Biostimulant effects of rhizobacteria on wheat growth and nutrient uptake depend on nitrogen application and plant development

The capacity of plant growth-promoting rhizobacteria (PGPR) – Bacillus amyloliquefaciens GB03 (BamGB03), B. megaterium SNji (BmeSNji), and Azospirillum brasilense 65B (Abr65B) – to enhance growth and nutrient uptake in wheat was evaluated under different mineral N fertilizer rates, in sterile and non-sterile soils, and at different developmental stages. In gnotobiotic conditions, the three strains significantly increased plant biomass irrespective of the N rates. Under greenhouse conditions using non-sterile soil, growth promotion was generally highest at a moderate N rate, followed by a full N dose, while no significant effect was observed for the inoculants in the absence of N fertilizer. At 50N, plant biomass was most significantly increased in roots (up to +45% with Abr65B) at stem-elongation stage and in the ears (+19–23% according to the strains) at flowering stages. For some nutrients (N, P, Mn, and Cu), the biomass increases in roots and ears were paralleled with reduced nutrient concentrations in the same organs. Nevertheless, growth stimulation resulted in a higher total nutrient uptake and higher nutrient uptake efficiency. Furthermore, Abr65B and BmeSNji counteracted the reduction of root development caused by a high N supply. Therefore, combining PGPR with a proper cultivated system, N rate, and plant stage could enhance their biostimulant effects.     

Release of phytosiderophores from roots of wheat and triticale under nickel‐deficient conditions

Despite numerous studies on phytosiderophores (PS) there is still an open question whether nickel (Ni) deficiency induces release of PS from graminaceous plant roots. Seedlings of two wheat cultivars (Triticum aestivum L. cvs. Rushan and Kavir) and a triticale cultivar (X. triticosecale) were grown in Ni‐free nutrient solution (Ni‐deficient, Ni–) and with 10 µM NiSO₄ (Ni‐sufficient, Ni+, control). Root exudates were collected weekly for 4 weeks and the amount of PS in the root exudates was measured. The response to Ni deficiency on the release of PS differed between species. Roots of Rushan and triticale exuded higher PS in response to Ni‐deficient conditions. Nickel deficiency significantly enhanced shoot Fe and Zn concentrations in wheat, while it decreased shoot Fe and Zn concentrations in triticale. In Kavir, PS exudation was decreased by Ni deficiency at weeks 3 and 4 and the reduced release of PS from roots of Kavir was accompanied by lower concentrations of Fe and Zn in plant roots but higher Fe and Zn concentrations in shoot tissue. The PS release by Kavir was triggered by a Ni‐induced Zn deficiency particularly in the shoots. According to the results, it is suggested that in the studies concerning the phytosiderophore release under Ni deficiency, special attention should be given to different responses among and within cereals and to the plant Zn or Fe nutritional status.     

Timely supplemental irrigation changed nitrogen use of wheat by regulating root vertical distribution

Crop nitrogen (N) uptake depends on the root absorption area and the soil N availability which are closely related to the soil water status. With the increasing water shortages in the North China Plain, supplemental irrigation (SI) to winter wheat is a promising technique. To clarify the relationships between water and nitrogen use, four SI regimes in Tritcum aestivum L. cv. Jimai 22 were set up: no‐irrigation after emergence (T1), SI at jointing and anthesis (T2), SI at sowing, jointing and anthesis (T3), and SI at pre‐wintering, jointing and anthesis (T4). The results indicate that T2 had higher root length density (RLD) and root surface area density (RAD) in the 0–20, 60–80, and 80–100 cm soil layers, as well as higher post‐anthesis N uptake from soil by 23–26% in 2012–2013 and 162–177% in 2013–2014, compared to T3 and T4. The grain yield under T2 was lower than T3 but was not significantly different from T4, whereas its water use efficiency (WUE) was higher relative to both T3 and T4. There were no significant differences among T2, T3, and T4 in N use efficiency (NUE). The N uptake after jointing and WUE were positively correlated with the RLD and RAD in the 0–20 cm soil layer. The NUE was positively correlated with the RLD and RAD in the 20–40 cm soil layer. These results indicate that timely SI at jointing and anthesis was dependent on a suitable water supply at sowing, which increased the soil water content in the upper soil layer after jointing and improved the absorption area of the roots in both the deep and surface soil layers; this further improved the post‐anthesis N uptake from the soil and the WUE. This approach can be a valuable way to maintain high grain yields and NUE in winter wheat while using less irrigation and achieving higher WUE in the North China Plain.     

