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Saline–alkaline (SA) stress is characterized by high salinity and high alkalinity (high pH), which severely inhibit plant growth and cause huge losses in crop yields worldwide. Here we show that a moderate elevation of endogenous abscisic acid (ABA) levels by RNAi-mediated suppression of OsABA8ox1 (OsABA8ox1-kd), a key ABA catabolic gene, significantly increased tolerance to SA stress in rice plants. We produced OsABA8ox1-kd lines in two different japonica cultivars, Dongdao 4 and Nipponbare. Compared with non-transgenic control plants (WT), the OsABA8ox1-kd seedlings accumulated 25.9%–55.7% higher levels of endogenous ABA and exhibited reduced plasmalemma injury, ROS accumulation and Na+/K+ ratio, and higher survival rates, under hydroponic alkaline conditions simulated by 10, 15, and 20 mmol L−1 of Na2CO3. In pot trials using SA field soils of different alkali levels (pH 7.59, 8.86, and 9.29), OsABA8ox1–kd plants showed markedly higher seedling survival rates and more vigorous plant growth, resulting in significantly higher yield components including panicle number (85.7%–128.6%), spikelets per panicle (36.9%–61.9%), branches (153.9%–236.7%), 1000–kernel weight (20.0%–28.6%), and percentage of filled spikelets (96.6%–1340.8%) at harvest time. Under severe SA soil conditions (pH = 9.29, EC = 834.4 μS cm−1), OsABA8ox1-kd lines showed an 194.5%–1090.8% increase in grain yield per plant relative to WT plants. These results suggest that suppression of OsABA8ox1 to increase endogenous ABA levels provides a new molecular approach for improving rice yield in SA paddies.

Abscisic acid (ABA) has been shown to prime rice seedlings for enhanced tolerance to alkaline stress under both greenhouse and field conditions. This study aimed to understand the mechanism of the ABA priming effect. Rice seedlings were grown hydroponically and pretreated with ABA (10 μM) for 24 h, and then subjected to alkaline stress simulated by 15 mM Na2CO3 solution (pH 10.87). Paraquat (2.5–25 μM) was used as an intracellular generator of reactive oxygen species (ROS). Compared to the control treatment, pretreatment with ABA significantly mitigated root damage and improved the survival rate of rice seedlings under alkaline conditions. Furthermore, ABA pretreatment increased the antioxidant enzyme activities of superoxide dismutase, catalase, peroxidase and ascorbate peroxidase, and reduced the alkalinity- and paraquat-induced ROS accumulation (O2·- and H2O2) and seedling injury and mortality. In addition, the expression of the ABA-responsive genes SalT and OsWsi18, as well as the stress tolerance-related genes OsJRL, OsPEX11, OsNAC9, OsAKT1 and OsHKT1, was superinduced by ABA pretreatment under alkaline conditions. ABA priming enhances tolerance to alkaline stress by upregulating the antioxidant defense system and stress tolerance-related genes in the roots of rice seedlings.

Soil sodicity is a major ecological problem in the western Songnen Plain of Northeast China and rice cultivation is the main approach used to mitigate saline-sodic soils. However, rice cultivation alone may not be the most effective practice. This study aimed to investigate the combined effects of annual tillage and rice cultivation with either phosphogypsum or farmyard manure on soil salinity, mineral status, and concentration of heavy metals in saline-sodic paddy fields. Treatments were: 1) untreated (no amendments), untilled, and uncultivated (no rice) saline-sodic native grasslands (UG); 2) untreated, tilled, rice-cultivated paddy fields (PFU); 3) tilled, rice-cultivated, amended paddy fields with phosphogypsum (PFPG); and 4) tilled, rice-cultivated, amended paddy fields with farmyard manure (PFFM). The effectiveness of these treatments on soil improvement was evaluated after a 10-year field experiment. Compared to the UG control, the 0–20 cm topsoil layer of PFU, PFPG, and PFFM had respective decreases in Na+ concentrations of 42.9%, 61.5%, and 60.9%; in CO32- + HCO3- concentrations of 18.9%, 63.2%, and 57.9%; in Cl- concentration of 64.6%, 75.7%, and 79.9%; in pH units of 0.57, 1.05, and 1.30; in soil electrical conductivity (EC1:5) of 18.3%, 49.1%, and 48.3%; and in exchange sodium percentage (ESP) of 47.2%, 66.9%, and 72.5%. Also, the 0–20 cm topsoil layer of PFPG and PFFM had its concentrations of soil organic matter (SOM), available nitrogen (AN), and available phosphorus (AP) significantly (P

