Effects of N-serve on soybeans and soil nitrogen transformations

Summary Plant dry weight, total N, and total Ca was increased at 0.1 and 1 ppm N-serve. At greater 10 ppm the plants showed visual symptoms of a stunted growth, stem elongation, flowers, and pods failed to form or were aborted, young leaves were curled, and roots were club shaped with many branches. These symptoms were increasingly evident with increasing N-serve application rates. The reason was attributed to an auxin effect. Dry wt and total N in the plant was less than the control at the higher N-serve applications. There was little effect on nitrogenase activity at less than 10 ppm N-serve. Nodulation tended to increase at 0.1 and 1 ppm N-serve.Nitrification was inhibited up to 104 days at 20 ppm N-serve. The soil pH of the high N-serve rates was decreased at 104 days probably due to nitrification. Generally there were little detectable differences among treatments in soil organic N. The average soil organic N from 0 to 104 days decreased by 0.01%. Average increase in total N within each pot at harvest was equivalent to about 138 kg N/ha.     

A non-destructive field assay for soybean nitrogen fixation by acetylene reduction

Summary A system for employing open-ended root chambers to measurein situ acetylene reduction rates under field conditions is described. Gas mixtures containing about 2 mbar acetylene were continuously flowed through the chambers providing a continuous record of acetylene reduction. These chambers have been used to measure acetylene reduction rates of soybeans during three growing seasons. The system has proved to be reliable with a high degree of precision. The large amount of plant-to-plant variability observed in N₂ fixation research has been confirmed by the data collected with this system. However, such variability in physiological studies can be reduced by using a non-destructive system to compare the response of an individual plant with its rates before treatment.     

Drought tolerance processes in soybeans

Fifteen soybean cultivars were evaluated in two water supply conditions, inducing or not a drought stress. Main canopy traits were measured several times during the reproductive period and, at maturity date, the yield components were estimated. Using principal components analysis, the main physiological functions involved in soybean drought tolerance are described: leaf cells enlargement and assimilates transport. These processes could be a good basis on which to define new selection criteria for soybean drought tolerance.     

Effect of plant genotype and nitrogen fertilizer on symbiotic nitrogen fixation by soybean cultivars

Summary Isotopic as well as non-isotopic methods were used to assess symbiotic nitrogen fixation within eight soybean [Glycine max (L.) Merr.] cultivars grown at 20 and 100 kg N/ha levels of nitrogen fertilizer under field conditions.The¹⁵N methodology revealed large differences between soybean cultivars in their abilities to support nitrogen fixation. In almost all cases, the application of 100 kg N/ha resulted in lower N₂ fixed in soybean than at 20 kg N/ha in the first year of the study. However, N₂ fixed in one cultivar, Dunadja, was not significantly affected by the higher rate of N fertilizer application. These results were confirmed by measurements of acetylene reduction activity, nodule dry weight and N₂ fixed as measured by the difference method. Further proof of differences in N₂ fixed within soybean cultivars and the ability of Dunadja to fix similar amounts of N₂ at 20 and 100 kg N/ha was obtained during a second year experiment. Dunadja yield was affected by N fertilizer and produced larger yield at 100 kg N/ha than at 20 kg N/ha. This type of cultivar could be particularly useful in situations where soil N levels are high or where there is need to apply high amounts of N fertilizer.The present study reveals the great variability between legume germplasms in the ability to fix N₂ at different inorganic N levels, and also the potential that exists in breeding for nitrogen fixation associative traits. The¹⁵N methodology offers a unique tool to evaluate germplasms directly in the field for their N₂ fixation abilities at different N fertilizer levels.     

