QTL mapping of root hair and acid exudation traits and their relationship to phosphorus uptake in common bean

The relationship between root-hair growth, acid exudation and phosphorus (P) uptake as well as the quantitative trait loci (QTLs) associated with these traits were determined for a recombinant inbred line (RIL) population derived from the cross of two contrasting common bean (Phaseolus vulgaris L.) genotypes, DOR364 and G19833, which were grown in solution culture and under field conditions with low-P availability. In the solution-culture study, root-hair density, root-hair length, H⁺ exudation and total acid exudation were measured. Substantial genotypic variability was observed for these traits and their response to P availability. The P-efficient parent G19833 had greater root-hair density, longer root-hair length, and greater exudation of H⁺ and total acid than the P-inefficient genotype DOR364. These traits segregated continuously in the RIL population, with obvious tendency of trait transgression. Genetic analysis revealed that the root traits measured had various heritabilities, with h _b ² ranging from 43.24 to 86.70%. Using an integrated genetic map developed for the population, a total of 19 QTLs associated with root hair, acid exudation and P-uptake traits were detected on 8 linkage groups. P uptake in the field was positively correlated with total acid exudation, basal root-hair length, and basal root-hair density. Acid-exudation traits were intercorrelated, as were root-hair traits. Total acid exudation was positively correlated with basal root-hair density and length. Linkage analysis revealed that some of the root-trait QTLs were closely linked with QTLs for P uptake in the field. We propose that marker-assisted selection (MAS) might be a feasible alternative to conventional screening of phenotypic root traits.     

Effect of phosphorus availability on basal root shallowness in common bean

Root gravitropism may be an important element of plant response to phosphorus availability because it determines root foraging in fertile topsoil horizons, and thereby phosphorus acquisition. In this study we seek to test this hypothesis in both two dimensional paper growth pouch and three-dimensional solid media of sand and soil cultures. Five common bean (Phaseolus vulgaris L.) genotypes with contrasting adaptation to low phosphorus availability were evaluated in growth pouches over 6 days of growth, and in sand culture and soil culture over 4 weeks of growth. In all three media, phosphorus availability regulated the gravitropic response of basal roots in a genotype-dependent manner. In pouches, sand, and soil, the phosphorus-inefficient genotype DOR 364 had deeper roots with phosphorus stress, whereas the phosphorus-efficient genotype G19833 responded to phosphorus stress by producing shallower roots. Genotypes were most responsive to phosphorus stress in sand culture, where relative root allocation to the 0–3- and 3–6-cm horizons increased 50% with phosphorus stress, and varied 300% (3–6 cm) to 500% (0–3 cm) among genotypes. Our results indicate that (1) phosphorus availability regulates root gravitropic growth in both paper and solid media, (2) responses observed in young seedlings continue throughout vegetative growth, (3) the response of root gravitropism to phosphorus availability varies among genotypes, and (4) genotypic adaptation to low phosphorus availability is correlated with the ability to allocate roots to shallow soil horizons under phosphorus stress.     

Potassium influx into maize root systems : influence of root potassium concentration and ambient ammonium

Potassium influx into roots of dark-grown decapitated maize seedling (Zea mays L., cv Pioneer 3369A) was examined in presence and absence of ambient ammonium and at various root potassium concentrations. Six-day old seedlings which were dependent on the endosperm reserves for their energy source were exposed to KCl (labeled with ⁸⁶Rb) ranging from 5 to 200 micromolar. At both low (13 micromoles per gram fresh weight) and high (100 micromoles per gram fresh weight) root potassium concentration, isotherms indicated two potassium influx systems, one approaching saturation at 50 to 100 micromolar potassium and an additional one tentatively considered to be linear. A mixed-type inhibition by ammonium for the low-concentration saturable system was indicated by a concomitant decrease in V_max and increase in K_m. High root potassium concentration decreased V_max but had little effect on K_m of this system. The rate constant for the second quasilinear system was decreased by ambient ammonium and by high root potassium status. Transfer of high potassium roots to potassium-free solutions resulted in an increase in influx within 2 hours; by 24 hours influx significantly exceeded that of roots not previously exposed to potassium. In roots of both low and high root potassium concentrations, potassium influx was restricted progressively as ambient ammonium increased to about 100 micromolar, but there was little further inhibition as ammonium concentrations increased beyond that to 500 micromolar. The data imply that potassium influx has two components, one subject to inhibition by ambient ammonium and one relatively resistant.     