Soil salinity and sodicity can not only directly restrain crop growth by osmotic and specific ion stresses, it also may reduce grain yield indirectly by impacting plant absorption of essential nutrients. Ensuring adequate nitrogen is an important management aspect of rice production in saline-sodic soils. The objective of this study was to investigate the interaction of soil pH, salinity and nitrogen application on rice yield and nutrient absorption. We conducted a rice experiment in containers in a greenhouse. The soils were first leached with 9 target salt solutions of pH 7, 8 and 9 and electrical conductivity (EC) of 2, 6 and 10 dS m −1 . Nitrogen application rates were 100, 200 and 300 kg N ha −1 . Rice grain yield and shoot weight significantly decreased with increasing soil pH and increasing soil EC, and significantly increased with increasing nitrogen application ( P P 2 = 0.565, P

Soil aggregation, an ecosystem function correlated with the concentration of glomalin-related soil protein (GRSP), is highly disturbed in saline soil. However, few studies have focused on differences in amount, composition, and ion binding capacity of GRSP in typical sodic-saline soils. In this study, a field study was performed in Songnen Plain. Combined indicators of soil salinity (Q value) were significant negatively correlated with GRSP concentration by Principal Component Analysis. Multiple linear regression models showed that soil salinity might account for 46%, 25% and 44% variation in total GRSP (T-GRSP), easily-extractable GRSP (EE-GRSP) and difficultly-extractable GRSP (DE-GRSP), respectively. Soil bulk density had most important impact on GRSP concentration, followed by the pH, soil EC had the weak influence. Comparative analysis was carried out between low-salinity and high-salinity soil. Purified T-GRSP of high-saline soil contained higher N content (13.13%), lower C content (43.41%) and lower functional groups relative content (e.g. CO and SiOSi). Purified T-GRSP of high-salinity soil had a greater binding capacity with calcium and phosphorus, the binding capacity could compensate the GRSP loss about 29.8% and 14.1%, respectively. Our findings suggested that sodic salinization of the soil led to a decrease in GRSP concentration and a change in the component percentages. This change in composition might be related to adaptation of fungi-plant systems to varied environments. The calcium and phosphorus binding capacity had a positive dependent of soil salinization, which was possible to develop ecological management or recovery technology in the future.

Saline-alkaline (SA) stress is one of the major constraints to crop production worldwide. It is characterized by high salinity and high alkalinity (high pH), with the latter being the primary inhibiting factor to rice growth. We previously reported that pretreatment with abscisic acid (ABA) primed rice seedlings for enhanced tolerance to alkaline stress. In the present study, we conducted a 3-year field experiment to further assess the ABA-priming effect on the stress tolerance of rice plants in two paddy fields with different SA levels (field-A: non-reclaimed field, pH = 10.13 +/- 0.009, EC1:5 = 0.50 +/- 0.004; field-B: reclaimed field, pH = 9.74 +/- 1 0.005, EC1:5 = 0.48 +/- 0.001). Rice seedlings were treated with ABA at concentrations of 0 (control), 1, and 10 mu M by root-drenching for 24 h prior to transplanting to the two paddy fields with different SA levels. Compared to the control treatment, ABA application substantially reduced the occurrence of leaf withering by 22%-50% and seedling death by 11%-17%. Further, ABA application also significantly improved subsequent plant growth and the final grain yield by 8%-55%; a more profound effect was observed in the harder SA field. Expressions of the ABA-responsive genes SalT and OsWsi18 were highly induced by ABA treatment, and the induced levels were sustained until at least 8 d after transplanting. Collectively, these results suggest a potent priming effect of ABA on the SA stress tolerance of rice plants, providing an effective and practical approach to improving rice production in SA paddy fields. (C) 2016 Elsevier B.V. All rights reserved.

