Relative growth rates of leaves from soybean grown under drought-stressed and irrigated field conditions

Abstract. The relative growth rates and leaf area were graphed against leaf area, normalized with respect to final leaf area, to assess the applicability of the Lockhart cell wall expansion equation to soybean, Glycine max (L.) Merr., leaf development under field conditions. For leaves that had completed more than 20% of their growth, relative growth rates decreased linearly with an increase in the normalized leaf area, indicating that these leaves were undergoing strictly expansive growth. Drought stress significantly decreased the relative growth rate of these larger leaves. Small leaves which had completed less than 20% of their growth, were found to have highly variable relative growth rates. The large variability in relative growth rates indicated that the Lockhart cell wall expansion equation was inadequate to evaluate the growth of these young leaves. Drought stress had virtually no influence on the relative growth rates observed in the small leaves.     

连作障碍因素对大豆养分吸收和固氮作用的影响

采用分室装置,利用不同孔径的膜研究大豆连作条件下,化感物质、土壤有害生物和大豆胞囊线虫等因素不断累加对植株生长、生物固氮作用和矿质养分吸收和分配的影响。结果表明,随着各因素不断累加,植株的地上部、根系和根瘤干重逐渐降低;除Ca元素外,植株组织的P、K等矿质元素单位含量下降,吸收总量下降,地上部分配的养分比例下降。在化感物质和土壤有害生物因素的基础上接种线虫,对生物固氮和矿质养 分的吸收和分配影响明显。     

Effects of drought on nitrogen fixation in soybean root nodules

Soybean plants [Glycine max (L.) Merr.] were grown in silica sand and were drought stressed for a 4 week period during reproductive development and without any mineral N supply in order to maximize demand for fixed nitrogen. A strain of Bradyrhizobium japonicum that forms large quantities of polysaccharide in nodules was used to determine whether or not the supply of reduced carbon to bacteroids limits nitrogenase activity. A depression of 30–40% in nitrogen content in leaves and pods of stressed plants indicated a marked decline in nitrogen fixation activity during the drought period. A 50% increase in the accumulation of bacterial polysaccharide in nodules accompanied this major decrease in nitrogen fixation activity and this result indicates that the negative impact of drought on nodules was not due to a depression of carbon supply to bacteroids. The drought treatment resulted in a statistically significant increase in N concentration in leaves and pods. Because N concentration and chlorophyll concentration in leaves were not depressed, there was no evidence of nitrogen deficiency in drought‐stressed plants, and this result indicates that the negative impact of drought on nodule function was not the cause of the depression of shoot growth. At the end of the drought period, the concentration of carbohydrates, amino nitrogen, and ureides was significantly increased in nodules on drought‐stressed plants. The overall results support the view that, under drought conditions, nitrogen fixation activity in nodules was depressed because demand for fixed N to support growth was lower.     

UV-B辐射对8个大豆品种种子萌发率和 幼苗生长的影响

在生长房5种(暗处、可见光、低、中、高强度紫外线-B)处理下,研究了8个大豆品种的种子萌发率和萌发后幼苗的生长状况。结果表明,暗处种子萌发率高于自然光和UV-B辐射的种子。UV-B辐射增强对大豆种子的萌发率没有显著影响,仅使部分品种的最大萌发率降低和导致部分品种达到最大萌发率的时间延长。幼苗的生长对增强的UV-B辐射非常敏感。使大部分品种的胚根变短增粗,这可能是植物激素作用的结果。大豆的叶绿素a、叶绿素b和总叶绿素含量明显受到UV-B辐射的抑制。UV-B作用能促进类黄酮在幼苗中的积累,紫外吸收色素的增设有利于提高对UV-B的抵抗力。UV-B辐射的这种效应及大豆品种间的差异在自然情况下会产生深远的生物学和生态学意义     

Microaerobic respiration and oxidative phosphorylation by soybean nodule mitochondria: implications for nitrogen fixation