不同磷水平下小麦-蚕豆间作根系形态的变化及其与内源激素的关系

采用水培方法,研究了不同磷水平下小麦-蚕豆间作体系根系形态变化及其与内源激素的相关关系。结果表明: 与单作小麦相比,在低磷(1/2P)水平下,小麦-蚕豆间作能显著增加小麦的根长,显著减少小麦根系的平均直径,显著增加根系的表面积;在常规磷(P)水平下,间作能显著降低小麦根系的平均直径,有增加小麦根长和根表面积的趋势;与单作蚕豆相比,间作能明显促进蚕豆根系的增长,同时增加蚕豆根表面积。在1/2P水平下,间作能显著提高小麦和蚕豆根系中的生长素(IAA)、脱落酸(ABA)、水杨酸(SA)和茉莉酸(JA)含量;在P水平下,间作能显著提高小麦根系中的IAA、ABA和JA含量,单、间作小麦根系中的SA含量没有显著差异,间作显著增加了蚕豆根系中ABA和SA含量,单、间作蚕豆根系中的IAA和JA含量无显著差异。单作条件下,小麦和蚕豆根系中的内源激素(IAA、ABA、SA和JA)含量与其根系形态(根长、根平均直径和根表面积)无显著相关性;间作条件下,小麦和蚕豆根系中的IAA含量与根长和根表面积之间存在明显的正相关关系。由此可见,小麦-蚕豆间作能够诱导小麦和蚕豆根系IAA的增加。这种变化可能是驱动间作系统根系形态变化的重要因子。     

The role of maize root size in phosphorus uptake and productivity of maize/faba bean and maize/wheat intercropping systems

Interspecific root/rhizosphere interactions affect phosphorus (P) uptake and the productivity of maize/faba bean and maize/wheat intercropping systems. The aim of these experiments was to determine whether manipulation of maize root growth could improve the productivity of the two intercropping systems. Two near isogenic maize hybrids (the larger-rooted T149 and smaller-rooted T222) were intercropped with faba bean and wheat, under conditions of high- and low-P availability. The larger-rooted T149 showed greater competitive ability than the smaller-rooted T222 in both maize/faba bean and maize/wheat intercropping systems. The higher competitive ability of T149 improved the productivity of the maize/faba bean intercropping system in P-sufficient conditions. In maize/wheat intercropping systems, root growth, shoot biomass, and P uptake of maize were inhibited by wheat, regardless of the P-supply. Compared with T222, the larger-rooted T149 suffered less in the intercropping systems. The total biomass of the maize/wheat intercropping system was higher for wheat/T149 than for wheat/T222 under low-P conditions. These data suggested that genetic improvement of maize root size could enhance maize growth and its ability to compete for P resources in maize/faba bean and maize/wheat intercropping systems. In addition, depending on the P availability, larger maize roots could increase the productivity of intercropping systems.     

小麦与蚕豆间作体系根系形态与磷吸收的定量解析

豆科与禾本科作物间作能够改变作物根系生长,但不同施磷水平下间作-根系形态-磷吸收之间的关系尚未明确.本研究通过田间定位试验和根箱模拟试验,研究不同种植模式(小麦单作、蚕豆单作和小麦-蚕豆间作)和不同磷水平下小麦和蚕豆的产量、生物量、磷吸收及根系形态特征,分析探讨不同施磷条件下小麦-蚕豆间作对根系形态和磷吸收的影响.结果...     

