Tolerance To Nitrogen Deficiency Research Articles

Nitrogen deficiency tolerance conferred by introgression of a QTL derived from wild emmer into bread wheat

Key messageGenetic dissection of a QTL from wild emmer wheat, QGpc.huj.uh-5B.2, introgressed into bread wheat, identified candidate genes associated with tolerance to nitrogen deficiency, and potentially useful for improving nitrogen-use efficiency.Nitrogen (N) is an important macronutrient critical to wheat growth and development; its deficiency is one of the main factors causing reductions in grain yield and quality. N availability is significantly affected by drought or flooding, that are dependent on additional factors including soil type or duration and severity of stress. In a previous study, we identified a high grain protein content QTL (QGpc.huj.uh-5B.2) derived from the 5B chromosome of wild emmer wheat, that showed a higher proportion of explained variation under water-stress conditions. We hypothesized that this QTL is associated with tolerance to N deficiency as a possible mechanism underlying the higher effect under stress. To validate this hypothesis, we introgressed the QTL into the elite bread wheat var. Ruta, and showed that under N-deficient field conditions the introgression IL99 had a 33% increase in GPC (p < 0.05) compared to the recipient parent. Furthermore, evaluation of IL99 response to severe N deficiency (10% N) for 14 days, applied using a semi-hydroponic system under controlled conditions, confirmed its tolerance to N deficiency. Fine-mapping of the QTL resulted in 26 homozygous near-isogenic lines (BC4F5) segregating to N-deficiency tolerance. The QTL was delimited from − 28.28 to − 1.29 Mb and included 13 candidate genes, most associated with N-stress response, N transport, and abiotic stress responses. These genes may improve N-use efficiency under severely N-deficient environments. Our study demonstrates the importance of WEW as a source of novel candidate genes for sustainable improvement in tolerance to N deficiency in wheat.

Nitrogen-deficient leaves and roots can keep high abilities to scavenge reactive oxygen species and methylglyoxal, and protect them against oxidative damage in Citrus sinensis seedlings

Nitrogen-deficiency (ND) usually occurs in some citrus orchard soils in China. The roles of reactive oxygen species (ROS) and methylglyoxal (MG) generation and their detoxification systems in ND tolerance of horticultural woody plants still need to be revealed. For the first time, we examined the effects of ND on ROS and MG generation and their detoxification systems in leaves and roots of Citrus sinensis seedlings. The objectives are to test the hypotheses that N-deficient leaves and roots can keep high abilities to scavenge ROS and MG, thereby protecting them from oxidative damage, and that ND-induced alterations of ROS and MG formation and their detoxification systems in leaves and roots are different. ND augmented superoxide anion production rate and MG concentrations, but it decreased malondialdehyde (MDA) concentrations and electrolyte leakage in leaves and roots. ND increased the activities of most enzymes involved in ROS (ascorbate-glutathione cycle-related enzymes, antioxidant enzymes, and sulfur metabolism-related enzymes) and MG (glyoxalases) detoxification expressed on a protein basis with a few exceptions, and the concentrations of ascorbate, phytochelatins, and total non-protein thiols in leaves and roots. These results suggested that nitrogen-deficient leaves and roots could keep high abilities to detoxify ROS and MG, and protect them from oxidative damage. Generally viewed, ND affected the production and removal of ROS and MG more in roots than in leaves.

Mining natural genetic variations for nitrogen use efficiency utilizing nested synthetic hexaploid wheat introgression libraries

Wheat is grown on more than 240 million hectares globally, more than any other commercial crop. Variation for nitrogen-use efficiency (NUE) in wheat germplasm is very important as efficiency of nitrogen fertilizers use can have significant effect on overall consumption of N. In the present study we used nested synthetic hexaploid wheat introgression libraries to detect genetic variation associated with NUE and related traits. The libraries were genotyped with 9474 SNP markers and used for genome-wide association mapping. Significant phenotypic variation was observed for all measured and derived traits. We detected 10 Quantitative trait locus (QTL) and 19 marker-trait associations (MTAs) possibly involved in improving NUE. Of these, 5 QTL and 8 MTAs detected under nitrogen-limited conditions have potential for use in breeding to increase nitrogen-deficiency tolerance. Nitrate transporter genes collocated with detected MTAs showed significant changes in expression of TaNRT2 genes in response to N-starvation and N-recovery. The identified promising breeding lines with stable yield, better NUE and acceptable protein content may constitute an important genetic resource in improving NUE of modern wheat varieties.