The infected cells of soybean (Glycine max) root nodules require ATP production for ammonia assimilation and purine synthesis under microaerobic conditions. It is likely that the bulk of this demand is supplied through mitochondrial oxidative phosphorylation. Mitochondria purified from root nodules respired and synthesized ATP in sub-micromolar oxygen concentrations as measured by leghaemoglobin spectroscopy and luciferase luminescence. Both oxygen uptake and the apparent ATP/O ratio declined significantly as the oxygen concentration fell below 100 μmol m^?3. Cytochrome-pathway respiration by root nodule mitochondria had a higher apparent affinity for oxygen (Km 50 μmol m^?3) than did mitochondria isolated from roots (Km 125 μmol m^?3). Electron micrographs showed that mitochondria predominated at the periphery of infected cells adjacent to gas-filled intercellular spaces, where the oxygen concentration is predicted to be highest. Calculations of oxygen concentration and nitrogen fixation rates on an infected cell basis suggest that the measured rates of ATP production by isolated mitochondria are sufficient for the quantifiable in vivo requirements of ammonia assimilation and purine synthesis. The possible roles of mitochondrial respiration in the control of infected cell metabolism are also discussed.     

Sensitivity of soybean leaf development to water deficits

Abstract. Drought effects on the final leaf area of individual leaves were hypothesized to depend on the leaf developmental stage at which drought occurred. To evaluate this hypothesis, final leaf area and cell number were measured for soybean ( Glycine max (L.) Merr.) leaves that were at different stages of development when single or cyclical drought treatment was imposed. Leaf emergence rate from the meristem, as depicted by changes in the plastochron index, was not as sensitive as leaf expansion to cyclical droughts. For leaf expansion, small leaves, once they emerged from the meristem, suffered larger decreases in growth than leaves undergoing rapid leaf area expansion. Decreases in final leaf area as a result of a cyclical drought were correlated with decreases in final cell number. Decreases resulting from a single 8-d drought were dependent on the age of the leaf at the time of drought, because small leaves were found to have proportionately larger decreases in final cell number and area than larger leaves. These results indicated that age-dependent leaf responses to drought are based on the relative activity of cell division and expansion at the time stress was imposed.     

Emergence and seedling growth of soybean cultivars and maturity groups under salinity

Soybean is an important agricultural crop and has, among its genotypes, a relatively wide variation in salt tolerance. As measured by vegetative growth and yield, however, the achievement or failure of a high emergence ratio and seedling establishment in saline soils can have significant economic implications in areas where soil salinity is a potential problem for soybean. This study was conducted to determine the effects of salinity, variety and maturation rate on soybean emergence and seedling growth. Included in the study were the variety ‘Manokin’; four near-isogenic sibling lines of the variety ‘Lee’ belonging to maturity groups IV, V, VI and VII; and the variety ‘Essex’ and two of its near-isogenic related lines representing maturity groups V, VI and VII, respectively. Field plots were salinized with sodium chloride and calcium chloride salts prior to planting. The soybeans were irrigated with furrow irrigation which redistributed the salts towards the tail ends of the field plots. Elevated soil salinity near the tail ends of the field significantly reduced soybean emergence rate, shoot height and root length. No significant reduction was found for emergence or seedling growth of variety ‘Manokin’ when the electrical conductivity of soil solution extract (ECe) was less than 3 dS m⁻¹. Soybean emergence and seedling growth was significantly reduced when soil ECe reached about 11 dS m⁻¹. Maturity groups V and VII of variety ‘Lee’ or V and VI of ‘Essex’ appeared to be more sensitive to salinity stress than other maturity groups. Salt tolerance of different genotypes and maturity groups should be considered, among other limiting factors, in minimizing salinity effects on soybean growth. This revised version was published online in June 2006 with corrections to the Cover Date.     

