Mapping QTLs for root traits under different nitrate levels at the seedling stage in maize (Zea mays L.)

Nitrogen (N) loss is a worldwide problem in crop production. Apart from reasonable N fertilizer application, breeding N efficient cultivars provides an alternative way. Root architecture is an important factor determining N acquisition. However, little is known about the molecular genetic basis for root growth in relation to N supply. In the present study, an F₈ maize (Zea may L.) recombinant inbred (RI) population consisting of 94 lines was used to identify the QTLs for root traits under different nitrate levels. The lateral root length (LRL), axial root length (ARL), maximal axial root length (MARL), axial root number (ARN) and average axial root length (AARL) were evaluated under low N (LN) and high N (HN) conditions in a hydroponics system. A total of 17 QTLs were detected among which 14 loci are located on the same chromosome region as published QTLs for root traits. A major QTL on chromosome 1 (between bnlg1025 and umc2029) for the AARL under LN could explain 43.7% of the phenotypic variation. This QTL co-localizes with previously reported QTLs that associate with root traits, grain yield, and N uptake. Our results indicate that longer axial roots are important for efficient N acquisition and the major QTL for AARL may be used as a marker in breeding N efficient maize genotypes.     

Application of NMR-based metabolomics to the investigation of salt stress in maize (Zea mays)

Introduction – High salinity, caused by either natural (e.g. climatic changes) or anthropic factors (e.g. agriculture), is a widespread environmental stressor that can affect development and growth of salt‐sensitive plants, leading to water deficit, the inhibition of intake of essential ions and metabolic disorders. Objective – The application of an NMR‐based metabolic profiling approach to the investigation of saline‐induced stress in Maize plants is presented. Methodology – Zea Maize seedlings were grown in either 0, 50 or 150 mM saline solution. Plants were harvested after 2, 4 and 6 days (n = 5 per class and time point) and ¹H NMR spectroscopy was performed separately on shoot and root extracts. Spectral data were analysed and interpreted using multivariate statistical analyses. Results – A distinct effect of time/growth was observed for the control group with relatively higher concentrations of acetoacetate at day 2 and increased levels of alanine at days 4 and 6 in root extracts, whereas concentration of alanine was positively correlated with the shoot extracts harvested at day 2 and trans‐aconitic acid increased at days 4 and 6. A clear dose‐dependent effect, superimposed on the growth effect, was observed for saline treated shoot and root extracts. This was correlated with increased levels of alanine, glutamate, asparagine, glycine‐betaine and sucrose and decreased levels of malic acid, trans‐aconitic acid and glucose in shoots. Correlation with salt‐load shown in roots included elevated levels of alanine, γ‐amino‐N‐butyric acid, malic acid, succinate and sucrose and depleted levels of acetoacetate and glucose. Conclusions – The metabolic effect of high salinity was predominantly consistent with osmotic stress as reported for other plant species and was found to be stronger in the shoots than the roots. Using multivariate data analysis it is possible to investigate the effects of more than one environmental stressor simultaneously. Copyright © 2010 John Wiley & Sons, Ltd.     

Evaluation of oxidative stress tolerance in maize (Zea mays L.) seedlings in response to drought

Seedlings of selected six genotypes of maize (Zea mays L.) differing in their drought sensitivity (LM5 and Parkash drought-tolerant and PMH2, JH3459, Paras and LM14 as drought-sensitive) were exposed to 72 h drought stress at two leaf stage. Alterations in their antioxidant pools combined with activities of enzymes involved in defense against oxidative stress were investigated in leaves. Activities of some reactive oxygen species (ROS)-scavenging enzymes, catalase (CAT) and ascorbate peroxidase (APX) were enhanced in tolerant genotypes in response to drought stress. Superoxide dismutase (SOD) activity was significantly decreased in sensitive genotypes, but remained unchanged in tolerant genotypes under stress. Peroxidase (POX) activity was significantly induced in tolerant, as well as sensitive genotypes. Imposition of stress led to increase in H2O2 and malondialdehyde (MDA, a marker for lipid peroxidation) content in sensitive genotypes, while in tolerant genotypes no change was observed. Significant increase in glutathione content was observed in sensitive genotypes. Ascorbic acid pool was induced in both tolerant and sensitive genotypes, but induction was more pronounced in tolerant genotypes. Significant activation of antioxidative defence mechanisms correlated with drought-induced oxidative stress tolerance was the characteristic of the drought tolerant genotypes. These studies provide a mechanism for drought tolerance in maize seedlings.     

