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chemotaxis elegans identification attractants analysis
Proc. Nat. Acad. Sci. USA
Vol. 70, No. 3, pp. 817-821, March 1973
Chemotaxis by the Nematode Caenorhabditis elegans: Identification
of Attractants and Analysis of the Response by Use of Mutants
(cAMP/anions/cations/hydroxyl ions/klinotaxis)
SAMUEL WARD*
Medical Research Council, Laboratory of Molecular Biology, Hills Road, Cambridge, CB2 2QH, England
Communicated by F. H. C. Crick, January 8, 1973
ABSTRACT The nematode Caenorhabditis elegans is Establishment of Gradients. The two methods used for
attracted by at least four classes of attractants: by cyclic studying chemosensory behavior depend upon the establish-
nucleotides, cAMP and cGMP; by anions, Cl-, Br-, 1; by ment of defined and reproducible gradients of attractants.
cations, Na+, Li+, K+, Mg+; and by alkaline pH values.
The nematodes behavioral response to gradients of these Radial gradients were established in a thin layer of agarose
attractants involves orientation and movement up the or Sephadex gel beads spread on petri-plates. The agarose or
gradient, accumulation, and then habituation. Compari- gel beads stabilize the liquid against convection and also
son of the tracks of wild-type and mutant animals re- provide a medium on or through which the worms move
sponding to gradients of attractants indicates that sensory
receptors in the head alone mediate the orientation re- rapidly. When the diffusion of 32PO4 and [14C]phenylalanine
sponse and that the direction of orientation is determined were measured, the gradients established at different times
by the lateral motion of the head. Therefore, the orienta- were found to decrease exponentially as predicted by theory
tion response is a klinotaxis. (ref. 2; Eq. 3, 10), and the diffusion coefficients of the
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