Plant proximity perception dynamically modulates hormone levels and sensitivity in Arabidopsis

Shade perception involves altered hormone synthesis and sensitivity. Here, we showed that several shade regulators act as positive and negative modulators of the hypocotyl auxin and/or brassinosteroid-induced elongation. The s hade a voidance s yndrome (SAS) refers to a set of plant responses initiated after perception by the phytochromes of light enriched in far-red colour reflected from or filtered by neighbouring plants. These varied responses are aimed at anticipating eventual shading from potential competitor vegetation. In Arabidopsis thaliana, the most obvious SAS response at the seedling stage is the increase in hypocotyl elongation. Here, we describe how plant proximity perception rapidly and temporally alters the levels of not only auxins but also active brassinosteroids and gibberellins. At the same time, shade alters the seedling sensitivity to hormones. Plant proximity perception also involves dramatic changes in gene expression that rapidly result in a new balance between positive and negative factors in a network of interacting basic helix-loop-helix proteins, such as HFR1, PAR1, and BIM and BEE factors. Here, it was shown that several of these factors act as auxin- and BR-responsiveness modulators, which ultimately control the intensity or degree of hypocotyl elongation. It was deduced that, as a consequence of the plant proximity-dependent new, dynamic, and local balance between hormone synthesis and sensitivity (mechanistically resulting from a restructured network of SAS regulators), SAS responses are unleashed and hypocotyls elongate

Análisis de la interconexión entre rutas transcripcionales hormonales y del síndrome de huida de la sombra de Arabidopsis

[spa] El crecimiento y desarrollo vegetal está fuertemente determinado por las condiciones ambientales, siendo la luz una de las señales ambientales más importantes. Las plantas perciben continuamente la información lumínica por los fotorreceptores. En concreto, la fracción del espectro correspondiente al rojo (R) y rojo lejano (FR) es registrada por los fitocromos. La proximidad de otras plantas (que ocurre en alta densidad vegetal) disminuye la relación R/FR de la luz percibida por una planta, desencadenando un conjunto de respuestas fisiológicas que se conoce como el síndrome de huida de la sombra (SAS). En el estadío de plántula de Arabidopsis thaliana, el alargamiento del hipocotilo es la respuesta más evidente. La longitud del hipocotilo también está influenciada por la acción de ciertas hormonas vegetales como las auxinas y las giberelinas, lo que sugiere que podrían estar participando en el SAS. En este trabajo se describe que los factores PHYTOCHROME RAPIDLY REGULATED 1 (PAR1), PAR2, LONG HYPOCOTYL IN FR1 (HFR1), BRENHANCED EXPRESSION (BEE) y BES1-INTERACTING MYC-LIKE (BIM), involucrados en la modulación del SAS, afectan a la sensibilidad a giberelinas. Se propone que dentro de esta interconexión entre factores del SAS y giberelinas también tendrían un rol importante el incremento en los niveles de auxinas. Con el fin de identificar el mecanismo molecular implicado en esta regulación, se ha caracterizado el funcionamiento de PAR1, un regulador negativo del SAS, así como también de sus posibles interactores (BEEs y BIMs), que actúan como reguladores positivos del SAS. La secuencia primaria de PAR1, una proteína nuclear que reprime la expresión génica, sugiere que esta bHLH atípica probablemente no une DNA para llevar a cabo su función. Mediante la sobrexpresión en planta de formas truncadas de PAR1 y fusiones traduccionales al dominio de activación de VP16 hemos concluido que PAR1 no uniría DNA para modular la transcripción. Hemos hipotetizado por tanto que probablemente heterodimeriza con factores BHLH típicos e inhibe su capacidad de unir DNA. Nuestros resultados confirmaron que los factores positivos del SAS, BEE2 y BIM1, unen DNA a secuencias específicas, unión que se ve atenuada en presencia de PAR1, resultados que apoyan nuestra hipótesis de que PAR1 actuaría heterodimerizando. Además, la caracterización fisiológica y molecular de líneas mutantes de pérdida de función sugiere que los genes BEEs y BIMs se organizan en módulos funcionales diferentes en la red regulatoria del SAS. Finalmente todos los análisis genéticos han permitido ver que los BEEs y BIMs no sólo estarían participando en el SAS si no que también serían reguladores del desarrollo y además de redundantes en el control de la viabilidad de la planta.[eng] The shade avoidance syndrome (SAS) refers to a set of plant responses initiated after perception by the phytochrome photoreceptors of light enriched in far-red colour reflected from or filtered by neighboring plants. Overall, these responses are aimed to anticipate eventual shading from potential competitor vegetation by overgrowing them or flowering, ensuring next generation. Specifically at the seedling stage, the SAS includes an increase in hypocotyl elongation, a response also regulated by changes in the levels of auxins and gibberellins. Here we describe that in Arabidopsis thaliana plant proximity perception rapidly alters sensitivity to gibberellins. In addition, it also involves a fast change in a network of interacting BHLH proteins, such as PAR1, BIM and BEE factors, that act as auxin and gibberellins responsiveness modulators, which ultimately control growth and possibly are some of the factors involved in the regulation of auxin over gibberellins. Although the molecular mechanism of this regulation is not known, we propose that PAR1 is a transcriptional cofactor that interacts with BRENHANCED EXPRESSION (BEE) and BES1-INTERACTING MYC-LIKE (BIM) factors. PAR1 directly inhibits these factors by forming non-DNA-binding heterodimers with BEE2 and BIM1. In addition, here we describe the identification of PAR1-interacting factors (POF), including the brassinosteroid signalling components BEEs and BIMs, and characterize their role as networked positive regulators of SAS responses. We provide genetic evidence that BEEs and BIMs not only control plant growth and development under shaded and non-shaded conditions but are also redundant in the control of plant viability. The results of this work suggest that SAS responses are unleashed as a consequence of a plant proximity-triggered new balance of transcriptional regulators within the pre-existing bHLH network, eventually causing hypocotyls to elongate

