Personality in animals has been investigated across a variety of different scientific fields including agricultural science, animal behaviour, anthropology, psychology, veterinary medicine, and zoology. [1] Thus, the definition for animal personality may vary according to the context and scope of study. However, there is recent consensus in the literature for a broad definition that describes animal personality as individual differences in behaviour that are consistent across time and ecological context. [2] Here, consistency refers to the repeatability of behavioural differences between individuals and not a trait that presents itself the same way in varying environments. [3] [4]
Animal personality traits are measurable and are described in over 100 species. [5] Personality in non-human animals has also been referred to as animal disposition, coping style, and temperament. [1] There are also personality norms through the species, often found between genders. [6] The diversity of animal personality can be compared in cross-species studies, demonstrating its pervasiveness in the evolutionary process of animals. [1] Research on animal personality variation has been burgeoning since the mid 1990s. [7] Recent studies have focused on its proximate causation and the ecological and evolutionary significance of personality in animals. [4]
The extent of personality phenomena considered when examining animal personality is significantly reduced compared to those studied in humans. Concepts such as personal objects, identity, attitudes and life stories are not considered relevant in animals. Similarly, any approach that requires the subject to explain motives, beliefs or feelings is not applicable to the study of animal behaviour. [1]
The study of animal personality is largely based on the observation and investigation of behavioural traits. In an ecological context, traits or ‘characters’ are attributes of an organism that are shared by members of a species. Traits can be shared by all or only a portion of individuals in a population. For example, studies in animal personality often examine traits such as aggressiveness, avoidance of novelty, boldness, exploration and sociality. [3]
The initial framework used to study animal personality was comparative psychology. The descriptive language used by comparative psychologists in the late nineteenth century often attributed disposition and behavioural tendencies to individual animals in their studies. Many of these reports are the result of researchers anthropomorphizing the animal subjects and did not explicitly examine what is now considered animal personality. However, these studies do represent some of the first instances of scientists reporting individual differences in animal behaviour. [8]
Russian physiologist Ivan Pavlov was one of the first researchers to integrate personality into his research of animal behaviour. In his seminal studies on conditional reflexes, he categorized the behaviour of dogs as Excitable, Lively, Quiet or Inhibited. He linked these personalities to learning ability. The Excitable type, for example, showed signs of strong excitatory conditioning, but a limited ability for the acquisition of inhibitory connections. The Lively type was the most balanced and displayed rapid associative learning, while the Quiet type exhibited consistent but slow learning. [9]
The first study that empirically examined animal personality was done in 1938. Meredith Crawford quantified individual differences in the behaviour of young chimpanzees using a behaviour rating scale. Crawford conducted his research out of the Yale Laboratories of Primate Biology. [8] Since then, psychologists have continued to investigate personality in animals across a wide range of taxa. Meanwhile, the incorporation of animal personality into the fields of ecology and evolution is a relatively new practice. Ecologists began to recognize the importance of individual differences in behaviour near the end of the twentieth century. [10] [3]
Rating traits involve a group of observers that scrutinize the behavioural traits of an individual animal. Observers are typically given a list of characters and/or descriptions to use as a reference for interpreting animal behaviour then use the list to rate the behaviours. [11] The rating method uses observers and data recording instruments. [1]
Researchers will observe how individual animals respond to behavioural tests and code their reactions. This method is typically used in experimental studies in which environmental conditions are heavily manipulated. [1]
As research in animal personality became more prevalent in ecological studies many behavioural ecologists were substituting behavioural plasticity for animal personality. [10] That is, attributing variation in behaviour to diverse personality types rather than plasticity in a single behavioural trait. Thus, behavioural reaction norms were introduced as a means of incorporating personality and individual plasticity in the study of animal behaviour. Behavioural reaction norms measure the behaviour of an individual over an environmental gradient, therefore providing information on how an animal behaves on average and how their behaviour changes across an environmental gradient. [12]
Repeatability refers to the fraction of variation in a population that is owed to differences between individuals. Repeatability estimates are one of the most widely used statistical tools that can quantify consistent individual differences in behaviour. [13] Formally,
where, is variance among individuals and is the variance within individuals over time. [14] In a meta-analysis of published, peer-reviewed repeatability estimates, reviewers found that, in general, approximately 35% of behavioural variation among individuals could be attributed to individual differences. [15]
The Five Factor Model , or the Big Five personality trait model, has been used to assign personality archetypes to some animals. [6] The 5 categories for the five factor model for personality are openness to experience, conscientiousness, extraversion, agreeableness, and neuroticism. Each of these categories identifies personality factors at the broadest level of abstraction. Because the assessment of personality using the five factor model is often a self-reported measure for humans, applying this model to animals can be difficult to standardize. Researchers often measure personality by assessing the behavior of the being over a period of time to establish the pattern. Naturally, some animals may not have as wide a range of personality as humans do. The species of the animal determines how the personality manifests itself. Likewise, a species may be predisposed to exhibit a category of personality more than other categories. So far, chimpanzees are the only animal shown to exhibit conscientiousness. [16] Chimpanzees are also the only non-human species shown to demonstrate a hierarchical structure of personality, with two dimensions of affect corresponding to negative and positive emotionality, and a third dimension of disinhibition (vs. constraint), which is thought to comprise a regulatory system that is known to play a role in the perception and interpretation of incoming stimuli.[ clarification needed ] [17]
