Neuronal Tuning
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Neuronal tuning refers to the hypothesized property of brain cells by which they selectively represent a particular type of sensory, association, motor, or cognitive information. Some neuronal responses have been hypothesized to be optimally tuned to specific patterns through experience. Neuronal tuning can be strong and sharp, as observed in
primary visual cortex The visual cortex of the brain is the area of the cerebral cortex that processes visual information. It is located in the occipital lobe. Sensory input originating from the eyes travels through the lateral geniculate nucleus in the thalamus and ...
(area V1) (but see Carandini et al 2005 Matteo Carandini, Jonathan B. Demb, Valerio Mante, David J. Tolhurst, Yang Dan, Bruno A. Olshausen, Jack L. Gallant and Nicole C. Rust. Do we know what the early visual system does? ''Journal of Neuroscience'' 25:10577-10597.), or weak and broad, as observed in
neural ensemble A neuronal ensemble is a population of nervous system Cell (biology), cells (or Cell culture, cultured neurons) involved in a particular neural computation. Background The concept of neuronal ensemble dates back to the work of Charles Sherring ...
s. Single neurons are hypothesized to be simultaneously tuned to several modalities, such as visual, auditory, and olfactory. Neurons hypothesized to be tuned to different signals are often hypothesized to integrate information from the different sources. In computational models called
neural network A neural network is a network or neural circuit, circuit of biological neurons, or, in a modern sense, an artificial neural network, composed of artificial neurons or nodes. Thus, a neural network is either a biological neural network, made up ...
s, such integration is the major principle of operation. The best examples of neuronal tuning can be seen in the visual, auditory, olfactory, somatosensory, and memory systems, although due to the small number of stimuli tested the generality of neuronal tuning claims is still an open question.


Visually Tuned System

Accepted neuronal tuning models suggest that
neuron A neuron, neurone, or nerve cell is an membrane potential#Cell excitability, electrically excitable cell (biology), cell that communicates with other cells via specialized connections called synapses. The neuron is the main component of nervous ...
s respond to different degrees based on the similarity between the optimal stimulus of the neuron and the given stimulus. (Teller (1984), however, has challenged the "detector" view of neurons on logical grounds) The first major evidence of neuronal tuning in the visual system was provided by
Hubel Hubel, Hübel or Huebel is a German language topographic surname, denoting a person who lived near a hill (Middle High German ''hübel'' "hill") and may refer to: * Allison Hubel, American mechanical engineer and cryobiologist *David H. Hubel (1926 ...
and Wiesel in 1959.Hubel, D. H.; Wiesel, T. N. Receptive Fields of Single Neurones in the Cat's Striate Cortex. ''J. Physiol.'' 1959, 148:574-591. They discovered that oriented slits of light were the most effective (of a very small set tested) stimuli for striate cortex “ simple cell” neurons. Other neurons, “ complex cells," responded best to lines of a certain orientation moving in a specific direction. Overall, the V1 neurons were found to be selectively tuned to certain orientations, sizes, positions, and forms. Hubel and Wiesel won th
Nobel Prize in Physiology or Medicine
in 1981 for their discoveries concerning information processing in the visual system. (More recently, Carandini et al (2005) have pointed out that the distinction between "simple" and "complex" cells may not be a valid one, observing that "simple and complex cells may not form a dichotomy at all.") While these simple cells in V1 respond to oriented bars through small
receptive field The receptive field, or sensory space, is a delimited medium where some physiological stimuli can evoke a sensory neuronal response in specific organisms. Complexity of the receptive field ranges from the unidimensional chemical structure of odo ...
s, the optimal visual stimulus becomes increasing complex as one moves toward the anterior of the brain.Riesenhuber, Maximilian; Poggio, Tomaso. Neural mechanisms of object recognition. ''Current Opinion in Neurobiology'' 2002, 12:162-168. Neurons in area V4 are selectively tuned to different wavelengths, hues, and saturations of color.Kolb, B.; Whishaw, I. Q. Fundamentals of Human Neuropsychology (2003). New York, NY: Worth Publishers. The middle temporal or area V5 is specifically tuned to the speed and direction of moving stimuli. At the apex of the ventral stream called the inferotemporal cortex, neurons became tuned to complex stimuli, such as faces. The specific tuning of intermediate neurons in the
ventral stream The two-streams hypothesis is a model of the neural processing of vision as well as hearing. The hypothesis, given its initial characterisation in a paper by David Milner and Melvyn A. Goodale in 1992, argues that humans possess two distinct visua ...
is less clear, because the range of form variety that can be utilized for probing is nearly infinite.Sakai, K.; Naya, Y.; Miyashita, Y. Neuronal tuning and associative mechanisms in form representation. ''Learning and Memory'' 1994, 1:83-105. In the anterior part of the
ventral stream The two-streams hypothesis is a model of the neural processing of vision as well as hearing. The hypothesis, given its initial characterisation in a paper by David Milner and Melvyn A. Goodale in 1992, argues that humans possess two distinct visua ...
, various regions appear to be tuned selectively to identify body parts ( extrastriate body area), faces ( fusiform face area) (according to a recent paper by Adamson and Troiani (2018) regions of the fusiform face area respond equally to "food"), moving bodies ( posterior superior temporal sulcus), or even scenes ( parahippocampal place area). Neuronal tuning in these areas requires fine discrimination among complex patterns in each relevant category for object recognition. Recent findings suggest that this fine discrimination is a function of
expertise An expert is somebody who has a broad and deep understanding and competence in terms of knowledge, skill and experience through practice and education in a particular field. Informally, an expert is someone widely recognized as a reliable sou ...
and the individual level of categorization with stimuli. Specifically, work has been done by Gauthier ''et al'' (2001) to show fusiform face area (FFA) activation for birds in bird experts and cars in car experts when compared to the opposing stimuli. Gauthier ''et al'' (2002) also utilized a new class of objects called Greebles and trained people to recognize them at individual levels. After training, the
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was tuned to distinguish between this class of objects as well as faces.Gauthier, I.; Tarr, M. J. Unraveling mechanisms for expert object recognition: Bridging brain activity and behavior. ''Journal of Experimental Psychology: Human Perception and Performance'' 2002, 28(2):431-446. Curran ''et al'' (2002) similarly trained people in a less structured class of objects called "blobs" and showed
FFA FFA may refer to: Aviation and military * First Flight Airport, near Kitty Hawk, North Carolina, United States * Free-fire area in U.S. military parlance * Flug- und Fahrzeugwerke Altenrhein, a Swiss aircraft and railway vehicle manufacturer * ...
selective activation for them.Curran, I.; Tanaka, J.; Weiskopf, D. An electrophysiological comparison of visual categorization and recognition memory. ''Cognitive, Affective, & Behavioral Neuroscience'' 2002, 18(2):1-18. Overall, neurons can be tuned selectively discriminate between certain sets of stimuli that are experienced regularly in the world.


