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In order to improve our understanding of the regulation of nitrogen assimilation and recycling in wheat (Triticum aestivum L.), we studied the localization of plastidic (GS2) and cytosolic (GS1) glutamine synthetase isoenzymes and of glutamate dehydrogenase (GDH) during natural senescence of the flag leaf and in the stem. In mature flag leaves, large amounts of GS1 were detected in the connections between the mestome sheath cells and the vascular cells, suggesting an active transfer of nitrogen organic molecules within the vascular system in the mature flag leaf. Parallel to leaf senescence, an increase of a GS1 polypeptide (GS1b) was detected in the mesophyll cytosol of senescing leaves, while the GS protein content represented by another polypetide (GS1a) in the phloem companion cells remained practically constant in both leaves and stems. Both GDH aminating activity and protein content were strongly induced in senescing flag leaves. The induction occurred both in the mitochondria and in the cytosol of phloem companion cells, suggesting that the shift in GDH cellular compartmentation is important during leaf nitrogen remobilization although the metabolic or sensing role of the enzyme remains to be elucidated. Taken together, our results suggest that in wheat, nitrogen assimilation and recycling are compartmentalized between the mesophyll and the vasculature, and are shifted in different cellular compartments within these two tissues during the transition of sink leaves to source leaves.  相似文献   

5.
A genetic study is presented for traits relating to nitrogen use in wheat. Quantitative trait loci (QTLs) were established for 21 traits relating to growth, yield and leaf nitrogen (N) assimilation during grain fill in hexaploid wheat (Triticum aestivum L.) using a mapping population from the cross Chinese Spring × SQ1. Glutamine synthetase (GS) isozymes and estimated locations of 126 genes were placed on the genetic map. QTLs for flag leaf GS activity, soluble protein, extract colour and fresh weight were found in similar regions implying shared control of leaf metabolism and leaf size. Flag leaf traits were negatively associated with days to anthesis both phenotypically and genetically, demonstrating the complex interactions of metabolism with development. One QTL cluster for GS activity co-localised with a GS2 gene mapped on chromosome 2A, and another with the mapped GSr gene on 4A. QTLs for GS activity were invariably co-localised with those for grain N, with increased activity associated with higher grain N, but with no or negative correlations with grain yield components. Peduncle N was positively correlated, and QTLs co-localised, with grain N and flag leaf N assimilatory traits, suggesting that stem N can be indicative of grain N status in wheat. A major QTL for ear number per plant was identified on chromosome 6B which was negatively co-localised with leaf fresh weight, peduncle N, grain N and grain yield. This locus is involved in processes defining the control of tiller number and consequently assimilate partitioning and deserves further examination. Electronic Supplementary Material The online version of this article () contains supplementary material, which is available to authorized users.  相似文献   

6.
New perspectives on glutamine synthetase in grasses   总被引:2,自引:0,他引:2  
Members of the glutamine synthetase (GS) gene family have now been characterized in many crop species such as wheat, rice, and maize. Studies have shown that cytosolic GS isoforms are involved in nitrogen remobilization during leaf senescence and emphasized a role in seed production particularly in small grain crop species. Data from the sequencing of genomes for model crops and expressed sequence tag (EST) libraries from non-model species have strengthened the idea that the cytosolic GS genes are organized in three functionally and phylogenetically conserved subfamilies. Using a bioinformatic approach, the considerable publicly available information on high throughput gene expression was mined to search for genes having patterns of expression similar to GS. Interesting new hypotheses have emerged from searching for co-expressed genes across multiple unfiltered experimental data sets in rice. This approach should inform new experimental designs and studies to explore the regulation of the GS gene family further. It is expected that understanding the regulation of GS under varied climatic conditions will emerge as an important new area considering the results from recent studies that have shown nitrogen assimilation to be critical to plant acclimation to high CO(2) concentrations.  相似文献   