Neuroticism, agreeableness and extraversion are the most commonly found personality traits among measured animals. For example, chimpanzees show emotional stability, agreeableness and surgency, [18] audiovisual reactivity, affect-extraversion, [19] excitability-agitation, aggression, affinity and social play. [20]
Some behaviors are correlated into a set of personality traits that remain constant throughout different situations and contexts. These traits are referred to as behavioral syndromes. For example, the aggressiveness-boldness syndrome refers to the correlation between an individual's aggressiveness with conspecifics and boldness in novel environments. [2]
Many correlated behaviours are species specific. For instance, a study in 2014 reported that for horses, personality (quantified using a validated questionnaire) and tolerance to pain (using a Likert scale) indicated that neuroticism is negatively related to stoicism whereas extroversion was positively related to levels of expressed lameness. This suggests that pain may be more easily identified in highly extrovert horses. [21] In zebrafish (Danio rerio), Proactive and Reactive personalities express different thermal preferences and general activity within the temperature gradient. Proactive fish (more aggressive, bold risk-takers, prone to routine formation) have a preference for higher temperature environments. Reactive fish (shy, less risk-prone, more flexible) favor medium colder temperatures. [22]
The degree of variation in a population has been determined to influence the direction and outcome of natural selection. Most scientific research has focused on genetic and phenotypic variation or differences in resource use; however, variation in consistent behaviours (i.e. personality) also has important evolutionary consequences. For example, personality in animals can affect the way individuals interact with their environment and with each other which can affect the relative fitness of individuals. [23] Therefore, personality can influence selection. Also, behavioural traits are more dynamic which may allow an animal to adapt more quickly which, in turn, can speed up the rate of evolution. [2]
Further, natural or artificial selection cannot act on personality unless there is a mechanism for its inheritance. In rhesus macaques (Maccaca mulatta), the personality traits of Meek, Bold, Aggressive, Passive, Loner and Nervous have heritability values of 0.14 to 0.35, thus indicated that there is some genetic basis to the expression of personality traits in animals. In apes, including humans, heritability estimates of personality dimensions range from 0.07 to 0.63. [24] In horses, heritability estimates range mostly between h2=0.15 and h2=0.40 for traits assessed in personality tests. Values at this level are considered as "promising" for artificial selection. [25]
Personality in animals has been studied across a wide array of taxa. Some of these studies have investigated personality in mammals, [26] elasmobranchs, [27] reptiles, [28] fish [29] and birds. [30]
There have been several studies that have either been notable in that they have changed scientific understandings of animal personality or have applications in the field of human personality. For example, the study of personality in chimpanzees by King and Figueredo in 1997 [31] was one of the first studies to apply the five-factor model in animal personality. It demonstrated the existence of personality traits in animals and provided a foundation for similar assessment strategies in future studies of personality in animals. [8] Similarly, zebrafish have been used as a neurobehavioral model species for studying personality using the trait approach in non-human animals. These studies can then be translated to study personality development and personality disorders in humans. [32]
Another general example is the spider Anelosimus studiosus. This spider forms groups in which some females show an aggressive personality type and engage more in colony defense and prey capture, while others are docile and engage more in brood care. Groups containing both these two different personalities have better fitness than groups of only one personality type. This is because aggressive females are more efficient at foraging, web construction and defense, while docile females are better at raising the young. When groups contain a mix of both personalities, overall group performance is improved benefiting all group members. [33] In the social spider Stegodyphus dumicola individuals differ in their boldness, with bolder individuals having a greater risk appetite. Boldness changes were found to relate to social interactions with nest mates, indicating that individual personality is more plastic in groups. [34]
Many researchers are critical of the lack of consistency in the terminology surrounding animal personality. For example, temperament, behavioral syndrome, disposition and animal personality have been used interchangeably by some while others maintain that each term has a unique meaning. [3] Additionally, there is some concern that researchers may be misinterpreting the relationship between personality and behavioural plasticity. Behavioural ecologist Niels Dingemanse illustrates that one could erroneously determine that variation in behaviour between individuals exists if the subjects have not been examined across a gradient of ecological contexts, and urges fellow researchers to apply behavioural reaction norms whenever possible. [12] In addition, because of critical anthropomorphism, ethology, and comparative psychology are relatively unknown concepts to the layman, there are critics that question the validity in the claim that animals have a personality schema. There are those that fear that, while assessing behaviors, researchers will project anthropomorphic ratings onto the animals. [16]
The chimpanzee, also simply known as the chimp, is a species of great ape native to the forests and savannahs of tropical Africa. It has four confirmed subspecies and a fifth proposed one. When its close relative the bonobo was more commonly known as the pygmy chimpanzee, this species was often called the common chimpanzee or the robust chimpanzee. The chimpanzee and the bonobo are the only species in the genus Pan. Evidence from fossils and DNA sequencing shows that Pan is a sister taxon to the human lineage and is thus humans' closest living relative. The chimpanzee is covered in coarse black hair, but has a bare face, fingers, toes, palms of the hands, and soles of the feet. It is larger and more robust than the bonobo, weighing 40–70 kg (88–154 lb) for males and 27–50 kg (60–110 lb) for females and standing 150 cm.
