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In plants, salicylic acid (SA) hydroxylation regulates SA homoeostasis, playing an essential role during plant development and response to pathogens. This reaction is catalysed by SA hydroxylase enzymes, which hydroxylate SA producing 2,3‐dihydroxybenzoic acid (2,3‐DHBA) and/or 2,5‐dihydroxybenzoic acid (2,5‐DHBA). Several SA hydroxylases have recently been identified and characterised from different plant species, but no such activity has yet been reported in maize. In this work, we describe the identification and characterisation of a new SA hydroxylase in maize plants. This enzyme, with high sequence similarity to previously described SA hydroxylases from Arabidopsis and rice, converts SA into 2,5‐DHBA; however, it has different kinetic properties to those of previously characterised enzymes, and it also catalysers the conversion of the flavonoid dihydroquercetin into quercetin in in vitro activity assays, suggesting that the maize enzyme may have different roles in vivo to those previously reported from other species. Despite this, ZmS5H can complement the pathogen resistance and the early senescence phenotypes of Arabidopsis s3h mutant plants. Finally, we characterised a maize mutant in the S5H gene (s5hMu) that has altered growth, senescence and increased resistance against Colletotrichum graminicola infection, showing not only alterations in SA and 2,5‐DHBA but also in flavonol levels. Together, the results presented here provide evidence that SA hydroxylases in different plant species have evolved to show differences in catalytic properties that may be important to fine tune SA levels and other phenolic compounds such as flavonols, to regulate different aspects of plant development and pathogen defence. Summary statement: Maize plants express an enzyme of the 2‐oxoglutarate‐dependent oxygenase family with unique kinetic properties, having salicylic acid hydroxylase activity but it can also convert flavonols from dihydroflavonols. Zm2‐ODD7 confers increased resistance against pathogens and regulates senescence. [ABSTRACT FROM AUTHOR]

SUMMARY: Mediator 17 (MED17) is a subunit of the Mediator complex that regulates transcription initiation in eukaryotic organisms. In yeast and humans, MED17 also participates in DNA repair, physically interacting with proteins of the nucleotide excision DNA repair system, but this function in plants has not been investigated. We studied the role of MED17 in Arabidopsis plants exposed to UV‐B radiation. Our results demonstrate that med17 and OE MED17 plants have altered responses to UV‐B, and that MED17 participates in various aspects of the DNA damage response (DDR). Comparison of the med17 transcriptome with that of wild‐type (WT) plants showed that almost one‐third of transcripts with altered expression in med17 plants were also changed by UV‐B exposure in WT plants. Increased sensitivity to DNA damage after UV‐B in med17 plants could result from the altered regulation of UV‐B responsive transcripts but MED17 also physically interacts with DNA repair proteins, suggesting a direct role of this Mediator subunit during repair. Finally, we show that MED17 is necessary to regulate the DDR activated by ataxia telangiectasia and Rad3 related (ATR), and that programmed cell death 5 (PDCD5) overexpression reverts the deficiencies in DDR shown in med17 mutants. Our data demonstrate that MED17 is an important regulator of DDR after UV‐B irradiation in Arabidopsis. Significance Statement: In Arabidopsis, MED17 regulates the DNA damage response after UV‐B exposure transcriptionally modulating the expression of genes and possibly also physically interacting with DNA repair proteins. [ABSTRACT FROM AUTHOR]

In Arabidopsis thaliana plants, exposure to UV-B induces an inhibition of primary root elongation. Different mutants have been isolated that are deficient in this response; however, little is known about the cellular and molecular mechanisms that regulate inhibition of root elongation in seedlings exposed to UV-B. In this work, we investigated the effect UV-B irradiation of different organs on primary root elongation. Our results demonstrate that irradiation of the leaves and shoots only induce a partial inhibition of primary root elongation, while when only roots are exposed to this radiation, primary root inhibition is similar as that measured when the complete seedling is irradiated. The consequences of exposure at different root developmental stages and times after the end of the treatment was also studied. We here show that inhibition of primary root elongation is a consequence of a decrease in cell proliferation in the meristematic zone of the primary roots, while the elongation zone size is not affected by the treatment. The decrease in cell number after UV-B exposure is partially compensated by an increase in cell length in the root meristem; however, this compensation is not enough to maintain the meristem size. We also here demonstrate that, similarly as what occurs in developing leaves, GROWTH REGULATING FACTOR 3 (GRF3) transcription factor regulates cell proliferation in UV-B irradiated roots; however, and in contrast to what occurs in the leaves, this response does not depend on the presence of MITOGEN ACTIVATED PROTEIN KINASE 3 (MPK3). Inhibition of primary root elongation by UV-B under our experimental conditions is also independent of the UV-B photoreceptor UV RESISTANT LOCUS 8 (UVR8) or ATAXIA TELANGIECTASIA MUTATED (ATM); but a deficiency in ATM AND RAD3-RELATED (ATR) expression increases UV-B sensitivity in the roots. Finally, our data demonstrate that UV-B affects primary root growth in various Arabidopsis accessions, showing different sensitivities to this radiation. [ABSTRACT FROM AUTHOR]

