Foliar Application of NH4 +/NO3 - Ratios Enhance the Lodging Resistance of Soybean Stem by Regulating the Physiological and Biochemical Mechanisms Under Shade Conditions.

Shading is one of the most chronic restrains which can lead to the lodging of intercropped plants. In order to increase the soybean stem lodging resistance, a 2-year field trial was conducted to evaluate the impact of different ratios and concentrations of NH4 +/NO3 - on the morpho-physiological and biochemical characteristics of soybean stem under shade conditions. The total 5 ratios of NH4 +/NO3 - were applied as follows: T0 = 0/0 (control), T1 = 0/100 (higher ratio), T2 = 25/75 (optimum), T3 = 50/50 (optimum), and T4 = 75/25 (higher ratio) as a nitrogen source. Our findings displayed that the T2 (25/75) and T3 (50/50) treatments alleviated the shading stress by improving the photosynthetic activity, biomass accumulation, carbohydrates contents, and lignin related enzymes (POD, CAD, and 4Cl) which led to improvement in stem lodging resistance. The correlation analysis (p ≤ 0.05, p ≤ 0.01) revealed the strong relationship between lodging resistance index and stem diameter, stem strength, lignin content, photosynthesis, and lignin related enzymes (POD, CAD, and 4CL) evidencing the strong contribution of lignin and its related enzymes in the improvement of lodging resistance of soybean stem under shade conditions. Collectively, we concluded that optimum NH4 +/NO3 - ratios (T2 and T3) can boost up the lodging resistance of soybean stem under shade stress.

Titanium Application Increases Phosphorus Uptake Through Changes in Auxin Content and Root Architecture in Soybean (Glycine Max L.).

Phosphorus (P) is an essential macronutrient needed for plant growth, development, and production. A deficiency of P causes a severe impact on plant development and productivity. Several P-based fertilizers are being used in agriculture but limited uptake of P by the plant is still a challenge to be solved. Titanium (Ti) application increases the nutrient uptake by affecting the root growth; however, the role of Ti in plant biology, specifically its application under low light and phosphorus stress, has never been reported. Therefore, a pot study was planned with foliar application of Ti (in a different concentration ranging from 0 to 1,000 mg L-1) under different light and P concentrations. The result indicated that under shade and low P conditions the foliar application of Ti in different concentrations significantly improves the plant growth parameters such as root length, root surface area, root dry matter, and shoot dry matters. The increase was observed to be more than 100% in shade and low P stressed soybean root parameter with 500 mg L-1 of Ti treatment. Ti was observed to improve the plant growth both in high P and low P exposed plants, but the improvement was more obvious in Low P exposed plants. Auxin concentration in stressed and healthy plant roots was observed to be slightly increased with Ti application. Ti application was also observed to decrease rhizosphere soil pH and boosted the antioxidant enzymatic activities with an enhancement in photosynthetic efficiency of soybean plants under shade and P stress. With 500 mg L-1 of Ti treatment, the photosynthetic rate was observed to improve by 45% under shade and P stressed soybean plants. Thus, this work for the first time indicates a good potential of Ti application in the low light and P deficient agricultural fields for the purpose to improve plant growth and development parameters.

Foliar application of silicon improves growth of soybean by enhancing carbon metabolism under shading conditions.

An experiment was set up to investigate physiological responses of soybeans to silicon (Si) under normal light and shade conditions. Two soybean varieties, Nandou 12 (shade resistant), and Nan 032-4 (shade susceptible), were tested. Our results revealed that under shading, the net assimilation rate and the plant growth were significantly reduced. However, foliar application of Si under normal light and shading significantly improved the net photosynthetic rate (Pn), stomatal conductance (Gs), transpiration rate (Tr), and decreased intercellular carbon dioxide concentration (Ci). The net photosynthetic rate of Nandou 12 under normal light and shading increased by 46.4% and 33.3% respectively with Si treatment (200 mg/kg) compared to controls. Si application also enhanced chlorophyll content, soluble sugars, fresh weight, root length, root surface area, root volume, root-shoot ratio, and root dry weight under both conditions. Si application significantly increased the accumulation of some carbohydrates such as soluble sugar and sucrose in stems and leaves ensuring better stem strength under both conditions. Si application significantly increased the yield by increasing the number of effective pods per plant, the number of beans per plant and the weight of beans per plant. After Si treatment, the yield increased 24.5% under mono-cropping, and 17.41% under intercropping. Thus, Si is very effective in alleviating the stress effects of shading in intercropped soybeans by increasing the photosynthetic efficiency and lodging resistance.

Foliar application of silicon improves stem strength under low light stress by regulating lignin biosynthesis genes in soybean (Glycine max (L.) Merr.).

In order to improve soybean's resistance to lodging, silicon (Si) solutions at concentrations of 0,100, 200,300 mg kg-1 were applied during the seedling stage. The Si accumulation in different parts of the plants, the photosynthetic parameters of leaves and chlorophyll content, the stem bending resistance, the expression of genes of lignin biosynthesis and associated enzyme activity and sap flow rates were measured at early and late growth stages. The potential mechanisms for how Si improve growth and shade tolerance, enhances lodging resistance and improves photosynthesis were analyzed to provide a theoretical basis for the use of Si amendments in agriculture. After application of Si at 200 mg kg-1, the net photosynthetic rate of soybeans increased by 46.4 % in the light and 33.3 % under shade. The application of Si increased chlorophyll content, and fresh weight of leaves, reduced leaf area and enhanced photosynthesis by increasing stomatal conductance. The activity of peroxidase (POD), 4-coumarate:CoA ligase (4CL), cinnamyl alcohol dehydrogenase (CAD) and phenylalanine ammonia-lyase (PAL) increased during pre-and post-growth periods, whereas Si also increased lignin accumulation and inhibited lodging. We concluded that Si affects the composition of plant cell walls components, mostly by altering linkages of non-cellulosic polymers and lignin. The modifications of the cell wall network through Si application could be a useful strategy to reduce shading stress in intercropping.

