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J. Biol. Chem., Vol. 276, Issue 24, 21714-21723, June 15, 2001

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Role of the Lewis^x Glycan Determinant in Corneal Epithelial Cell Adhesion and Differentiation^*

Zhiyi Cao, Zheng Zhao, Royce Mohan, Joseph Alroy^§, Pamela Stanley^¶, and Noorjahan Panjwani^**

From the  New England Eye Center and Departments of Ophthalmology,  Biochemistry, and ^§ Pathology, Tufts University School of Medicine, Boston, Massachusetts 02111 and the ^¶ Department of Cell Biology, Albert Einstein College of Medicine, New York, New York 10461

In this study we demonstrate that in corneal epithelium there is cell-cell contact-regulated expression of a 145-kDa glycoprotein (GP) bearing the glycan determinant Lewis^x (Le^x) (Gal(1,4)[Fuc(1,3)]GlcNAc). This glycoprotein (Le^x-GP) was expressed in confluent/contact-inhibited cultures but not in sparse cultures of corneal epithelium. In contrast, a 135-kDa glycoprotein bearing precursor, unfucosylated, lactosamine-containing glycans (Gal1-4GlcNAc1-R) was expressed in sparse cultures. Immunofluorescence staining and confocal microscopy of confluent cultures revealed that in corneal epithelium, Le^x antigen is located in high density at sites of cell-cell adhesion. In in vitro cell-cell adhesion assays, anti-Le^x, but not anti-sialyl-Le^x monoclonal antibodies, inhibited the formation of corneal epithelial cell-cell adhesion. Also, when added to confluent cultures, antibodies to Le^x disrupted the monolayer and caused tightly packed polygonal cells to round up. Analysis of the expression of Fut genes that encode -1,3-fucosyltransferases, the enzymes that generate the Le^x determinant, revealed that confluent/contact-inhibited cultures of rabbit corneal epithelium contain markedly elevated levels of Fut4 and Fut3/5/6 gene transcripts compared with sparse cultures. These data suggest that the Fut4 and Fut3/5/6 genes are targets of cell-cell contact-regulated signals and that Fut gene products direct cell-cell contact-associated expression of Le^x on the Le^x-GP in corneal epithelium. Immunohistochemical analysis revealed that the expression of Le^x antigen in the epithelium of adult and developing corneas is related to the stage of differentiation of the cells. Although early differentiated cells robustly expressed Le^x, relatively undifferentiated cells did not, and the expression level was relatively low in terminally differentiated cells. Overall, these data provide evidence that a Le^x-bearing glycoprotein plays a role through the Le^x determinant in corneal epithelial cell-cell adhesion, and these data suggest that Le^x-mediated cell-cell interactions contribute to mechanisms that mediate corneal epithelial cell differentiation.

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