RGA1 alleviates low-light-repressed pollen tube elongation by improving the metabolism and allocation of sugars and energy.

Low-light stress compromises photosynthetic and energy efficiency and leads to spikelet sterility; however, the effect of low-light stress on pollen tube elongation in the pistil remains poorly understood. The gene RGA1, which encodes a Gα-subunit of the heterotrimeric G-protein, enhanced low-light tolerance at anthesis by preventing the cessation of pollen tube elongation in the pistil of rice plants. In this process, marked increases in the activities of acid invertase (INV), sucrose synthase (SUS) and mitochondrial respiratory electron transport chain complexes, as well as the relative expression levels of SUTs (sucrose transporter), SWEETs (sugars will eventually be exported transporters), SUSs, INVs, CINs (cell-wall INV 1), SnRK1A (sucrose-nonfermenting 1-related kinase 1) and SnRK1B, were observed in OE-1 plants. Accordingly, notable increases in contents of ATP and ATPase were presented in OE-1 plants under low-light conditions, while they were decreased in d1 plants. Importantly, INV and ATPase activators (sucrose and Na2 SO3 , respectively) increased spikelet fertility by improving the energy status in the pistil under low-light conditions, and the ATPase inhibitor Na2 VO4 induced spikelet sterility and decreased ATPase activity. These results suggest that RGA1 could alleviate the low-light stress-induced impairment of pollen tube elongation to increase spikelet fertility by promoting sucrose unloading in the pistil and improving the metabolism and allocation of energy.

How Rice Plants Response to Abiotic Stresses.

With the combustion of fossil fuels, unequal and unsustainable energy and land use, and irrational human activities, greenhouse gas emissions remain high, which leads to global warming [...].

Comparative Transcriptome Analysis Reveals OsBGs and OsGSLs Influence Sugar Transport through Callose Metabolism under Heat Stress in Rice.

Heat or high temperature stress have caused huge damage to many crops and have become the largest threat in terms of the future. Although a huge amount of research has been conducted to explore the mechanisms of heat tolerance and many achievements were accomplished, the mechanism by which how heat stress (HS) influences the yield is still unclear. In this study, RNA-seq analysis indicated that nine 1,3-ß-glucanases (BGs) belonging to the carbohydrate metabolic pathway were expressed differently during heat treatment. Therefore, we identified the BGs and glucan-synthase-likes (GSLs) in three rice ecotypes and processed the analyses of gene gain and loss, phylogenetic relationship, duplication, and syntenic relationship. We found the possibility of an environmental adaption based on BGs and GSLs during evolution. Submicrostructure and dry matter distribution analysis confirmed that HS might block the endoplasmic sugar transport pathway by increasing callose synthesis, which may lead to decreased yield and quality in rice production. This study provides a new clue regarding rice yield and quality under HS and provides guidance to rice cultivation and heat tolerance breeding.

Abscisic Acid Improves Rice Thermo-Tolerance by Affecting Trehalose Metabolism.

Heat stress that occurs during the flowering stage severely decreases the rice (Oryza sativa L.) seed-setting rate. This damage can be reversed by abscisic acid (ABA), through effects on reactive oxygen species, carbohydrate metabolism, and heat shock proteins, but the exact role of trehalose and ATP in this process remains unclear. Two rice genotypes, namely, Zhefu802 (heat-resistant plant, a recurrent parent) and its near-isogenic line (faded green leaf, Fgl, heat-sensitive plant), were subjected to 38 °C heat stress after being sprayed with ABA or its biosynthetic inhibitor, fluridone (Flu), at the flowering stage. The results showed that exogenous ABA significantly increased the seed-setting rate of rice under heat stress, by 14.31 and 22.40% in Zhefu802 and Fgl, respectively, when compared with the H2O treatment. Similarly, exogenous ABA increased trehalose content, key enzyme activities of trehalose metabolism, ATP content, and F1Fo-ATPase activity. Importantly, the opposite results were observed in plants treated with Flu. Therefore, ABA may improve rice thermo-tolerance by affecting trehalose metabolism and ATP consumption.

The Evolution and Functional Roles of miR408 and Its Targets in Plants.

