Light

As with auditory stimuli, light travels in waves. The compression waves that compose sound must travel in a medium—a gas, a liquid, or a solid. In contrast, light is composed of electromagnetic waves and needs no medium; light can travel in a vacuum (Figure 1). The behavior of light can be discussed in terms of the behavior of waves and also in terms of the behavior of the fundamental unit of light—a packet of electromagnetic radiation called a photon. A glance at the electromagnetic spectrum shows that visible light for humans is just a small slice of the entire spectrum, which includes radiation that we cannot see as light because it is below the frequency of visible red light and above the frequency of visible violet light.

Certain variables are important when discussing perception of light. Wavelength (which varies inversely with frequency) manifests itself as hue. Light at the red end of the visible spectrum has longer wavelengths (and is lower frequency), while light at the violet end has shorter wavelengths (and is higher frequency). The wavelength of light is expressed in nanometers (nm); one nanometer is one billionth of a meter. Humans perceive light that ranges between approximately 380 nm and 740 nm. Some other animals, though, can detect wavelengths outside of the human range. For example, bees see near-ultraviolet light in order to locate nectar guides on flowers, and some non-avian reptiles sense infrared light (heat that prey gives off).

Figure 1. In the electromagnetic spectrum, visible light lies between 380 nm and 740 nm. (credit: modification of work by NASA)

Wave amplitude is perceived as luminous intensity, or brightness. The standard unit of intensity of light is the candela, which is approximately the luminous intensity of one common candle.

Light waves travel 299,792 km per second in a vacuum, (and somewhat slower in various media such as air and water), and those waves arrive at the eye as long (red), medium (green), and short (blue) waves. What is termed “white light” is light that is perceived as white by the human eye. This effect is produced by light that stimulates equally the color receptors in the human eye. The apparent color of an object is the color (or colors) that the object reflects. Thus a red object reflects the red wavelengths in mixed (white) light and absorbs all other wavelengths of light.

Anatomy of the Eye

The eye itself is a hollow sphere composed of three layers of tissue. The outermost layer is the fibrous tunic, which includes the white sclera and clear cornea. The sclera accounts for five sixths of the surface of the eye, most of which is not visible, though humans are unique compared with many other species in having so much of the “white of the eye” visible (Figure 2). The transparent cornea covers the anterior tip of the eye and allows light to enter the eye.

The middle layer of the eye is the vascular tunic, which is mostly composed of the choroid, ciliary body, and iris. The choroid is a layer of highly vascularized connective tissue that provides a blood supply to the eyeball. The choroid is posterior to the ciliary body, a muscular structure that is attached to the lens by zonule fibers. These two structures bend the lens, allowing it to focus light on the back of the eye. The lens is dynamic, focusing and re-focusing light as the eye rests on near and far objects in the visual field. The ciliary body stretches the lens flat or allows it to thicken, changing the focal length of light coming through it to focus it sharply on the retina. With age comes the loss of the flexibility of the lens, and a form of farsightedness called presbyopia results. Presbyopia occurs because the image focuses behind the retina. Presbyopia is a deficit similar to a different type of farsightedness called hyperopia caused by an eyeball that is too short. For both defects, images in the distance are clear but images nearby are blurry. Myopia (nearsightedness) occurs when an eyeball is elongated and the image focus falls in front of the retina. In this case, images in the distance are blurry but images nearby are clear.

Figure 2: Correcting abnormalities in light refraction in the eye. This work by Cenveo is licensed under a Creative Commons Attribution 3.0 United States (http://creativecommons.org/licenses/by/3.0/us/).

Overlaying the ciliary body, and visible in the anterior eye, is the iris—the colored part of the eye. The iris is a smooth muscle that opens or closes the pupil, which is the hole at the center of the eye that allows light to enter. The iris constricts the pupil in response to bright light and dilates the pupil in response to dim light.

