SCI 200 Physical Science Lecture 9 Color & Color Vision Rob Daniell July 21, 2011 Physical vs. Psychological Color Psychological Color • Physical color • Objective • Directly measurable • Based on wavelength • Any “color” can be defined by the relative intensity of light at each wavelength revised 20 Jul 2011 Subjective Indirectly measurable Based on the response of cones and subsequent processing NEiA SCI 200 Lecture 9 Ganglions Brain 2 Physical Color Electromagnetic Spectrum Color vs. wavelength revised 20 Jul 2011 NEiA SCI 200 Lecture 9 3 Physical Color Electromagnetic Spectrum Intensity vs. wavelength revised 20 Jul 2011 NEiA SCI 200 Lecture 9 4 Physical Color Spectroscope Light source Entrance slit Dispersive element (prism or grating) Screen or detector revised 20 Jul 2011 NEiA SCI 200 Lecture 9 5 Physical Color Simplified Grating Spectroscope Project STAR Spectrometer Transmission grating Adjustable scale Do not point directly at sun revised 20 Jul 2011 NEiA SCI 200 Lecture 9 6 Physical Color Diffraction Grating revised 20 Jul 2011 NEiA SCI 200 Lecture 9 7 Physical Color Simplified Grating Spectroscope Project STAR Spectrometer Transmission grating Adjustable scale Do not point directly at sun revised 20 Jul 2011 NEiA SCI 200 Lecture 9 8 Physical Color Wavelength spectra of various light sources Intensity units are relative Gilbert and Haeberli [2007] Am. J. Phys., 75, 313-319. revised 20 Jul 2011 NEiA SCI 200 Lecture 9 9 Physical Color Wavelength spectra of fluorescent light bulbs As seen through Project STAR Spectrometer revised 20 Jul 2011 NEiA SCI 200 Lecture 9 10 Physical Color Discrete spectrum Helium revised 20 Jul 2011 NEiA SCI 200 Lecture 9 11 Physical Color Discrete spectrum: more examples Hydrogen, Sodium, Helium, Neon, Mercury revised 20 Jul 2011 NEiA SCI 200 Lecture 9 12 Physical Color Continuous spectrum White light, sunlight, etc. revised 20 Jul 2011 NEiA SCI 200 Lecture 9 13 Monochromatic vs. Non-monochromatic Colors Monochromatic colors: Consist of a single wavelength Sometimes called “spectral colors” Non-monochromatic colors: 1. A discrete spectrum several discrete wavelengths 2. A continuous spectrum Most colors in nature are nonmonochromatic Example: sunlight revised 20 Jul 2011 NEiA SCI 200 Lecture 9 14 Psychological Color • Physical color as perceived by the human eye and brain • Color perception is mediated by the cones in the retina • There are (usually) three kinds of cones operating – Each cone type responds differently to a specific physical color – The signals from the cones are processed in a non-intuitive way to produce the sensation of color revised 20 Jul 2011 NEiA SCI 200 Lecture 9 15 Psychological Color • Color specification systems: – HSV: Hue, Saturation, Value • Also: – HSL (hue, saturation, lightness) – HSB (hue, saturation, brightness) • Corresponds most closely to human color perception • Preferred by many artists – RGB: (Red, Green, Blue) • Used in additive color systems • Used in many digital graphics applications – Displays – Software – CMYK: (Cyan-Magenta-Yellow-blacK) • Used in subtractive color systems • Used for printing inks, etc. – “Four Color Printing” revised 20 Jul 2011 NEiA SCI 200 Lecture 9 16 Color Vision • HSV: cylinder Hue: perceived color 0°-240° 240°-360° (“purples”) Saturation: Purity of color 0-1 Value: Light intensity 0-1 or black to white (brightest) revised 20 Jul 2011 NEiA SCI 200 Lecture 9 17 Color Vision Another representation of HSL revised 20 Jul 2011 NEiA SCI 200 Lecture 9 18 Trichromacy History Thomas Young (1773-1829) Observed that it only takes three quantities (Hue, Saturation, Value) to specify a color Three output quantities require three input quantities Postulated three kinds of photoreceptors revised 20 Jul 2011 NEiA SCI 200 Lecture 9 19 Trichromacy History (continued) Hermann von Helmholtz (1821-1894) Suggested that Young’s three photoreceptors were Short wavelength Intermediate wavelength Long wavelength Must overlap Monochromatic light of different wavelengths have different colors revised 20 Jul 2011 NEiA SCI 200 Lecture 9 20 Trichromacy Monochromatic light would appear to consist of exactly three colors Response Suppose there were no overlap: 400 S I 500 L 700 600 For example, (above) any monochromatic light source between 400 and 500 nm would appear blue. Yet we know that 450 nm light is a very different shade of blue