MicroRNA408 (miR408) is an ancient and highly conserved miRNA, which is involved in the regulation of plant growth, development and stress response. However, previous research results on the evolution and functional roles of miR408 and its targets are relatively scattered, and there is a lack of a systematic comparison and comprehensive summary of the detailed evolutionary pathways and regulatory mechanisms of miR408 and its targets in plants. Here, we analyzed the evolutionary pathway of miR408 in plants, and summarized the functions of miR408 and its targets in regulating plant growth and development and plant responses to various abiotic and biotic stresses. The evolutionary analysis shows that miR408 is an ancient and highly conserved microRNA, which is widely distributed in different plants. miR408 regulates the growth and development of different plants by down-regulating its targets, encoding blue copper (Cu) proteins, and by transporting Cu to plastocyanin (PC), which affects photosynthesis and ultimately promotes grain yield. In addition, miR408 improves tolerance to stress by down-regulating target genes and enhancing cellular antioxidants, thereby increasing the antioxidant capacity of plants. This review expands and promotes an in-depth understanding of the evolutionary and regulatory roles of miR408 and its targets in plants.

Acid invertase confers heat tolerance in rice plants by maintaining energy homoeostasis of spikelets.

Heat stress impairs both pollen germination and pollen tube elongation, resulting in pollination failure caused by energy imbalance. Invertase plays a critical role in the maintenance of energy homoeostasis; however, few studies investigated this during heat stress. Two rice cultivars with different heat tolerance, namely, TLY83 (heat tolerant) and LLY722 (heat susceptible), were subjected to heat stress. At anthesis, heat stress significantly decreased spikelet fertility, accompanied by notable reductions in pollen germination on stigma and pollen tube elongation in ovule, especially in LLY722. Acid invertase (INV), rather than sucrose synthase, contributed to sucrose metabolism, which explains the different tolerances of both cultivars. Under heat stress, larger enhancements in NAD(H), ATP, and antioxidant capacity were found in TLY83 compared with LLY722, whereas a sharp reduction in poly(ADP-ribose) polymerase (PARP) activity was found in the former compared with the latter. Importantly, exogenous INV, 3-aminobenzamide (a PARP inhibitor), sucrose, glucose, and fructose significantly increased spikelet fertility under heat stress, where INV activity was enhanced and PARP activity was inhibited. Therefore, INV can balance the energy production and consumption to provide sufficient energy for pollen germination and pollen tube growth under heat stress.

Non-Photochemical Quenching Involved in the Regulation of Photosynthesis of Rice Leaves under High Nitrogen Conditions.

Excess and deficient nitrogen (N) inhibit photosynthesis in the leaves of rice plants, but the underlying mechanism is still unclear. N can improve the chlorophyll content and thus affect photon absorption, but the photosynthetic rate does not increase accordingly. To investigate this mechanism, three concentrations of N treatments were applied to two rice varieties, Zhefu802 and Fgl. The results indicated increased chlorophyll content of leaves with an increased N supply. Little discrepancy was detected in Rubisco enzyme activity and Non-photochemical quenching (NPQ) in the high nitrogen (HN) and moderate nitrogen (MN) treatments. The model that photoinhibition occurs in Zhefu802 due to a lack of balance of light absorption and utilization is supported by the higher malondialdehyde (MDA) content, higher H2O2 content, and photoinhibitory quenching (qI) in HN treatment compared with MN treatment. A lower proportion of N in leaf was used to synthesize chlorophyll for Fgl compared with Zhefu802, reducing the likelihood of photoinhibition under HN treatment. In conclusion, HN supply does not allow ideal photosynthetic rate and increases the likelihood of photoinhibition because it does not sustain the balance of light absorption and utilization. Apart from Rubisco enzyme activity, NPQ mainly contributes to the unbalance. These results of this study will provide reference for the effective N management of rice.

Abscisic acid synergizes with sucrose to enhance grain yield and quality of rice by improving the source-sink relationship.

