A new mutant called psi2 (for p hytochrome si gnaling) was isolated by screening for elevated activity of a chlorophyll a / b binding protein-luciferase ( CAB2-LUC ) transgene in Arabidopsis. This mutant exhibited hypersensitive induction of CAB1 , CAB2 , and the small subunit of ribulose-1,5-bisphosphate carboxylase ( RBCS ) promoters in the very low fluence range of red light and a hypersensitive response in hypocotyl growth in continuous red light of higher fluences. In addition, at high-but not low-light fluence rates, the mutant showed light-dependent superinduction of the pathogenrelated protein gene PR-1a and developed spontaneous necrotic lesions in the absence of any pathogen. Expression of genes responding to various hormone and environmental stress pathways in the mutant was not significantly different from that of the wild type. Analysis of double mutants demonstrated that the effects of the psi2 mutation are dependent on both phytochromes phyA and phyB. The mutation is recessive and maps to the bottom of chromosome 5. Together, our results suggest that PSI2 specifically and negatively regulates both phyA and phyB phototransduction pathways. The induction of cell death by deregulated signaling pathways observed in psi2 is reminiscent of retinal degenerative diseases in animals and humans.
INTRODUCTIONBecause light provides the energy source for photosynthesis, plants must adapt to changes in ambient light quantity and quality to optimize the efficiency of this important process. The results of this adaptation are apparent: almost every step of plant development, from seed germination to fruit maturation, is dependent on light. Moreover, light regulates the circadian rhythm, the adaptative size modification of tissue and organs, and the expression of photosynthetic genes and many other known and unknown genes (Terzaghi and Cashmore, 1995).To date, what we know about light information processing and signal transduction concerns the nature and function of photoreceptors. A family of photoreceptors called phytochromes has been shown to be involved in the perception of red and far-red light. Their biochemical and molecular properties and physiological roles are now well described (reviewed in Quail et al., 1995;Smith, 1995). In addition, a longpostulated UV and blue light receptor (cryptochrome) was recently characterized by Ahmad and Cashmore (1993). Mutants lacking one or several of these receptors have provided useful information concerning the role of individual photoreceptors and their interaction in transducing light signals (reviewed in Barnes et al., 1997).Much less is known concerning molecules that link light perception to gene activation. The CONSTITUTIVE PHOTO-MORPHOGENIC/DEETIOLATED/FUSCA (COP/DET/FUS) class of proteins has been suggested to play an important role in regulating the repression of light-inducible genes and the maintenance of skotomorphogenesis (Chory et al., 1989). For some of these proteins, the genes have been cloned and include COP1 (Deng et al., 1992), DET1 (Pepper et al., ...