Development of protein bodies and accumulation of carbohydrates in a soybean (Leguminosae) shriveled seed mutant

The soybean seed mutant T311, when grown under specific environmental conditions, produces shriveled seed. This research investigated changes in development of protein bodies and accumulation of carbohydrates during seed development by comparing the mutant with P2180 seeds. The shriveled seeds contained larger protein bodies but fewer protein bodies per cell than round seeds. Protein bodies in T311 seeds included more dispersed crystals and less globoid regions than P2180 seeds. The elemental compositions of the crystals and of whole seeds in T311 were different from that in P2180 seeds. Starch breakdown was reduced with concomitant lower soluble sugar content in T311 seeds after the D11 stage (10.0-11.9 mm long seeds). The reduced starch breakdown and lowered soluble sugar content were consistent with lower a-amylase activity and earlier and greater water loss in T311 seeds. Changes in development of protein bodies and accumulation of carbohydrates were associated with the development of the shriveled seeds.     

Nodule gas exchange and water potential response to rapid imposition of water deficit

The permeability (P) of the gaseous diffusion barrier in the nodules of soybean [Glycine max (L.) Merr.] decreases when water deficits are extended over a 7 to 10 d period. The mechanism controlling P changes is unclear, but may result from the release of water to intercellular pathways, and an associated change in the nodule water potential. The purpose of these experiments was to impose water deficit treatments rapidly in order to determine the early sequence of the responses of nodule water potential and nodule gas exchange without the complications that arise from long-term water deficit treatments. A vertical, split-root system was used to separate nodule drying effects from plant water deficits by replacing humidified air that was passed over upper root nodules in well-watered plants with dry air, or by replacing the nutrient solution that surrounded lower roots with -1.0 MPa polyethylene glycol (PEG) solution, or by a combination of the dry air and PEG treatments. The PEG treatment caused large decreases in both the components of nodule water potential and nodule relative water content, but there was no indication that these factors had immediate, direct effects on either nitrogenase activity or P. After 7 h of the PEG treatment a significant decrease in nitrogenase activity was found but no decrease in P was detected. These results indicate that changes in nitrogenase activity in response to water deficits precede decreases in P. Exposure of nodules to dry air in well-watered plants had no significant effect on either nitrogenase activity or P during the 7 h treatment.     

Vascular transport and soybean nodule function: nodule xylem is a blind alley, not a throughway

Abstract. The only published consideration of product removal from the soybean root nodule hypothesizes that the peripheral xylem circuit of this determinate nodule structure is flushed by the transpiration stream. However, dyes fed to the transpiration stream through a cut root distal to the nodule do not enter the nodule, and the observed movement of radio-tracers from the root into the nodule can be explained by simple diffusion, Also, there are few xylem elements in the nodule, and these elements are of a small diameter, such that this path can not act as a functional loop of the root system. Further, in this study, nodule vascular strands were never observed to be continuous about the nodule, but were observed to end at the nodule tip in a loop within an intact, closed endodermal sac. Nodule vascular tissue was invested in a pericycle of at least three cell layers. These cells are suggested to be active in the loading of the xylem apoplast with ureides, such that the xylem of the nodule always operates in an export role. Nodule water requirements may be supplied via the phloem or the root cortex apoplasm.     

Responses of soybean to oxygen deficiency and elevated root-zone carbon dioxide concentration

Root flooding is damaging to the growth of crop plants such as soybean (Glycine max L.). Field flooding for 3 d often results in leaf chlorosis, defoliation, cessation of growth and plant death. These effects have been widely attributed solely to a lack of oxygen in the root-zone. However, an additional damaging factor may be CO(2), which attains levels of 30 % (v/v) of total dissolved gases. Accordingly, the effects of root-zone CO(2) on oxygen-deficient soybean plants were investigated in hydroponic culture. Soybean plants are shown to be very tolerant of excess water and anaerobiosis. No oxygen (100 % N(2) gas) and low oxygen (non-aerated) treatments for 14 d had no effect on soybean survival or leaf greenness, but plants became severely chlorotic and stunted when the roots were exposed to no oxygen together with CO(2) concentrations similar to those in flooded fields (equilibrium concentrations of 30 %). When root-zone CO(2) was increased to 50 %, a quarter of soybean plants died. Those plants that survived showed severe symptoms of chlorosis, necrosis and root death. In contrast, rice (Oryza sativa L.) plants were not affected by the combination of no oxygen and elevated root-zone CO(2.) A concentration of 50 % CO(2) did not affect rice plant survival or leaf colour. These results suggest that the high susceptibility of soybean to soil flooding, compared with that of rice, is an outcome of its greater sensitivity to CO(2).     

