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First published online 30 August 2006
doi: 10.1242/dev.02554


Development 133, 3755-3766 (2006)
Published by The Company of Biologists 2006


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The HVE/CAND1 gene is required for the early patterning of leaf venation in Arabidopsis

María Magdalena Alonso-Peral1,*, Héctor Candela1,*,{dagger}, Juan Carlos del Pozo2, Antonio Martínez-Laborda3, María Rosa Ponce1 and José Luis Micol1,{ddagger}

1 División de Genética and Instituto de Bioingeniería, Universidad Miguel Hernández, Campus de Elche, 03202 Elche, Alicante, Spain.
2 Instituto Nacional de Investigación y Tecnología Agraria y Alimentaria, Departamento de Biotecnología, Carretera de la Coruña Km. 7, 28040 Madrid, Spain.
3 División de Genética, Universidad Miguel Hernández, Campus de San Juan, 03550 Alicante, Spain.

{ddagger} Author for correspondence (e-mail: jlmicol{at}umh.es)

Accepted 24 July 2006

The hemivenata-1 (hve-1) recessive allele was isolated in a search for natural variations in the leaf venation pattern of Arabidopsis thaliana, where it was seen to cause extremely simple venation in vegetative leaves and cotyledons, increased shoot branching, and reduced root waving and fertility, traits that are reminiscent of some mutants deficient in auxin signaling. Reduced sensitivity to exogenous auxin was found in the hve-1 mutant, which otherwise displayed a wild-type response to auxin transport inhibitors. The HVE gene was positionally cloned and found to encode a CAND1 protein. The hve-1 mutation caused mis-splicing of the transcripts of the HVE/CAND1 gene and a vein phenotype indistinguishable from that of hve-2 and hve-3, two putatively null T-DNA alleles. Inflorescence size and fertility were more affected by hve-2 and hve-3, suggesting that hve-1 is hypomorphic. The simple venation pattern of hve plants seems to arise from an early patterning defect. We found that HVE/CAND1 binds to CULLIN1, and that the venation patterns of axr1 and hve mutants are similar, which suggest that ubiquitin-mediated auxin signaling is required for venation patterning in laminar organs, the only exception being cauline leaves. Our analyses of double mutant and transgenic plants indicated that auxin transport and perception act independently to pattern leaf veins, and that the HVE/CAND1 gene acts upstream of ATHB-8 at least in higher order veins, in a pathway that involves AXR1, but not LOP1, PIN1, CVP1 or CVP2.

Key words: Arabidopsis, TIP120, CAND1, Venation pattern formation, Natural variation


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