Retinotopy (from Greek τόπος, place) is the mapping of visual input from 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 the ...

to neurons, particularly those neurons within the visual stream. For clarity, 'retinotopy' can be replaced with 'retinal mapping', and 'retinotopic' with 'retinally mapped'. Visual field maps (retinotopic maps) are found in many amphibian and mammalian species, though the specific size, number, and spatial arrangement of these maps can differ considerably. Sensory topographies can be found throughout the brain and are critical to the understanding of one's external environment. Moreover, the study of sensory topographies and retinotopy in particular has furthered our understanding of how neurons encode and organize sensory signals. Retinal mapping of the visual field is maintained through various points of the visual pathway including but not limited to the retina, the dorsal

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 ...

, the optic tectum, 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 ...

(V1), and higher visual areas (V2-V4). Retinotopic maps in cortical areas other than V1 are typically more complex, in the sense that adjacent points of the visual field are not always represented in adjacent regions of the same area. For example, in the second visual area ( V2), the map is divided along an imaginary horizontal line across the visual field, in such a way that the parts of the retina that respond to the upper half of the visual field are represented in cortical tissue that is separated from those parts that respond the lower half of the visual field. Even more complex maps exist in the third and fourth visual areas V3 and V4, and in the dorsomedial area (V6). In general, these complex maps are referred to as second-order representations of the visual field, as opposed to first-order (continuous) representations such as V1. Additional retinotopic regions include ventral occipital (VO-1, VO-2), lateral occipital (LO-1, LO-2), dorsal occipital (V3A, V3B), and posterior parietal cortex (IPS0, IPS1, IPS2, IPS3, IPS4).

History

The discovery of visual field maps in humans can be traced to neurological cases, arising from war injuries, described and analyzed independently by Tatsuji Inouye (a Japanese ophthalmologist) and Gordon Holmes (a British neurologist). They observed correlations between the position of the entry wound and visual field loss (see Fishman, 1997 for an historical review).

Development

Molecular Cues

The "

Chemoaffinity Hypothesis The Chemoaffinity hypothesis states that neurons make connections with their targets based on interactions with specific molecular markers and, therefore, that the initial wiring diagram of an organism is (indirectly) determined by its genotype. Th ...

" was established by Sperry et al in 1963 in which it is thought that molecular gradients in both presynaptic and postsynaptic partners within the optic tectum organize developing axons into a coarse retinotopic map. This was established after a series of seminal experiments in fish and amphibians showed that retinal ganglion axons were already retinotopically organized within the optic tract and if severed, would regenerate and project back to retinotopically appropriate locations. Later, it was identified that

receptor tyrosine kinase Receptor tyrosine kinases (RTKs) are the high- affinity cell surface receptors for many polypeptide growth factors, cytokines, and hormones. Of the 90 unique tyrosine kinase genes identified in the human genome, 58 encode receptor tyrosine kin ...

s family EphA and a related EphA binding molecule referred to as ephrin-A family are expressed in complementary gradients in both the retina and the tectum. More specifically in the mouse, Ephrin A5 is expressed along the rostral-caudal axis of the optic tectum whereas the EphB family is expressed along the medio-lateral axis. This bimodal expression suggests a mechanism for the graded mapping of the temporal-nasal axis and the dorsoventral axis of the retina.

Target Space

While molecular cues are thought to guide axons into a coarse retinotopic map, the resolution of this map is thought to be influenced by available target space on postsynaptic partners. In wild type mice, it is thought that competition of target space is important for ensuring continuous retinal mapping, and that if perturbed, this competition may lead to the expansion or compression of the map depending on the available space. If the available space is altered, such as lesioning or ablating half of the retina, the healthy axons will expand their arbors in the tectum to fill the space. Similarly, if part of the tectum is ablated, the retinal axons will compress the topography to fit within the available tectal space.

Neural Activity

While neural activity in the retina is not necessary for the development of retinotopy, it seems to be a critical component for the refinement and stabilization of connectivity. Dark reared animals (no external visual cues) develop a normal retinal map in the tectum with no marked changes in receptive field size or laminar organization. While these animals may not have received external visual cues during development, these experiments suggest that spontaneous activity in the retina may be sufficient for retinotopic organization. In the goldfish, no neural activity (no external visual cues, and no spontaneous activity) did not prevent the formation of the retinal map but the final organization showed signs of lower resolution refinement and more dynamic growth (less stable). Based on Hebbian mechanisms, the thought is that if neurons are sensitive to similar stimuli (similar area of the visual field, similar orientation or direction selectivity) they will likely fire together. This patterned firing will result in stronger connectivity within the retinotopic organization through NMDAR synapse stabilization mechanisms in the post synaptic cells.