BACKGROUND: Abscisic acid (ABA) and sucrose act as molecular signals in response to abiotic stress. However, how their synergy regulates the source-sink relationship has rarely been studied. This study aimed to reveal the mechanism underlying the synergy between ABA and sucrose on assimilates allocation to improve grain yield and quality of rice. The early indica rice cultivar Zhefu802 was selected and planted in an artificial climate chamber at 32/24 °C (day/night) under natural sunlight conditions. Sucrose and ABA were exogenously sprayed (either alone or in combination) onto rice plants at flowering and 10 days after flowering. RESULTS: ABA plus sucrose significantly improved both the grain yield and quality of rice, which was mainly a result of the higher proportion of dry matter accumulation and non-structural carbohydrates in panicles. These results were mainly ascribed to the large improvement in sucrose transport in the sheath-stems in response to the ABA plus sucrose treatment. In this process, ABA plus sucrose significantly enhanced the contents of starch, gibberellic acids, and zeatin ribosides as well as the activities and gene expression of enzymes involved in starch synthesis in grains. Additionally, remarkable increases in trehalose content and expression levels of trehalose-6-phosphate synthase1, trehalose-6-phosphate phosphatase7, and sucrose non-fermenting related protein kinase 1A were also found in grains treated with ABA plus sucrose. CONCLUSION: The synergy between ABA and sucrose increased grain yield and quality by improving the source-sink relationship through sucrose and trehalose metabolism in grains.

Salicylic acid reverses pollen abortion of rice caused by heat stress.

BACKGROUND: Extremely high temperatures are becoming an increasingly severe threat to crop yields. It is well documented that salicylic acid (SA) can enhance the stress tolerance of plants; however, its effect on the reproductive organs of rice plants has not been described before. To investigate the mechanism underlying the SA-mediated alleviation of the heat stress damage to rice pollen viability, a susceptible cultivar (Changyou1) was treated with SA at the pollen mother cell (PMC) meiosis stage and then subjected to heat stress of 40 °C for 10 d until 1d before flowering. RESULTS: Under control conditions, no significant difference was found in pollen viability and seed-setting rate in SA treatments. However, under heat stress conditions, SA decreased the accumulation of reactive oxygen species (ROS) in anthers to prevent tapetum programmed cell death (PCD) and degradation. The genes related to tapetum development, such as EAT1 (Eternal Tapetum 1), MIL2 (Microsporeless 2), and DTM1 (Defective Tapetum and Meiocytese 1), were found to be involved in this process. When rice plants were exogenously sprayed with SA or paclobutrazol (PAC, a SA inhibitor) + H2O2 under heat stress, a significantly higher pollen viability was found compared to plants sprayed with H2O, PAC, or SA + dimethylthiourea (DMTU, an H2O2 and OH· scavenger). Additionally, a sharp increase in H2O2 was observed in the SA or PAC+ H2O2 treatment groups compared to other treatments. CONCLUSION: We suggest that H2O2 may play an important role in mediating SA to prevent pollen abortion caused by heat stress through inhibiting the tapetum PCD.

Novel roles of hydrogen peroxide (H2O2) in regulating pectin synthesis and demethylesterification in the cell wall of rice (Oryza sativa) root tips.

Hydrogen peroxide (H2O2) has been reported to increase lignin formation, enhance cell wall rigidification, restrict cell expansion and inhibit root elongation. However, our results showed that it not only inhibited rice (Oryza sativa) root elongation, but also increased root diameter. No study has reported how and why H2O2 increases cell expansion and root diameter. Exogenous H2O2 and its scavenger 4-hydroxy-Tempo were applied to confirm the roles of H2O2. Immunofluorescence, fluorescence probe, ruthenium red staining, histological section and spectrophotometry were used to monitor changes in the degree of pectin methylesterification, pectin content, pectin methylesterase (PME) activity and H2O2 content. Exogenous H2O2 inhibited root elongation, but increased cell expansion and root diameter significantly. H2O2 not only increased the region of pectin synthesis and pectin content in root tips, but also increased PME activity and pectin demethylesterification. The scavenger 4-hydroxy-Tempo reduced root H2O2 content and recovered H2O2-induced increases in cell expansion and root diameter by inhibiting pectin synthesis, PME activity and pectin demethylesterification. H2O2 plays a novel role in the regulation of pectin synthesis, PME activity and pectin demethylesterification. H2O2 increases cell expansion and root diameter by increasing pectin content and demethylesterification.

RGA1 Negatively Regulates Thermo-tolerance by Affecting Carbohydrate Metabolism and the Energy Supply in Rice.