Soil alkalization triggers ion toxicity and osmotic and alkaline (high pH) stresses in plants, damaging their growth and productivity. Therefore, we investigated whether priming with abscisic acid (ABA) increases the tolerance of alfalfa seedlings to alkaline stress, and then examined the underlying molecular mechanisms. Alfalfa seedlings were pretreated with ABA (10 μM) for 16 h and then subjected to alkaline stress using a 15 mM Na2CO3 solution (pH 10.87). Compared with the control, ABA pretreatment significantly alleviated leaf damage and improved the fresh weight, water content, and survival rate of alfalfa seedlings under alkaline conditions. Abscisic acid pretreatment reduced accumulation of reactive oxygen species (ROS), increased activities of the antioxidant enzymes superoxide dismutase (SOD) and peroxidase (POD), maintained higher ratios of K+/Na+, Ca2+/Na+, and Mg2+/Na+, and increased accumulation of proline. In addition, ABA upregulated the expression of genes involved in proline biosynthesis (P5CS) and the sequestration of Na+ in vacuoles (NHX1 and AVP) under alkaline conditions. Abscisic acid priming increased tolerance to alkaline stress by maintaining homeostasis of ROS and metal ions and upregulating osmoprotection and the expression of stress tolerance-related genes.

Seed priming is a widely used technique in crops to obtain uniform germination and high-quality seedlings. In this study, we found a long-term effect of seed priming with gibberellic acid-3 (GA3) on plant growth and production in Leymus chinensis. Seeds were germinated on agar plates containing 0–200 μM GA3, and the germinated seedlings were transplanted to clay planting pots and grown for about one year. The clonal tillers grown from the mother plants were transplanted to field conditions in the second year. Results showed that GA3 treatment significantly increased seed germination rate by 14–27%. GA3 treatment also promoted subsequent plant growth and biomass production, as shown by a significant increase in plant height, tiller number, and fresh and dry weight in both pot (2016) and field (2017) conditions. It is particularly noteworthy that the growth-promoting effect of a single seed treatment with GA3 lasted for at least two years. In particular, GA3 treatment at 50 μM increased aboveground fresh and dry weight by 168.2% and 108.9% in pot-grown conditions, and 64.5% and 126.2% in field-grown conditions, respectively. These results imply a transgenerational transmission mechanism for the GA-priming effect on clonal offspring growth and biomass production in L. chinensis.

Background and aims Alkaline soils, characterized by high pH, are representative of degraded regions throughout the world. Studying germination in relation to alkalinity can contribute to understanding how species cope with such conditions. Although the effects of pH have been widely studied, it is unknown whether germination response to pH gradients created with buffer solutions is representative of the conditions experienced inalkalinesoils.Our aimswereto(1) determineif high pH gives an accurate assessment of the effects of alkaline soils on germination, and (2) identify the inhibitory factors for germination in alkaline soils. Methods Using Leymus chinensis seeds, germination was tested over a gradient of pH solutions prepared using Tris (50 mM and 100 mM) and H2O buffers and eight germination media prepared from non-alkaline and alkaline soils with different pH and electrical conductivities (EC). Additionally, solutions of 10–100 mM NaCl, Na2SO4 ,N a2CO3 and NaHCO3 were used to determine the main ions inhibiting seed germination. Results H2O-buffered pH had no effect on seed germination, and seed germination was much lower at all pH levels in 50 mM Tris–HCl solutions (pH 7.0–10.35) than in the H2O control (pH 7.05). No seeds germinated in 100 mM Tris–HCl buffers irrespective of the pH. In alkalinegermination media (pH 10.04–10.61), highgermination was obtained only at low EC. The rank order of the inhibitory effect of salts was Na2CO3>NaHCO3> NaCl>Na2SO4. Conclusions Buffer solutions used to simulate alkaline environments did not provide a reliable indicator of the effects of alkalinesoils onseedgermination.High pH of alkaline soil had no negative effects, and results suggest that salt composition and concentration, especially CO3 2� and HCO3 � , are key inhibitors.

Saline-alkaline stress is characterized by high salinity and high alkalinity (high pH); alkaline stress has been shown to be the primary factor inhibiting rice seedling growth. In this study, we investigated the potential priming effect of abscisic acid (ABA) on tolerance of rice seedlings to alkaline stress simulated by Na2CO3. Seedlings were pretreated with ABA at concentrations of 0 (control), 10, and 50 μM by root-drench for 24 h and then transferred to a Na2CO3 solution that did not contain ABA. Compared to control treatment, pretreatment with ABA substantially improved the survival rate of rice seedlings and increased biomass accumulation after 7 days under the alkaline condition. ABA application at 10 μM also alleviated the inhibitory effects of alkaline stress on the total root length and root surface area. Physiologically, ABA increased relative water content (RWC) and decreased cell membrane injury degree (MI) and Na(+)/K(+) ratios. In contrast, fluridone (an ABA biosynthesis inhibitor) decreased the RWC and increased MI in shoots under the alkaline conditions. These data suggest that ABA has a potent priming effect on the adaptive response to alkaline stress in rice and may be useful for improving rice growth in saline-alkaline paddy fields.