The innermost layer of the eye is the neural tunic, or retina, which contains the nervous tissue responsible for photoreception. The eye is also divided into two cavities: the anterior cavity and the posterior cavity. The anterior cavity is the space between the cornea and lens, including the iris and ciliary body. It is filled with a watery fluid called the aqueous humor. The posterior cavity is the space behind the lens that extends to the posterior side of the interior eyeball, where the retina is located. The posterior cavity is filled with a more viscous fluid called the vitreous humor. The retina is composed of several layers and contains specialized cells for the initial processing of visual stimuli. The photoreceptors (rods and cones) change their membrane potential when stimulated by light energy. The change in membrane potential alters the amount of neurotransmitter that the photoreceptor cells release onto bipolar cells in the outer synaptic layer. It is the bipolar cell in the retina that connects a photoreceptor to a retinal ganglion cell (RGC) in the inner synaptic layer. There, amacrine cells additionally contribute to retinal processing before an action potential is produced by the RGC. The axons of RGCs, which lie at the innermost layer of the retina, collect at the optic disc and leave the eye as the optic nerve (see Figure 3). Because these axons pass through the retina, there are no photoreceptors at the very back of the eye, where the optic nerve begins. This creates a “blind spot” in the retina, and a corresponding blind spot in our visual field.

Note that the photoreceptors in the retina (rods and cones) are located behind the axons, RGCs, bipolar cells, and retinal blood vessels. A significant amount of light is absorbed by these structures before the light reaches the photoreceptor cells. However, at the exact center of the retina is a small area known as the fovea. At the fovea, the retina lacks the supporting cells and blood vessels, and only contains photoreceptors. Therefore, visual acuity, or the sharpness of vision, is greatest at the fovea. This is because the fovea is where the least amount of incoming light is absorbed by other retinal structures (see Figure 3).

Figure 3. Structure of the Eye The sphere of the eye can be divided into anterior and posterior chambers. The wall of the eye is composed of three layers: the fibrous tunic, vascular tunic, and neural tunic. Within the neural tunic is the retina, with three layers of cells and two synaptic layers in between. The center of the retina has a small indentation known as the fovea.

As one moves in either direction from this central point of the retina, visual acuity drops significantly. In addition, each photoreceptor cell of the fovea is connected to a single RGC. Therefore, this RGC does not have to integrate inputs from multiple photoreceptors, which reduces the accuracy of visual transduction. Toward the edges of the retina, several photoreceptors converge on RGCs (through the bipolar cells) up to a ratio of 50 to 1.

The difference in visual acuity between the fovea and peripheral retina is easily evidenced by looking directly at a word in the middle of this paragraph. The visual stimulus in the middle of the field of view falls on the fovea and is in the sharpest focus. Without moving your eyes off that word, notice that words at the beginning or end of the paragraph are not in focus. The images in your peripheral vision are focused by the peripheral retina, and have vague, blurry edges and words that are not as clearly identified. As a result, a large part of the neural function of the eyes is concerned with moving the eyes and head so that important visual stimuli are centered on the fovea. Light falling on the retina causes chemical changes to pigment molecules in the photoreceptors, ultimately leading to a change in the activity of the RGCs.

Figure 4. (a) The human eye is shown in cross section. (b) A blowup shows the layers of the retina.

 

There are two types of photoreceptors in the retina: rods and cones, named for their general appearance as illustrated in Figure 5. Rods are strongly photosensitive and are located in the outer edges of the retina. They detect dim light and are used primarily for peripheral and nighttime vision. Cones are weakly photosensitive and are located near the center of the retina. They respond to bright light, and their primary role is in daytime, color vision.

Figure 5. Rods and cones are photoreceptors in the retina. Rods respond in low light and can detect only shades of gray. Cones respond in intense light and are responsible for color vision. (credit: modification of work by Piotr Sliwa)

The fovea has a high density of cones. When you bring your gaze to an object to examine it intently in bright light, the eyes orient so that the object’s image falls on the fovea. However, when looking at a star in the night sky or other object in dim light, the object can be better viewed by the peripheral vision because it is the rods at the edges of the retina, rather than the cones at the center, that operate better in low light. In humans, cones far outnumber rods in the fovea.

Transduction of Light

Photoreceptor cells have two parts, the inner segment and the outer segment (Figure 6). The inner segment contains the nucleus and other common organelles of a cell, whereas the outer segment is a specialized region in which photoreception takes place. There are two types of photoreceptors—rods and cones—which differ in the shape of their outer segment. The rod-shaped outer segments of the rod photoreceptor contain a stack of membrane-bound discs that contain the photosensitive pigment rhodopsin. The cone-shaped outer segments of the cone photoreceptor contain their photosensitive pigments in infoldings of the cell membrane. There are three cone photopigments, called opsins, which are each sensitive to a particular wavelength of light. The wavelength of visible light determines its color. The pigments in human eyes are specialized in perceiving three different primary colors: red, green, and blue.