SUMMARY: Ultraviolet (UV)B radiation affects plant growth inhibiting cell proliferation. This inhibition is in part controlled by the activity of transcription factors from the E2F family. In particular, the participation of E2Fc and E2Fe in UV‐B responses in Arabidopsis plants was previously reported. However, the E2Fa and E2Fb contribution to these processes has still not been investigated. Thus, in this work, we provide evidence that, in Arabidopsis, both E2Fa and E2Fb control leaf size under UV‐B conditions without participating in the repair of cyclobutane pyrimidine dimers in the DNA. Nevertheless, in UV‐B‐exposed seedlings, E2Fa, but not E2Fb, regulates primary root elongation, cell proliferation, and programmed cell death in the meristematic zone. Using e2fa mutants that overexpress E2Fb, we showed that the role of E2Fa in the roots could not be replaced by E2Fb. Finally, our results show that E2Fa and E2Fb differentially regulate the expression of genes that activate the DNA damage response and cell cycle progression, both under conditions without UV‐B and after exposure. Overall, we showed that both E2Fa and E2Fb have different and non‐redundant roles in developmental and DNA damage responses in Arabidopsis plants exposed to UV‐B. [ABSTRACT FROM AUTHOR]

Flavonoids are plant specialized metabolites that consist of one oxygenated and two aromatic rings. Different flavonoids are grouped according to the oxidation degree of the carbon rings; they can later be modified by glycosylations, hydroxylations, acylations, methylations, or prenylations. These modifications generate a wide collection of different molecules which have various functions in plants. All flavonoids absorb in the UV wavelengths, they mostly accumulate in the epidermis of plant cells and their biosynthesis is generally activated after UV exposure. Therefore, they have been assumed to protect plants against exposure to radiation in this range. Some flavonoids also absorb in other wavelengths, for example anthocyanins, which absorb light in the visible part of the solar spectrum. Besides, some flavonoids show antioxidant properties, that is, they act as scavengers of reactive oxygen species that could be produced after high fluence UV exposure. However, to date most reports were based on in vitro studies, and there is very little in vivo evidence of how their roles are carried out. In this review we first summarize the biosynthetic pathway of flavonoids and their characteristics, and we describe recent advances on the investigation of the role of three of the most abundant flavonoids: flavonols, flavones, and anthocyanins, protecting plants against UV exposure and high light exposure. We also present examples of how using UV‐B supplementation to increase flavonoid content, is possible to improve plant nutritional and pharmaceutical values. [ABSTRACT FROM AUTHOR]

Abstract Background Plants living at high altitudes are typically exposed to elevated UV-B radiation, and harbor mechanisms to prevent the induced damage, such as the accumulation of UV-absorbing compounds. The maize R2R3-MYB transcription factor P1 controls the accumulation of several UV-B absorbing phenolics by activating a subset of flavonoid biosynthetic genes in leaves of maize landraces adapted to high altitudes. Results Here, we studied the UV-B regulation of P1 in maize leaves of high altitude landraces, and we investigated how UV-B regulates P1binding to the CHS promoter in both low and high altitude lines. In addition, we analyzed whether the expansion in the P1 expression domain between these maize landraces and inbred lines is associated to changes in the molecular structure of the proximal promoter, distal enhancer and first intron of P1. Finally, using transient expression experiments in protoplasts from various maize genotypes, we investigated whether the different expression patterns of P1 in the high altitude landraces could be attributed to trans- or cis-acting elements. Conclusions Together, our results demonstrate that, although differences in cis-acting elements exist between the different lines under study, the different patterns of P1 expression are largely a consequence of effects in trans.

Abstract Background Under normal solar fluence, UV-B damages macromolecules, but it also elicits physiological acclimation and developmental changes in plants. Excess UV-B decreases crop yield. Using a treatment twice solar fluence, we focus on discovering signals produced in UV-B-irradiated maize leaves that translate to systemic changes in shielded leaves and immature ears. Results Using transcriptome and proteomic profiling, we tracked the kinetics of transcript and protein alterations in exposed and shielded organs over 6 h. In parallel, metabolic profiling identified candidate signaling molecules based on rapid increase in irradiated leaves and increased levels in shielded organs; pathways associated with the synthesis, sequestration, or degradation of some of these potential signal molecules were UV-B-responsive. Exposure of just the top leaf substantially alters the transcriptomes of both irradiated and shielded organs, with greater changes as additional leaves are irradiated. Some phenylpropanoid pathway genes are expressed only in irradiated leaves, reflected in accumulation of pathway sunscreen molecules. Most protein changes detected occur quickly: approximately 92% of the proteins in leaves and 73% in immature ears changed after 4 h UV-B were altered by a 1 h UV-B treatment. Conclusions There were significant transcriptome, proteomic, and metabolomic changes under all conditions studied in both shielded and irradiated organs. A dramatic decrease in transcript diversity in irradiated and shielded leaves occurs between 0 h and 1 h, demonstrating the susceptibility of plants to short term UV-B spikes as during ozone depletion. Immature maize ears are highly responsive to canopy leaf exposure to UV-B.

