A sense is a biological system used by an organism for sensation, the process of gathering information about the surroundings through the detection of stimuli. Although, in some cultures, five human senses were traditionally identified as such (namely sight, smell, touch, taste, and hearing), many more are now recognized.^[1] Senses used by non-human organisms are even greater in variety and number. During sensation, sense organs collect various stimuli (such as a sound or smell) for transduction, meaning transformation into a form that can be understood by the brain. Sensation and perception are fundamental to nearly every aspect of an organism's cognition, behavior and thought.

In organisms, a sensory organ consists of a group of interrelated sensory cells that respond to a specific type of physical stimulus. Via cranial and spinal nerves (nerves of the Central and Peripheral nervous systems that relay sensory information to and from the brain and body), the different types of sensory receptor cells (such as mechanoreceptors, photoreceptors, chemoreceptors, thermoreceptors) in sensory organs transduct sensory information from these organs towards the central nervous system, finally arriving at the sensory cortices in the brain, where sensory signals are processed and interpreted (perceived).

Sensory systems, or senses, are often divided into external (exteroception) and internal (interoception) sensory systems. Human external senses are based on the sensory organs of the eyes, ears, skin, nose, mouth and the vestibular system. Internal sensation detects stimuli from internal organs and tissues. Internal senses possessed by humans include spatial orientation, proprioception (body position) and nociception (pain). Further internal senses lead to signals such as hunger, thirst, suffocation, and nausea, or different involuntary behaviors, such as vomiting.^[2][3][4] Some animals are able to detect electrical and magnetic fields, air moisture, or polarized light, while others sense and perceive through alternative systems, such as echolocation. Sensory modalities or sub modalities are different ways sensory information is encoded or transduced. Multimodality integrates different senses into one unified perceptual experience. For example, information from one sense has the potential to influence how information from another is perceived.^[5] Sensation and perception are studied by a variety of related fields, most notably psychophysics, neurobiology, cognitive psychology, and cognitive science.

Definitions

Sensory organs

Sensory organs are organs that sense and transduce stimuli. Humans have various sensory organs (i.e. eyes, ears, skin, nose, and mouth) that correspond to a respective visual system (sense of vision), auditory system (sense of hearing), somatosensory system (sense of touch), olfactory system (sense of smell), and gustatory system (sense of taste). Those systems, in turn, contribute to vision, hearing, touch, smell, and the ability to taste.^[5][6] Internal sensation, or interoception, detects stimuli from internal organs and tissues. Many internal sensory and perceptual systems exist in humans, including the vestibular system (sense of balance) sensed by the inner ear and providing the perception of spatial orientation; proprioception (body position); and nociception (pain). Further internal chemoreception- and osmoreception-based sensory systems lead to various perceptions, such as hunger, thirst, suffocation, and nausea, or different involuntary behaviors, such as vomiting.^[2][3][4]

Nonhuman animals experience sensation and perception, with varying levels of similarity to and difference from humans and other animal species. For example, other mammals in general have a stronger sense of smell than humans. Some animal species lack one or more human sensory system analogues and some have sensory systems that are not found in humans, while others process and interpret the same sensory information in very different ways. For example, some animals are able to detect electrical fields^[7] and magnetic fields,^[8] air moisture,^[9] or polarized light.^[10] Others sense and perceive through alternative systems such as echolocation.^[11][12] Recent theory suggests that plants and artificial agents such as robots may be able to detect and interpret environmental information in an analogous manner to animals.^[13][14][15]

Sensory modalities

Sensory modality refers to the way that information is encoded, which is similar to the idea of transduction. The main sensory modalities can be described on the basis of how each is transduced. Listing all the different sensory modalities, which can number as many as 17, involves separating the major senses into more specific categories, or submodalities, of the larger sense. An individual sensory modality represents the sensation of a specific type of stimulus. For example, the general sensation and perception of touch, which is known as somatosensation, can be separated into light pressure, deep pressure, vibration, itch, pain, temperature, or hair movement, while the general sensation and perception of taste can be separated into submodalities of sweet, salty, sour, bitter, spicy, and umami, all of which are based on different chemicals binding to sensory neurons.^[16]

