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Plant genotypes shape root-associated microbiota that affect plant nutrient acquisition and productivity. It is unclear how maize hybrids modify root-associated microbiota and their functions and relationship with nitrogen use efficiency (NUE) by regulating rhizosphere soil metabolites. Here, two N-efficient (NE) (ZD958, DMY3) and two N-inefficient (NIE) maize hybrids (YD9953, LY99) were used to investigate this issue under low N (60 kg N ha

Appropriate root system architecture (RSA) can improve maize yields in densely planted fields, but little is known about its genetic basis in maize. Here we performed root phenotyping of 14,301 field-grown plants from an association mapping panel to study the genetic architecture of maize RSA. A genome-wide association study identified 81 high-confidence RSA-associated candidate genes and revealed that 28 (24.3%) of known root-related genes were selected during maize domestication and improvement. We found that modern maize breeding has selected for a steeply angled root system. Favourable alleles related to steep root system angle have continuously accumulated over the course of modern breeding, and our data pinpoint the root-related genes that have been selected in different breeding eras. We confirm that two auxin-related genes, ZmRSA3.1 and ZmRSA3.2, contribute to the regulation of root angle and depth in maize. Our genome-wide identification of RSA-associated genes provides new strategies and genetic resources for breeding maize suitable for high-density planting.

Aims A sound root system architecture (RSA) is important for efficient nutrient and water acquisition. The adaptation of root growth and RSA to the special soil structure under conservative strip tillage (ST) system is largely unknown. Methods We describe three-year field experimentation on a meadow black soil in the Northeast China to investigate the RSA and the dynamic root growth of maize under ST by comparison with the conventional tillage (CT). Results Grain yield in ST and CT were not significantly different, but their yield components differed. Compared to CT, grain number per ear was reduced by 4.4%, while 1000-grain weight was increased by 6.6% in ST. Root growth in ST plants was inhibited in the vegetative stage, as indicated by the reduced total root length (by 27.7-40.1%) compared to CT. During post-silking stage, however, the total root length was not different between ST and CT plants. Accordingly, the root xylem bleeding rate in ST plants was 70.7%-449.9% greater than CT plants. Because ST practice caused uneven horizontal distribution of soil bulk density, the RSA became steeper in ST plants compared to CT. Moreover, the D95 of ST plant roots was greater, indicating their roots grew deeper. Conclusions In conclusion, although root growth is inhibited at vegetative stage, the slow-aging and steep, deep roots in ST plants may contribute to larger grain weight by favoring water and nutrient acquisition during grain filling stage. As a result, grain yield of ST plants is maintained though grain number per ear is reduced.

Nitrogen (N) deficiency is one of the main factors limiting maize (Zea mays L.) productivity. Genetic improvement of root traits could improve nitrogen use efficiency. An association panel of 461 maize inbred lines was assayed for root growth at seedling emergence under high-nitrate (HN, 5 mmol L−1) and low-nitrate (LN, 0.05 mmol L−1) conditions. Twenty-one root traits and three shoot traits were measured. Under LN conditions, the root-to-shoot ratio, root dry weight, total root length, axial root length, and lateral root length on the primary root were all increased. Under LN conditions, the heritability of the plant traits ranged from 0.43 to 0.82, a range much wider than that of 0.27 to 0.55 observed under HN conditions. The panel was genotyped with 542,796 high-density single-nucleotide polymorphism (SNP) markers. Totally 328 significant SNP markers were identified using either mixed linear model (MLM) or general linear model analysis, with 34 detected by both methods. In the 100-kb intervals flanking these SNP markers, four candidate genes were identified. Under LN conditions, the protoporphyrinogen IX oxidase 2 gene was associated with total root surface area and the DELLA protein-encoding gene was associated with the length of the visible lateral root zone of the primary root. Under HN conditions, a histone deacetylase gene was associated with plant height. Under both LN and HN conditions, the gene encoding MA3 domain-containing protein was associated with the first whorl crown root number. The phenotypic and genetic information from this study may be exploited for genetic improvement of root traits aimed at increasing NUE in maize.

