Being unable to move away from their places of germination, in order to avoid excess metal-induced damages, plants have to evolve different strategies and complex regulatory mechanisms to survive harsh conditions. While both ROS and auxin are documented to be important in plant response to metal stress, the mechanisms underlying the crosstalk between ROS and auxin in metal stress are poorly understood. In this review, we provide an update on the regulation of plant responses to metal-stress by ROS and auxin signaling pathways, primarily, with a focus on the copper, aluminum and cadmium stress. We aim at surveying the mechanisms underlying how metal stress modulates the changes in auxin distribution and the network of ROS and auxin in plant response to metal stress based on recent studies.
abiotic stress; aluminium; auxin; cadmium; metal stress; copper; heavy metal stress; ROS
Aposematism (warning) signaling is a common defensive mechanism toward predatory or herbivorous animals, i.e., interactions between different trophic levels. I propose that it should be considered at least as a working hypothesis that chemical aposematism operates between certain host plants and their plant predators, parasitic plants, and that although they are also plants, they belong to a higher trophic level. Specific host plant genotypes emit known repelling chemical signals toward parasitic plants, which reduce the level of, slow the directional parasite growth (attack) toward the signaling hosts, or even cause parasitic plants to grow away from them in response to these chemicals. Chemical host aposematism toward parasitic plants may be a common but overlooked defense from parasitic plants.
aposematic; defense; parasitic plants; signaling; trophic levels
Phytohormones are essential regulators of various processes in plant growth and development. Several phytohormones are also known to regulate plant responses to environmental stress and pathogens. Only recently, cytokinins have been demonstrated to play an important role in plant immunity. Increased levels of cytokinins such as trans-zeatin, which are considered highly active, induced resistance against mainly (hemi)biotrophic pathogens in different plant species. In contrast, cis-zeatin is commonly regarded as a cytokinin exhibiting low or no activity. Here we comparatively study the impact of both zeatin isomers on the infection of Nicotiana tabacum by the (hemi)biotrophic microbial pathogen Pseudomonas syringae. We demonstrate a biological effect of cis-zeatin and a differential effect of the two zeatin isomers on symptom development, defense responses and bacterial multiplication.
cytokinin; phytohormone; plant immunity; plant pathogen interaction; Pseudomonas syringae; plant defense; plant resistance; tobacco; zeatin
In Arabidopsis thaliana the CPC-like MYB transcription factors [CAPRICE (CPC), TRIPTYCHON (TRY), ENHANCER OF TRY AND CPC 1, 2, 3/CPC-LIKE MYB 3 (ETC1, ETC2, ETC3/CPL3), TRICHOMELESS 1, 2/CPC-LIKE MYB 4 (TCL1, TCL2/CPL4)] and the bHLH transcription factors [GLABRA3 (GL3) and ENHANCER OF GLABRA 3 (EGL3)] are central regulators of trichome initiation and root-hair differentiation. By transforming the tomato orthologous genes SlTRY (TRY) and SlGL3 (GL3) into Arabidopsis, we demonstrated that SlTRY inhibited trichome initiation and enhanced root-hair differentiation. These results suggest that tomato and Arabidopsis partially use similar transcription factors for epidermal cell differentiation, and that a CPC-like R3 MYB may be a key common regulator of plant trichome and root-hair development. CPC and GL3 are also known to be involved in anthocyanin biosynthesis. Here, we show that anthocyanin accumulation was repressed in the CPC::SlTRY and GL3::SlGL3 transgenic plants, suggesting that SlTRY and SlGL3 can influence anthocyanin pigment synthesis.
