Weixing Cao

Effects of salt and waterlogging stresses and their combination on leaf photosynthesis, chloroplast ATP synthesis, and antioxidant capacity in wheat

The objective of this study was to investigate the effects of salt (ST) and waterlogging (WL) stresses and their combination (SW) on leaf photosynthesis, chloroplast ATP synthesis, and antioxidant capacity in wheat (Triticum aestivum L.). Two winter wheat cultivars, Huaimai 17 and Yangmai 12, differing in their tolerance to ST and WL stresses were used. The plants were grown in pots and were subjected to ST, WL, and SW from 7 days after anthesis (DAA). The WL and SW treatments lasted for 5 days, while the ST treatment was continuously imposed during the grain filling stage. Significant decrease in net photosynthetic rate (PN) of the flag leaf was observed under the ST and SW treatments from 10 DAA in Yangmai 12 and at 18 DAA in both cultivars, which could be stomatal closure related. At 18 DAA, clear reduction in PN under the ST and SW treatments was observed, which was associated with chlorosis, damages to the photosystem II (PSII), enhanced lipid peroxidation, and depressed ATP synthesis in the chloroplasts of the flag leaf. Whereas, WL treatment alone had slightly negative effect on PN, which was mainly attributed to leaf chlorosis and waste in harvested energy by the PSII reaction center dispersed via non-photochemical approaches.

Alterations in photosynthesis and antioxidant enzyme activity in winter wheat subjected to post-anthesis water-logging

Winter wheat (Triticum aestivum L.) cultivars Yangmai 9 (water-logging tolerant) and Yumai 34 (water-logging sensitive) were subjected to water-logging (WL) from 7 d after anthesis to determine the responses of photosynthesis and anti-oxidative enzyme activities in flag leaf. At 15 d after treatment (DAT), net photosynthetic rate under WL was only 3.7 and 8.9 µmol(CO2) m−2 s−1 in Yumai 34 and Yangmai 9, respectively, which was much lower than in the control. Ratios of variable to maximum and variable to initial fluorescence, actual photosynthetic efficiency, and photochemical quenching were much lower, while initial fluorescence and non-photochemical quenching were much higher under WL than in control, indicating damage to photosystem 2. WL decreased activities of superoxide dismutase and catalase in both cultivars, and activity of peroxidase (POD) in Yumai 34, while POD activity in Yangmai 9 was mostly increased. The obvious decrease in the amount of post-anthesis accumulated dry matter, which was redistributed to grains, also contributed to the grain yield loss under WL.

Pre-anthesis high-temperature acclimation alleviates damage to the flag leaf caused by post-anthesis heat stress in wheat.

The objective of this study was to investigate the effect of pre-anthesis high-temperature acclimation on leaf physiology of winter wheat in response to post-anthesis heat stress. The results showed that both pre- and post-anthesis heat stresses significantly depressed flag leaf photosynthesis and enhanced cell membrane peroxidation, as exemplified by increased O₂⁻(·) production rate and reduction in activities of antioxiditave enzymes. However, under post-anthesis heat stress, plants with pre-anthesis high-temperature acclimation (HH) showed much higher photosynthetic rates than those without pre-anthesis high-temperature acclimation (CH). Leaves of HH plants exhibited a higher Chl a/b ratio and lower chlorophyll/carotenoid ratio and superoxide anion radical release rate compared with those of the CH plants. In addition, antioxidant enzyme activities in HH plants were significantly higher than in CH. Coincidently, expressions of photosythesis-responsive gene encoding Rubisco activase B (RcaB) and antioxidant enzyme-related genes encoding mitochondrial manganese superoxide dismutase (Mn-SOD), chloroplastic Cu/Zn superoxide dismutase (Cu/Zn-SOD), catalase (CAT) and cytosolic glutathione reductase (GR) were all up-regulated under HH, whereas a gene encoding a major chlorophyll a/b-binding protein (Cab) was up-regulated by post-anthesis heat stress at 10 DAA, but was down-regulated at 13 DAA. The changes in the expression levels of the HH plants were more pronounced than those for the CH. Collectively, the results indicated that pre-anthesis high-temperature acclimation could effectively alleviate the photosynthetic and oxidative damage caused by post-anthesis heat stress in wheat flag leaves, which was partially attributable to modifications in the expression of the photosythesis-responsive and antioxidant enzymes-related genes.

