Phytochrome A (phyA) is the photolabile plant light receptor that mediates broad spectrum very low-fluence responses and high irradiance responses to continuous far-red light (FR c ). An Arabidopsis mutant laf3-1 (long after far-red 3) was recovered from a screen for transposon-tagged mutants that exhibit reduced inhibition of hypocotyl elongation in FR c . The laf phenotype correlated well with a strongly attenuated disappearance of XTR7 transcript in FR c . The effects of laf3-1 on phyA-controlled CAB, CHS, and PET H expression were more subtle, and the mutation had no clear effects on PET E and ASN1 transcript levels in FR c . The use of two alternative transcription initiation sites in the LAF3 gene generates two isoforms that differ only at their N termini. Transcripts encoding both isoforms were induced during germination and were present at slightly higher levels in de-etiolated seedlings than in those grown in darkness. No significant differential regulation of the two isoforms was observed upon exposure to either FR c or continuous red light. Transcripts encoding the shorter isoform (LAF3 ISF2 ) always appear to be more abundant than those encoding the longer isoform (LAF3 ISF1 ). However, both isoforms were capable of full complementation of the laf3-1 hypocotyl phenotype in FR c . When fused to a yellow fluorescent protein, both isoforms localize to the perinuclear region, suggesting that LAF3 encodes a product that might regulate nucleo-cytoplasmic trafficking of an intermediate(s) involved in phyA signal transduction.Phytochromes are soluble chromoproteins that regulate plant growth and development by their ability to interconvert between two stable spectral forms. Red light (R) converts the R-absorbing Pr form (the form synthesized in darkness) to the Pfr (far-red absorbing form) and far-red light (FR) reconverts Pfr to Pr. Two important features distinguish phytochrome A (phyA) from the other four phytochromes in Arabidopsis. First, although the Arabidopsis phytochromes phyB to phyE are activated exclusively by R and inactivated by irradiation with FR, phyA can be activated by FR and low fluences of R and blue light (B). Secondly, although phyA levels decrease rapidly after exposure to light as a result of both down-regulation of PHYA gene transcription (Cantó n and Quail, 1999) and far greater photolability of phyA than other phytochromes (Sharrock and Clack, 2002), phyA is by far the most abundant phytochrome in etiolated seedlings. These features enable phyA to perform a seminal role in triggering the shift between skotomorphogenesis and photomorphogenesis. Together, they ensure that the inhibition of hypocotyl elongation, and the activation of physiological changes needed to ensure photosynthetic competence are already underway when seedlings emerge from the soil surface. Despite its lability in light-grown plants, the influence of phyA throughout the life cycle is evidenced by its involvement in sensing photoperiod to ensure that flowering is initiated at the proper time (Johnson et al., 199...