Chapter 8: Color Vision
Introduction to Color Perception
Our perception of color involves complex physiological and psychological processes that enable us to differentiate between various wavelengths of light, even though light itself does not inherently possess color. In this chapter, we examine the intriguing world of color vision, exploring the major concepts, physiological mechanisms, and color mixing theories that underlie our ability to perceive and appreciate different colors.
The Nature of Color
Visible light, a small portion of the electromagnetic spectrum, is what we perceive as color. The diverse colors we encounter, such as red, green, and blue, are our psychological interpretations of different wavelengths of light. For example, when we see a 750-nanometer wavelength of light, we label it as "red," but in reality, it's simply a specific wavelength of light with no intrinsic color. Despite this, color plays a significant role in our lives, influencing various aspects, including art, design, psychology, and even tourism.
Color’s Impact on Perception and Behavior
Color has a profound psychological impact on our decisions and behaviors (Elliot & Maier, 2014). For instance, it affects our choices in food and beverages; we tend to be wary of consuming foods with unusual or unexpected colors. This phenomenon was notably exemplified in the 1990s when the market introduced clear drinks like "Crystal Pepsi" and clear alcoholic beverages like "Zima," both of which were met with skepticism, in part because their appearance violated consumers' expectations. Even today, unusual colors in food and beverages can be off-putting to many individuals.
Color constancy illustrates our tendency to perceive the color of an object as relatively consistent, regardless of the lighting conditions (Land & McCann, 1971). For example, a banana under green- tinted lighting will still appear yellow to us due to our brain's ability to account for the color shifts caused by the surrounding illumination.
Early Research on Light and Color Vision
Isaac Newton's groundbreaking experiments revealed that white light is composed of various wavelengths of light (Newton, 1704/1952). This discovery challenged the prevailing notion that white light was pure and sent from the heavens. Instead, Newton demonstrated that white light could be broken down into its constituent colors when passed through a prism. He also showed that these colors could be recombined to form white light. This work laid the foundation for our understanding of color vision.
Trichromatic Theory of Color Vision
One of the key theories explaining color vision is the trichromatic theory, which suggests that we have three types of color receptors, or cones, in our eyes (Young, 1802; Helmholtz, 1867/1962). These cones are sensitive to specific portions of the electromagnetic spectrum: short-wavelength cones for blue, medium-wavelength cones for green, and long-wavelength cones for red. By combining the signals from these three types of cones, our brain creates the perception of a wide range of colors.
Trichromatic theory is supported by the fact that people can match colors by adjusting the intensities of three primary colors: red, green, and blue (Guild, 1931). This concept is central to additive color mixing, where colors are created by adding different wavelengths of light together. When red, green, and blue light are combined in various proportions, we can produce a broad spectrum of colors.
Although trichromatic theory explains how cone photoreceptors encode wavelength, it does not explain several perceptual phenomena, including negative afterimages and the fact that people do not perceive reddish-green or bluish-yellow colors. These observations motivated the development of opponent-process theory.
Additive and Subtractive Color Mixing
Additive Color Mixing
Additive color mixing occurs when colored lights are combined to produce new colors. It is commonly used in digital displays, such as computer screens and television monitors. In additive color mixing, different wavelengths of light are added together, and the result becomes lighter as more light is added. This process is analogous to how colors are mixed with flashlights or projectors (you might have experience with this if you’ve worked on lighting in a theater).
The primary colors used in additive color mixing are red, green, and blue (RGB). By varying the intensities of these three colors, we can create a wide range of colors, including white. Notably, when red and green light are mixed at full intensity, they produce yellow, demonstrating the additive nature of this color mixing process. You can observe this effect by looking at red and green dots from a distance (see Figure 8.1). When the two colors are intermixed, the resulting display appears yellow. Artists like Seurat have used this effect in a technique called pointillism.
Figure 8.1
Small red, green, and blue dots are presented separately and in different combinations to illustrate additive color mixing. When viewed from a sufficient distance, the visual system spatially integrates the individual dots, causing mixtures of red and green to appear yellow, red and blue to appear magenta, green and blue to appear cyan.
"Additive color mixing." by Kahan, T.A. is licensed under CC BY-NC-SA 4.0
Subtractive Color Mixing:
Subtractive color mixing involves mixing pigments or dyes to create colors. This method is commonly used in color printing. Unlike additive color mixing, subtractive color mixing starts with a surface that reflects all colors (usually white) and subtracts specific colors by absorbing them.