Tuning in Other Systems

Neuron A neuron, neurone, or nerve cell is an membrane potential#Cell excitability, electrically excitable cell (biology), cell that communicates with other cells via specialized connections called synapses. The neuron is the main component of nervous ...
s in other systems also become selectively tuned to stimuli. In the
auditory system The auditory system is the sensory system for the sense of hearing. It includes both the sensory organs (the ears) and the auditory parts of the sensory system. System overview The outer ear funnels sound vibrations to the eardrum, increasin ...
, different neurons may respond selectively to the frequency (pitch), amplitude (loudness), and/or complexity (uniqueness) of sounds. In the
olfactory system The olfactory system, or sense of smell, is the sensory system used for smelling (olfaction). Olfaction is one of the special senses, that have directly associated specific organs. Most mammals and reptiles have a main olfactory system and an ac ...
, neurons may be tuned to certain kinds of smells. In the
gustatory system The gustatory system or sense of taste is the sensory system that is partially responsible for the perception of taste (flavor). Taste is the perception produced or stimulated when a substance in the mouth reacts chemically with taste receptor ...
, different neurons may respond selectively to different components of food: sweet, sour, salty, and bitter. In the
somatosensory system In physiology, the somatosensory system is the network of neural structures in the brain and body that produce the perception of touch ( haptic perception), as well as temperature ( thermoception), body position ( proprioception), and pain. I ...
, neurons may be selectively tuned to different types of pressure, temperature, bodily position, and pain. This tuning in the
somatosensory system In physiology, the somatosensory system is the network of neural structures in the brain and body that produce the perception of touch ( haptic perception), as well as temperature ( thermoception), body position ( proprioception), and pain. I ...
also provides feedback to the
motor system The motor system is the set of central and peripheral structures in the nervous system that support motor functions, i.e. movement. Peripheral structures may include skeletal muscles and neural connections with muscle tissues. Central structur ...
so that it may selectively tune neurons to respond in specific ways to given stimuli. Finally, the encoding and storage of information in both short-term and long-term memory requires the tuning of neurons in complex ways such that information may be later retrieved.


References

{{reflist Neurophysiology