Effects of drought stress on legume symbiotic nitrogen fixation: physiological mechanisms

Drought stress is one of the major factors affecting nitrogen fixation by legume-rhizobium symbiosis. Several mechanisms have been previously reported to be involved in the physiological response of symbiotic nitrogen fixation to drought stress, i.e. carbon shortage and nodule carbon metabolism, oxygen limitation, and feedback regulation by the accumulation of N fixation products. The carbon shortage hypothesis was previously investigated by studying the combined effects of CO2 enrichment and water deficits on nodulation and N2 fixation in soybean. Under drought, in a genotype with drought tolerant N2 fixation, approximately four times the amount of 14C was allocated to nodules compared to a drought sensitive genotype. It was found that an important effect of CO2 enrichment of soybean under drought was an enhancement of photo assimilation, an increased partitioning of carbon to nodules, whose main effect was to sustain nodule growth, which helped sustain N2 rates under soil water deficits. The interaction of nodule permeability to O2 and drought stress with N2 fixation was examined in soybean nodules and led to the overall conclusion that O2 limitation seems to be involved only in the initial stages of water deficit stresses in decreasing nodule activity. The involvement of ureides in the drought response of N2 fixation was initially suspected by an increased ureide concentration in shoots and nodules under drought leading to a negative feedback response between ureides and nodule activity. Direct evidence for inhibition of nitrogenase activity by its products, ureides and amides, supported this hypothesis. The overall conclusion was that all three physiological mechanisms are important in understanding the regulation of N2 fixation and its response of to soil drying.     

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.     

Relationship between carbohydrate partitioning and drought resistance in common bean

Drought is a major yield constraint in common bean (Phaseolus vulgaris L.). Pulse-chase (14)C-labelling experiments were performed using Pinto Villa (drought resistant) and Canario 60 (drought sensitive) cultivars, grown under optimal irrigation and water-deficit conditions. Starch and the radioactive label incorporated into starch were measured in leaves and pods at different time points, between the initiation of pod development and the production of mature pods. The water-stress treatment induced a higher starch accumulation in the drought-resistant cultivar pods than in those of the drought-sensitive cultivar. This effect was more noticeable during the early stages of pod development. Consistently, a reduction of starch content occurred in the leaves of the drought-resistant cultivar during the grain-filling stage. Furthermore, a synchronized accumulation of sucrose was observed in immature pods of this cultivar. These data indicate that carbohydrate partitioning is affected by drought in common bean, and that the modulation of this partitioning towards seed filling has been a successful strategy in the development of drought-resistant cultivars. In addition, our results suggest that, in the drought-resistant cultivar, the efficient carbon mobilization towards the seeds in response to water limitation is favoured by a mechanism that implies a more effective sucrose transport.     

Review article. Symbiotic N2 fixation response to drought

Symbiotic nitrogen fixation is highly sensitive to drought, which results in decreased N accumulation and yield of legume crops. The effects of drought stress on N2 fixation usually have been perceived as a consequence of straightforward physiological responses acting on nitrogenase activity and involving exclusively one of three mechanisms: carbon shortage, oxygen limitation, or feedback regulation by nitrogen accumulation. The sensitivity of the nodule water economy to the volumetric flow rate of the phloem into the nodule offers a common framework to understand each of these mechanism. As these processes are sensitive to volumetric phloem flow into the nodules, variations in phloem flow as a result of changes in turgor pressure in the leaves are likely to cause rapid changes in nodule activity. This could explain the special sensitivity of N2 fixation to drying soils. It seems likely that N feedback may be especially important in explaining the response mechanism in nodules. A number of studies have indicated that a nitrogenous signal(s), associated with N accumulation in the shoot and nodule, exists in legume plants so that N2 fixation is inhibited early in soil drying. The existence of genetic variation in N2 fixation response to water deficits among legume cultivars opens the possibility for enhancing N2 fixation tolerance to drought through selection and breeding.     

Comparison of inhibition of N2 fixation and ureide accumulation under water deficit in four common bean genotypes of contrasting drought tolerance

Background and Aims

Drought is the principal constraint on world production of legume crops. There is considerable variability among genotypes in sensitivity of nitrogen fixation to drought, which has been related to accumulation of ureides in soybean. The aim of this study was to search for genotypic differences in drought sensitivity and ureide accumulation in common bean (Phaseolus vulgaris) germplasm that may be useful in the improvement of tolerance to water deficit in common bean.