7.
外源钙离子对小麦幼苗氮素代谢的影响   总被引:4,自引:1,他引:3  
王志强  王春丽  林同保 《生态学报》2008,28(8):3662-3667
以普通小麦豫麦34为材料,研究了不同浓度的外源Ca2 对小麦幼苗氮素代谢的影响.在小麦第一片叶完全展开后,开始外源Ca2 处理,设0 (对照)、2、4 mmol · L-1 和8 mmol · L-1 4个Ca2 浓度梯度.处理5d后,测定氮同化酶活性、氮同化量及其它相关代谢物含量.结果表明,小麦幼苗叶片中硝酸还原酶(NR)和谷氨酰胺合成酶(GS)在2 mmol · L-1 Ca2 处理下活性比对照有显著增加,4 mmol · L-1 Ca2 处理的NR活性增加明显,但GS活性增加不显著;8 mmol · L-1 Ca2 处理下NR和GS活性比对照均明显降低.谷氨酸脱氢酶(NADH-GDH)活性在2 mmol · L-1 Ca2 处理下活性增加不明显,而在4、8 mmol · L-1 Ca2 处理下活性显著增加.小麦幼苗氮同化量以4 mmol · L-1处理最大,2 mmol · L-1处理与4 mmol · L-1之间差异不显著;Ca2 浓度为8 mmol · L-1时,氮素同化量明显降低.结果揭示了小麦幼苗不同氮同化途径对Ca2 的响应不同,GS途径比GDH途径对小麦氮素同化量的增加作用更大;4 mmol · L-1对小麦幼苗的氮素利用可能是最有效的Ca2 浓度.  相似文献   

8.
Field-grown winter wheat (Triticum aestivum L. cv. Castell) was used to study changes in the free amino acid pools of different plant parts and related enzyme activities in the flag leaf throughout the grain-filling period in three consecutive growing seasons. Amino acid analysis data indicated that, during senescence, the nitrogen flow in the flag leaf was directed towards the synthesis of glutamine as a specific nitrogen transport form. Of the enzymes involved, total glutamine synthetase (GS; EC 6.3.1.2) and especially ferredoxin-dependent glutamate synthase (Fd-GOGAT; EC 1.4.7.1) activities declined continuously as senescence progressed. Unlike (chloroplastic) GS2, (cytosolic) GS1 was shown to be very persistent suggesting a special role for this isoenzyme in the N-reallocation process. Glutamate-oxaloacetate transaminase (GOT; EC 2.6.1.1), glutamate-pyruvate transaminase (GPT; EC 2.6.1.2) and isocitrate dehydrogenase (IDH; EC 1.1.1.42) showed a characteristic biphasic activity profile after anthesis. It is proposed that these enzymes, for each of which at least two isoenzymes were demonstrated, are involved in glutamate synthesis at the later stages of leaf senescence. Ammonium levels were fairly constant throughout the flag leafs life span, an ultimate rise often following peak values of glutamate dehydrogenase (GDH; EC 1.4.1.4) activity. The enzymology of flag leaf amino acid metabolism during grain development is further discussed in relation to observations of NH3-volatilization from naturally senescing wheat plants.  相似文献   

9.
Changes in the levels of cytosolic glutamine synthetase (GS1) and chloroplastic glutamine synthetase (GS2) polypeptides and of corresponding mRNAs were determined in leaves of hydroponically grown rice (Oryza sativa) plants during natural senescence. The plants were grown in the greenhouse for 105 days at which time the thirteenth leaf was fully expanded. This was counted as zero time for senescence of the twelfth leaf. The twelfth leaf blade on the main stem was analyzed over a time period of −7 days (98 days after germination) to +42 days (147 days after germination). Total GS activity declined to less than a quarter of its initial level during the senescence for 35 days and this decline was mainly caused by a decrease in the amount of GS2 polypeptide. Immunoblotting analyses showed that contents of other chloroplastic enzymes, such as ribulose-1,5-bisphosphate carboxylase/oxygenase and Fd-glutamate synthase, declined in parallel with GS2. In contrast, the GS1 polypeptide remained constant throughout the senescence period. Translatable mRNA for GS1 increased about fourfold during the senescence for 35 days. During senescence, there was a marked decrease in content of glutamate (to about one-sixth of the zero time value); glutamate is the major form of free amino acid in rice leaves. Glutamine, the major transported amino acid, increased about threefold compared to the early phase of the harvest in the senescing rice leaf blades. These observations suggest that GS1 in senescing leaf blades is responsible for the synthesis of glutamine, which is then transferred to the growing tissues in rice plants.  相似文献   

