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Nitrogen is one of the important nutrients required for plant growth and development. There is increasing evidences that almost all types of nitrogen metabolites affect, at least to some extent, auxin content and/or signaling in plants, which in turn affects seed germination, plant root elongation, gravitropism, leaf expansion and floral transition. This opinion focuses on the roles of nitrogen metabolites, NO3−, NH4+, tryptophan and NO and their synergistic effects with auxin on plant growth and development. Nitrate reductase (NR) converts nitrate into nitrite, and was roughly positive-correlated with the root auxin level, suggesting a crosstalk between nitrate signaling and auxin signaling. Abscisic Acid Responsive Element Binding Factor 3 (AFB3) and Tryptophan Aminotransferase of Arabidopsis 1 (TAA1) are also the key enzymes involved in nitrogen metabolite-regulated auxin biosynthesis. Recent advances in the crosstalk among NO3−, NH4+, tryptophan and NO in regulation to NR, AFB3 and TAA1 are also summarized.

It has been well demonstrated that melatonin plays an important protective role in photosynthesis of plants under various environmental stresses, while the detailed mechanisms by which melatonin protects photosystem II (PSII) under environmental stress are still unclear. In the study, the effects of melatonin on photosynthetic efficiency, energy dissipation, PSII protein composition, and reversible phosphorylation of thylakoid proteins were investigated in wheat plants under osmotic stress. The results showed that osmotic stress significantly reduced pigment content, photochemical efficiency of PSII, oxygen-evolving activity, and dissipation of excess excitation energy, while 25 μM melatonin applications greatly alleviated their decline under osmotic stress. Western blot data of PSII proteins revealed that melatonin upregulated the levels of D1, Lhcb5, Lhcb6, PsbQ, and PsbS proteins in wheat exposed to osmotic stress. In addition, thylakoid membrane proteins were strongly phosphorylated in wheat under osmotic stress with or without melatonin. Furthermore, the results from PSII protein dephosphorylation showed that exogenous melatonin promoted the dephosphorylation of LCHII, CP43, and D1 under osmotic stress. Therefore, our findings suggest that melatonin can provide an effective protection for the photosynthetic apparatus by the regulation of PSII proteins and the reversible phosphorylation of thylakoid proteins under drought stress.

Bundle sheath cells play a crucial role in photosynthesis in C4 plants, but the structure and function of photosystem II (PSII) in these cells is still controversial. Photoprotective roles of bundle sheath chloroplasts at the occurrence of environmental stresses have not been investigated so far. Non-photochemical quenching (NPQ) of chlorophyll a fluorescence is the photoprotective mechanism that responds to a changing energy balance in chloroplasts. In the present study, we found a much higher NPQ in bundle sheath chloroplasts than in mesophyll chloroplasts under a drought stress. This change was accompanied by a more rapid dephosphorylation of light-harvesting complex II (LHCII) subunits and a greater increase in PSII subunit S (PsbS) protein abundance than in mesophyll cell chloroplasts. Histochemical staining of reactive oxygen species (ROS) suggested that the high NPQ may be one of the main reasons for the lower accumulation of ROS in bundle sheath chloroplasts. This may maintain the stable functioning of bundle sheath cells under drought condition. These results indicate that the superior capacity for dissipation of excitation energy in bundle sheath chloroplasts may be an environmental adaptation unique to C4 plants.

After nitrogen treatments, plant leaves become narrower and thicker, and the chlorophyll content increases. However, the molecular mechanisms behind these regulations remain unknown. Here, we found that the changes in leaf width and thickness were largely compromised in the shade avoidance 3 (sav3) mutant. The SAV3 gene encodes an amino-transferase in the auxin biosynthesis pathway. Thus, the crosstalk between shade and nitrogen in Arabidopsis leaf development was investigated. Both hypocotyl elongation and leaf expansion promoted by the shade treatment were reduced by the high-N treatment; high-N-induced leaf narrowing and thickening were reduced by the shade treatment; and all of these developmental changes were largely compromised in the sav3 mutant. Shade treatment promoted SAV3 expression, while high-N treatment repressed SAV3 expression, which then increased or decreased auxin accumulation in cotyledons/leaves, respectively. SAV3 also regulates chlorophyll accumulation and nitrogen assimilation and thus may function as a master switch responsive to multiple environmental stimuli.