In plants, nitrogen remobilization from source to sink organs is an important process regulated by complex transcriptional regulatory networks. However, the relationship between nitrogen remobilization and leaf senescence and the molecular regulatory network that controls them are unknown in maize. Here, using 15N labeling and a transcriptome approach, a dynamic analysis of the nitrogen remobilization process was conducted in two elite maize inbred lines (PH4CV and PH6WC) with contrasting leaf senescence. PH4CV showed higher nitrogen remobilization efficiency (NRE) than PH6WC, mainly in the middle and lower leaves from 15 d to 35 d after silking. The co-expression network analysis revealed that ethylene and cytokinin metabolism-related genes triggered the onset of nitrogen remobilization, while abscisic acid and jasmonic acid biosynthesis-related genes controlled the progression of nitrogen remobilization. By integrating genetic analysis, functional annotation, and gene expression, two candidate genes underlying a major quantitative trait locus of NRE were identified, namely an early senescence acting gene (ZmASR6) and an ATP-dependent Clp protease gene (GRMZM2G172230). Hormone-coupled transcription factors and downstream target genes reveal a gene regulatory network for the nitrogen remobilization process after silking in maize. These results uncovered a sophisticated regulatory mechanism for nitrogen remobilization, and further provided characterization of valuable genes for genetic improvement of nitrogen use efficiency in maize.

To test the hypothesis that stay-green genotypes help to increase stalk lodging resistance under high planting density by reducing dry matter remobilization from stalk, a two-year field study with three planting densities (60,000, 75,000, and 90,000 plants per ha) and six hybrids was conducted. When planting density changed from 75,000 to 90,000, stalk dry matter remobilization efficiency increased by 4.1–191%. Stem dry weight per length, rind penetration strength, average crushing strength for the first to fifth internode, and average bending strength for the third to fifth internode decreased by 25.4–47.2%, 5.1–24.6%, 1.4–29.9%, and 14.0–36.0%, respectively. Stalk lodging rate increased by 17–275%. Stalk dry matter remobilization efficiency was negatively correlated to either stem dry weight per length (r = 0.23, p

A MADS-box transcription factor transcriptionally up-regulated and modulating lateral root development in response to local nitrate supply exists as a truncated form unique to grass species., Plants can develop root systems with distinct anatomical features and morphological plasticity to forage nutrients distributed heterogeneously in soils. Lateral root proliferation is a typical nutrient-foraging response to a local supply of nitrate, which has been investigated across many plant species. However, the underlying mechanism in maize roots remains largely unknown. Here, we report on identification of a maize truncated MIKC-type MADS-box transcription factor (ZmTMM1) lacking K- and C-domains, expressed preferentially in the lateral root branching zone and induced by the localized supply of nitrate. ZmTMM1 belongs to the AGL17-like MADS-box transcription factor family that contains orthologs of ANR1, a key regulator for root nitrate foraging in Arabidopsis. Ectopic overexpression of ZmTMM1 recovers the defective growth of lateral roots in the Arabidopsis anr1 agl21 double mutant. The local activation of glucocorticoid receptor fusion proteins for ZmTMM1 and an artificially truncated form of AtANR1 without the K- and C-domains stimulates the lateral root growth of the Arabidopsis anr1 agl21 mutant, providing evidence that ZmTMM1 encodes a functional MADS-box that modulates lateral root development. However, no phenotype was observed in ZmTMM1-RNAi transgenic maize lines, suggesting a possible genetic redundancy of ZmTMM1 with other AGL17-like genes in maize. A comparative genome analysis further suggests that a nitrate-inducible transcriptional regulation is probably conserved in both truncated and non-truncated forms of ZmTMM1-like MADS-box transcription factors found in grass species.

Maize growth in Northeast China is suffering from climate change (seasonal drought, cold springs) and low nutrient use efficiency caused by one-time fertilization. Drip fertigation is widely used in vegetable and fruit plant production, yet an efficient, practicable and cost-effective drip fertigation system is lacking for maize production. A two-year field experiment was conducted to evaluate the potential of different drip fertigation methods for increasing maize yield, and water and nutrient use efficiency in sandy and clay soil. Five irrigation methods were applied in each soil: conventional (rain-fed, CK), drip irrigation (DI), surface drip fertigation (SDF), fertigation plus plastic film mulching (SDFP), and subsurface fertigation (SSDF). Compared with rain-fed method (CK), water optimization by DI increased grain yield by (28% in sandy soil and 12% in clay soil), partial fertilizer productivity (PFP) and nitrogen (N), phosphorus (P) and potassium (K) uptake, without effect on water productivity (WP) in both soils. The optimization of both water and nutrient management by SDF increased grain yield by (41% in sandy soil and 17% in clay soil), PFP and NPK uptake, at greater extent than DI. Furthermore, SDF also increased the water productivity in both soils. Compared with DI, SDF increased post-silking N in both soil, and K accumulation in sandy soil. There was no significant difference in yield and PFP between SDF, SSDF and SDFP methods in both soils. In sandy soil, the net profit of DI, SDF, SSDF and SDFP was 13%, 28%, 31% and 10% higher than that of CK, respectively. In clay soil, However, No obvious advantage in net income was found in either DI or fertigation treatments. SDF and SSDF are recommended to increase maize yield, water and nutrient use efficiency, as well as economic benefit synchronously in sandy soil.