In genetics, the phenotype is the set of observable characteristics or traits of an organism. The term covers the organism's morphology, its developmental processes, its biochemical and physiological properties, its behavior, and the products of behavior. An organism's phenotype results from two basic factors: the expression of an organism's genetic code and the influence of environmental factors. Both factors may interact, further affecting the phenotype. When two or more clearly different phenotypes exist in the same population of a species, the species is called polymorphic. A well-documented example of polymorphism is Labrador Retriever coloring; while the coat color depends on many genes, it is clearly seen in the environment as yellow, black, and brown. Richard Dawkins in 1978 and then again in his 1982 book The Extended Phenotype suggested that one can regard bird nests and other built structures such as caddisfly larva cases and beaver dams as "extended phenotypes".
Aggression is a behavior aimed at opposing or attacking something or someone. Though often done with the intent to cause harm, it can be channeled into creative and practical outlets for some. It may occur either reactively or without provocation. In humans, aggression can be caused by various triggers. For example, built-up frustration due to blocked goals or perceived disrespect. Human aggression can be classified into direct and indirect aggression; whilst the former is characterized by physical or verbal behavior intended to cause harm to someone, the latter is characterized by behavior intended to harm the social relations of an individual or group.
The genus Pan consists of two extant species: the chimpanzee and the bonobo. Taxonomically, these two ape species are collectively termed panins. The two species were formerly collectively called "chimpanzees" or "chimps"; if bonobos were recognized as a separate group at all, they were referred to as "pygmy" or "gracile chimpanzees". Together with humans, gorillas, and orangutans they are part of the family Hominidae. Native to sub-Saharan Africa, chimpanzees and bonobos are currently both found in the Congo jungle, while only the chimpanzee is also found further north in West Africa. Both species are listed as endangered on the IUCN Red List of Threatened Species, and in 2017 the Convention on Migratory Species selected the chimpanzee for special protection.
In the zoological field of ethology, a dominance hierarchy is a type of social hierarchy that arises when members of animal social groups interact, creating a ranking system. A dominant higher-ranking individual is sometimes called an alpha, and a submissive lower-ranking individual is called a beta. Different types of interactions can result in dominance depending on the species, including ritualized displays of aggression or direct physical violence. In social living groups, members are likely to compete for access to limited resources and mating opportunities. Rather than fighting each time they meet, individuals of the same sex establish a relative rank, with higher-ranking individuals often gaining more access to resources and mates. Based on repetitive interactions, a social order is created that is subject to change each time a dominant animal is challenged by a subordinate one.
Fluctuating asymmetry (FA), is a form of biological asymmetry, along with anti-symmetry and direction asymmetry. Fluctuating asymmetry refers to small, random deviations away from perfect bilateral symmetry. This deviation from perfection is thought to reflect the genetic and environmental pressures experienced throughout development, with greater pressures resulting in higher levels of asymmetry. Examples of FA in the human body include unequal sizes (asymmetry) of bilateral features in the face and body, such as left and right eyes, ears, wrists, breasts, testicles, and thighs.
Richard Walter Wrangham is an English anthropologist and primatologist; he is Professor of Biological Anthropology at Harvard University. His research and writing have involved ape behavior, human evolution, violence, and cooking.