Determination of stress responses induced by aluminum in maize (Zea mays)

To assess the alternative responses to aluminum toxicity, maize (Zea mays L. cv Karadeniz y?ld?z?) roots were exposed to different concentrations of AlCl3 (150, 300 and 450 μM). Aluminum reduced the root elongation by 39.6% in 150 μM, 44.1% in 300 μM, 50.1% in 450 μM AlCl3 after 96 h period. To correlate the root elongation with the alternative stress responses including aluminum accumulation, lipid peroxidation, mitotic abnormalities, reduction of starch content, intracellular Ca2+ accumulation, callose formation, lignin deposition and peroxidase activity, cytochemical and biochemical tests were performed. The results indicated that aluminum accumulation and lipid peroxidation were observed more densely on the root cap and the outer cortex cells. In addition to morphological deformations, cytochemical analysis displayed cellular deformations. Furthermore, mitotic abnormalities were observed such as c-mitosis, micronuclei, bi- and trinucleated cells in aluminum treated root tips. Aluminum treatment induced starch reduction, callose formation, lignin accumulation and intracellular Ca2+ increase. Moreover, the peroxidase activity increased significantly by 3, 4.4 and 7.7 times higher than in that of control after 96 h, respectively. In conclusion, aluminum is significantly stressful in maize culminating in morphological and cellular alterations.     

Genotypic variation between maize (Zea mays L.) single cross hybrids in response to drought stress

Effects of soil drought on growth and productivity of 16 single cross maize hybrids were investigated under field and greenhouse experiments. The Drought Susceptibility Index (DSI) was evaluated in a three year field experiment by the determination of grain loss in conditions of two soil moisture levels (drought and irrigated) and in a pot experiment by the effects of periodical soil drought on seedling dry matter. In the greenhouse experiment response to drought in maize genotypes was also evaluated by root to shoot dry mater ratio, transpiration productivity index, indexes of kernel germination and index of leaf injury by drought and heat temperature. The obtained values of DSI enabled the ranking of the tested genotypes with respect to their drought tolerance. The values of DSI obtained in the field experiment allow to divide the examined genotypes into three, and in the greenhouse experiment into two groups of drought susceptibility. The correlation coefficients between the DSI of maize hybrids in the field and the greenhouse experiments was high and statistically significant, being equal to 0.876. The ranking of hybrids drought tolerance, identified on the basis of field experiments was generally in agreement with the ranking established on the basis of the greenhouse experiment. In the greenhouse experiment statistically significant coefficients of correlation with DSI values in hybrids were obtained for the ratio of dry matter of overground parts to dry matter of roots, both for control and drought treatments, whereas in the estimation of the transpiration productivity coefficient and total dry matter the correlation coefficients were not statistically significant. In this study several laboratory tests were carried out for the drought tolerance of plants (kernel germination, leaf injury) on 4 drought resistant and 4 drought sensitive maize hybrids. Statistically significant correlation coefficients between DSI and the examined parameter of grain germination and leaf injury were obtained for the determination of promptness index (PI), seedling survival index (SS) and leaf injuries indexes (IDS, ITS) as a result of exposure to 14 days of soil drought, osmotic drought −0.9 MPa and exposure to high temperature 45 ° or 50 °C. The results of laboratory tests show that in maize the genetic variation in the degree of drought tolerance is better manifested under severe conditions of water deficit in the soil.     

Genomic prediction of seedling root length in maize (Zea mays L.)