In response to pathogen attack, plants prioritize defense reactions generally at the expense of plant growth. In this work, we report that changes in phytohormone signaling pathways are associated with the stunted plant growth caused by blast disease in rice seedlings. Infection of rice seedlings with blast fungus Magnaporthe oryzae (race 007.0) at the four-leaf stage (three true leaves) resulted in considerable inhibition of the growth of the upper uninfected distal leaves; the length of leaf blade and leaf sheath of the sixth and seventh leaf was reduced by 27 and 82%, and 88 and 72%, respectively, compared to that in the uninoculated plant control. Interestingly, cutting off the blast-infected fourth leaf blade within 2 days post inoculation (dpi) significantly rescued the inhibition of leaf growth, implying that an inhibitory substance(s) and/or signal was generated in the blast-infected leaves (fourth leaf) and transmitted to the upper distal leaves (sixth and seventh) during the 2-dpi period that induced growth inhibition. Expression analysis of marker genes for phytohormone pathways revealed acute activation of the jasmonate (JA) and abscisic acid (ABA) signaling pathways, and repression of auxin, gibberellic acid (GA) and salicylic acid (SA) signaling pathways, in the sixth leaf. The genes related to cell wall expansion were also significantly downregulated. In the blast-infected fourth leaf, JA pathway was activated within 2 dpi, followed by activation of ABA pathway 3 dpi. Further, leaf inhibition caused by blast infection was partially rescued in the rice mutant line coleoptile photomorphogenesis 2 (cpm2), which is defective in the gene encoding allene oxide cyclase (OsAOC). These results indicate that the JA signaling pathway is at least partly involved in the growth inhibition processes. Collectively, our data suggest that, upon pathogen attack, rice seedlings prioritize defense reactions against the infecting pathogen by temporarily ceasing plant growth through the systemic control of phytohormone pathways.

Saline–alkaline (SA) stress is a complex effect of three main factors: ion toxicity from high salinity, alkalinity, and high osmotic pressure. In this study, we dissected response of rice (Oryza sativa L.) to above-mentioned SA stress factors simulated by using NaCl, Na₂CO₃, and PEG6000, respectively. Seed germination was most strongly affected by osmotic stress, followed by alkaline and salt stresses. Meanwhile, the seedling growth was most reduced under alkaline stress. The alkaline treatment significantly reduced total biomass, total root length (TRL), root surface area (RSA), root numbers (RN), and root volume (RV), while it increased root diameter (RD) and Na⁺/K⁺ ratio. Further, it caused severe damages to the root system as shown by a remarkable increase in degree of cell injury and expression of the cell death-related gene OsNAC4. On the other hand, salt stress reduced the total biomass, RSA and RV, and increased the Na⁺/K⁺ ratio, while to less extent than those under alkaline stress. These results suggest that the deleterious effect of alkaline stress on rice seedlings is due mainly to high alkalinity that induces cell injury in the root system. In contrast, the response to osmotic stress exhibited the typical adaptive responses to dehydration: reduced relative water content (RWC), RD, RV, and shoot biomass; whereas increased TRL, RSA, RN, and root biomass. In addition, we show that proline was accumulated most abundantly in response to alkaline treatment, and which was corroborated by the transcriptional induction of OsP5CS1 and OsP5CS2, the genes encoding proline carboxylate synthase.

The objectives of this study were to evaluate the effects of different fertilizer types and application rates on ammonia volatilization loss and to explore nitrogen distribution and nitrogen use efficiency using the 15N isotope tracing technique in different alkaline salt-affected conditions in the Songnen Plain, Northeast China. The results showed a decreasing trend in ammonia volatilization loss from ammonium nitrate and ammonium sulfate, but not that from urea, as the electrical conductivity gradient increased, whereas the reverse trend was found as the pH gradient increased. Ammonia volatilization loss increased in moderately salt-affected soil compared with that in slightly salt-affected soil, particularly during the tillering stage, regardless of the N fertilizer rate. The percentage of N absorbed by rice plants increased from urea but decreased from the soil as the amount of nitrogen was increased. Interestingly, the N retention rate in soil decreased and rice grain yield and nitrogen agronomic efficiency increased as the amount of nitrogen increased in both salt-affected soil conditions. The nitrogen application amount of highest N physiological efficiency was 225 kg·N/ha. Considering high rice production and a minimal environmental threat, we should fully consider controlling ammonia volatilization losses by adjusting the fertilizer type and the crop stage when the fertilizer is applied.

