钙对镉胁迫下花生生理特性、产量和品质的影响水

通过盆栽试验,选用高油品种豫花15和高蛋白品种XB023,研究了不同浓度钙对镉胁迫下不同类型花生品种营养生长、叶片叶绿素含量、光合速率、保护酶活性等生理特性及产量和品质的影响.结果表明:施钙可以缓解镉胁迫对花生植株主茎高和侧枝长的抑制作用,增加花生植株干物质量,提高叶片叶绿素含量和光合速率,提高叶片超氧化物歧化酶( SOD)、过氧化氢酶(CAT)、过氧化物酶(POD)活性和可溶性蛋白质含量,降低丙二醛(MDA)的积累量,减轻镉胁迫对花生叶片的伤害;施钙可以缓解镉胁迫对花生的减产作用,增加花生荚果和籽仁产量,其增产的主要原因是增加了单株结荚数和出仁率;施钙可以促使籽仁中可溶性糖向粗脂肪和蛋白质转化,增加籽仁中脂肪和蛋白质含量,改善镉胁迫下花生籽仁品质.施钙可以降低两花生品种籽仁中镉含量,对豫花15的降低效果好于XB023.     

Growth and nitrogen fixation ofAzolla pinnata andAzolla caroliniana as affected by urea fertilizer and their influence on rice yield

Soils from avocado (Persea americana Mill.) orchards in Israel (IS) and California (CA), both sites with a Mediterranean climate, were sampled and analyzed for the species and quantities of vesicular-arbuscular mycorrhizal fungus (VAMF) spores in them, and for soil physical and chemical characteristics.Numbers of spores were similar in soil from IS and CA but the dominant VAMF species were very different. In IS the most common fungi were Sclerocystis sinuosa and Glomus macrocarpum. In CA, Gl. constrictum was present in every orchard examined and Gl. fasciculatum was nearly as widespread. Acaulospora spp. and other Glomus spp. also were found, including A. elegans which has never before been reported from CA.The differences in VAMF populations and species constituents found on two continents but in areas with similar climates and soil types may be due to host or edaphic factors. Different avocado rootstocks are used in the two countries and lower pH and higher soil fertility levels were present in CA soils.The total VAMF spore populations in each orchard was about 275 per 100 mL soil. The population level was not correlated with any of the soil physical or chemical characteristics examined nor with avocado cultivar or age. In IS no fungus spores were found in three orchards; available P, Ca, Mg and Cu levels were high in these soils.     

施钙对干旱胁迫下花生生理特性、产量和品质的影响

以花生品种606为试材,在旱棚池栽人工控水条件下,研究了钙肥不同用量对花针期和结荚期干旱胁迫下花生的营养生长、生理特性、产量及品质的影响.结果表明:干旱胁迫下施钙,可以促进花生的营养生长,提高叶片的叶绿素含量、净光合速率和根系活力,提高干旱后复水过程中花生的恢复能力,缓解干旱对花生的不利影响;增加了花生荚果和籽仁的产量,尤其是增加了单株结果数和出仁率.施钙提高了籽仁中的脂肪和蛋白质含量,改善了干旱胁迫下花生的籽仁品质.在本试验条件下,施钙量为300 kg·hm-2时效果最佳.     

Differences between cereal crop cultivars in root-associated nitrogen fixation,possible causes of variable yield response to seed inoculation

Large differences in N₂-ase activity with fractions of active plants from 3–67% and maximal activities from 3–35 nmol C₂H₄.h^-1 were found between sterile, Azospirillum-inoculated seedlings of 14 German cereal cultivars. Examples of similar cultivar differences in gnotobiotic or unsterile cereals in response to Azospirillum inoculation, in root exudation and the specificity of bacteria-root interactions are reviewed. As possible causes of yield responses to bacterial seed inoculation N₂-fixation, plant growth regulating metabolites and bacterial interaction with root pathogens are discussed. The need for suitable screening methods to select and breed cultivars with desirable responses to beneficial rhizosphere bacteria is pointed out.     