Agro-Techniques for Lodging Stress Management in Maize-Soybean Intercropping System-A Review.

Lodging is one of the most chronic restraints of the maize-soybean intercropping system, which causes a serious threat to agriculture development and sustainability. In the maize-soybean intercropping system, shade is a major causative agent that is triggered by the higher stem length of a maize plant. Many morphological and anatomical characteristics are involved in the lodging phenomenon, along with the chemical configuration of the stem. Due to maize shading, soybean stem evolves the shade avoidance response and resulting in the stem elongation that leads to severe lodging stress. However, the major agro-techniques that are required to explore the lodging stress in the maize-soybean intercropping system for sustainable agriculture have not been precisely elucidated yet. Therefore, the present review is tempted to compare the conceptual insights with preceding published researches and proposed the important techniques which could be applied to overcome the devastating effects of lodging. We further explored that, lodging stress management is dependent on multiple approaches such as agronomical, chemical and genetics which could be helpful to reduce the lodging threats in the maize-soybean intercropping system. Nonetheless, many queries needed to explicate the complex phenomenon of lodging. Henceforth, the agronomists, physiologists, molecular actors and breeders require further exploration to fix this challenging problem.

Acclimation strategy and plasticity of different soybean genotypes in intercropping.

In response to shading, plant leaves acclimate through a range of morphological, physiological and biochemical changes. Plants produce a myriad of structurally and functionally diverse metabolites that play many important roles in plant response to continually changing environmental conditions as well as abiotic and biotic stresses. To develop a clearer understanding of the effects of shade on soybeans at different growth stages, a comprehensive, three-year, stage-wise study was conducted. Leaf area, leaf thickness, stem diameter, chlorophyll contents, photosynthetic characteristics and other morphological and physiological features were measured along with biochemical assays for antioxidants such as superoxide dismutase, peroxidase and caralase and yield attributes of different soybean genotypes (Guixia 2, Nandou12, Nandong Kang-22, E61 and C103) under shading nets with 50% light transmittance. It was observed that early shading (VER1 and VER2) significantly decreased main stem length and main stem length/stem diameter. Later shading (R1R8 and R2R8) had significant effects on morphological characters such as branch number and pod height. In Nandou 12, the protein contents in plants shaded at R1R8, R2R8 and R5R8 were 9.20, 8.98 and 6.23% higher than in plants grown under normal light levels (CK), respectively, and the crude fat content was 9.31, 10.74 and 4.28% lower. The influence of shading in the later period on anatomy was greater than that in the earlier period. Shading reduced the light saturation point (LSP), the light compensation point (LCP) and the maximum photosynthetic rate (Pnmax), and increased the apparent quantum yield (AQ). Shading also increased the antioxidant enzyme activity in the plants, and this increase was greater with early shading than late. The variability in the chlorophyll (a + b) content and the chlorophyll a/b ratio in R2 stage plants was less than in R5 stage (VER5) plants. Similarly, the activity of antioxidant enzymes in R2 after returning the plants to normal light levels (VER2) was lower than in R5 after relighting (VER5). Compared with later shading, the early shading had a greater effect on the photosynthetic and related characteristics. The longer the shading time, the greater the adverse effects and the less able the plants' were to recover. The data collected in this study contribute to an understanding of the physiological mechanisms underlying the early and late growth stage acclimation strategies in different soybean genotypes subjected to shade stress.

Effects of lignin, cellulose, hemicellulose, sucrose and monosaccharide carbohydrates on soybean physical stem strength and yield in intercropping.

Soybean (Glycine max L.) has been extensively cultivated in maize-soybean relay intercropping systems in southwest China. However, during the early co-growth period, soybean seedlings suffer from severe shading by maize resulting in lodging and significant yield reduction. The purpose of the present research was to investigate the reasons behind severe lodging and yield loss. Therefore, four different soybean genotypes (B3, B15, B23, and B24) having different agronomic characteristics were cultivated in intercropping and monocropping planting patterns. The results showed that under different planting patterns, the stem resistance varied among genotypes (P < 0.01). The lodging resistance index of B3, B15, B23, and B24 genotypes was 70.9%, 60.5%, 65.2%, and 57.4%, respectively, under intercropping, among which the B24 genotype was less affected by the shade environment as there was little decrease in the lodging resistance index of this genotype under intercropping. The lignin content of B23 and B24 was significantly higher than that of B3 and B15 under both planting patterns. Under intercropping, the hemicellulose content of B23 and B24 stems was significantly higher than that of B3 and B15. Compared to the monocropping, the content of mannose in the structural carbohydrate of soybean stems was decreased in all genotypes except B23, but the difference was not significant. The content of xylose in the structural carbohydrate of soybean stems was significantly higher than that in B3 and B15. Mannose content showed no significant difference among genotypes. The arabinose content of B24 was significantly higher than that of B3, B15, and B23. The effective pod number, seed number per plant, seed weight per plant and yield of soybean plants were significantly decreased under intercropping. Conclusively, manipulation of structural and nonstructural carbohydrate rich soybean genotypes in intercropping systems could alleviate the yield loss due to lodging.