H1 neuron

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The H1 neuron is located in the visual cortex of true flies of the order Diptera and mediates motor responses to visual stimuli. H1 is sensitive to horizontal motion in the visual field and enables the fly to rapidly and accurately respond to optic flow with motor corrections to stabilize flight. [1] It is particularly responsive to horizontal forward motion associated with movement of the fly's own body during flight. [2] Damage to H1 impairs the fly's ability to counteract disturbances during flight, suggesting that it is a necessary component of the optomotor response. H1 is an ideal system for studying the neural basis of information processing due to its highly selective and predictable responses to stimuli. [3] Since the initial anatomical and physiological characterizations of H1 in 1976, study of the neuron has greatly benefited the understanding of neural coding in a wide range of organisms, especially the relationship between the neural code and behavior.

Contents

Anatomy

Fly brain Fly Brain Neural Anatomy.gif
Fly brain

Flies possess two H1 neurons, one in each hemisphere of the brain. H1 is a lobula plate tangential cell (LPTC) located in the lobula plate of the optic lobe, the final destination of visual information originating from photoreceptors of the eye. [4] The lobula plate forms the posterior part of the lobula complex where the LPTCs are located. The large process diameter of these neurons allowed them to be amongst the first visual neurons to be intracellularly recorded in the fly. H1 axons are about 5 μm diameter, 1200 μm long, and myelinated with a spike conduction velocity of 1 meter/second. [5]

Phylogeny

Neurons sensitive to motion during flight are not specific to flies, and have been found in numerous nondipterous insect groups including Odonata, Lepidoptera, and Hymenoptera. [6] As in flies, these neurons receive input from both eyes and are sensitive to optic flow rotations corresponding to movement of the flying insect's body, suggesting motion sensitive neurons are an essential component of optomotor responses throughout the insect kingdom.

Connectivity

Fly eyes are composed of many individual ommatidia that possess their own lenses and photoreceptors. [5] The dendritic arbor of the H1 neuron covers the anterior surface of the lobula plate, where it receives retinotopic input from interneurons of the medulla lobula. To respond to image motion, the H1 neuron sends action potentials of varying frequency to the contralateral lobula plate. [5]

Hardwiring

Unlike human brains that rely on experience-dependent neuronal plasticity, the brain of the fly is hardwired for particular tasks in the visual sensory system. The H1 neuron and related tangential neurons are suggested to be genetically determined, meaning that these neurons are unaffected by visual stimuli during early development. [7] Parts of the fly brain have neuroplasticity but the H1 and other tangential neurons are hardwired neuronal machinery. Genetic hardwiring is likely an adaptation strategy that allow the flies to navigate in flight soon after hatching, actions largely mediated by the H1 and related tangential neurons. [7]

Function

Flies are agile flyers and strongly depend on vision during flight. [8] For visual course control, flies optic flow field is analyzed by a set of ~60 motion-sensitive neurons, each present in the third visual neuropil of the left and right eyes. [9] A subset of these neurons is thought to be involved in using the optic flow to estimate the parameters of self-motion, such as yaw, roll, and sideward translation. [10] Other neurons are thought to be involved in analyzing the content of the visual scene itself, for example, to separate figure from ground using motion parallax. [11] [12] The H1 neuron is responsible for detecting horizontal motion across the entire visual field of the fly, allowing the fly to generate and guide stabilizing motor corrections mid-flight with respect to yaw. [2]

Exploring the Neural Code

Three characteristics of H1, reliability, specificity, and sensitivity, make it exceptionally well suited for testing proposed models of neural encoding.

Reliability

Visual information in optical systems is inhibited by the temporal and spatial attributes of the sensory input, and by the biophysical properties of the neuronal circuits. How neural circuits encode behaviorally relevant information is dependent on the computational capacity of the nervous system with relation to the ambient conditions the organisms normally operate in. [13] H1 neurons are proven to be very efficient encoders of information via their high resilience to stimulus noise from external sources. [14] The operational and encoding processes of sensory pathways are often negatively affected by both external noise (relating to the stimulus) and internal noise (imperfect physiological processes); however, the activity of H1 is unaffected by photon noise. Instead, neuronal noise intrinsic to the H1 neural architecture is the limiting factor for accurate responses to stimuli. This dramatically reduces the noise of H1 electrophysiological readings, and provides the reliability necessary for accurate study conclusions.