Modeling long-term crop response to fertilizer and soil nitrogen

A simple nitrogen balance model to calculate long-term changes in soil organic nitrogen, nitrogen uptake by the crop and recovery of applied nitrogen, is presented. It functions with time intervals of one year or one growing season. In the model a labile and a stable pool of soil organic nitrogen are distinguished. Transfer coefficients for the various inputs of nitrogen are established that specify the fractions taken up by the crop, lost from the system, and incorporated in soil organic nitrogen. It is shown how input data, model parameters and initial pool sizes can be derived and how the model can be used for calculating long-term changes in total soil organic nitrogen and uptake by the crop. For nitrogen applied annually as fertilizer or organic material the time course of nitrogen uptake and recovery of applied nitrogen is calculated. To test the sensitivity of the model, calculations have been performed for different environmental conditions with higher or lower risks for losses. The model has also been applied to establish fertilizer recommendations for a certain target nitrogen uptake by the crop. Finally, for agricultural systems where periods of cropping alternate with peroids of green fallow the time course of nitrogen uptake by the crop is calculated.     

Influence of lime on nitrogen fixation by tropical and temperate legumes

Summary A nitrogen deficient Oxisol which had been fertilized with P, K, Zn and Mo received CaCO₃ at rates which increased continuously from zero to 22 tons/ha. Liming produced a range of pH in the saturation paste from 4.7 to 7.1; a range of calcium in the saturation extract from 0.3 to 3 meq/l; and a significant decline in available manganese. Responses of 9 tropical and 7 temperate legumes were compared across the pH variable.For Stylosanthes spp. nodule numbers and weight and plant yield declined above pH 5.5. Arachis hypogaea and Vigna sinensis, whose yield increased by only 30%, showed no clear improvement in nodulation or nodule effectiveness (acetylene reduction rate per unit nodule weight).Increased nodule effectiveness could account for most of the growth increase in Dolichos axillaris and Glycine max var. Kahala. Both the number and effectiveness of nodules increased for Desmodium intortum, Glycine wightii, Medicago sativa, and Trifolium subterraneum. Nodule size and effectiveness increased for G. max var. Kanrich. Only in one species, Coronilla varia, could increase in nodule numbers alone account for the increased growth associated with liming, although increased numbers of nodules probably accounted for much of the response by Lotus corniculatus, and by Desmodium canum and D. intortum up to pH 5.3.Increased nodulation and plant N contents were consistent with the conclusion that for most species improved N₂-fixation was the cause of growth improvement associated with liming. However, percent N was high in leaves of Vigna and Phaseolus vulgaris at all lime levels. In Phaseolus, variation in nodulation and growth were unrelated. The growth improvement was associated with decline in leaf N, suggesting that something other than N nutrition was limiting.Journal Series No. 1957 of the Hawaii Agricultural Experiment Station.Department of Soils and Plant Nutrition, University of California, Davis.Department of Agronomy and Soil Science, University of Hawaii, Honolulu, Hawaii.     

Comparative effects of organic and inorganic nitrogen sources applied to a flooded soil on rice yield and availability of N

A pot experiment was conducted to study the effect of organic and inorganic nitrogen (N) sources on the yield and N uptake of rice from applied and native soil-N. The residual effect of these N sources on a succeeding wheat crop was also studied. Organic N was applied in the form of ¹⁵N-labelled Sesbania aculeata L., a legume, and inorganic N in the form of ¹⁵N-labelled ammonium sulphate. The two sources were applied to the soil separately or together at the time of transplanting rice. Recovery of N by rice from both the applied sources was quite low but both sources caused significant increases in biomass and N yield of rice. Maximum increase was recorded in soil treated with organic N. The residual value of the two materials as source of N for wheat was not significant; the wheat took up only a small fraction of the N initially applied. Loss of N occurred from both applied N sources, the losses being more from inorganic N. Both applied N sources caused a substantial increase in the availability of soil-N to rice and wheat; most of this increase was due to organic N and was attributed to the so-called ‘priming’ effect or ANI (added nitrogen interaction) of the applied material.     

