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A neural circuit is a population of
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 interconnected by
synapse In the nervous system, a synapse is a structure that permits a neuron (or nerve cell) to pass an electrical or chemical signal to another neuron or to the target effector cell. Synapses are essential to the transmission of nervous impulses fr ...
s to carry out a specific function when activated. Neural circuits interconnect to one another to form large scale brain networks. Biological neural networks have inspired the design of artificial neural networks, but artificial neural networks are usually not strict copies of their biological counterparts.


Early study

Early treatments of neural networks can be found in Herbert Spencer's ''Principles of Psychology'', 3rd edition (1872), Theodor Meynert's ''
Psychiatry Psychiatry is the medical specialty devoted to the diagnosis, prevention, and treatment of mental disorders. These include various maladaptations related to mood, behaviour, cognition, and perceptions. See glossary of psychiatry. Initial p ...
'' (1884),
William James William James (January 11, 1842 – August 26, 1910) was an American philosopher, historian, and psychologist, and the first educator to offer a psychology course in the United States. James is considered to be a leading thinker of the la ...
' ''Principles of
Psychology Psychology is the scientific study of mind and behavior. Psychology includes the study of conscious and unconscious phenomena, including feelings and thoughts. It is an academic discipline of immense scope, crossing the boundaries betwe ...
'' (1890), and
Sigmund Freud Sigmund Freud ( , ; born Sigismund Schlomo Freud; 6 May 1856 – 23 September 1939) was an Austrian neurologist and the founder of psychoanalysis, a clinical method for evaluating and treating psychopathology, pathologies explained as originatin ...
's Project for a Scientific Psychology (composed 1895). The first rule of neuronal learning was described by
Hebb Hebb is a surname. Notable people with the surname include: * Bobby Hebb Robert Von Hebb (July 26, 1938 – August 3, 2010) was an American R&B and soul singer, musician, songwriter, recording artist, and performer known for his 1966 hit ent ...
in 1949, in the Hebbian theory. Thus, Hebbian pairing of pre-synaptic and post-synaptic activity can substantially alter the dynamic characteristics of the synaptic connection and therefore either facilitate or inhibit signal transmission. In 1959, the
neuroscientist A neuroscientist (or neurobiologist) is a scientist who has specialised knowledge in neuroscience, a branch of biology that deals with the physiology, biochemistry, psychology, anatomy and molecular biology of neurons, Biological neural network, n ...
s, Warren Sturgis McCulloch and Walter Pitts published the first works on the processing of neural networks. They showed theoretically that networks of artificial neurons could implement
logic Logic is the study of correct reasoning. It includes both formal and informal logic. Formal logic is the science of deductively valid inferences or of logical truths. It is a formal science investigating how conclusions follow from premis ...
al,
arithmetic Arithmetic () is an elementary part of mathematics that consists of the study of the properties of the traditional operations on numbers—addition, subtraction, multiplication, division, exponentiation, and extraction of roots. In the 19th c ...
, and
symbol A symbol is a mark, sign, or word that indicates, signifies, or is understood as representing an idea, object, or relationship. Symbols allow people to go beyond what is known or seen by creating linkages between otherwise very different conc ...
ic functions. Simplified models of biological neurons were set up, now usually called perceptrons or artificial neurons. These simple models accounted for neural summation (i.e., potentials at the post-synaptic membrane will summate in the cell body). Later models also provided for excitatory and inhibitory synaptic transmission.