Maize plants grown in the presence of a mixed nitrate and ammonium supply exhibited rapid biomass accumulation associated with increased auxin biosynthesis and up-regulation of auxin-responsive transcripts, including those known to regulate cell expansion., The use of mixed nitrate and ammonium as a nitrogen source can improve plant growth. Here, we used metabolomics and transcriptomics to study the underlying mechanisms. Maize plants were grown hydroponically in the presence of three forms of nitrogen (nitrate alone, 75%/25% nitrate/ammonium, and ammonium alone). Plants grown with mixed nitrogen had a higher photosynthetic rate than those supplied only with nitrate, and had the highest leaf area and shoot and root biomass among the three nitrogen treatments. In shoot and root, the concentration of nitrogenous compounds (ammonium, glutamine, and asparagine) and carbohydrates (sucrose, glucose, and fructose) in plants with a mixed nitrogen supply was higher than that with nitrate supply, but lower than that with ammonium supply. The activity of the related enzymes (glutamate synthase, asparagine synthase, phosphoenolpyruvate carboxylase, invertase, and ADP-glucose pyrophosphorylase) changed accordingly. Specifically, the mixed nitrogen source enhanced auxin synthesis via the shikimic acid pathway, as indicated by the higher levels of phosphoenolpyruvate and tryptophan compared with the other two treatments. The expression of corresponding genes involving auxin synthesis and response was up-regulated. Supply of only ammonium resulted in high levels of glutamine and asparagine, starch, and trehalose hexaphosphate. We conclude that, in addition to increased photosynthesis, mixed nitrogen supply enhances leaf growth via increasing auxin synthesis to build a large sink for carbon and nitrogen utilization, which, in turn, facilitates further carbon assimilation and nitrogen uptake.

Frequent spring droughts and nitrogen (N) fertilizer overuse limit rain-fed maize production in Northeast China. However, the interactions between water-stress and N rates on maize seedling growth and root development remain elusive. In this study, a pot experiment was conducted with maize grown under three N rates (N omission, low and high N rates, i.e., N0, LN and HN) and severe water-stress (SS), moderate water-stress (MS) and well-watered (WW) conditions for 2-week from 4-leaf stage. Dry biomass (DM), leaf rolling performance, water use efficiency (WUE), N fertilizer use efficiency (NUE), root morphology and distribution were evaluated. It was showed that soil water levels and N rates had significant individual and interactive effects on most measured parameters. The dual stress of drought and N deficiency severely limited maize growth and N uptake. Nitrogen fertilization improved maize growth, N uptake and WUE under water-stress. However, compared with HN treatment, LN treatment had less water consumption, higher leaf relative water content and less leaf rolling symptom both under the MS and SS conditions. The reason is that LN enhanced root growth and elongation with more fine roots, especially in deep soil. Notably, maize plants in MS-LN treatment had an optimal root distribution characterized by higher root length density (0.30 cm cm−3), larger and deeper penetration scale throughout the soil layers, and thus showed fewer drought responses and obtained the highest WUE (4.1 g L−1) and NUE (17.2%) among all the water-stress treatments. In contrast, HN limited root growth and extension in deep soil and in turn increased water and N depletion, resulting in more serve leaf rolling and lower WUE and NUE. Therefore, reducing basal N rate is recommended to optimize root growth, morphology and distribution at the seedling stage in rain-fed maize production, to enhance drought resistance and improve WUE and NUE.