Figure 6. Photoreceptor (a) All photoreceptors have inner segments containing the nucleus and other important organelles and outer segments with membrane arrays containing the photosensitive opsin molecules. Rod outer segments are long columnar shapes with stacks of membrane-bound discs that contain the rhodopsin pigment. Cone outer segments are short, tapered shapes with folds of membrane in place of the discs in the rods. (b) Tissue of the retina shows a dense layer of nuclei of the rods and cones. LM × 800. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)

At the molecular level, visual stimuli cause changes in the photopigment molecule that lead to changes in membrane potential of the photoreceptor cell. A single unit of light is called a photon, which is described in physics as a packet of energy with properties of both a particle and a wave. The energy of a photon is represented by its wavelength, with each wavelength of visible light corresponding to a particular color. Visible light is electromagnetic radiation with a wavelength between 380 and 720 nm. Longer wavelengths of less than 380 nm fall into the infrared range, whereas shorter wavelengths of more than 720 nm fall into the ultraviolet range. Light with a wavelength of 380 nm is blue whereas light with a wavelength of 720 nm is dark red. All other colors fall between red and blue at various points along the wavelength scale.

Opsin pigments are actually transmembrane proteins that contain a cofactor known as retinal. Retinal is a hydrocarbon molecule related to vitamin A. When a photon hits retinal, the long hydrocarbon chain of the molecule is biochemically altered. Specifically, photons cause some of the double-bonded carbons within the chain to switch from a cis to a trans conformation. This process is called photoisomerization. Before interacting with a photon, retinal’s flexible double-bonded carbons are in the cis conformation. This molecule is referred to as 11-cis-retinal. A photon interacting with the molecule causes the flexible double-bonded carbons to change to the trans– conformation, forming all-trans-retinal, which has a straight hydrocarbon chain (Figure 7).

Figure 7. Retinal Isomers The retinal molecule has two isomers, (a) one before a photon interacts with it and (b) one that is altered through photoisomerization.

The shape change of retinal in the photoreceptors initiates visual transduction in the retina. Activation of retinal and the opsin proteins result in activation of a G protein. The G protein changes the membrane potential of the photoreceptor cell, which then releases less neurotransmitter into the outer synaptic layer of the retina. Until the retinal molecule is changed back to the 11-cis-retinal shape, the opsin cannot respond to light energy, which is called bleaching. When a large group of photopigments is bleached, the retina will send information as if opposing visual information is being perceived. After a bright flash of light, afterimages are usually seen in negative. The photoisomerization is reversed by a series of enzymatic changes so that the retinal responds to more light energy.

Figure 8. Comparison of Color Sensitivity of Photopigments Comparing the peak sensitivity and absorbance spectra of the four photopigments suggests that they are most sensitive to particular wavelengths.

The opsins are sensitive to limited wavelengths of light. Rhodopsin, the photopigment in rods, is most sensitive to light at a wavelength of 498 nm. The three color opsins have peak sensitivities of 564 nm, 534 nm, and 420 nm corresponding roughly to the primary colors of red, green, and blue (Figure 8). The absorbance of rhodopsin in the rods is much more sensitive than in the cone opsins; specifically, rods are sensitive to vision in low light conditions, and cones are sensitive to brighter conditions.

In normal sunlight, rhodopsin will be constantly bleached while the cones are active. In a darkened room, there is not enough light to activate cone opsins, and vision is entirely dependent on rods. Rods are so sensitive to light that a single photon can result in an action potential from a rod’s corresponding RGC.

The three types of cone opsins, being sensitive to different wavelengths of light, provide us with color vision. By comparing the activity of the three different cones, the brain can extract color information from visual stimuli. For example, a bright blue light that has a wavelength of approximately 450 nm would activate the “red” cones minimally, the “green” cones marginally, and the “blue” cones predominantly. The relative activation of the three different cones is calculated by the brain, which perceives the color as blue. However, cones cannot react to low-intensity light, and rods do not sense the color of light. Therefore, our low-light vision is—in essence—in grayscale. In other words, in a dark room, everything appears as a shade of gray. If you think that you can see colors in the dark, it is most likely because your brain knows what color something is and is relying on that memory.