Effect of zinc‐biofortified seeds on grain yield of wheat,rice, and common bean grown in six countries

Seeds enriched with zinc (Zn) are ususally associated with better germination, more vigorous seedlings and higher yields. However, agronomic benefits of high‐Zn seeds were not studied under diverse agro‐climatic field conditions. This study investigated effects of low‐Zn and high‐Zn seeds (biofortified by foliar Zn fertilization of maternal plants under field conditions) of wheat (Tritcum aestivum L.), rice (Oryza sativa L.), and common bean (Phaseolus vulgaris L.) on seedling density, grain yield and grain Zn concentration in 31 field locations over two years in six countries. Experimental treatments were: (1) low‐Zn seeds and no soil Zn fertilization (control treatment), (2) low‐Zn seeds + soil Zn fertilization, and (3) Zn‐biofortified seeds and no soil Zn fertilization. The wheat experiments were established in China, India, Pakistan, and Zambia, the rice experiments in China, India and Thailand, and the common bean experiment in Brazil. When compared to the control treatment, soil Zn fertilization increased wheat grain yield in all six locations in India, two locations in Pakistan and one location in China. Zinc‐biofortified seeds also increased wheat grain yield in all four locations in Pakistan and four locations in India compared to the control treatment. Across all countries over 2 years, Zn‐biofortified wheat seeds increased plant population by 26.8% and grain yield by 5.37%. In rice, soil Zn fertilization increased paddy yield in all four locations in India and one location in Thailand. Across all countries, paddy yield increase was 8.2% by soil Zn fertilization and 5.3% by Zn‐biofortified seeds when compared to the control treatment. In common bean, soil Zn application as well as Zn‐biofortified seed increased grain yield in one location in Brazil. Effects of soil Zn fertilization and high‐Zn seed on grain Zn density were generally low. This study, at 31 field locations in six countries over two years, revealed that the seeds biofortfied with Zn enhanced crop productivity at many locations with different soil and environmental conditions. As high‐Zn grains are a by‐product of Zn biofortification, use of Zn‐enriched grains as seed in the next cropping season can contribute to enhance crop productivity in a cost‐effective manner.     

Regulation of mycorrhiza development in durum wheat by P fertilization: Effect on plant nitrogen metabolism

The aim of this work was to study the effect of arbuscular mycorrhizal fungus Glomus mosseae on growth and nitrogen (N) metabolism of durum wheat (Tritcum durum) under various P soil contents. The analyses were extended to macro and micronutrient tissue concentrations, nitrate reductase and glutamine synthetase activities, as well as protein, aminoacids, pyridine dinucleotides and adenine nucleotides. Arbuscular mycorrhiza increased wheat growth in soil in which P availability was low and nitrate was the dominant N form. The root colonization occurred at the highest level in plants grown in limiting soil P and was inversely related to soil P content. The micorrhizal wheat plants contained also the highest concentrations of macro (P, K, Ca, N) and micronutrients (Fe, Zn, Mn) as well as free amino acids, protein, NAD, NADP, AMP, ADP, ATP in roots and leaves. In particular, the micronutrient tissue concentrations (Zn, Mn) supported that mycorrhiza actively modulated their uptake limiting interferences and optimizing growth better than the plant roots, like a very efficient “rootstock”. Control plants grown at the highest soil P did not reach the same concentration as the mycorrhizal plants. Nitrate reductase activities in the roots of mycorrhizal plants were higher than in the control ones, while glutamine synthetase activities were highest in the leaves. Protein and amino acids concentrations, as well as AMP, ADP, ATP, NAD(P), and NAD(P)H were also higher than in the control. Among the free amino acids in the roots, the high levels of glutamine, asparagine, arginine, support the view that ammonium was transferred through the arbuscules to the root cells where it was re‐assimilated in the cortical cells, forming high N : C ratio‐amino acids. They were transferred to the leaves where all the other N compounds could be largely synthesized using the carbon skeletons supplied by photosynthesis.     

Impact of post‐flowering nitrate availability on nitrogen remobilization in hydroponically grown durum wheat

The nitrogen (N) utilization efficiency in hydroponically grown durum wheat was dissected during nitrate depletion at post‐flowering stage. Feeding plants with low nitrate dose after anthesis lowered photosynthetic activity, triggered senescence, but induced N remobilization from vegetative parts. At harvest, tissue N concentration was 50% and 80% less important in leaves and culms, respectively, but similar in grains of N‐deficient plants compared with control ones.     