tomato; Arabidopsis; R3 MYB; CPC; SlTRY; anthocyanin
Salt impairs cellular morphology and photosynthetic pigment accumulation in the cyanobacterium Fremyella diplosiphon. Recent findings indicated that the impact of salt on cellular morphology was attributable to salt-associated effects on osmotic regulation, as the impact on morphology was reversible when cells were treated with an osmoticum in the presence of salt. The impact of salt on photosynthetic pigment accumulation was associated with ionic effects of salt on the cells, as pigment levels remained low when salt-treated cells were incubated together with an osmoticum or an antioxidant, the latter to mitigate the impact of a salt-associated accumulation of reactive oxygen species. Here, we provide evidence that the transcripts for genes encoding the phycobiliproteins are not reduced in the presence of salt. These results suggest that the negative impact of salt-mediated changes on pigment accumulation occurs post-transcriptionally. A greater understanding of the mechanisms which impact growth of strains such as F. diplosiphon, which harbor pigments that allow low-light and shade-tolerated growth, may facilitate the development or adaptation of such strains as useful for remediation of salt-impacted soils or biofuel production.
cellular morphology; cyanobacteria; osmotic stress; oxidative stress; phycobiliprotein; reactive oxygen species; salinity
Alternative splicing (AS) gives rise to multiple mRNA isoforms from the same gene, providing possibilities to regulate gene expression beyond the level of transcription. In a recent paper in Nucleic Acids Research we used a high resolution RT-PCR based panel to study changes in AS patterns in plants with altered levels of an hnRNP-like RNA-binding protein in Arabidopsis thaliana. Furthermore, we detected significant changes in AS patterns between different Arabidopsis ecotypes. Here we investigated how small changes in ambient temperature affect AS. We found significant changes in AS for 12 of 28 investigated events (43%) upon transfer of Arabidopsis plants from 20°C to 16°C and for 6 of the 28 investigated events (21%) upon transfer from 20°C to 24°C.
alternative splicing; RNA-binding proteins; temperature; posttranscriptional regulation
Post-translational attachment of small ubiquitin-like modifier (SUMO), defined as SUMOylation, has emerged as a new mechanism of protein regulation in plant biology. In plant, SUMOylation has been shown to play crucial roles in a variety of biotic and abiotic stress responses. Recent work using viable mutants with defective SUMOylation have indicated an important role for SUMOylation in a wide range of developmental processes, such as cell division, expansion, survival and differentiation, vegetative growth and reproductive development. This review will summarize the currently emerging information regarding the function of SUMOylation in plant development.
SUMOylation; SUMO proteases; AtMMS21; AtSIZ1; development; Arabidopsis
Brassinosteroids (BRs) regulate various physiological processes, such as tolerance to stresses and root growth. Recently, a connection was reported between BRs and nitric oxide (NO) in plant responses to abiotic stress. Here we present evidence supporting NO functions in BR signaling during root growth process. Arabidopsis seedlings treated with BR 24-epibrassinolide (BL) show increased lateral roots (LR) density, inhibition of primary root (PR) elongation and NO accumulation. Similar effects were observed adding the NO donor GSNO to BR-receptor mutant bri1-1. Furthermore, BL-induced responses in the root were abolished by the specific NO scavenger c-PTIO. The activities of nitrate reductase (NR) and nitric oxide synthase (NOS)-like, two NO generating enzymes were involved in BR signaling. These results demonstrate that BR increases the NO concentration in root cells, which is required for BR-induced changes in root architecture.
Arabidopsis; brassinosteroids; nitric oxide; root morphology
The stability and completeness of male sterility is still a challenge in some male sterile rice lines, especially those of photoperiod/thermo-sensitive genic male sterility (P/TGMS). Leaf color marker is a widely practiced approach to reduce the impact of self-pollinated seeds of male sterile lines. The papst1 is a leaf color mutant. The newly emerged leaves of papst1 are chlorosis and have an impaired photosynthesis. But the other agronomic traits, such as germination rate, duration of maturation and seed weight, are not changed. The papst1/PAPST1 F1 showed the wild-type leaf phenotype. The papst1/PAPST1 F2 progenies displayed an approximately 3:1 segregation ratio of WT phenotype:mutant phenotype (72: 28, χ2 = 0.48, p > 0.05), suggesting that papst1 mutant phenotype is caused by a single repressive gene. Map-based cloning and sequencing analysis revealed that a point mutation was occurred in Os01 g16040 (OsPAPST1). Given these results, the Ospapst1 mutant is a useful mutant for identifying seed purity and authenticity in hybrid rice.