Land plants live in a crisis-filled environment and the fluctuation of sunlight intensity often causes damage to photosynthetic apparatus. Phyto-melatonin is an effective bioactive molecule that helps plants to resist various biotic and abiotic stresses. In order to explore the role of melatonin under high light stress, we investigated the effects of melatonin on anti-oxidative system and photosynthesis of Arabidopsis thaliana under high light. Results showed that exogenous melatonin increased photosynthetic rate and protected photosynthetic proteins under high light. This was mainly owing to the fact that exogenous melatonin effectively decreased the accumulation of reactive oxygen species and protected integrity of membrane and photosynthetic pigments, and reduced cell death. Taken together, our study promoted more comprehensive understanding in the protective effects of exogenous melatonin under high light.

Leaf is an important organ for higher plants, and the shape of it is one of the crucial traits of crops. In this study, we investigated a unique aberrant leaf morphology trait in a mutational rapeseed material, which displayed ectopic blade-like outgrowths on the adaxial side of leaf. The abnormal line 132000B-3 was crossed with the normal line 827-3. Based on the F2:3 family, we constructed two DNA pools (normal pool and abnormal pool) by the bulked segregant analysis (BSA) method and performed whole genome re-sequencing (WGR), obtaining the single-nucleotide polymorphism (SNP) and insertion/deletion (InDel) data. The SNP-index method was used to calculate the Δ(SNP/InDel-index), and then an association region was identified on chromosome A10 with a length of 5.5 Mbp, harboring 1048 genes totally. Subsequently, the fine mapping was conducted by using the penta-primer amplification refractory mutation system (PARMS), and the associated region was narrowed down to a 35.1-kbp segment, containing only seven genes. These seven genes were then analyzed according to their annotations and finally, BnA10g0422620 and BnA10g0422610, orthologs of LATE MERISTEM IDENTITY1 (LMI1) gene from Arabidopsis and REDUCED COMPLEXITY (RCO) gene from its relative Cardamine hirsuta, respectively, were identified as the candidate genes responding to this blade-like outgrowth trait in rapeseed. This study provides a novel perspective into the leaf formation in Brassica plants.

The protective role of melatonin in plants against various abiotic stresses have been widely demonstrated, but poorly explored in organ-specific responses and the transmission of melatonin signals across organs. In this study, the effects of melatonin with the root-irrigation method and the leaf-spraying method on the antioxidant system and photosynthetic machinery in maize seedlings under drought stress were investigated. The results showed that drought stress led to the rise in reactive oxygen species (ROS), severe cell death, and degradation of D1 protein, which were mitigated by the melatonin application. The application of melatonin improved the photosynthetic activities and alleviated the oxidative damages of maize seedlings under the drought stress. Compared with the leaf-spraying method, the root-irrigation method was more effective on enhancing drought tolerance. Moreover, maize seedlings made organ-specific physiological responses to the drought stress, and the physiological effects of melatonin varied with the dosage, application methods and plant organs. The signals of exogenous melatonin received by roots could affect the stress responses of leaves, and the melatonin signals perceived by leaves also led to changes in physiological metabolisms in roots under the stress. Consequently, the whole seedlings coordinated the different parts and made a systemic acclimation against the drought stress. Melatonin as a protective agent against abiotic stresses has a potential application prospect in the agricultural industry.

Higher plants contain a small, 5-member family of Rieske non-heme oxygenases that comprise the inner plastid envelope protein TIC55, phaeophorbide a oxygenasee (PAO), chlorophyllide a oxygenase (CAO), choline monooxygenase, and a 52 kDa protein (PTC52) associated with the precursor NADPH:protochlorophyllide (Pchlide) oxidoreductase A (pPORA) A translocon (PTC). Some of these chloroplast proteins have documented roles in chlorophyll biosynthesis (CAO) and degradation (PAO and TIC55), whereas the function of PTC52 remains unresolved. Biochemical evidence provided here identifies PTC52 as Pchlide a oxygenase of the inner plastid envelope linking Pchlide b synthesis to pPORA import. Protochlorophyllide b is the preferred substrate of PORA and its lack no longer allows pPORA import. The Pchlide b-dependent import pathway of pPORA thus operates in etiolated seedlings and is switched off during greening. Using dexamethasone-induced RNA interference (RNAi) we tested if PTC52 is involved in controlling both, pPORA import and Pchlide homeostasis in planta. As shown here, RNAi plants deprived of PTC52 transcript and PTC52 protein were unable to import pPORA and died as a result of excess Pchlide a accumulation causing singlet oxygen formation during greening. In genetic studies, no homozygous ptc52 knock-out mutants could be obtained presumably as a result of embryo lethality, suggesting a role for PTC52 in the initial greening of plant embryos. Phylogenetic studies identified PTC52-like genes amongst unicellular photosynthetic bacteria and higher plants, suggesting that the biochemical function associated with PTC52 may have an ancient evolutionary origin. PTC52 also harbors conserved motifs with bacterial oxygenases such as the terminal oxygenase component of 3-ketosteroid 9-alpha-hydroxylase (KshA) from Rhodococcus rhodochrous. 3D-modeling of PTC52 structure permitted the prediction of amino acid residues that contribute to the substrate specificity of this enzyme. In vitro-mutagenesis was used to test the predicted PTC52 model and provide insights into the reaction mechanism of this Rieske non-heme oxygenase.