Relationships of endogenous plant hormones to accumulation of grain protein and starch in winter wheat under different post-anthesis soil water statusses

Accumulation of protein and starch in grain is a key process determining grain yield and quality in wheat. Under drought or waterlogging, endogenous plant hormone levels will change and may have an impact on the yield and quality of wheat. In a greenhouse experiment, four winter wheat (Triticum aestivum L.) varieties differing in grain protein content, Heimai 76, Wanmai 38, Yangmai 10 and Yangmai 9, were subjected to drought (SRWC = 45∼50%, DR), waterlogging (WL) and moderate water supply (SRWC = 75∼80%, CK), beginning from 4 days post-anthesis (DPA) to maturity. On the 10 (grain enlargement stage) and 20 (grain filling stage) DPA, endogenous abscisic acid (ABA), gibberellins (GA1+3), indole-3-acetic acid (IAA) and zeatin riboside (ZR) were determined in sink and source organs of wheat plants by enzyme linked immunosorbent assay (ELISA). The patterns of hormonal changes were similar in four varieties. The ABA levels were much higher under DR and WL than under CK. Compared with CK, GA1+3 levels in whole-plant under DR and WL changed a little at 10 DPA, but markedly decreased under DR and WL at 20 DPA. Changes of endogenous IAA level under DR and WL exhibited a complicated pattern, depending on organs and growth stages. Particularly at the 20 DPA, the mean levels of IAA in roots, leaves and grains decreased significantly under DR and WL. In comparison with CK, ZR levels in all organs significantly decreased under DR and WL at both stages. The correlation analyses between yields and contents of starch and protein in grains and levels and ratios of four hormones in source and sink organs indicated that the changes in yield and content of grain starch and protein under DR and WL were associated with the reduced IAA, ZR and GA1+3 levels and elevated ABA level in plants, especially in grains. It was proposed that the changed levels of endogenous hormones under post-anthesis DR and WL might indirectly affect protein and starch accumulation in grains by influencing the regulatory enzymes and processes.

Activities of key enzymes for starch synthesis in relation to growth of superior and inferior grains on winter wheat (Triticum aestivum L.) spike

Weight of individual grains is a major yield component in wheat. The non-uniform distribution of single grain weight on a wheat spike is assumed to be closely associated with starch synthesis in grains. The present study was undertaken to determine if the enzymes involved in starch synthesis cause the differences in single grain weight between superior and inferior grains on a wheat spike. Using two high-yield winter wheat (Triticum aestivum L.) varieties differing in grain weight and three nitrogen rates for one variety, the contents of amylose and amylopectin, and activities of enzymes involved in starch synthesis in both superior and inferior grains were investigated during the entire period of grain filling. Superior grains showed generally higher starch accumulation rates and activities of enzymes including SS (sucrose synthase), UDPGPPase (UDP-glucose pyrophosphorylase), ADPGPPase (ADP-glucose pyrophosphorylase), SSS (soluble starch synthase) and GBSS (starch granule bound starch synthase) and subsequently produced much higher single grain weight than inferior grains. Nitrogen increased enzyme activities and starch accumulation rates, and thus improved individual grain weight, especially for inferior grains. The SS, ADPGPPase and SSS were significantly correlated to amylopectin accumulation, while SS, ADPGPPase, SSS and GBSS were significantly correlated to amylose accumulation. This infers that SS, ADPGPPase and starch synthase play key roles in regulating starch accumulation and grain weight in superior and inferior grains on a wheat spike.