Ospapst1; leaf color; hybrid rice; seed purity
The involvement of potassium (K+)-selective, Shaker-type channels, particularly AKT1, in primary K+ acquisition in roots of higher plants has long been of interest, particularly in the context of low-affinity K+ uptake, at high K+ concentrations, as well as uptake from low-K+ media under ammonium (NH4+) stress. We recently demonstrated that K+ channels cannot mediate K+ acquisition in roots of intact barley (Hordeum vulgare L.) seedlings at low (22.5 µM) external K+ concentrations ([K+]ext) and in the presence of high (10 mM) external NH4+, while the model species Arabidopsis thaliana L. utilizes channels under comparable conditions. However, when external NH4+ was suddenly withdrawn, a thermodynamic shift to passive (channel-mediated) K+ influx was observed in barley and both species demonstrated immediate and dramatic stimulations in K+ influx, illustrating a hitherto unexplored magnitude and rapidity of K+-uptake capacity and plasticity. Here, we expand on our previous work by offering further characterization of channel-mediated K+ fluxes in intact barley, with particular focus on anion effects, root respiration and pharmacological sensitivity and highlight key additions to the current model of K+ acquisition.
barley; Arabidopsis; roots; ammonium; radiotracer; influx; transporters; respiration; anion
The senescence delaying effect of cytokinin is well known, however, the details behind how this process occurs remain unclear. Efforts to improve understanding of this phenomenon have led to the identification in Arabidopsis of specific cytokinin signaling components through which senescence signal responses are regulated. These include the cytokinin receptor (AHK3), the type-B response regulator (ARR2) and the recently identified cytokinin response factor (CRF6). At the mechanistic end of this process, it was found that increased cell-wall invertase activity which occurs in response to cytokinin is both necessary and sufficient for the inhibition of senescence. Yet, a direct link between the signaling and mechanistic steps of a cytokinin regulated senescence process has yet to be demonstrated. This may be in part because the relationship between senescence and primary metabolism implied by the key role of cell-wall invertase is the subject of two apparently opposing bodies of evidence. Here we briefly summarize and propose a model in which cytokinin mediated changes in sink/source relationships leads to delayed senescence which is consistent with existing evidence both for and against sugars as a trigger for developmental senescence.
senescence; cytokinin; leaf; cell wall invertase; carbon allocation; sink/source relationships
Cysteine-rich proteins seem to play important regulatory roles in Medicago truncatula/Sinorhizobium meliloti symbiosis. In particular, a large family of nodule-specific cysteine-rich (NCR) peptides is crucial for the differentiation of nitrogen-fixing bacteroids. The Medicago truncatula N5 protein (MtN5) is currently the only reported non-specific lipid transfer protein necessary for successful rhizobial symbiosis; in addition, MtN5 shares several characteristics with NCR peptides: a small size, a conserved cysteine-rich motif, an N-terminal signal peptide for secretion and antimicrobial activity. Unlike NCR peptides, MtN5 expression is not restricted to the root nodules and is induced during the early phases of symbiosis in root hairs and nodule primordia. Recently, MtN5 was determined to be involved in the regulation of root tissue invasion; while, it was dispensable for nodule primordia formation. Here, we discuss the hypothesis that MtN5 participates in linking the progression of bacterial invasion with restricting the competence of root hairs for infection.