The primary colors used in subtractive color mixing are cyan, magenta, and yellow (CMY). When these pigments are mixed together in various proportions, they subtract or absorb certain wavelengths of light, resulting in different colors. For example, mixing cyan and magenta pigments subtracts red and green wavelengths, leaving blue light to be reflected, creating a blue color. When all three primary pigments are mixed together in equal amounts, they subtract all colors, resulting in a dark or black appearance. If you have an inkjet printer then you might be more familiar with subtractive color mixing.
Figure 8.2
In subtractive color mixing, blue and yellow pigments combine to produce the perception of green. Blue pigment absorbs primarily red wavelengths, while yellow pigment absorbs primarily blue wavelengths. When the pigments are mixed, only the wavelengths that both pigments reflect—primarily green—remain to be reflected to the eye. Unlike additive color mixing, combining pigments removes (subtracts) portions of the visible spectrum rather than adding light.
"Subtractive color mixing." by Kahan, T.A. is licensed under CC BY-NC-SA 4.0
Opponent Process Theory
While trichromatic theory explains how we perceive a wide range of colors, another theory, known as opponent process theory, complements our understanding of color vision (Hering, 1878/1964). Opponent process theory posits that opponent color coding occurs in retinal ganglion cells, cells of the lateral geniculate nucleus (LGN), and throughout the visual cortex. In this theory, red and green are considered opposing colors, as are blue and yellow.
To understand what "opponent" means, imagine a seesaw with red on one end and green on the other. When red light is present, the seesaw tips toward red and away from green. When green light is present, it tips in the opposite direction. The same idea applies to blue and yellow. In other words, a single opponent cell does not independently signal both colors in a pair. Instead, it signals which color is stronger. For example, a red–green opponent cell increases its firing rate when red light is detected but decreases its firing rate when green light is detected. Likewise, a blue–yellow opponent cell increases its firing to blue and decreases its firing to yellow, or vice versa. Opponent processing does not replace trichromatic theory (Hurvich & Jameson, 1957). Rather, the visual system first detects light using three types of cone photoreceptors (L, M, and S cones) and then combines those cone signals into opponent channels that compare red versus green and blue versus yellow. This second stage of processing helps the brain distinguish colors and explains phenomena such as color afterimages.
Figure 8.3
Examples of single-opponent color receptive fields. In the left receptive field, blue light presented in the center excites the neuron (+B), whereas yellow light presented in the surrounding region inhibits its activity (−Y). In the right receptive field, red light in the center excites the neuron (+R), while green light in the surrounding region inhibits its activity (−G).
"Receptive fields of ganglion cells.” by Kahan, T.A. is licensed under CC BY-NC-SA 4.0
Physiology and opponent process theory
This theory is supported by the fact that the receptive fields of many ganglion cells and LGN cells are opponent for color in this same arrangement (De Valois et al., 1966). For example, blue light might increase the firing rate of a cell when presented in a central region while yellow light inhibits the activity in a surrounding region (i.e.,+B/-Y). Similarly, red light might increase the firing rate of a cell when presented in a central region while green light inhibits the activity in a surrounding region (i.e., +R/-G). Opponent color responses arise from neural circuits that compare the activity of different cone types through excitatory and inhibitory connections. This comes about because of the way in which a three- cone system is wired (Dacey, 2000). When viewing the wiring diagram in the figure an open Y connection is facilitatory and a perpendicular T connection is inhibitory. It is also important to understand that red and green sum in an additive manner to create yellow. Here, the amacrine layer of cells (A) is hypothesized to play a critical role in summing the activity from mid and long wavelength cones (to create the perception of yellow).
Figure 8.4
Long- (L), medium- (M), and short-wavelength (S) cone photoreceptors combine through retinal bipolar, amacrine, and ganglion cells to create opponent color channels. Red–green opponency is produced by reciprocal L- and M-cone inputs, generating +L/−M and −L/+M ganglion cells. Blue–yellow opponency is produced by combining L- and M-cone signals into a yellow channel before comparing it with S-cone activity , which then interacts antagonistically with S-cone input to generate +Y/−B and −Y/+B ganglion cells. Green arrows indicate excitatory connections, and red Ts indicate inhibitory connections. The diagram is intended to illustrate the computational logic of opponent processing rather than depict the exact anatomical circuitry of the retina.
