Scotopic vision

{{Short description|Visual perception under low-light conditions}}

In the study of visual perception, '''scotopic vision''' (or '''scotopia''') is the vision of the eye under low-light conditions.<ref>{{Cite book |title=A Dictionary of Biology |publisher=Oxford University Press |year=2019 |isbn=978-0-19-882148-9 |editor-last=Hine |editor-first=Robert |edition=8th |location=New York, NY |doi=10.1093/acref/9780198821489.001.0001}}</ref> The term comes from the Greek ''skotos'', meaning 'darkness', and ''-opia'', meaning 'a condition of sight'.<ref>{{cite Dictionary.com|scotopia}}</ref> In the human eye, cone cells are nonfunctional in low visible light. Scotopic vision is produced exclusively through rod cells, which are most sensitive to wavelengths of around 498 nm (blue-green)<ref name="Bowmaker1980"}>{{Cite journal | last1 = Bowmaker | first1= J K | first2= H J | last2= Dartnall. | title = Visual pigments of rods and cones in a human retina | journal =The Journal of Physiology | volume = 298 | year = 1980 | issue= 1 |pages= 501–11 | doi = 10.1113/jphysiol.1980.sp013097 | pmid= 7359434 | pmc = 1279132 | bibcode= 1980JPhsg.298..501B }}</ref> and are insensitive to wavelengths longer than about 640 nm.<ref>{{Cite book |last1=Frisby |first1=John P. |title=Seeing: The Computational Approach to Biological Vision |last2=Stone |first2=James V. |publisher=MIT Press |year=2010 |isbn=978-0-262-51427-9 |edition=2nd |location=Cambridge, Mass |oclc=430192600 }}</ref> Under scotopic conditions, light incident on the retina is not encoded in terms of the spectral power distribution. Higher visual perception occurs under scotopic vision as it does under photopic vision.<ref name="McKyton Elul Levin 2024 p. 108929" />

==Retinal circuitry==

Of the two types of photoreceptor cells in the retina, rods dominate scotopic vision. This dominance is due to the increased sensitivity of the photopigment molecule expressed in rods, as opposed to those in cones. Rods signal light increments to rod bipolar cells which, unlike most types of bipolar cells, do not form direct connections with retinal ganglion cells – the output neurons of the retina. Instead, two types of amacrine cell – AII and A17 – allow lateral information flow from rod bipolar cells to cone bipolar cells, which in turn contact ganglion cells. Thus, rod signals, mediated by amacrine cells, dominate scotopic vision.<ref>{{Citation |last=Kolb |first=Helga |title=Roles of Amacrine Cells |date=1995 |work=Webvision: The Organization of the Retina and Visual System |editor-last=Kolb |editor-first=Helga |url=https://www.ncbi.nlm.nih.gov/books/NBK11539/ |access-date=2024-04-16 |place=Salt Lake City (UT) |publisher=University of Utah Health Sciences Center |pmid=21413397 |editor2-last=Fernandez |editor2-first=Eduardo |editor3-last=Nelson |editor3-first=Ralph}}</ref>

==Occurrence== Scotopic vision occurs at luminance levels of 10−3<ref>{{Cite book |editor-last=Pokorny |editor-first=Joel |display-editors=etal |year=1979 |series=Current Ophthalmology Monographs |title=Congenital and Acquired Color Vision Defects |chapter=Chapter 2 |url=http://faculty.washington.edu/sbuck/545ColorClass/PokornyCh2.1979b.PDF |access-date=2016-01-14 |archive-date=2016-03-04 |archive-url=https://web.archive.org/web/20160304083808/http://faculty.washington.edu/sbuck/545ColorClass/PokornyCh2.1979b.PDF }}</ref> to 10−6{{citation needed|date=January 2016}} cd/m2. Other species are not universally color blind in low-light conditions. The elephant hawk-moth (''Deilephila elpenor'') displays advanced color discrimination even in dim starlight.<ref>{{cite journal|title=Scotopic colour vision in nocturnal hawkmoths|first1=Almut|last1=Kelber|first2=Anna|last2=Balkenius|first3=Eric J.|last3=Warrant|date=31 October 2002|journal=Nature|volume=419|issue=6910|pages=922–925|doi=10.1038/nature01065|pmid = 12410310|bibcode = 2002Natur.419..922K|s2cid=4303414}}</ref>

Mesopic vision occurs in intermediate lighting conditions (luminance level 10−3 to 100.5 cd/m2){{Citation needed|date=May 2021}} and is effectively a combination of scotopic and photopic vision. This gives inaccurate visual acuity and color discrimination.

In normal light (luminance level 10 to 108 cd/m2), the vision of cone cells dominates and is photopic vision. There is good visual acuity (VA) and color discrimination.