Receptors

Sensory receptors are the cells or structures that detect sensations. Stimuli in the environment activate specialized receptor cells in the peripheral nervous system. During transduction, physical stimulus is converted into action potential by receptors and transmitted towards the central nervous system for processing.^[17] Different types of stimuli are sensed by different types of receptor cells. Receptor cells can be classified into types on the basis of three different criteria: cell type, position, and function. Receptors can be classified structurally on the basis of cell type and their position in relation to stimuli they sense. Receptors can further be classified functionally on the basis of the transduction of stimuli, or how the mechanical stimulus, light, or chemical changed the cell membrane potential.^[16]

Structural receptor types

Location

One way to classify receptors is based on their location relative to the stimuli. An exteroceptor is a receptor that is located near a stimulus of the external environment, such as the somatosensory receptors that are located in the skin. An interoceptor is one that interprets stimuli from internal organs and tissues, such as the receptors that sense the increase in blood pressure in the aorta or carotid sinus.^[16]

Cell type

The cells that interpret information about the environment can be either (1) a neuron that has a free nerve ending, with dendrites embedded in tissue that would receive a sensation; (2) a neuron that has an encapsulated ending in which the sensory nerve endings are encapsulated in connective tissue that enhances their sensitivity; or (3) a specialized receptor cell, which has distinct structural components that interpret a specific type of stimulus. The pain and temperature receptors in the dermis of the skin are examples of neurons that have free nerve endings (1). Also located in the dermis of the skin are lamellated corpuscles, neurons with encapsulated nerve endings that respond to pressure and touch (2). The cells in the retina that respond to light stimuli are an example of a specialized receptor (3), a photoreceptor.^[16]

A transmembrane protein receptor is a protein in the cell membrane that mediates a physiological change in a neuron, most often through the opening of ion channels or changes in the cell signaling processes. Transmembrane receptors are activated by chemicals called ligands. For example, a molecule in food can serve as a ligand for taste receptors. Other transmembrane proteins, which are not accurately called receptors, are sensitive to mechanical or thermal changes. Physical changes in these proteins increase ion flow across the membrane, and can generate an action potential or a graded potential in the sensory neurons.^[16]

Functional receptor types

A third classification of receptors is by how the receptor transduces stimuli into membrane potential changes. Stimuli are of three general types. Some stimuli are ions and macromolecules that affect transmembrane receptor proteins when these chemicals diffuse across the cell membrane. Some stimuli are physical variations in the environment that affect receptor cell membrane potentials. Other stimuli include the electromagnetic radiation from visible light. For humans, the only electromagnetic energy that is perceived by our eyes is visible light. Some other organisms have receptors that humans lack, such as the heat sensors of snakes, the ultraviolet light sensors of bees, or magnetic receptors in migratory birds.^[16]

Receptor cells can be further categorized on the basis of the type of stimuli they transduce. The different types of functional receptor cell types are mechanoreceptors, photoreceptors, chemoreceptors (osmoreceptor), thermoreceptors, electroreceptors (in certain mammals and fish), and nociceptors. Physical stimuli, such as pressure and vibration, as well as the sensation of sound and body position (balance), are interpreted through a mechanoreceptor. Photoreceptors convert light (visible electromagnetic radiation) into signals. Chemical stimuli can be interpreted by a chemoreceptor that interprets chemical stimuli, such as an object's taste or smell, while osmoreceptors respond to a chemical solute concentrations of body fluids. Nociception (pain) interprets the presence of tissue damage, from sensory information from mechano-, chemo-, and thermoreceptors.^[18] Another physical stimulus that has its own type of receptor is temperature, which is sensed through a thermoreceptor that is either sensitive to temperatures above (heat) or below (cold) normal body temperature.^[16]

Thresholds

Absolute threshold

Each sense organ (eyes or nose, for instance) requires a minimal amount of stimulation in order to detect a stimulus. This minimum amount of stimulus is called the absolute threshold.^[5] The absolute threshold is defined as the minimum amount of stimulation necessary for the detection of a stimulus 50% of the time.^[6] Absolute threshold is measured by using a method called signal detection. This process involves presenting stimuli of varying intensities to a subject in order to determine the level at which the subject can reliably detect stimulation in a given sense.^[5]

Differential threshold

Differential threshold or just noticeable difference (JDS) is the smallest detectable difference between two stimuli, or the smallest difference in stimuli that can be judged to be different from each other.^[6] Weber's Law is an empirical law that states that the difference threshold is a constant fraction of the comparison stimulus.^[6] According to Weber's Law, bigger stimuli require larger differences to be noticed.^[5]

