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Neural decoding is a
neuroscience Neuroscience is the scientific study of the nervous system (the brain, spinal cord, and peripheral nervous system), its functions and disorders. It is a multidisciplinary science that combines physiology, anatomy, molecular biology, development ...
field concerned with the hypothetical reconstruction of sensory and other stimuli from information that has already been encoded and represented in the
brain A brain is an organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It is located in the head, usually close to the sensory organs for senses such as vision. It is the most complex organ in a v ...
by
networks Network, networking and networked may refer to: Science and technology * Network theory, the study of graphs as a representation of relations between discrete objects * Network science, an academic field that studies complex networks Mathematics ...
of
neurons 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. N ...
. Reconstruction refers to the ability of the researcher to predict what sensory stimuli the subject is receiving based purely on neuron
action potential An action potential occurs when the membrane potential of a specific cell location rapidly rises and falls. This depolarization then causes adjacent locations to similarly depolarize. Action potentials occur in several types of animal cells, ...
s. Therefore, the main goal of neural decoding is to characterize how the
electrical activity This is a list of electrical phenomena. Electrical phenomena are a somewhat arbitrary division of electromagnetic phenomena. Some examples are: * Biefeld–Brown effect — Thought by the person who coined the name, Thomas Townsend Brown, to ...
of neurons elicit activity and responses in the brain. This article specifically refers to neural decoding as it pertains to the
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
ian
neocortex The neocortex, also called the neopallium, isocortex, or the six-layered cortex, is a set of layers of the mammalian cerebral cortex involved in higher-order brain functions such as sensory perception, cognition, generation of motor commands, sp ...
.


Overview

When looking at a picture, people's brains are constantly making decisions about what object they are looking at, where they need to move their eyes next, and what they find to be the most salient aspects of the input stimulus. As these images hit the back of the retina, these stimuli are converted from varying wavelengths to a series of neural spikes called
action potential An action potential occurs when the membrane potential of a specific cell location rapidly rises and falls. This depolarization then causes adjacent locations to similarly depolarize. Action potentials occur in several types of animal cells, ...
s. These pattern of action potentials are different for different objects and different colors; we therefore say that the neurons are encoding objects and colors by varying their spike rates or temporal pattern. Now, if someone were to probe the brain by placing
electrode An electrode is an electrical conductor used to make contact with a nonmetallic part of a circuit (e.g. a semiconductor, an electrolyte, a vacuum or air). Electrodes are essential parts of batteries that can consist of a variety of materials de ...
s in the
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 ...
, they may find what appears to be random electrical activity. These neurons are actually firing in response to the lower level features of visual input, possibly the edges of a picture frame. This highlights the crux of the neural decoding hypothesis: that it is possible to reconstruct a stimulus from the response of the ensemble of neurons that represent it. In other words, it is possible to look at spike train data and say that the person or animal being recorded is looking at a red ball. With the recent breakthrough in large-scale neural recording and decoding technologies, researchers have begun to crack the neural code and already provided the first glimpse into the real-time neural code of memory traces as memory is formed and recalled in the hippocampus, a brain region known to be central for memory formation. Neuroscientists have initiated large-scale brain activity mapping or brain decoding projectThe Brain Decoding Project. http://braindecodingproject.org/ to construct the brain-wide neural codes.