Abscisic acid mimics effects of dehydration on area expansion and photosynthetic partitioning in young soybean leaves

Abstract. Leaf area expansion, photosynthetic carbon dioxide uptake and leaf dry mass accumulation were compared for expanding leaves of well-watered soybean ( Glycine max [L.] Merr.) plants, mildly dehydrated plants and well-watered plants treated with abscisic acid (ABA). Both ABA treatment and dehydration reduced area expansion in the light and over a 24 h period without decreasing the photosynthetic rates of expanding leaves. Dry mass accumulation during the light was less in ABA-treated and water-stressed leaves than in control leaves, with no differences among treatments in leaf mass per unit of area. ABA treatment and water stress both increased export of carbon from expanding leaves in the light. ABA applied near the end of the light period also increased export of carbon during the following dark period. However, it is unlikely that decreased availability of photosynthate caused slow expansion in the ABA and dehydration treatments, because expansion rates were not slowed in plants kept in dim light, even though photosynthetic rates and dry mass accumulation rates were greatly reduced. The data suggest that ABA may mediate the effects of mild dehydration on leaf area expansion and partitioning of photosynthate.     

A small heat shock protein,GmHSP17.9, from nodule confers symbiotic nitrogen fixation and seed yield in soybean

Legume–rhizobia symbiosis enables biological nitrogen fixation to improve crop production for sustainable agriculture. Small heat shock proteins (sHSPs) are involved in multiple environmental stresses and plant development processes. However, the role of sHSPs in nodule development in soybean remains largely unknown. In the present study, we identified a nodule-localized sHSP, called GmHSP17.9, in soybean, which was markedly up-regulated during nodule development. GmHSP17.9 was specifically expressed in the infected regions of the nodules. GmHSP17.9 overexpression and RNAi in transgenic composite plants and loss of function in CRISPR-Cas9 gene-editing mutant plants in soybean resulted in remarkable alterations in nodule number, nodule fresh weight, nitrogenase activity, contents of poly β-hydroxybutyrate bodies (PHBs), ureide and total nitrogen content, which caused significant changes in plant growth and seed yield. GmHSP17.9 was also found to act as a chaperone for its interacting partner, GmNOD100, a sucrose synthase in soybean nodules which was also preferentially expressed in the infected zone of nodules, similar to GmHSP17.9. Functional analysis of GmNOD100 in composite transgenic plants revealed that GmNOD100 played an essential role in soybean nodulation. The hsp17.9 lines showed markedly more reduced sucrose synthase activity, lower contents of UDP-glucose and acetyl coenzyme A (acetyl-CoA), and decreased activity of succinic dehydrogenase (SDH) in the tricarboxylic acid (TCA) cycle in nodules due to the missing interaction with GmNOD100. Our findings reveal an important role and an unprecedented molecular mechanism of sHSPs in nodule development and nitrogen fixation in soybean.     

缺硼对大豆根瘤结构和功能的影响

在营养液培养条件下以普通结结瘤大豆Ｂｒａｇｇｃｖ．「Ｇｌｙｃｉｎｅｍａｘ（Ｌ．）Ｍｅｒｒ」及其超结瘤突变体ｎｔｓ３８２为实验材料,运用光学显微方法研究了硼对大豆根瘤结构的影响,并测定了根瘤固氮酶活性结果表明,缺硼使根瘤结构受到严重破坏,并使固氮酶活性显著下降,缺硼使根瘤结构受到破坏是导致固氮酶活性下降的可能原因。     