Effects of earthworms and plant growth–promoting rhizobacteria (PGPR) on availability of nitrogen,phosphorus, and potassium in soil

Both earthworms and plant growth–promoting rhizobacteria (PGPR) are ubiquitous and important for promoting circulation of plant macronutrients. Two series of laboratory experiments were conducted to investigate the effects of earthworm casts and activities on the growth of PGPR, and the inoculation of earthworms and PGPR on the availability of N, P, and K in soils, respectively. During a short incubation period (0–34 h), the extracts of earthworm (Pheretima guillelmi)‐worked soil significantly (p < 0.05) increased the abundance of the three species of PGPR, including N‐fixing bacteria (NFB) (Azotobacter chroococcum HKN‐5), phosphate‐solubilizing bacteria (PSB) (Bacillus megaterium HKP‐1), and K‐solubilizing bacteria (KSB) (B. mucilaginous HKK‐1), in Luria‐Bertani (LB) broth. There were synergistic effects of dual inoculation of earthworms and PGPR on increasing the concentrations of NH$ _4^+ $ ‐N, (NO$ _3^- $ + NO$ _2^- $ )‐N, NaHCO₃‐extractable P, and NH₄OAc‐extractable K in the corresponding soils. Bioavailable N (the sum of NH$ _4^+ $ ‐N and [NO$ _3^- $ + NO$ _2^- $ ]‐N) in the dual inoculation was 4 to 24 times those inoculated with earthworms or NFB alone, respectively. The significantly higher concentrations of bioavailable N and P in the dual inoculation of earthworms and NFB or PSB may be due to the higher abundance of PGPR and/or higher activities of urease and acid phosphatase than those of single inoculation of NFB or PSB, respectively. Dual inoculation of earthworms and PGPR would be most effective in reducing the need for chemical fertilizers in agriculture.     

基于土壤-作物系统模拟模型的冬小麦田间水氮优化管理

将作物生长模型与土壤水氮管理过程模块相结合,构建了土壤-作物系统水分养分模拟模型。以灌水、施氮总量为决策变量,冬小麦生物量、水分和氮肥利用效率为优化目标,将冬小麦按生长发育时期分为6个阶段,建立了多目标的动态规划模型。在土壤-作物系统过程模型的基础上,用动态规划的方法对田间水、氮资源管理措施进行优化。通过对作物水分胁迫系数和氮肥胁迫系数的模拟计算,可获得最佳的灌水、施肥时间及用量。算例结果表明：在养分供应充足仅水分胁迫的条件下,优化方案的灌水量较对照处理平均节约了25%,水分利用效率比对照处理平均高出约1     

Effects of 15N-labelled ammonium nitrate and urea on soil nitrogen during growth of wheat (Triticum aestivum L.) under field conditions

We studied the effects of ¹⁵N-labelled ammonium nitrate and urea on the yield and uptake of labelled and unlabelled N by wheat (Triticum aestivum L., cv. Mexi-Pak-65) in a field experiment. The dry matter and N yields were significantly increased with fertilizer N application compared to those from unfertilized soil. The wheat crop used 33.6–51.5 and 30.5–40.9% of the N from ammonium nitrate and urea, respectively. Splitting the fertilizer N application had a significant effect on the uptake of fertilizer N by the wheat. The fertilizer N uptake showed that ammonium nitrate was a more available source of N for wheat than urea. The effective use of fertilizer N (ratio of fertilizer N in grain to fertilizer N in whole plant) was statistically similar for the two N fertilizers. The application of fertilizer N increased the uptake of unlabelled soil N by wheat, a result attributed to a positive added N interaction, which varied according to the fertilizer N split; six split applications gave the highest added N interaction compared to a single application or two split applications for both fertilizers. Ammonium nitrate gave 90.5, 33.5, and 48.5% more added N interaction than urea with one, two, and six split N applications. A values were not significantly correlated with the added N interaction (r=0.557). The observed added N interaction may have been the result of pool substitution, whereby added labelled fertilizer N replaced unlabelled soil N.     

Priming effects of 15N-labelled ammonium nitrate on uptake of soil N by wheat (Triticum aestivum L.) under field conditions

Summary The uptake of labelled and unlabelled N by wheat was measured in a field experiment using ¹⁵N-labelled ammonium nitrate fertilizer. The dry matter yield and N yields were significantly increased with fertilizer N application compared to those from unfertilized soil. The uptake of applied N by wheat ranged between 25 and 34%. Fertilizer N application increased the uptake of unlabelled soil N which was attributed to a positive priming effect or added N interaction. The added N interaction observed by applying 20, 60, and 120 kg fertilizer N was 11.4, 19.1, and 27.9 kg, corresponding to 26, 44 and 64%, respectively of the N taken up from unfertilized soil. The A values did not alter with the increase in fertilizer N application. The observed added N interaction may have been the result of pool substitution whereby added labelled fertilizer N stood proxy for unlabelled soil N. A significant correlation coefficient (r=0.996^**) between the uptake of soil N and the dry matter yield showed that soil N was more important than fertilizer N in wheat production.     