==Wavelength sensitivity== [[File:CIE 1951 scotopic luminosity function.svg|thumb|300px|The CIE 1951 scotopic luminosity function. The horizontal axis is wavelength in nm.]] The normal human observer's relative wavelength sensitivity will not change due to background illumination under scotopic vision. The wavelength sensitivity is determined by the rhodopsin photopigment. This is a red pigment seen at the back of the eye in animals that have a white background to their eye called Tapetum lucidum. The pigment is not noticeable under photopic and mesopic conditions. The principle that the wavelength sensitivity does not change during scotopic vision led to the ability to detect two functional cone classes in individuals. If two cone classes are present, then their relative sensitivity will change the behavioral wavelength sensitivity. Therefore, experimentation can determine "the presence of two cone classes by measuring wavelength sensitivity on two different backgrounds and noting a change in the observer's relative wavelength sensitivity."<ref name="auto"/>

The behavior of the rhodopsin photopigment explains why the human eye cannot resolve lights with different spectral power distributions under low light. The reaction of this single photopigment will give the same quanta for 400 nm light and 700 nm light. Therefore, this photopigment only maps the rate of absorption and does not encode information about the relative spectral composition of the light.<ref name="auto"/>

In scientific literature, one occasionally encounters the term ''scotopic lux'' which corresponds to photopic lux, but uses instead the scotopic visibility weighting function.<ref>[https://books.google.com/books?id=GUVHtJCtM0UC Photobiology: The Science of Light and Life] {{Webarchive|url=https://web.archive.org/web/20240409024207/https://books.google.com/books?id=GUVHtJCtM0UC |date=2024-04-09 }} (2002), Lars Olof Björn, [https://books.google.com/books?id=GUVHtJCtM0UC&pg=PA43 p.43] {{Webarchive|url=https://web.archive.org/web/20240409024229/https://books.google.com/books?id=GUVHtJCtM0UC&pg=PA45&source=gbs_toc_r&cad=0_0&sig=xLVz68xyrQB3DzosDDO9upHfsdU#v=onepage&q&f=false#PPA43,M1 |date=2024-04-09 }}, {{ISBN|1-4020-0842-2}}</ref> The scotopic luminosity function is a standard function established by the Commission Internationale de l'Éclairage (CIE) and standardized in collaboration with the ISO.<ref>{{cite book |title=ISO/CIE 23539:2023 CIE TC 2-93 Photometry — The CIE system of physical photometry |date=2023 |publisher=ISO/CIE |url=https://www.iso.org/standard/83178.html |language=en |access-date=2023-04-14 |archive-date=2023-04-08 |archive-url=https://web.archive.org/web/20230408225321/https://www.iso.org/standard/83178.html |url-status=live }}</ref>

The maximum scotopic efficacy is 1700 lm/W at 507 nm (compared with 683 lm/W at 555 nm for maximum photopic efficacy).<ref name="scotopic efficacy">{{cite web|url=http://hyperphysics.phy-astr.gsu.edu/hbase/vision/bright.html|title=Brightness and Night/Day Sensitivity|access-date=2018-12-03|archive-date=2014-02-21|archive-url=https://web.archive.org/web/20140221094605/http://hyperphysics.phy-astr.gsu.edu/hbase/vision/bright.html|url-status=live}}</ref> While the ratio between scotopic and photopic efficacies is only around 2.5 counted at peak sensitivity the ratio increases strongly below 500 nm.

== Resolution == For adaptation to occur at very low levels, the human eye needs to have a large sample of light across the signal in order to get a reliable image. This leads to the human eye being unable to resolve high spatial frequencies in low light since the observer is spatially averaging the light signal.<ref name="auto">{{cite web|url=http://foundationsofvision.stanford.edu/|title=Foundations of Vision|website=foundationsofvision.stanford.edu|access-date=2013-03-13|archive-date=2023-06-17|archive-url=https://web.archive.org/web/20230617205219/https://foundationsofvision.stanford.edu/|url-status=live}}</ref>

Another reason that vision is poor under scotopic vision is that rods, which are the only cells active under scotopic vision, converge to a smaller number of neurons in the retina. This many-to-one ratio leads to poor spatial frequency sensitivity.<ref name="auto"/>

== General perception == High-level visual perception is similar with scotopic as with photopic sight with people reading with unimpaired accuracy (though twice as long fixations), able to recognize faces, and show a face inversion effect.<ref name="McKyton Elul Levin 2024 p. 108929">{{cite journal |last1=McKyton |first1=Ayelet |last2=Elul |first2=Deena |last3=Levin |first3=Netta |date=2024 |title=Seeing in the dark: High-order visual functions under scotopic conditions |journal=iScience |volume=27 |issue=2 |article-number=108929 |doi=10.1016/j.isci.2024.108929 |pmc=10844829 |pmid=38322984|bibcode=2024iSci...27j8929M }}</ref>

== See also == {{col div}} * {{annotated link|Photopic vision}} * {{annotated link|Mesopic vision}} * {{annotated link|Adaptation (eye)}} * {{annotated link|Averted vision}} * {{annotated link|Night vision}} * {{annotated link|Purkinje effect}} * {{annotated link|Scotopic stilb}} * {{annotated link|Skot (unit)}} * {{annotated link|Spatial frequency}} * {{annotated link|Spectral sensitivity}} {{colend}}

==References== {{reflist}}

== Further reading == {{refbegin}} * {{Cite journal | pmid = 25237297| pmc = 4154443| year = 2014| last1 = Marc| first1 = R. E.| title = The AII amacrine cell connectome: A dense network hub| journal = Frontiers in Neural Circuits| volume = 8| page = 104| last2 = Anderson| first2 = J. R.| last3 = Jones| first3 = B. W.| last4 = Sigulinsky| first4 = C. L.| last5 = Lauritzen| first5 = J. S.| doi = 10.3389/fncir.2014.00104| doi-access = free}} * {{cite journal |last1=Zele |first1=Andrew J. |last2=Cao |first2=Dingcai |title=Vision under mesopic and scotopic illumination |journal=Frontiers in Psychology |date=22 January 2015 |volume=5 |page=1594 |doi=10.3389/fpsyg.2014.01594|doi-access=free|pmid=25657632|pmc=4302711 }} (Review) {{refend}}

{{DEFAULTSORT:Scotopic Vision}} Category:Eye Category:Vision