Nitrogen accumulation and redistribution in soybean genotypes with variation in seed protein concentration

Breeding for high seed protein concentration in soybean [Glycine max (L.) Merrill] often results in lower yield, but the basis for this negative relationship is not well understood. To address this question, we evaluated the N acquisition characteristics of three high protein and three normal soybean genotypes in the field for 3 years. Plants were grown in 0.76 m rows following conventional cultural practices and water stress was minimized with sprinkler irrigation. We determined the mass and N concentration of leaves, petioles and stems at the beginning of seed filling (growth stage R5) and of stems at maturity. The N concentration of abscised leaves and petioles was also determined. There was significant variation among genotypes in total seed N (g m⁻²) at maturity (range from 14.7 to 24.4 g N m⁻²) as a result of variation in seed N concentration and yield. There was no evidence that the larger amounts of mature seed N were associated with a larger vegetative N reservoir at growth stage R5 as determined by vegetative mass at R5 or the concentration of N in vegetative tissues. Increasing seed N at maturity did not lower the N concentration in abscised leaves and petioles, or in the stems at maturity. The rate and timing of leaf senescence (loss of chlorophyll) was essentially the same for all genotypes. With no increase in the contribution from redistributed N, increases in N uptake or fixation during seed filling must have been responsible for the higher levels of seed N at maturity in high-protein genotypes. These data suggest that increasing total seed N at maturity by selecting for higher seed protein concentration or higher yield in soybean does not require, as some models suggest, a larger vegetative N reservoir at the beginning of seed filling or more rapid senescence.     

Phosphorus fertilization and tillage effect on dinitrogen fixation in soybeans

Summary Tillage has been shown to affect the uptake of phosphorus (P) and yield of soybeans, [Glycine max (L.) Merr.], but there is little information concerning the effects of P fertilization on nitrogen (N₂) fixation in soybeans under no-tillage. Two field experiments were conducted in 1980 and 1981 to determine the effects of soil P on N₂ fixation under no-tillage and to study the interaction of P fertilization and tillage of N₂ fixation, nutrient uptake, and yield of soybeans. In Exp. I, P was applied in 1977 at five rates up to 384 kg P ha⁻¹ and the effects of residual soil P were evaluated in 1980 and 1981 under no-tillage management. Nitrogen fixation rates, as measured by acetylene reduction assay, were significantly affected by soil P in Exp. I, but the assay proved to be a poor technique for estimating total plant N in these tests. Acetylene reduction rates and plant P increased rapidly as soil P increased from 2 to 20 mg kg⁻¹, with little additional increase above 20 mg P kg⁻¹. In Exp. II, rates (0, 32, 64, and 128 kg P ha⁻¹) and time (fall, spring and fall plus spring) of P application were compared under conventional tillage and no tillage. However, plant P increased with increasing levels of applied P. Applied P had no affect on acetylene reduction rates but rates were greater for no-tillage than conventional tillage at the V9 and R5 stages of growth in 1981. Plant uptake of P was more efficient under no-tillage than under conventional tillage in 1980 and 1981. Application of 64 kg P ha⁻¹ under no-tillage resulted in equivalent plant P levels as the 128 kg P ha⁻¹ applied under conventional tillage.     

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.     

Dry matter and nitrogen accumulation and partitioning in selected soybean genotypes of different derivation

Summary Experiments were conducted to determine if changes in the accumulation and partitioning of dry matter (DM) and nitrogen (N) in soybean [Glycine max (L.) Merr.] were associated with agronomic improvements and to assess the degree of genetic variation present for these traits. Fifteen maturity group II soybean genotypes including three ancestral cultivars, three modern cultivars, and nine agronomically superior plant introductions (PI's) were grown in replicated tests at four locations in the eastern U.S. The DM and N of stems, pod walls, and seeds were determined at maturity, and the apparent harvest indices (HI) and the apparent nitrogen harvest indices (NHI) were calculated. Pod DM partitioning was calculated as the ratio of seed DM to total pod DM and pod N partitioning was the ratio of seed N to total pod N. The mean DM accumulation of the modern cultivars was significantly greater than that of the ancestral cultivars and PI's. The apparent HI and the pod DM partitioning of both the modern and ancestral cultivars were significantly higher than that of the PI's. The three modern cultivars demonstrated the highest N accumulation. As a group, the modern cultivars consistently showed maximal accumulation and partitioning of DM and N suggesting that these physiological traits are associated with agronomic improvement. No individual PI was found to possess DM or N accumulation or partitioning which significantly exceeded the best modern cultivar or ancestral cultivar, indicating that genotypes with accumulation or partitioning characteristics which exceed available germplasm may be difficult to identify. Seed yield was correlated (P<0.05) with both DM (r=0.61) and N (r=0.57) accumulation.     