Encoding to decoding

Implicit about the decoding hypothesis is the assumption that neural spiking in the brain somehow represents stimuli in the external world. The decoding of neural data would be impossible if the neurons were firing randomly: nothing would be represented. This process of decoding neural data forms a loop with
neural encoding Neural coding (or Neural representation) is a neuroscience field concerned with characterising the hypothetical relationship between the Stimulus (physiology), stimulus and the individual or Neuronal ensemble, ensemble neuronal responses and the re ...
. First, the organism must be able to perceive a set of stimuli in the world – say a picture of a hat. Seeing the stimuli must result in some internal learning: the encoding stage. After varying the range of stimuli that is presented to the observer, we expect the neurons to adapt to the statistical properties of the
signals In signal processing, a signal is a function that conveys information about a phenomenon. Any quantity that can vary over space or time can be used as a signal to share messages between observers. The ''IEEE Transactions on Signal Processing'' ...
, encoding those that occur most frequently:Barlow, H. (1961). Possible principles underlying the transformation of sensory messages. Sensory communication. the efficient-coding hypothesis. Now neural decoding is the process of taking these statistical consistencies, a
statistical model A statistical model is a mathematical model that embodies a set of statistical assumptions concerning the generation of Sample (statistics), sample data (and similar data from a larger Statistical population, population). A statistical model repres ...
of the world, and reproducing the stimuli. This may map to the process of thinking and acting, which in turn guide what stimuli we receive, and thus, completing the loop. In order to build a model of neural spike data, one must both understand how information is originally stored in the brain and how this information is used at a later point in time. This
neural coding Neural coding (or Neural representation) is a neuroscience field concerned with characterising the hypothetical relationship between the stimulus and the individual or ensemble neuronal responses and the relationship among the electrical activity o ...
and decoding loop is a symbiotic relationship and the crux of the brain's learning algorithm. Furthermore, the processes that underlie neural decoding and encoding are very tightly coupled and may lead to varying levels of representative ability.


Spatial resolutions

Much of the neural decoding problem depends on the
spatial resolution In physics and geosciences, the term spatial resolution refers to distance between independent measurements, or the physical dimension that represents a pixel of the image. While in some instruments, like cameras and telescopes, spatial resolutio ...
of the data being collected. The number of neurons needed to reconstruct the stimulus with reasonable accuracy depends on the means by which data is collected and the area being recorded. For example,
rods and cones A photoreceptor cell is a specialized type of neuroepithelial cell found in the retina that is capable of visual phototransduction. The great biological importance of photoreceptors is that they convert light (visible electromagnetic radiati ...
(which respond to colors of small visual areas) in the retina may require more recordings than
simple cell A simple cell in the visual cortex, primary visual cortex is a cell that responds primarily to oriented edges and gratings (bars of particular orientations). These cells were discovered by Torsten Wiesel and David Hubel in the late 1950s. Such ...
s (which respond to orientation of lines) in the primary visual cortex. Previous recording methods relied on stimulating single neurons over a repeated series of tests in order to generalize this neuron's behavior. New techniques such as high-density multi-electrode array recordings and multi-photon calcium imaging techniques now make it possible to record from upwards of a few hundred neurons. Even with better recording techniques, the focus of these recordings must be on an area of the brain that is both manageable and qualitatively understood. Many studies look at spike train data gathered from the
ganglion cells {{stack, A ganglion cell is a cell found in a ganglion. Examples of ganglion cells include: * Retinal ganglion cell (RGC) found in the ganglion cell layer of the retina * Cells that reside in the adrenal medulla, where they are involved in the s ...
in the retina, since this area has the benefits of being strictly feedforward,
retinotopic Retinotopy (from Greek τόπος, place) is the mapping of visual input from the retina to neurons, particularly those neurons within the visual stream. For clarity, 'retinotopy' can be replaced with 'retinal mapping', and 'retinotopic' with 'r ...
, and amenable to current recording granularities. The duration, intensity, and location of the stimulus can be controlled to sample, for example, a particular subset of ganglion cells within a structure of the visual system. Other studies use spike trains to evaluate the discriminatory ability of non-visual senses such as rat facial whiskers and the olfactory coding of moth pheromone receptor neurons. Even with ever-improving recording techniques, one will always run into the limited sampling problem: given a limited number of recording trials, it is impossible to completely account for the error associated with noisy data obtained from stochastically functioning neurons (for example, a neuron's
electric potential The electric potential (also called the ''electric field potential'', potential drop, the electrostatic potential) is defined as the amount of work energy needed to move a unit of electric charge from a reference point to the specific point in ...
fluctuates around its
resting potential A relatively static membrane potential which is usually referred to as the ground value for trans-membrane voltage. The relatively static membrane potential of quiescent cells is called the resting membrane potential (or resting voltage), as oppo ...
due to a constant influx and efflux of
sodium Sodium is a chemical element with the symbol Na (from Latin ''natrium'') and atomic number 11. It is a soft, silvery-white, highly reactive metal. Sodium is an alkali metal, being in group 1 of the periodic table. Its only stable iso ...
and
potassium Potassium is the chemical element with the symbol K (from Neo-Latin ''kalium'') and atomic number19. Potassium is a silvery-white metal that is soft enough to be cut with a knife with little force. Potassium metal reacts rapidly with atmosphe ...
ions). Therefore, it is not possible to perfectly reconstruct a stimulus from spike data. Luckily, even with noisy data, the stimulus can still be reconstructed within acceptable error bounds.