Connections between neurons

The connections between neurons in the brain are much more complex than those of the artificial neurons used in the connectionist neural computing models of artificial neural networks. The basic kinds of connections between neurons are
synapse In the nervous system, a synapse is a structure that permits a neuron (or nerve cell) to pass an electrical or chemical signal to another neuron or to the target effector cell. Synapses are essential to the transmission of nervous impulses fr ...
s: both chemical and
electrical synapse Electricity is the set of physical phenomena associated with the presence and motion of matter that has a property of electric charge. Electricity is related to magnetism, both being part of the phenomenon of electromagnetism, as descri ...
s. The establishment of synapses enables the connection of neurons into millions of overlapping, and interlinking neural circuits. Presynaptic proteins called neurexins are central to this process. One principle by which neurons work is neural summationpotentials at the postsynaptic membrane will sum up in the cell body. If the depolarization of the neuron at the axon hillock goes above threshold an action potential will occur that travels down the
axon An axon (from Greek ἄξων ''áxōn'', axis), or nerve fiber (or nerve fibre: see spelling differences), is a long, slender projection of a nerve cell, or neuron, in vertebrates, that typically conducts electrical impulses known as action ...
to the terminal endings to transmit a signal to other neurons. Excitatory and inhibitory synaptic transmission is realized mostly by excitatory postsynaptic potentials (EPSPs), and inhibitory postsynaptic potentials (IPSPs). On the electrophysiological level, there are various phenomena which alter the response characteristics of individual synapses (called synaptic plasticity) and individual neurons (
intrinsic plasticity Nonsynaptic plasticity is a form of neuroplasticity that involves modification of ion channel function in the axon, dendrites, and cell body that results in specific changes in the integration of excitatory postsynaptic potentials and inhibitory ...
). These are often divided into short-term plasticity and long-term plasticity. Long-term synaptic plasticity is often contended to be the most likely
memory Memory is the faculty of the mind by which data or information is encoded, stored, and retrieved when needed. It is the retention of information over time for the purpose of influencing future action. If past events could not be remembered ...
substrate. Usually, the term "
neuroplasticity Neuroplasticity, also known as neural plasticity, or brain plasticity, is the ability of neural networks in the brain to change through growth and reorganization. It is when the brain is rewired to function in some way that differs from how it ...
" refers to changes in the brain that are caused by activity or experience. Connections display temporal and spatial characteristics. Temporal characteristics refers to the continuously modified activity-dependent efficacy of synaptic transmission, called spike-timing-dependent plasticity. It has been observed in several studies that the synaptic efficacy of this transmission can undergo short-term increase (called facilitation) or decrease ( depression) according to the activity of the presynaptic neuron. The induction of long-term changes in synaptic efficacy, by long-term potentiation (LTP) or depression (LTD), depends strongly on the relative timing of the onset of the excitatory postsynaptic potential and the postsynaptic action potential. LTP is induced by a series of action potentials which cause a variety of biochemical responses. Eventually, the reactions cause the expression of new receptors on the cellular membranes of the postsynaptic neurons or increase the efficacy of the existing receptors through phosphorylation. Backpropagating action potentials cannot occur because after an action potential travels down a given segment of the axon, the m gates on voltage-gated sodium channels close, thus blocking any transient opening of the h gate from causing a change in the intracellular sodium ion (Na+) concentration, and preventing the generation of an action potential back towards the cell body. In some cells, however, neural backpropagation does occur through the dendritic branching and may have important effects on synaptic plasticity and computation. A neuron in the brain requires a single signal to a neuromuscular junction to stimulate contraction of the postsynaptic muscle cell. In the spinal cord, however, at least 75
afferent Afferent may refer to: Anatomical structures Meaning "conveying towards a center": * Afferent arterioles, blood vessels that supply the nephrons * Afferent lymphatic vessels, lymph vessels that carry lymph to a lymph node * Afferent nerve fiber ...
neurons are required to produce firing. This picture is further complicated by variation in time constant between neurons, as some cells can experience their EPSPs over a wider period of time than others. While in synapses in the developing brain synaptic depression has been particularly widely observed it has been speculated that it changes to facilitation in adult brains.