Genotypes with extreme phenotypes are valuable for studying ‘difficult’ quantitative traits. Genomic prediction (GP) might allow the identification of such extremes by phenotyping a training population of limited size and predicting genotypes with extreme phenotypes in large sequences of germplasm collections. We tested this approach employing seedling root traits in maize and the extensively genotyped Ames Panel. A training population made up of 384 inbred lines from the Ames Panel was phenotyped by extracting root traits from images using the software program aria . A ridge regression best linear unbiased prediction strategy was used to train a GP model. Genomic estimated breeding values for the trait ‘total root length’ (TRL) were predicted for 2431 inbred lines, which had previously been genotyped by sequencing. Selections were made for 100 extreme TRL lines and those with the predicted longest or shortest TRL were validated for TRL and other root traits. The two predicted extreme groups with regard to TRL were significantly different (P = 0.0001). The difference in predicted means for TRL between groups was 145.1 cm and 118.7 cm for observed means, which were significantly different (P = 0.001). The accuracy of predicting the rank between 1 and 200 of the validation population based on TRL (longest to shortest) was determined using a Spearman correlation to be ρ = 0.55. Taken together, our results support the idea that GP may be a useful approach for identifying the most informative genotypes in sequenced germplasm collections to facilitate experiments for quantitative inherited traits.     

Genome-wide association analysis of seedling root development in maize (Zea mays L.)

Background

Plants rely on the root system for anchorage to the ground and the acquisition and absorption of nutrients critical to sustaining productivity. A genome wide association analysis enables one to analyze allelic diversity of complex traits and identify superior alleles. 384 inbred lines from the Ames panel were genotyped with 681,257 single nucleotide polymorphism markers using Genotyping-by-Sequencing technology and 22 seedling root architecture traits were phenotyped.

Results

Utilizing both a general linear model and mixed linear model, a GWAS study was conducted identifying 268 marker trait associations (p ≤ 5.3×10^-7). Analysis of significant SNP markers for multiple traits showed that several were located within gene models with some SNP markers localized within regions of previously identified root quantitative trait loci. Gene model GRMZM2G153722 located on chromosome 4 contained nine significant markers. This predicted gene is expressed in roots and shoots.

Conclusion

This study identifies putatively associated SNP markers associated with root traits at the seedling stage. Some SNPs were located within or near (<1 kb) gene models. These gene models identify possible candidate genes involved in root development at the seedling stage. These and respective linked or functional markers could be targets for breeders for marker assisted selection of seedling root traits.

Electronic supplementary material

The online version of this article (doi:10.1186/s12864-015-1226-9) contains supplementary material, which is available to authorized users.     

弱光胁迫影响夏玉米光合效率的生理机制初探

大田条件下, 以普通夏玉米(Zea mays) ‘泰玉2号’为材料, 于授粉后1-20天遮光55% (+S), 以大田自然光照条件下生长的玉米作为对照(-S), 研究了遮光及恢复过程中玉米植株的光合性能、叶绿体荧光参数、叶黄素循环以及光能分配的变化, 初步揭示夏玉米开花后弱光条件下光适应的生理机制, 为玉米高产稳产提供理论依据。结果表明, 遮光后玉米穗位叶叶绿素含量及可溶性蛋白含量均减少, RuBP羧化酶和PEP羧化酶活性显著降低, 导致穗位叶净光合速率(P_n)迅速下降, 光饱和点也明显降低; 恢复初期P_n迅速升高, 光合关键酶活性有所增强。遮光后植株的最大光化学效率(F_v/F_m)、实际光化学效率(Ф_PSII)显著降低, 非光化学淬灭(NPQ)则显著升高, 而恢复初期植株穗位叶Ф_PSII有所升高, 表明突然暴露在自然光下的光合电子传递速率明显加快, 这与其光合速率及光合酶活性的趋势保持一致; 遮光处理对穗位叶叶黄素循环库的大小(紫黄质+花药黄质+玉米黄质(V + A + Z))影响不显著, 但使叶黄素循环的脱环氧化状态(A + Z)/(V + A + Z)增加; 遮光后植株分配于光化学反应的光能明显减少, 天线耗散光能比率显著增加, 恢复过程中植株主要以过剩非光化学反应的形式耗散过剩的光能。遮光后及恢复初期, 玉米植株的PSII原初光化学活性明显下降, 限制了光合碳代谢的电子供应从而抑制了光合作用, 主要依赖叶黄素循环途径进行能量耗散, 而在光照转换后遮光的玉米叶片在适应自然光过程中的光保护机制不断完善, 光合能力逐渐得到 恢复。     

