Difference between revisions of "W3 Cell"

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[[Image:w3_connections.png|thumb|right|320px|A W3 cell in connection with a few different types of [[Amacrine Cell|amacrine cells]].]]
 
[[Image:w3_connections.png|thumb|right|320px|A W3 cell in connection with a few different types of [[Amacrine Cell|amacrine cells]].]]
Until relatively recently, it was thought that the receptive fields of retinal ganglion cells were one of two center-surround receptive field types (either OFF-center, ON-surround or ON-center, OFF-surround). Within the last decade, however, it has become increasingly clear that the notion that only two types of receptive fields exist in photoreceptors is a gross oversimplification. Scientists now know that ganglion cells come in at least 15 or 20 types, each of which has a distinct shape and physiological function, and which correspondingly has connections with different types of cells in the rest of the retina. Further, each of these different ganglion cell types has a distinctive receptive field. Together, the receptive fields of all of the different cell types form an array that allows individuals to perceive all properties of the entire visual field. Specific cell types are distinct from one another in the sense that they respond to different visual properties than other cells: it is therefore the summation of input coming from all of the different types of retinal ganglion cells that allows an individual to completely visually perceive something . In order to develop a more comprehensive understanding of the retina, researchers are trying not only to identify the different retinal cell types (i.e. to explicitly classify the 15 or 20 types), but also to identify what image operation each type of retinal ganglion cell reports.
 
  
One example of a retinal ganglion cell type with specific visual response properties is the W3 retinal ganglion cell, which has been identified in the mouse retina. W3 is considered the equivalent to the object motion selective (OMS) ganglion cell in salamanders. Because relatively little work has been done specifically on W3 and a fair amount has been done regarding connections and properties of OMS, some of the information in the article will be about OMS.
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Retinal ganglion cells (RGCs) are the cells which are responsible for transferring information from the eye to the brain. These cells come with distinct functional signature, size, and morphology, and their dendritic arbors have been shown to be confined to specific sublayers of the inner plexiform layer (IPL) of the retina. However, despite this heterogeneity in structure, connectivity and function, to this point, many studies on the development of the visual system and its pathways have regarded all RGC's as being part of a single group. Studies are now emerging that show that different subclasses of RGC's are maximally responsive to particular visual features, and that they arborize selectively within particular layers of the IPL. The number of RGC subtypes and the extent to which they differ from one another remains to be seen, but based upon studies on dendritic morphology, it is estimated that, in mammals, there are approximately 20 subtypes of RCG's.  
  
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One example is the W3 retinal ganglion cell, which has been identified in the mouse retina. W3 is considered to be the equivalent to the object motion selective (OMS) ganglion cell in salamanders. Because relatively little work has been done specifically on W3 and a fair amount has been done regarding connections and properties of OMS, some of the information in the article will be about OMS. The W3 class is named after the transgenic line of mice that was used to mark it, and is referred to as such because there is not yet an accepted classification or scheme for the nomenclature of RGC subtypes.
  
 
== Physiology ==
 
== Physiology ==
  
OMS and W3 both respond sensitively to differential motion between the receptive field center and surround, as produced by an object moving over the background, but are strongly suppressed by global image motion, as produced by the observer’s head or eye movements.
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OMS and W3 both respond sensitively to differential motion between the receptive field center and surround, as produced by an object moving over the background. However, both are strongly suppressed by global image motion, as produced by the observer’s head or eye movements. While it is clear that W3 is responsive to moving stimuli, there is no evidence of preference for motion in a particular direction.  
  
 
== Anatomy ==
 
== Anatomy ==
W3 cells have small cell body and small arbors that are densely branched. Typically, the diameter of their dendritic field is about 120 microns. The arbors occupy a thick swath in the middle of the IPL from sublayer4 (SL4) through SL6 with minor sprouts arborization in SL1, when the IPL is divided into 10 imaginary strata. The region from SL4 to SL6 lies between the ON and OFF ChAT bands.
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W3 cells have small cell body and small arbors that are densely branched. Typically, the diameter of their somatic field is about 120 microns (as compared with other RGC types such as W7, which has an average dendritic field diameter of nearly 300 microns). If the IPL is divided into 10 arbitrary and evenly-spaced strata, dendritic arbors of W3 occupy a thick swath in the middle of the IPL (from sublayer4 (SL4) through SL6)) with minor arborization in SL1. The region that is most populated with W3 dendritic arbors (SL4 to SL6)f lies sandwiched between 2 ChAT (choline acetyltransferase) positive bands.
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The dendritic arbors of W3 have been shown to form in a step-wise manner at different times during the development of the mouse visual system. Initially, the arbors are completely restricted between sublayers 4 and 6. However, over the next few days in development, dendritic arbors expand to reach sublayers 1 and 2. Later still, the proximal processes between SL4 and SL6 expand, and the distal ones become isolated to SL1 and expand there as well. In adults, there is therefore a bistratified dendritic arbor distribution.  
  