Methods

Changes in response to water deficit of nitrogen fixation rates, ureide content and the expression and activity of key enzymes for ureide metabolism were measured in four P. vulgaris genotypes differing in drought tolerance.

Key Results

A variable degree of drought-induced nitrogen fixation inhibition was found among the bean genotypes. In addition to inhibition of nitrogen fixation, there was accumulation of ureides in stems and leaves of sensitive and tolerant genotypes, although this was higher in the leaves of the most sensitive ones. In contrast, there was no accumulation of ureides in the nodules or roots of stressed plants. In addition, the level of ureides in the most sensitive genotype increased after inhibition of nitrogen fixation, suggesting that ureides originate in vegetative tissues as a response to water stress, probably mediated by the induction of allantoinase.

Conclusions

Variability of drought-induced inhibition of nitrogen fixation among the P. vulgaris genotypes was accompanied by subsequent accumulation of ureides in stems and leaves, but not in nodules. The results indicate that shoot ureide accumulation after prolonged exposure to drought could not be the cause of inhibition of nitrogen fixation, as has been suggested in soybean. Instead, ureides seem to be produced as part of a general response to stress, and therefore higher accumulation might correspond to higher sensitivity to the stressful conditions.     

Soybean genotypic difference in growth, nutrient accumulation and ultrastructure in response to manganese and iron supply in solution culture

* BACKGROUND AND AIMS: The objective of this research was to characterize the physiology and cell ultrastructure of two soybean genotypes subjected to nutrient solutions with increasing concentrations of manganese (Mn) at two contrasting iron (Fe) concentrations. Genotypes 'PI227557' and 'Biloxi' were selected based on their distinctly different capacities to accumulate Mn and Fe. * METHODS: Bradyrhizobium-inoculated plants were grown in hydroponic cultures in a greenhouse. Nutrient solutions were supplied with Mn concentrations ranging from 0.3 to 90 microm, at either 5 or 150 microm Fe as FeEDTA. * KEY RESULTS: For both genotypes and at both Fe concentrations, Mn concentrations from 6.6 to 50 microm did not affect shoot, root and nodule mass, or leaf and nodule ureide concentration. Mn concentrations of 70 and 90 microm did not result in visible toxicity symptoms, but hindered growth and nodulation of 'Biloxi'. An Mn concentration of 0.3 microm was, however, deleterious to growth and nodulation for both genotypes, and caused an accumulation of ureides in leaves and major alterations in the ultrastructure of chloroplasts, nuclei and mitochondria, regardless of the Fe concentration. In 'PI227557', there was also a proliferation of Golgi apparatus and endoplasmic reticulum in the cytoplasm of leaf cells, and nodules showed disrupted symbiosomes lacking poly-beta-hydroxybutirate grains concomitantly with a proliferation of endoplasmic reticulum as well as arrested bacterial division. At 15 microm Fe, ferritin-like crystals were formed in the lumen of chloroplasts of 'PI227557' plants. For both genotypes, there was an antagonism between the Fe and Mn concentrations in leaves, the higher values of both microelements being detected in 'PI227557'. The absence of any detectable relationship between Fe or Mn and zinc, phosphorus and copper concentrations in leaves ruled out those micronutrients as relevant for Mn and Fe nutrition in soybeans. * CONCLUSIONS: The results confirmed the greater capacity of 'PI227557' for Mn and Fe accumulation than 'Biloxi' for most nutrient treatments. Hence, 'PI227557' may be a very useful genetic resource both in developing soybean cultivars for growth on low nutrient soils and in physiological studies to understand differing approaches to nutrient accumulation in plants.     

Soybean cultivar variability for nodule formation and growth under drought

Both nodulation and nitrogen fixation in soybean [Glycine max (L.) Merr.] are sensitive to soil drying, which can have important negative effects on yield. An exception to this general response has been the identification of the cultivar Jackson as being drought tolerant for N₂ fixation. The objectives of this research were to examine nodule formation and growth in Jackson among other soybean cultivars in response to soil drying under field conditions. Two field experiments were conducted to examine the genetic variation in the sensitivity of nodule numbers and dry weights to soil drying. Substantial variation among soybean lines was found, and the drought-tolerance trait was demonstrated again in Jackson. Greenhouse experiments were conducted to further analyze the variation of nodulation response to soil water content. The differences among cultivars observed in the field were confirmed in the greenhouse. Importantly, the relative drought insensitivity of N₂ fixation in cultivar Jackson was associated with high individual nodule dry weight under drought conditions, relative to well-watered plants. It was concluded that large variation in nodulation sensitivity to water deficit exists among soybean cultivars and that the response of N₂ fixation rates to drought is related in part to nodule formation and growth.     