Waterlogging effect on xylem sap glutamine of nodulated soybean

Waterlogging of soybean plants (Glycine max L.) led to impaired symbiotic N₂ fixation and a marked decline in glutamine (Gln) concentration in xylem bleeding sap. Xylem Gln concentration increased during the growth cycle of the plant and was correlated with nodule formation. Treatments known to impair N₂ fixation, such as exposing the root system to pure N₂ gas or a mixture of Ar and O₂ (80:20; v/v), led to specific declines in xylem sap Gln. The decrease in Gln observed during waterlogging was also seen on transfer of nodulated plants to aerated hydroponics, where the decline was highly correlated with ureide content in the xylem sap. Upon flooding the nodulated root system, the specific decline in xylem sap Gln could be detected within 10 min and reached a minimum within 60 min, indicating that waterlogging has an immediate effect on N₂ fixation. It is concluded that xylem Gln arises directly from N₂-fixation and is a useful indicator of N₂ fixation activity of symbiotic soybean plants.     

Drought-avoidant soybean germplasm maintains nitrogen-fixation capacity under water stress

Inoculated soybeans (Glycine max L. (Merrill)) were grown in controlled environments to evaluate the relationship between genotype and plant water status on nodule function, nitrogen assimilation, growth rates, and seed yield. Plants were grown under well-watered (WW) and water-stressed (WS) conditions during the linear pod-filling growth stage in sand culture using N-free nutrient solution. Dry matter and N accumulation were greater for the drought-adapted Plant Introduction 416937 (PI) than for Forrest, a commercially adapted genotype of similar phenology. These differences are attributed to: (i) more favorable internal water balance throughout the pod-filling period (higher total leaf water potential), (ii) higher photosynthetic function (more total leaf area and higher net carbon exchange rates), and (iii) stronger nodule function (larger nodule mass, greater specific and total nodule activity, and thus more nitrogen assimilation) for the PI than for Forrest. While Forrest out yielded the PI under WW conditions, the percentage reduction in seed mass per plant was less for the PI than for Forrest when both genotypes were exposed to desiccating conditions. The inference is that soybean germplasm with the capacity to maintain tissue turgidity, and thus leaf and nodule function, during reproductively-imposed desiccation may reduce the extent to which yield is compromised during drought. These findings have implications for the role of symbiotic nitrogen fixation in conserving yield under dry weather conditions.Abbreviations DAE Days After Emergence - NCE Net CO₂ Exchange - PI PI 416937 - SNA Specific Nodule Activity - TNA Total Nodule Activity - WS Water Stressed - WW Well Watered     

Soil temperature effects on competitiveness and growth ofRhizobium japonicum and on Rhizobium-induced chlorosis of soybeans

Summary The effects of temperature on growth in broth and soil and on competition for nodule formation betweenRhizobium japonicum serotypes USDA 76 and 94 compared to 6 and 110 were studied. Increasing root temperatures of Lee soybean from 20 to 35°C increased the competitiveness of 76 and 94 relative to 6 and 110 for all inoculum ratios such that at 30 and 35°C symptoms ofRhizobium-induced chlorosis appeared. Tolerance to elevated temperatures was exhibited by 76 and 110, but not 94 and 6 in broth and soil which suggested that increased competitiveness of 76 and 94 at high soil temperatures was not dependent upon growth at elevated temperatures. Nodulation and vegetative growth of Lee soybeans were at a minimum at 20°C and optimum at 30°C. Differences in competitiveness of 6 to previous studies indicated the need to standardize temperatures of assays. Differences in growth responses of 76 and 94 to temperature from a previous study suggested a confounding effect on different carbon sources in growth media. Scientific Article No. A-3721 Contribution No. 6697 of the Maryland Agric Exp Sta, Dept of Agronomy, College Park, MD 20742 and the USDA, ARS, Beltsville, MD 20705. Part of a thesis submitted by the senior author in partial fulfillment of the requirements for the M.S. Degree.     