Maize (Zea mays L.) has undergone profound changes in root anatomy for environmental adaptation during domestication. However, the genetic mechanism of plasticity of maize root anatomy during the domestication process remains unclear. In this study, high-resolution mapping was performed for nine root anatomical traits using a maize-teosinte population (mexicana × Mo17) across three environments. Large genetic variations were detected for different root anatomical traits. The cortex, stele, aerenchyma areas, xylem vessel number, and cortical cell number had large variations across three environments, indicating high plasticity. Sixteen quantitative trait loci (QTL) were identified, including seven QTL with QTL × environment interaction (EIQTL) for high plasticity traits and nine QTL without QTL × environment interaction (SQTL). Most of the root loci were consistent with shoot QTL depicting domestication signals. Combining transcriptome and genome-wide association studies revealed that AUXIN EFFLUX CARRIER PIN–FORMED LIKE 4 (ZmPILS4) serves as a candidate gene underlying a major QTL of xylem traits. The near-isogenic lines (NILs) with lower expression of ZmPILS4 had 18–24% more auxin concentration in the root tips and 8–15% more xylem vessels. Nucleotide diversity values analysis in the promoter region suggested that ZmPILS4 was involved in maize domestication and adaptation. These results revealed the potential genetic basis of root anatomical plasticity during domestication.

High yield and efficient use of nitrogen is one of the major target for maize breeding worldwide. Both yield potential and nitrogen use efficiency (NUE) of maize is lower in China compared to that in the US. In this study, we analyzed the change of the plant traits related to high yield and NUE of maize cultivars registered and planted in China and US during the past 40 decades. Compared to the cultivar released in the US, Chinese cultivar are less tolerant to high planting density, higher ratio of ear height to plant height, low grain filling rate, more stay-green and longer growth period. We suggest an ideal plant architecture of maize for high yield and NUE. To achieve high yield and high NUE under intensive production conditions (medium and high nitrogen input, high density planting), an ideotype plant architecture is proposed which includes compact plant architecture, low ratio of ear height to plant height (around 0.36–0.38, close to the golden rate.), medium ear, small tassel, medium root size, one-layer brace root, half stay-green, efficient N remobilization, fast grain filling rate etc. The parameters can be utilized to guide future breeding.

As a primary food crop, maize is widely grown around the world. However, the deficiency of essential amino acids, such as lysine, tryptophan, and methionine, results in poor nutritional quality of maize. In addition, the protein concentration of maize declines with the increase in yield, which further reduces the nutritional quality. Here, the photosynthesis of leaves, grain amino acid composition, and stoichiometry of N and S are explored. The results show that N and S maintained the redox balance by increasing the content of glutathione in maize leaves, thereby enhancing the photosynthetic rate and maize yield. Simultaneously, the synergy of N and S increased the grain protein concentration and promoted amino acid balance by increasing the cysteine concentration in maize grains. The maize yield, grain protein concentration, and concentration of essential amino acids, such as lysine, tryptophan, and methionine, could be simultaneously increased in the N:S ratio range of 11.0 to 12.0. Overall, the synergy of N and S simultaneously improved the maize yield and nutritional quality by regulating the redox balance of maize leaves and the amino acids balance of grains, which provides a new theoretical basis and practical method for sustainable production of maize.

The dynamic interactions between soil, weather and crop management have considerable influences on crop yield within a region, and should be considered in optimizing nitrogen (N) management. The objectives of this study were to determine the influence of soil type, weather conditions and planting density on economic optimal N rate (EONR), and to evaluate the potential benefits of site-specific N management strategies for maize production. The experiments were conducted in two soil types (black and aeolian sandy soils) from 2015 to 2017, involving different N rates (0 to 300 kg ha&minus, 1) with three planting densities (55,000, 70,000, and 85,000 plant ha&minus, 1) in Northeast China. The results showed that the average EONR was higher in black soil (265 kg ha&minus, 1) than in aeolian sandy soil (186 kg ha&minus, 1). Conversely, EONR showed higher variability in aeolian sandy soil (coefficient of variation (CV) = 30%) than in black soil (CV = 10%) across different weather conditions and planting densities. Compared with farmer N rate (FNR), applying soil-specific EONR (SS-EONR), soil- and year-specific EONR (SYS-EONR) and soil-, year-, and planting density-specific EONR (SYDS-EONR) would significantly reduce N rate by 25%, 30% and 38%, increase net return (NR) by 155 $ ha&minus, 1, 176 $ ha&minus, 1, and 163 $ ha&minus, 1, and improve N use efficiency (NUE) by 37&ndash, 42%, 52%, and 67&ndash, 71% across site-years, respectively. Compared with regional optimal N rate (RONR), applying SS-EONR, SYS-EONR and SYDS-EONR would significantly reduce N application rate by 6%, 12%, and 22%, while increasing NUE by 7&ndash, 8%, 16&ndash, 19% and 28&ndash, 34% without significantly affecting yield or NR, respectively. It is concluded that soil-specific N management has the potential to improve maize NUE compared with both farmer practice and regional optimal N management in Northeast China, especially when each year&rsquo, s weather condition and planting density information is also considered. More studies are needed to develop practical in-season soil (site)-specific N management strategies using crop sensing and modeling technologies to better account for soil, weather and planting density variation under diverse on-farm conditions.