BACKGROUND: Signal transduction mediated by heterotrimeric G proteins, which comprise the α, ß, and γ subunits, is one of the most important signaling pathways in rice plants. RGA1, which encodes the Gα subunit of the G protein, plays an important role in the response to various types of abiotic stress, including salt, drought, and cold stress. However, the role of RGA1 in the response to heat stress remains unclear. RESULTS: The heat-resistant mutant ett1 (enhanced thermo-tolerance 1) with a new allele of the RGA1 gene was derived from an ethane methyl sulfonate-induced Zhonghua11 mutant. After 45 °C heat stress treatment for 36 h and recovery for 7 d, the survival rate of the ett1 mutants was significantly higher than that of wild-type (WT) plants. The malondialdehyde content was lower, and the maximum fluorescence quantum yield of photosystem II, peroxidase activity, and hsp expression were higher in ett1 mutants than in WT plants after 12 h of exposure to 45 °C. The RNA-sequencing results revealed that the expression of genes involved in the metabolism of carbohydrate, nicotinamide adenine dinucleotide, and energy was up-regulated in ett1 under heat stress. The carbohydrate content and the relative expression of genes involved in sucrose metabolism indicated that carbohydrate metabolism was accelerated in ett1 under heat stress. Energy parameters, including the adenosine triphosphate (ATP) content and the energy charge, were significantly higher in the ett1 mutants than in WT plants under heat stress. Importantly, exogenous glucose can alleviate the damages on rice seedling plants caused by heat stress. CONCLUSION: RGA1 negatively regulates the thermo-tolerance in rice seedling plants through affecting carbohydrate and energy metabolism.

Tungstate: is it really a specific nitrate reductase inhibitor in plant nitric oxide research?

Nitrate reductase (NR) is an enzymatic source of nitric oxide (NO) in plants, and it needs Mo for the Mo-cofactor to be activated. Because NR-deficient mutants are not always available in some species, a cheap and simple pharmacological application of tungstate, which substitutes for Mo in the Mo-cofactor as a competitive antagonist, is widely used as a NR inhibitor in plant NO research. However, evidence indicates that tungstate not only inactivates NR but also inhibits other molybdate-dependent enzymes in plants. In addition, a number of investigations have shown that tungstate also inhibits root growth, affects cortical microtubule formation, and induces programmed cell death (PCD) in plants, just like other heavy metals do. Therefore, tungstate has been shown to exert many other effects that are not connected with the inhibition of NR activity. The origin and mechanism of using tungstate as a NR inhibitor in plants is reviewed here and the progress regarding tungstate toxicity to plants and the possible problems involved in using tungstate as a NR inhibitor in plant NO research are analysed. In summary, the use of tungstate as a NR inhibitor in plant NO research must be treated with caution, keeping in mind that it is not completely specific. It is necessary to search for more NR-deficient mutants and new, specific NR inhibitors. A combination of pharmacological and biochemical analysis with a genetic approach will be necessary in order to investigate the roles of NO in plants.

Drought-induced proline accumulation is uninvolved with increased nitric oxide, which alleviates drought stress by decreasing transpiration in rice.

Accumulation of proline is trusted to be an adaptive response of plants against drought stress, and exogenous application of nitric oxide (NO) enhances proline accumulation in Cu-treated algae. In order to investigate whether NO works as a necessary signaling molecule in drought-induced proline accumulation in rice leaves, effects of drought stress on endogenous NO content and proline accumulation were studied in rice leaves, using sodium nitroprusside (SNP, a NO donor) and 2-(4-carboxyphenyl)-4,4,5,5-tetramethylimidazoline-1-oxyl-3-oxide (cPTIO, a NO scavenger). The results showed that drought treatment increased both endogenous NO and proline contents in rice leaves, while foliar spray of various concentrations of SNP failed to induce proline accumulation in the leaves of well-watered rice and foliar spray of cPTIO failed to inhibit proline accumulation in the leaves of drought-stressed rice. These results indicate that increase of endogenous NO is dispensable for proline accumulation in the leaves of rice under drought stress. Further studies indicate that exogenous application of NO alleviates drought-induced water loss and ion leakage by decreasing transpiration rate of rice leaves.