Trifluralin-induced chlorosis in soybeans (Glycine max (L.) Merr.) grown on clayey,high pH soils

Summary The cause of leaf chlorosis, frequently observed on soybeans (Glycine max (L.) Merr.) grown on high pH soils of the Mississippi Blackland Prairie, is thought to be low Fe availability and restricted rooting. Three greenhouse experiments were conducted using two soils, Sumter, a Rendollic Eutrocrept and Okolona, a Typic Chromudert; nine soybean cultivars differing in Feefficiency; and trifluralin (α-α-α-trifluoro-2,6-dinitro-N, N-di-propyl-p-toludine). Trifluralin at rates greater than 0.56 kg/ha caused chlorosis which was more severe on the Sumter, a soil low in available Fe. Fe-efficient cultivars were more resistant to the chlorosis induced by trifluralin than the Fe-inefficient cultivars. It was concluded that the chlorosis is an Fe deficiency caused by reduced uptake. The herbicide-induced chlorosis can be avoided by proper dosage and placement of the herbicide.     

Nodulation and nitrogen fixation by fast- and slow-growing rhizobia strains of soybean on several temperate and tropical legumes

Three slow-growingBradyrhizobium japonicum (G₃, USDA-110 and KUL-150) of diverse origins and two fast-growing strains ofRhizobium fredii (USDA-192 and USDA-193) were tested with a cropped soybean (Glycine max L. Merrill) cultivar, two cowpeas (Vigna unguiculata), one mung-bean (Phaseolus radiata), one winged-bean (Psophocarpus tetragonolobus) and one field bean (Phaseolus vulgaris) varieties.TheR. fredii strains nodulated and fixed Nitrogen as effectively as the strains ofB. japonicum in a modern european soybean cultivar, namely Fiskeby V. The other western bred soybeans tested were not nodulated by theseR. fredii strains. All of the soybean rhizobia produced nodules in both cowpeas and in mung-bean; theR. fredii strains showed effective N₂-fixation in the cowpeas, particularly USDA-193, yielding shoot dry weights greater than those from theB. japonicum. The symbiotic performance of theR. fredii strains with soybean and other legumes indicated that they should be placed in an intermediate group between the slow-growingB. japonicum and cowpearhizobium sp.The hydrogen uptake activites suggested a possible host effect on the expression of such genes in one out of theB. japonicum strains tested. Furthermore, the slow-growing rhizobia showed significantly higher nitrate-reduction than theR. fredii in the nodules.     

Seasonal accumulation and partitioning of nitrogen by lentil

Although wheat (Triticum aestivum L.) is the dominant crop of the semi-arid plains of Canada and the western United States, lentil (Lens culinaris Medik.) has become an important alternative crop. Sources and seasonal accumulation of N must be understood in order to identify parameters that can lead to increased N₂-fixing activity and yield. Inoculated lentil was grown in a sandy-loam soil at an irrigated site in Saskatchewan, Canada. Wheat was used as the reference crop to estimate N₂ fixation by the A-value approach. Lentil and wheat received 10 and 100 kg N ha⁻¹ of ammonium nitrate, respectively. Crops were harvested six times during the growing season and plant components analyzed. During the first 71 days after planting the wheat had a higher daily dry matter and N accumulation compared to lentil. However, during the latter part of the growing season, daily dry matter and N accumulation were greater for lentil. The maximum total N accumulation for lentil at maturity was 149 kg ha⁻¹. In contrast, wheat had a maximum N accumulation of 98 kg ha⁻¹ in the Feekes 11.1 stage, or 86 days after planting. The maximum daily rates of N accumulation were 3.82 kg N ha⁻¹ day⁻¹ for lentil and 2.21 kg N ha⁻¹ day⁻¹ for wheat. The percentage of N derived from N₂ fixation (% Ndfa) ranged from 0 at the first harvest to 92 % at final harvest. Generative plant components had higher values for % Ndfa than the vegetative components which indicates that N in the reproductive plant parts was derived largely from current N₂ fixation and lentil continued to fix N until the end of the pod fill stage. At final harvest, lentil had derived 129 kg N ha⁻¹ from N₂ fixation with maximum N₂-fixing activity (4.4 kg N ha⁻¹ day⁻¹) occurring during the early stages of pod fill. Higher maximum rates of N₂-fixing activity than net N accumulation (3.82 kg N ha⁻¹ day⁻¹) may have been caused by N losses like volatilization. In addition, lentil provided a net N contribution to the soil of 59 kg ha⁻¹ following the removal of the grain.     

