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Behavioural responses to stress are evoked from underlying complex physiological changes that arise consequently from stress. [1]
Real or perceived threat in the environment elicits stress response in animals, which disrupts internal homeostasis. [2] Physiological changes cause behavioural responses in animals, including: impairment of response inhibition and lack of motivation, [3] as well as changes in social, sexual, [4] [5] aggression [6] and nurture [7] [8] behaviour in animals. [9] [10] The extent of the impact is dependent upon the type and duration of the stress, as well as the animal's past experiences. Behavioural responses to prolonged stress can also be transferred across generations. [11]
A stress, as defined to Walter Cannon (1871–1945), is any disturbance that imbalances the internal environment of an organism (i.e. their homeostasis). [2] There are two major types of stressors that cause stress to animals: abiotic stressors and biotic stressors. [12] Abiotic stressors are any ecological, geological, or climate changes that causes stress to the animal, such as increased temperatures and natural disasters. [13] Biotic stressors are living things related complications that causes stress, such as dominance, pollution, infection, social pressures, and competition. [10] [14] Animals can respond with physiological responses, [10] behavioural responses, psychological responses, [9] or physical responses (fight-or-flight). [2] Canon argued that there are two possible choices that an organism may choose when stress is encountered: fight responses or flight responses. [2]
Behavioural responses to stress are evoked from some underlying complex physiological changes that arise consequently from stress. [15]
According to a study conducted by Mika and his colleagues, prolonged stress in rats causes response inhibition. It was evident through their experiment that stressed rats had inhibited premature responses (decreased timing of intervals to food), along with decreased intrinsic motivation to initiate a response. [10] They link the decreased motivation to the stress-associated reduction in incentive motivation, as presented by another study conducted by Kleen and his colleagues. [3] Decreased motivation was also seen in a study conducted by Beery and Kaufer, where they explained that stressed rodents are less likely to be motivated to interact with one another. [9]
Beery and Kaufer noted that social withdrawal and general reduction in social interaction after an exposure to a stressor are evident in rodents. They argue that this is due to the underlying physiological changes that the rodent goes through in response to stress. For instance, the changes that occur to the hypothalamic-pituitary-adrenal (HPA) hormonal axis is directly related to the changes in social behaviour. Social avoidance is another consequence of stress that can be seen in rodents. Rodents are more likely to avoid dominant rats and avoid social interactions amongst each other after the exposure to a stressor. [9]
Sexual interests change in many species when exposed to stressors. For instance, stressed male and female rats express inhibited mating behaviour, which is evident through the clear increase in the inhibitory hormone RF-amide. [9] Another study suggests that masculine sexual behaviour in male rats is subject to changes in accordance to the type of stressors that the rats were subjected to. [9] The female zebra finch's mating choice is determined by the stressors that they are exposed to early in life, which remain consistent throughout adulthood. [4] A study about stress effects on female songbird’s response to sexual signal for mating indicated that the response to this specific signal can be impaired if the female is exposed to developmental stress. Behavioural changes as a result from developmental stress impairs neural responses to sexual signals, which reduces mating. [5]
Stressed animals would choose to avoid a novel situation rather than confront it. Aggressive behaviour is associated with sex hormones, such as testosterone, and specific brain regions and systems, such as the medial preoptic nucleus, prefrontal cortex-dependent response inhibition, and anterior hypothalamus. Stress negatively impacts sex hormones, which results in an imbalance and reduction in aggression related hormones and function. Also, chronic stress results in prefrontal cortex-dependent response inhibition. This results in reduction in aggression, thus promoting anti-predator responses. [16] [10]
Prolonged stress alters parental behaviour toward offspring and promotes parental neglect. According to a study performed by Tilgar and associates, predation stress alters parents’ behaviours, such as the reduction in provisioning rates, which negatively impacts the offspring’s performance. [8] The hormones oxytocin and vasopressin are generally responsible for affiliative and pair-bonding behaviours in many species. Stress alters the level of both hormones, resulting in an abnormal behaviour from parents towards offspring. For instance, levels of oxytocin decrease as a result of prolonged stress, which has been shown to reduce pair-bonding behaviour and increase withdrawal behaviour. Prolactin is another important hormone that is associated with nurture by parents to offspring, and levels of this hormone can be altered as a result of stress. Reduced levels of prolactin as a result of stress decreases behaviours such as suckling, licking, and brooding. [16]
Behavioural response to stress can be transmitted from parents to offspring. When an offspring observes the expression of behaviour such as fear in response to a stress stimulus, the same response would be elicited in the offspring when the stimulus is presented. Debeik and associates illustrated in rodents that fear is transferred from mother to infant in response to a certain odour stimulus. They further proved that the neural activity, such as the basal amygdala activity of the offspring’s brain, illustrated fear response. [11]
Fear is an intensely unpleasant emotion in response to perceiving or recognizing a danger or threat. Fear causes physiological changes that may produce behavioral reactions such as mounting an aggressive response or fleeing the threat. Fear in human beings may occur in response to a certain stimulus occurring in the present, or in anticipation or expectation of a future threat perceived as a risk to oneself. The fear response arises from the perception of danger leading to confrontation with or escape from/avoiding the threat, which in extreme cases of fear can be a freeze response or paralysis.