Effects and oxygen-regulated mechanisms of water management on cadmium (Cd) accumulation in rice (Oryza sativa).

Irrigation has been considered an effective approach for decreasing cadmium (Cd) uptake and accumulation in rice (Oryza sativa), but increasing evidence shows that the effects of different water management strategies on Cd accumulation in rice are contradictory in different studies, and the detailed regulatory mechanisms remain unconfirmed. Most previous studies have shown that irrigation regulates Cd accumulation in rice mainly by affecting Cd bioavailability, pH and redox potential (Eh) in soil, and few reports have focused on the function of oxygen (O2) in regulating the physiological mechanisms of rice on Cd tolerance or accumulation. Here, we concluded that irrigation affects Cd bioavailability, pH and Eh in soil mainly by regulating O2 content. In addition, recent studies have also shown that irrigation-regulated O2 also affects Cd accumulation in rice by affecting iron plaque (IP), the radial oxygen loss (ROL) barrier, the cell wall and mass flow in rice roots. All these results indicate that O2 is the key factor in irrigation-regulated Cd accumulation in rice, and dramatic result variations from different irrigation experiments are due to the different rhizosphere O2 conditions. This review will help clarify the effects and regulatory mechanisms of irrigation on Cd accumulation in rice and reveal the roles of O2 in this process.

Increased ATPase activity promotes heat-resistance, high-yield, and high-quality traits in rice by improving energy status.

Heat stress during the reproductive stage results in major losses in yield and quality, which might be mainly caused by an energy imbalance. However, how energy status affected heat response, yield and quality remains unclear. No relationships were observed among the heat resistance, yield, and quality of the forty-nine early rice cultivars under normal temperature conditions. However, two cultivars, Zhuliangyou30 (ZLY30) and Luliangyou35 (LLY35), differing in heat resistance, yield, and quality were detected. The yield was higher and the chalkiness degree was lower in ZLY30 than in LLY35. Decreases in yields and increases in the chalkiness degree with temperatures were more pronounced in LLY35 than in ZLY30. The accumulation and allocation (ratio of the panicle to the whole plant) of dry matter weight and non-structural carbohydrates were higher in ZLY30 than in LLY35 across all sowing times and temperatures. The accumulation and allocation of dry matter weight and non-structural carbohydrates in panicles were higher in ZLY30 than in LLY35. Similar patterns were observed in the relative expression levels of sucrose unloading related genes SUT1 and SUT2 in grains. The ATP content was higher in the grains of LLY35 than in ZLY30, whereas the ATPase activity, which determined the energy status, was significantly lower in the former than in the latter. Thus, increased ATPase activity, which improved the energy status of rice, was the factor mediating the balance among heat-resistance, high-yield, and high-quality traits in rice.

Oligomeric Proanthocyanidins Confer Cold Tolerance in Rice through Maintaining Energy Homeostasis.

Oligomeric proanthocyanidins (OPCs) are abundant polyphenols found in foods and botanicals that benefit human health, but our understanding of the functions of OPCs in rice plants is limited, particularly under cold stress. Two rice genotypes, named Zhongzao39 (ZZ39) and its recombinant inbred line RIL82, were subjected to cold stress. More damage was caused to RIL82 by cold stress than to ZZ39 plants. Transcriptome analysis suggested that OPCs were involved in regulating cold tolerance in the two genotypes. A greater increase in OPCs content was detected in ZZ39 than in RIL82 plants under cold stress compared to their respective controls. Exogenous OPCs alleviated cold damage of rice plants by increasing antioxidant capacity. ATPase activity was higher and poly (ADP-ribose) polymerase (PARP) activity was lower under cold stress in ZZ39 than in RIL82 plants. Importantly, improvements in cold tolerance were observed in plants treated with the OPCs and 3-aminobenzamide (PARP inhibitor, 3ab) combination compared to the seedling plants treated with H2O, OPCs, or 3ab alone. Therefore, OPCs increased ATPase activity and inhibited PARP activity to provide sufficient energy for rice seedling plants to develop antioxidant capacity against cold stress.

Respiration, Rather Than Photosynthesis, Determines Rice Yield Loss Under Moderate High-Temperature Conditions.