Modified gibberellin (GA) signaling leads to semi-dwarfism with low nitrogen (N) use efficiency (NUE) in crops. An understanding of GA-mediated N uptake is essential for the development of crops with improved NUE. The function of GA in modulating N uptake capacity and nitrate (NO3&minus, ) transporters (NRTs) was analyzed in the GA synthesis-deficient mutant zmga3ox grown under low (LN) and sufficient (SN) N conditions. LN significantly suppressed the production of GA1, GA3, and GA4, and the zmga3ox plants showed more sensitivity in shoots as well as LN stress. Moreover, the higher anthocyanin accumulation and the decrease of chlorophyll content were also recorded. The net NO3&minus, fluxes and 15N content were decreased in zmga3ox plants under both LN and SN conditions. Exogenous GA3 could restore the NO3&minus, uptake in zmga3ox plants, but uniconazole repressed NO3&minus, uptake. Moreover, the transcript levels of ZmNRT2.1/2.2 were downregulated in zmga3ox plants, while the GA3 application enhanced the expression level. Furthermore, the RNA-seq analyses identified several transcription factors that are involved in the GA-mediated transcriptional operation of NRTs related genes. These findings revealed that GAs influenced N uptake involved in the transcriptional regulation of NRTs and physiological responses in maize responding to nitrogen supply.

Doxorubicin hydrochloride (DOX) is an effective anthracycline anticancer drug. However, the exceeded taken up could induce several side-effects such as cardiotoxicity, alopecia. Therefore, the level of DOX needs to be closely monitored to avoid the occurrence of its side-effects. Herein, we report a novel core CuInSe

To test the hypothesis that plants could remobilize more nutrients from vegetative organs to fulfill the requirement of grain formation under high planting density, eight maize cultivars with different leaf senescence characters were chosen to study the relationship between stay-green character, nutrient (N, P, K) accumulation, post-silking nutrient remobilization and grain yield under three planting densities (60,000, 75,000, and 90,000 plants per ha) in 2 years. We found that increased plant density aggravated leaf senescence, reduced concentration of N, P and K in all organs, reduced post-silking N, P, K accumulation per ha, increased nutrient remobilized efficiency. Stay-green degree was positively correlated to yield reduction at high planting density (R2 = 0.35, P 2 = 0.29, P 2 = 0.51, P 2 = 0.17, P

Accurate leaf area simulation is critical for the performance of crop growth models. Area of fully expanded individual leaves of maize hybrids released before 1995 (defined as old hybrids) has been simulated using a bell-shaped function (BSF) and the relationship between its parameters and total leaf number (TLNO). However, modern high-yielding maize hybrids show different canopy architectures. The function parameters calibrated for old hybrids will not accurately represent modern hybrids. In this study, we evaluated these functions using a dataset including old and modern hybrids that have been widely planted in China in recent years. Maximum individual leaf area (Y0) and corresponding leaf position (X0) were not predicted well by TLNO (R2 = 0.56 and R2 = 0.70) for modern hybrids. Using recalibrated shape parameters a and b with values of Y0 and X0 for modern hybrids, the BSF accurately predicted individual leaf area (R2 = 0.95–0.99) and total leaf area of modern hybrids (R2 = 0.98). The results show that the BSF is still a robust way to predict the fully expanded leaf area of maize when parameters a and b are modified and Y0 and X0 are fitted. Breeding programs have led to increases in TLNO of maize but have not altered Y0 and X0, reducing the correlation between Y0, X0, and TLNO. For modern hybrids, the values of Y0 and X0 are hybrid-specific. Modern hybrids tend to have less-negative values of parameter a and more-positive values of parameter b in the leaf profile. Growth conditions, such as plant density and environmental conditions, also affect the fully expanded leaf area but were not considered in the original published equations. Thus, further research is needed to accurately estimate values of Y0 and X0 of individual modern hybrids to improve simulation of maize leaf area in crop growth models. Keywords: Hybrids, Leaf area, Leaf number, Plant density, Environmental conditions