不同品质类型花生品质性状及相关酶活性差异

在大田栽培条件下,以高蛋白花生品种KB008、高脂肪品种花17(H17)和高油酸/亚油酸(O/L)品种农大818(818)为试验材料,研究了3种类型花生品种籽仁中蛋白质、脂肪含量及与品质合成相关的碳、氮代谢酶活性差异.结果表明:KB008的蛋白质含量显著高于H17和818,而可溶性糖含量和O/L值显著低于其他两品种.KB008籽仁中氨基酸组分含量均高于其他两品种,特别是谷氨酸和赖氨酸含量显著高于后两者;油酸含量显著低于、而亚油酸含量显著高于其他两品种.3种类型花生在整个生育期中叶片的硝酸还原酶(NR)、谷氨酰胺合成酶(GS)、谷氨酸脱氢酶(GDH)、谷氨酸合成酶(GOGAT)和谷丙转氨酶(GPT)活性均以KB008最高,其次为H17.3种类型花生在结荚期的叶片磷酸烯醇式丙酮酸羧化酶(PEPCase)和1,5-二磷酸核酮糖羧化酶(RuBPCase)活性大小均表现为KB008 ＞H17 ＞818,说明较高的PEPCase和RuBPCase活性有利于蛋白质合成与积累.叶片中蔗糖合成酶(SS)活性大小表现为H17＞818＞KB008,KB008的磷酸蔗糖合成酶(SPS)活性显著低于其他两品种,而H17的SPS活性在花后60d时仍保持较高活性,说明较高的叶片SPS、SS活性有利于花生籽仁脂肪的形成.     

Estimation of biological nitrogen fixation associated withBrachiaria andPaspalum grasses using15N labelled organic matter and fertilizer

Summary Six pasture grasses,Paspalum notatum cv batatais,P. notatum cv pensacola,Brachiaria radicans, B. ruziziensis, B. decumbens andB. humidicola, were grown in concrete cylinders (60 cm diameter) in the field for 31 months. The soil was amended with either a single addition of¹⁵N labelled organic matter or frequent small (2 kg N. ha^–1) additions of¹⁵N enriched (NH₄)₂SO₄. In the labelled fertilizer treatment soil analysis revealed that there was a very drastic change in¹⁵N enrichment in plant-available nitrogen (NO ₃ ^– +NH ₄ ⁺ ) with depth. The different grass cultivars recovered different quantities of applied labelled N, and evidence was obtained to suggest that the roots exploited the soil to different depths thus obtaining different¹⁵N enrichments in soil derived N. This invalidated the application of the isotope dilution technique to estimate the contribution of nitrogen fixation to the grass cultivars in this treatment. In the labelled organic matter treatment the¹⁵N label in the plant-available N declined at a decreasing rate during the experiment until in the last 12 months the decrease was only from 0.274 to 0.222 atom % excess. There was little change in¹⁵N enrichment of available N with depth, hence it was concluded that although the grasses recovered different quantities of labelled N, they all obtained virtually the same¹⁵N enrichment in soil derived N. Data from the final harvests of this treatment indicated thatB. humidicola andB. decumbens obtained 30 and 40% respectively of their nitrogen from N₂ fixation amounting to an input of 30 and 45 kg N.ha^–1 year^–1 respectively.     

Growth, yield and grain quality of barley (Hordeum vulgare L.) in response to nitrogen uptake. II. Plant development and rate of germination

A nutrient culture system supporting stands of barley was used to examinethe effect of nitrogen supply on plant development and germination rate.Two contrasting cultivars of barley were grown for two seasons withnitrogen supplied at an optimal rate and at one-third that rate. Maturegrain was dissected from ears by grain position; stored until out ofdormancy, weighed, and the time from inhibition to germination estimatedfor different ear positions. A parallel set of samples was analysed forgrain nitrogen concentrations. A systematic increase in germination timewas observed from the bottom to the top of the ear. This effect was inaddition to and of the same order as the effect of grain size ongermination time. Although there was significant variation in nitrogenconcentration by grain position, these effects were smaller than thevariations in grain weight within the ear and were not related togermination rate. These results are discussed in relation to plantdevelopment and events at the onset of germination.Keywords:Barley, germination, grain size, ear position, nitrogen.     