Temporal resolutions

Timescales and frequencies of stimuli being presented to the observer are also of importance to decoding the neural code. Quicker timescales and higher frequencies demand faster and more precise responses in neural spike data. In humans, millisecond precision has been observed throughout the
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 ...
, the
retina The retina (from la, rete "net") is the innermost, light-sensitive layer of tissue of the eye of most vertebrates and some molluscs. The optics of the eye create a focused two-dimensional image of the visual world on the retina, which then ...
, and the
lateral geniculate nucleus In neuroanatomy, the lateral geniculate nucleus (LGN; also called the lateral geniculate body or lateral geniculate complex) is a structure in the thalamus and a key component of the mammalian visual pathway. It is a small, ovoid, ventral projec ...
, so one would suspect this to be the appropriate measuring frequency. This has been confirmed in studies that quantify the responses of neurons in the
lateral geniculate nucleus In neuroanatomy, the lateral geniculate nucleus (LGN; also called the lateral geniculate body or lateral geniculate complex) is a structure in the thalamus and a key component of the mammalian visual pathway. It is a small, ovoid, ventral projec ...
to white-noise and naturalistic movie stimuli. At the cellular level,
spike-timing-dependent plasticity Spike-timing-dependent plasticity (STDP) is a biological process that adjusts the strength of connections between neurons in the brain. The process adjusts the connection strengths based on the relative timing of a particular neuron's output and in ...
operates at millisecond timescales; therefore, models seeking biological relevance should be able to perform at these temporal scales.


Probabilistic decoding

When decoding neural data, arrival times of each spike t_1,\textt_2,\text...,\textt_n\text=\text\, and the
probability Probability is the branch of mathematics concerning numerical descriptions of how likely an Event (probability theory), event is to occur, or how likely it is that a proposition is true. The probability of an event is a number between 0 and ...
of seeing a certain stimulus, P (t)/math> may be the extent of the available data. The
prior distribution In Bayesian statistical inference, a prior probability distribution, often simply called the prior, of an uncertain quantity is the probability distribution that would express one's beliefs about this quantity before some evidence is taken int ...
P (t)/math> defines an ensemble of signals, and represents the
likelihood The likelihood function (often simply called the likelihood) represents the probability of random variable realizations conditional on particular values of the statistical parameters. Thus, when evaluated on a given sample, the likelihood funct ...
of seeing a stimulus in the world based on previous experience. The spike times may also be drawn from a
distribution Distribution may refer to: Mathematics *Distribution (mathematics), generalized functions used to formulate solutions of partial differential equations * Probability distribution, the probability of a particular value or value range of a vari ...
P /math>; however, what we want to know is the
probability distribution In probability theory and statistics, a probability distribution is the mathematical function that gives the probabilities of occurrence of different possible outcomes for an experiment. It is a mathematical description of a random phenomenon i ...
over a set of stimuli given a series of spike trains P \/math>, which is called the response-conditional ensemble. What remains is the characterization of the neural code by translating stimuli into spikes, P s(t)/math>; the traditional approach to calculating this probability distribution has been to fix the stimulus and examine the responses of the neuron. Combining everything using
Bayes' Rule In probability theory and statistics, Bayes' theorem (alternatively Bayes' law or Bayes' rule), named after Thomas Bayes, describes the probability of an event, based on prior knowledge of conditions that might be related to the event. For examp ...
results in the simplified probabilistic characterization of neural decoding: P \ = P s(t) * (P (t)P ). An area of active research consists of finding better ways of representing and determining P s(t) /math>.Rieke, F. (1999). Spikes: exploring the neural code. exploring the neural code (p. 395). The MIT Press. The following are some such examples.