Ureide degradation pathways in intact soybean leaves

Ureides dramatically accumulate in shoots of N(2)-fixing soybean (Glycine max L. Merr.) under water deficit and this accumulation is higher in cultivars that have N(2) fixation that is sensitive to water deficit. One possible explanation is that ureide accumulation is associated with a feedback inhibition of nitrogenase activity. A critical factor involved in ureide accumulation is likely to be the rate of ureide degradation in the leaves. There exists, however, a controversy concerning the pathway of allantoic acid degradation in soybean. Allantoate amidinohydrolase was reported to be the pathway of degradation in studies using the cultivar Maple Arrow and allantoate amidohydrolase was the pathway that existed in the cultivar Williams. This investigation was undertaken to resolve the existence of these two pathways. An in situ technique was developed to examine the response of ureide degradation in leaf tissue to various treatments. In addition, the response of ureide accumulation and N(2) fixation activity was measured for intact plants in response to treatments that differentially influenced the two degradation pathways. The results from these studies confirmed that Maple Arrow and Williams degraded allantoic acid by different pathways as originally reported. The existence of two degradation pathways within the soybean germplasm opens the possibility of modifying ureide degradation to minimize the influence of soil water deficits on N(2) fixation activity.     

Drought survival,summer dormancy and dehydrin accumulation in contrasting cultivars of Dactylis glomerata

To study survival under prolonged and severe drought in the perennial grass Dactylis glomerata we compared dormant, resistant and sensitive cultivars (cvs.) in both field and glasshouse experiments. Water status, membrane stability and expression of dehydrins were assessed in the immature leaf bases, which are the last surviving organs. Analysis of leaf elongation and senescence of aerial tissues showed that dormancy was exhibited by the potentially dormant cultivar (cv.) only in the field. This cultivar exhibited a high survival rate, similar levels of dehydration and expression of a low-molecular weight (22–24 kDa) dehydrin in both drought and irrigated plants, whether fully dormant or not. At the same level of soil water deficit, there were no differences between the non-dormant drought resistant and drought sensitive cultivars in plant water status and membrane stability. However, the accumulation of dehydrins as drought progressed was markedly different between these cultivars and was associated with their contrasting survival. The possible role of the major low-molecular dehydrins in maintenance of cell integrity under dehydration is discussed with reference to both summer dormancy and survival under severe drought.     

Expression of symbiotic genes of Rhizobium japonicum USDA 191 in other rhizobia.

A 200-megadalton plasmid was mobilized from Rhizobium japonicum USDA 191 to other Rhizobium strains either that cannot nodulate soybeans or that form Fix- nodules on certain cultivars. The symbiotic properties of the transconjugants indicate that both soybean specificity for nodulation and cultivar specificity for nitrogen fixation are plasmid encoded.     

NolX of Sinorhizobium fredii USDA257, a Type III-Secreted Protein Involved in Host Range Determination, Is Localized in the Infection Threads of Cowpea (Vigna unguiculata [L.] Walp) and Soybean (Glycine max [L.] Merr.) Nodules

Sinorhizobium fredii USDA257 forms nitrogen-fixing nodules on soybean (Glycine max [L.] Merr.) in a cultivar-specific manner. This strain forms nodules on primitive soybean cultivars but fails to nodulate agronomically improved North American cultivars. Soybean cultivar specificity is regulated by the nolXWBTUV locus, which encodes part of a type III secretion system (TTSS). NolX, a soybean cultivar specificity protein, is secreted by TTSS and shows homology to HrpF of the plant pathogen Xanthomonas campestris pv. vesicatoria. It is not known whether NolX functions at the bacterium-plant interface or acts inside the host cell. Antibodies raised against S. fredii USDA257 NolX were used in immunocytochemical studies to investigate the subcellular localization of this protein. Immunostaining of paraffin-embedded sections of developing soybean and cowpea (Vigna unguiculata [L.] Walp) nodules revealed localization of NolX in the infection threads. Protein A-gold immunocytochemical localization studies utilizing affinity-purified NolX antibodies revealed specific deposition of gold particles in the fibrillar material inside infection threads. Similar immunogold localization studies failed to detect NolX in thin sections of mature soybean and cowpea nodules. The results from this study indicate that NolX is expressed in planta only during the early stages of nodule development.     