Circuitry

An example of a neural circuit is the
trisynaptic circuit The trisynaptic circuit, or trisynaptic loop is a relay of Neurotransmission, synaptic transmission in the hippocampus. The circuit was initially described by the neuroanatomist Santiago Ramón y Cajal, Santiago Ramon y Cajal, in the early twentiet ...
in the
hippocampus The hippocampus (via Latin from Greek , ' seahorse') is a major component of the brain of humans and other vertebrates. Humans and other mammals have two hippocampi, one in each side of the brain. The hippocampus is part of the limbic system, ...
. Another is the Papez circuit linking the hypothalamus to the limbic lobe. There are several neural circuits in the cortico-basal ganglia-thalamo-cortical loop. These circuits carry information between the cortex,
basal ganglia The basal ganglia (BG), or basal nuclei, are a group of subcortical nuclei, of varied origin, in the brains of vertebrates. In humans, and some primates, there are some differences, mainly in the division of the globus pallidus into an extern ...
, thalamus, and back to the cortex. The largest structure within the basal ganglia, the striatum, is seen as having its own internal microcircuitry. Neural circuits in the
spinal cord The spinal cord is a long, thin, tubular structure made up of nervous tissue, which extends from the medulla oblongata in the brainstem to the lumbar region of the vertebral column (backbone). The backbone encloses the central canal of the spin ...
called central pattern generators are responsible for controlling motor instructions involved in rhythmic behaviours. Rhythmic behaviours include walking, urination, and
ejaculation Ejaculation is the discharge of semen (the ''ejaculate''; normally containing sperm) from the male reproductory tract as a result of an orgasm. It is the final stage and natural objective of male sexual stimulation, and an essential compone ...
. The central pattern generators are made up of different groups of spinal interneurons. There are four principal types of neural circuits that are responsible for a broad scope of neural functions. These circuits are a diverging circuit, a converging circuit, a reverberating circuit, and a parallel after-discharge circuit. In a diverging circuit, one neuron synapses with a number of postsynaptic cells. Each of these may synapse with many more making it possible for one neuron to stimulate up to thousands of cells. This is exemplified in the way that thousands of muscle fibers can be stimulated from the initial input from a single motor neuron. In a converging circuit, inputs from many sources are converged into one output, affecting just one neuron or a neuron pool. This type of circuit is exemplified in the respiratory center of the
brainstem The brainstem (or brain stem) is the posterior stalk-like part of the brain that connects the cerebrum with the spinal cord. In the human brain the brainstem is composed of the midbrain, the pons, and the medulla oblongata. The midbrain is ...
, which responds to a number of inputs from different sources by giving out an appropriate breathing pattern. A reverberating circuit produces a repetitive output. In a signalling procedure from one neuron to another in a linear sequence, one of the neurons may send a signal back to initiating neuron. Each time that the first neuron fires, the other neuron further down the sequence fire again sending it back to the source. This restimulates the first neuron and also allows the path of transmission to continue to its output. A resulting repetitive pattern is the outcome that only stops if one or more of the synapses fail, or if an inhibitory feed from another source causes it to stop. This type of reverberating circuit is found in the respiratory center that sends signals to the respiratory muscles, causing inhalation. When the circuit is interrupted by an inhibitory signal the muscles relax causing exhalation. This type of circuit may play a part in
epileptic seizure An epileptic seizure, informally known as a seizure, is a period of symptoms due to abnormally excessive or neural oscillation, synchronous neuronal activity in the brain. Outward effects vary from uncontrolled shaking movements involving much o ...
s. In a parallel after-discharge circuit, a neuron inputs to several chains of neurons. Each chain is made up of a different number of neurons but their signals converge onto one output neuron. Each synapse in the circuit acts to delay the signal by about 0.5 msec so that the more synapses there are will produce a longer delay to the output neuron. After the input has stopped, the output will go on firing for some time. This type of circuit does not have a feedback loop as does the reverberating circuit. Continued firing after the stimulus has stopped is called ''after-discharge''. This circuit type is found in the reflex arcs of certain reflexes.