Spike train number

The simplest coding strategy is the spike train number coding. This method assumes that the spike number is the most important quantification of spike train data. In spike train number coding, each stimulus is represented by a unique firing rate across the sampled neurons. The color red may be signified by 5 total spikes across the entire set of neurons, while the color green may be 10 spikes; each spike is pooled together into an overall count. This is represented by: P(r, s) = \prod_ P(n_ , s) where r = n = the number of spikes, n_ is the number of spikes of neuron i at stimulus presentation time j, and s is the stimulus.


Instantaneous rate code

Adding a small temporal component results in the spike timing coding strategy. Here, the main quantity measured is the number of spikes that occur within a predefined
window A window is an opening in a wall, door, roof, or vehicle that allows the exchange of light and may also allow the passage of sound and sometimes air. Modern windows are usually glazed or covered in some other transparent or translucent materia ...
of time T. This method adds another dimension to the previous. This timing code is given by: P(/Sr.)=2 we index hr eg where t_ is the jth spike on the lth presentation of neuron i, v_i(t, s) is the firing rate of neuron i at time t, and 0 to T is the start to stop times of each trial.


Temporal correlation

Temporal correlation code, as the name states, adds
correlations In statistics, correlation or dependence is any statistical relationship, whether causal or not, between two random variables or bivariate data. Although in the broadest sense, "correlation" may indicate any type of association, in statistics ...
between individual spikes. This means that the time between a spike t_i and its preceding spike t_ is included. This is given by: P(r, s) = \prod_ \left s)dt \right exp \left s) \right where \tau(t) is the time interval between a neurons spike and the one preceding it.


Ising decoder

Another description of neural spike train data uses the
Ising model The Ising model () (or Lenz-Ising model or Ising-Lenz model), named after the physicists Ernst Ising and Wilhelm Lenz, is a mathematical model of ferromagnetism in statistical mechanics. The model consists of discrete variables that represent ...
borrowed from the physics of magnetic spins. Because neural spike trains are effectively binarized (either on or off) at small time scales (10 to 20 ms), the
Ising model The Ising model () (or Lenz-Ising model or Ising-Lenz model), named after the physicists Ernst Ising and Wilhelm Lenz, is a mathematical model of ferromagnetism in statistical mechanics. The model consists of discrete variables that represent ...
is able to effectively capture the present pairwise correlations, and is given by: P(r, s) = \frac exp \left ( \sum_ h_i(s)r_i + \frac \sum_ J_(s)r_ir_j \right ) where r = (r_1, r_2, . . . , r_n )^T is the set of binary responses of neuron i, h_i is the external fields function, J_ is the pairwise couplings function, and \Zeta(s) is the partition function


Agent-based decoding

In addition to the probabilistic approach,
agent-based model An agent-based model (ABM) is a computational model for simulating the actions and interactions of autonomous agents (both individual or collective entities such as organizations or groups) in order to understand the behavior of a system and wha ...
s exist that capture the spatial dynamics of the neural system under scrutiny. One such model is
hierarchical temporal memory Hierarchical temporal memory (HTM) is a biologically constrained machine intelligence technology developed by Numenta. Originally described in the 2004 book ''On Intelligence'' by Jeff Hawkins with Sandra Blakeslee, HTM is primarily used today for ...
, which is a
machine learning Machine learning (ML) is a field of inquiry devoted to understanding and building methods that 'learn', that is, methods that leverage data to improve performance on some set of tasks. It is seen as a part of artificial intelligence. Machine ...
framework that organizes visual perception problem into a
hierarchy A hierarchy (from Greek: , from , 'president of sacred rites') is an arrangement of items (objects, names, values, categories, etc.) that are represented as being "above", "below", or "at the same level as" one another. Hierarchy is an important ...
of interacting nodes (neurons). The connections between nodes on the same levels and a lower levels are termed
synapses 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 ...
, and their interactions are subsequently learning. Synapse strengths modulate learning and are altered based on the temporal and spatial firing of nodes in response to input patterns.Hawkins, J., Ahmad, S., & Dubinsky, D. (2006). Hierarchical temporal memory: Concepts, theory and terminology. Whitepaper.Hawkins, J., & Blakeslee, S. (2005). On intelligence. Owl Books. While it is possible to take the firing rates of these modeled neurons, and transform them into the probabilistic and mathematical frameworks described above, agent-based models provide the ability to observe the behavior of the entire population of modeled neurons. Researchers can circumvent the limitations implicit with lab-based recording techniques. Because this approach does rely on modeling biological systems, error arises in the assumptions made by the researcher and in the data used in
parameter estimation Estimation theory is a branch of statistics that deals with estimating the values of parameters based on measured empirical data that has a random component. The parameters describe an underlying physical setting in such a way that their value ...
.