Retinal Processing

Visual signals leave the cones and rods, travel to the bipolar cells, and then to ganglion cells. A large degree of processing of visual information occurs in the retina itself, before visual information is sent to the brain.

Photoreceptors in the retina continuously undergo tonic activity. That is, they are always slightly active even when not stimulated by light. In neurons that exhibit tonic activity, the absence of stimuli maintains a firing rate at a baseline; while some stimuli increase firing rate from the baseline, and other stimuli decrease firing rate. In the absence of light, the bipolar neurons that connect rods and cones to ganglion cells are continuously and actively inhibited by the rods and cones. Exposure of the retina to light hyperpolarizes the rods and cones and removes their inhibition of bipolar cells. The now active bipolar cells in turn stimulate the ganglion cells, which send action potentials along their axons (which leave the eye as the optic nerve). Thus, the visual system relies on change in retinal activity, rather than the absence or presence of activity, to encode visual signals for the brain. Sometimes horizontal cells carry signals from one rod or cone to other photoreceptors and to several bipolar cells. When a rod or cone stimulates a horizontal cell, the horizontal cell inhibits more distant photoreceptors and bipolar cells, creating lateral inhibition. This inhibition sharpens edges and enhances contrast in the images by making regions receiving light appear lighter and dark surroundings appear darker. Amacrine cells can distribute information from one bipolar cell to many ganglion cells.

You can demonstrate this using an easy demonstration to “trick” your retina and brain about the colors you are observing in your visual field. Look fixedly at Figure 9 for about 45 seconds. Then quickly shift your gaze to a sheet of blank white paper or a white wall. You should see an afterimage of the Norwegian flag in its correct colors. At this point, close your eyes for a moment, then reopen them, looking again at the white paper or wall; the afterimage of the flag should continue to appear as red, white, and blue. What causes this? According to an explanation called opponent process theory, as you gazed fixedly at the green, black, and yellow flag, your retinal ganglion cells that respond positively to green, black, and yellow increased their firing dramatically. When you shifted your gaze to the neutral white ground, these ganglion cells abruptly decreased their activity and the brain interpreted this abrupt downshift as if the ganglion cells were responding now to their “opponent” colors: red, white, and blue, respectively, in the visual field. Once the ganglion cells return to their baseline activity state, the false perception of color will disappear.

Figure 9. View this flag to understand how retinal processing works. Stare at the center of the flag (indicated by the white dot) for 45 seconds, and then quickly look at a white background, noticing how colors appear.

Higher Processing

The myelinated axons of ganglion cells make up the optic nerves. Within the nerves, different axons carry different qualities of the visual signal. Some axons constitute the magnocellular (big cell) pathway, which carries information about form, movement, depth, and differences in brightness. Other axons constitute the parvocellular (small cell) pathway, which carries information on color and fine detail. Some visual information projects directly back into the brain, while other information crosses to the opposite side of the brain. This crossing of optical pathways produces the distinctive optic chiasma (Greek, for “crossing”) found at the base of the brain and allows us to coordinate information from both eyes.

Once in the brain, visual information is processed in several places, and its routes reflect the complexity and importance of visual information to humans and other animals. One route takes the signals to the thalamus, which serves as the routing station for all incoming sensory impulses except olfaction. In the thalamus, the magnocellular and parvocellular distinctions remain intact, and there are different layers of the thalamus dedicated to each. When visual signals leave the thalamus, they travel to the primary visual cortex at the rear of the brain. From the visual cortex, the visual signals travel in two directions. One stream that projects to the parietal lobe, in the side of the brain, carries magnocellular (“where”) information. A second stream projects to the temporal lobe and carries both magnocellular (“where”) and parvocellular (“what”) information.

Another important visual route is a pathway from the retina to the superior colliculus in the midbrain, where eye movements are coordinated and integrated with auditory information. Finally, there is the pathway from the retina to the suprachiasmatic nucleus (SCN) of the hypothalamus. The SCN is a cluster of cells that is considered to be the body’s internal clock, which controls our circadian (day-long) cycle. The SCN sends information to the pineal gland, which is important in sleep/wake patterns and annual cycles.