10.
The genetic variability of the duration of leaf senescence during grain filling has been shown to affect both carbon and nitrogen acquisition. In particular, maintaining green leaves during grain filling possibly leads to increased grain yield, but its associated effect on grain protein concentration has not been studied. The aim of this study was to dissect the genetic factors contributing to correlations observed at the phenotypic level between leaf senescence during grain filling, grain protein concentration, and grain yield in winter wheat. With this aim in view, an analysis of quantitative trait locus (QTL) co-locations for these traits was carried out on a doubled haploid mapping population grown in a large multienvironment trial network. Pleiotropic QTLs affecting leaf senescence and grain yield and/or grain protein concentration were identified on chromosomes 2D, 2A, and 7D. These were associated with QTLs for anthesis date, showing that the phenotypic correlations with leaf senescence were mainly explained by flowering time in this wheat population. Study of the allelic effects of these pleiotropic QTLs showed that delaying leaf senescence was associated with increased grain yield or grain protein concentration depending on the environments considered. It is proposed that this differential effect of delaying leaf senescence on grain yield and grain protein concentration might be related to the nitrogen availability during the post-anthesis period. It is concluded that the benefit of using leaf senescence as a selection criterion to improve grain protein concentration in wheat cultivars may be limited and would largely depend on the targeted environments, particularly on their nitrogen availability during the post-anthesis period.  相似文献   

11.
Gisela Mäck 《Planta》1995,196(2):231-238
One cytosolic glutamine synthetase (GS, EC 6.3.1.2) isoform (GS 1a) was active in the germinating seeds of barley (Hordeum vulgare L.). A second cytosolic GS isoform (GS 1b) was separated from the leaves as well as the roots of 10-d-old seedlings. The chloroplastic isoform (GS 2) was present and active only in the leaves. The three GS isoforms were active in N-supplied (NH+ 4 or NO 3 ) as well as in N-free-grown seedlings. This indicates (i) that a supply of nitrogen to the germinating seeds was not necessary for the induction of the GS isoforms and (ii) that no nitrogen-specific isoforms appeared during growth of seedlings with different nitrogen sources. The activity of GS, however, depended on the seedlings' nitrogen source: the specific activity was much higher in the leaves and much lower in the roots of NH+ 4-grown barley than in the respective organs of NO 3 -fed or N free-grown plants. With increasing concentrations of NH+ 4 (supplied hydroponically during growth), the specific activity of GS 1b increased in the leaves, but decreased in the roots. The activity of GS 2 (leaf) also increased with increasing NH+ 4 supply, whereas GS 1a activity (leaf and root) was not affected. The changes in the activities of GS 1b and GS 2 were correlated with changes in the subunit compositions of the active holoenzymes: growth at increased levels of external NH+ 4 resulted in an increased abundance of one of the four GS subunits, and of two of the five GS 1b subunits in the leaves. In the roots, however, the abundance of these two GS 1b subunits was decreased under the same growth conditions, indicating an organ-specific difference either in the expression of the genes coding for the respective GS 1b subunits or in the assembly of the GS 1b holoenzymes. Furthermore, growth at different levels of NH+ 4 resulted in changes in the substrate affinities of the isoforms GS 1b (root and leaf) and GS 2 (leaf), presumably due to the changes in the subunit compositions of the active holoenzymes.Abbreviations FPLC fast protein liquid chromatography - GHA -glutamyl hydroxamate - GS glutamine synthetase Dr. Roger Wallsgrove's (Rothamsted Experimental Station, Harpenden, UK) generous gift of GS antiserum is greatly appreciated.  相似文献   

12.
To investigate the role of stress in nitrogen management in plants, the effect of pathogen attack, elicitors, and phytohormone application on the expression of the two senescence-related markers GS1 (cytosolic glutamine synthetase EC 6.3.1.2) and GDH (glutamate dehydrogenase, EC 1.4.1.2) involved in nitrogen mobilization in senescing leaves of tobacco (Nicotiana tabacum L.) plants, was studied. The expression of genes involved in primary nitrogen assimilation such as GS2 (chloroplastic glutamine synthetase) and Nia (nitrate reductase, EC 1.6.1.1) was also analysed. The Glubas gene, coding a beta-1,3-glucanase, was used as a plant-defence gene control. As during natural senescence, the expression of GS2 and Nia was repressed under almost all stress conditions. By contrast, GS1 and GDH mRNA accumulation was increased. However, GS1 and GDH showed differential patterns of expression depending on the stress applied. The expression of GS1 appeared more selective than GDH. Results indicate that the GDH and GS1 genes involved in leaf senescence are also a component of the plant defence response during plant-pathogen interaction. The links between natural plant senescence and stress-induced senescence are discussed, as well as the potential role of GS1 and GDH in a metabolic safeguard process.  相似文献   

