Color Vision:

Color sense is the ability of the eye to discriminate between colors excited by light of the
different wavelengths. Or it is the ability to discriminate a light stimulus as a function of its
wavelength.
Facts about Color Vision:
 Colour vision is a function of cones and thus better appreciated in photopic vision.
 There are three different types of cones viz. red sensitive, green sensitive and blue
sensitive which combined perform the function of colour vision.
 Colours have three attributes: hue, Intensity & saturation.
 For any colour there is a complementary colour that, when properly mixed with it,
produces a sensation of white.
 The colour perceived depends in part on the colour of other objects in the visual field.
 A normal person can see all wavelengths between violets to red.
 Blue cones retain some sensitivity at around 10 nm, but crystalline lens blocks all UV
rays.
 In dim light all the colours are seen as gray, this is called Purkinje shift phenomenon.

Theories of Color Vision:

Theory of trichomacy:
 Originally suggested by Young (1802) and subsequently modified by Helmholtz
(1866). Hence it is called Young-Helmholtz theory.
 It postulates the existence of three kinds of cones.
  The sensation of any give- colour is determined by the relative frequency of the impulse
from each of the three cone systems.

 The red sensitive (Erythrolab) or long wave length sensitive (LWS)

 The green sensitive (Chlorolab) or middle wave length sensitive (MWS)
 The blue sensitive (Cyanolab) or short wave length sensitive (SWS)

 The Young-Helmholtz theory conclude that the blue, green and red are primary

 colour It has been studied that the gene for:

1) Human rhodopsin : chromosome 3

2) Blue cone: chromosome 7.
3) Red and green cones: q arm of X chromosome.

Opponent Color Theory:

 Hering proposed opponent color theory in 1892.
 He noted that there are some color combinations that we never see, such as reddish-
green or yellowish- blue.
 Hering hypothesized that trichromatic signals from the cones fed into subsequent neural
stages and exhibited two major opponent classes of processing:
1) Spectrally opponent processes which were red vs. green and yellow vs. blue.
2) Spectrally non-opponent processes which was black vs. white.
 Stage Theory:

Photochemistry of color Vision:

 The cone pigments like rhodopsin also consist of 11-cis-rentinal and an opsin part Known as
photopsin.
 It is different than the opsin part of the rhodospin.
 The green sensitive and red sensitive cone pigments are very similar in Structure.
 Each of these pigments has only about 43% homology with the opsin of blue sensitive
cone pigment.
 All the three cone pigments have about 41% homology with rod pigment rhodospin.
 The principles of photochemistry of rhodopsin can be applied to the cone pigments.
 The only difference being that the three different types of cones are bleached by light of
different wavelength.
Neurophysiology of Color Vision:

Similar to photochemical changes, the physiological ‘process ‘concerned with color vision are
also the same as for the vision in generals.
Genesis of Visual Signals in Photoreceptors:

Cone V/S Rod receptor Potential:

 The rods are much more sensitive.
 Cone receptor potential has a sharp onset and offset; whereas the rod receptor
potential has a sharp onset but slow offset.
 Rod responses are proportionate to stimulus intensity at levels of illumination that are
below the threshold for cones.
 cone responses are proportionate to stimulus intensity at high levels of illumination
when the rod responses are maximal and cannot change.
Processing and Transmission of Color Vision Signal in the retina:
 The receptor potential generated in the photoreceptors is transmitted by electronic
conduction.
 The ganglion cells transmit the visual signal by means of action potential.
Horizontal Cells:
 Transmit signals horizontally in the OPL from rods and cones to the bipolar cells.
 Their main function is to enhance the visual contrast by causing lateral inhibitions.
 When a minute spot of light strikes the retina, the central most area is excited but the
area around (called as surround) is inhibited.
 Horizontal cells showed two completely different kinds of response:
1) First, there was a hyperpolarizing response with a broad spectral function
termed as luminosity response.
2) Second, a chromatic (c) response which was hyperpolarizing for part of the
spectrum and depolarizing for the remainder.

Bipolar Cell:
 The bipolar cells are the first order neurons of visual pathway.
 Recordings made from goldfish bipolar cells showed a 'center-surround'
spatial pattern.
 The two different types of bipolar cells provide opposing excitatory and
inhibitory signals in the visual pathway.
 Receptive fields of the bipolar cell is also circular in configuration but has
got a center- surround antagonism.
 The importance is, it provides a second mechanism for lateral inhibition
in addition to horizontal cell mechanism.
Amacrine Cell:
 The exact role of these cells in colour vision is not clear.
 Some workers have speculated that they may act as an 'automatic colour
control'
Ganglion Cell:
 It is at this level that we see first direct evidence in the visual system for
colour coding.