Long term nitrogen deficiency alters expression of miRNAs and alters nitrogen metabolism and root architecture in Indian dwarf wheat (Triticum sphaerococcum Perc.) genotypes

The important roles of plant microRNAs (miRNAs) in adaptation to nitrogen (N) deficiency in different crop species especially cereals (rice, wheat, maize) have been under discussion since last decade with little focus on potential wild relatives and landraces. Indian dwarf wheat (Triticum sphaerococcum Percival) is an important landrace native to the Indian subcontinent. Several unique features, especially high protein content and resistance to drought and yellow rust, make it a very potent landrace for breeding. Our aim in this study is to identify the contrasting Indian dwarf wheat genotypes based on nitrogen use efficiency (NUE) and nitrogen deficiency tolerance (NDT) traits and the associated miRNAs differentially expressed under N deficiency in selected genotypes. Eleven Indian dwarf wheat genotypes and a high NUE bread wheat genotype (for comparison) were evaluated for NUE under control and N deficit field conditions. Based on NUE, selected genotypes were further evaluated under hydroponics and miRNome was compared by miRNAseq under control and N deficit conditions. Among the identified, differentially expressed miRNAs in control and N starved seedlings, the target gene functions were associated with N metabolism, root development, secondary metabolism and cell-cycle associated pathways. The key findings on miRNA expression, changes in root architecture, root auxin abundance and changes in N metabolism reveal new information on the N deficiency response of Indian dwarf wheat and targets for genetic improvement of NUE.

Understanding the physiological, genetic and molecular basis of nitrogen deficiency tolerance and their application in rice improvement

Nitrogen (N) is a major nutrient required for growth and yield of rice plants. Several factors including plant, edapic and climate conditions influence the criticle yield response curve of the plants. Apart from breeding for N responsive rice varieties, excessive use of nitrogenous fertilizers have become a general farmers practice to boost rice productivity under intensive cropping system. Now, it is imperative to orient the crop improvement programme for sustainable crop production strategy as well as to achieve the evergreen revolution through improving nitrogen use efficiency (NUE) under global climate change condition. To develop N-efficient rice varieties under crop breeding programs, it is crucial to comprehend the physiological, genetic and molecular features associated with tolerance to nitrogen deprivation. It has always been challenging for a rice breeders to develop rice varieties with high nitrogen use efficiency (NUE), as it is highly complex physiological trait involving several component traits and its dynamic interaction with environemental factor. NUE is a polygenic traits controlled by number of quantitative trait loci's at genomic level. Till date, researchers targeted component traits for increasing NUE such as, nitrogen uptake/absorption, transport from root to shoot, assimilation, utilisation, remobilisation, reasssssmilation and partitioning /redistribution. Here, we described a short summary of the physiological, genetic and molecular underpinnings of nitrogen deficit tolerance and how these prior art information can be used for improving NUE in rice. Insight from our discussions may facilitate the breeders to improve the NUE of rice plants in future.

Nitrogen Deficiency Tolerance and Responsiveness of Durum Wheat Genotypes in Ethiopia

Development of low-nitrogen (N) tolerant and N-responsive durum wheat genotypes is required since nitrogen efficiency has emerged as a highly desirable trait from economic and environmental perspectives. Two hundred durum wheat genotypes were evaluated at three locations under optimum (ON) and low (LN) nitrogen conditions to screen genotypes for low-nitrogen tolerance and responsiveness to an optimum N supply. The results showed significant variations among the durum wheat genotypes for low-N tolerance and responsiveness. The average reduction in grain yield under the LN condition was 48.03% across genotypes. Only 17% of the genotypes tested performed well (grain yield reduction &lt;40%) under LN conditions. Based on the absolute grain yield, biomass yield, and normalized difference vegetative index value, on average, 32, 14, 17, and 37% of the tested genotypes were classified as efficient and responsive, efficient and nonresponsive, inefficient and responsive, and inefficient and nonresponsive, respectively. Considering the absolute and relative grain yield, biomass yield, normalized difference vegetative index values, and stress tolerance indices as selection criteria, 17 genotypes were chosen for subsequent breeding. Among the screening indices, geometric mean productivity, stress tolerance index, yield index, and stress susceptibility index exhibited positive and significant correlations with grain yield under both N conditions; hence, either of these traits can be used to select low-N-tolerant genotypes. The common genotypes identified as LN-tolerant and responsive to N application in this study could be used as parental donors for developing N-efficient and responsive durum wheat varieties.

