavies. Physiol Behav.
116. Coppens CM, De Boer SF, Koolhaas JM. Coping styles and behavioural (2015) 145:22–8. doi: 10.1016/j.physbeh.2015.03.026
flexibility: towards underlying mechanisms. Philos Trans R Soc B. (2010) 138. Diener E, Suh EM, Lucas RE, Smith HL. Subjective well-being:
365:4021–8. doi: 10.1098/rstb.2010.0217 three decades of progress. Psychol Bull. (1999) 125:276–302.
117. Koolhaas JM, De Boer SF, Coppens CM, Buwalda B. Neuroendocrinology doi: 10.1037/0033-2909.125.2.276
of coping styles: towards understanding the biology of individual 139. Weiss A, Bates TC, Luciano M. Happiness is a personal(ity) thing: the
variation. Front Neuroendocrinol. (2010) 31:307–21. doi: 10.1016/j.yfrne. genetics of personality and well-being in a representative sample. Psychol Sci.
2010.04.001 (2008) 19:205–10. doi: 10.1111/j.1467-9280.2008.02068.x
118. Uher J. Individual behavioural phenotypes: an integrative meta-theoretical 140. DeNevre KM, Cooper H. Origins and functions of positive and negative
framework. Why “Behavioral Syndromes” are not analogs of “personality”. affect: a control-process view. Psychol Rev. (1998) 124:197–229.
Dev Psychobiol. (2011) 53:521–48. doi: 10.1002/dev.20544 141. Tetley CL, O’Hara SJ. Ratings of animal personality as a tool for improving
119. Koolhaas JM. Coping style and immunity in animals: making the breeding management and welfare of zoo animals. Animal Welf. (2012)
sense of individual variation. Brain Behav Immun. (2008) 22:662–7. 21:463–76. doi: 10.7120/09627286.21.4.463
doi: 10.1016/j.bbi.2007.11.006 142. Dawkins MS. Evolution and animal welfare. Q Rev Biol. (1998) 73:305–28.
120. Careau V, Thomas D, Humphries M, Réale D. Energy metabolism and animal doi: 10.1086/420307
personality. Oikos (2008) 117:641–53. doi: 10.1111/j.0030-1299.2008.16513.x 143. Dawkins MS. Using behaviour to assess animal welfare. Anim Welf. (2004)
121. Trompf L, Brown C. Personality affects learning and trade-offs between 13:3–7.
private and social information in guppies, Poecilia reticulate. Anim Behav. 144. Fureix C, Menguy H, Hausberger M. Partners with bad temper: reject or
(2014) 88:99–106. doi: 10.1016/j.anbehav.2013.11.022 cure? A study of chronic pain and aggression in horses. PLoS ONE (2008)
122. Brust V, Wuerz Y, Krüger O. Behavioural flexibility and personality in zebra 5:e12434. doi: 10.1371/journal.pone.0012434
finches. Ethology (2013) 119:559–69. doi: 10.1111/eth.12095 145. Burrow HM, Dillon RD. Relationship between temperament and growth in
123. Silva PIM, Martins CIM, Engrola S, Marino G, Øverli Ø, Conceição LEC. a feedlot and commercial carcass traits in Bos indicus crossbreeds. Aust J Exp
Individual differences in cortisol levels and behaviour of Senegalese sole Agric. (1997) 37:407–11. doi: 10.1071/EA96148
(Solea senegalensis) juveniles: evidence for coping styles. Appl Anim Behav 146. Voisinet BD, Grandin T, Tatum JD, O’Connor SF, Struthers JJ.
Sci. (2010) 124:75–81. doi: 10.1016/j.applanim.2010.01.008 Feedlot cattle with calm temperaments have higher average daily