non-specific lipid transfer proteins; MtN5; rhizobial symbiosis; root infection; Medicagotruncatula
The proliferating cell nuclear antigen (PCNA) is a key component of the eukaryotic DNA replication machinery. It also plays an important role in DNA repair mechanisms. Despite the intense scientific research on yeast and human PCNA, information describing the function of this protein in plants is still very limited. In the previous study Arabidopsis PCNA2 but not PCNA1 was proposed to be functionally important in DNA polymerase η-dependent postreplication repair. In addition to the above study, PCNA2 but not PCNA1 was also shown to be necessary for Arabidopsis DNA polymerase λ-dependent oxidative DNA damage bypass. Taking into account the reported differences between PCNA1 and PCNA2, we tested the idea of a possible cooperation between PCNA1 and PCNA2 in the plant cell. In a bimolecular fluorescence complementation assay an interaction between PCNA1 and PCNA2 was observed in the nucleus, as well as in the cytoplasm. This finding, together with our previous results, indicates that PCNA1 and PCNA2 may cooperate in planta by forming homo- and heterotrimeric rings. The observed interaction might be relevant when distinct functions for PCNA1 and PCNA2 are considered.
Arabidopsis thaliana; PCNA; DNA replication; DNA repair; cell cycle
Trehalases are enzymes that carry out the degradation of the non-reducing disaccharide trehalose. Trehalase phylogeny unveiled three major branches comprising those from bacteria; plant and animals; and those from fungal origin. Comparative analysis between several deduced trehalase structures and the crystallographic structure of bacterial trehalase indicated that these enzyme’s structures are highly conserved in spite of the marked differences found at the sequence level. These results suggest a bacterial origin for the trehalases in contrast to an eukaryotic origin, as previously proposed. Trehalases structural analysis showed that they contain six discrete motifs which are characteristic of each phylogenetic group, suggesting a positive evolutionary selection pressure for the structural conservation. Interestingly, trehalases are involved in multiple regulatory functions: In the response against pathogens (plant-pathogen interactions); the regulation of bacterial viability in symbiotic interactions (legume-Rhizobium); carbon partitioning in plants; regulating chitin biosynthesis, as well as energy supply in the hemolymph for flight, in insects. In summary, trehalases seem to have a prokaryotic origin and play an active role in carbon metabolism and other diverse regulatory effects on cell physiology.
Trehalases; structural conservation and distribution; carbon-metabolism
The plant cytoskeleton plays a crucial role in the cells’ growth and development during different developmental stages and it undergoes many rearrangements. In order to describe the arrangements of the F-actin cytoskeleton in root epidermal cells of Arabidopsis thaliana, the recently developed software MicroFilament Analyzer (MFA) was exploited. This software enables high-throughput identification and quantification of the orientation of filamentous structures on digital images in a highly standardized and fast way. Using confocal microscopy and transgenic GFP-FABD2-GFP plants the actin cytoskeleton was visualized in the root epidermis. MFA analysis revealed that during the early stages of cell development F-actin is organized in a mainly random pattern. As the cells grow, they preferentially adopt a longitudinal organization, a pattern that is also preserved in the largest cells. In the evolution from young to old cells, an approximately even distribution of transverse, oblique or combined orientations is always present besides the switch from random to a longitudinal oriented actin cytoskeleton.
actin cytoskeleton; Arabidopsis; development; microfilament analyzer; root epidermis
Recognition of pathogens by plants initiates defense responses including activation of defense-related genes and production of antimicrobial compounds. Recently, we reported that Phytophthora capsici can successfully infect Arabidopsis and revealed interaction specificity among various accession-isolate combinations. We used this novel pathosystem to demonstrate that camalexin, indole glucosinolates (iGS) and salicylic acid (SA) have a role in defense against P. capsici. To further investigate the role of camalexin-, iGS- and SA-related pathways in the differential interaction between Arabidopsis and P. capsici, we monitored expression of marker genes over time during infection. In both compatible and incompatible interactions, induction of expression was detected, but in compatible interactions transcript levels of camalexin and iGS marker genes were higher.