Partitioning of nitrogen from biological fixation and fertilizer in Phaseolus vulgaris

Nitrogen uptake, distribution and remobilization in the vegetative and reproductive parts of the plant were studied in bean (Phaseolus vulgaris L.) cultivars Negro Argel and Rio Tibagi inoculated with either Rhizobium strain C05 or 127 K-17. Greenhouse grown plants were supplied with 2.5 mg N (plant)⁻¹ day⁻¹ as KNO₃ or K¹⁵NO₃ and the relative contribution to total plant nitrogen of mineral and symbiotically fixed nitrogen was determined. Control plants included those entirely dependent on fixed nitrogen as well as uninoculated plants supplied with 10 mg N (plant)⁻¹ day⁻¹. No differences were observed between inoculated treatments in total nitrate reductase activity and in the amount of mineral nitrogen absorbed, but there were considerable differences in the contribution of fixed nitrogen. Nitrogen fixation supplied from 58 to 72% of the total nitrogen assimilated during the bean growth cycle and the symbiotic combinations fixed most of their nitrogen (66 to 78% of total nitrogen) after flowering. Maximum uptake of mineral nitrogen was in the 15-day-period between flowering and mid-podfill (47 to 58% of total mineral nitrogen). Nitrogen partitioning varied with Rhizobium strains, and inoculation with strain C05 increased the nitrogen harvest index of both cultivars. Applied mineral nitrogen had a variable effect and in cv. Negro Argel was more beneficial to vegetative growth, resulting in smaller nitrogen harvest indices. Seed yield was not increased by heavy nitrogen fertilization. In contrast, cv. Rio Tibagi always benefited from nitrogen applications. Among the various nitrogen sources supplying the grain, the most important one was the fixed nitrogen translocated directly from nodules or after a rapid transfer through leaves, representing from 60 to 64% of the total nitrogen incorporated into the seeds.     

Growth, nutrition, and nitrogen fixation in grey alder at varied rate of nitrogen addition

Seedlings of grey alder (Alnus incana Moench), nodulated or unnodulated, were investigated at varied relative addition rate of nitrogen. Nitrogen fixation alone, without addition of mineral nitrogen, resulted in an almost optimum nitrogen status but only about half the maximum relative growth rate, probably mainly because of energy costs of nodulation and fixation. The growth deficit due to nodulation was much more than can be explained by the theoretical energy requirement for the amount of nitrogen fixed. Thus, the nitrogen fixation process was not very efficiently used. The nitrogen fixation rate was strongly stimulated by increasing nitrogen addition rate up to high levels. The fixation rate decreased rapidly close to optimum (maximum relative growth rate) and was negligible at maximum growth. A feed-back of mineral nitrogen on photosynthesis increased fixation rate with time, and the relative importance of fixation over mineral nitrogen nutrition increased. However, nitrogen fixation, also at maximum rate, supplied only a small proportion of the nitrogen amount required for maximum growth. The optimum nutrient solutions contained comparatively high nitrogen concentrations to secure free access to nitrogen. The nodules were damaged by this treatment, and it is concluded that the nitrogen additions must be adjusted to the current consumption of the plants to avoid an increased external nitrogen concentration. Strong linear regressions were found between relative growth rate, nitrogen status expressed as percentage content of fresh weight, and relative growth rate in unnodulated seedlings. There was a greater variability in nodulated seedlings than in unnodulated ones, because of the nitrogen fixation. The reactions of unnodulated grey alder were largely the same as previously reported for birch seedlings, but the maximum growth capacity was lower in grey alder. During an initial period of change in the internal nitrogen status, deficiency symptoms appeared, especially in unnodulated seedlings. As in birch, the leaves turned green again at stable nitrogen status, independent of level. The results are in sharp contrast to data from the literature where the external nitrogen concentration was used as the driving variable for the internal nitrogen status. The measured fixation rates for grey alder are much higher than those previously reported. Still, the maximum fixation rate observed is small compared to the total nitrogen uptake rate required for maximum growth, in contrast to reported relationships. These comparisons indicate that increased external nitrogen concentration obscures the real relations between mineral and fixed nitrogen, on one hand because of rapid inhibition of nitrogen fixation and, on the other hand, because of failure to obtain stable optimum nutrition and maximum growth by means of this treatment variable.     