Multiple heat priming enhances thermo-tolerance to a later high temperature stress via improving subcellular antioxidant activities in wheat seedlings.

Seedlings of winter wheat (Triticum aestivum L.) were firstly twice heat-primed at 32/24 °C, and subsequently subjected to a more severe high temperature stress at 35/27 °C. The later high temperature stress significantly decreased plant biomass and leaf total soluble sugars concentration. However, plants experienced priming (PH) up-regulated the Rubisco activase B encoding gene RcaB, which was in accordance with the higher photosynthesis rate in relation to the non-primed plants (NH) under the later high temperature stress. In relation to NH, the major chlorophyll a/b-binding protein gene Cab was down-regulated in PH plants, implying a reduction of the light absorption to protect the photosystem II from excitation energy under high temperature stress. At the same time, under the later high temperature stress PH plants showed significantly higher actual photochemical efficiency, indicating an improvement of light use efficiency due to the priming pre-treatment. Under the later high temperature stress, PH could be maintained a better redox homeostasis than NH, as exemplified by the higher activities of superoxide dismutase (SOD) in chloroplasts and glutathione reductase (GR), and of peroxidase (POD) in mitochondria, which contributed to the lower superoxide radical production rate and malondialdehyde concentration in both chloroplasts and mitochondria. The improved antioxidant capacity in chloroplasts and mitochondria was related to the up-regulated expressions of Cu/Zn-SOD, Mn-SOD and GR in PH. Collectively, heat priming effectively improved thermo-tolerance of wheat seedlings subjected to a later high temperature stress, which could be largely ascribed to the enhanced anti-oxidation at the subcellular level.

Induction of chilling tolerance in wheat during germination by pre-soaking seed with nitric oxide and gibberellin

Chilling depresses seed germination and seedling establishment, and is one major constraint to grain yield formation in late sown winter wheat. Seeds of winter wheat (Triticum aestivum L.) were separately pre-soaked with sodium nitroprusside (SNP, as nitric oxide donor) and Gibberellic acid (GA3) before germination and then germinated under low temperature. SNP and GA3 pre-treatment increased seed germination rate, germination index, weights and lengths of coleoptile and radicle, while they decreased mean germination time and weight of seeds germinating under low temperature. Exogenous NO and GA3 increased seed respiration rate and promoted starch degradation along with increased amylase activities. In addition, efficient antioxidant systems were activated by NO, and which effectively reduced concentrations of malondialdehyde and hydrogen peroxide (H2O2). Seedling growth was also enhanced by exogenous NO and GA3 as a result of improved seed germination and maintenance of better reactive oxygen species homeostasis in seedling growing under chilling temperatures. It is indicated that exogenous NO was more effective than GA3 in alleviating chilling stress during seed germination and seedling establishment in wheat.

Cold priming drives the sub-cellular antioxidant systems to protect photosynthetic electron transport against subsequent low temperature stress in winter wheat

Low temperature seriously depresses the growth of wheat through inhibition of photosynthesis, while earlier cold priming may enhance the tolerance of plants to subsequent low temperature stress. Here, winter wheat plants were firstly cold primed (5.2xa0°C lower temperature than the ambient temperature, viz., 10.0xa0°C) at the Zadoks growth stage 28 (i.e.xa0re-greening stage, starting on 20th of March) for 7xa0d, and after 14xa0d of recovery the plants were subsequently subjected to a 5xa0d low temperature stress (8.4xa0°C lower than the ambient temperature, viz., 14.1xa0°C) at the Zadoks growth stage 31 (i.e.xa0jointing stage, starting on 8th April). Compared to the non-primed plants, the cold-primed plants possessed more effective oxygen scavenging systems in chloroplasts and mitochondria as exemplified by the increased activities of SOD, APX and CAT, resulting in a better maintenance in homeostasis of ROS production. The trapped energy flux (TRO/CSO) and electron transport (ETO/CSO) in the photosynthetic apparatus were found functioning well in the cold-primed plants leading to higher photosynthetic rate during the subsequent low temperature stress. Collectively, the results indicate that cold priming activated the sub-cellular antioxidant systems, depressing the oxidative burst in photosynthetic apparatus, hereby enhanced the tolerance to subsequent low temperature stress in winter wheat plants.

