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{{Infobox neuron
 
|neuron_name = Horizontal Cell
</ul>
|image_neuron = Gray882.png
|caption_neuron = Plan of retinal neurons.
|location =
|function =
|neurotransmitter =
|morphology =
|afferents =
|efferents =
|NeuroLex = Horizontal Cell
|NeuroLexID = nifext_40
}}
 
'''Horizontal cells''' are the laterally interconnecting neurons having cell bodies in the [[inner nuclear layer]] of the  [[retina]] of [[vertebrate]] eyes. They help integrate and regulate the input from multiple [[photoreceptor cell]]s. Among their functions, horizontal cells are responsible for allowing eyes to adjust to see well under both bright and dim light conditions.
 
==Organization==
There are three basic types of horizontal cells, designated HI, HII and HIII.  The selectivity of these three horizontal cells, towards one of the three [[Cone cell#Types|cone types]], is a matter of debate.  According to studies conducted by Boycott and Wassle neither HI cells nor HII cells were selective towards S,M, or L cones. By contrast, Anhelt and Kolb claim that in their observations HI cells connected to all three cone types indiscriminantly, however, HII cells tended to contact S cones the most.  They also identified a third type of horizontal cell, HIII, which was identical to HI but did not make contact with S cones.
 
The HII cells also make connections with rods, but do so far enough away from the horizontal cell's soma such that they do not interfere with the activities of the cones.
 
They span across cones and summate inputs from them all to control the amount of [[GABA]] released back onto the photoreceptor cells, which hyperpolarizes them. Their arrangement together with the on-center and off-center [[Retina bipolar cell|bipolar cells]] that receive input from the photoreceptors constitutes a form of [[lateral inhibition]], increasing spatial resolution at the expense of some information on absolute intensity. The eye is thus more sensitive to contrast and differences in intensity.
 
There is a greater density of horizontal cells towards the central region of the retina. In cat retina it is observed that there is a density of
225 cells/mm<sup>2</sup> near the center of the retina and a density of 120 cells/mm<sup>2</sup> in the peripherals. <ref>{{cite journal|last=Wassle|first=H.|coauthors=Riemann, H. J.|title=The Mosaic of Nerve Cells in the Mammalian Retina|journal=Proceedings of the Royal Society B: Biological Sciences|date=22 March 1978|volume=200|issue=1141|pages=441–461|doi=10.1098/rspb.1978.0026}}</ref>
Horizontal cells and other retinal interneuron cells are less likely to be near neighbours of the same subtype than would occur by chance, resulting in ‘exclusion zones’ that separate them. Mosaic arrangements provide a mechanism to distribute each cell type evenly across the retina, ensuring that all parts of the visual field have access to a full set of processing elements. <ref>{{cite journal|last=Wassle|first=H.|coauthors=Riemann, H. J.|title=The Mosaic of Nerve Cells in the Mammalian Retina|journal=Proceedings of the Royal Society B: Biological Sciences|date=22 March 1978|volume=200|issue=1141|pages=441–461|doi=10.1098/rspb.1978.0026}}</ref> [[MEGF10]] and [[MEGF11]] transmembrane proteins have critical roles in the formation of the mosaics by horizontal cells and starburst [[Retina amacrine cell|amacrine cells]] in mice.<ref>{{cite journal|last=Kay|first=Jeremy N.|coauthors=Chu, Monica W., Sanes, Joshua R.|title=MEGF10 and MEGF11 mediate homotypic interactions required for mosaic spacing of retinal neurons|journal=Nature|date=March 2012|doi=10.1038/nature10877}}</ref>
 
==Functional properties==
Horizontal cells are depolarized by the release of glutamate from photoreceptors, which happens in the absence of light. Depolarization of a horizontal cell causes it to release the inhibitory neurotransmitter GABA on an adjacent photoreceptor. GABA inhibits this photoreceptor, effectively hyperpolarizing it. Conversely, in the light a photoreceptor does not release or it releases less glutamate onto the horizontal cell, which hyperpolarizes it and prevents or lowers the release of inhibitory GABA on the adjecent photoreceptor. We therefore have the following negative feedback.
 