"Blue, green, and red cones.” by Kahan, T.A. is licensed under CC BY-NC-SA 4.0
Figure 8.5
If you stare at the eye of the frog in the upper image for an entire minute and then look quickly at the eye of the frog in the lower image, what do you see? Does it look, very briefly, like the frog is green and the banana is yellow? This happens because the upper image is shown in red(ish) and blue(ish) and when you look at the black and white image you perceive the opponent colors (green and yellow). Note: banana for scale.
"Frog with Banana.” by Kahan, T.A. is licensed under CC BY-NC-SA 4.0
If a ganglion cell is sensitive to red and green, it increases its firing when exposed to one color and decreases its firing when exposed to the opposing color. After staring at one color for a prolonged period, the opponent cells responding to that color become temporarily less responsive (they adapt) (Webster, 2015). When you then look at a white or achromatic surface, all cone types are stimulated approximately equally. However, because one member of the opponent pair has adapted, the balance shifts toward the opposite channel. As a result, you briefly perceive the opponent color as an afterimage. For example, staring at a red image produces a green afterimage, while staring at a blue image produces a yellow afterimage (see Figure 8.5) (Hurvich & Jameson, 1957).
Color Processing in the Primary Visual Cortex
Now, let's explore how the human brain processes color information. We've already seen that cone photoreceptors detect different ranges of wavelengths of light. The next stage of color processing begins in the retina, where signals from the cones are transmitted by retinal ganglion cells through the optic nerve to the lateral geniculate nucleus (LGN) of the thalamus. From there, the information is relayed to the primary visual (striate) cortex and then to higher visual areas, where it contributes to our conscious perception of color.
Many retinal ganglion cells and LGN neurons exhibit color opponency, meaning that they respond in opposite ways to different colors. Single-unit recording studies showed that these neurons increase their firing rate when stimulated by one color and decrease their firing rate when stimulated by its opponent color (De Valois et al., 1966; Derrington et al., 1984). For example, a neuron may be excited by red light but inhibited by green light, or excited by blue light but inhibited by yellow light. These opponent responses arise because the neurons compare signals from different cone types rather than simply measuring the amount of light (De Valois et al., 1966; Dacey, 2000).
Most color-opponent neurons also have center-surround receptive fields (Wiesel & Hubel, 1966). For example, the center of a receptive field may be excited by red light while the surrounding region is inhibited by green light. This center-surround organization enhances color contrast and helps the visual system distinguish boundaries between objects. As color-opponent signals are transmitted through the LGN and processed by the visual cortex, they are integrated with information about form, depth, and motion to produce our rich perception of the colorful world around us (Livingstone & Hubel, 1988; Gegenfurtner & Kiper, 2003).
The information is then relayed to the primary visual cortex (striate cortex), where complex neural processing occurs. Within the striate cortex, we find the so-called "blobs" of cells (Livingstone & Hubel, 1984). These blobs are specialized regions where there are many neurons that are selective for color. Within the blob regions two classes of cells have been identified that respond to colors.
The first class, denoted as single-opponent cells, exhibit response patterns that closely resemble those observed in ganglion cells and cells within the lateral geniculate nucleus (LGN). single-opponent cells exhibit a center-surround arrangement, where certain color pairs are in opposition. For example, in one scenario, the presence of red in a central region is excitatory and green in a surrounding region is
inhibitory (+R/-G).
In contrast, double-opponent cells exhibit a slightly different response profile and there are two types of double-opponent cells (Johnson et al., 2001). For one of these, a specific color, such as red, may evoke excitatory responses in a particular region, while the opposing color, green, elicits inhibitory responses within the same spatial region, rather than in the surrounding area. For the other type, known as "double opponent" cells, the central region responds in an excitatory manner to one color while inhibiting responses to its opposing color (e.g., +R-G). Then, in the surrounding region, double opponent cells respond to light in the opposite manner (e.g., +G-R).
Figure 8.6
Example of a double-opponent receptive field. In the central region, red light excites the neuron (+R) while green light inhibits it (−G). In the surrounding region, the response is reversed: green light excites the neuron (+G) and red light inhibits it (−R). Because both the center and surround exhibit chromatic opponency, double-opponent neurons respond strongly to color contrast at edges and borders. They are thought to play an important role in detecting color boundaries.
