Haldan Keffer Hartline

One of the scientists who took advantage of this remarkable animal was H. Keffer Hartline. He was able to isolate and study the activity of single nerve fibers as they relayed signals from single ommatidia to the brain in Limulus. In 1932 he published a paper titled "Nerve Impulses from Single Receptors in the Eye" describing this work. It was his work in clarifying how ommatidia interact with each other, however, that led to understanding the mechanisms of lateral inhibition and won him the Nobel Prize.

Web sites which feature lateral inhibition

http://www.exploratorium.edu/xref/phenomena/lateral_inhibition.html

http://serendip.brynmawr.edu/bb/latinhib.html

Lateral inhibition is a process that animals, including humans, use to better distinguish borders. When you look at the ocean horizon the ocean appears darker at the horizon, at the boundary between sea and sky. This apparent difference in light intensity is not actually there but is created by our visual receptors and is known as lateral inhibition. This process increases contrast and results in a sharpening of vision. In fact, computers sharpen images by almost the same process process. (the illustration below illustrates both lateral inhibition and a sharpening filter from a graphics program.) What this means is that the signals coming from the outside are actually altered before being sent to the brain so that what we see isn't necessarily there.

The way this works is as follows. Impulses originate in the eccentric cell when the cell is stimulated by light . This signal is transmitted through the axon then to the optic nerve to the brain.

The ability of an ommatidia to discharge impulses is related to the amount of light that neighboring ommatidia are receiving. Hartline found that if one ommatidia is receiving bright light and a neighbor is receiving dim light, the first ommatidia will inhibit the signal from it's neighbor. The result is that the dimmer signal gets even dimmer and the result is an increased difference between the two which the eye would perceive as an increase in contrast.

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References used in the Vision sections of this site.

Hartline., H.K., Wagner, H.G., Ratcliff, Floyd, Inhibition in the Eye of Limulus, Journal of General Physiology, 1956, 39:5 pp.651-673

Westerman, L.A., Barlow, R.B., Ultraviolet responses of the Limulus mediann ocellus, Biological Bulletin, 1981 161 352-353

Barlow, R.B., Ireland, C.I., Kass, L., Vision in Limulus mating behavior, Biological Bulletin, 1981 161 339-340

Powers, M.K., Barlow, R.B., Circadian changes in visual sensitivity of Limulus: behavioral evidence, Biological Bulletin, 1981 161 350-351

Hubbard, Ruth. Retinene Isomerase, Journal of General Physiology, Vol 39, No. 6 pp.935-962

Wald, G., Human Vision and the Spectrum, Science, 1945, 101, 653

Wald, G., Life and Light, Scientific American, Oct. 1959, pp 92-108

Invertebrate Photoreceptors, A Comparative Analysis, Jerome J. Wolken, Academic Press, NY, 1971

Kimbel, R.L., Poincelot, R.P., Abrahamson, E.W., Chromophore Transfer from Lipid to Protein in Bovine Rhodopsin, Biochemistry 1970 9:8 1817

Westerman, L.A, Barlow, R.B, Ultraviolet responses of the Limulus median ocellus, Biological Bulletin General Scientific Meetings. 161:3 352-353

Barlow, R.B, Ireland, L.C., Cass, L., Viision in Limulus mating behavior, Biological Bulletin General Scientific Meetings. 161:3 339-340

Sargent, William., The Year of the Crab., W.W. Norton & Company 1987