'''Illumination<math>\to</math> Center photoreceptor hyperpolarization <math>\to</math> horizontal cell hyperpolarization <math>\to</math> Surround photoreceptor depolarization'''
 
One proposed theory for facilitation by the horizontal cells proceeds as follows{{Citation needed|reason=|date=January 2014}}. Assume we have 10 photoreceptors, one hyperpolarizing (H) bipolar cell, and one horizontal cell. All ten photoreceptors connect to the horizontal cell, and the middle photoreceptor (<math>P_m</math>) connects to the bipolar cell. The surrounding cells, which represent the outer receptive field, will be designated <math>P_o</math> then we can explain an off-centre arrangement as follows. If light is shone onto the <math>P_m</math> then
 
#<math>P_m</math> is activated by light and therefore hyperpolarizes
#<math>P_m</math> reduces its release of glutamate
#Reduction of glutamate hyperpolarizes the H bipolar cell
#Reduction of glutamate hyperpolarizes the horizontal cell and it reduces release of GABA
#Since <math>P_o</math>is still releasing glutamate, reduction in GABA is marginal
 
If the light is shone onto only <math>P_o</math> but not <math>P_m</math> then:
 
#<math>P_o</math> is activated and therefore hyperpolarizes
#<math>P_o</math> reduce their release of glutamate
#The reduction of glutamate hyperpolarizes the horizontal cell
#The horizontal cell reduces its release of GABA
#The reduction of GABA generally promotes the depolarization of adjacent photoreceptors
#However, because <math>P_o</math> are illuminated they are strongly hyperpolarized by phototransduction - therefore a reduction in GABA is insufficient to make <math>P_o</math> depolarise
#But, because <math>P_m</math> is already in darkness, the reduction in GABA levels is effective at causing <math>P_m</math> to depolarise
#<math>P_m</math> releases glutamate
#H Bipolar cell is depolarized
 
To explain diffuse light, then we consider both cases together, and as it turns out, the two effects cancel each other out, and we get little or no net effect. See http://en.wikipedia.org/wiki/Center_surround_antagonism
 
==See also==
*[[Photoreceptor cell|Photoreceptors]]
*[[Bipolar cell of the retina|Bipolar cells]]
*[[Amacrine cells]]
*[[Ganglion cells]]
 
== References ==
{{reflist}}
* {{cite book
  | last =Nicholls
  | first =John G.
  | coauthors =A. Robert Martin, Bruce G. Wallace, Paul A. Fuchs
  | title =From Neuron to Brain
  | publisher =Sinauer Associates, Inc
  | year =2001
  | location =Boston, Massachusetts
  | isbn = 0-87893-439-1
}}
 
* {{cite journal |author=Masland RH |title=The fundamental plan of the retina |journal=Nat. Neurosci. |volume=4 |issue=9 |pages=877–86 |year=2001 |pmid=11528418 |doi=10.1038/nn0901-877}}
 
* {{cite journal
  | last =Dacey
  | first =Dennis M
  | title =Primate Retina: Cell Types, Circuits and Color Opponency
  | journal =Progress in Retinal and Eye Research
  | volume =18
  | pages =737–763
  | year =1999
  | doi =10.1016/S1350-9462(98)00013-5
  | pmid =10530750
  | issue =6
}}
 
== External links ==
*[http://webvision.med.utah.edu/book/part-ii-anatomy-and-physiology-of-the-retina/horizontal-cells/#horizontal Webvision: Horizontal Cell]
* [https://www.neuinfo.org/mynif/search.php?q=Horizontal%20Cell&t=data&s=cover&b=0&r=20 NIF Search - Horizontal Cell] via the [[Neuroscience Information Framework]]
 
{{Eye anatomy}}
 
[[Category:Human eye anatomy]]
[[Category:Human cells]]
[[Category:Neurons]]

Revision as of 07:41, 7 January 2014

Template:Incomplete

Template:Infobox neuron

Horizontal cells are the laterally interconnecting neurons having cell bodies in the inner nuclear layer of the retina of vertebrate eyes. They help integrate and regulate the input from multiple photoreceptor cells. Among their functions, horizontal cells are responsible for allowing eyes to adjust to see well under both bright and dim light conditions.

Organization

There are three basic types of horizontal cells, designated HI, HII and HIII. The selectivity of these three horizontal cells, towards one of the three cone types, is a matter of debate. According to studies conducted by Boycott and Wassle neither HI cells nor HII cells were selective towards S,M, or L cones. By contrast, Anhelt and Kolb claim that in their observations HI cells connected to all three cone types indiscriminantly, however, HII cells tended to contact S cones the most. They also identified a third type of horizontal cell, HIII, which was identical to HI but did not make contact with S cones.