Magnitude estimation is a psychophysical method in which subjects assign perceived values of given stimuli. The relationship between stimulus intensity and perceptive intensity is described by Steven's power law.^[6]

Signal detection theory

Signal detection theory quantifies the experience of the subject to the presentation of a stimulus in the presence of noise. There is internal noise and there is external noise when it comes to signal detection. The internal noise originates from static in the nervous system. For example, an individual with closed eyes in a dark room still sees something—a blotchy pattern of grey with intermittent brighter flashes—this is internal noise. External noise is the result of noise in the environment that can interfere with the detection of the stimulus of interest. Noise is only a problem if the magnitude of the noise is large enough to interfere with signal collection. The nervous system calculates a criterion, or an internal threshold, for the detection of a signal in the presence of noise. If a signal is judged to be above the criterion, thus the signal is differentiated from the noise, the signal is sensed and perceived. Errors in signal detection can potentially lead to false positives and false negatives. The sensory criterion might be shifted based on the importance of the detecting the signal. Shifting of the criterion may influence the likelihood of false positives and false negatives.^[6]

Private perceptive experience

Subjective visual and auditory experiences appear to be similar across humans subjects. The same cannot be said about taste. For example, there is a molecule called propylthiouracil (PROP) that some humans experience as bitter, some as almost tasteless, while others experience it as somewhere between tasteless and bitter. There is a genetic basis for this difference between perception given the same sensory stimulus. This subjective difference in taste perception has implications for individuals' food preferences, and consequently, health.^[6]

Sensory adaptation

When a stimulus is constant and unchanging, perceptual sensory adaptation occurs. During that process, the subject becomes less sensitive to the stimulus.^[5]

Fourier analysis

Biological auditory (hearing), vestibular and spatial, and visual systems (vision) appear to break down real-world complex stimuli into sine wave components, through the mathematical process called Fourier analysis. Many neurons have a strong preference for certain sine frequency components in contrast to others. The way that simpler sounds and images are encoded during sensation can provide insight into how perception of real-world objects happens.^[6]

Sensory neuroscience and the biology of perception

Perception occurs when nerves that lead from the sensory organs (e.g. eye) to the brain are stimulated, even if that stimulation is unrelated to the target signal of the sensory organ. For example, in the case of the eye, it does not matter whether light or something else stimulates the optic nerve, that stimulation will results in visual perception, even if there was no visual stimulus to begin with. (To prove this point to yourself (and if you are a human), close your eyes (preferably in a dark room) and press gently on the outside corner of one eye through the eyelid. You will see a visual spot toward the inside of your visual field, near your nose.)^[6]

Sensory nervous system

All stimuli received by the receptors are transduced to an action potential, which is carried along one or more afferent neurons towards a specific area (cortex) of the brain. Just as different nerves are dedicated to sensory and motors tasks, different areas of the brain (cortices) are similarly dedicated to different sensory and perceptual tasks. More complex processing is accomplished across primary cortical regions that spread beyond the primary cortices. Every nerve, sensory or motor, has its own signal transmission speed. For example, nerves in the frog's legs have a 90 ft/s (99 km/h) signal transmission speed, while sensory nerves in humans, transmit sensory information at speeds between 165 ft/s (181 km/h) and 330 ft/s (362 km/h).^[6]

Multimodal perception

Perceptual experience is often multimodal. Multimodality integrates different senses into one unified perceptual experience. Information from one sense has the potential to influence how information from another is perceived.^[5] Multimodal perception is qualitatively different from unimodal perception. There has been a growing body of evidence since the mid-1990s on the neural correlates of multimodal perception.^[20]

Philosophy

The philosophy of perception is concerned with the nature of perceptual experience and the status of perceptual data, in particular how they relate to beliefs about, or knowledge of, the world. Historical inquiries into the underlying mechanisms of sensation and perception have led early researchers to subscribe to various philosophical interpretations of perception and the mind, including panpsychism, dualism, and materialism. The majority of modern scientists who study sensation and perception take on a materialistic view of the mind.^[6]

Human sensation

General

Absolute threshold

Some examples of human absolute thresholds for the nine to 21 external senses.^[21]