Identification of shoot and root genomic regions in response to nitrogen deficiency tolerance in rice through the use of rice back cross recombinant inbred lines (BRILs)

Identification of shoot and root genomic regions in response to nitrogen deficiency tolerance in rice through the use of rice back cross recombinant inbred lines (BRILs)

Screening of Wheat Genotypes for Nitrogen Deficiency Tolerance Using Stress Screening Indices

An increased awareness of environmental protection and sustainable production raise the necessity of incorporating the selection of low nitrogen-tolerant winter wheat cultivars for high yield and quality in the breeding process. This selection can be assisted by using stress screening indices. Our study aimed to evaluate and compare a number of stress screening indices and to determine and select the most nitrogen deficiency-tolerant winter wheat cultivars for further breeding. The experiment included forty-eight winter wheat cultivars from eight different countries that were grown for two consecutive years at three different locations under low-nitrogen (LN) and high-nitrogen (HN) conditions. The results emphasized the importance of applying the appropriate stress screening indices in evaluating and selecting nitrogen deficiency-tolerant wheat cultivars. The promising stress screening indices were the mean productivity index (MP), geometric mean productivity index (GMP), harmonic mean index (HM), stress tolerance index (STI) and yield index (YI). They identified cultivars Sofru, BC Opsesija and MV-Nemere as the most tolerant cultivars to LN conditions for grain yield. The same indices classified U-1, OS-Olimpija, Forcali, Viktoria and BC Tena cultivars as the most tolerant to LN conditions for the grain protein content. Using the tolerance index (TOL), yield stability index (YSI) and relative stress index (RSI), the Katarina and Ficko cultivars were denoted as LN-tolerant cultivars in terms of the grain yield and Isengrain, Tosunbey, Vulkan and BC Darija in terms of the grain protein content.

QTL Mapping and Favorable Allele Mining of Nitrogen Deficiency Tolerance Using an Interconnected Breeding Population in Rice.

Nitrogen is one of the most important nutrients for rice growth and development. Breeding of nitrogen deficiency tolerance (NDT) variety is considered to be the most economic measure to solve the constrain of low nitrogen stress on grain yield in rice. An interconnected breeding (IB) population of 497 lines developed using Huanghuazhan (HHZ) as the recurrent parent and eight elite lines as the donor parents were tested for five traits including grain yield, biomass, harvest index, thousand grain weight, and spikelet fertility under two nitrogen treatments in three growing seasons. Association analysis using 7,388 bins generated by sequencing identified a total of 14, 14, and 12 QTLs for the five traits under low nitrogen (LN), normal nitrogen (NN), and LN/NN conditions, respectively, across three seasons. Favorable alleles were dissected for the 40 QTLs at the 10 NDT regions, and OM1723 was considered as the most important parent with the highest frequency of favorable alleles contributing to NDT-related traits. Six superior lines all showed significantly higher GY in LN environments and similar GY under NN environments except for H10. Substitution mapping using near-isogenic introgression lines delimited the qTGW2-1, which was identified on chromosome 2 under LN, NN, and LN/NN conditions into two QTLs, which were located in the two regions of about 200 and 350 kb with different favorable alleles. The bins 16, 1301, 1465, 1486, 3464, and 6249 harbored the QTLs for NDT detected in this study, and the QTLs/genes previously identified for NDT or nitrogen use efficiency (NUE) could be used for enhancing NDT and NUE by marker-assisted selection (MAS).

Cyanobacterial NOS expression improves nitrogen use efficiency, nitrogen-deficiency tolerance and yield in Arabidopsis

Developing strategies to improve nitrogen (N) use efficiency (NUE) in plants is a challenge to reduce environmental problems linked to over‐fertilization. The nitric oxide synthase (NOS) enzyme from the cyanobacteria Synechococcus PCC 7335 (SyNOS) has been recently identified and characterized. SyNOS catalyzes the conversion of arginine to citrulline and nitric oxide (NO), and then approximately 75 % of the produced NO is rapidly oxidized to nitrate by an unusual globin domain in the N-terminus of the enzyme. In this study, we assessed whether SyNOS expression in plants affects N metabolism, NUE and yield. Our results showed that SyNOS-expressing transgenic Arabidopsis plants have greater primary shoot length and shoot branching when grown under N-deficient conditions and higher seed production both under N-sufficient and N-deficient conditions. Moreover, transgenic plants showed significantly increased NUE in both N conditions. Although the uptake of N was not modified in the SyNOS lines, they showed an increase in the assimilation/remobilization of N under conditions of low N availability. In addition, SyNOS lines have greater N-deficiency tolerance compared to control plants. Our results support that SyNOS expression generates a positive effect on N metabolism and seed production in Arabidopsis, and it might be envisaged as a strategy to improve productivity in crops under adverse N environments.

