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A neural circuit is a population of
neuron A neuron, neurone, or nerve cell is an electrically excitable cell that communicates with other cells via specialized connections called synapses. The neuron is the main component of nervous tissue in all animals except sponges and placozoa ...
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 from ...
s to carry out a specific function when activated. Neural circuits interconnect to one another to form
large scale brain networks Large-scale brain networks (also known as intrinsic brain networks) are collections of widespread brain regions showing functional connectivity by statistical analysis of the fMRI BOLD signal or other recording methods such as EEG, PET and MEG. A ...
. Biological
neural network A neural network is a network or 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 of biological ...
s have inspired the design of
artificial neural network Artificial neural networks (ANNs), usually simply called neural networks (NNs) or neural nets, are computing systems inspired by the biological neural networks that constitute animal brains. An ANN is based on a collection of connected unit ...
s, 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 Herbert Spencer (27 April 1820 – 8 December 1903) was an English philosopher, psychologist, biologist, anthropologist, and sociologist famous for his hypothesis of social Darwinism. Spencer originated the expression " survival of the f ...
'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 lat ...
' ''Principles of
Psychology Psychology is the science, scientific study of mind and behavior. Psychology includes the study of consciousness, conscious and Unconscious mind, unconscious phenomena, including feelings and thoughts. It is an academic discipline of immens ...
'' (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 pathologies explained as originating in conflicts i ...
's Project for a Scientific Psychology (composed 1895). The first rule of neuronal learning was described by Hebb 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 telecommunications, transmission is the process of sending or propagating an analog or digital signal via a medium that is wired, wireless, or fiber-optic. Transmission technologies typically refer to physical layer protocol duties ...
. In 1959, the neuroscientists, 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 prem ...
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 ...
, 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 The soma (pl. ''somata'' or ''somas''), perikaryon (pl. ''perikarya''), neurocyton, or cell body is the bulbous, non-process portion of a neuron or other brain cell type, containing the cell nucleus. The word 'soma' comes from the Greek '' σῶμ ...
). 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 network Artificial neural networks (ANNs), usually simply called neural networks (NNs) or neural nets, are computing systems inspired by the biological neural networks that constitute animal brains. An ANN is based on a collection of connected unit ...
s. 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 from ...
s: both chemical and electrical synapses. The establishment of synapses enables the connection of neurons into millions of overlapping, and interlinking neural circuits. Presynaptic proteins called
neurexin Neurexins (NRXN) are a family of presynaptic cell adhesion proteins that have roles in connecting neurons at the synapse. They are located mostly on the presynaptic membrane and contain a single transmembrane domain. The extracellular domai ...
s are central to this process. One principle by which neurons work is neural summation
potentials Potential generally refers to a currently unrealized ability, in a wide variety of fields from physics to the social sciences. Mathematics and physics * Scalar potential, a scalar field whose gradient is a given vector field * Vector potentia ...
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 to the terminal endings to transmit a signal to other neurons. Excitatory and inhibitory synaptic transmission is realized mostly by
excitatory postsynaptic potentials In neuroscience, an excitatory postsynaptic potential (EPSP) is a postsynaptic potential that makes the postsynaptic neuron more likely to fire an action potential. This temporary depolarization of postsynaptic membrane potential, caused by the f ...
(EPSPs), and
inhibitory postsynaptic potentials An inhibitory postsynaptic potential (IPSP) is a kind of synaptic potential that makes a postsynaptic neuron less likely to generate an action potential.Purves et al. Neuroscience. 4th ed. Sunderland (MA): Sinauer Associates, Incorporated; 2008. I ...
(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). 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 remember ...
substrate. Usually, the term " neuroplasticity" 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 In chemistry, phosphorylation is the attachment of a phosphate group to a molecule or an ion. This process and its inverse, dephosphorylation, are common in biology and could be driven by natural selection. Text was copied from this source, wh ...
. 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 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 synaptic transmission in the hippocampus. The circuit was initially described by the neuroanatomist Santiago Ramon y Cajal, in the early twentieth century, using the Golgi staining method. ...
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 syste ...
. Another is the Papez circuit linking the
hypothalamus The hypothalamus () is a part of the brain that contains a number of small nuclei with a variety of functions. One of the most important functions is to link the nervous system to the endocrine system via the pituitary gland. 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 ext ...
, thalamus, and back to the cortex. The largest structure within the basal ganglia, the
striatum The striatum, or corpus striatum (also called the striate nucleus), is a nucleus (a cluster of neurons) in the subcortical basal ganglia of the forebrain. The striatum is a critical component of the motor and reward systems; receives gluta ...
, 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 sp ...
called
central pattern generator Central pattern generators (CPGs) are self-organizing biological neural circuits that produce rhythmic outputs in the absence of rhythmic input. They are the source of the tightly-coupled patterns of neural activity that drive rhythmic and stereo ...
s 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 compo ...
. 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 A motor neuron (or motoneuron or efferent neuron) is a neuron whose cell body is located in the motor cortex, brainstem or the spinal cord, and whose axon (fiber) projects to the spinal cord or outside of the spinal cord to directly or indirect ...
. 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 The respiratory center is located in the medulla oblongata and pons, in the brainstem. The respiratory center is made up of three major respiratory groups of neurons, two in the medulla and one in the pons. In the medulla they are the dorsal ...
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 seizures. 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
reflex In biology, a reflex, or reflex action, is an involuntary, unplanned sequence or action and nearly instantaneous response to a stimulus. Reflexes are found with varying levels of complexity in organisms with a nervous system. A reflex occurs ...
es.


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 Functional magnetic resonance imaging or functional MRI (fMRI) measures brain activity by detecting changes associated with blood flow. This technique relies on the fact that cerebral blood flow and neuronal activation are coupled. When an area o ...
(fMRI), brain positron emission tomography (brain PET), and computed axial tomography (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 Electroencephalography (EEG) is a method to record an electrogram of the spontaneous electrical activity of the brain. The biosignals detected by EEG have been shown to represent the postsynaptic potentials of pyramidal neurons in the neocorte ...
(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 network Artificial neural networks (ANNs), usually simply called neural networks (NNs) or neural nets, are computing systems inspired by the biological neural networks that constitute animal brains. An ANN is based on a collection of connected unit ...
s, 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 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 ...
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 Artificial neural networks (ANNs), usually simply called neural networks (NNs) or neural nets, are computing systems inspired by the biological neural networks that constitute animal brains. An ANN is based on a collection of connected units ...
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 beco ...
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 ext ...
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 Systems neuroscience is a subdiscipline of neuroscience and systems biology that studies the structure and function of neural circuits and systems. Systems neuroscience encompasses a number of areas of study concerned with how nerve cells behave ...


References


Further reading


Intrinsic plasticity
Robert H. Cudmore, Niraj S. Desai
Scholarpedia ''Scholarpedia'' is an English-language wiki-based online encyclopedia with features commonly associated with open-access online academic journals, which aims to have quality content in science and medicine. ''Scholarpedia'' articles are writ ...
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