than 475 nm light revised 20 Jul 2011 NEiA SCI 200 Lecture 9 21 A. Overlap of Response Curves Example: six monochromatic emission lines from atomic Helium Each a different color Conclusion: There must be at least two overlapping cones at each wavelength in the visible region revised 20 Jul 2011 Line spectrum of helium (He) Blue-violet: Blue: Green: Orange: Red-orange: Dark red: NEiA SCI 200 Lecture 9 447.1 nm 471.3 nm 501.5 nm 587.5 nm 706.5 nm 728.1 nm 22 Trichromacy Where do the curves cross? This requires exploring the properties of psychological color revised 20 Jul 2011 NEiA SCI 200 Lecture 9 23 Trichromacy • Complementary colors: • R+CW • G+MW • B+YW • White can be produced by • Broadband light (e.g., sunlight) • Pairs of complementary colors • Stimulate the three kinds of photoreceptors “equally” • An infinite variety of other combinations revised 20 Jul 2011 C = Cyan, M = Magenta, Y = Yellow W = White NEiA SCI 200 Lecture 9 24 Color Perception Mechanisms If Helmholtz is right, how can we determine the actual response curves? A. B. C. D. Overlap of response curves Spectral complementaries Hue discrimination Microspectrophotometry revised 20 Jul 2011 NEiA SCI 200 Lecture 9 25 Trichromacy Where do the curves cross? Consider a monochromatic color at about 430 nm. Stimulates S with a little I Another monochromatic color near 610 nm could stimulate some I and more L to produce white. Also, vice versa revised 20 Jul 2011 NEiA SCI 200 Lecture 9 26 Trichromacy Where do the curves cross? Note that in the region where the Intermediate photoreceptors dominate, no single complementary spectral (monochromatic) color exists No one spectral color can stimulate both the S and the L photoreceptors equally. Empirically, this is the region from 490 nm to 565 nm revised 20 Jul 2011 NEiA SCI 200 Lecture 9 27 Trichromacy So 490 nm and 565 nm represent the crossover points between S and I and between I and L, respectively Between these wavelengths, it takes two additional monochromatic sources to combine with a “green” source to produce white A “blue” source and a “red” source - hence “purple” (or magenta) This has consequences for color mixing (Lecture 10) revised 20 Jul 2011 NEiA SCI 200 Lecture 9 28 Trichromacy Hue discrimination: The difference in wavelength (Δλ, pronounced “delta lambda”) at which two monochromatic sources are barely distinguishable Varies with wavelength Where Δλ is small, the photoreceptor response must be changing rapidly Further Details of the spectral response curves required microspectrophotometry The physical measurement of the amount of light of each wavelength absorbed by each kind of cone Although many cones have been measured this way only three basic types have been found revised 20 Jul 2011 NEiA SCI 200 Lecture 9 29 Trichromacy Cone Mosaic: Simulation based on measured cone densities No “blue” cones in the central fovea! Visual acuity in blue light is less than in green and red light Over the entire retina There are about 100 “red” and “green” cones for every “blue” cone There are about 150 “red” cones for every 100 “green” cones However: Much variation among individuals revised 20 Jul 2011 NEiA SCI 200 Lecture 9 30 Trichromacy Spectral sensitivity of the three types of cones in the human eye Intensity of each wavelength is the same There is considerable overlap among the three cone types Type II & Type III cones have the same sensitivity at about 560 nm Figure 6.4 from text revised 20 Jul 2011 NEiA SCI 200 Lecture 9 31 Trichromacy Spectral sensitivity of Type II (green) cones Two different wavelengths can produce the same response Figure 6.5 from text revised 20 Jul 2011 Using all three types of cones, the four colors can be distinguished. Figure 6.6 from text NEiA SCI 200 Lecture 9 32 Trichromacy • Color vision: – : 3 kinds of cones • Type I: Short (S), beta (β), or blue (B) • Type II: Intermediate (I), gamma (γ), or green (G) • Type III: Long (L), rho (ρ), or red (R) Note that the three kinds of cones do not actually correspond to blue, green, and red. The RGB model is merely a convenient means of representing color. revised 20 Jul 2011 NEiA SCI 200 Lecture 9 33 Color Vision • RGB color system: • Based (loosely) on the three cones of the human eye • Z ~ blue, Y ~ green, X ~ red (even though it peaks shortward of red) revised 20 Jul 2011 NEiA SCI 200 Lecture 9 34 Color Vision • Additive