Remobilization of leaf nitrogen (N) contributes greatly to grain N in maize, but leads to low photosynthetic rate (Pn). Pn is determined by various N components involving in light harvest and CO2 reduction. However, it is less clear which N component is the major contributor for the reduction of photosynthesis in modern stay-green maize hybrids. In this study, we analyzed the relationship between remobilization of different N components and Pn during grain filling stage under low N (no N application) and high N (180 kg N ha−1) in a field experiment. The remobilization efficiency of photosynthetic enzymes (PEPc, PPDK and Rubisco) in the leaf was much higher than that of thylakoid N and other N components. Low N supply increased the remobilization efficiency of all the leaf N components. During grain filling stage, the amount of all the N components decreased together with Pn. The ratio of Pn to the N in the PEPc, PPDK and Rubisco kept increase in the whole grain filling stage, while the ratio of Pn to chlorophyll and thylakoid-N decreased. Correlation analysis indicated that Pn was more related to the content of photosynthetic enzymes than to chlorophyll and thylakoid N. It is concluded that photosynthetic enzymes serve as an N storage reservoir at early grain filling stage and their degradation is critical in the reduction of Pn during later grain filling stage. Future breeding targets may be focused on enhancing the efficiency of photosynthetic enzymes during late grain filling stage.

This paper aims to investigate the adaptation of maize root system architecture (RSA) in response to increasing planting densities. A three-year field study was conducted with three planting densities (40,000, 70,000, and 90,000 plants per ha, which are abbreviated as D40000, D70000 and D90000, respectively). The dynamic change of root morphological traits and the 3-dimensional RSA were quantified. The grain yield per ha increased with increasing plant density from D40000 to D70000, and then decreased at D90000. Compared to D70000, high planting density of D90000 did not changed the total root biomass per ha but increased shoot biomass per ha by 4 to 8% in two of the three experimental years. Grain yield per plant and plant NPK concentration decreased with increasing planting density. Total accumulation of P and K per ha also decreased at D90000 compared to D70000. Root to shoot ratio was reduced at high planting density beginning 50 days after emergence. Compared to the control (D70000), total root length (TRL) per plant was reduced by 18 to 30% at D90000 and increased by 43 to 56% at D40000, root biomass per plant was reduced by 23 to 34% at D90000 and increased by 66 to 75% at D40000. High plant density reduced the number of nodal roots, lateral root density (LRD) and the average lateral root (LR) length, but with less effect on the length of axial roots. The RSA is characteristic of “intra-row contraction and inter-row extension”. Vertically, root growth in top soil layer (0- to 36- cm) was enhanced under supra-optimal plant density, but had a negligible effect in deep soil layers (36- to 60- cm). To adapt to the limited photosynthesis capacity in the roots under high planting density, maize plants tend to reduce nodal root number and inhibit lateral root growth. They maintain nodal root length to explore a larger soil space, and adjust root growth in the intra-row and inter-row direction to avoid root-to-root competition.

Although nitrate represents an important nitrogen (N) source for maize, a major crop of dryland areas, the molecular mechanisms of nitrate uptake and assimilation remain poorly understood. Here, we identified nine maize NIN-like protein (ZmNLP) genes and analyzed the function of one member, ZmNLP3.1, in nitrate nutrition and signaling. The NLP family genes were clustered into three clades in a phylogenic tree. Comparative genomic analysis showed that most ZmNLP genes had collinear relationships to the corresponding NLPs in rice, and that the expansion of the ZmNLP family resulted from segmental duplications in the maize genome. Quantitative PCR analysis revealed the expression of ZmNLP2.1, ZmNLP2.2, ZmNLP3.1, ZmNLP3.2, ZmNLP3.3, and ZmNLP3.4 was induced by nitrate in maize roots. The function of ZmNLP3.1 was investigated by overexpressing it in the Arabidopsis nlp7-1 mutant, which is defective in the AtNLP7 gene for nitrate signaling and assimilation. Ectopic expression of ZmNLP3.1 restored the N-deficient phenotypes of nlp7-1 under nitrate-replete conditions in terms of shoot biomass, root morphology and nitrate assimilation. Furthermore, the nitrate induction of NRT2.1, NIA1, and NiR1 gene expression was recovered in the 35S::ZmNLP3.1/nlp7-1 transgenic lines, indicating that ZmNLP3.1 plays essential roles in nitrate signaling. Taken together, these results suggest that ZmNLP3.1 plays an essential role in regulating nitrate signaling and assimilation processes, and represents a valuable candidate for developing transgenic maize cultivars with high N-use efficiency.

