Vision

Vision is the ability to detect light patterns from the outside environment and interpret them into images. Animals are bombarded with sensory information, and the sheer volume of visual information can be problematic. Fortunately, the visual systems of species have evolved to attend to the most-important stimuli. The importance of vision to humans is further substantiated by the fact that about one-third of the human cerebral cortex is dedicated to analyzing and perceiving visual information.

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 (abbreviated 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).

The illustration shows the electromagnetic spectrum, which consists of different wavelengths of electromagnetic radiation. Radio waves have the longest wavelength, about 103 meters. Wavelength gets increasingly shorter for microwave, infrared, visible, ultraviolet, x-rays and gamma rays. Gamma rays have a wavelength of about 10-12 meters. Frequency is inversely proportional to wavelength.

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 a 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 photoreceptive cells of the eye, where transduction of light to nervous impulses occurs, are located in the retina (shown in Figure 2) on the inner surface of the back of the eye. But light does not impinge on the retina unaltered. It passes through other layers that process it so that it can be interpreted by the retina (Figure 2b). The cornea, the front transparent layer of the eye, and the crystalline lens, a transparent convex structure behind the cornea, both refract (bend) light to focus the image on the retina. The iris, which is conspicuous as the colored part of the eye, is a circular muscular ring lying between the lens and cornea that regulates the amount of light entering the eye. In conditions of high ambient light, the iris contracts, reducing the size of the pupil at its center. In conditions of low light, the iris relaxes and the pupil enlarges.

The left illustration shows a human eye, which is round and filled with vitreous humour. The optic nerve and retinal blood vessels exit the back of the eye. At the front of the eye is the lens with a pupil in the middle. The lens is covered by the iris, which in turn is covered by the cornea. The aqueous humour is a gel-like substance between the cornea and iris. The retina is the lining of the inner eye. A second illustration is a blowup which shows that the optic nerve is at the surface of the retina. Beneath the optic nerve is a layer of ganglion cells, and beneath this is a layer of bipolar cells. Both ganglia and bipolar cells are nerve cells with root-like appendages. Beneath the bipolar cell layer are the rods and cones. Rods and cones are similar in structure and column-like.

Figure 2. (a) The human eye is shown in cross section. (b) A blowup shows the layers of the retina.

Practice Question

Which of the following statements about the human eye is false?

  1. Rods detect color, while cones detect only shades of gray.
  2. When light enters the retina, it passes the ganglion cells and bipolar cells before reaching photoreceptors at the rear of the eye.
  3. The iris adjusts the amount of light coming into the eye.
  4. The cornea is a protective layer on the front of the eye.

The main function of the lens is to focus light on the retina and fovea centralis. The lens is dynamic, focusing and re-focusing light as the eye rests on near and far objects in the visual field. The lens is operated by muscles that stretch it flat or allow 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.

This illustration shows that rods and cones are both long, column-like cells with the nucleus located in the bottom portion. The rod is longer than the cone. The outer segment of the rod contains rhodopsin. The outer segment of the rod contains other photo-pigments. An oil droplet is located beneath the outer segment.

Figure 3. 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)

There are two types of photoreceptors in the retina: rods and cones, named for their general appearance as illustrated in Figure 3. 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. There are 3 types of cones each detecting a different wavelength of light (red, blue and green).

The fovea is the region in the center back of the eye that is responsible for acute vision. 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.

Review the anatomical structure of the eye, clicking on each part to practice identification.

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 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.  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.

View this interactive presentation to review what you have learned about how vision functions.

In Summary: Vision

Vision is the only photo responsive sense. Visible light travels in waves and is a very small slice of the electromagnetic radiation spectrum. Light waves differ based on their frequency (wavelength = hue) and amplitude (intensity = brightness).

In the vertebrate retina, there are two types of light receptors (photoreceptors): cones and rods. Cones, which are the source of color vision, exist in three formsand they are differentially sensitive to different wavelengths (red, blue and green). Cones are located in the retina, along with the dim-light, achromatic receptors (rods). Cones are found in the fovea, the central region of the retina, whereas rods are found in the peripheral regions of the retina.

Visual signals travel from the eye over the axons of retinal ganglion cells, which make up the optic nerves. Ganglion cells come in several versions. Some ganglion cell axons carry information on form, movement, depth, and brightness, while other axons carry information on color and fine detail.