The shade avoidance syndrome in Arabidopsis: a fundamental role for atypical basic helix–loop–helix proteins as transcriptional cofactors

The shade avoidance syndrome (SAS) refers to a set of plant responses aimed at anticipating eventual shading by potential competitors. The SAS is initiated after perception of nearby vegetation as a reduction in the red to far-red ratio (R:FR) of the incoming light. Low R:FR light is perceived by the phytochromes, triggering dramatic changes in gene expression that, in seedlings, eventually result in an increased hypocotyl elongation to overgrow competitors. This response is inhibited by genes such as PHYTOCHROME RAPIDLY REGULATED 1 (PAR1), PAR2 and LONG HYPOCOTYL IN FR 1 (HFR1), which are transcriptionally induced by low R:FR. Although PAR1/PAR2 and HFR1 proteins belong to different groups of basic helix–loop–helix (bHLH) transcriptional regulators, they all lack a typical basic domain required for binding to E-box and G-box motifs in the promoter of target genes. By overexpressing derivatives of PAR1 and HFR1 we show that these proteins are actually transcriptional cofactors that do not need to bind DNA to directly regulate transcription. We conclude that protein–protein interactions involving the HLH domain of PAR1 and HFR1 are a fundamental aspect of the mechanism by which these proteins regulate gene expression, most likely through interaction with true transcription factors that do bind to the target genes and eventually unleash the observed SAS responses.Fellowships or contracts came from Ministerio de Educación (AG), Gobierno de Chile (NC-E) and CSIC (JB-T). Research in the lab is supported by grants from the Generalitat de Catalunya (Xarba, 2009-SGR697) and Ministerio de Ciencia e Innovación – FEDER funds (BIO2005-00154, CSD2007-00036, BIO2008-00169).Peer reviewe

Identification and characterization of genes differentially expressed in cherimoya (Annona cherimola mill) after exposure to chilling injury conditions

Cherimoyas (Annona cherimola), like other subtropical/tropical fruits, are susceptible to damage from exposure to temperatures between 0 and 5 °C (chilling injury, CI), which may affect fruit quality. To increase our understanding of the molecular mechanisms involved in the CI response, a forward suppression subtractive hybridization (SSH) cDNA library was constructed. In this work, we obtained 75 genes that could potentially be involved in the CI response. The CI induced activation of genes that are involved in a range of metabolic pathways, such as primary metabolism, transport, and endomembrane traffic, among others. We also characterized the expression of 12 selected genes in different A. cherimola tissues by polymerase chain reaction (PCR), and we confirmed the differential expression of a subset in CI fruits by real-time quantitative PCR (qPCR). The expression of six A. cherimola genes: annexin (AcAnex), UDP-glucose pyrophosphorylase (AcUGP), syntaxin of plants 71 (AcSyp71), 1-aminocyclopropane-1-carboxylic-acid synthase (AcACS), ubiquitin carrier-like protein (AcUCP), and enolase (AcEnol), was up-regulated after cold storage for 12 days at 0 °C. These results imply that selected genes could be related to the development of internal browning observed in cherimoyas after exposure to CI conditions. The information generated in this study provides new clues that may aid in understanding the cherimoya ripening process.M.G.-A. gratefully acknowledges the PBCT-Conicyt (PSD03) project for financial support.Peer reviewe