Genetic Architecture of Nitrogen-Deficiency Tolerance in Wheat Seedlings Based on a Nested Association Mapping (NAM) Population.

Genetic divergence for nitrogen utilization in germplasms is important in wheat breeding programs, especially for low nitrogen input management. In this study, a nested association mapping (NAM) population, derived from “Yanzhan 1” (a Chinese domesticated cultivar) crossed with “Hussar” (a British domesticated cultivar) and another three semi-wild wheat varieties, namely, “Cayazheda 29” (Triticum aestivum ssp. tibetanum Shao), “Yunnan” (T. aestivum ssp. yunnanense King), and “Yutian” (T. aestivum petropavloski Udats et Migusch), was used to detect quantitative trait loci (QTLs) for nitrogen utilization at the seedling stage. An integrated genetic map was constructed using 2,059 single nucleotide polymorphism (SNP) markers from a 90 K SNP chip, with a total coverage of 2,355.75 cM and an average marker spacing of 1.13 cM. A total of 67 QTLs for RDW (root dry weight), SDW (shoot dry weight), TDW (total dry weight), and RSDW (root to shoot ratio) were identified under normal nitrogen conditions (N+) and nitrogen deficient conditions (N−). Twenty-three of these QTLs were only detected under N− conditions. Moreover, 23 favorable QTLs were identified in the domesticated cultivar Yanzhan 1, 15 of which were detected under N+ conditions, while only four were detected under N− conditions. In contrast, the semi-wild cultivars contributed more favorable N−−specific QTLs (eight from Cayazheda 29; nine from Yunnan), which could be further explored for breeding cultivars adapted to nitrogen-deficient conditions. In particular, QRSDW-5A.1 from YN should be further evaluated using high-resolution mapping.

Identification of QTLs associated with agronomic performance under nitrogen-deficient conditions using chromosome segment substitution lines of a wild rice relative, Oryza rufipogon

Improved root system architecture can enhance agronomic performance by increasing water and nitrogen (N) acquisition efficiency. However, little is known about interaction between root system architecture and agronomic performance under field environments. To gain a better understanding about the genetic basis of these relationships, we evaluated a set of chromosome segment substitution lines (CSSLs) derived from crosses between a tropical japonica rice cultivar ‘Curinga’ and a wild species Oryza rufipogon accession IRGC105491. Root system architectural traits were investigated using the CSSLs at 40 days old seedlings using the root basket method under hydroponic conditions, and agronomic performances were also tested under field conditions with different N treatments. Agronomic performances were computed as the ratio of a trait value under low to high N treatments, including grain yield and biomass yield as nitrogen-deficiency tolerance (NDT) traits. Root architecture and NDT trait QTLs were mapped using 238 SNP marker loci. A total of 13 QTLs for root system architectural, NDT and morpho-physiological traits were identified on chromosomes 1, 3, 4, 5, 7, 8, 9, 10 and 12. Interestingly, a QTL for deeper root number was identified the region of SNP markers between id1012330 and id1021697 on chromosome 1 under hydroponic conditions overlapped with a QTL for NDT trait of relative grain yield (qRGY1). These results suggest that deeper root trait is helpful to maintain grain yield under nitrogen-deficient conditions. The QTL associated root architecture could potentially be used in future rice-breeding efforts to increase agronomic performance under nitrogen-deficient conditions.

QTL mapping for nitrogen-use efficiency and nitrogen-deficiency tolerance traits in rice

Aims The improvement of nitrogen-deficiency tolerance (NDT) and nitrogen-use efficiency (NUE) traits is an important objective of many rice breeding programs. A better understanding of their relationship is required for more efficient breeding. The objectives of this study were to map quantitative trait loci (QTLs) for NDT and NUE traits, and to gain a better understanding about the genetic basis of the relationships between NUE and NDT traits.

