How Do We Perceive Colour?

Dionne Angela Donnelly Tutorial Essay Year One

Perception has been defined as the acquisition and processing of sensory information in order to see, hear, taste, or feel objects in the world; [perception] also guides an organisms actions with respect to those objects (Sekuler & Blake, 2002, p.621 cited in Eysenck & Keane, 2005). Perception provides each of us with an individual representation of the outside world. It is commonly assumed that perception occurs solely within the brain. However, since the 19th century theories have focused on the processes that occur before information about stimuli reaches the brain. The two most significant of these are; the trichromatic and opponent-process theories. When first developed they were considered to be contradictory, but it is now accepted that the two are synthesised in an attempt to give a complete picture of human colour perception.

To understand these theories it is necessary to define what is meant by colour. Colours have three physical characteristics (attributes of the stimulus itself) and three perceptual characteristics (what we actually see). Each physical quality is essentially synonymous with its perceptual counterpart. The first of these are wavelength/hue. A wavelength refers to the distance between two oscillating light waves (measured in nanometres). Altogether they form the electromagnetic scale. Humans can only see a limited part of this scale, ranging from 380 nanometres (nm) to 760 nm. Wavelengths between these two extremes are all associated with a specific hue. For example, a wavelength of 560 nm is equal to a yellow-green colour. The second sets of characteristics are intensity/brightness, they depend upon the amount of energy that is being emitted or reflected by the object. The third sets of characteristics are purity/saturation. A colour is fully saturated if it contains light of only one wavelength e.g. pure yellow. The level of saturation depends upon how intense the dominant wavelength is in relation to the total amount of energy emitted/reflected. A pure or fully saturated colour can be desaturated by adding white light. This is because white light is made up of the 2

wavelengths of all possible hues. As there is no dominant wavelength, white light is described as being completely desaturated (Buskist, Carlson & Martin, 2004). However, not all of the 7 million colours we can potentially see can be made in this fashion, as the spectrum only contains a limited number of potential hues.

In order to create colours that do not appear on the spectrum it is necessary to mix spectral colours. There are two types of mixing. Colour mixing refers to mixing lights i.e. adding two or more wavelengths together. Pigment mixing refers to mixing physical substances e.g. paints. When we mix lights there is a lighter resulting colour e.g. red + green = yellow. When we mix pigments there is a darker result e.g. red + blue = purple. This is because light mixing is an additive process; the two lights can be separated again, by a prism for example. The two colours do not seem separate because our visual faculty synthesises them. The mixing effect here occurs within our own perception rather than in the physical world. In contrast, pigment mixing is a subtractive process. If you mix two different paints they become one because the light waves that were previously reflected by the object are now absorbed, meaning that less waves are actually reflected. Consequently, if red, green and blue lights are mixed together it creates white light, but if we mix red, green and blue paint it creates a dark grey or black. It is also possible to create almost any hue from these three primary colours, by varying their intensity. This can be seen in everyday life in television and computer screens, which use red, green and blue dots to create the images we see.

In order for us to see such colours, light passes through the cornea and lens of the eye and hits the back of the retina. It is here that the process of colour perception begins. Situated at the back of the retina are photoreceptors sensitive to different kinds of light. Rods are adapted for 3

dim light (scotopic) vision and also deal with achromatic (colourless vision). Cones are lightadapted (photopic) and play a vital role in colour discrimination. Cones are most prevalent in and around the fovea, which is a small depression in the centre of the retina.

Young (1802) made them the central feature of his trichromatic theory of vision (updated by Helmholtz, 1866). He identified that there were three different kinds of cone, each containing a different photopigment (chemical sensitive to photons (particles of light) which when hit by the photons it is most responsive to, breaks down and starts a chemical reaction). Cones are sensitive to either short wavelengths (e.g. blue), medium wavelengths (e.g. yellow-green) or long wavelengths (e.g. red). Thus causing them to be labelled as S-cones, M-cones and L-cones respectively (according to Cicerone and Nerger (1989, cited in Eysenck & Keane, 2005) there are approximately 4 million L-cones, 2 million M-cones and 1 million S-cones). Each cone is responsive to a wide range of wavelengths, but each cones sensitivity peaks at a different stage (See Figure 1).

S-cones

L-cones

M-cones

Figure 1: graph showing the spectral sensitivity of cones (and rods). (Schubert, n.d. Adapted from Dowling, 1987.)