 
=== Location ===
 
=== Location ===

Revision as of 19:43, 6 April 2012

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A W3 cell in connection with a few different types of amacrine cells.

Retinal ganglion cells (RGCs) are the cells which are responsible for transferring information from the eye to the brain. These cells come with distinct functional signature, size, and morphology, and their dendritic arbors have been shown to be confined to specific sublayers of the inner plexiform layer (IPL) of the retina. However, despite this heterogeneity in structure, connectivity and function, to this point, many studies on the development of the visual system and its pathways have regarded all RGC's as being part of a single group. Studies are now emerging that show that different subclasses of RGC's are maximally responsive to particular visual features, and that they arborize selectively within particular layers of the IPL. The number of RGC subtypes and the extent to which they differ from one another remains to be seen, but based upon studies on dendritic morphology, it is estimated that, in mammals, there are approximately 20 subtypes of RCG's.

One example is the W3 retinal ganglion cell, which has been identified in the mouse retina. W3 is considered to be the equivalent to the object motion selective (OMS) ganglion cell in salamanders. Because relatively little work has been done specifically on W3 and a fair amount has been done regarding connections and properties of OMS, some of the information in the article will be about OMS. The W3 class is named after the transgenic line of mice that was used to mark it, and is referred to as such because there is not yet an accepted classification or scheme for the nomenclature of RGC subtypes.

Physiology

OMS and W3 both respond sensitively to differential motion between the receptive field center and surround, as produced by an object moving over the background. However, both are strongly suppressed by global image motion, as produced by the observer’s head or eye movements. While it is clear that W3 is responsive to moving stimuli, there is no evidence of preference for motion in a particular direction.

Anatomy

W3 cells have small cell body and small arbors that are densely branched. Typically, the diameter of their somatic field is about 120 microns (as compared with other RGC types such as W7, which has an average dendritic field diameter of nearly 300 microns). If the IPL is divided into 10 arbitrary and evenly-spaced strata, dendritic arbors of W3 occupy a thick swath in the middle of the IPL (from sublayer4 (SL4) through SL6)) with minor arborization in SL1. The region that is most populated with W3 dendritic arbors (SL4 to SL6)f lies sandwiched between 2 ChAT (choline acetyltransferase) positive bands.

The dendritic arbors of W3 have been shown to form in a step-wise manner at different times during the development of the mouse visual system. Initially, the arbors are completely restricted between sublayers 4 and 6. However, over the next few days in development, dendritic arbors expand to reach sublayers 1 and 2. Later still, the proximal processes between SL4 and SL6 expand, and the distal ones become isolated to SL1 and expand there as well. In adults, there is therefore a bistratified dendritic arbor distribution.

Location

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Rough visualization of the stratification of W3 cell, occupying approximately 2.5/10 sublayers from the INL side.

Shape

Connections

Molecules

Until recently, few if any molecular markers were available to identify these RGC subtypes, so most analyses depended on nonselective labeling methods. Likewise, many developmental studies have treated RGCs as a single population. This limitation severely compromises analysis of RGC projections and development. For example, it is difficult to learn whether subtypes develop in distinct ways if they can be identified only after they have matured.This problem is now being circumvented in mice by generation of genetically engineered lines in which RGC subsets are marked with reporter genes (Hattar et al., 2002; Kim et al., 2008[1]; Yonehara et al., 2008; Badea et al., 2009; Huberman et al., 2009; Siegert et al., 2009). Here, we characterize the structure and function of RGCs marked in four transgenic lines, then use them to address a set of open questions about patterning and development of axonal and dendritic arbors:

History

Open Questions and Relevance to the EyeWire Project

References

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