Cell elongation during seedling development is antagonistically regulated by light and gibberellins (GAs) 1,2 . Light induces photomorphogenesis, leading to inhibition of hypocotyl growth, whereas GAs promote etiolated growth, characterized by increased hypocotyl elongation. The mechanism underlying this antagonistic interaction remains unclear. Here we report on the central role of the Arabidopsis thaliana nuclear transcription factor PIF4 (encoded by PHYTOCHROME INTERACTING FACTOR 4) 3 in the positive control of genes mediating cell elongation and show that this factor is negatively regulated by the light photoreceptor phyB (ref. 4) and by DELLA proteins that have a key repressor function in GA signalling 5 . Our results demonstrate that PIF4 is destabilized by phyB in the light and that DELLAs block PIF4 transcriptional activity by binding the DNA-recognition domain of this factor. We show that GAs abrogate such repression by promoting DELLA destabilization, and therefore cause a concomitant accumulation of free PIF4 in the nucleus. Consistent with this model, intermediate hypocotyl lengths were observed in transgenic plants over-accumulating both DELLAs and PIF4. Destabilization of this factor by phyB, together with its inactivation by DELLAs, constitutes a protein interaction framework that explains how plants integrate both light and GA signals to optimize growth and development in response to changing environments.Seedlings undergo alternative developmental programmes depending on whether they are germinated in the dark or in the light. Dark-grown seedlings exhibit etiolated growth, characterized by long hypocotyls, small and closed cotyledons with undifferentiated chloroplasts, and the repression of light-regulated genes 1 . During photomorphogenesis, light inhibits hypocotyl growth and promotes cotyledon opening and expansion, chloroplast differentiation and the activation of light-regulated genes. phyB is the main photoreceptor mediating de-etiolation in red light 4,6 . Absorption of red light converts this photoreceptor into a Pfr active form that is translocated into the nucleus 7,8 ; Pfr interacts there with members of the bHLH family of phytochrome-interacting factors (PIFs), involved in modulation of light-regulated genes with a role in photomorphogenesis 1,4 . Gibberellins (GAs) exert an opposite effect to light on photomorphogenesis 2 . GAs promote etiolated growth, whereas GA-deficiency induces a partially de-etiolated phenotype in the dark, which is reverted by a lack of DELLA function 2,9 . DELLAs function as key repressors of GA-responsive growth, by inhibiting GA-regulated gene expression 5 . These repressors accumulate in the nucleus and are rapidly degraded in response to GA 10,11 . In Arabidopsis, RGA (encoded by repressor of ga1-3) and GAI (encoded by GA insensitive) are the main repressors controlling hypocotyl growth and stem elongation 12,13 . Mutations within the DELLA domain render these proteins resistant to degradation, and result in a GA-insensitive dwarf phenotype 12,14 . This ...
Production of phytohormones is one of the main mechanisms to explain the beneficial effects of plant growth-promoting rhizobacteria (PGPR) such as Azospirillum sp. The PGPRs induce plant growth and development, and reduce stress susceptibility. However, little is known regarding the stress-related phytohormone abscisic acid (ABA) produced by bacteria. We investigated the effects of Azospirillum brasilense Sp 245 strain on Arabidopsis thaliana Col-0 and aba2-1 mutant plants, evaluating the morphophysiological and biochemical responses when watered and in drought. We used an in vitro-grown system to study changes in the root volume and architecture after inoculation with Azospirillum in Arabidopsis wild-type Col-0 and on the mutant aba2-1, during early growth. To examine Arabidopsis development and reproductive success as affected by the bacteria, ABA and drought, a pot experiment using Arabidopsis Col-0 plants was also carried out. Azospirillum brasilense augmented plant biomass, altered root architecture by increasing lateral roots number, stimulated photosynthetic and photoprotective pigments and retarded water loss in correlation with incremented ABA levels. As well, inoculation improved plants seed yield, plants survival, proline levels and relative leaf water content; it also decreased stomatal conductance, malondialdehyde and relative soil water content in plants submitted to drought. Arabidopsis inoculation with A. brasilense improved plants performance, especially in drought.
SummaryBlue light inhibits elongation of etiolated Arabidopsis thaliana hypocotyls during the ®rst 30 min of irradiation by a mechanism that depends on the phototropin 1 (phot1) photoreceptor. The cryptochrome 1 (cry1) photoreceptor begins to exert control after 30 min. To identify genes responsible for the cry1 phase of growth inhibition, mRNA expression pro®les of cry1 and wild-type seedlings were compared using DNA microarrays. Of the roughly 420 genes found to be differentially expressed at the point of cry1 response incipience, approximately half were expressed higher and half lower in cry1 relative to the wild type. Many of the cry1-dependent genes encoded kinases, transcription factors, cell cycle regulators, cell wall metabolism enzymes, gibberellic acid (GA) biosynthesis enzymes, and auxin response factors. High-resolution growth studies supported the hypothesis that genes in the last two categories were indeed relevant to cry1-mediated growth control. Inhibiting GA 4 biosynthesis with a 3b-hydroxylase inhibitor (Ca-prohexadione) restored wild-type response kinetics in cry1 and completely suppressed its long-hypocotyl phenotype in blue light. Co-treatment of cry1 seedlings with Ca-prohexadione plus GA 4 completely reversed the effects of the inhibitor, restoring the long-hypocotyl phenotype typical of the mutant. Treatment of wild-type seedlings with GA 4 was not suf®cient to phenocopy cry1 seedlings, but cotreatment with IAA plus GA 4 produced cry1-like growth kinetics for a period of approximately 5 h. The genomic and physiological data together indicate that blue light acting through cry1 quickly affects the expression of many genes, a subset of which suppresses stem growth by repressing GA and auxin levels and/or sensitivity.
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