 There are three distinct groups of ganglion cells designated as W, X and Y

cells.

 It has been observed that colour sensation is mediated by the 'X' ganglion cells
 When all the three types of cones stimulate the same ganglion cell the resultant
signal is white
 Opponent colour cell : some of the ganglion cells are excited by one
colour type cone (e.g. red) and are inhibited by other (i.e. green) or vice
versa.
 It is concerned in the ‘successive colour contrast’.
 Double opponent colour cell: these ganglion cells have a system which is
opponent for both colour and space.
 This system is called ‘double opponent cell’ system and is concerned with
the ‘simultaneous colour contrast’.
 The double opponent cells have a receptive field with a centre and
surround.
 The response may be ‘on’ to red colour in the centre and ‘off’ to it in the
surround, while the response may be ‘off’ to green in the centre and ‘on’
to it in surround.
Distribution of Color Vision in the Retina:
 Trichromatic colour vision mechanism extends 20-30 degrees from the point of fixation.
 Peripheral to this red and green become indistinguishable, and
 In the far periphery all colour sense is lost.
 The very centre of fovea (1/8 degree) is blue blind.
 It is attributed to chromatic aberration
Processing of Color Signals in lateral geniculate body (LGB):
 All lateral geniculate body neurons carry information from
more than one cone cell.
 Colour information carried by the ganglion cells is relayed to
the parvocellular portion of the LGB.
 Spectrally nonopponent cells which give the same type of
response to any monochromatic light constitute about 30 per
cent of all the LGB neurons.
 Spectrally opponent cells make 60% of LGB neurons.
 These cells are excited by some wavelengths and inhibited by
others and thus appear to carry colour information.
 These have been classified into 4 types:

1) Cells having red and green antagonism with +R/-G.

2) Cells having red and green antagonism with +G/-R.
3) Cells having blue and yellow antagonism with +B/-Y.
4) Cells having blue and yellow antagonism with +Y/-B.

Analysis of Color Signals in the Visual Cortex:

 Colour information from the parvocelluar portion of the LGB is relayed to the
layer IV c of the striate cortex.
 From there, the information passes to the blobs in layers II and III.
 The neurons in the Blobs lack orientation specificity but respond to colours.
 Like the ganglion cells and LGB cells they are centre-surround cells.
  Colour information is then relayed to thin strips in the visual association area
and from there to a specialized area concerned with colour.

Phenomena Associated with Color Sense:

 Simultaneous colour contrast:

1) This phenomenon refers to perception of particular

coloured spot against the coloured background.
2) Ex: a grey spot appears greenish in a red surround and
redish in a green surround.
3) It is function of double opponent cell
 Successive colour contrast:

1) It is a phenomenon of coloured after images,

2) The colour of the after image tends to be near the complementary of the
primary image.
3) Ex: when one sees at a green spot for several seconds and then looks at a
grey card, one sees a red spot on the card.

 Phenomenon of colour constancy:

1) It refers to a phenomenon, in which the human eye
continues to perceive the colour of a particular object
unchanged even after the spectral composition of the
 light falling on it is markedly altered.

Hierarchy of Color Coded Cell:

The phenomenon of hierarchy of colour coded cells suggests a system of serial
analysis of colour sense. The colour coded cells have been reported to be
arranged in a hierarchy manner as follows:

 The opponent colour cells being located in ganglion cells and lateral
geniculate neurons and

 The double opponent cells with 'center-surround' receptive fields in the layer IV of
striate cortex.

 Complex and hyper complex colour coded cells have been described in the layers II, III, V
and VI of the striate cortex in the form of 'blobs'.

The Color Metric:

 They have been evolved on a psychological basis and have no
connection with the physiology of eye.

 The two important colour metric systems used internationally in industry, in

printing and in the graphic arts.

The CIE Color Space System:

 This system was developed by the International Commission of Illumination
(CIE – Commission Internal de Eclairage

 CIE colour space system is based on the amounts of three primary colours necessary to
match a specified colour.
The Muncell Color System:
 In this system all the colours are represented in a cylinder in terms of hue, value and chroma
(HVC).
 This system covers a wide range of colours and is thus widely used in medicine and industry.
Normal Color Attributes:
 Hue:
i. i.e.dominant spectral colour
ii. It is determined by the wavelength of the particular colour.
iii. Munsell defined hue as "the quality by which we distinguish
one color from another."

 Saturation:

It can be estimated by measuring how much of a particular wavelength

must be added to white before it is distinguishable from white.
 Lightness or Brightness: The lightness or brightness of a colour depends upon the luminosity
of the component wavelength. In photopic vision normal eye has a peak luminosity function at
approximately 555 nm and in scotopic vision at about 507 nm.