Chlorophyll fluorescence measurements have been mainly applied to investigate the functioning of the photosynthetic apparatus in the diagnosis of environmental stress. Moss is sensitive to several abiotic stresses and is considered an environmental indicator. Therefore, moss chlorophyll fluorescence can be as a visual parameter applicable for monitoring heavy metal contaminants in water. Different from previous studies with value changes of chlorophyll fluorescence in mosses, we suggest that phenotypes with anthocyanin accumulation pattern and chlorosis pattern and colors of chlorophyll fluorescence images of the maximum efficiency of PSII photochemistry (Fv/Fm) and the quantum yield of PSII electron transport (ΦPSII) could reflect metal species groups and concentrations roughly. And we further indicated that Cr(III) and Cr(VI) could be monitored distinguishably according to the non-photochemical quenching (NPQ) fluorescence of sporadic purple and sporadic lavender images, respectively. It is interesting that the fluorescence color patterns were nearly the same for all treatment concentrations. This perspective provides additional data of chlorophyll fluorescence changes in moss under cold, heat, salinity, high light or osmotic stress. Only heat stress and high light have significant effects on the fluorescence parameters of Fv/Fm and ΦPSII. In contrast, mosses are less sensitive to short-term cold, salinity, and osmotic stress. While NPQ decreases rapidly under the osmotic stress. Nevertheless, heat stress, high light or osmotic stress does not usually co-occur in the place where the moss grows. Estimation through moss chlorophyll fluorescence color patterns is still a rapid and non-invasive method to monitor heavy metal pollutions in water.

In intercropping systems shading conditions significantly impair the seed yield and quality of soybean, and rarely someone investigated the minimum amount of light requirement for soybean growth and development. Therefore, it is an urgent need to determine the threshold light intensity to ensure sustainable soybean production under these systems. An integrated approach combining morphology, physiology, biochemistry and genetic analysis was undertaken to study the light intensity effects on soybean growth and development. A pot experiment was set up in a growth chamber under increasing light intensity treatments of 100 (L100), 200 (L200), 300 (L300), 400 (L400), and 500 (L500) μmol m−2 s−1. Compared with L100, plant height, hypocotyl length, and abaxial leaf petiole angle were decreased, biomass, root:shoot ratio, and stem diameter were increased, extremum was almost observed in L400 and L500. Leaf petiole movement and leaf hyponasty in each treatment has presented a tendency to decrease the leaf angle from L500 to L100. In addition, the cytochrome content (Chl a, Chl b, Car), net photosynthetic rate, chlorophyll fluorescence values of Fv/Fm, Fv′/Fm′, ETR, ΦPSII, and qP were increased as the light intensity increased, and higher values were noted under L400. Leaf microstructure and chloroplast ultrastructure positively improved with increasing light intensity, and leaf-thickness, palisade, and spongy tissues-thickness were increased by 105, 90, and 370%, under L500 than L100. Moreover, the cross-sectional area of chloroplast (C) outer membrane and starch grains (S), and sectional area ratio (S:C) was highest under L400 and L500, respectively. Compared to L100, the content of starch granules increased by 35.5, 122.0, 157.6, and 145.5%, respectively in L400. The same trends were observed in the enzyme activity of sucrose-synthase, sucrose phosphate synthase, starch synthase, rubisco, phosphoenol pyruvate carboxykinase, and phosphoenol pyruvate phosphatase. Furthermore, sucrose synthesis-related genes were also up-regulated by increasing light intensity, and the highest seed yield and yield related parameters were recorded in the L400. Overall, these results suggested that 400 and 500 μmol m−2 s−1 is the optimum light intensity which positively changed the leaf orientation and adjusts leaf angle to perpendicular to coming light, consequently, soybean plants grow well under prevailing conditions.