13.
Wheat leaf non-sequential senescence at the late grain-filling stage involves the early senescence of younger flag leaves compared to that observed in older second leaves. On the other hand, sequential senescence involves leaf senescence that follows an age-related pattern, in which flag leaves are the latest to undergo senescence. The characteristics of sugar metabolism in two sequential senescence cultivars and two non-sequential senescence cultivars under both natural and drought conditions were studied to elucidate the underlying mechanism of drought tolerance in two different senescence modes. The results showed that compared to sequential senescence wheat cultivars, under natural and drought conditions, non-sequential senescence wheat cultivars showed a higher leaf net photosynthetic rate, higher soluble sugar levels in leaves, leaf sheaths, and internodes, higher leaf sucrose synthase (SS) and sucrose phosphate synthase (SPS) activity, and higher grain SS activity, thereby suggesting that non-sequential senescence wheat cultivars had stronger source activity. Spike weight, grain weight per spike, and 100-grain weight of non-sequential senescence cultivars at maturity were significantly higher than those of sequential senescence cultivars under both natural and drought conditions. These findings indicate that the higher rate of accumulation and the higher mobilization of soluble sugar in the leaves, leaf sheaths and internodes of non-sequential senescence cultivars improve grain weight and drought tolerance. At the late grain-filling stage, drought conditions adversely affected leaf chlorophyll content, net photosynthetic rate, soluble sugar and sucrose content, SS and SPS activity, gain SS activity, and weight. This study showed that higher rates of soluble sugar accumulation in the source was one of the reasons of triggering leaf non-sequential senescence, and higher rates of soluble sugar mobilization during leaf non-sequential senescence promoted high and stable wheat yield and drought tolerance.  相似文献   

14.
Leaf senescence is characterised by a massive degradation of proteins in order to recycle nitrogen to other parts of the plant, such as younger leaves or developing grain/seed. Protein degradation during leaf senescence is a highly regulated process and it is suggested that proteins to be degraded are marked by an oxidative modification (carbonylation) that makes them more susceptible to proteolysis. However, there is as yet no evidence of an increase in protein carbonylation level during natural leaf senescence. The aim of our study was thus to monitor protein carbonylation level during the process of natural senescence in the flag leaf of field‐grown winter wheat plants. For this purpose, we adapted a fluorescence‐based method using fluorescein‐5‐thiosemicarbazide (FTC) as a probe for detecting protein carbonyl derivatives. As used for the first time on plant material, this method allowed the detection of both quantitative and qualitative modifications in protein carbonyl levels during the last stages of wheat flag leaf development. The method described herein represents a convenient, sensitive and reproducible alternative to the commonly used 2,4‐dinitrophenylhydrazine (DNPH)‐based method. In addition, our analysis revealed changes in protein carbonylation level during leaf development that were associated with qualitative changes in protein abundance and carbonylation profiles. In the senescing flag leaf, protein carbonylation increased concomitantly with a stimulation of endoproteolytic activity and a decrease in protein content, which supports the suggested relationship between protein oxidation and proteolysis during natural leaf senescence.  相似文献   

15.
16.
Over‐expression of glutamine synthetase (GS, EC 6.3.1.2), a key enzyme in nitrogen assimilation, may be a reasonable approach to enhance plant nitrogen use efficiency. In this work phenotypic and biochemical characterizations of young transgenic poplars showing ectopic expression of a pine cytosolic GS transgene in photosynthetic tissue (Gallardo et al., Planta 210, 19–26, 1999) are presented. Analysis of 22 independent transgenic lines in a 6 month greenhouse study indicated that expression of the pine GS transgene affects early vegetative growth and leaf morphology. In comparison with non‐transgenic controls, transgenic trees exhibited significantly greater numbers of nodes and leaves (12%), and higher average leaf length and width resulting in an increase in leaf area (25%). Leaf shape was not altered. Transgenic poplars also exhibited increased GS activity (66%), chlorophyll content (33%) and protein content (21%). Plant height was correlated with GS content in young leaves, suggesting that GS can be considered a marker for vegetative growth. Molecular and kinetic characterization of GS isoforms in leaves indicated that poplar GS isoforms are similar to their counterparts in herbaceous plants. A new GS isoenzyme that displayed molecular and kinetic characteristics corresponding to the octomeric pine cytosolic GS1 was identified in the photosynthetic tissues of transgenic poplar leaves. These results indicate that enhanced growth and alterations in biochemistry during early growth are the consequence of transgene expression and assembly of pine GS1 subunits into a new functional holoenzyme in the cytosol of photosynthetic cells.  相似文献   