Photosynthesis is an important biophysical and biochemical reaction that provides food and oxygen to maintain aerobic life on earth. Recently, increasing photosynthesis has been revisited as an approach for reducing rice yield losses caused by high temperatures. We found that moderate high temperature causes less damage to photosynthesis but significantly increases respiration. In this case, the energy production efficiency is enhanced, but most of this energy is allocated to maintenance respiration, resulting in an overall decrease in the energy utilization efficiency. In this perspective, respiration, rather than photosynthesis, may be the primary contributor to yield losses in a high-temperature climate. Indeed, the dry matter weight and yield could be enhanced if the energy was mainly allocated to the growth respiration. Therefore, we proposed that engineering smart rice cultivars with a highly efficient system of energy production, allocation, and utilization could effectively solve the world food crisis under high-temperature conditions.

Roles of nitric oxide in alleviating heavy metal toxicity in plants.

Nitric oxide (NO) is involved in the regulation of multiple plant responses to a variety of abiotic and biotic stresses. Recently, an increasing number of articles have reported the effects of exogenous NO on alleviating heavy metal toxicity in plants. However, compared with the current understanding of the relationships between NO and other abiotic stresses, knowledge of the molecular and physiological mechanisms of NO in alleviating heavy metal toxicity is quite limited, and some results contradict one another. Therefore, to help clarify the roles of NO in heavy metal tolerance, it is valuable to review and discuss the recent advances on this research topic. In this mini-review, the latest advances in understanding the effects of heavy metals on endogenous NO content and the mechanisms and signaling pathways of exogenous NO in alleviating heavy metal toxicity in plants are summarized and discussed. A basic scheme for the roles of NO in alleviating heavy metal toxicity is also proposed.

Regulatory mechanisms of nitrogen (N) on cadmium (Cd) uptake and accumulation in plants: A review.

Cadmium (Cd) is a heavy metal that is toxic to plants and animals. Nitrogen (N), the most significant macro-nutrient and a common input for crop production, is often excessively applied than plants' demands by farmers to obtain more economic benefits. Understanding the regulatory mechanisms of N that control Cd uptake, translocation, and accumulation may enable the development of solutions regarding Cd pollution in the trophic chain, a major and global threat to agricultural sustainability and human health. In this review, we clarified that an increased amount of N, regardless of its form, enhances Cd uptake, translocation, and accumulation in plants, and nitrate promotes Cd uptake more than any other N form. We also described that N fertilizer alters the Cd exchange capacity and the bio-available Cd content in soil; regulates nitric oxide induced divalent cation gene expression of Nramp1, HMA2, and IRT1; and changes cell wall isolation, chelation capacity, and oxidative resistance to regulate Cd accumulation in plants. By revealing the integrated interaction effects between Cd accumulation and N fertiliser use, we propose new challenges to investigate the functions and mechanisms of N in Cd-contaminated croplands and develop suitable N-fertilisation protocols to practically reduce food health risks in agricultural food production.

Abscisic Acid Negatively Modulates Heat Tolerance in Rolled Leaf Rice by Increasing Leaf Temperature and Regulating Energy Homeostasis.

BACKGROUND: Abscisic acid (ABA) acts as a signaling hormone in plants against abiotic stress, but its function in energy homeostasis under heat stress is unclear. RESULTS: Two rice genotypes, Nipponbare (wild-type, WT) with flat leaves and its mutant high temperature susceptibility (hts) plant with semi-rolled leaves, were subjected to heat stress. We found significantly higher tissue temperature, respiration rate, and ABA and H2O2 contents in leaves as well as a lower transpiration rate and stomatal conductance in hts than WT plants. Additionally, increased expression of HSP71.1 and HSP24.1 as well as greater increases in carbohydrate content, ATP, NAD (H), and dry matter weight, were detected in WT than hts plants under heat stress. More importantly, exogenous ABA significantly decreased heat tolerance of hts plants, but clearly enhanced heat resistance of WT plants. The increases in carbohydrates, ATP, NAD (H), and heat shock proteins in WT plants were enhanced by ABA under heat stress, whereas these increases were reduced in hts plants. CONCLUSION: It was concluded that ABA is a negative regulator of heat tolerance in hts plants with semi-rolled leaves by modulating energy homeostasis.