color rules: • • • • R+G+B=W R+G=Y G+B=C R+B=M • Complementary colors: • R+C=W • G+M=W • B+Y=W • Can any 3 colors be combined to produce any other color? • Can R, G, & B be combined to produce any other color? revised 20 Jul 2011 C = Cyan, M = Magenta, Y = Yellow W = White NEiA SCI 200 Lecture 9 35 Color Vision • Red, Green, & Blue can be combined to produce most colors, but some saturated (or nearly saturated) colors cannot be reproduced. • Will be considered in more detail in Lecture 10 C = Cyan, M = Magenta, Y = Yellow W = White revised 20 Jul 2011 NEiA SCI 200 Lecture 9 36 Color Vision • Subtractive color combination: • Filters that absorb or block light of certain colors • Ink or pigments that reflect only certain colors and absorb the others • Primary Subtractive Colors: • Cyan, Magneta, Yellow • Supplemented by Black in “four color printing” • Will be considered in more detail in Lecture 10 C = Cyan, M = Magenta, Y = Yellow K = Black revised 20 Jul 2011 NEiA SCI 200 Lecture 9 37 Trichromacy • Where does “yellow” come from? revised 20 Jul 2011 NEiA SCI 200 Lecture 9 38 Trichromacy or Opponent Colors? Statements: Magenta looks like a mixture of Red & Blue Cyan looks like a mixture of Green & Blue Yellow looks nothing like a mixture of Red & Green revised 20 Jul 2011 NEiA SCI 200 Lecture 9 39 Trichromacy or Opponent Colors? Based on the trichromacy theory We should expect an additive mixture of red and green to give a reddish green (or a greenish red). Instead it gives yellow In fact, it takes four psychological primaries to verbally describe any color Blue, green, yellow, and red Orange looks yellowish red Cyan looks bluish green Purple looks reddish blue Etc. revised 20 Jul 2011 NEiA SCI 200 Lecture 9 40 Opponent Processing When asked to name the color of a spot of spectral (i.e., monochromatic) light, most people give responses similar to those at right Note that there is no “reddish green” or “yellowish blue” revised 20 Jul 2011 NEiA SCI 200 Lecture 9 41 Opponent Processing Yellow and blue seem to oppose each other Red and green also seem to oppose each other How can the three kinds of cones be wired together to produce this kind of color opposition? revised 20 Jul 2011 NEiA SCI 200 Lecture 9 42 Opponent Processing S inhibits y-b and stimulates r-g & w-bk I inhibits r-g and stimulates y-b & w-bk L stimulates all three opponent systems revised 20 Jul 2011 NEiA SCI 200 Lecture 9 43 Opponent Processing Net stimulation of y-b makes the light appear yellowish; net inhibition, bluish Net stimulation of r-g makes the light appear reddish; net inhibition, greenish The w-bk channel conveys brightness information revised 20 Jul 2011 NEiA SCI 200 Lecture 9 44 Opponent Processing There are at least two rival theories for the details of how the three kinds of cones get processed to produce the opponent activity. One theory makes use of lateral inhibition in the form of center-surround antagonism among the various cones Another assumes some kind of filter that narrows the wavelength range accessible to some cones but not others. revised 20 Jul 2011 NEiA SCI 200 Lecture 9 45 Genetics of Color Vision Review: basics of human genetics Each cell in the human body contains 23 pairs of chromosomes The chromosomes are numbered 1 through 22 plus the X and/or Y chromosome In each pair, one comes from the mother, the other from the father. The gender is (mostly) determined by the X and Y chromosomes Females have 2 X chromosomes, one from each parent Males have an X chromosome from their mothers and a Y chromosome from their fathers revised 20 Jul 2011 Other primate species have differing numbers of chromosomes The Great Apes all have 24 pairs Gender is generally determined in the same was as for humans The genes controlling color vision differ among primate species NEiA SCI 200 Lecture 9 46 Genetics of Color Vision Color Vision The number of cone types varies dramatically throughout the Animal Kingdom Mammals Most mammals have only two types of cones – dichromats Short vs. long wavelength: Yellow vs. Blue Red-Green color blind Primates All new world primates are dichromats But see next slide Many old world primates are trichromats Especially monkeys, apes, and humans revised 20 Jul 2011 NEiA SCI 200 Lecture 9 47 Genetics of Color Vision Scientific American, April 2009, The Evolution of Primate Color Vision, pp. 56-63. Some