17.
The relationship of copper supply to the content and movementof copper among organs of wheat plants was examined at sevenstages in their growth from seedlings to maturity on a copperdeficient sand. In the absence of copper (Cu0), plants becameseverely copper deficient and produced no grain; developmentof tillers, leaves, stems, and inflorescences was delayed andgrowth of roots strongly depressed; leaf senescence was retardedand tiller growth was prolonged. Application of a marginal supplyof copper (Cu1) overcame all symptoms and promoted growth andgrain production. Increasing copper supply eightfold (Cu2) didnot change vegetative or grain production. Copper concentrations in stems, individual leaves, and wholetops were highest and responded most strongly to copper supplywhen they were young. As they aged, Cu1 and Cu2 leaves lostcopper rapidly; the first Cu0 leaves retained their copper andremained healthy for more than 7 weeks even though younger leavesdeveloped severe copper deficiency. In all treatments, lossof copper from the oldest leaf paralleled senescence and theloss of nitrogen. It is suggested that copper does not move out of plant leavesuntil they lose organic nitrogen compounds. As a result, copperbehaves in non-senescent leaves as if it is not mobile in plantphloem. But under conditions favouring senescence, copper ishighly mobile: in the present experiment, 67 per cent of thecopper present in vegetative organs of the Cu2 primary shootat flowering moved from them during grain development and thiscould account for all of the copper found in the grain at maturity. The retention of copper by leaves before senescence, its rapidloss during senescence, and the effect of copper deficiencyin delaying senescence resulted in the oldest leaf of severelydeficient Cu0 plants in the present experiment having a highercopper concentration than that of copper adequate Cu1 and Cu2plants. This behaviour could account for the many reports ofanomalous C-shaped ‘Piper-Steenbjerg’ curves inthe relationship of yield to copper concentrations in planttops. The coupling of copper movement from leaves to nitrogenmovement can also account for the unusually high values reportedfor critical concentrations of copper in tops of plants givenhigh levels of nitrogen fertilizers. Old organs should not be included in samples for diagnosis ofcopper deficiency. Only young organs should be used. In thepresent experiment, the copper concentration of young leavesgave a good indication of the copper status of wheat: a valueof 1 µg g–1 in young leaves indicated copper deficiency. copper, nitrogen, phloem transport, mineral transport, deficiency diagnosis, wheat, Triticum aestivum L.  相似文献   

18.
G. Mäck  R. Tischner 《Planta》1994,194(3):353-359
In extracts from the primary leaf blade of sugar beet (Beta vulgaris L.) we separated a chloroplastic isoform (GS 2) of glutamine synthetase (GS, EC 6.3.1.2) and one or two (depending on leaf age) cytosolic isoforms (GS 1a and GS 1b). The latter were prominent in the early (GS 1a) and late stages of leaf ontogeny (GS 1a and GS 1b), whereas during leaf maturation GS 2 was the predominantly active GS isoform. The GS 1 isoforms were active exclusively in the octameric state although tetrameric GS 1 protein was detected immunologically. Their activity stayed at a relatively constant level during leaf ontogeny; an increase was observed only in the senescent leaf. The activity of GS 2, however, changed drastically during primary leaf ontogeny and was modulated by changes in the oligomeric state of the active enzyme. In the early and late stages of leaf ontogeny when GS 2 activity was low (lower than that of the GS 1 isoforms), GS 2 was active only in the octameric state. In the maturing leaf, when GS 2 activity had reached its maximum level (much higher than that of the GS 1 isoforms), 80 of total GS 2 activity was due the activity of the tetrameric form of the enzyme and 20 was due to octameric GS 2. Tetrameric GS 2 was a hetero-tetramer and thus not the unspecific dissociation product of homo-octameric GS 2. In addition, GS 2 activity was modulated by an activation/inactivation of the tetrameric GS 2 protein. Due to an activation of the GS 2 tetramer, the activity of tetrameric GS 2 increased during leaf maturation from zero level 23-fold compared with that of GS 1a and 18-fold compared with that of GS 1b. Possible activators of tetrameric GS 2 are thiol-reactive substances. During leaf senescence, GS 2 activity decreased to zero; this decrease was due to an inactivation of the tetrameric GS 2 protein probably caused by oxidation.Abbreviations FLL final lamina length - FPLC fast protein liquid chromatography - GS glutamine synthetase - GHA -glutamyl hydroxamate - Rubisco ribulose-1,5-bisphosphate carboxylase/oxygenase Dr. Roger Wallsgrove's (Rothamsted Experimental Station, Harpenden, UK) generous gift of GS antiserum is greatly appreciated.  相似文献   