"Example receptive field for a double opponent cell.” by Kahan, T.A. is licensed under CC BY-NC-SA 4.0
Color Vision Deficiencies
Color vision deficiencies, commonly known as color blindness, can result from various genetic factors. The most common types are red-green deficiencies, which affect the ability to distinguish between these two colors accurately. These deficiencies are more prevalent in males because they involve genes located on the X chromosome (Nathans et al., 1986).
Color vision deficiencies, often referred to as color blindness, are usually inherited genetic conditions. These conditions stem from anomalies in the X chromosome. Because males have one X chromosome (XY) and females have two (XX), the way color vision deficiency is inherited and expressed differs between genders.
For a male (XY) to exhibit color vision deficiency, the single X chromosome he possesses must carry the gene responsible for the condition. Since males have only one X chromosome, there's no second X chromosome to provide a normal gene that could override the deficient one.
For a female (XX) to show color vision deficiency, both of her X chromosomes need to carry the gene for the condition. If only one of the X chromosomes has the gene, the other X chromosome can compensate, and she typically will not exhibit a color vision deficiency. This scenario makes color vision deficiencies less common in females than in males, as a female would need to inherit a color vision deficiency from both parents, while a male only needs to inherit this from his mother.
Punnett Square Example: A Punnett square is a diagram that is used to predict the genetic makeup of the offspring from two parents.
Let's consider a scenario to illustrate this: The gene for normal color vision on the X chromosome will be represented as "XN" (where "N" denotes not problematic for color vision). The gene for color vision deficiency on the X chromosome will be represented as "Xp" (where "p" denotes problematic for color vision deficiency). The Y chromosome will be represented as "Y" since it does not carry a gene for this trait.
Scenario: A woman who is a carrier for a color vision deficiency ("XNXp") has children with a man who has normal color vision (XNY). Mother's genotype: XNXp Father's genotype: XNY. To predict the possible genetic outcomes for their children, we can set up a Punnett square.
Figure 8.7
Punnett square illustrating the inheritance of an X-linked recessive red–green color vision deficiency when the mother is a carrier (XᴺXᶜ) and the father has normal color vision (XᴺY). Each pregnancy has an equal probability of producing one of four genotypes: a daughter with normal color vision (XᴺXᴺ), a carrier daughter (XᴺXᶜ), a son with normal color vision (XᴺY), or a son with a color vision deficiency (XᶜY). Thus, none of the daughters are expected to have a color vision deficiency, although half are carriers, while half of the sons are expected to inherit the condition.
"Punnet square for color vision deficiency" by Kahan, T.A. is licensed under CC BY-NC-SA 4.0
From this Punnett square, we can infer:
There's a 25% chance (1 out of 4) the child will be a girl with normal color vision (top left of table). There's a 25% chance (1 out of 4) the child will be a girl who is a carrier like her mother (bottom left of table). There's a 25% chance (1 out of 4) the child will be a boy with normal color vision (top right of table). There's a 25% chance (1 out of 4) the child will be a boy with color vision deficiency (bottom right of table).
The most common types of red-green color vision deficiencies are deuteranopia (lack of functional green cones) and protanopia (lack of functional red cones). Individuals with these deficiencies see the world with altered color perception compared to those with normal color vision (Neitz & Neitz, 2011). For them, red and green objects may appear similar or indistinguishable.
Figure 8.8
Illustration of how an Ishihara color vision test plate may be perceived differently depending on an individual's color vision (Ishihara, 1917/1972). A person with typical color vision might perceive the left plate as containing the number 8, whereas a person with a red–green color vision deficiency might perceive the number 3 (shown on the right). These images are intended to demonstrate how color vision deficiencies can alter the appearance of Ishihara test plates. They are provided for illustrative purposes only and should not be used for diagnosis, as the colors displayed may vary across monitors, mobile devices, printers, and viewing conditions.
"Eight Ishihara charts for testing color blindness, Europe" by Wellcome Trust is licensed under CC BY 4.0
Color Phenomena and Illusions
Our perception of color is influenced not only by the physical properties of objects but also by contextual factors. Some of these phenomena include:
- Color Constancy: Our brain adjusts for changes in lighting conditions to maintain a consistent perception of an object's color (Land & McCann, 1971; Brainard & Freeman, 1997).
Figure 8.9
A white T-shirt might appear orange while inside a translucent orange bag because sunlight passing through the bag colors the illumination reaching the shirt. Although the light reflected from the shirt is strongly orange, observers may still perceive the shirt as white because the visual system compensates for the color of the illumination (color constancy).