Effects of Long-Term Fertilization on Leaf Photosynthetic Characteristics and Grain Yield in Winter Wheat

Based on a 20-year fertilization experiment with wheat-maize double cropping system, the effects of different long-term fertilization treatments on leaf photosynthetic characteristics and grain yield in different winter wheat (Triticum aestivum L.) cultivars were studied in the growing seasons of 2000–2001 and 2001–2002. A total of nine fertilization treatments were implemented, i.e. no fertilizer (CK), N fertilizer (N), N and P fertilizers (NP), N and K fertilizers (NK), N, P, and K fertilizers (NPK), only organic manure (M), organic manure and N fertilizer (MN), organic manure and N and P fertilizers (MNP), and organic manure and N, P, and K fertilizers (MNPK). With the treatments of combined organic manure and inorganic fertilizers (TMI), net photosynthetic rate (PN), maximal activity of photosystem 2, PS2 (Fv/Fm), and chlorophyll content (SPAD value) of flag leaves and leaf area index (LAI) were much higher at the mid grain filling stage (20 or 23 d post anthesis, DPA), and exhibited slower declines at the late grain filling stage (30 DPA), compared with the treatments of only inorganic fertilizers (TI). The maximal canopy photosynthetic traits expressed as PN×LAI and SPAD×LAI at the mid grain filling stage were also higher in TMI than those in TI, which resulted in different grain yields in TMI and TI. Among the treatments of TMI or among the treatments of TI, both flag leaf and canopy photosynthetic abilities and yield levels increased with the supplement of inorganic nutrients (N, P, and K fertilizers), except for the treatment of NK. Under NK, soil contents of N and K increased while that of P decreased. Hence the unbalanced nutrients in soil from the improper input of nutrients in NK treatment were probably responsible for the reduced flag leaf and canopy photosynthetic characteristics and LAI, and for the fast declining of flag leaf photosynthetic traits during grain filling, resulting in the reduced yield of NK similar to the level of CK.

Effects of exogenous ABA and cytokinin on leaf photosynthesis and grain protein accumulation in wheat ears cultured in vitro

A detached culture system and steady-state 15N labeling technique were used to study the effects of exogenous ABA and ZR on photosynthetic characteristics, nitrogen remobilization and the activities of key enzymes for nitrogen metabolism in detached wheat parts during grain protein accumulation. The differences in net photosynthetic rate, chlorophyll content (SPAD value) and soluble protein content in the flag leaves of detached culture system between the treatments of ABA and ZR showed that ABA facilitates the post-anthesis senescence course compared to the ZR treatment. The differences in the changes of 15N amount in different organs in the detached culture system between the ABA and ZR treatments showed that nitrogen remobilization from vegetative organs to the grain is accelerated by the ABA treatment but is delayed by ZR. The activities of GS and GPT in grains treated with ABA were significantly higher than those with the control treatment at 5 DAC, but reduced significantly compared with control at 11 DAC. The two enzyme activities in grains were reduced significantly by ZR at 5 DAC and increased significantly at 11 DAC, compared with those treated with ABA. The above changes of enzyme activity showed that the ABA treatment hastens amino acid conversion into grains and protein accumulation in grains, whereas the ZR treatment delays these processes. A significant reduction in grain weight with ABA treatment is associated with the reduction of net photosynthesis, chlorophyll content, and soluble protein content in flag leaves. Compared with the control and ZR treatments, a significant increase in grain protein content with the ABA treatment may result from the accelerating effects of ABA on N remobilization, amino acid conversion into grains and protein accumulation in grains.