Study methods

Different
neuroimaging Neuroimaging is the use of quantitative (computational) techniques to study the structure and function of the central nervous system, developed as an objective way of scientifically studying the healthy human brain in a non-invasive manner. Incr ...
techniques have been developed to investigate the activity of neural circuits and networks. The use of "brain scanners" or functional neuroimaging to investigate the structure or function of the brain is common, either as simply a way of better assessing brain injury with high-resolution pictures, or by examining the relative activations of different brain areas. Such technologies may include functional magnetic resonance imaging (fMRI),
brain positron emission tomography The brain is an organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It consists of nervous tissue and is typically located in the head ( cephalization), usually near organs for special se ...
(brain PET), and
computed axial tomography A computed tomography scan (CT scan; formerly called computed axial tomography scan or CAT scan) is a medical imaging technique used to obtain detailed internal images of the body. The personnel that perform CT scans are called radiographers ...
(CAT) scans. Functional neuroimaging uses specific brain imaging technologies to take scans from the brain, usually when a person is doing a particular task, in an attempt to understand how the activation of particular brain areas is related to the task. In functional neuroimaging, especially fMRI, which measures hemodynamic activity (using BOLD-contrast imaging) which is closely linked to neural activity, PET, and electroencephalography (EEG) is used. Connectionist models serve as a test platform for different hypotheses of representation, information processing, and signal transmission. Lesioning studies in such models, e.g. artificial neural networks, where parts of the nodes are deliberately destroyed to see how the network performs, can also yield important insights in the working of several cell assemblies. Similarly, simulations of dysfunctional neurotransmitters in neurological conditions (e.g., dopamine in the basal ganglia of Parkinson's patients) can yield insights into the underlying mechanisms for patterns of cognitive deficits observed in the particular patient group. Predictions from these models can be tested in patients or via pharmacological manipulations, and these studies can in turn be used to inform the models, making the process iterative. The modern balance between the connectionist approach and the single-cell approach in neurobiology has been achieved through a lengthy discussion. In 1972, Barlow announced the ''single neuron revolution'': "our perceptions are caused by the activity of a rather small number of neurons selected from a very large population of predominantly silent cells." This approach was stimulated by the idea of grandmother cell put forward two years earlier. Barlow formulated "five dogmas" of neuron doctrine. Recent studies of ' grandmother cell' and sparse coding phenomena develop and modify these ideas. The single cell experiments used intracranial electrodes in the medial temporal lobe (the hippocampus and surrounding cortex). Modern development of concentration of measure theory (stochastic separation theorems) with applications to artificial neural networks give mathematical background to unexpected effectiveness of small neural ensembles in high-dimensional brain.


Clinical significance

Sometimes neural circuitries can become pathological and cause problems such as in
Parkinson's disease Parkinson's disease (PD), or simply Parkinson's, is a long-term degenerative disorder of the central nervous system that mainly affects the motor system. The symptoms usually emerge slowly, and as the disease worsens, non-motor symptoms becom ...
when the
basal ganglia The basal ganglia (BG), or basal nuclei, are a group of subcortical nuclei, of varied origin, in the brains of vertebrates. In humans, and some primates, there are some differences, mainly in the division of the globus pallidus into an extern ...
are involved. Problems in the Papez circuit can also give rise to a number of neurodegenerative disorders including Parkinson's.


See also

*
Feedback Feedback occurs when outputs of a system are routed back as inputs as part of a chain of cause-and-effect that forms a circuit or loop. The system can then be said to ''feed back'' into itself. The notion of cause-and-effect has to be handled ...
*
List of regions in the human brain The human brain anatomical regions are ordered following standard neuroanatomy hierarchies. Functional, connective, and developmental regions are listed in parentheses where appropriate. Hindbrain (rhombencephalon) Myelencephalon *M ...
* Network science * Neural coding * Neural engineering * Neural oscillation *
Pulse-coupled networks Pulse-coupled networks or pulse-coupled neural networks (PCNNs) are neural models proposed by modeling a cat's visual cortex, and developed for high-performance biomimetic image processing. In 1989, Eckhorn introduced a neural model to emulate the ...
* Systems neuroscience


References


Further reading


Intrinsic plasticity
Robert H. Cudmore, Niraj S. Desai Scholarpedia 3(2):1363. doi:10.4249/scholarpedia.1363


External links


Comparison of Neural Networks in the Brain and Artificial Neural Networks

Lecture notes at MIT OpenCourseWare



Biological Neural Network Toolbox
- A free Matlab toolbox for simulating networks of several different types of neurons

- C. elegans, a nematode with 302 neurons, is the only organism for whom the entire neural network has been uncovered. Use this site to browse through the network and to search for paths between any 2 neurons.

''Neuroscience Online'' (electronic neuroscience textbook)

{{Authority control * Cognition