Basal root whorl number: a modulator of phosphorus acquisition in common bean (Phaseolus vulgaris)

Background and Aims

Root architectural phenes enhancing topsoil foraging are important for phosphorus acquisition. In this study, the utility of a novel phene is described, basal root whorl number (BRWN), that has significant effects on topsoil foraging in common bean (Phaseolus vulgaris).

Methods

Whorls are defined as distinct tiers of basal roots that emerge in a tetrarch fashion along the base of the hypocotyl. Wild and domesticated bean taxa as well as two recombinant inbred line (RIL) populations were screened for BRWN and basal root number (BRN). A set of six RILs contrasting for BRWN was evaluated for performance under low phosphorus availability in the greenhouse and in the field. In the greenhouse, plants were grown in a sand–soil media with low or high phosphorus availability. In the field, plants were grown in an Oxisol in Mozambique under low and moderate phosphorus availability.

Key Results

Wild bean accessions tended to have a BRWN of one or two, whereas cultivated accessions had BRWN reaching four and sometimes five. BRWN and BRN did not vary with phosphorus availability, i.e. BRWN was not a plastic trait in these genotypes. Greater BRWN was beneficial for phosphorus acquisition in low phosphorus soil. Genotypes with three whorls had almost twice the shoot biomass, greater root length and greater leaf area than related genotypes with two whorls. In low phosphorus soil, shoot phosphorus content was strongly correlated with BRWN (R² = 0·64 in the greenhouse and R² = 0·88 in the field). Genotypes with three whorls had shallower root systems with a greater range of basal root growth angles (from 10 to 45 ° from horizontal) than genotypes with two whorls (angles ranged from 60 to 85 ° from horizontal).

Conclusions

The results indicate that BRWN is associated with increased phosphorus acquisition and that this trait may have value for selection of genotypes with better performance in low phosphorus soils.     

Fractal geometry of root systems: Field observations of contrasting genotypes of common bean (Phaseolus vulgaris L.) grown under different phosphorus regimes

Root growth and architecture are important for phosphorus acquisition due to the relative immobility of P in the soil. Fractal geometry is a potential new approach to the analysis of root architecture. Substantial genetic variation in root growth and architecture has been observed in common bean. Common bean (Phaseolus vulgaris L.) genotypes with contrasting root architecture were grown under moderate and low P conditions in a field experiment. Linear and planar fractal dimension were measured by tracing root intercepts with vertical planes. Linear fractal dimension increased over time in efficient genotypes, but remained fairly constant over time in inefficient genotypes. Planar fractal dimension increased over time for all genotypes, but was higher in efficient than inefficient genotypes at the end of the experiment. Planar fractal dimension of medium P plants was found to correlate with shoot P content indicating fractal dimension to be a possible indicator for root P uptake. The increasing fractal dimension over time indicates that fractal analysis is a sensitive measure of root branching intensity. A less destructive method for acquisition of data that allows for continuous analysis of fractal geometry and thereby screening for more P efficient genotypes in the field is suggested. This method will allow the researcher to conduct fractal analysis and still complete field trials with final yield evaluation.     

Fractal geometry of bean root systems: correlations between spatial and fractal dimension

An obstacle to the study of root architecture is the difficulty of measuring and quantifying the three-dimensional configuration of roots in soil. The objective of this work was to determine if fractal geometry might be useful in estimating the three-dimensional complexity of root architecture from more accessible measurements. A set of results called projection theorems predict that the fractal dimension (FD) of a projection of a root system should be identical to the FD of roots in three-dimensional space (three-dimensional FD). To test this prediction we employed SimRoot, an explicit geometric simulation model of root growth derived from empirical measurements of common bean (Phaseolus vulgaris L.). We computed the three-dimensional FD, FD of horizontal plane intercepts (planar FD), FD of vertical line intercepts (linear FD), and FD of orthogonal projections onto planes (projected FD). Three-dimensional FD was found to differ from corresponding projected FD, suggesting that the analysis of roots grown in a narrow space or excavated and flattened prior to analysis is problematic. A log-linear relationship was found between FD of roots and spatial dimension. This log-linear relationship suggests that the three-dimensional FD of root systems may be accurately estimated from excavations and tracing of root intersections on exposed planes.     

