Ethylene Induces Epidermal Cell Death at the Site of Adventitious Root Emergence in Rice

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Abstract
In deepwater rice (Oryza sativa), adventitious root primordia initiate at the nodes as part of normal development. Emergence of the roots is dependent on flooding of the plant and is mediated by ethylene action. Root growth was preceded by the induced death of epidermal cells of the node external to the tip of the root primordium. Cell death proceeded until the epidermis split open. Through this crack the root eventually emerged. Induced death was confined to nodal epidermal cells covering the tip of the primordia. Our results suggest that this process facilitates adventitious root emergence and prevents injury to the growing root. Cell death was inducible not only by submergence but also by application of 1-aminocyclopropane-1-carboxylic acid, the natural precursor of ethylene and it was suppressed in the presence of 2,5-norbornadiene (bicyclo[2.2.1]hepta-2,5-diene), an inhibitor of ethylene action. Adventitious root growth and epidermal cell death are therefore linked to the ethylene signaling pathway, which is activated in response to low oxygen stress.