Specificity

H1 exhibits very specific and predictable responses to directional stimuli, characteristics that are greatly beneficial for exploring the neural code because they allow for confident correlations between neural activity and stimuli. H1 neurons are known as Horizontally Sensitive (HS) cell, meaning HS cells depolarize most strongly in response to horizontal stimuli, and hyperpolarize when the direction of motion is opposite. HS cells, and their counterpart Vertically Sensitive (VS) cells, respond to a fixed direction regardless of the color or contrast of the background or the stimulus, making these neuronal systems ideal for testing. H1 exhibits a response to the stimulation of a single ommatidium, and can discriminate between translational motion of 2-3˚ in the visual field. [5]

Sensitivity

The response amplitude of H1 decreases during high velocity flight, thus becoming more sensitive to changes in optic flow speed and image contrast, [15] and increasing the dynamic range over which H1 operates. Changes in H1 axonal membrane potential is proportional to the amplitude of optic image velocity. However, medullary interneurons that synapse with H1 exhibit periodic, local changes in membrane potential as optic image velocities increases. To rectify this discrepancy, the dendrites of H1 temporally integrate these local fluctuations, resulting in a linear relationship between H1 axon membrane potential and stimulus intensity. This adaptation allows flies to rapidly transition between stationary, hovering, and swift modes of flight.

Related Research Articles

Visual cortex Region of the brain that processes visual information

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 then reaches the visual cortex. The area of the visual cortex that receives the sensory input from the lateral geniculate nucleus is the primary visual cortex, also known as visual area 1 (V1), Brodmann area 17, or the striate cortex. The extrastriate areas consist of visual areas 2, 3, 4, and 5.

The development of the nervous system, or neural development, or neurodevelopment, refers to the processes that generate, shape, and reshape the nervous system of animals, from the earliest stages of embryonic development to adulthood. The field of neural development draws on both neuroscience and developmental biology to describe and provide insight into the cellular and molecular mechanisms by which complex nervous systems develop, from nematodes and fruit flies to mammals.

Interneuron Neurons that are not motor or sensory

Interneurons are neurons that connect two brain regions, i.e. not direct motor neurons or sensory neurons. Interneurons are the central nodes of neural circuits, enabling communication between sensory or motor neurons and the central nervous system (CNS). They play vital roles in reflexes, neuronal oscillations, and neurogenesis in the adult mammalian brain.

Periaqueductal gray Nucleus surrounding the cerebral aqueduct

The periaqueductal gray is a brain region that plays a critical role in autonomic function, motivated behavior and behavioural responses to threatening stimuli. PAG is also the primary control center for descending pain modulation. It has enkephalin-producing cells that suppress pain.

Motion perception

Motion perception is the process of inferring the speed and direction of elements in a scene based on visual, vestibular and proprioceptive inputs. Although this process appears straightforward to most observers, it has proven to be a difficult problem from a computational perspective, and difficult to explain in terms of neural processing.

A gamma wave or gamma Rhythm is a pattern of neural oscillation in humans with a frequency between 25 and 140 Hz, the 40-Hz point being of particular interest. Gamma rhythms are correlated with large scale brain network activity and cognitive phenomena such as working memory, attention, and perceptual grouping, and can be increased in amplitude via meditation or neurostimulation. Altered gamma activity has been observed in many mood and cognitive disorders such as Alzheimer's disease, epilepsy, and schizophrenia.

Neural adaptation or sensory adaptation is a gradual decrease over time in the responsiveness of the sensory system to a constant stimulus. It is usually experienced as a change in the stimulus. For example, if a hand is rested on a table, the table's surface is immediately felt against the skin. Subsequently, however, the sensation of the table surface against the skin gradually diminishes until it is virtually unnoticeable. The sensory neurons that initially respond are no longer stimulated to respond; this is an example of neural adaptation.