Phenotypic plasticity refers to some of the changes in an organism's behavior, morphology and physiology in response to a unique environment. Fundamental to the way in which organisms cope with environmental variation, phenotypic plasticity encompasses all types of environmentally induced changes that may or may not be permanent throughout an individual's lifespan.
Evolutionary developmental psychology (EDP) is a research paradigm that applies the basic principles of evolution by natural selection, to understand the development of human behavior and cognition. It involves the study of both the genetic and environmental mechanisms that underlie the development of social and cognitive competencies, as well as the epigenetic processes that adapt these competencies to local conditions.
Animal culture can be defined as the ability of non-human animals to learn and transmit behaviors through processes of social or cultural learning. Culture is increasingly seen as a process, involving the social transmittance of behavior among peers and between generations. It can involve the transmission of novel behaviors or regional variations that are independent of genetic or ecological factors.
Dog behavior is the internally coordinated responses of individuals or groups of domestic dogs to internal and external stimuli. It has been shaped by millennia of contact with humans and their lifestyles. As a result of this physical and social evolution, dogs have acquired the ability to understand and communicate with humans. Behavioral scientists have uncovered a wide range of social-cognitive abilities in domestic dogs.
Mate choice is one of the primary mechanisms under which evolution can occur. It is characterized by a "selective response by animals to particular stimuli" which can be observed as behavior. In other words, before an animal engages with a potential mate, they first evaluate various aspects of that mate which are indicative of quality—such as the resources or phenotypes they have—and evaluate whether or not those particular trait(s) are somehow beneficial to them. The evaluation will then incur a response of some sort.
Evolutionary physiology is the study of the biological evolution of physiological structures and processes; that is, the manner in which the functional characteristics of individuals in a population of organisms have responded to natural selection across multiple generations during the history of the population. It is a sub-discipline of both physiology and evolutionary biology. Practitioners in the field come from a variety of backgrounds, including physiology, evolutionary biology, ecology, and genetics.
Behavioural genetics, also referred to as behaviour genetics, is a field of scientific research that uses genetic methods to investigate the nature and origins of individual differences in behaviour. While the name "behavioural genetics" connotes a focus on genetic influences, the field broadly investigates the extent to which genetic and environmental factors influence individual differences, and the development of research designs that can remove the confounding of genes and environment. Behavioural genetics was founded as a scientific discipline by Francis Galton in the late 19th century, only to be discredited through association with eugenics movements before and during World War II. In the latter half of the 20th century, the field saw renewed prominence with research on inheritance of behaviour and mental illness in humans, as well as research on genetically informative model organisms through selective breeding and crosses. In the late 20th and early 21st centuries, technological advances in molecular genetics made it possible to measure and modify the genome directly. This led to major advances in model organism research and in human studies, leading to new scientific discoveries.
A behaviour mutation is a genetic mutation that alters genes that control the way in which an organism behaves, causing their behavioural patterns to change.
The biological basis of personality is a collection of brain systems and mechanisms that underlie human personality. Human neurobiology, especially as it relates to complex traits and behaviors, is not well understood, but research into the neuroanatomical and functional underpinnings of personality are an active field of research. Animal models of behavior, molecular biology, and brain imaging techniques have provided some insight into human personality, especially trait theories.
In behavioral ecology, a behavioral syndrome is a correlated suite of behavioral traits, often measured across multiple contexts. The suite of traits that are correlated at the population or species level is considered the behavioral syndrome, while the phenotype of the behavioral syndrome an individual shows is their behavioral type. For example, a population may show a behavioral syndrome that includes a positive correlation between foraging behavior and mating behavior. An individual may be more or less aggressive than another individual within this behavioral syndrome, and this aggressive or passive phenotype is that individual's behavioral type.
Behavioral plasticity refers to a change in an organism's behavior that results from exposure to stimuli, such as changing environmental conditions. Behavior can change more rapidly in response to changes in internal or external stimuli than is the case for most morphological traits and many physiological traits. As a result, when organisms are confronted by new conditions, behavioral changes often occur in advance of physiological or morphological changes. For instance, larval amphibians changed their antipredator behavior within an hour after a change in cues from predators, but morphological changes in body and tail shape in response to the same cues required a week to complete.
Domestication syndrome refers to two sets of phenotypic traits that are common to either domesticated plants or domesticated animals.
Pathogen avoidance, also referred to as, parasite avoidance or pathogen disgust, refers to the theory that the disgust response, in humans, is an adaptive system that guides behavior to avoid infection caused by parasites such as viruses, bacteria, fungi, protozoa, helminth worms, arthropods and social parasites. Pathogen avoidance is a psychological mechanism associated with the behavioral immune system. Pathogen avoidance has been discussed as one of the three domains of disgust which also include sexual and moral disgust.
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