The genome of plants is organized into chromatin. The chromatin structure regulates the rates of DNA metabolic processes such as replication, transcription, DNA recombination, and repair. Different aspects of plant growth and development are regulated by changes in chromatin status by the action of chromatin-remodeling activities. Recent data have also shown that many of these chromatin-associated proteins participate in different aspects of the DNA damage response, regulating DNA damage and repair, cell cycle progression, programmed cell death, and entry into the endocycle. In this review, we present different examples of proteins and chromatin-modifying enzymes with roles during DNA damage responses, demonstrating that rapid changes in chromatin structure are essential to maintain genome stability. [ABSTRACT FROM AUTHOR]

Plant ribosomal proteins play universal roles in translation, although they are also involved in developmental processes and hormone signaling pathways. Among Arabidopsis RPL10 family members, RPL10A exhibits the highest expression during germination and early development, suggesting that RPL10A is the main contributor to these processes. In this work, we first analyzed RPL10A expression pattern in Arabidopsis thaliana using transgenic RPL10Apro:GUS plants. The gene exhibits a ubiquitous expression pattern throughout the plant, but it is most strongly expressed in undifferentiated tissues. Interestingly, gene expression was also detected in stomatal cells. We then examined protein function during seedling establishment and abscisic acid (ABA) response. Heterozygous rpl10A mutant plants show decreased ABA-sensitivity during seed germination, are impaired in early seedling and root development, and exhibit reduced ABA-inhibition of stomatal aperture under light conditions. Overexpression of RPL10A does not affect the germination and seedling growth, but RPL10A -overexpressing lines are more sensitive to ABA during early plant development and exhibit higher stomatal closure under light condition both with and without ABA treatment than wild type plants. Interestingly, RPL10A expression is induced by ABA. Together, we conclude that RPL10A could act as a positive regulator for ABA-dependent responses in Arabidopsis plants. [ABSTRACT FROM AUTHOR]

CURLY LEAF (CLF) encodes the methyltransferase subunit of the Polycomb Repressor Complex 2 (PRC2), which regulates the expression of target genes through H3K27 trimethylation. We isolated a new CLF mutant allele (clf-78) using a genetic screen designed to identify microRNA (miRNA) deficient mutants. CLF mutant plants showed impaired miRNA activity caused by increased ubiquitination and enhanced degradation of ARGONAUTE 1 (AGO1) in specific tissues. Such CLF-mediated AGO1 regulation was evident when plants were exposed to UV radiation, which caused increased susceptibility of clf mutants to some UV-induced responses. Furthermore, we showed that CLF directly regulates FBW2 , which in turn triggers AGO1 degradation in the clf mutants. Interestingly, AGO1 bound to a target appeared particularly prone to degradation in the mutant plants, a process that was exacerbated when the complex bound a non-cleavable target. Thus, prolonged AGO1–target interaction seems to favor AGO1 degradation, suggesting that non-cleavable miRNA targets may overcome translation inhibition by modulating AGO1 stability in plants. This study found that CURLY LEAF regulates AGO1 degradation and thus impairs microRNA activity in plants. This effect is observed in specific cellular niches during normal conditions as well as during UV stress. It was also found that prolonged AGO1-target interaction favors protein degradation, suggesting a potential pathway that releases translation-inhibited and non-cleavable targets of microRNA-mediated silencing. [ABSTRACT FROM AUTHOR]

Chromatin assembly factor‐1 (CAF‐1) is a histone H3/H4 chaperone that participates in DNA and chromatin interaction processes. In this manuscript, we show that organs from CAF‐1 deficient plants respond differently to ultraviolet‐B (UV‐B) radiation than to other genotoxic stresses. For example, CAF‐1 deficient leaves tolerate better UV‐B radiation, showing lower cyclobutane pyrimidine dimer (CPD) accumulation, lower inhibition of cell proliferation, increased cell wall thickness, UV‐B absorbing compounds, and ploidy levels, whereas previous data from different groups have shown that CAF‐1 mutants show shortening of telomeres, loss of 45S rDNA, and increased homologous recombination, phenotypes associated to DNA breaks. Interestingly, CAF‐1 deficient roots show increased inhibition of primary root elongation, with decreased meristem size due to a higher inhibition of cell proliferation after UV‐B exposure. The decrease in root meristem size in CAF‐1 mutants is a consequence of defects in programmed cell death after UV‐B exposure. Together, we provide evidence demonstrating that root and shoot meristematic cells may have distinct protection mechanisms against CPD accumulation by UV‐B, which may be linked with different functions of the CAF‐1 complex in these different organs. We provide evidence that demonstrate that ultraviolet‐B (UV‐B) radiation that produce cyclobutane pyrimidine dimers activate different DNA damage response mechanisms than other DNA‐damaging agents, which require chromatin assembly factor‐1 activity, and that root and shoot meristematic cells may have distinct protection mechanisms against DNA damage by UV‐B. [ABSTRACT FROM AUTHOR]