Hypoxic stress and the transport systems of the peribacteroid membrane of bean root nodules

When the roots of Vicia faba L. beans were subjected to hypoxic stress, the activity of H⁺-ATPase on the peribacteroid membrane, as well as the transport of dicarboxylates (malate and succinate) mediated by this enzyme, decreased. Since malate and succinate are the main carbon-containing metabolites involved in the energy supply to bacteroids, this caused a change of the relation type from mutualism to commensalism, and the domination of the eukaryotes over the prokaryotes consequently increased.     

Modeling nitrogen uptake in oilseed rape cv Capitol during a growth cycle using influx kinetics of root nitrate transport systems and field experimental data

The use of kinetic equations of NO3- transport systems in oilseed rape (Brassica napus), determined by 15NO3- labeling under controlled conditions, combined with experimental field data from the INRA-Chalons rape database were used to model NO3- uptake during the plant growth cycle. The quantitative effects of different factors such as day/night cycle, ontogenetic stages, root temperature, photosynthetically active radiation, and soil nitrate availability on different components of the constitutive high-affinity transport systems, constitutive low-affinity transport systems, inducible low-affinity transport systems, and inducible high-affinity transport systems of nitrate were then determined to improve the model's predictions. Simulated uptake correlated well with measured values of nitrogen (N) uptake under field conditions for all N fertilization rates tested. Model outputs showed that the high-affinity transport system accounted for about 89% of total NO3- uptake (18% and 71% for constitutive high-affinity transport systems and inducible high-affinity transport systems, respectively) when no fertilizer was applied. The low-affinity transport system accounted for a minor proportion of total N uptake, and its activity was restricted to the early phase of the growth cycle. However, N fertilization in spring increased the duration of its contribution to total N uptake. Overall, data show that this mechanistic and environmentally regulated approach is a powerful means to simulate total N uptake in the field with the advantage of taking both physiologically regulated processes at the overall plant level and specific nitrate transport system characteristics into account.     

Partitioning of shoot and root dry matter and carbohydrates in bean plants suffering from phosphorus, potassium and magnesium deficiency

The influence of varied supply of phosphorus (10 and 250 mmolP m^–3) potassium (50 and 2010 mmol K m^–3) and magnesium(20 and 1000 mmol Mg m^–3) on the partitioning of dry matterand carbohydrates (reducing sugars, sucrose and starch) betweenshoots and roots was studied in bean (Phaseolus vulgaris) plantsgrown in nutrient solution over a 12 d period. Shoot and rootgrowth were quite differently affected by low supply of P, K,and Mg. The shoot/root dry weight ratios were 4.9 in the control(sufficient plants), 1.8 in P-deficient, 6.9 in K-deficientand 10.2 in Mg-deficient plants. In primary (source) leaves,but not in trifoliate leaves, concentrations of reducing sugars,sucrose and starch were also differently affected by low nutrientsupply. In primary leaves under K deficiency and, particularlyMg deficiency, the concentrations of sucrose and reducing sugarswere much higher than in control and P-deficient plants. Magnesiumdeficiency also distinctly increased the starch concentrationin the primary leaves. In contrast, in roots, the lowest concenfrationsof sucrose, reducing sugars and starch were found in Mg-deficientplants, whereas the concentrations of sucrose and starch wereparticularly high in P-deficient plants. There was a close relationshipbetween shoot/root dry weight ratios and relative distributionof total carbohydrates (sugars and starch) in shoot and roots.Of the total amounts of carbohyd rates per plant, the followingproportions were parti tioned to the roots: 22.7% in P-deficient,15.7% in control, 3.4% in K-deficient and 0.8% in Mg-deficientplants. The results indicate a distinct role of Mg and K in the exportof photosynthates from leaves to roots and suggest that alterationin photosynthate partitioning plays a major role in the differencesin dry matter distribution between shoots and roots of plantssuffering from mineral nutrient deficiency. Key words: Bean, carbohydrates, magnesium nutrition, phosphorus nutrition, potassium nutrition, shoot/root growth     