Dynamic Growth

Another important factor in the development of retinotopy is the potential for structural plasticity even after neurons are morphologically mature. One interesting hypothesis is that axons and dendrites are continuously extending and retracting their axons and dendrites. Several factors alter this dynamic growth including the

, the presence of developed synapses, and neural activity. As the nervous system develops and more cells are added, this structural plasticity allows for axons to gradually refine their place within the retinotopy. This plasticity is not specific to retinal ganglion axons, rather it's been shown that dendritic arbors of tectal neurons and filopodial processes of

radial glial cell Radial glial cells, or radial glial progenitor cells (RGPs), are bipolar-shaped progenitor cells that are responsible for producing all of the neurons in the cerebral cortex. RGPs also produce certain lineages of glia, including astrocytes and ...

s are also highly dynamic.

Description

In many locations within the brain, adjacent

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 have

receptive fields The receptive field, or sensory space, is a delimited medium where some physiological stimuli can evoke a sensory neuronal response in specific organisms. Complexity of the receptive field ranges from the unidimensional chemical structure of od ...

that include slightly different, but overlapping portions of the

visual field The visual field is the "spatial array of visual sensations available to observation in introspectionist psychological experiments". Or simply, visual field can be defined as the entire area that can be seen when an eye is fixed straight at a poin ...

. The position of the center of these receptive fields forms an orderly sampling mosaic that covers a portion of the visual field. Because of this orderly arrangement, which emerges from the spatial specificity of connections between neurons in different parts of the visual system, cells in each structure can be seen as contributing to a map of the visual field (also called a retinotopic map, or a visuotopic map). Retinotopic maps are a particular case of

topographic Topography is the study of the forms and features of land surfaces. The topography of an area may refer to the land forms and features themselves, or a description or depiction in maps. Topography is a field of geoscience and planetary scie ...

organization. Many brain structures that are responsive to visual input, including much of 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 visual nuclei of the

brain stem 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 co ...

(such as the

superior colliculus In neuroanatomy, the superior colliculus () is a structure lying on the roof of the mammalian midbrain. In non-mammalian vertebrates, the homologous structure is known as the optic tectum, or optic lobe. The adjective form '' tectal'' is common ...

) and

thalamus The thalamus (from Greek θάλαμος, "chamber") is a large mass of gray matter located in the dorsal part of the diencephalon (a division of the forebrain). Nerve fibers project out of the thalamus to the cerebral cortex in all direct ...

(such as the

and the pulvinar), are organized into retinotopic maps, also called visual field maps. Areas of the

are sometimes defined by their retinotopic boundaries, using a criterion that states that each area should contain a complete map of the visual field. However, in practice the application of this criterion is in many cases difficult. Those visual areas of the brainstem and cortex that perform the first steps of processing the retinal image tend to be organized according to very precise retinotopic maps. The role of retinotopy in other areas, where neurons have large receptive fields, is still being investigated. Retinotopy mapping shapes the folding of the

cerebral cortex The cerebral cortex, also known as the cerebral mantle, is the outer layer of neural tissue of the cerebrum of the brain in humans and other mammals. The cerebral cortex mostly consists of the six-layered neocortex, with just 10% consisting o ...

. In both the V1 and V2 areas of

macaque The macaques () constitute a genus (''Macaca'') of gregarious Old World monkeys of the subfamily Cercopithecinae. The 23 species of macaques inhabit ranges throughout Asia, North Africa, and (in one instance) Gibraltar. Macaques are principall ...

s and

human Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, cultu ...

s the vertical meridian of their visual field tends to be represented on the cerebral cortex's convex

gyri In neuroanatomy, a gyrus (pl. gyri) is a ridge on the cerebral cortex. It is generally surrounded by one or more sulci (depressions or furrows; sg. ''sulcus''). Gyri and sulci create the folded appearance of the brain in humans and other ...

folds whereas the horizontal meridian tends to be represented in their concave

sulci Sulci or Sulki (in Greek , Steph. B., Ptol.; , Strabo; , Paus.), was one of the most considerable cities of ancient Sardinia, situated in the southwest corner of the island, on a small island, now called Isola di Sant'Antioco, which is, how ...

folds.

Methods

Retinotopy mapping in humans is done with

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). The subject inside the fMRI machine focuses on a point. Then the

is stimulated with a circular image or angled lines about focus point.> The radial map displays the distance from the center of vision. The angular map shows angular location using rays angled about the center of vision. Combining the radial and angular maps, you can see the separate regions of the

and the smaller maps in each region.

References

{{reflist Visual system