Purkinje effect
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The Purkinje effect (; sometimes called the Purkinje shift, often incorrectly pronounced ) is the tendency for the peak
luminance Luminance is a photometric measure of the luminous intensity per unit area of light travelling in a given direction. It describes the amount of light that passes through, is emitted from, or is reflected from a particular area, and falls with ...
sensitivity of the eye to shift toward the
blue Blue is one of the three primary colours in the RYB colour model (traditional colour theory), as well as in the RGB (additive) colour model. It lies between violet and cyan on the spectrum of visible light. The eye perceives blue when ...
end of the
color spectrum The visible spectrum is the portion of the electromagnetic spectrum that is visible to the human eye. Electromagnetic radiation in this range of wavelengths is called ''visible light'' or simply light. A typical human eye will respond to wavel ...
at low illumination levels as part of
dark adaptation Darkness, the direct opposite of lightness, is defined as a lack of illumination, an absence of visible light, or a surface that absorbs light, such as black or brown. Human vision is unable to distinguish colors in conditions of very low lum ...
. In consequence, reds will appear darker relative to other colors as light levels decrease. The effect is named after the
Czech Czech may refer to: * Anything from or related to the Czech Republic, a country in Europe ** Czech language ** Czechs, the people of the area ** Czech culture ** Czech cuisine * One of three mythical brothers, Lech, Czech, and Rus' Places * Czech, ...
anatomist Anatomy () is the branch of biology concerned with the study of the structure of organisms and their parts. Anatomy is a branch of natural science that deals with the structural organization of living things. It is an old science, having it ...
Jan Evangelista Purkyně Jan Evangelista Purkyně (; also written Johann Evangelist Purkinje) (17 or 18 December 1787 – 28 July 1869) was a Czech anatomist and physiologist. In 1839, he coined the term '' protoplasm'' for the fluid substance of a cell. He was one of ...
. While the effect is often described from the perspective of the human eye, it is well established in a number of animals under the same name to describe the general shifting of spectral sensitivity due to pooling of rod and cone output signals as a part of dark/light adaptation. This effect introduces a difference in color contrast under different levels of illumination. For instance, in bright
sunlight Sunlight is a portion of the electromagnetic radiation given off by the Sun, in particular infrared, visible, and ultraviolet light. On Earth, sunlight is scattered and filtered through Earth's atmosphere, and is obvious as daylight when ...
,
geranium ''Geranium'' is a genus of 422 species of annual, biennial, and perennial plants that are commonly known as geraniums or cranesbills. They are found throughout the temperate regions of the world and the mountains of the tropics, but mostly i ...
flower A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechanis ...
s appear bright red against the dull
green Green is the color between cyan and yellow on the visible spectrum. It is evoked by light which has a dominant wavelength of roughly 495570 nm. In subtractive color systems, used in painting and color printing, it is created by a combin ...
of their
leaves A leaf ( : leaves) is any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, st ...
, or adjacent blue flowers, but in the same scene viewed at
dusk Dusk occurs at the darkest stage of twilight, or at the very end of astronomical twilight after sunset and just before nightfall.''The Random House College Dictionary'', "dusk". At predusk, during early to intermediate stages of twilight, enou ...
, the contrast is reversed, with the red
petal Petals are modified leaves that surround the reproductive parts of flowers. They are often brightly colored or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the ''corolla''. Petals are usuall ...
s appearing a dark red or black, and the leaves and blue petals appearing relatively bright. The sensitivity to light in
scotopic vision In the study of human visual perception, scotopic vision (or scotopia) is the vision of the eye under low-light conditions. The term comes from Greek ''skotos'', meaning "darkness", and ''-opia'', meaning "a condition of sight". In the human eye, ...
varies with wavelength, though the perception is essentially
black-and-white Black-and-white (B&W or B/W) images combine black and white in a continuous spectrum, producing a range of shades of grey. Media The history of various visual media began with black and white, and as technology improved, altered to color. ...
. The Purkinje shift is the relation between the absorption maximum of
rhodopsin Rhodopsin, also known as visual purple, is a protein encoded by the RHO gene and a G-protein-coupled receptor (GPCR). It is the opsin of the rod cells in the retina and a light-sensitive receptor protein that triggers visual phototransductio ...
, reaching a maximum at about 500 nm, and that of the
opsin Animal opsins are G-protein-coupled receptors and a group of proteins made light-sensitive via a chromophore, typically retinal. When bound to retinal, opsins become Retinylidene proteins, but are usually still called opsins regardless. Most ...
s in the longer-wavelength cones that dominate in
photopic vision Photopic vision is the vision of the eye under well-lit conditions (luminance levels from 10 to 108  cd/m2). In humans and many other animals, photopic vision allows color perception, mediated by cone cells, and a significantly higher visu ...
, about 555 nm (green). In visual
astronomy Astronomy () is a natural science that studies celestial objects and phenomena. It uses mathematics, physics, and chemistry in order to explain their origin and evolution. Objects of interest include planets, moons, stars, nebulae, g ...
, the Purkinje shift can affect visual estimates of
variable star A variable star is a star whose brightness as seen from Earth (its apparent magnitude) changes with time. This variation may be caused by a change in emitted light or by something partly blocking the light, so variable stars are classified as e ...
s when using comparison stars of different colors, especially if one of the
star A star is an astronomical object comprising a luminous spheroid of plasma (physics), plasma held together by its gravity. The List of nearest stars and brown dwarfs, nearest star to Earth is the Sun. Many other stars are visible to the naked ...
s is red.