The hypothalamic–pituitary–adrenal axis is a complex set of direct influences and feedback interactions among three components: the hypothalamus, the pituitary gland, and the adrenal glands. These organs and their interactions constitute the HPA axis.
The Coolidge effect is a biological phenomenon seen in animals, whereby males exhibit renewed sexual interest whenever a new female is introduced, even after sex with prior but still available sexual partners. To a lesser extent, the effect is also seen among females with regard to their mates.
In biology, 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 the submissive lower-ranking individual 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, relative rank is established between individuals of the same sex, 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.
An apical dendrite is a dendrite that emerges from the apex of a pyramidal cell. Apical dendrites are one of two primary categories of dendrites, and they distinguish the pyramidal cells from spiny stellate cells in the cortices. Pyramidal cells are found in the prefrontal cortex, the hippocampus, the entorhinal cortex, the olfactory cortex, and other areas. Dendrite arbors formed by apical dendrites are the means by which synaptic inputs into a cell are integrated. The apical dendrites in these regions contribute significantly to memory, learning, and sensory associations by modulating the excitatory and inhibitory signals received by the pyramidal cells.
The stria terminalis is a structure in the brain consisting of a band of fibers running along the lateral margin of the ventricular surface of the thalamus. Serving as a major output pathway of the amygdala, the stria terminalis runs from its centromedial division to the ventromedial nucleus of the hypothalamus.
Reproductive suppression is the prevention or inhibition of reproduction in otherwise healthy adult individuals. It includes delayed sexual maturation (puberty) or inhibition of sexual receptivity, facultatively increased interbirth interval through delayed or inhibited ovulation or spontaneous or induced abortion, abandonment of immature and dependent offspring, mate guarding, selective destruction and worker policing of eggs in some eusocial insects or cooperatively breeding birds, and infanticide, and infanticide in carnivores) of the offspring of subordinate females either by directly killing by dominant females or males in mammals or indirectly through the withholding of assistance with infant care in marmosets and some carnivores. The Reproductive Suppression Model argues that "females can optimize their lifetime reproductive success by suppressing reproduction when future conditions for the survival of offspring are likely to be greatly improved over present ones”. When intragroup competition is high it may be beneficial to suppress the reproduction of others, and for subordinate females to suppress their own reproduction until a later time when social competition is reduced. This leads to reproductive skew within a social group, with some individuals having more offspring than others. The cost of reproductive suppression to the individual is lowest at the earliest stages of a reproductive event and reproductive suppression is often easiest to induce at the pre-ovulatory or earliest stages of pregnancy in mammals, and greatest after a birth. Therefore, neuroendocrine cues for assessing reproductive success should evolve to be reliable at early stages in the ovulatory cycle. Reproductive suppression occurs in its most extreme form in eusocial insects such as termites, hornets and bees and the mammalian naked mole rat which depend on a complex division of labor within the group for survival and in which specific genes, epigenetics and other factors are known to determine whether individuals will permanently be unable to breed or able to reach reproductive maturity under particular social conditions, and cooperatively breeding fish, birds and mammals in which a breeding pair depends on helpers whose reproduction is suppressed for the survival of their own offspring. In eusocial and cooperatively breeding animals most non-reproducing helpers engage in kin selection, enhancing their own inclusive fitness by ensuring the survival of offspring they are closely related to. Wolf packs suppress subordinate breeding.
The reward system is a group of neural structures responsible for incentive salience, associative learning, and positively-valenced emotions, particularly ones involving pleasure as a core component. Reward is the attractive and motivational property of a stimulus that induces appetitive behavior, also known as approach behavior, and consummatory behavior. A rewarding stimulus has been described as "any stimulus, object, event, activity, or situation that has the potential to make us approach and consume it is by definition a reward". In operant conditioning, rewarding stimuli function as positive reinforcers; however, the converse statement also holds true: positive reinforcers are rewarding.