Regulatory components of shade avoidance syndrome

Competition for light has an important impact on plant development. Plants sense the presence of nearby competitor vegetation as a change in the light quality, i.e. a reduced red to far-red ratio. The responses to shade are generally referred to as the shade avoidance syndrome (SAS), and involve various developmental changes aimed to outgrow the neighbouring plants, and are characterized by enhanced elongation, reduced leaf expansion, decreased branching and ultimately early flowering. These responses can be detrimental in agriculture, because they induce reallocation of resources into elongation growth at the expense of harvestable organs, hence lowering the crop yield. Genetic analyses performed on the SAS response of seedlings have shown the involvement of several transcription factors in the regulation of this response. At least in a few cases, it has been shown that phytochrome rapidly regulates the expression levels of several modulators of hormone responsiveness, rapidly linking shade perception, massive changes in gene expression and modification of hormone sensitivity of the responsive tissues. Here we develop our view on how shade-modulated changes in the transcriptional profiles result in complex SAS responses.Financial support of JB-T and MS-M came from the CSIC (JAEdoc and JAEpre Programmes, respectively). AG and MG received predoctoral fellowships from FPU and FPI programmes, respectively, of the Spanish Ministry of Science and Innovation (MICINN). NC-E received a predoctoral fellowship from the Gobierno de Chile. Our research is supported by grants from the Generalitat de Catalunya (Xarxa de Referència en Biotecnologia and 2009SGR-697) and MICINN–FEDER (BIO2008-00169 and CSD2007-00036).Peer reviewe

The bHLH proteins BEE and BIM positively modulate the shade avoidance syndrome in Arabidopsis seedlings

The shade avoidance syndrome (SAS) refers to a set of plant responses initiated after perception by the phytochromes of light with a reduced red to far-red ratio, indicative of vegetation proximity or shade. These responses, including elongation growth, anticipate eventual shading from potential competitor vegetation by overgrowing neighboring plants or flowering to ensure production of viable seeds for the next generation. In Arabidopsis thaliana seedlings, the SAS includes dramatic changes in gene expression, such as induction of PHYTOCHROME RAPIDLY REGULATED 1 (PAR1), encoding an atypical basic helix-loop-helix (bHLH) protein that acts as a transcriptional co-factor to repress hypocotyl elongation. Indeed, PAR1 has been proposed to act fundamentally as a dominant negative antagonist of conventional bHLH transcription factors by forming heterodimers with them to prevent their binding to DNA or other transcription factors. Here we report the identification of PAR1-interacting factors, including the brassinosteroid signaling components BR-ENHANCED EXPRESSION (BEE) and BES1-INTERACTING MYC-LIKE (BIM), and characterize their role as networked positive regulators of SAS hypocotyl responses. We provide genetic evidence that these bHLH transcriptional regulators not only control plant growth and development under shade and non-shade conditions, but are also redundant in the control of plant viability. Our results suggest that SAS responses are initiated as a consequence of a new balance of transcriptional regulators within the pre-existing bHLH network triggered by plant proximity, eventually causing hypocotyls to elongate.Fellowships or contracts were provided by CSIC (J.B.–T. and M.S.–M.), the Ministerio de Educación (A.G.), the Ministerio de Economía y Competitividad (I.R.–V, and M.G.) and the Gobierno de Chile (N.C.–E.). Research in our laboratories is supported by grants to I.R. from the Italian Ministry of Economy and Finance (Project FaReBio di Qualità) and to J.F.M.–G.'s laboratory (XRB, 2009-SGR697 from the Generalitat de Catalunya and CSD2007-00036, BIO2005-00154, BIO2008-00169 and BIO2011-23489 from Ministerio de Economía y Competitividad/Fondo Europeo de Desarrollo Regional funds).Peer reviewe

Plant proximity perception dynamically modulates hormone levels and sensitivity in Arabidopsis

Shade perception involves altered hormone synthesis and sensitivity. Here, we showed that several shade regulators act as positive and negative modulators of the hypocotyl auxin and/or brassinosteroid-induced elongation. The s hade a voidance s yndrome (SAS) refers to a set of plant responses initiated after perception by the phytochromes of light enriched in far-red colour reflected from or filtered by neighbouring plants. These varied responses are aimed at anticipating eventual shading from potential competitor vegetation. In Arabidopsis thaliana, the most obvious SAS response at the seedling stage is the increase in hypocotyl elongation. Here, we describe how plant proximity perception rapidly and temporally alters the levels of not only auxins but also active brassinosteroids and gibberellins. At the same time, shade alters the seedling sensitivity to hormones. Plant proximity perception also involves dramatic changes in gene expression that rapidly result in a new balance between positive and negative factors in a network of interacting basic helix-loop-helix proteins, such as HFR1, PAR1, and BIM and BEE factors. Here, it was shown that several of these factors act as auxin- and BR-responsiveness modulators, which ultimately control the intensity or degree of hypocotyl elongation. It was deduced that, as a consequence of the plant proximity-dependent new, dynamic, and local balance between hormone synthesis and sensitivity (mechanistically resulting from a restructured network of SAS regulators), SAS responses are unleashed and hypocotyls elongate