It has been known that PSI and PSII supercomplexes are involved in the linear and cyclic electron transfer, dynamics of light capture, and the repair cycle of PSII under environmental stresses. However, evolutions of photosystem (PS) complexes from evolutionarily divergent species are largely unknown. Here, we improved the blue native polyacrylamide gel electrophoresis (BN-PAGE) separation method and successfully separated PS complexes from all terrestrial plants. It is well known that reversible D1 protein phosphorylation is an important protective mechanism against oxidative damages to chloroplasts through the PSII photoinhibition-repair cycle. The results indicate that antibody-detectable phosphorylation of D1 protein is the latest event in the evolution of PS protein phosphorylation and occurs exclusively in seed plants. Compared to angiosperms, other terrestrial plant species presented much lower contents of PS supercomplexes. The amount of light-harvesting complexes II (LHCII) trimers was higher than that of LHCII monomers in angiosperms, whereas it was opposite in gymnosperms, pteridophytes, and bryophytes. LHCII assembly may be one of the evolutionary characteristics of vascular plants. In vivo chloroplast fluorescence measurements indicated that lower plants (bryophytes especially) showed slower changes in state transition and nonphotochemical quenching (NPQ) in response to light shifts. Therefore, the evolution of PS supercomplexes may be correlated with their acclimations to environments.

Nitrate reductase (NR) is the key enzyme for nitrogen assimilation in plant cells and also works as an important enzymatic source of nitric oxide (NO), which then regulates plant growth and resistance to biotic and abiotic stresses. However, how NR activities are finely tuned to modulate these biological processes remain largely unknown. Here we present a SWISSPROT 3D analysis of different NR from plant sources indicating the possible sites of S-nitrosylation, and show some evidence of immunoblottings to S-nitrosated (SNO-) proteins. We also found that S-nitrosylation status of NR is negatively correlated with the enzyme activity. The production of NO via NR in vitro represents only 1% of its nitrate reduction activity, possibly due to NO generated through NR reaction may deactivate the enzyme by this S-nitrosylation-mediated negative-feedback regulation. NR-mediated NO generation also plays a key role in protecting plants from abiotic stresses through activating antioxidant enzymes and increasing antioxidants. Putative connections between NR S-nitrosylation and NO biosynthesis under pathogen attacks and abiotic stresses are discussed in this Perspective.

Plants use shade avoidance strategy to escape the canopy shade when grown under natural conditions. Previous studies showed that the Arabidopsis receptor-like kinase ERECTA (ER) is involved in shade avoidance syndrome. However, the mechanisms of ER in modulating SAR by promoting hypocotyl elongation are unknown yet. Here, we report that ER regulated hypocotyl elongation in shade avoidance requires auxin and gibberellins (GAs). The T-DNA insertional ER mutant er-3 shows a less hypocotyl length than that in Col-0 wild type. Promoter::GUS staining analysis shows that ER and its paralogous genes ERECTA-LIKE1 (ERL1) and ERECTA-LIKE2 (ERL2) are differentially expressed in the seedlings, of which only ER is most obviously upregulated in the hypocotyl by shade treatment. Exogenous feeding assay by using media-application with vertical-grown of Arabidopsis seedlings showed that the hypocotyl length of er-3 is partially promoted by indol-3-acetic acid (IAA), while it is relatively insensitive of er-3 to various concentrations of IAA than that of Col-0. Hypocotyl elongation of er-3 is promoted similar to that of Col-0 by high temperature in the white light condition, but the elongation was not significantly affected by the treatment of the auxin transport inhibitor 1-N-naphthylphthalamic acid (NPA). Exogenous GA3 increased the hypocotyl elongation of both er-3 and the wild type in the shade condition, and the GA3 biosynthesis inhibitor paclobutrazol (PAC) severely inhibits the hypocotyl elongation of Col-0 and er-3. Further analysis showed that auxin biosynthesis inhibitors yucasin and L-kynurenine remarkably inhibited the hypocotyl elongation of er-3 while yucasin shows a more severe inhibition to er-3 than Col-0. Relative expression of genes regulating auxin homeostasis and signaling, and GA homeostasis is less in er-3 than that in Col-0. Furthermore, genetic evidences show that ER regulated hypocotyl elongation is dependent of PHYTOCHROME B (PHYB). Overall, we propose that ER regulated shade avoidance by promoting hypocotyl elongation is PHYB-dependent and requires auxin and GAs.

