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 its disorders. It is a multidisciplinary science that combines physiology, anatomy, molecular biology, ...
field concerned with the hypothetical reconstruction of sensory and other stimuli from information that has already been encoded and represented in the
brain
The brain is an organ (biology), 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 ...
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 (American English), neurone (British English), or nerve cell, is an membrane potential#Cell excitability, excitable cell (biology), cell that fires electric signals called action potentials across a neural network (biology), neural net ...
. 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 (also known as a nerve impulse or "spike" when in a neuron) is a series of quick changes in voltage across a cell membrane. An action potential occurs when the membrane potential of a specific Cell (biology), cell rapidly ri ...
s. Therefore, the main goal of neural decoding is to characterize how the
electrical activity
Electricity is the set of physical phenomena associated with the presence and motion of matter possessing an electric charge. Electricity is related to magnetism, both being part of the phenomenon of electromagnetism, as described by Maxwel ...
of neurons elicit activity and responses in the brain.
This article specifically refers to neural decoding as it pertains to the
mammal
A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
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, ...
.
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 (also known as a nerve impulse or "spike" when in a neuron) is a series of quick changes in voltage across a cell membrane. An action potential occurs when the membrane potential of a specific Cell (biology), cell rapidly ri ...
s. These patterns 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 patterns. 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 a gas). In electrochemical cells, electrodes are essential parts that can consist of a varie ...
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 ...
, 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 a large-scale brain activity mapping or brain decoding project
[The Brain Decoding Project. http://braindecodingproject.org/] to construct 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 and the neuronal responses, and the relationship among the electrical activities of the neurons in t ...
. 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
A signal is both the process and the result of Signal transmission, transmission of data over some transmission media, media accomplished by embedding some variation. Signals are important in multiple subject fields including signal processin ...
, 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 repre ...
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 (physiology), stimulus and the neuronal responses, and the relationship among the Electrophysiology, e ...
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 resoluti ...
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 which is 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 radiatio ...
(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). Torsten Wiesel and David Hubel discovered these cells in the late 1950s.
Such cells are ...
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
Introduction
In neurophysiology, a ganglion cell is a cell found in a ganglion (a cluster of neurons in the peripheral nervous system). Depending on their location and function, ganglion cells can be categorized into several major groups:
* ...
in the retina, since this area has the benefits of being strictly
feedforward
Feedforward is the provision of context of what one wants to communicate prior to that communication. In purposeful activity, feedforward creates an expectation which the actor anticipates. When expected experience occurs, this provides confirmato ...
,
retinotopic
Retinotopy () is the mapping of visual input from the retina to neurons, particularly those neurons within the Two-streams hypothesis, visual stream. For clarity, 'retinotopy' can be replaced with 'retinal mapping', and 'retinotopic' with 'retin ...
, 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
Electric potential (also called the ''electric field potential'', potential drop, the electrostatic potential) is defined as electric potential energy per unit of electric charge. More precisely, electric potential is the amount of work (physic ...
fluctuates around its
resting potential
The relatively static membrane potential of quiescent cells is called the resting membrane potential (or resting voltage), as opposed to the specific dynamic electrochemical phenomena called action potential and graded membrane potential. The re ...
due to a constant influx and efflux of
sodium
Sodium is a chemical element; it has Symbol (chemistry), symbol Na (from Neo-Latin ) and atomic number 11. It is a soft, silvery-white, highly reactive metal. Sodium is an alkali metal, being in group 1 element, group 1 of the peri ...
and
potassium
Potassium is a chemical element; it has Symbol (chemistry), symbol K (from Neo-Latin ) and atomic number19. It is a silvery white metal that is soft enough to easily cut with a knife. Potassium metal reacts rapidly with atmospheric oxygen to ...
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 thalam ...
, the
retina
The retina (; or retinas) is the innermost, photosensitivity, light-sensitive layer of tissue (biology), tissue of the eye of most vertebrates and some Mollusca, molluscs. The optics of the eye create a focus (optics), focused two-dimensional ...
,
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, Anatomical ter ...
. 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 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 synaptic connections between neurons based on the relative timing of their action potentials (or spikes). It is a temporally sensitive form of synaptic p ...
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
, and the
probability
Probability is a branch of mathematics and statistics concerning events and numerical descriptions of how likely they are to occur. The probability of an event is a number between 0 and 1; the larger the probability, the more likely an e ...
of seeing a certain stimulus,