Spatial changes and influencing factors of fine root carbon,nitrogen and phosphorus stoichiometry of plants in China

为了研究中国陆地植物细根碳(C)、氮(N)、磷(P)的空间变化模式,揭示细根在"温度-植物生理假说"及"生长速率假说"等方面的规律,该文收集已发表的有关中国陆地植物细根研究的文献,从中提取细根C、N、P元素含量及其相关数据,分析了细根C、N、P含量及其比例与经纬度之间的关系。结果表明:细根N、P元素含量均随纬度增加而增加,P含量随经度增加而降低,N:P随经度增加而增加。细根N、P含量与年平均气温、年降水量均呈负相关关系,与土壤养分呈正相关关系。在土壤养分、温度、降水量3个非生物因素中,土壤养分对细根N、P含量的影响最大。该文中细根和粗根的C:P、N:P差异变化不完全支持"生长速率假说"。根系和叶片一样,N、P含量与纬度呈正相关关系,支持"温度-植物生理假说",反映了植物对自然环境的适应策略。     

中国植物细根碳、氮、磷化学计量学的空间变化及其影响因子

为了研究中国陆地植物细根碳(C)、氮(N)、磷(P)的空间变化模式, 揭示细根在“温度-植物生理假说”及“生长速率假说”等方面的规律, 该文收集已发表的有关中国陆地植物细根研究的文献, 从中提取细根C、N、P元素含量及其相关数据, 分析了细根C、N、P含量及其比例与经纬度之间的关系。结果表明: 细根N、P元素含量均随纬度增加而增加, P含量随经度增加而降低, N:P随经度增加而增加。细根N、P含量与年平均气温、年降水量均呈负相关关系, 与土壤养分呈正相关关系。在土壤养分、温度、降水量3个非生物因素中, 土壤养分对细根N、P含量的影响最大。该文中细根和粗根的C:P、N:P差异变化不完全支持“生长速率假说”。根系和叶片一样, N、P含量与纬度呈正相关关系, 支持“温度-植物生理假说”, 反映了植物对自然环境的适应策略。     

Tracking sulfur and phosphorus within single starch granules using synchrotron X-ray microfluorescence mapping

Background

Native starch accumulates as granules containing two glucose polymers: amylose and amylopectin. Phosphate (0.2–0.5%) and proteins (0.1–0.7%) are also present in some starches. Phosphate groups play a major role in starch metabolism while granule-bound starch synthase 1 (GBSS1) which represents up to 95% of the proteins bound to the granule is responsible for amylose biosynthesis.

Methods

Synchrotron micro-X-ray fluorescence (μXRF) was used for the first time for high-resolution mapping of GBSS1 and phosphate groups based on the XRF signal of sulfur (S) and phosphorus (P), respectively. Wild-type starches were studied as well as their related mutants lacking GBSS1 or starch-phosphorylating enzyme.

Results

Wild-type potato and maize starch exhibited high level of phosphorylation and high content of sulfur respectively when compared to mutant potato starch lacking glucan water dikinase (GWD) and mutant maize starch lacking GBSS1. Phosphate groups are mostly present at the periphery of wild-type potato starch granules, and spread all over the granule in the amylose-free mutant. P and S XRF were also measured within single small starch granules from Arabidopsis or Chlamydomonas not exceeding 3–5 μm in diameter.

Conclusions

Imaging GBSS1 (by S mapping) in potato starch sections showed that the antisense technique suppresses the expression of GBSS1 during biosynthesis. P mapping confirmed that amylose is mostly present in the center of the granule, which had been suggested before.

General significance

μXRF is a potentially powerful technique to analyze the minor constituents of starch and understand starch structure/properties or biosynthesis by the use of selected genetic backgrounds.