Melatonin (N-acetyl-5-methoxytryptamine) plays important role in multiple plant developmental processes and stress responses. We investigated the possible mediatory role of melatonin in growth, photosynthesis, and the response to cold stress in rice by using three different experiments: soaking seed; immersing roots, and spraying to leaves with 0, 20, or 100 μM melatonin. After 6 days of cold stress, the growth of rice seedlings was significantly inhibited, but this inhibition was alleviated by exogenous melatonin. Furthermore, exogenous melatonin pretreatment alleviated the accumulation of reactive oxygen species, malondialdehyde and cell death induced by cold stress. Melatonin pretreatment also relieved the stress-induced inhibitions to photosynthesis and photosystem II activities. Further investigations showed that, antioxidant enzyme activities and non-enzymatic antioxidant levels were increased by melatonin pretreatments. The treatment methods of seed soaking and root immersion were more effective in improving cold stress resistance than the spraying method. The results also indicated the dose-dependent response of melatonin on rice physiological, biochemical, and photosynthetic parameters.

The relationship between Mg-protoporphyrin IX (Mg-Proto IX) signals and plant’s tolerance to cold stress is investigated. Arabidopsis seedlings grown for 3 weeks were pretreated with 2 mM glutamate and 2 mM MgCl2 for 48 h at room temperature to induce Mg-Proto IX accumulation. Then cold stress was performed at 4 °C for additional 72 h. Glutamate + MgCl2 pre-treatments alleviated the subsequent cold stress significantly by rising the leaf temperature through inducing Mg-Proto IX signals. The protective role of glutamate + MgCl2 treatment was greatly compromised in the mutants of Mg-Proto IX synthesis, Mg-Proto IX signaling and cyanide-resistant respiration. And the enhancement of cold-responsive gene expression was greatly compromised in the mutants of Mg-Proto IX synthesis, Mg-Proto IX signaling and ABA signaling, but not in the mutant of cyanide-resistant respiration. Cold stress promoted cyanide-resistant respiration and leaf total respiration exponentially, which could be further induced by the glutamate + MgCl2 treatment. Mg-Proto IX signals also activate antioxidant enzymes and increase non-enzymatic antioxidants (glutathione but not ascorbic acid) to maintain redox equilibrium during the cold stress.

Photoperiod and nutrient nitrogen (N) supply influence the growth, development, and productivity of crops. This study examined the physiological, biochemical, and morphoanatomical traits of NA5 and NA9, two barley cultivars with contrasting photoperiod lengths, under the combined treatment of photoperiod regime and N supply. Under long photoperiod, high N supply decreased net photosynthesis; decreased chlorophyll a and chlorophyll a/b; decreased ascorbate peroxidase (APX), catalase (CAT), and superoxide dismutase (SOD) activities; decreased ascorbate, glutathione, soluble protein, and soluble sugar; destroyed mesophyll cell integrity; and increased O2•-, malondialdehyde, and proline in both NA5 and NA9. Under short photoperiod, high N content increased net photosynthesis; increased chlorophyll a and chlorophyll a/b; increased APX, CAT, and SOD activities; and increased antioxidants, soluble protein, and soluble sugar in NA9 but decreased the same parameters in NA5. These results indicated that N supply strongly affected photosynthetic capacity and the balance of reactive oxygen species in response to short and long photoperiod. High N supply enhanced the sensitivity of long-day barley to photoperiod change by inhibiting photosynthesis and decreasing antioxidant defense ability. High N mitigated the undesirable effects of shortened photoperiod in short-day barley. Therefore, the data from this study revealed that N status affects adaptation to photoperiod changes by maintaining redox homeostasis and photosynthetic capacity. [ABSTRACT FROM AUTHOR]

Wheat stripe rust (Puccinia striiformis f. sp. tritici, Pst), is one of the most serious diseases of wheat (Triticum aestivum L.) worldwide. To gain a better understanding of the protective mechanism against stripe rust at the adult plant stage, the differences in photosystem II and antioxidant enzymatic systems between susceptible and resistant wheat in response to stripe rust disease (P. striiformis) were investigated. We found that chlorophyll fluorescence and the activities of the antioxidant enzymes were higher in resistant wheat than in susceptible wheat after stripe rust infection. Compared with the susceptible wheat, the resistant wheat accumulated a higher level of D1 protein and a lower level of reactive oxygen species after infection. Furthermore, our results demonstrate that D1 and light-harvesting complex II (LHCII) phosphorylation are involved in the resistance to stripe rust in wheat. The CP29 protein was phosphorylated under stripe rust infection, like its phosphorylation in other monocots under environmental stresses. More extensive damages occur on the thylakoid membranes in the susceptible wheat compared with the resistant wheat. The findings provide evidence that thylakoid protein phosphorylation and antioxidant enzyme systems play important roles in plant responses and defense to biotic stress. [ABSTRACT FROM AUTHOR]