Neural oscillation Brainwaves, repetitive patterns of neural activity in the central nervous system

Neural oscillations, or brainwaves, are rhythmic or repetitive patterns of neural activity in the central nervous system. Neural tissue can generate oscillatory activity in many ways, driven either by mechanisms within individual neurons or by interactions between neurons. In individual neurons, oscillations can appear either as oscillations in membrane potential or as rhythmic patterns of action potentials, which then produce oscillatory activation of post-synaptic neurons. At the level of neural ensembles, synchronized activity of large numbers of neurons can give rise to macroscopic oscillations, which can be observed in an electroencephalogram. Oscillatory activity in groups of neurons generally arises from feedback connections between the neurons that result in the synchronization of their firing patterns. The interaction between neurons can give rise to oscillations at a different frequency than the firing frequency of individual neurons. A well-known example of macroscopic neural oscillations is alpha activity.

A neuronal ensemble is a population of nervous system cells involved in a particular neural computation.

Neuronal noise Random electric fluctuations in neurons

Neuronal noise or neural noise refers to the random intrinsic electrical fluctuations within neuronal networks. These fluctuations are not associated with encoding a response to internal or external stimuli and can be from one to two orders of magnitude. Most noise commonly occurs below a voltage-threshold that is needed for an action potential to occur, but sometimes it can be present in the form of an action potential; for example, stochastic oscillations in pacemaker neurons in suprachiasmatic nucleus are partially responsible for the organization of circadian rhythms.

Synaptic gating

Synaptic gating is the ability of neural circuits to gate inputs by either suppressing or facilitating specific synaptic activity. Selective inhibition of certain synapses has been studied thoroughly, and recent studies have supported the existence of permissively gated synaptic transmission. In general, synaptic gating involves a mechanism of central control over neuronal output. It includes a sort of gatekeeper neuron, which has the ability to influence transmission of information to selected targets independently of the parts of the synapse upon which it exerts its action.

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Optomotor response

The optomotor response is an innate orienting behavior evoked by whole-field visual motion and is common to fish and insects during locomotion, such as swimming, walking and flying. The optomotor response has algorithmic properties such that the direction of the whole-field coherent motion dictates the direction of the behavioral output, as such, leftward visual stimuli lead to turning left and rightward visual stimuli lead to turning right. For instance, when zebrafish larvae are presented with a sinusoidal black and white grating pattern, the larvae will turn and swim in the direction of the perceived motion.

Models of neural computation are attempts to elucidate, in an abstract and mathematical fashion, the core principles that underlie information processing in biological nervous systems, or functional components thereof. This article aims to provide an overview of the most definitive models of neuro-biological computation as well as the tools commonly used to construct and analyze them.

Feature detection is a process by which the nervous system sorts or filters complex natural stimuli in order to extract behaviorally relevant cues that have a high probability of being associated with important objects or organisms in their environment, as opposed to irrelevant background or noise.

Non-spiking neuron

Non-spiking neurons are neurons that are located in the central and peripheral nervous systems and function as intermediary relays for sensory-motor neurons. They do not exhibit the characteristic spiking behavior of action potential generating neurons.

An autapse is a chemical or electrical synapse from a neuron onto itself. It can also be described as a synapse formed by the axon of a neuron on its own dendrites, in vivo or in vitro.

Biological motion perception is the act of perceiving the fluid unique motion of a biological agent. The phenomenon was first documented by Swedish perceptual psychologist, Gunnar Johansson, in 1973. There are many brain areas involved in this process, some similar to those used to perceive faces. While humans complete this process with ease, from a computational neuroscience perspective there is still much to be learned as to how this complex perceptual problem is solved. One tool which many research studies in this area use is a display stimuli called a point light walker. Point light walkers are coordinated moving dots that simulate biological motion in which each dot represents specific joints of a human performing an action.

Laura Busse German neuroscientist

Laura Busse is a German neuroscientist and professor of Systemic Neuroscience within the Division of Neurobiology at the Ludwig Maximilian University of Munich. Busse's lab studies context-dependent visual processing in mouse models by performing large scale in vivo electrophysiological recordings in the thalamic and cortical circuits of awake and behaving mice.

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