Effect of water deficit on carbohydrate status and enzymes of carbohydrate metabolism in seedlings of wheat cultivars

The effect of water deficit on carbohydrate status and enzymes of carbohydrate metabolism (alpha and beta amylases, sucrose phosphate synthase, sucrose synthase, acid and alkaline invertases) in wheat (Triticum aestivum L.) was investigated in the seedlings of drought-sensitive (PBW 343) and drought-tolerant (C 306) cultivars. The water deficit was induced by adding 6% mannitol (water potential -0.815 Mpa) in the growth medium. The water deficit reduced starch content in the shoots of tolerant seedlings as compared to the sensitive ones, but increased sucrose content in the shoots and roots of tolerant seedlings, indicating their protective role during stress conditions. It also decreased the alpha-amylase activity in the endosperm of seedlings of both the cultivars, but increased alpha and beta amylase activities in the shoots of tolerant ones. Sucrose phosphate synthase (SPS) activity showed a significant increase at 6 days of seedling growth (DSG) in the shoots of stressed seedlings of tolerant cultivar. However, SPS activity in the roots of stressed seedlings of sensitive cultivar was very low at 4 DSG and appeared significantly only at day 6. Sucrose synthase (SS) activity was lower in the shoots and roots of stressed seedlings of tolerant cultivar than sensitive ones at early stage of seedling growth. Higher acid invertase activity in the shoots of seedlings of tolerant cultivar appeared to be a unique characteristic of this cultivar for stress tolerance. Alkaline invertase activity, although affected under water deficit conditions, but was too low as compared to acid invertase activity to cause any significant affect on sucrose hydrolysis. In conclusion, higher sucrose content with high SPS and low acid invertase and SS activities in the roots under water deficit conditions could be responsible for drought tolerance of C 306.     

The influence of ozone on the stomatal and non-stomatal limitation of photosynthesis in Norway spruce,Picea abies (L.) Karst,exposed to soil moisture deficit

Summary Norway spruce, Picea abies (L.) Karst., was exposed to charcoal-filtered air (CF) and non-filtered air + ozone (NF+) and periods of soil moisture deficit from 1985 to 1988 in open-top chambers. Net photosynthesis, stomatal conductance, needle water potential and various shoot properties were measured on 1-year-old shoots during a period of soil moisture deficit. The gas exchange was measured at saturating photosynthetic photon flux density and across a range of CO₂ concentrations. The soil moisture deficit induced a mild drought stress in the plants, expressed by a pre-dawn needle water potential of approximately-0.9 MPa and a substantial reduction in net photosynthesis and gas phase conductance. In the CF treatment, intercellular CO₂ concentration was reduced, but was unaffected in the NF+ treatment. Furthermore, net photosynthesis declined more in response to the soil moisture deficit in the NF+ treatment than in the CF treatment. This is suggested to be attributed to the carboxylation efficiency at the operating point, which was decreased by 47% and 64% in shoots from the CF and the NF+ treatments, respectively. Stomatal limitation of net photosynthesis was increased by drought by 24–45% in the CF treatment, while it was unaffected in the NF+ treatment. Thus, our results imply that the coupling between the stomatal conductance and the photosynthetic rate was changed and that the marginal cost of water per given amount of carbon gain will increase in trees exposed to ozone, during periods of drought.     