"Color constancy.” by Kahan, T.A. is licensed under CC BY-NC-SA 4.0
- Dress Illusion: The infamous dress photograph (see Figure 8.10) highlights how different assumptions about the scene's illumination can lead to variations in color perception among individuals (Brainard & Hurlbert, 2015). Under normal viewing conditions, the visual system maintains color constancy by estimating the color of the light illuminating an object and mentally discounting (or subtracting) that illumination to recover the object's true color. The dress photograph is ambiguous because the lighting cues are unclear, leading different observers to make different assumptions about the illumination (Lafer-Sousa et al., 2015). If an observer assumes the dress is illuminated by warm, yellowish light, the visual system discounts the yellow illumination, making the dress appear blue and black. In contrast, if the observer assumes the dress is illuminated by cool, bluish light, the visual system discounts the blue illumination, causing the dress to appear white and gold. Thus, the same image can produce dramatically different color perceptions depending on how the brain interprets the lighting in the scene. You can see this same effect in Kahan's Bates Bobcat Illusion (see Figure 8.11). To try the demonstration, click the link to open a Google document. Then go to File and select Make a copy. Once you've opened your own copy in Google Drive, you'll be able to interact with the bobcat and see for yourself how changing the apparent illumination changes the perceived colors of the image.
Figure 8.10
The famous "dress" image, which illustrates that color perception depends not only on the wavelengths reaching the eye but also on the brain's interpretation of the scene's illumination. Some viewers perceive the dress as white and gold, whereas others perceive it as blue and black. These differing perceptions arise because the visual system makes different assumptions about the color of the ambient lighting and discounts that illumination when estimating the object's true colors.
"The Dress" by Swiked
Figure 8.11
Example of Kahan's Bates Bobcat illusion. The rectangular region of the bobcat on the left is identical in color to the rectangular region of the bobcat on the right but should appear black and blue on the left (because the background lighting is golden) and should appear gold and white on the right (because the background lighting is blue). You can prove this to yourself using this online demonstration.
"Example of Kahan's Bates Bobcat illusion.” by Kahan, T.A. is licensed under CC BY-NC-SA 4.0
- Benham's Top: A rotating monochromatic pattern can create the illusion of colors (Benham, 1894). The most common explanation for this is that the rotation stimulates the three types of cones to fire at slightly different rates.
Figure 8.12
Top view of Benham's top, a black-and-white pattern that produces the illusion of color when rotated. Although the disk contains only black and white, many observers perceive faint red, green, blue, or yellow hues while it is spinning. These illusory colors are thought to arise from differences in the timing of neural responses to changing patterns of light rather than from colored light reaching the eye. The specific colors perceived vary among observers and depend on the speed and direction of rotation.
A depiction of the surface of Benham’s top. "Benham disc" by Smartse is licensed under CC BY-SA 3.0
- Illusory Color Contours: Specific visual patterns can create the perception of colors where none exist, illustrating the complexity of our visual processing (Day, 1983).
Figure 8.13
Four circular ring patterns are arranged so that their cyan segments define the corners of an invisible square. Although no cyan color or edges are physically present within the central region, many observers perceive a faint cyan square with distinct boundaries. This illusion demonstrates that the visual system can interpolate both color and contours from surrounding visual information, creating the perception of surfaces and edges that are not actually present.
"Illusory color contours.” by Kahan, T.A. is licensed under CC BY-NC-SA 4.0
V4 and Color Perception
A large body of research indicates that the V4 region of the brain plays a pivotal role in color perception (Zeki, 1973). Studies utilizing neuroimaging techniques, such as functional magnetic resonance imaging (fMRI) and electrophysiological recordings, have consistently shown heightened activity in V4 when individuals are exposed to different colors, indicating its involvement in processing and discriminating various hues. Furthermore, experiments involving selective V4 lesions in animal models have demonstrated significant impairments in color discrimination tasks, providing compelling evidence for the critical role of V4 in the neural underpinnings of color perception (Heywood et al., 1992).
Figure 8.14
A side view of the human brain. V4 appears in the bottom right of this image.
"Cortex functional areas" by Drking1234 is licensed under CC BY-SA 4.0
Conclusions
Color vision is a complex interplay between the eye's cone cells, the brain's processing, and external environmental factors. While most people experience color vision in a similar way, variations in cone sensitivities and visual context can lead to diverse and sometimes surprising color perceptions.
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