BIO254:ReceptiveField

Introduction

Receptive fields are regions of space where a stimulus causes a particular sensory neuron to respond. Receptive fields are known to exist in the somatosensory, visual and auditory system. Each piece of sensory information corresponds to a specific location in their respective receptive field. A stimulus that affects an area larger than its corresponding receptive field will also affect receptive fields in adjacent receptors. Dense population of receptors have small receptive fields and as a result, finer resolution of detail. The resolution of a sensory system is not uniform everywhere.

Receptors project to first order neurons, which then project to second order neurons that again project to higher order neurons. The receptive fields of neurons in each relay nucleus depend on the cells that converge on it. As a consequence, the receptive fields of higher order nucleus are larger and more complex. The receptive fields of higher-order sensory neurons in the somatosensory and visual system have both excitory and inhibitory regions.

Somatosensory System

The mechanoreceptors in the skin have receptive fields that overlap with each other. When a point on the skin is touched, the tactile stimulus generates spikes at the node of Ranvier which are conducted to reach the nerve terminals.

Schematic diagram of receptive fields of mechanoreceptors in somatosensory system. Each tactile stimulus projects to its corresponding receptive field. The receptive fields may overlap. Source: Moore, Nelson and Sur.

There are three relay sites between mechanoreceptors in the skin and the cerebral cortex. Cortical neuron responses are processed in the dorsal column nuclei, the thalamus, and the cortex. Cortical neurons are studied using extracellular recording.

Every point on the skin is represented in the cortex by cortical cells connected to the receptors that are activated when that point on the skin is touched. Cortical neurons are grouped by function, with receptive fields arranged in an orderly topographic sequence that forms a map of the body. Receptive fields in higher cortical regions are larger. In relay neurons, cortical receptive fields depend on divergent presynaptic connections and convergent postsynaptic connections.

Schematic diagram of pathway from whiskers to cortex of rodents. Whiskers are tactile receptors of rats. The cortex consists of barrels. Each barrel represents a distinct group of tactile receptors. Source:Mirabella, Battiston and Diamond.

When the skin is touched at two or more points simultaneously, regions of skin surrounding the excitatory region of a receptive field of a cortical neuron will suppress excitation to another stimulus because the area surrounding the excitatory region is inhibitory. This is also known as the inhibitory surround.

Visual System

Ganglion cells transmit information from the photoreceptors in the retina to higher order neurons using action potentials. Bipolar, horizontal and amacrine cells lie between the photoreceptors and the ganglion cells.

Schematic diagram of on-center and off-center ganglion cells. On-center ganglion cells are excited when stimulated by light on the center and inhibited when stimulated in the surround. Off-center ganglion cells are excited when stimulated by light in the surround and inhibited when stimulated on the center. Source: Hubel and Wiesel.

The receptive areas of ganglion cells are circular, and divided into two parts, the receptive field center and the surround. On-center ganglion cells are excited when light is at the center of the receptive field, and are inhibited when light is on the surround. Off-center ganglion cells are inhibited by light at the center of their receptive field, and are excited when light is on the surround.

Ganglion cells have two parallel pathways from on-center cells and off-center cells for processing information. The receptive fields of ganglion cells vary in size. Receptive fields are small in the foveal region of the retina and large at the periphery of the retina. The firing rate of a ganglion cell determines constrast in light between the center and surround.

Schematic diagram of simple cells. Receptive fields of simple cells have narrow elongated areas with excitatory and inhibitory regions. When the bar of light is vertical, the response of the cell to the stimulus is the strongest. When the bar is light is horizonal , the response of the cell to the stimulus is the weakest. Source: Worgotter and Holt

Ganglion cells have two functional classes, M and P. M cells have large receptive fields, respond to large objects, and have rapid changes in stimulus. P cells have small receptive fields, are in larger numbers, respond to specific wavelengths, and are involved in the perception of form and color.

In the primary visual cortex, the receptive fields of cells are different from those in the retina. Cells respond to stimuli with linear properties. The cells in the primary visual cortex are known as simple and complex cells. Simple cells have excitatory and inhibitory regions in their receptive fields, which are usually rectilinear. Complex cells have larger receptive fields than simple cells, with no clear on and off areas.

Photoreceptors in the retina are connected to bipolar cells that have connections to ganglion cells, which then project to geniculate cells. Geniculate cells project to simple cells that are connected to complex cells.

Auditory System

Receptors in the auditory system are spatially distributed according to the sound frequencies that they respond to. High frequencies are located at the base of the cochlea and low frequencies at the apex.

References

1. Hubel, D.H., Wiesel, T.N., "Receptive fields, binocular interaction and functional architecture in the cat's visual cortex" (Journal of Physiology, 160, 106-154)

2. Kandel, E.R, Schwartz, J.H, and Jessel, T.M., " Principles of Neural Science, Fourth Edition" (McGraw Hill, 2000)

3. Moore, C.I., Nelson, S.B., Sur, M. "Dynamics of Neuronal Processing in Rat Somatosensory Cortex" (Trends in Neurosciences 22, 1999, 513-520)

4. Mirabella, G., Battiston, S., and Diamond, M.E. "Integration of multiple-whisker inputs in rat somatosensory cortex" (Cerebral Cortex 11, 164-170)

5. Worgotter, F., and Holt, G. "Spatial-temporal mechanisms in receptive fields of visual cortical simple cells" (J. Neurophysiol, 1991, 494-510)

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