不同水氮水平下小麦品种对光、水和氮利用效率的权衡

通过再裂区设计田间试验,以3个春小麦品种(和尚头、西旱2号和宁春4号)为材料,设置两个灌溉水平(充分灌水4500 m³·hm^-2和有限灌水3000 m³·hm^-2)和5个施氮水平(0、75、150、225、300 kg N·hm^-2),研究小麦光能利用效率(LUE)、水分利用效率(WUE)、氮素利用效率(NUE)对水氮的响应特性及其相互关系.结果表明: 3个小麦品种间LUE、WUE和NUE差异显著.在一定范围内增加灌水和施氮量则LUE升高,过量施氮则LUE下降.强抗旱和中等抗旱品种(和尚头和西旱2号)WUE受灌水量的影响比不抗旱品种(宁春4号)小.施氮可以调节小麦WUE,中等施氮水平(和尚头和西旱2号在150 kg N·hm^-2时,宁春4号在225 kg N·hm^-2时)有最高的WUE.随施氮量增加,植株氮素累积量先增后减,氮素干物质生产效率(NUE_b)、氮素收获指数(NHI)、氮肥农学利用效率(NAE)和氮肥偏生产力(PFP)均显著降低.灌溉水平对NHI无显著影响;随灌水量增加,小麦氮素积累量显著增加,强抗旱和中等抗旱品种NUE_b和NAE显著降低,不抗旱品种 NUE_b和PFP显著升高,对其他指标无显著影响.3个小麦品种氮素获取能力与氮素利用效率呈极显著负相关,NUE_b与LUE、WUE呈显著负相关,LUE与WUE呈显著正相关,春小麦氮素利用效率与光能利用效率、水分利用效率间存在明显的权衡关系.当灌水量为3000 m³·hm^-2,强抗旱和中等抗旱品种在150 kg N·hm^-2,不抗旱品种在225 kg N·hm^-2时,有较高的资源利用效率.     

Comparative study of nitrogen fixation and carbon metabolism in two chick-pea (Cicer arietinum L.) cultivars under salt stress

Two cultivars of Cicer arietinum with differential tolerance to salinity have been compared by analysing growth, photosynthesis, nodulation, nitrogenase activity, and carbon metabolism in the nodule cytosol. The aim was to help elucidate the relationships between, on the one hand, sucrose and malate metabolism in nodules and, on the other, the inhibition of nitrogen fixation under salt stress. Chick-pea cultivars Pedrosillano (sensitive) and ILC1919 (tolerant) inoculated with Mesorhizobium ciceri strain Ch-191 were grown in a controlled environmental chamber and were treated with salt (0, 50, 75, and 100 mM NaCl) from sowing to harvest time (28 d). Plant growth and photosynthesis were more affected by salt in Pedrosillano than in ILC1919. Also the effect of salt on nodulation and nitrogen fixation was much more pronounced in Pedrosillano. The increase in nodular mass in ILC1919 can partially counteract the inhibition of nitrogenase activity. The enzymes of sucrose breakdown were inhibited by NaCl, but in ILC1919 a rise in alkaline invertase was observed with salinity, which could compensate for the lack of the sucrose synthase hydrolytic activity. The activity of PEPC was stimulated by salt in ILC1919. Also, this cultivar showed higher malate concentrations in root nodules.     

Soybean nodulation and N2 fixation response to drought under carbon dioxide enrichment

The combined effects of carbon dioxide (CO₂) enrichment and water deficits on nodulation and N₂ fixation were analysed in soybean [Glycine max (L.) Merr.]. Two short-term experiments were conducted in greenhouses with plants subjected to soil drying, while exposed to CO₂ atmospheres of either 360 or 700 μmol CO₂ mol^–1. Under drought-stressed conditions, elevated [CO₂] resulted in a delay in the decrease in N₂ fixation rates associated with drying of the soil used in these experiments. The elevated [CO₂] also allowed the plants under drought to sustain significant increases in nodule number and mass relative to those under ambient [CO₂]. The total non-structural carbohydrate (TNC) concentration was lower in the shoots of the plants exposed to drought; however, plants under elevated CO₂ had much higher TNC levels than those under ambient CO₂. For both [CO₂] treatments, drought stress induced a substantial accumulation of TNC in the nodules that paralleled N₂ fixation decline, which indicates that nodule activity under drought may not be carbon limited. Under drought stress, ureide concentration increased in all plant tissues. However, exposure to elevated [CO₂] resulted in substantially less drought-induced ureide accumulation in leaf and petiole tissues. A strong negative correlation was found between ureide accumulation and TNC levels in the leaves. This relationship, together with the large effect of elevated [CO₂] on the decrease of ureide accumulation in the leaves, indicated the importance of ureide breakdown in the response of N₂ fixation to drought and of feedback inhibition by ureides on nodule activity. It is concluded that an important effect of CO₂ enrichment on soybean under drought conditions is an enhancement of photoassimilation, an increased partitioning of carbon to nodules and a decrease of leaf ureide levels, which is associated with sustained nodule growth and N₂ rates under soil water deficits. We suggest that future [CO₂] increases are likely to benefit soybean production by increasing the drought tolerance of N₂ fixation.     

