Phase change and the regulation of trichome distribution in Arabidopsis thaliana

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Abstract
Higher plants pass through several phases of shoot growth during which they may produce morphologically distinct vegetative structures. In Arabidopsis thaliana this phenomenon is apparent in the distribution of trichomes on the leaf surface. Leaves produced early in rosette development lack trichomes on their abaxial (lower) surface, leaves produced later have trichomes on both surfaces, and leaves in the inflorescence (bracts) may have few or no trichomes on their adaxial (upper) surface. Here we describe some of the factors that regulate this distribution pattern. We found that the timing of abaxial trichome production and the extent to which bracts lack adaxial trichomes varies in different ecotypes. The production of abaxial trichomes appears to be regulated by the age, rather than the size of the plant. This conclusion is based on the observation that mutations that affect either the rate (altered meristem programming1) or onset (paused) of leaf initiation respectively increase or decrease the number of leaves that lack abaxial trichomes, but have only a minor effect on the time at which the first leaf with abaxial trichomes is produced. The production of abaxial trichomes is coordinated with the reproductive development of the shoot as this trait is delayed by photoperiodic conditions and some mutations that delay flowering. The loss of adaxial trichomes is likely to be a consequence of floral induction, and is accelerated by terminal flower1-10, a mutation that accelerates inflorescence development. We demonstrate that gibberellins promote trichome production in Arabidopsis and present evidence indicating that abaxial trichome production is regulated by both the level of a trichome inducer and the competence of the abaxial epidermis to respond to this inducer.