The HII cells also make connections with rods, but do so far enough away from the horizontal cell's soma such that they do not interfere with the activities of the cones.

They span across cones and summate inputs from them all to control the amount of GABA released back onto the photoreceptor cells, which hyperpolarizes them. Their arrangement together with the on-center and off-center bipolar cells that receive input from the photoreceptors constitutes a form of lateral inhibition, increasing spatial resolution at the expense of some information on absolute intensity. The eye is thus more sensitive to contrast and differences in intensity.

There is a greater density of horizontal cells towards the central region of the retina. In cat retina it is observed that there is a density of 225 cells/mm2 near the center of the retina and a density of 120 cells/mm2 in the peripherals. [1]

Horizontal cells and other retinal interneuron cells are less likely to be near neighbours of the same subtype than would occur by chance, resulting in ‘exclusion zones’ that separate them. Mosaic arrangements provide a mechanism to distribute each cell type evenly across the retina, ensuring that all parts of the visual field have access to a full set of processing elements. [2] MEGF10 and MEGF11 transmembrane proteins have critical roles in the formation of the mosaics by horizontal cells and starburst amacrine cells in mice.[3]

Functional properties

Horizontal cells are depolarized by the release of glutamate from photoreceptors, which happens in the absence of light. Depolarization of a horizontal cell causes it to release the inhibitory neurotransmitter GABA on an adjacent photoreceptor. GABA inhibits this photoreceptor, effectively hyperpolarizing it. Conversely, in the light a photoreceptor does not release or it releases less glutamate onto the horizontal cell, which hyperpolarizes it and prevents or lowers the release of inhibitory GABA on the adjecent photoreceptor. We therefore have the following negative feedback.

Illumination Center photoreceptor hyperpolarization horizontal cell hyperpolarization Surround photoreceptor depolarization

One proposed theory for facilitation by the horizontal cells proceeds as followsPotter or Ceramic Artist Truman Bedell from Rexton, has interests which include ceramics, best property developers in singapore developers in singapore and scrabble. Was especially enthused after visiting Alejandro de Humboldt National Park.. Assume we have 10 photoreceptors, one hyperpolarizing (H) bipolar cell, and one horizontal cell. All ten photoreceptors connect to the horizontal cell, and the middle photoreceptor (Pm) connects to the bipolar cell. The surrounding cells, which represent the outer receptive field, will be designated Po then we can explain an off-centre arrangement as follows. If light is shone onto the Pm then

  1. Pm is activated by light and therefore hyperpolarizes
  2. Pm reduces its release of glutamate
  3. Reduction of glutamate hyperpolarizes the H bipolar cell
  4. Reduction of glutamate hyperpolarizes the horizontal cell and it reduces release of GABA
  5. Since Pois still releasing glutamate, reduction in GABA is marginal

If the light is shone onto only Po but not Pm then:

  1. Po is activated and therefore hyperpolarizes
  2. Po reduce their release of glutamate
  3. The reduction of glutamate hyperpolarizes the horizontal cell
  4. The horizontal cell reduces its release of GABA
  5. The reduction of GABA generally promotes the depolarization of adjacent photoreceptors
  6. However, because Po are illuminated they are strongly hyperpolarized by phototransduction - therefore a reduction in GABA is insufficient to make Po depolarise
  7. But, because Pm is already in darkness, the reduction in GABA levels is effective at causing Pm to depolarise
  8. Pm releases glutamate
  9. H Bipolar cell is depolarized

To explain diffuse light, then we consider both cases together, and as it turns out, the two effects cancel each other out, and we get little or no net effect. See http://en.wikipedia.org/wiki/Center_surround_antagonism

See also

References

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    A vendor's stamp duty has been launched on industrial property for the primary time, at rates ranging from 5 per cent to 15 per cent. The Authorities might be trying to reassure the market that they aren't in opposition to foreigners and PRs investing in Singapore's property market. They imposed these measures because of extenuating components available in the market." The sale of new dual-key EC models will even be restricted to multi-generational households only. The models have two separate entrances, permitting grandparents, for example, to dwell separately. The vendor's stamp obligation takes effect right this moment and applies to industrial property and plots which might be offered inside three years of the date of buy. JLL named Best Performing Property Brand for second year running