In evolutionary psychology, people often speak of the four Fs which are said to be the four basic and most primal drives that animals are evolutionarily adapted to have, follow, and achieve: fighting, fleeing, feeding and fornicating.
Nesting behavior refers to an instinct or urge in pregnant animals associated with an increase of estradiol (E2) to prepare a home for the upcoming newborn(s). Nest building provides protection against predators and competitors that mean to exploit or kill infants. It also aids in family structure and is therefore influenced by different mating behaviours and social settings. It is found in a variety of animals such as birds, fish, squirrels, mice and pigs, and is said to be done by humans as well. However, recent research into nesting tendencies in women has questioned whether there is really any biological basis for nesting in humans, positing a social explanation instead.
Infanticide is the termination of a neonate after it has been born, and in zoology this is often the termination or consumption of newborn animals by either a parent or an unrelated adult. In rodents, it is not uncommon for the mother to commit infanticide shortly after parturition under conditions of extreme stress, or for an unrelated male to kill neonates.
Spontaneous Alternation Behavior (SAB) describes the tendency to alternate in their pursuit of different stimuli in consecutive trials despite a lack of training or reinforcement. The Behavior emerged from experiments using animals, mainly rodents, who naturally demonstrated the behavioral pattern when placed in previously unexplored maze shapes.
The challenge hypothesis outlines the dynamic relationship between testosterone and aggression in mating contexts. It proposes that testosterone promotes aggression when it would be beneficial for reproduction, such as mate guarding, or strategies designed to prevent the encroachment of intrasexual rivals. The positive correlation between reproductive aggression and testosterone levels is seen to be strongest during times of social instability. The challenge hypothesis predicts that seasonal patterns in testosterone levels are a function of mating system, paternal care, and male-male aggression in seasonal breeders.
Research into the mental disorder of schizophrenia, involves multiple animal models as a tool, including in the preclinical stage of drug development.
Sexual arousal describes the physiological and psychological responses in preparation for sexual intercourse or when exposed to sexual stimuli. A number of physiological responses occur in the body and mind as preparation for sexual intercourse, and continue during intercourse. Male arousal will lead to an erection, and in female arousal the body's response is engorged sexual tissues such as nipples, vulva, clitoris, vaginal walls, and vaginal lubrication. Mental stimuli and physical stimuli such as touch, and the internal fluctuation of hormones, can influence sexual arousal.
Sexual motivation is influenced by hormones such as testosterone, estrogen, progesterone, oxytocin, and vasopressin. In most mammalian species, sex hormones control the ability and motivation to engage in sexual behaviours.
Parental experience, as well as changing hormone levels during pregnancy and postpartum, cause changes in the parental brain. Displaying maternal sensitivity towards infant cues, processing those cues and being motivated to engage socially with her infant and attend to the infant's needs in any context could be described as mothering behavior and is regulated by many systems in the maternal brain. Research has shown that hormones such as oxytocin, prolactin, estradiol and progesterone are essential for the onset and the maintenance of maternal behavior in rats, and other mammals as well. Mothering behavior has also been classified within the basic drives. Less is known about the paternal brain, but changes in the father's brain occur alongside the mother once the offspring is born.
Social monogamy in mammals is defined as a long term or sequential living arrangement between an adult male and an adult female.
Endocrinology of parenting has been the subject of considerable study with focus both on human females and males and on females and males of other mammalian species. Parenting as an adaptive problem in mammals involves specific endocrine signals that were naturally selected to respond to infant cues and environmental inputs. Infants across species produce a number of cues to inform caregivers of their needs. These include visual cues, like facial characteristics, or in some species smiling, auditory cues, such as vocalizations, olfactory cues, and tactile stimulation. A commonly mentioned hormone in parenting is oxytocin, however many other hormones relay key information that results in variations in behavior. These include estrogen, progesterone, prolactin, cortisol, and testosterone. While hormones are not necessary for the expression of maternal behavior, they may influence it.
In zoology, copulation is animal sexual behavior in which sperm is introduced into a partner's body, especially directly into the reproductive tract. This is an aspect of mating. Many animals that live in water use external fertilization, whereas internal fertilization may have developed from a need to maintain gametes in a liquid medium in the Late Ordovician epoch. Internal fertilization with many vertebrates occurs via cloacal copulation, known as cloacal kiss, while mammals copulate vaginally, and many basal vertebrates reproduce sexually with external fertilization.