gains than cattle with excitable temperaments. J Anim Sci. (1997) 75: 158. Hellbrügge B, Tölle KH, Bennewitz J, Henze C, Presuhn U, Krieter J. Genetic
892–6. aspects regarding piglet losses and the maternal behaviour of sows. Part
147. Rauw WM., Johnson AK, Gomez-Raya L, Dekkers JCM. A hypothesis 2. Genetic relationship between maternal behaviour on sows and piglet
and review of the relationship between selection for improved production mortality. Animal (2008) 2:1281–8. doi: 10.1017/S1751731108002516
efficiency, coping behavior and domestication. Front Genet. (2017) 8:134. 159. Stukenborg A, Traulsen I, Stamer E, Puppe B, Presuhn U, Krieter J.
doi: 10.3389/fgene.2017.00134 Heritabilities of agonistic behavioural traits in pigs and their relationships
148. Drent PJ, van Oers K, Noordwijk AJ. Realized heritability of personalities within and between different age groups. Livest Sci. (2012) 149:25–32.
in the great tit (Parus major). Proc Royal Soc B. (2003) 270:45–51. doi: 10.1016/j.livsci.2012.06.020
doi: 10.1098/rspb.2002.2168 160. Hoppe S, Brandt HR, König S, Erhardt G, Gauly M. Temperament traits
149. van Oortmerssen GA, Bakker TC. Artificial selection for short and long of beef calves measured under field conditions and their relationships to
attack latencies in wild Mus musculus domesticus. Behav Genet. (1981) performance. J Anim Sci. (2010) 88:1982–9. doi: 10.2527/jas.2008-1557
11:115–26. doi: 10.1007/BF01065622 161. Hedlund L, Løvlie H. Personality and production: nervous cows produce less
150. van Oers K, Drent PJ, Jong GD, Noordwijk AJ. Additive and nonadditive milk. J Dairy Sci. (2015) 98:5819–28. doi: 10.3168/jds.2014-8667
genetic variation in avian personality traits. Heredity (2004) 93:496–503. 162. Hawken PAR, Williman M, Milton J, Kelly R, Nowak R, Blache D. Nutritional
doi: 10.1038/sj.hdy.6800530 supplementation during the last week of gestation increased the volume
151. Henderson CR. Applications of Linear Models in Animal Breeding. Guelph, and reduced the viscosity of colostrum produced by twin bearing ewes
ON: University of Guelph (1984). selected for nervous temperament. Small Rumin Res. (2012b) 105:308–14.
152. Kruuk LE. Estimating genetic parameters in natural populations using the doi: 10.1016/j.smallrumres.2012.01.011
‘animal model’. Proc R Soc B. (2004) 359:873–90. doi: 10.1098/rstb.2003.1437 163. Boersma GJ, Benthem L, van Beek AP, van Dijk G, Scheurink AJW.
153. Laine VN, van Oers K. The quantitative and molecular genetics of individual Personality, a key factor in personalized medicine? Eur J Pharmacol. (2011)
differences in animal personality. In: Vonk J, Weiss A, Kuczaj S, editors. 667:23–25. doi: 10.1016/j.ejphar.2011.05.079
Personality in Nonhuman Animals. Cham: Springer (2017). p. 55–72. 164. Clark B, Stewart GB, Panzone LA, Kyriazakis I, Frewer LJ. Citizens,
154. Careau V, Thomas D, Pelletier F, Turki L, Landry F, Garant D, et al. Genetic consumers and farm animal welfare: a meta-analysis of willingness-to-pay
correlation between resting metabolic rate and exploratory behaviour studies. Food Policy (2017) 68:112–27. doi: 10.1016/j.foodpol.2017.01.006
in deer mice (Peromyscus maniculatus). J Evol Biol. (2011) 24:2153–63.
doi: 10.1111/j.1420-9101.2011.02344.x Conflict of Interest Statement: The authors declare that the research was
155. Haskell MJ, Simm G, Turner SP. Genetic selection for temperament conducted in the absence of any commercial or financial relationships that could
traits in dairy and beef cattle. Front Genet. (2014) 5:368. be construed as a potential conflict of interest.
doi: 10.3389/fgene.2014.00368
156. Stirling DG, Réale D, Roff DA. Selection, structure and Copyright © 2018 Finkemeier, Langbein and Puppe. This is an open-access article
the heritability of behaviour. J Evol Biol. (2002) 15:277–89. distributed under the terms of the Creative Commons Attribution License (CC
doi: 10.1046/j.1420-9101.2002.00389.x BY). The use, distribution or reproduction in other forums is permitted, provided
157. D’Eath RB, Rohe R, Turner SP, Ison SH, Farish M, Jack MC, et al. Genetics the original author(s) and the copyright owner are credited and that the original
of animal temperament: aggressive behaviour at mixing is genetically publication in this journal is cited, in accordance with accepted academic practice.
associated with the response to handling in pigs. Animal (2009) 3:1544–54. No use, distribution or reproduction is permitted which does not comply with these
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published: 28 June 2018

Personality Research in Mammalian

Measuring and understanding personality in animals is a rising scientific field. Much

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TABLE 1 | Different personality-related terms in alphabetical order used to

Personality- Definition References

Behavioral Behavioral response to handling (17)

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Personality-related Test Measures Study

(A): CATTLE (Bos taurus)

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Personality-related Test Measures Study

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Some studies have shown evidence that emotionality, and

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