Mapping QTLs for nitrogen‐deficiency tolerance at seedling stage in rice (Oryza sativaL.)

With 2 figures and 4 tablesAbstractA recombinant inbred line (RIL) population, consisting of 238 F12lines derived from ‘R9308’/’Xieqingzao B’, the parents of a super hybrid rice in China, was developed for mapping quantitative trait locus (QTL) for nitrogen deficiency tolerance. A genetic linkage map with 198 simple sequence repeat (SSR) markers was constructed. Six traits of shoot dry weight (SW), root dry weight (RW), plant dry weight (PW), maximum root length (RL), chlorophyll content (Chl), plant height (PH) were used to assess the N‐deficiency tolerance. The ratio of the parameters from the two treatments was calculated as derived parameters. QTL analysis detected seven QTLs on chromosomes 1, 2, 3 and 8 associated with N‐deficiency tolerance in this mapping population. These QTLs explained 9.07–14.45% of the phenotypic variation. The ‘R9308’ alleles increased the relative traits except that of locus on chromosome 8. One of the major QTL clusters was detected in the interval RM5385–RM7192 on chromosome 1, which was associated with N recycling in rice. This chromosomal region may be enriched with the key N metabolism genes.

Phenylalanine ammonia-lyase activity and phenolic compounds accumulation in nitrogen-deficient Matricaria chamomilla leaf rosettes

Phenylalanine ammonia-lyase (PAL) activity, accumulation of 11 phenolic acids and 3 coumarin-related compounds (herniarin and its glucosidic precursors ( Z)- and ( E)-2- β- d-glucopyranosyloxy-4-methoxycinnamic acids; umbelliferone), biomass production, plant water content and lipid peroxidation status of Matricaria chamomilla L. leaf rosettes subjected to nitrogen (N) deficiency over 12 days were studied. Both biomass and water content of N-deficient rosettes decreased in comparison to controls, while quantities of most benzoic acids ( p-hydroxybenzoic, syringic, vanillic), cinnamic acids (caffeic, chlorogenic, o- and p-coumaric, ferulic), herniarin and ( Z)- and ( E)-2- β- d-glucopyranosyloxy-4-methoxycinnamic acids increased with prolonged nitrogen deficiency. However, three benzoic derivatives (gallic acid, protocatechuic acid and protocatechuicaldehyde) did not differ significantly from the controls. The content of umbelliferone, a stress metabolite in the leaves of chamomile, was not affected by N deficiency. It thus appears that synthesis of benzoic and cinnamic acids and of coumarins is probably regulated independently. PAL activity was significantly higher in N-deficient leaf rosettes after 4 and 8 days (by 61 and 23%, respectively), but decreased to control level by the end of the experiment. Nevertheless, it can be concluded that this is important biochemical factor contributing to the observed increase of phenolic compounds accumulation by producing nitrogen-free skeletons of t-cinnamate for subsequent pathways of phenylpropanoid metabolism. We also observed that the mechanism of nitrogen deficiency tolerance due to enhanced PAL activity is temporally limited. The non-altered extent of lipid peroxidation expressed as malondialdehyde amounts revealed that membrane integrity was not affected by N deficiency. This phenomenon in correlation to the antioxidative properties of phenolic metabolites is also discussed.

Differential change in root protein patterns of two wheat varieties under high and low nitrogen nutrition levels

Nitrogen deficiency tolerance is necessary to breed for wheat varieties adapted to low input management systems. The aim of this study was to identify root proteins differentially expressed in response to a nitrogen stress. Two wheat varieties were experimented at low (0.5 mM nitrate) and at high (3.0 mM nitrate) nitrogen levels. After 8 weeks, root samples were taken to assess enzyme activities and to perform two dimensional gel electrophoresis. A total of 860 root protein spots were qualitatively or quantitatively analysed. A significant variety effect existed for 74 spots, while a significant nitrate supply-dependent variation was observed for 56 spots. Moreover 20 spots showed a significant variety × N-supply effect. Correlations were made between physiological parameters (nitrogen content and enzyme activities) and the intensity of variable spots. Positive correlations were observed for nitrate reductase and glutamate dehydrogenase activities and the intensity of a number of root proteins. This approach may be a way to identify new structural or regulatory proteins involved in the response to the level of nitrogen fertilisation in different wheat varieties.