Physiology

The Purkinje effect occurs at the transition between primary use of the
photopic Photopic vision is the vision of the eye under well-lit conditions (luminance levels from 10 to 108  cd/m2). In humans and many other animals, photopic vision allows color perception, mediated by cone cells, and a significantly higher visua ...
(cone-based) and
scotopic In the study of human visual perception, scotopic vision (or scotopia) is the vision of the eye under low-light conditions. The term comes from Greek ''skotos'', meaning "darkness", and ''-opia'', meaning "a condition of sight". In the human eye, ...
(rod-based) systems, that is, in the
mesopic Mesopic vision, sometimes also called twilight vision, is a combination of photopic and scotopic vision under low-light (but not necessarily dark) conditions. Mesopic levels range approximately from 0.01 to 3.0  cd/m2 in luminance. Most ni ...
state: as intensity dims, the rods take over, and before color disappears completely, it shifts towards the rods' top sensitivity. The effect occurs because in mesopic conditions the outputs of cones in 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 ...
, which are generally responsible for the perception of color in daylight, are pooled with outputs of rods which are more sensitive under those conditions and have peak sensitivity in blue-green wavelength of 507 nm.


Use of red lights

The insensitivity of rods to long-wavelength light has led to the use of red lights under certain special circumstances—for example, in the control rooms of submarines, in research laboratories, aircraft, or during naked-eye astronomy. Red lights are used in conditions where it is desirable to activate both the photopic and scotopic systems. Submarines are well lit to facilitate the vision of the crew members working there, but the control room must be lit differently to allow crew members to read instrument panels yet remain dark adjusted. By using red lights or wearing red goggles, the cones can receive enough light to provide photopic vision (namely the high-acuity vision required for reading). The rods are not saturated by the bright red light because they are not sensitive to long-wavelength light, so the crew members remain dark adapted. Similarly, airplane cockpits use red lights so pilots can read their instruments and maps while maintaining night vision to see outside the aircraft. Red lights are also often used in research settings. Many research animals (such as rats and mice) have limited photopic vision, as they have far fewer cone photoreceptors. The animal subjects do not perceive red lights and thus experience darkness (the active period for
nocturnal Nocturnality is an animal behavior characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnal meaning the opposite. Nocturnal creatures generally have highly developed sens ...
animals), but the human researchers, who have one kind of cone (the "L cone") that is sensitive to long wavelengths, are able to read instruments or perform procedures that would be impractical even with fully dark adapted (but low acuity) scotopic vision. For the same reason, zoo displays of nocturnal animals often are illuminated with red light.


History

The effect was discovered in 1819 by
Jan Evangelista Purkyně Jan Evangelista Purkyně (; also written Johann Evangelist Purkinje) (17 or 18 December 1787 – 28 July 1869) was a Czech anatomist and physiologist. In 1839, he coined the term '' protoplasm'' for the fluid substance of a cell. He was one of ...
. Purkyně was a
polymath A polymath ( el, πολυμαθής, , "having learned much"; la, homo universalis, "universal human") is an individual whose knowledge spans a substantial number of subjects, known to draw on complex bodies of knowledge to solve specific pro ...
who would often meditate at dawn during long walks in the blossomed
Bohemia Bohemia ( ; cs, Čechy ; ; hsb, Čěska; szl, Czechy) is the westernmost and largest historical region of the Czech Republic. Bohemia can also refer to a wider area consisting of the historical Lands of the Bohemian Crown ruled by the Bohem ...
n fields. Purkyně noticed that his favorite flowers appeared bright red on a sunny afternoon, while at dawn they looked very dark. He reasoned that the eye has not one but two systems adapted to see colors, one for bright overall light intensity, and the other for dusk and dawn. Purkyně wrote in his ''Neue Beiträge:''As quoted in:
Objectively, the degree of illumination has a great influence on the intensity of color quality. In order to prove this most vividly, take some colors before daybreak, when it begins slowly to get lighter. Initially one sees only black and grey. Particularly the brightest colors, red and green, appear darkest. Yellow cannot be distinguished from a rosy red. Blue became noticeable to me first. Nuances of red, which otherwise burn brightest in daylight, namely carmine, cinnabar and orange, show themselves as darkest for quite a while, in contrast to their average brightness. Green appears more bluish to me, and its yellow tint develops with increasing daylight only.


See also

*
Kruithof curve The Kruithof curve describes a region of illuminance levels and color temperatures that are often viewed as comfortable or pleasing to an observer. The curve was constructed from psychophysical data collected by Dutch physicist Arie Andries K ...
*
Dark adaptation Darkness, the direct opposite of lightness, is defined as a lack of illumination, an absence of visible light, or a surface that absorbs light, such as black or brown. Human vision is unable to distinguish colors in conditions of very low lum ...
*
Dark adaptor goggles Dark adaptor goggles, also called red adaptation goggles, are used in the field of meteorology and astronomy for adapting the eyes to the dark prior to an observation at night. They also aid with the identification of clouds during bright sunshi ...


References


External links


Color Optical Illusions, Purkinje Effect
{{DEFAULTSORT:Purkinje Effect Color appearance phenomena Color vision Visual perception