Therefore, colour perception is not based upon which type of cone is responding; it is based upon the way the cones are responding in relation to one another. For example, if a wavelength of 480 nm hits the retina, the S-cones and M-cones will both respond equally. 4

This will be roughly double the response of the L-cones. Other patterns will represent different wavelengths (Fridlund, Gleitman & Reisberg, 2004). Also, stimulation of a certain cone is not essential for sensation of a specific hue, as colours such as blue can be perceived without blue light actually hitting the S-cones (Hofer, Singer & Williams, 2005, cited in Kingdom & Shevell, 2008). The Young-Helmholtz trichromatic theory has been supported by Dartnall, Bowmaker and Mollons research (1983, cited in Eysenck & Keane, 2005). They used microspectrophotometry (measures the amount of light absorbed at various wavelengths by individual cone receptors) to investigate whether the cones do respond differently to different wavelengths. They found that this is the case and that there are actually three different types of cone as stated by the trichromatic theory.

However, despite there being such evidence for the validity of the trichromatic theory, there are also a number of limitations. The first of these is that the theory does not account for the fact that yellow appears to be a pure colour (along with red, green and blue) unlike other colours such as purple which is quite clearly a mix of colours. A second, more important limitation is that it does not explain why colours come in pairs. These are known as complementary colours and are linked in such a way that if an individual stares at one of the pair then looks at a white background, an image of its counterpart appears. This is referred to as negative after-imaging e.g. if we look at an image of a red rose then a white background we will instead see an image of a green rose. Another limitation similar to negative afterimaging is that it cannot explain the phenomenon of simultaneous colour contrast. This occurs when we look at a colour and its complementary colour appears in an area beside the original colour e.g. if a small grey square is placed inside a large blue square the grey square will appear yellow. It is for these reasons that trichromacy cannot account wholly for how colour perception occurs. 5

In response, Hering (1878) postulated a more comprehensive theory of colour perception: the opponent-process theory. Herings theory states that there are three different opponentprocesses at work in the visual system. These are antagonistic pairings of red-green, blueyellow and black-white. If one side of the pair is stimulated then the other is inhibited, therefore we can never see a reddish-green or a yellowish-blue. Recent evidence by Abramov and Gordon (1994, cited in Eysenck & Keane, 2005) supports this assumption. When they asked participants to state the amount of blue, green, yellow and red they could see, no-one saw mixtures of blue and yellow or red and green.

However, Herings theory is only a basic interpretation of what occurs within the early stages of the visual system. Hurvich and Jamesons (1957) development of the theory to include contemporary evidence states that the opponent processes are present in the later stages of visual perception. There are two types of ganglion cell (which receive information from the cones in the retina, and transfer it towards the brain), these are as Hering described: blueyellow and red-green. When not stimulated they fire axons at a steady rate. The brain realises that a coloured stimulus is present when there is a change in the rate of firing. Such changes include an increase in firing if a red or yellow stimulus is present or a decrease in firing if a green or blue light is present. This theory is able to explain negative after-imaging by assuming that if a certain coloured stimulus is looked at for a long time then the relevant process is activated. Then when the stimulus has gone there is a rebound effect (Buskist, Carlson & Martin, 2004). The opposite process is then activated, resulting in a negative afterimage. This assumption was supported by research from DeValois and DeValois (1975, cited in Eysenck & Keane, 2005). They studied monkeys (who share a similar visual system to

humans). They found opponent-cells in the lateral geniculate nucleus (LGN a relay station between the eye and the brain). These cells behaved in a way consistent with the opponentprocess theory, they were excited by certain wavelengths and inhibited by others. This study not only shows that the opponent-processes exist, but also that they occur later on in the perception process rather than just the early stages (this is also in accordance with Lennies (1998, cited in Eysenck & Keane, 2005) hierarchical model of visual perception).

As the opponent-process theory is concerned with the later stages rather than the earlier stages of colour perception, it makes sense to synthesise the two in order to provide a fuller picture of how we perceive colour. Signals from the three types of cone are sent to the opponent cells via three channels. The black-white channel receives the inhibitory signals from the L- and M-cones. The blue-yellow channel receives inhibitory signals from the Land M-cones and the excitatory signals from the S-cones. Finally, the red-green channel receives inhibitory signals from the S-cones and L-cones and excitatory signals from the Mcones (Eysenck & Keane, 2005).