19.
Nitrogen availability has a strong influence on developmental processes in plants. We show that the time of nitrogen supply regulates the course of leaf senescence in flag leaves of Hordeum vulgare . The senescence-specific decrease in chlorophyll content and photosystem II efficiency is clearly delayed when plants are fertilised with nitrate at the onset of leaf senescence. Concurrently, the additional supply of nitrate affects expression patterns of two marker genes of nitrogen metabolism. As shown by quantitative RT-PCR analyses, senescence-specific downregulation of plastidic glutamine synthetase ( GS2 ) and senescence-specific upregulation of lysine-ketoglutarate reductase/saccharopine dehydrogenase ( LKR/SDH ) are both clearly retarded. Depletion of nitrogen in experiments using hydroponic growth systems results in premature primary leaf senescence. The already started senescence processes can be even reversed by later nitrogen addition, as proved by a further increase in photosystem II efficiency and chlorophyll content, returning to the high values of controls which had not been deprived of nitrogen. Although both addition of nitrate or ammonium effectively reversed nitrogen depletion-induced primary leaf senescence, addition of urea did not. Additionally, effects of nitrogen supply on the course of leaf senescence were analysed in the model plant Arabidopsis thaliana. Leaves of A. thaliana show the same reversion of senescence processes after receiving additional nitrogen supply, indicating that the nitrogen response of leaf development is conserved in different plant species.  相似文献   

20.
硫氮配施对持绿型小麦氮素运转及叶片衰老的影响   总被引:4,自引:0,他引:4  
以持绿型小麦品种‘豫麦66号’、‘潍麦8号’及非持绿型品种‘小偃6号’为材料,采用以氮肥为主区硫肥为副区的田间裂区试验,研究了2个施氮水平[纯氮120kg/hm2(N120)、220kg/hm2(N220)]和3个施硫水平[纯硫0kg/hm2(S0)、20kg/hm2(S20)、60kg/hm2(S60)]下植株各部位的含氮量、叶片干鲜重及叶片叶绿素含量的变化,探讨硫氮配施对不同类型小麦氮吸收及衰老的影响。结果表明:在相同处理下,持绿型小麦植株的总含氮量、氮素转运量、叶绿素含量、叶片含水量及穗粒数和千粒重均高于非持绿型小麦。N120处理条件下,不同硫肥处理时持绿型小麦与非持绿型小麦变化趋势相同,开花期茎和叶含氮量及叶绿素含量在S60处理下均低于其他2个硫肥处理,生育后期叶片含水量下降幅度也明显高于其他处理;在N220处理条件下,3个品种开花期叶含氮量、收获期总氮累积量、氮收获指数、叶绿素含量及叶片含水量在S60处理下均高于其他2个处理,其中非持绿型小麦在高硫处理条件下灌浆期的叶绿素含量的增长率明显高于持绿型小麦,而灌浆中后期叶片含水量的下降幅度则明显低于持绿型小麦。研究发现,施用硫肥在氮肥不足时会对小麦植株氮素的吸收利用及叶片衰老等方面产生负面影响,但在氮肥充足时却在氮素的吸收利用、延缓叶片衰老及最终籽粒产量和总生物量等方面表现出正面效应;本实验条件下,220kg/hm2左右施氮量和60kg/hm2左右施硫量有利于各品种小麦生长发育和产量提高;高N和高S水平对于延缓小麦的衰老而言,非持绿性小麦比持绿型小麦更明显。  相似文献   

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