A Comparison of Soybean Agglutinin in Cultivars Resistant and Susceptible to Phytophthora megasperma var. sojae (Race 1)

The amount of soybean agglutinin (SBA) detectable by radioimmunoassay in seeds of resistant cultivars to Phytophthora megasperma var. sojae was approximately twice that of susceptible cultivars. SBA was preferentially released at earlier times (6-9 hours) and in higher amounts in the imbibate from resistant cultivars as compared to susceptible cultivars. The lectin in the imbibate was immunologically identical to the seed lectin, indicating little or no proteolysis had occurred, and was active in hemagglutination. Binding of fluorescein isothiocyanate-labeled SBA to mycelial cell walls could be abolished by adding N-acetyl galactosamine or galactose. Purified SBA at concentrations of 150 to 300 micrograms inhibited mycelial growth by 50%, and the imbibate from Govan (resistant) cultivar was more inhibitory than the imbibate from Shore (susceptible) cultivar. Removal of SBA from the imbibate by affinity chromatography abolished the inhibition of mycelial growth, but the inhibition could be recovered from the eluant containing lectin.     

干旱胁迫对大豆鼓粒期叶片光合能力和根系生长的影响

以晋大70(抗旱型)和晋豆26(敏感型)2个大豆品种为材料,采用盆栽方法,在鼓粒期设置充分供水、轻度干旱和重度干旱3种水分处理,研究了干旱胁迫对大豆鼓粒期叶片光合能力和根系生长的影响.结果表明： 随着干旱程度加剧,2个品种的叶面积、叶绿素含量、净光合速率、叶片气孔导度、蒸腾速率、胞间CO₂浓度、株质量、株高、籽粒产量和收获指数均降低;根长和根质量在轻度干旱胁迫下增加,在重度干旱胁迫下减少;根冠比随干旱程度加剧而升高.重度干旱胁迫下,抗旱型品种晋大70根冠比的增幅达到135.7%,大于敏感型品种晋豆26根冠比的增幅(116.7%),晋大70的叶面积和叶绿素含量分别为对照的69.3%和85.5%,均优于晋豆26,晋大70的气孔导度和净光合速率分别下降了67.9%和77.9%,降幅均小于晋豆26,晋大70的收获指数的降幅为43.8%,小于晋豆26 (78.8%).不同干旱处理下,2个品种的叶面积、叶绿素含量、净光合速率、气孔导度、蒸腾速率、胞间CO₂浓度两两之间均呈显著正相关;株质量、株高、根长、根质量、籽粒产量、收获指数两两之间均呈显著正相关;根冠比与根质量呈显著正相关,而与其他5个指标呈显著负相关.     

The accumulation of trehalose in nodules of several cultivars of common bean (Phaseolus vulgaris) and its correlation with resistance to drought stress

Nine cultivars of common bean were grown in the presence of a natural microflora without exogenous rhizobial inoculation. Nodules were harvested 30 days post planting (early flowering stage) and the presence of trehalose determined. Amounts present varied according to cultivar and were between 0.20 and 1.63 mg g⁻¹ nodule dry weight. Rhizobial strains were isolated from the nodules of three selected cultivars (Canario 101, Flor de Mayo Bajio and Flor de Mayo 38). Trehalose levels in nodules produced after either mixed strain reinfection, or after axenic homologous reinfection or after axenic cross‐reinfection could be manipulated by applying drought stress. Mixed reinfection nodules from stressed plants accumulated between two and six times the trehalose concentration found in non‐stressed control plants. After axenic cross‐reinfection up to 48‐fold increases in nodule trehalose content were recorded during drought stress. Those cultivars exhibiting high nodule trehalose levels and/or a high degree of trehalose stimulation in response to drought stress also exhibited a high leaf relative water content and were also the most drought resistant. During drought stress nodule trehalase levels rose only slightly.