Growth, nitrogen fixation and ion distribution in Medicago truncatula subjected to salt stress

This study compared the growth, nodulation, N₂ fixation, and ion distribution in three Medicago truncatula lines, in response to salt in nutrient solution. Two local lines (TN8.20 and TN6.18) and a reference line (Jemalong 6) were inoculated with a reference strain Sinorhizobium meliloti 2011, a very tolerant strain to salinity (700 mM NaCl) and grown in a controlled glasshouse with or without 75 mM NaCl. A genotypic variation in tolerance to salt was found: TN6.18 was the most sensitive line whereas TN8.20 was the most tolerant. The relative tolerance of TN8.20 was concomitant with the lowest leaf Na⁺ concentration and the highest nodule biomass production. However, nodule efficiency (amount of nitrogen fixed per g dry weight nodule) decreased in all lines. Results suggest that the tolerance to salt seems to depend on the host plant ability to protect its leaves against an excessive Na⁺ (and Cl^?) accumulation, and its ability to maintain the development of an abundant nodular system, which in turn determines an important rate of nitrogen fixation and allows the plants to conserve their growth potentialities. The loss of the nodular efficiency under salt stress seems to be compensated by a large nodule biomass.     

Growth, yield and grain quality of barley (Hordeum vulgare L.) in response to nitrogen uptake. I. A low cost, controlled nutrient supply system

This paper describes the development of an inexpensive, perlite culturesystem with recirculating nutrient solutions which required simplemaintenance and provided aerial conditions approximating those of a cropstand. A range of repeatable nitrogen experiments provided sufficient plantmaterial for sequential, destructive harvests, grown under the sameclimatic conditions and protected from rainfall by a polythene tunnel. Astudy of the effects of nitrogen uptake on development, growth, grainyield, and quality was done in two contrasting cultivars of malting barley.Two nitrogen treatments were imposed, High N where nitrogen was supplied toplant demand and Low N where nitrogen supply was restricted to one-third ofthat demand. The development of the controlled nutrient supply system isdescribed and the possible improvements and test performance according tothree criteria is discussed: (1) the uniformity of growth and nitrogenuptake within a treatment bed, (2) the ability to achieve the desirednitrogen uptakes (3 : 1, High N:Low N) and (3) that the stand of plantsbehaves as a crop rather than a collection of isolated or semi-isolatedplants are discussed. This third criterion was evaluated using Greenwood's(1982) definition of optimal nitrogen uptake for maximal growth andcompared with the results obtained in the High Ntreatments.     

Moss-specific changes in nitrogen fixation following two decades of warming,shading, and fertilizer addition

Climate warming will induce changes in Arctic ecosystem carbon balance, but besides climate, nitrogen availability is a critical controlling factor of carbon cycling. It is therefore essential to obtain knowledge on the influence of a changing climate on nitrogen fixation, as this process is the main source of new nitrogen to arctic ecosystems. In order to gain information on future nitrogen fixation rates in a changing climate, we studied the effects of two decades of warming with passive greenhouses, shading with sackcloth, and fertilization with NPK fertilizer on nitrogen fixation rates. To expand the knowledge on species-specific responses, we measured nitrogen fixation associated with two moss species: Hylocomium splendens and Aulacomnium turgidum. Our expectations of decreased nitrogen fixation rates in the fertilizer and shading treatments were met. However, contrary to our expectation of increased nitrogen fixation in the warming treatment, we observed either no change (Hylocomium) or a decrease (Aulacomnium) in fixation in the warmed plots. We hypothesize that this could be due to moss-specific responses or to long-term induced effects of the warming. For example, we observed that the soil temperature increase induced by the warming treatment was low and insignificant as vegetation height and total vascular plant cover of the warmed plots increased, and moss cover decreased. Hence, truly long-term studies lasting more than two decades provide insights on changes in key biogeochemical processes, which differ from more transient responses to warming in the Arctic.     

