The high degree of development of the sense of temperature in mammals provides them with the capacity to use temperature information not only as a signal of the condition of the body but also as a sense useful for recognizing objects and exploring the environment. For example, comparative experiments show that the nocturnal owl monkey, Aotus nancymaae, has a highly developed, specialized neural pathway for thermal sensation near and inside its nose. This pathway probably has enormous survival value by enabling these animals to determine the temperature (or freshness) of scent markings on their arboreal trails in the darkness of their native rainforest habitat in Colombia. Cats have a similar but rudimentary thermoreceptive-specific pathway in their forebrains, and they can be trained to respond behaviorally to thermal stimulation (e.g., by pressing a bar or choosing a door to open). Such experiments reveal that cats are relatively incapable of discriminating warm and cold stimuli applied to the furred skin of the trunk or the legs. However, they are sensitive on their noses and paws, responding to temperature differences of several degrees. This response corresponds to the level of thermal sensitivity on the face of humans and also accords with neurophysiological evidence regarding the properties of peripheral thermoreceptors and central thermoreceptive neurons in cats. Damage to the thermoreceptive pathway at the level of the thalamus or cortex in cats transiently reduces their ability to respond behaviorally to thermal stimuli; in contrast, similar damage in humans causes thermanesthesia (inability to feel hot or cold). This observation indicates that the integrative (homeostatic) processing of thermoreceptive activity in the brainstem is sufficient to motivate a cat’s behaviour.

In humans thermosensory activity causes emotional (affective) experiences of thermal comfort and discomfort. Such emotions motivate behaviour, and this enhances survival since these behaviours help maintain an optimal core body temperature, which is the goal of the internal homeostatic process known as thermoregulation. Temperature sensations in humans provide a measure of the activity of warm and cold receptors in the skin; however, thermal comfort or discomfort reflects the general state of the thermoregulatory system, involving signals not only from thermoreceptors in the skin but also from the integrative centres in the brainstem and other regions. Thus, the same temperature at the skin can be experienced as comfortable or uncomfortable, depending on the thermal condition of the person’s whole body. For example, if one is overheated, cold is perceived as pleasant, but if the core body temperature is low, and one feels generally chilled, then the same cold stimulus is distinctly unpleasant.

The evolutionary role of thermoreception is to subserve the process of thermoregulation. Thermoregulatory responses, such as shivering or panting, can be initiated by local temperature changes in the spinal cord or hypothalamus, and physiological experiments using microelectrode recordings from neurons in these regions also indicate that these thermoreceptive elements are directly involved in thermoregulation. In contrast to reptiles and fish, which are cold-blooded (poikilothermic) and regulate their body temperature mainly behaviorally, mammals are warm-blooded (endothermic) and maintain a constant body temperature (homeothermic) using active neural, physiological, and behavioral processes. Signals from thermosensors in the hypothalamus, spinal cord, deep body tissues, and skin are integrated in multiple thermoregulatory centres located mainly in the mammalian brainstem and hypothalamus. The temperatures of the inner body (core) and the peripheral skin (shell) are integrated with other systemic information, such as the water and salt content of the body, the level of available energy stores, and cardiovascular and immune system function. Such information serves to activate internal physiological and behavioral mechanisms that maintain body temperature within the normal range of values. These internal mechanisms include regulating the relative blood flow to skin and deep tissues, the release of metabolic activators (such as cortisol), and thermogenesis by brown adipose tissue.

When signals from warm thermoreceptors prevail over signals from cold thermoreceptors, heat-loss mechanisms, such as sweating, panting, and widening of blood vessels (vasodilation) in the skin, act to reduce body temperature. Cool-seeking behaviours are motivated by emotions of thermal discomfort. When signals from cold receptors predominate, heat conservation and production mechanisms are initiated. Thus, muscles expend energy in shivering and through other metabolic reactions (nonshivering thermogenesis), cutaneous blood vessels narrow (vasoconstriction), hairs fluff out to enhance thermal insulation, and appropriate warm-seeking behaviours are stimulated. Intervening elements in the nervous system (e.g., in the medulla oblongata) have been identified that integrate thermoregulatory signals from the hypothalamus and provide output links to produce changes in vascular tone or in the activity of brown adipose tissue. All these autonomic, or involuntary, regulatory functions continue even without the involvement of the cerebral cortex; thus, they do not require consciousness and persist during sleep and, to a limited extent, during anesthesia.

Arthur D. Craig

sensory neuron, nerve cell that carries information about changes in external and internal environments to the central nervous system (CNS). Such neurons are part of the peripheral nervous system, which lies outside the brain and spinal cord. They collect information from so-called sensory receptors, which are located in specialized tissues of the ears, eyes, mouth, nose, skin, and internal organs. In general, sensory neurons are described as afferent (carrying information to the CNS), whereas motor neurons are described as efferent (carrying information away from the CNS).

Information from a sensory neuron is transmitted to the CNS in the form of an action potential, a brief reversal of electric polarization of the membrane of a neuron or a muscle cell. The information flows across a synapse, or junction between neurons. In a few cases, sensory neurons communicate directly with motor neurons via synapses, allowing for a very fast reflex response. In most cases, however, sensory neurons communicate with interneurons in the CNS before a response is sent back to the body.

Sensory neurons may be categorized as peripheral or visceral. Peripheral sensory neurons are activated by stimuli external to the body, such as light, touch, sound, scent, or taste. Visceral sensory neurons respond to stimuli that originate within the body, such as pain, blood pressure, hunger, or inflammation. The body’s response to visceral sensory information allows it to maintain homeostasis (the self-regulation of physical systems that are necessary for survival).

Like other types of neurons, each sensory neuron has a cell body and projections (dendrites and axons) that gather and transmit information. The cell bodies of sensory neurons are often clustered into ganglia, which are located outside the CNS. The specific shapes and sizes of sensory neurons vary according to their function. Many sensory neurons are pseudounipolar; that is, each has one projection from the cell body that branches into two axons—one axon projecting to the periphery of the body and the other toward the CNS. Other sensory neurons are bipolar, each having two projections departing the cell body—one gathering information and the other passing information to other cells. In addition, many sensory neurons are enclosed in myelin, a coating consisting primarily of fatty materials that increases the speed of signaling along the axon. The layer of myelin varies in thickness, and it may be absent altogether.

Sensory neurons can be affected by diseases and disorders, such that affected individuals lose access to information about their external or internal environment. For example, humans rely on three types of cones (the light-sensitive cells in the retina of the eye that function in the perception of colour) to sense the full range of colours. In certain forms of colour blindness, however, only one or two types of cones are functional, resulting in a reduction of sensory information about colour in affected individuals’ environment. Another example of sensory impairment is hearing loss caused by repeated exposure to extremely loud noise that damages sensory receptors in the inner ear. Damage to auditory sensory neurons or to the temporal lobes of the brain, which normally process auditory information, can also result in hearing loss.

When sensory neurons become nonfunctional, the brain may adapt through a process known as neuroplasticity. For example, individuals who are blind from an early age can learn to use biosonar, or echolocation, to sense objects (similarly to bats). In this case, echoes are detected by auditory receptors and sensory neurons, but they are processed in the occipital lobe of the brain, which normally integrates visual, rather than auditory, information. Thus, individuals who are blind but who learn to employ biosonar can use auditory information to create mental images of their surroundings.

The Editors of Encyclopaedia Britannica Karin Akre