Nature 431, 647-652 (7 October 2004) | doi:10.1038/nature02952; Received 25 May 2004; Accepted 19 August 2004

Surface mechanics mediate pattern formation in the developing retina

Takashi Hayashi1,2 & Richard W. Carthew1

  1. Department of Biochemistry, Molecular Biology and Cell Biology, Northwestern University, Evanston, Illinois 60208, USA
  2. Department of Biophysics and Biochemistry, Graduate School of Science, University of Tokyo, Tokyo 113-0033, Japan

Correspondence to: Takashi Hayashi1,2Richard W. Carthew1 Correspondence and requests for materials should be addressed to T.H. (Email: or R.W.C. (Email:


Pattern formation of biological structures involves organizing different types of cells into a spatial configuration. In this study, we investigate the physical basis of biological patterning of the Drosophila retina in vivo. We demonstrate that E- and N-cadherins mediate apical adhesion between retina epithelial cells. Differential expression of N-cadherin within a sub-group of retinal cells (cone cells) causes them to form an overall shape that minimizes their surface contact with surrounding cells. The cells within this group, in both normal and experimentally manipulated conditions, pack together in the same way as soap bubbles do. The shaping of the cone cell group and packing of its components precisely imitate the physical tendency for surfaces to be minimized. Thus, simple patterned expression of N-cadherin results in a complex spatial pattern of cells owing to cellular surface mechanics.


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