Vision A Recommended literature 1. Kandel ER, Schwartz JH, Jessel TM (2000) Principles of Neural Science, McGraw-Hill, Ch. xx. 2. Berne EM, Levy MN, Koeppen BM, Stanton BA (2004) Physiology, Mosby, Ch. 8. 3. Schmidt RF, Lang F, Thews G (2005) Physiologie des Menschen, Springer, Ch. 18. Q1 How do the photoreceptors work? Photoreceptor function Light is a form of electromagnetic irradiation f I Photons move in a straight way with a velocity of aprox. 3x105 km/s The visible light only comprises a small range of wavelenghts (400 to 700 nm). The basic measure of light intensity is candela (cd). The light density (cd/m2) of the human environment varies from10-6 cd/m2 to 107 cd/m2. Photoreceptors are activated by the absorption of photons. We have two classes of photoreceptors- the rods and the cones, each with different photopigments Rods - rhodopsin Cones – 3 types of iodopsins rhodopsin STL2004-18-11 Photoreceptor function KS-2001-24-7 Photoreceptor function The retina and the following visual representation areas all have clear cut layers Photoreceptor function KS2001-24-9 In contrast to other receptors, light does not depolarize, but it hyperpolarizes the receptor cell membrane icCa icCa Hyperpolarization increases with light intensity When photopigments absorb light, they are „boosted“ to a higher energy level, which is associated with a series of chemical changes. The photopigments are bleached. Their amount decreases with light intensity. STL2004-18-11 Photoreceptor function The adjustment to decreasing light intensity is determined by measuring the intensity threshold of vision. The reciprocal of the intensity threshold is a measure os sensitivity of perception The dark adaption curve reflects, first of all, the recovery of the receptor pigments. In cases of vitamin A deficiency one observes night blindness due to a rhodopsin deficit. STL-2004-18-16 Photoreceptor function Q2 How does visual acuity relate to the different regions of the retina? Visual acuity With the help of a mirror you can see the retina vessels fovea centralis blind spot Visual acuity The spatial resolution (visus) reflects the ability to discriminate two stimulation points as being separate (= spatial threshold). The visus can also be expressed as the reciprocal of the respective spatial angle. Visual acuity STL2004-18-21 The visus (visual acuity) differs in dependence on the coordinates of the image on the retina 1‘=1/60o (~1.5 mm at a distance of 5 m) The visual acuity is higher if an object is imaged on the fovea, but in this case the light intensity must be higher KS2001 -24-14 Visual acuity D = spatial resolution (visus) = 1/α Visual acuity One can separate 2 stimuli, if 2 receptors have another receptor in between that absorbs less light Between the fovea and the retinal periphery there are substantial differences in: - receptor density - degree of convergence Significance for spatial resolution Lower receptor density, higher convergence Higher receptor density, lower convergence 3 degrees 0.3 degrees Q3 How do retinal ganglion cells contribute to the perception of contrast? Contrast perception By analyzing sensory illusions one can better understand the mechanisms of contrast perception Contrast perception Contrast perception is based on the antagonistic organization of the receptive fields of retinal ganglion cells On-center RGC Off-center RGC RGCs are classified according to the response to center stimulation Contrast perception The retina contains 7 classes of cells that differ in their structure, position, connectivity, transmitters and transmitter receptors Signaling path of on-center RGCs Note: only ganglion cells and some amacrine cells in the ganglion cell layer have axons and generate full action potentials; all other cells generate gradual potentials Contrast perception The antagonistic organization of receptive fields requires inhibitory activity of horizontal cells In the absence of this inhibition, the excitatory center expands and the spatial threshold increases KS2001 -24-11 Contrast perception The antagonistic receptive field organization of the retinal ganglion cells is the prerequisite for the perception of contrast KS2001 -24-12 Contrast perception Q4 How are colors discriminated? Color vision Color vision largely contributes to the perception of contrast and the recognition of visual objects Color vision False color coding of biological parameters Control Pulse 3 ms, 2 μ A Example: ic Ca-concentration in individual inhibitory synapses Red: Ca concentration is high (10-6 mol/l) Blue: Ca concentration is low (10-7 mol/l) Kirischuk, Veselovsky, Grantyn, Pflüg. Arch. (Neurotechniques) 1999, 438, 716-724 Color vision Color vision 400 500 600 wavelenght The human visual system discriminates about 200 wavelenghts (λ) as separate colors 700nm Color vision The perception of color not only varies with hue (λ), but also with saturation and darkness 200 Hue (wavelenght λ) 20 Saturation (weighting factor) 500 Darkness (fraction of black) Discrimination of a total of 2x106 color values Color vision Any color can be produced (A) by mixing light of defined wavelenght or (B) by removing light of defined wavelenght from white light Colored projected light: monochromatic light sources Additive color mixture Colorless: When all are λ are offered simultaneously (white) Mixing of colors for painting: color pigments as l-specific filters Subtractive color mixture Colorless: When filter excludes the entire visible spectrum (black) Color vision Science, media and industry require defined colors, that can be composed from defined primary colors. The commonly used color models for digital imaging are RGB (additive, screen) and CMYK (subtractive, printed media) Color vision Red, Green, Blue Cyan, Magenta, Yellow, BlacK Color vision Color charts (LUT, look-up tables) provide a set of defined colors Example: Corel-Green STL2004 -1 Color vision The human eye is not equally transparent for all colors; and the visual system is not equally sensitive for different wavelenghts Note: there is an age dependency of transmission of blue Color vision Two theories of color vision: A - trichromatic theory (Helmholtz, Young) B – opponent color theory (Hering) These theories apply to particular components of the visual system: A – for level of photoreceptors B – for all following levels Color vision Concerning A: Humans are equipped with 3 types of clones, each containing one of 3 photopigments Color vision In a color mixing test, the color hypochromat adds more of the color that he sees worse Anomalous trichromat: has three photopigments, but one of them with changed absorbtion characteristics Dichromat: one pigment is missing - Protanopy (red pigment is missing) - Deuteranopy (green pigment is missing) - Tritanopyy (red pigment is missing) Red-green-defect: (8% of US population) More frequent in men, since responsible genes are in the X-chromosome Color vision Concerning B: Green-red and blue –yellow are opponent colors. They induce an opposite responses in ganglion cells Red-on-ganglion cell From configuration A follows B-D Please note: With regard to color vision there are big differences among animal species! Color vision The perception of contrast is already enhanced by dichromatic vision, because the 2 types of cones can respond to the stimuli at different wavelenghts as to stimuli of different intensity The functionality of the retina can be examined by recording an electroretinogram (ERG) KS2001 -24-13 Methods
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