Together, these theories provide such a comprehensive account of how we perceive colour, that they can also help us to understand deficiencies of colour vision. Protanopia is a lowered sensitivity to long-wavelength light (Reber & Reber, 2001). It can be explained in terms of the L-cones being damaged; therefore the red-green channel cannot be activated. Deuteranopia and tritanopia are similar conditions occurring within the M-cones/red-green channel and S-cones/blue-yellow channel respectively. Despite the theories being accurate enough to explain such conditions, there is one aspect of colour vision that they cannot explain. 7

Colour constancy occurs when we view an object and the wavelengths from the light-source change, but the object appears to remain the same colour. According to the trichromatic and opponent-process theories, a change in wavelength would lead to a change in perceived colour. Theories of colour constancy suggest that context is important when we perceive colour. Research into colour perception is not ecologically valid as often the stimulus presented to participants is a single isolated object. In order to understand how we perceive colours it is necessary to take into account that neighbouring stimuli affect what we see (Finger, 1994).

After consideration of the evidence it appears that a synthesis of the trichromatic and opponent-process theories is the most logical way for us to understand how we perceive colour. However, not only has research into the area been both too artificial and simplistic, but both theories also have serious limitations. From this it seems that in order to understand how we perceive colour, we would benefit from a theory that can encompass all aspects of colour perception, from the first photon reaching the retina to more complex problems such as colour constancy.

References Abramov, I., & Gordon, J. (1994). Colour appearance: On seeing red, or yellow, or green,

or blue. Annual Review of Psychology, 36, 715-729. In Eysenck, M. W., & Keane, M. T. (2005) Cognitive Psychology A Students Handbook (Fifth Ed.). New York: Psychology Press. Blake, R., & Sekuler, R., (2002). Perception (Fourth Ed.). New York: McGraw Hill. p. 621. In Eysenck, M. W., & Keane, M. T. (2005) Cognitive Psychology A Students Handbook (Fifth Ed.). New York: Psychology Press, p. 31. Bowmaker, J. K., Dartnall, H. J. A., & Mollon, J. D. (1983). Human visual pigments: Microspectrophotometric results from the eyes of seven persons. Proceedings of the Royal Society of London Series B, 220, 115-130. In Eysenck, M. W., & Keane, M. T. (2005) Cognitive Psychology A Students Handbook (Fifth Ed.). New York: Psychology Press. Buskist, W., Carlson, N. R., & Martin, G. N., (2004) Psychology (Second Ed.). UK: Pearson Educated Limited. p. 166. Cicerone, C.M., & Nerger, J.L., (1989). The relative number of long-wavelength-sensitive to middle-wavelength-sensitive cones in the human fovea centralis. Vision Research, 29, pp. 115-128. In Eysenck, M. W., & Keane, M. T. (2005) Cognitive Psychology A Students Handbook (Fifth Ed.). New York: Psychology Press. p. 49. DeValois, K. K, & DeValois, R. L. (1975). Neural coding of colour. In Carterette, E. C., &Friedman, M. P. (Eds.), Handbook of Perception, Vol. 5. New York: Academic Press. In Eysenck, M. W., & Keane, M. T. (2005) Cognitive Psychology A Students Handbook (Fifth Ed.). New York: Psychology Press.

Finger, S., (1994). Origins of Neuroscience. USA: Oxford University Press. Fridlund, A. J., Gleitman, H., & Reisberg, D. (2004) Psychology (Sixth Ed.). USA: W. W. Norton & Company, Inc. Helmholtz, H. von (1866) Treatise on physiological optics, Vol. III. New York: Dover [translation published in 1962]. In Eysenck, M. W., & Keane, M. T. (2005) Cognitive Psychology A Students Handbook (Fifth Ed.). New York: Psychology Press. Hofer, H., Singer, B., Williams, D.R., (2005). Different sensations from cones with the same photopigment. J. Vision 5: 44454 . In Kingdom, F. A. A., & Shevell, S.K. (2008) Annual Review of Psychology, Vol. 59: pp. 143-166 Lennie, P. (1998). Single units and visual cortical organisation. Perception, 27, 889-935. In Eysenck, M. W., & Keane, M. T. (2005) Cognitive Psychology A Students Handbook (Fifth Ed.). New York: Psychology Press. Reber, A. S., & Reber, E. (2001). Dictionary of Psychology (Third Ed.). London: Penguin Group. p. 574. Schubert, E.F., (n.d). Adapted from Dowling (1987), Light Emitting Diodes, Chapter 16, Retrieved October 30, 2008 from http://www.ecse.rpi.edu/~schubert/Light-EmittingDiodes-dot-org/chap16/

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