Yield and composition of soybean seed as a function of potassium supply

Summary A reduction in K supply to soybean plants to deficiency levels during both vegetative and reproductive development resulted in reductions not only in yield, but also in oil and K concentrations in the seed and a concomittant increase in seed protein concentration. Correlations between mean fruit yield and oil, protein and K concentrations, over a wide range of K regimes, were 0.97, −0.94 and 0.98, respectively. When K supply was increased well above the level necessary to produce maximum yields,i.e. luxury consumption, there was no significant change in K concentration in the seed, indicating a high degree of control in the movement of K to the developing seed under high K regimes. When the K supply to the plant was limiting, the rate of accumulation of oil and carbohydrate fractions, but not of seed protein, declined during the latter part of podfilling. This resulted in a fall in the C/N ratio in the non-structural seed components during this part of seed development. Depriving plants of K only during seed development had no effect on seed composition or yield, whereas resupplying K to deficient plants after anthesis resulted in almost the same seed composition and yield as that which occurred with control plants. Possible mechanisms whereby K deficiency influences soybean seed composition and yield are discussed in terms of movement of carbohydrate and nitrogen to the seed. We suggest that potassium-deficient soils are likely to produce crops with low yields and low seed oil levels; the crop may respond to K fertilizers as late as anthesis.     

The influence of soybean planting density on dinitrogen fixation and yield

We investigated the effect of planting density on soybean (Glycine max (L.) Merr.) yield in glasshouse and field experiments. Because net canopy photosynthesis increases with increasing plant density, we hypothesized that increasing planting density would result in increasing rates of dinitrogen fixation in soybeans and higher yields per unit land area.In glasshouse studies, Wayne variety soybeans were planted in 10-cm diameter pots, 1 plant pot^-1 in matrices of 10-, 15-, 20-, 25-, or 30-cm equidistant intervals. Bradyrhizobium japonicum inoculum was added to half of the plants in each treatment. Replicate measurements of total stem height, internode lengths, leaf mass, stem mass, root mass, nodule number, nodule mass, and nitrogenase activity were obtained at 3, 6, and 9 weeks post-emergence. Fruits were harvested and counted at week 14. As planting density increased there were (1) altered morphology and growth rates, (2) increased apparent specific nodule activity (SNA), (3) decreased nodule number and mass, and (4) nearly constant fruit and seed production/plant. Expressed on a unit area basis, nitrogen influx and yield increased geometrically as planting density increased, with maximum values observed for 10-cm plantings.Field studies of Wayne, Stein, Williams, and Gold Harvest soybean varieties were made in 1985. Plots were established containing 100 plants spaced at 10-, 20-, and 30-cm distances. Measurements made during the growing season and at harvest established the same relative trends identified from the glasshouse studies. Increasing plant densities resulted in higher yields per unit land. Varietal differences were almost significant.     

Soybean genotypic differences in sensitivity of symbiotic nitrogen fixation to soil dehydration

Nitrogen fixation activity by soybean (Glycine max (L.) Merr.) nodules has been shown to be especially sensitive to soil dehydration. Specifically, nitrogen fixation rates have been found to decrease in response to soil dehydration preceding alterations in plant gas exchange rates. The objective of this research was to investigate possible genetic variation in the sensitivity of soybean cultivars for nitrogen fixation rates in response to soil drying. Field tests showed substantial variation among cultivars with Jackson and CNS showing the least sensitivity in nitrogen accumulation to soil drying. Glasshouse experiments confirmed a large divergence among cultivars in the nitrogen fixation response to drought. Nitrogen fixation in Jackson was again found to be tolerant of soil drying, but the other five cultivars tested, including CNS, were found to be intolerant. Experiments with CNS which induced localized soil drying around the nodules did not result in decreases in nitrogen fixation rates, but rather nitrogen fixation responded to drying of the entire rooting volume. The osmotic potential of nodules was found to decrease markedly upon soil drying. However, the decrease in nodule osmotic potential occurred after significant decreases in nitrogen fixation rates had already been observed. Overall, the results of this study indicate that important genetic variations for sensitivity of nitrogen fixation to soil drying exist in soybean, and that the variation may be useful in physiology and breeding studies.