Crop rotation and seasonal effects on fatty acid profiles of neutral and phospholipids extracted from no‐till agricultural soils

Analysis of phospholipids (PLFA) and neutral lipids fatty acids (NLFA) was used to characterize no‐till productive agricultural soils associated with different crop rotation levels, replicated across a 400 km transect in the Argentinean pampas, during two sampling seasons, summer and winter. High rotation (HR) management consisted in maize–wheat–soybean intense rotation including cover crops. Low rotation (LR) management trend to soybean monocultures. Soils from nearby natural environments (NEs) were used as references. Fatty acids concentration in soils (nmol/g) decreased c.a. 50% from summer to winter differentially according to soil treatment being the smallest decrease in HR management 35%. Both PLFA and NLFA profiles showed strong potential to discriminate between different land uses. In winter samples, some rare or unknown fatty acids were relevant for the discrimination of agricultural practices while NLFA 20:0 appears to be a good marker of HR soils despite season or location. The PLFA‐based taxonomic biomarkers for total bacteria, Gram‐negative bacteria and arbuscular mycorrhiza showed a significant trend NE>HR>LR in the winter sampling. HR management was also characterized by high levels of NLFA in winter samples as if high crop rotation improves lipids reserves in soil during winter more than in monocropping soil management. In conclusion, PLFA and particularly NLFA profiles appear to provide useful and complementary information to obtain a footprint of different soil use and managements, improving soil biochemistry characterization tools.     

Differential effects of soil disturbance and plant residue retention on function of arbuscular mycorrhizal (AM) symbiosis are not reflected in colonization of roots or hyphal development in soil

The effects of soil disturbance and residue retention on the functionality of the symbiosis between medic (Medicago truncatula L.) and arbuscular mycorrhizal fungi (AMF) were assessed in a two-stage experiment simulating a crop rotation of wheat (Triticum aestivum L.) followed by medic. Plants were inoculated or not with the AMF, Glomus intraradices and Gigaspora margarita, separately or together. The contribution of the arbuscular mycorrhizal (AM) pathway for P uptake was determined using ³²P-labeled soil in a small hyphal compartment accessible only to hyphae of AMF. In general AM colonization was not affected by soil disturbance or residue application and disturbance did not affect hyphal length densities (HLDs) in soil. At 4 weeks disturbance had a negative effect on growth and phosphorus (P) uptake of plants inoculated with G. margarita, but not G. intraradices. By 7 weeks disturbance reduced growth of plants inoculated with G. margarita or AMF mix and total P uptake in all inoculated plants. With the exception of plants inoculated with G. margarita in disturbed soil at 4 weeks, the AM pathway made a significant contribution to P uptake in all AM plants at both harvests. Inoculation with both AMF together eliminated the negative effects of disturbance on AM P uptake and growth, showing that a fungus insensitive to disturbance can compensate for loss of contribution of a sensitive one. Application of residue increased growth and total P uptake of plants but decreased ³²P in plants inoculated with the AMF mix in disturbed soil, compared with plants receiving no residue. The AMF responded differently to disturbance and G. intraradices, which was insensitive to disturbance, compensated for lack of contribution by the sensitive G. margarita when they were inoculated together. Colonization of roots and HLDs in soil were not good predictors of the outcomes of AM symbioses on plant growth, P uptake or P delivery via the AM pathway.     

The effects of compost in a rice–wheat cropping system on aggregate size,carbon and nitrogen content of the size–density fraction and chemical composition of soil organic matter,as shown by 13C CP NMR spectroscopy

We investigated whether the long‐term application of compost from agricultural waste improved soil physical structure, fertility and soil organic matter (SOM) storage. In 2006, we began a long‐term field experiment based on a rice–wheat rotation cropping system, having a control without fertilizer (NF) and three treatments: chemical fertilizers (CF), pig manure compost (PMC) and a prilled mixture of PMC and inorganic fertilizers (OICF). Following the harvest of wheat in 2010, the mean‐weight diameter (MWD) of water‐stable aggregates and the concentration of C and N in bulk soil (0–20 cm; <2 mm fraction) were significantly greater (P < 0.05) in PMC and NF plots than in CF or OICF plots. Pig manure compost significantly increased the proportion of >5‐mm aggregates, whereas CF significantly increased the proportion of 0.45‐ to 1‐mm aggregates. The C and N contents of all density fractions were greater in PMC than in other treatments with levels decreasing in the following order: free particulate organic matter (fPOM) >occluded particulate organic matter (oPOM) > mineral‐combined SOM (mineral–SOM). Solid‐state ¹³C CPMAS NMR spectra showed that alkyl C/O‐alkyl C ratios and aromatic component levels of SOM were smaller in PMC and OICF plots than in CF plots, suggesting that SOM in PMC and OICF plots was less degraded than that in CF plots. Nevertheless, yields of wheat in PMC and NF plots were smaller than those in CF and OICF plots, indicating that conditions for producing large grain yields did not maintain soil fertility.