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.     

Growth response of Pinus radiata to multiple applications of nitrogen fertilizer and evaluation of the quantity of added nitrogen remaining in the forest system

Over a period of nine years, 922 kg ha^-1 of N was added in eight applications to a 16 year old P. radiata stand in a low rainfall area in north-east Tasmania, Australia. Fertilizing lifted current annual increment from 8.5 m³ha^-1 for the unfertilized plots, to 31 m³ha^-1. Increased growth was associated with improved health of the trees. Biomass measurements showed that there was a large increase in needle retention as well as needle mass on the fertilized trees. Concentration of N was also higher in fertilized trees. Fertilized plots contained 467 kg ha^-1 more N than unfertilized plots. This represented about 50 percent of the N applied. Most of this extra N was in the forest crop and in the surface 10 cm of the soil. There was no increase in organic carbon in the surface soil with the result that the C/N ratio was reduced from a very high 28 to 17.Despite the high growth rates attained in the N fertilized plots, the failure to increase soil organic matter and the loss from the site of much of the applied N indicated that long term improvement of growth of these low rainfall sites was unlikely. Growth would only be maintained with continued N fertilizer additions.     

有机-无机态肥氮在微型农业生态系统的转移和循环研究

本文应用~(15)N示踪法研究了有机态和无机态肥料氮在微型农业生态系统中循环的机理,以及配施下各自的功能和相互影响。初步结果表明,在土壤-黑麦草-兔亚系统中,草对硫铵~(15)N利用率达59.36％;以草饲兔,所收集兔粪~(15)N回收率为11.78％,尿为17.98％。兔粪尿N与硫铵N以1:0,0.5:0.5,0.3:0.7和0.1配施种稻,结果表明,在土壤-水稻亚系统中:(1)0.5:0.5配施促进穗头和根系发育;(2)稻株吸收有机态和无机态肥料N的比例,受配施比例的影响;(3)与单施无机肥料氮比较,配施下无机肥料氮利用率未提高或甚至下降;(4)0.3:0.7配比使有机态肥料N利用率提高;(5)0.5:0.5配比,有机肥能明显地促进无机态肥料N从秆向谷运输;(6)有机肥使无机态肥N在土壤中的固化作用增加,从而使无机肥料氮向环境转移量下降。农业生态系统具有多组分和牧食-碎屑复合食物链,氮素再循环指数达0.5,生态稳定性和发育程度优于单一种植系统。     

Understanding cassava yield response to soil and fertilizer nutrient supply in West Africa

Plant and Soil - Carbon distribution between root production and rhizodeposition represents the plant strategy for growth and nutrient capture. It can shift in response to changed availability of...     

Growth and nitrogen assimilation in nodules in response to nitrate levels in Vicia faba under salt stress

This study analyses the effects of salt on the effective symbiosisof faba bean (Vicia faba L. var. minor cv. Alborea) and salt-tolerantRhizobium leguminosarum biovar. viciae strain GRA19 grown withtwo KNO₃ levels (2 and 8 mM). The addition of 8 mM KNO₃ to thegrowth medium increases plant tolerance to salinity even witha concentration of 100 mM NaCl. This KNO₃ level in control plantsreduced the N₂ fixation. For 2 and 8 mM KNO₃ the plants treatedwith NaCl reduced N₂ fixation to identical values. The activityof the enzymes mediating ammonium assimilation in nodules (GS,NADH-GOGAT and NADH-GDH) was decreased by high KNO₃ levels.The results show that NADH-GOGAT activity was more markedlyinhibited than was GS activity by salinity, therefore NADH-GOGATlimits the ammonium assimilation by nodules in V. faba undersalt stress. The total proline content in the nodule was notrelated to salt tolerance and thus does not serve as a salttoleranceindex for V. faba. Key words: Glutamate synthase, glutamine synthetase, N₂ fixation, nitrate, salinity