One strategy for plants to adapt to low nitrogen is to reduce shoot growth and allocate more nitrogen and carbon for root growth. The mechanism underlying the response of leaf growth to low-nitrogen remains unknown. In this study, we investigated cell division and elongation in maize leaf growth in response to low-nitrogen by using an integrated approach. Kinematic analysis revealed that low-nitrogen inhibited leaf elongation mainly by shortening length of division zone, reducing cell flux and elemental growth rate. Hormone analysis revealed that changes of gibberellins caused by low-nitrogen correlate with the observed changes in division zone size and elemental growth rate in response to low-nitrogen, suggesting role of gibberellin in low-nitrogen induced inhibition of leaf elongation. RNA-Seq identified that GA20ox4 (GRMZM2G060940), a key enzyme for synthesis of gibberellins, was down-regulated in both division and elongation zone of leaf under low-nitrogen supply. Furthermore, exogenous GA3 application on low-nitrogen plants restored leaf growth. However, application of gibberellin biosynthesis inhibitor reduced leaf growth. It is concluded that low-nitrogen reduces cell flux and elemental growth rate in maize leaf via reducing gibberellin synthesis. As a result, leaf elongation rate was slower and leaf area was smaller, freeing nitrogen for export to roots.

Root system architecture (RSA) plays an important role in the acquisition of mineral nutrients. Nevertheless, RSA has seldom been selected as an important agricultural trait in conventional breeding programs. Little is known about the response of RSA and phosphorus use efficiency (PUE) in regards to phosphorus (P) availability between parental inbred lines and their F1. In this study, 6 maize inbred lines and their 15 F1 generated by a diallel mating system, were used to analyze the genetic associations between RSA and PUE. Heterosis for PUE-related traits were comparatively greater under high P condition and reduced significantly under low P condition. Relative mid-parent heterosis for root traits were expressed more under the low P condition. Low P supply had a significant effect on heterosis, GCA and SCA of RSA- and PUE- related traits. The hybrid C3 (7922 × 8703-2), which had the highest PUE, showed an average yield with a lower P uptake under the both P conditions. Results from this study suggested breeding for a relatively high grain yield with reducing aboveground P demand and grain P concentration should be sufficient to reduce P fertilizer input and improve P efficiency.

In order to understand the regulation of source-sink relationship on leaf senescence in maize, we investigated the effect of pollination prevention on leaf senescence, post-silking dry matter and nitrogen accumulation and remobilization in different vegetative organs in one landrace and ten representative maize hybrids released between 1973 and 2000 in China. When pollination was prevented by covering silk with paper bags at silking stage, leaf senescence of Nongda 60, Yedan 13, Shendan 7 and Zhengdan 958 was delayed as shown by the remaining green leaf area per plant at maturity. However, pollination-prevention did not have obvious effect on the leaf senescence of Baimaya, Danyu 6, Yedan 2 and Xianyu 335. By contrast, pollination-prevention accelerated the leaf senescence of maize hybrids Zhongdan 2, Danyu13 and Yedan 4. It was found that leaf senescence of the late-released (since 1985) Chinese hybrids tended to be delayed by pollination-prevention, except XY335. From silking to physiological maturity, nitrogen content increased in the stem (plus the sheath, cob, husk, and tassel) and root of the non-pollinated plants. However, there was still a net reduction in leaf nitrogen content. We also found that pollination-prevention reduced leaf nitrogen remobilization efficiency, with genotypic difference and variation between the two years of testing. The results suggested that the response of leaf senescence to pollination-prevention is at least partially due to the change of leaf nitrogen remobilization efficiency. Leaf senescence tended to be delayed if leaf nitrogen remobilization efficiency is highly reduced by pollination-prevention.