The effect of nitrogen (N) application on seed yields and yield components in Leymus chinensis (Trin.) Tzvel., a perennial rhizomatous grass, was measured in a field experiment with two saline-sodic soils at Da’an Sodic Land Experiment Station during 2010–11. Two grassland field sites were classified as moderately saline–sodic (MSSL) and severely saline–sodic (SSSL). Application rates of N at each site were 0, 30, 60, 90, 120, 150, 180 and 210 kg ha–1. Application of N significantly improved seed yield mainly through increased spike number (R2 = 0.96, P ≤ 0.001). Compared with nil N, seed yield increased 7.4–10.9 times with N application of 150 kg ha–1 at MSSL, and 5.3–7.5 times with N application of 120 kg ha–1 at SSSL. However, absolute increases at SSSL were relatively small. Some significant differences (P ≤ 0.01) in seed yield occurred between 2010 and 2011 with different N application rates in the same soil, and between MSSL and SSSL in the same year. Increasing N application rate significantly decreased N physiological efficiency (NPE) but increased N apparent-recovery fraction (NRF) and N partial-factor productivity (NPP) at both sites. Seed yield and NPP indicated that the optimal N application rates to increase yield were 150 kg ha–1 at MSSL and 120 kg ha–1 at SSSL. High soil pH was the major factor adversely impacting seed yield, and pH and soil salinity were major factors negative affecting NPE, NRF and NPP as well as decreasing the positive effect of N application. Nitrogen application is a practical and effective method to increase seed yield of L. chinensis in saline-sodic grasslands of Northeast China, particularly when soil pH and salinity are not limiting.

Aims Leymus chinensis is an original dominant plant in the Songnen grass- land, and it has great value for restoration of severely degraded land. However, seeds are dormant, and low germination percentage is a problem for restoringL.chinensis grassland. The mechanism of seed dormancy is not been well understood. The primary aims of the pres- ent study were to investigate the dormancy mechanism ofL.chinensis seeds (caryopses) with reference to the role of embryo-covering layers, endogenous hormones and temperature. Methods Changes in concentration of the endogenous hormones GA3, indole- acetic acid (IAA), zeatin riboside (ZR) and abscisic acid (ABA) inL.chi- nensis seeds from anthesis to maturity were measured by the enzyme- linked immunosorbent assay method. Germination at different stages of maturity were tested at 16/28C, 5/28C and 5/35C for intact seeds with glumes (control), intact seeds with glumes removed (naked- whole seeds) and intact seeds with glumes and one-half of the endo- sperm removed (naked-half seeds). Important Findings Of the four endogenous hormones monitored, only the concentra- tion of ZR differed significantly between the beginning and the end of seed development (increased); the GA3/ABA ratio also did not differ. Rank of germination percentage of control at the three tem- perature regimens was 5/28C > 16/28C > 5/35C. Germination per- centage of the naked-half seeds reached 100% under the three temperature regimens. We concluded that dormancy of L.chinensis seeds is not mainly controlled by endogenous hormones. Germina- tion temperature, mechanical resistance of glumes and inhibition of endosperm are the main factors controlling dormancy and germina- tion of L.chinensis seeds.

The net photosynthetic rate of flag leaves and influencing factors under saline sodic soil conditions were investigated at the full heading stage of rice. The net photosynthetic rate of rice leaves showed a double-peak curve in a day in both non-saline sodic and saline sodic soil treatments. The first peak of the net photosynthetic rate appeared at 9:00–10:00 and 9:00 in the saline sodic and non-saline sodic soil treatments, respectively, whereas the second peak both at 14:00. The midday depression of the net photosynthetic rate always appeared regardless of non-saline sodic or saline sodic soil conditions. In addition, the net photosynthetic rate significantly decreased in all day under saline sodic conditions compared with that under non-saline sodic conditions. Some differences were observed in correlation characters between the net photosynthetic rate and all influencing factors during 9:00–13:00. Under non-saline sodic conditions, the diurnal changes of the net photosynthetic rate in a day were mainly caused by stomatal conductance, and the limitation value and the stomatal factors served as determinants; whereas under saline sodic stress, the diurnal changes of the net photosynthetic rate in a day were mainly caused by non stomatal factors including light intensity and air temperature.