Abstract
The Arabidopsis transcription factor HY5 controls light-induced gene expression downstream of photoreceptors and plays an important role in the switch of seedling shoots from dark-adapted to light-adapted development. In addition, HY5 has been implicated in plant hormone signaling, accounting for the accelerated root system growth phenotype of hy5 mutants. Mutants in the close HY5 homolog HYH resemble wild-type, despite the largely similar expression patterns and levels of HY5 and HYH, and the functional equivalence of the respective proteins. Moreover, the relative contribution of HYH to the overall activity of the gene pair is increased by an alternative HYH transcript, which encodes a stabilized protein. Consistent with the enhanced root system growth observed in hy5 loss-of-function mutants, constitutively overexpressed alternative HYH inhibits root system growth. Paradoxically, however, in double mutants carrying hy5 and hyh null alleles, the hy5 root growth phenotype is suppressed rather than enhanced. Even more surprisingly, compared to wild-type, root system growth is diminished in hy5 hyh double mutants. In addition, the double mutants display novel shoot phenotypes that are absent from either single mutant. These include cotyledon fusions and defective vasculature, which are typical for mutants in genes involved in the transcriptional response to the plant hormone auxin. Indeed, many auxin-responsive and auxin signaling genes are misexpressed in hy5 mutants, and at a higher number and magnitude in hy5 hyh mutants. Therefore, auxin-induced transcription is constitutively activated at different levels in the two mutant backgrounds. Our data support the hypothesis that the opposite root system phenotypes of hy5 single and hy5 hyh double mutants represent the morphological response to a quantitative gradient in the same molecular process, that is gradually increased constitutive auxin signaling. The data also suggest that HY5 and HYH are important negative regulators of auxin signaling amplitude in embryogenesis and seedling development.
Highlights
Homologous genes of the same family display genetic redundancy to varying degrees if their expression pattern and their function overlap
The discrepancy between the hy5 and hyh mutant phenotypes could be explained by differential expression patterns of the two genes
HY5 HOMOLOG (HYH) is expressed throughout the seedling as well, but at a clearly higher level than HYPOCOTYL 5 (HY5) (Figure 1C and 1G)
Summary
Homologous genes of the same family display genetic redundancy to varying degrees if their expression pattern and their function overlap. Loss-of-function mutations of redundantly acting genes are expected to result in no phenotype in the case of full redundancy, or similar phenotypes in the case of partial redundancy. If the mutations in partially redundant genes are combined, an enhancement of the single mutant phenotypes is expected. We investigated the genetic redundancy between two functionally equivalent Arabidopsis transcription factors. Their combined loss-of-function leads to a phenotype that is opposite to what would be expected from the single mutant phenotypes. These two genes have been originally identified because of their role in light signaling
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