    The data offered is for normal info purposes only and isn't supposed to be personalised investment or monetary advice. Motley Fool Singapore contributor Stanley Lim would not personal shares in any corporations talked about. Singapore private home costs increased by 1.eight% within the fourth quarter of 2012, up from 0.6% within the earlier quarter. Resale prices of government-built HDB residences which are usually bought by Singaporeans, elevated by 2.5%, quarter on quarter, the quickest acquire in five quarters. And industrial property, prices are actually double the levels of three years ago. No withholding tax in the event you sell your property. All your local information regarding vital HDB policies, condominium launches, land growth, commercial property and more

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  2. One of the biggest reasons investing in a Singapore new launch is an effective things is as a result of it is doable to be lent massive quantities of money at very low interest rates that you should utilize to purchase it. Then, if property values continue to go up, then you'll get a really high return on funding (ROI). Simply make sure you purchase one of the higher properties, reminiscent of the ones at Fernvale the Riverbank or any Singapore landed property Get Earnings by means of Renting

    In its statement, the singapore property listing - website link, government claimed that the majority citizens buying their first residence won't be hurt by the new measures. Some concessions can even be prolonged to chose teams of consumers, similar to married couples with a minimum of one Singaporean partner who are purchasing their second property so long as they intend to promote their first residential property. Lower the LTV limit on housing loans granted by monetary establishments regulated by MAS from 70% to 60% for property purchasers who are individuals with a number of outstanding housing loans on the time of the brand new housing purchase. Singapore Property Measures - 30 August 2010 The most popular seek for the number of bedrooms in Singapore is 4, followed by 2 and three. Lush Acres EC @ Sengkang

    Discover out more about real estate funding in the area, together with info on international funding incentives and property possession. Many Singaporeans have been investing in property across the causeway in recent years, attracted by comparatively low prices. However, those who need to exit their investments quickly are likely to face significant challenges when trying to sell their property – and could finally be stuck with a property they can't sell. Career improvement programmes, in-house valuation, auctions and administrative help, venture advertising and marketing, skilled talks and traisning are continuously planned for the sales associates to help them obtain better outcomes for his or her shoppers while at Knight Frank Singapore. No change Present Rules

    Extending the tax exemption would help. The exemption, which may be as a lot as $2 million per family, covers individuals who negotiate a principal reduction on their existing mortgage, sell their house short (i.e., for lower than the excellent loans), or take part in a foreclosure course of. An extension of theexemption would seem like a common-sense means to assist stabilize the housing market, but the political turmoil around the fiscal-cliff negotiations means widespread sense could not win out. Home Minority Chief Nancy Pelosi (D-Calif.) believes that the mortgage relief provision will be on the table during the grand-cut price talks, in response to communications director Nadeam Elshami. Buying or promoting of blue mild bulbs is unlawful.

    A vendor's stamp duty has been launched on industrial property for the primary time, at rates ranging from 5 per cent to 15 per cent. The Authorities might be trying to reassure the market that they aren't in opposition to foreigners and PRs investing in Singapore's property market. They imposed these measures because of extenuating components available in the market." The sale of new dual-key EC models will even be restricted to multi-generational households only. The models have two separate entrances, permitting grandparents, for example, to dwell separately. The vendor's stamp obligation takes effect right this moment and applies to industrial property and plots which might be offered inside three years of the date of buy. JLL named Best Performing Property Brand for second year running

    The data offered is for normal info purposes only and isn't supposed to be personalised investment or monetary advice. Motley Fool Singapore contributor Stanley Lim would not personal shares in any corporations talked about. Singapore private home costs increased by 1.eight% within the fourth quarter of 2012, up from 0.6% within the earlier quarter. Resale prices of government-built HDB residences which are usually bought by Singaporeans, elevated by 2.5%, quarter on quarter, the quickest acquire in five quarters. And industrial property, prices are actually double the levels of three years ago. No withholding tax in the event you sell your property. All your local information regarding vital HDB policies, condominium launches, land growth, commercial property and more