The inheritance of a urease-null trait in soybeans

Summary Four soybean seed urease nulls (lacking both the activity and antigen of the embryo-specific urease) were intermated and the F₁ and F₂ seed examined for urease activity. Both generations were without urease activity, and the nulls were therefore considered noncomplementing. In crosses of each null line to cultivars homozygous for the allelic, codominantly inherited urease slow or fast isozyme, the F₁ seed expressed the embryo-specific urease isozyme of the urease-expressing parent. A 3 1 segregation for presence and absence of urease was observed in progeny from F₁ and heterozygous F₂ plants. The F₂ and F₃ from fastXnull combinations revealed that urease-positive seed were all phenotypically urease fast, while the same seed from slowXnull combinations showed a segregation of one seed containing a fast urease, either exclusively or in a heterozygous state with the slow isozyme, for every 69 phenotypic slows. Data pooled from F₂ plants which segregate for both the presence (Sun) and absence (Sun) of urease and for the fast (Eu1-b) or slow (Eu1-a) urease allele indicate that the null lesion (Sun) is linked to Eu1 by approximately one map unit. The evidence is consistent with two models: (1) sun is an allele at the embryo-specific urease isozyme locus (Eu1) and that a high degree of exchange (and/or conversion) within the locus results in a 1% recombination frequency between the null trait and urease allozyme; (2) sun is at a distinct locus which is separated by one map unit from the embryo-specific urease isozyme locus (Eu1) upon which it acts in the cis position. Polyadenylated embryo RNA from one of the null lines, PI 229324, exhibited no urease template activity in vitro. Thus, the lack of urease antigen is due to lack of accumulation of translatable urease mRNA. The availability of soybeans lacking seed urease should be extremely useful to breeders as a trait for linkage studies and to geneticists as a transformation marker.Portions of this work were funded by the Illinois and Missouri Agricultural Experiment Stations, the SOHIO-University of Illinois Center of Excellence in Crop Molecular Genetics and Genetic Engineering and by grants PCM-8219652 from the National Science Foundation and USDA/SEA-CRCR-1-1374 from the USDA Competitive Grants Office     

The role of active Bradyrhizobium japonicum in iron stress response of soybeans

The objective of this study was to identify the sites of H-ion exudation and Fe(III) reduction along both inoculated and non-inoculated roots of A7 and T203 soybeans. A split-root system was used in which half the roots of each plant were inoculated and actively fixing nitrogen and the other half were not. Expectedly, the Fe-stress response was strong on both sides of the split-root system in the +N-Fe treatment of variety A7 (inactive nodules) but not of variety T203. The Fe-stress response of A7 was enhanced by the presence of active nodules. Variety T203 is Fe inefficient and normally fails to produce any Fe-stress response, but in the absence of nitrogen and iron (–N–Fe), inoculated roots responded to Fe stress with exudation of both H-ions and reductants. Intact split-root systems were embedded in agar to determine the location of H-ion exudation and Fe(III) reduction. On the inoculated side of the –N–Fe and –N+Fe treatments (active nodules) of both soybean varieties, H-ion production was associated mainly with the active nodules. However, quantities of H-ion release were much greater under Fe stress (–N–Fe) than with adequate Fe (–N+Fe). Reduction of Fe(III) to Fe(II) was found only on the nodulated side with T203, but on both sides with A7. In variety T203 the Fe reduction was associated with younger roots located just below the nodule clusters on the inoculated side of the –N treatments. Active nodules appear to play a key role in the Fe-deficiency stress response of T203 soybean.     