Applicability

The advancement in our understanding of neural decoding benefits the development of brain-machine interfaces,
prosthetics In medicine, a prosthesis (plural: prostheses; from grc, πρόσθεσις, prósthesis, addition, application, attachment), or a prosthetic implant, is an artificial device that replaces a missing body part, which may be lost through trau ...
and the understanding of neurological disorders such as
epilepsy Epilepsy is a group of non-communicable neurological disorders characterized by recurrent epileptic seizures. Epileptic seizures can vary from brief and nearly undetectable periods to long periods of vigorous shaking due to abnormal electrical ...
.


See also

*
Brain-reading Brain-reading or thought identification uses the responses of multiple voxels in the brain evoked by stimulus then detected by fMRI in order to decode the original stimulus. Advances in research have made this possible by using human neuroimaging ...
* Bursting * Correlation coding *
Grandmother cell The grandmother cell, sometimes called the "Jennifer Aniston neuron", is a hypothetical neuron that represents a complex but specific concept or object. It activates when a person "sees, hears, or otherwise sensibly discriminates" a specific entit ...
* Independent-spike coding *
Multielectrode array Microelectrode arrays (MEAs) (also referred to as multielectrode arrays) are devices that contain multiple (tens to thousands) microelectrodes through which neural signals are obtained or delivered, essentially serving as neural interfaces that co ...
*
Nervous system network models Network of human nervous system comprises nodes (for example, neurons) that are connected by links (for example, synapses). The connectivity may be viewed anatomically, functionally, or electrophysiologically. These are presented in several Wikipedi ...
*
Neural coding Neural coding (or Neural representation) is a neuroscience field concerned with characterising the hypothetical relationship between the stimulus and the individual or ensemble neuronal responses and the relationship among the electrical activity o ...
*
Neural synchronization Neural oscillations, or brainwaves, are rhythmic or repetitive patterns of neural activity in the central nervous system. Neural tissue can generate oscillation, oscillatory activity in many ways, driven either by mechanisms within individual ne ...
*
NeuroElectroDynamics Neural coding (or Neural representation) is a neuroscience field concerned with characterising the hypothetical relationship between the stimulus and the individual or ensemble neuronal responses and the relationship among the electrical activity ...
*
Patch clamp The patch clamp technique is a laboratory technique in electrophysiology used to study ionic currents in individual isolated living cells, tissue sections, or patches of cell membrane. The technique is especially useful in the study of excitabl ...
* Phase-of-firing code * Population coding *
Rate coding Neural coding (or Neural representation) is a neuroscience field concerned with characterising the hypothetical relationship between the stimulus and the individual or ensemble neuronal responses and the relationship among the electrical activity ...
*
Sparse coding Neural coding (or Neural representation) is a neuroscience field concerned with characterising the hypothetical relationship between the stimulus and the individual or ensemble neuronal responses and the relationship among the electrical activit ...
*
Temporal coding Neural coding (or Neural representation) is a neuroscience field concerned with characterising the hypothetical relationship between the stimulus and the individual or ensemble neuronal responses and the relationship among the electrical activity o ...


References

{{Neuroscience Computational neuroscience Neural circuits