Phytophthora capsici; Arabidopsis; defense signaling; salicylic acid; jasmonic acid; ethylene; camalexin; indole glucosinolates
Abiotic stress factors can interfere with the emission of herbivore-induced plant volatile organic compounds (VOCs) and thus disrupt chemical communication channels between plants and other organisms. We investigated whether copper (Cu) stress alone or in conjunction with insect damage modifies the kinetics of (1) VOCs, (2) the VOC-inducing phytohormone jasmonic acid (JA) and (3) its putative antagonist salicylic acid (SA). Hydroponically grown Zea mays exposed to 10 and 80 µM of Cu showed no increases in JA or VOC levels in the absence of herbivory. However when challenged by herbivores, Cu (80 µM) caused ROS generation in root tissues and primed for increased JA accumulation and VOC emission in leaves. SA synthesis was equally primed but higher concentrations were also apparent before insects started feeding. In contrast, plants grown at 10 µM Cu did not differ from controls. These results show that abiotic and biotic stresses result in concentration-dependent, non-additive defense responses. Further support is given to the notion that JA-SA antagonism is absent in Z. mays.
heavy metals; Zea mays; induced indirect defense; herbivory; crosstalk
Jasmonic acid (JA) and salicylic acid (SA) play important roles in plant defense systems. JA and SA signaling pathways interact antagonistically in dicotyledonous plants, but, the status of crosstalk between JA and SA signaling is unknown in monocots. Our rice microarray analysis showed that more than half of the genes upregulated by the SA analog BTH are also upregulated by JA, suggesting that a major portion of the SA-upregulated genes are regulated by JA-dependent signaling in rice. A common defense system that is activated by both JA and SA is thus proposed which plays an important role in pathogen defense responses in rice.
common defense system, induced resistance; jasmonic acid; salicylic acid; rice
Ascorbic acid (AsA) is a major antioxidant and plays an important role in plant growth and development. There are two aspects to improve AsA content, including de novo synthesis and recycling from its oxidized form. However, the knowledge of regulatory mechanisms of AsA synthesis pathways and metabolism still remains limited. We have determined that AsA synthesis process was modulated on both transcriptional and post-transcriptional levels in Arabidopsis. GDP-mannose pyrophosphorylase (VTC1) is the initial AsA biosynthetic enzyme in L-galactose pathway, we previously showed that Arabidopsis ERF98 transcriptionally activates gene expression of VTC1 to improve AsA content and respond salt stress; recent discovery of the interaction between photomorphogenic factor COP9 signalosome (CSN) subunit CSN5B and VTC1 indicates that CSN5B promotes VTC1 degradation in the dark, which keeps the change of AsA content from day to night. This mini-review integrates previous reports and recent evidence to better understand the regulatory mechanisms involved in AsA synthesis.
ascorbic acid; L-galactose pathway; regulation; VTC1; ERF98; CSN5B
The deposition of callose, a (1,3)-β-glucan cell wall polymer, can play an essential role in the defense response to invading pathogens. We could recently show that Arabidopsis thaliana lines with an overexpression of the callose synthase gene PMR4 gained complete penetration resistance to the adapted powdery mildew Golovinomyces cichoracearum and the non-adapted powdery mildew Blumeria graminis f. sp hordei. The penetration resistance is based on the transport of the callose synthase PMR4 to the site of attempted fungal penetration and the subsequent formation of enlarged callose deposits. The deposits differed in their total diameter comparing both types of powdery mildew infection. In this study, further characterization of these callose deposits revealed that size differences were especially pronounced in the core region of the deposits. This suggests that specific, pathogen-dependent factors exist, which might regulate callose synthase transport to the core region of forming deposits.
(1,3)-β-glucan; Arabidopsis thaliana; AtGSL5; PMR4; callose synthase; cell wall; papillae; plant defense; powdery mildew
RNAs fold into intricate structures that are determined by specific base pairing interactions encoded within their primary sequences. Recently, a number of transcriptome-wide studies have suggested that RNA secondary structure is a potent cis-acting regulator of numerous post-transcriptional processes in viruses and eukaryotes. However, the need for experimentally-based structure determination methods has not been well addressed. Here, we show that the regulatory significance of Arabidopsis RNA secondary structure is revealed specifically through high-throughput, sequencing-based, structure mapping data, not by computational prediction. Additionally, we find that transcripts with similar levels of secondary structure in their UTRs (5' or 3') or CDS tend to encode proteins with coherent functions. Finally, we reveal that portions of mRNAs encoding predicted protein domains are significantly more structured than those specifying inter-domain regions. In total, our findings show the utility of high-throughput, sequencing-based, structure-mapping approaches and suggest that mRNA folding regulates protein maturation and function.