Old World primates (including humans) are trichromats Gene for the short wavelength (“blue”) cone resides on chromosome 7 Genes for the medium wavelength (“green”) cone and the long wavelength (“red”) cone both reside on the X chromosome revised 20 Jul 2011 NEiA SCI 200 Lecture 9 48 Genetics of Color Vision Scientific American, April 2009, The Evolution of Primate Color Vision, pp. 56-63. New World primates are mostly dichromats Gene for the short wavelength (“blue”) cone resides on chromosome 7 Gene for one of the longer wavelength (“green”, “yellow”, or “red”) cones resides on the X chromosome revised 20 Jul 2011 NEiA SCI 200 Lecture 9 49 Genetics of Color Vision Scientific American, April 2009, The Evolution of Primate Color Vision, pp. 56-63. Some female New World primates are trichromats One X chromosome has one of the green, yellow, or red cones The other X chromosome has a different “long wavelength” cone These females can distinguish colors that their dichromat brothers and sisters cannot revised 20 Jul 2011 NEiA SCI 200 Lecture 9 50 Genetics of Color Vision It appears that “new world” dichromacy is the ancestral condition: Among old world primates, a recombination error resulted in both “green” and “red” genes appearing on every X chromosome. Both males and females became trichromats Strong selective advantage, so this system became the norm in Old World primates. revised 20 Jul 2011 NEiA SCI 200 Lecture 9 51 Genetics of Color Vision Every cone cell contains the genes for every cone type Dichromats have two types Trichromats have three types In any particular cone cell, only one gene is actually expressed Mechanism for selection of which gene to express is not known For the genes on the X chromosome, it appears that the choice is random Matrix of “red” and “green” cones is a random distribution So an individual with both a red and a green cone gene on the X chromosome would have both red and green cones. revised 20 Jul 2011 NEiA SCI 200 Lecture 9 52 Genetics of Color Vision Having both red and green cones would give the individual a strong survival advantage It would be much easier to distinguish ripe fruits (yellow, orange, etc.) from unripe fruits (green) It would be much easier to distinguish some predators (e.g., a leopard with a tawny coat) from the leaves or bushes (green) in which it was hiding. The selection pressure was so strong that trichromats have completely displace dichromats among Old World monkeys and apes – and, of course, humans revised 20 Jul 2011 NEiA SCI 200 Lecture 9 53 Genetics of Color Vision Implications for “opponent processing” If the ancestral color processing was dichromacy: It probably involved the opposition of blue cones and the “yellow” (i.e., longer wavelength) cones The advent of trichromacy with the simultaneous appearance of red and green cones on the X chromosome made a second color opposition possible Red vs. green Thus, the psychological colors - blue, green, yellow, and red - may have arisen naturally from the basic distinction between blue cones on the one hand and red, green, and yellow cones on the other. Yellow being a synthesis of the red and green cones revised 20 Jul 2011 NEiA SCI 200 Lecture 9 54 Genetics of Color Vision The ability to distinguish between “blue” cones and the other kinds of cones appears to be “hardwired” into the brain The ability to distinguish between “red” and “green” cones appears to be “learned.” Female mice that have been genetically engineered to have a “green” cone on one X chromosome and a “red” cone on the other can learn to distinguish hues that are indistinguishable to their dichromatic relatives There is evidence that the neural circuitry for distinguishing “red” and “green” cones is the same as that used for spatial vision Detecting boundaries, etc. If so, then “trichromacy can be viewed as a “hobby” of the preexisting spatial vision system.” revised 20 Jul 2011 NEiA SCI 200 Lecture 9 55 Genetics of Color Vision • The trichromatic theory of color vision is based on the three types of cones • However, it has recently been discovered that some people have a rare “yellow” cone – Similar (identical?) to the “yellow” cone in New World Monkeys revised 20 Jul 2011 NEiA SCI 200 Lecture 9 56 Genetics of Color Vision • For males, with only one X chromosome – Standard trichromat: red, green, blue – Non-standard trichromat (rare) • Red, yellow, blue • Yellow, green, blue • For females with two X chromosomes – Standard trichromat: red, green, blue – Tetrachromat: red, green, yellow, and blue • Still rare, but different revised 20 Jul 2011 NEiA SCI 200 Lecture 9 57 Genetics of Color Vision Implications for “tetrachromacy” in some women. If distinguishing between “red” and “green” cones is learned, perhaps distinguishing among “red, yellow, and green” cones is also learned Unfortunately, so far vision tests have not produced conclusive evidence for true tetrachromatic vision, but research is ongoing. revised 20 Jul 2011 NEiA SCI 200 Lecture 9 58 Genetics of Color Vision Genetics of and evolution of color vision Subject of ongoing research Very complex system with lots of threads to unravel revised 20 Jul 2011 NEiA SCI 200 Lecture 9 59 Color Vision Problems • There are various kinds of color vision “anomalies” or “deficiencies” • Some are sex linked, since they involve the red and green (and yellow?) cones on the Xchromosome • Some are more general genetic anomalies revised 20 Jul 2011 NEiA SCI 200 Lecture 9 60 Color Vision Problems • Monochromats: People who see only one color • Relatively rare • Two main types: • Cone monochromats have cones, but only one type is actually functional • Can see under photopic conditions • Rod monochromats lack all cone function • Have difficulty seeing in bright light • Poor visual acuity (no foveal rods) revised 20 Jul 2011 NEiA SCI 200 Lecture 9 61 Color Vision Problems • Dichromats: People who see only two colors (and their combinations) • Two subtypes: • People with only two kinds of functional cones • Three classes, depending on which cone type is nonfunctional • Protanopia: lacking L (red) cones • Deuteranopia: lacking I (green) cones • Tritanopia: lacking S (blue) cones • People for whom one of the opponent color systems is not working • Two classes, depending on which of the two opponent color systems is nonfunctional revised 20 Jul 2011 NEiA SCI 200 Lecture 9 62 Color Vision Problems • Trichromats: People who see all three colors (and their combinations) • Normal trichromats • Slight variations in the cone pigments • Anomalous trichromats • Large variations in cone pigments • Connections between one type of cone and the nerve cells is defective • Protanomalous, deuteranomalous, tritanomalous variations recognized • No sharp boundaries, however revised 20 Jul 2011 NEiA SCI 200 Lecture 9 63 Color Vision Problems • • • Ishihara “Test for Colour-Blindness” (2 sample plates) On left: “Normals” see “26”; Protanopes and some protanomalous observers see only the “6”. Deuteranopes and some deuteranomalous observers see only the “2” On right: Many color deficients can see a serpentine path between the two x’s; Normals cannot. revised 20 Jul 2011 NEiA SCI 200 Lecture 9 64 Summary Three variables or quantities are sufficient to describe any color Trichromacy theory: Developed during 19th century Confirmed and quantified in the 20th century Three kinds of cones: Long, Intermediate, Short Red, Green, and Blue Not the whole story: Opponent Color processing Signals from the three kinds of cones are processed to produce Yellow-Blue and Red-Green opponents revised 20 Jul 2011 NEiA SCI 200 Lecture 9 65 Summary Original Opponent Color processing seems to involve Blue vs. Yellow (Short vs. “Long”) Still the dominant form of color vision in New World monkeys Also most mammals In Old World Primates A recombination error enabled Red-Green opponent colors Apparently “learned” behavior Also enabled three color processing and the rich colors visible to humans There appear to be three kinds of “intermediate” and “long” cones “yellow” cones are relatively rare Some women have four kinds of cones May be able to sense an even richer range of colors revised 20 Jul 2011 NEiA SCI 200 Lecture 9 66 Summary Various kinds of color vision “anomalies” or “deficiencies” Trichromats: normal color vision Except some people have deficient cones of one or more colors Dichromats: can see only two colors Monochromats: only a single color Some have no cones, so have poor visual acuity and difficulty seeing in bright light revised 20 Jul 2011 NEiA SCI 200 Lecture 9 67 Homework & Lab Read Chapter 6 in textbook Homework Packet 9: Due Thursday, July 28 Next Lab: Lab 6: Water Prism Thursday, July 21 revised 20 Jul 2011 NEiA SCI 200 Lecture 9 68