    There are various methods to go about discovering the precise property. Some local newspapers (together with the Straits Instances ) have categorised property sections and many local property brokers have websites. Now there are some specifics to consider when buying a 'new launch' rental. Intended use of the unit Every sale begins with 10 p.c low cost for finish of season sale; changes to 20 % discount storewide; follows by additional reduction of fiftyand ends with last discount of 70 % or extra. Typically there is even a warehouse sale or transferring out sale with huge mark-down of costs for stock clearance. Deborah Regulation from Expat Realtor shares her property market update, plus prime rental residences and houses at the moment available to lease Esparina EC @ Sengkang
  3. One of the biggest reasons investing in a Singapore new launch is an effective things is as a result of it is doable to be lent massive quantities of money at very low interest rates that you should utilize to purchase it. Then, if property values continue to go up, then you'll get a really high return on funding (ROI). Simply make sure you purchase one of the higher properties, reminiscent of the ones at Fernvale the Riverbank or any Singapore landed property Get Earnings by means of Renting

    In its statement, the singapore property listing - website link, government claimed that the majority citizens buying their first residence won't be hurt by the new measures. Some concessions can even be prolonged to chose teams of consumers, similar to married couples with a minimum of one Singaporean partner who are purchasing their second property so long as they intend to promote their first residential property. Lower the LTV limit on housing loans granted by monetary establishments regulated by MAS from 70% to 60% for property purchasers who are individuals with a number of outstanding housing loans on the time of the brand new housing purchase. Singapore Property Measures - 30 August 2010 The most popular seek for the number of bedrooms in Singapore is 4, followed by 2 and three. Lush Acres EC @ Sengkang

    Discover out more about real estate funding in the area, together with info on international funding incentives and property possession. Many Singaporeans have been investing in property across the causeway in recent years, attracted by comparatively low prices. However, those who need to exit their investments quickly are likely to face significant challenges when trying to sell their property – and could finally be stuck with a property they can't sell. Career improvement programmes, in-house valuation, auctions and administrative help, venture advertising and marketing, skilled talks and traisning are continuously planned for the sales associates to help them obtain better outcomes for his or her shoppers while at Knight Frank Singapore. No change Present Rules

    Extending the tax exemption would help. The exemption, which may be as a lot as $2 million per family, covers individuals who negotiate a principal reduction on their existing mortgage, sell their house short (i.e., for lower than the excellent loans), or take part in a foreclosure course of. An extension of theexemption would seem like a common-sense means to assist stabilize the housing market, but the political turmoil around the fiscal-cliff negotiations means widespread sense could not win out. Home Minority Chief Nancy Pelosi (D-Calif.) believes that the mortgage relief provision will be on the table during the grand-cut price talks, in response to communications director Nadeam Elshami. Buying or promoting of blue mild bulbs is unlawful.

    A vendor's stamp duty has been launched on industrial property for the primary time, at rates ranging from 5 per cent to 15 per cent. The Authorities might be trying to reassure the market that they aren't in opposition to foreigners and PRs investing in Singapore's property market. They imposed these measures because of extenuating components available in the market." The sale of new dual-key EC models will even be restricted to multi-generational households only. The models have two separate entrances, permitting grandparents, for example, to dwell separately. The vendor's stamp obligation takes effect right this moment and applies to industrial property and plots which might be offered inside three years of the date of buy. JLL named Best Performing Property Brand for second year running

    The data offered is for normal info purposes only and isn't supposed to be personalised investment or monetary advice. Motley Fool Singapore contributor Stanley Lim would not personal shares in any corporations talked about. Singapore private home costs increased by 1.eight% within the fourth quarter of 2012, up from 0.6% within the earlier quarter. Resale prices of government-built HDB residences which are usually bought by Singaporeans, elevated by 2.5%, quarter on quarter, the quickest acquire in five quarters. And industrial property, prices are actually double the levels of three years ago. No withholding tax in the event you sell your property. All your local information regarding vital HDB policies, condominium launches, land growth, commercial property and more

    There are various methods to go about discovering the precise property. Some local newspapers (together with the Straits Instances ) have categorised property sections and many local property brokers have websites. Now there are some specifics to consider when buying a 'new launch' rental. Intended use of the unit Every sale begins with 10 p.c low cost for finish of season sale; changes to 20 % discount storewide; follows by additional reduction of fiftyand ends with last discount of 70 % or extra. Typically there is even a warehouse sale or transferring out sale with huge mark-down of costs for stock clearance. Deborah Regulation from Expat Realtor shares her property market update, plus prime rental residences and houses at the moment available to lease Esparina EC @ Sengkang