RNA genomics; mRNA secondary structure; protein domains; post-transcriptional regulation
The accumulation of unfolded proteins in the ER lumen induces intracellular signaling mediated by the ER stress sensor protein IRE1. Our recent study identified a new common cis-element of ER stress-responsive genes (such as rice BiP paralogs and WRKY45) that were regulated via an IRE1-dependent pathway. ER stress-responsive cis-elements had been expected to be conserved between plants and mammals. However, contrary to expectations, sequences of the plant cis-element, pUPRE-II, were not identical to those of its mammalian counterpart. Additionally, pUPRE-II also interacted with another ER stress sensor protein and mediated multiple signaling pathways. Here, we provide a summary of the results that suggest the complicated mechanism underlying the regulation of ER stress-responsive gene expression in plants.
endoplasmic reticulum stress; unfolded protein response; transcription factors; cis-elements; plants; Oryza sativa
Cross-talk between hormones is required for plant response to developmental cues and environmental stresses. This cross-talk is achieved through several regulators located in convergence point of distinct hormonal signaling. In plant defense responses, salicylic acid and jasmonic acid affect each other in antagonistic manner. In a recent study we showed that AtMYB44 transcription factor positively regulates SA-mediated defense expression and enhanced resistance to Pst DC3000. On the other hand, AtMYB44 negatively regulates expression of JA-mediated defense gene expression and downregulated resistance to Alternaria brassicicola. Effects of AtMYB44 in SA- and JA-mediated defense responses were achieved through direct regulation of WRKY70 expression which acts as an integrator of cross-talk between SA and JA in plant defense responses. Here we provide further evidence that AtMYB44 regulates defense responses by transcriptional activation of downstream gene, WRKY70. This result shows that AtMYB44 is an integrator of cross-talk between SA and JA in plant defense responses.
AtMYB44; WRKY70; jasmonic acid; salicylic acid; MYB transcription factor
The GARP domain is a single Myb-related DNA-binding domain found in plant transcription factors. Proteins containing the GARP domain (GARP family proteins) are suggested to be involved in the regulation of various physiological processes through their interactions with ostensibly different DNA sequences. Our recent study on a nitrate-inducible gene encoding a GARP family protein, referred to as NIGT1 (Nitrate-Inducible, GARP-type Transcriptional Repressor 1), not only suggests a previously unidentified role for the GARP family proteins in higher plants but also provides a hypothesis for why NIGT1 can show dual specificity on DNA binding and why respective GARP family proteins can recognize very different DNA sequences.
GARP transcription factor; dual DNA recognition; DNA binding specificity; nitrate induction
We recently reported that the Medicago WOX gene, STENOFOLIA (STF), acts as a transcriptional repressor in regulating leaf blade outgrowth. By using the Nicotiana sylvestris bladeless lam1 mutant as a genetic tool, we showed that the WUS-box, which is conserved among WUS clade WOX genes, is partly responsible for the repressive activity of STF. All members of the modern/WUS clade genes (WUS, WOX1-WOX7) in Arabidopsis that contain intact WUS-box can substitute for STF/LAM1 function while the intermediate and ancient clade members including WOX9,WOX11 and WOX13 cannot, due to lack of the intact WUS-box. Taken together, our results reveal a conserved repression mechanism playing a central role in cell proliferation conferred to the evolutionarily dynamic WOX gene family with acquisition of a repressor domain.
WOX family; STF; lam1; repressor; WUS-box; leaf blade outgrowth; cell proliferation