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Abstract
Ethylene is an important regulator of plant growth. We identified an Arabidopsis mutant,
responsive-to-antagonist1 (ran1), that shows ethylene phenotypes in response to treatment
with trans-cyclooctene, a potent receptor antagonist. Genetic epistasis studies revealed
an early requirement for RAN1 in the ethylene pathway. RAN1 was cloned and found to
encode a protein with similarity to copper-transporting P-type ATPases, including
the human Menkes/Wilson proteins and yeast Ccc2p. Expression of RAN1 complemented
the defects of a ccc2delta mutant, demonstrating its function as a copper transporter.
Transgenic CaMV 35S::RAN1 plants showed constitutive expression of ethylene responses,
due to cosuppression of RAN1. These results provide an in planta demonstration that
ethylene signaling requires copper and reveal that RAN1 acts by delivering copper
to create functional hormone receptors.