Identification and characterization of common bean (Phaseolus vulgaris L.) lines well nodulated in the presence of high nitrate

Common bean,Phaseolus vulgaris L., is known to be ‘inefficient’ in nodulation and N₂ fixation although it responds to applied nitrogen. An experiment was conducted to identify and to characterize bean cultivars nodulating in the presence of a high level of nitrogen. Sixteen cultivars and a check for inefficient nodulation, OAC Seaforth, were inoculated and grown for 40 days in replicated pots supplied with zero, 3.5 and 10.5 mM combined nitrogen as NO ₃ ⁻ and NH ₄ ⁺ . Seven traits relating to nodulation and N₂ fixation were all significantly affected by N level (N), cultivar (Cv) and N × Cv interactions (except for root dry weight), indicating that cultivars responded differently to the N treatments. Total dry weight (W) and shoot to root ratio (S/R) increased with the increased N levels. Nodule dry weight (Wn), visual nodulation score (Nv) and nodulation index (Nx) decreased as the N increased. Percent N and N content per plant increased with the increased N level. Plant weight (W) was positively correlated with Wn, Nv and N content and negatively correlated with %N. Nodulation score was positively associated with Wn and plant N content. Genotypes superior in nodulation and N₂ fixation in the presence of N were identified. Cultivars Italian Barlotti, California Light Red Kidney, Kentucky Wonder A and Pueblo 152 were selected for further testing and use in improving the nitrate tolerant nodulating characteristic of bean.     

Effect of vesicular-arbuscular mycorrhizae on the growth and yield of rice-stubble cultured soybeans

Summary Inoculation with vesicular-arbuscular (VA) mycorrhizal fungiGlomus fasciculatus, G. mosseae, G. etunicatus orAcaulospora scrobiculatus, increased plant dry weight and seed yields of pot-grown soybean plants in sterilized soil. Inoculation with a mixture ofG. fasciculatus, G. mosseae andG. etunicatus, orG. fasciculatus alone, increased seed yields and other agronomic traits of soybean plants grown in a no-tillage, rice-stubble field.     

Effects of plant breeding and selection on yields and nitrogen fixation in soybeans under two soil nitrogen regimes

Summary Soybeans (Glycine max (L.) Merr.) have a high N requirement which is fulfilled by soil N uptake and N₂-fixation. This study was concerned with the effects of past yield selection on N₂-fixation in soybeans.The soybean cultivars, Lincoln, Shelby, and Williams, which represent successive improvements in the Lincoln germplasm, and a non-nodulating control were planted in a soil containing¹⁵N labelled organic matter. Two replications occurred on soil previously cropped to alfalfa and two on soil previously cropped to soybeans. Plants were harvested at five growth stages and leaf area, plant weight, total N, and atom percent¹⁵N were determined. Mature grain was harvested and yield components were also determined, as well as the total N and¹⁵N content.Cultivar differences in total dry matter were only evident at physiological maturity, when Williams contained the greatest dry matter. Williams exhibited the longest period of seed formation and seed fill and also had the highest grain yield which resulted from a larger weight per seed.The N content of the cultivars did not vary until physiological maturity when Williams contained the highest percent N. The quantity of N fixed at physiological maturity was highest for Williams and lowest for Lincoln. Fixed N contained in the harvested grain was greater for Williams than for the other two cultivars. The fraction of the total plant N derived from fixation was not greatly affected by cultivar and all cultivars acquired an average of 50% of their total N through N₂-fixation.Previous cropping history greatly affected the quantity of N fixed and the fraction of the total plant N derived from fixation. Soybeans following soybeans were more dependent upon N₂-fixation than soybeans following alfalfa with the former deriving 65% of the total plant N from fixation and the latter only 32%. These soybean cultivars apparently utilized soil N first and then used N₂-fixation to satisfy their N requirement.The past selection for higher yield has resulted in soybean cultivars with improved capacities to fix atmospheric N₂ and an improved ability to take up available soil N.