Zinc deficiency and pollen fertility in maize (Zea mays)

Zinc deficiency decreased pollen viability in maize (Zea mays L. cv. G2) grown in sand culture. On restoring normal zinc supply to zinc-deficient plants before the pollen mother cell stage of anther development, the vegetative yield of plants and pollen fertility could be recovered to a large extent, but the recovery treatment was not effective when given after the release of microspores from the tetrads. If zinc deficiency was induced prior to microsporogenesis it did not significantly affect vegetative yield and ovule fertility, but decreased the fertility of pollen grains, even of those which visibly appeared normal. If the deficiency was induced after the release of microspores from the tetrads, not only vegetative yield and ovule fertility but pollen fertility also remained unaffected.     

Brassinolide interacts with auxin and ethylene in the root gravitropic response of maize (Zea mays)

This study examines how brassinolide (BL) and ethylene interact in the gravitropic response mechanism of maize ( Zea mays ) primary roots. When applied exogenously, ethylene increases the rate of gravitropic curvature in a dose-dependent manner. This effect of ethylene was confirmed by the fact that AVG, a specific action inhibitor of ACC synthase, reduces the gravitropic curvature in the presence and absence of BL. Since AVG did not inhibit BL-increased gravitropic curvature completely, we investigated the possibility that BL may act on the gravitropic response by ways other than simply through enhanced ethylene production. We show that BL exhibits some of its stimulatory effect in the absence of ethylene. In addition, BL reduces the presentation time and lag period for the gravitropic response, whereas ethylene increases them. One possible mechanism of such action is that BL affects protein kinase activity, since the protein kinase inhibitors, staurosporine and H89, reduce BL-increased gravitropic curvature. In summary, BL is involved in the gravitropic response in maize primary roots via ethylene production, but it acts in a way that differs somewhat from that of ethylene.     

Mapping the anther culture response genes in maize (Zea mays L.).

In order to map the genes conditioning the induction of embryos during our anther culture process, we evaluated F2 plants from three different crosses for their anther culture ability and also performed RFLP analysis on these plants. The results showed that six chromosomal regions appear to be associated with the ability to induce embryo-like structures from maize microspores. These regions are located on chromosomes 1 (two regions), 3, 5, 7, and 8. Some of these chromosomes are identical to those found in previous studies and we have localized the regions more precisely. Notably, all chromosome regions identified, except one, are near viviparous mutant loci. Since the viviparous mutations are known to involve the plant hormone abscisic acid (ABA), these results suggest that ABA or its antagonist, gibberellic acid (GA3), might somehow be related to anther culture ability. We also propose some combinations of probes to screen for anther culture ability in the three genotypes studied.     

The response of maize (Zea mays) to Azospirillum inoculation in various types of soils in the field

Application of a peat-based powder inoculant of Azospirillum brasilense, as well as a granular inoculant (each containing 0.5–1.0×10⁷ Azospirillum/g moist peat), in the seed furrows of Zea mays resulted in significantly increased yields (11 to 14%) in light soils at low rates of N fertilization. In general, there was no effect of inoculation on plant yields in heavier soils nor when N fertilization was high. Pre-emergence application of granular inoculant and inoculation associated with irrigation were more efficient in increasing yield than inoculation post-emergence or seed coating.E. Fallik is with ARO-The Volcani Centre, Department of Postharvest Science of Fresh Produce, Bet-Dagan 50250, Israel. Y. Okon is with The Hebrew University of Jerusalem, Faculty of Agriculture, Department of Plant Pathology and Microbiology, Rehovot 76100, Israel     

基于分期播种的气候变化对东北地区玉米(Zea mays)生长发育和产量的影响

利用在东北地区中部开展玉米(Zea mays)分期播种试验资料,分析气候变化对玉米出苗速度、生长发育速度、灌浆过程、植株干物质积累和产量的影响,进而分析未来气候变化对东北地区玉米生产的影响及适应性对策.结果表明,气候变化对玉米生长发育和产量的影响十分明显,在水分基本适宜的条件下,东北地区气候变暖导致玉米生长季气温升高、积温增加,使玉米生长发育和灌浆速度加快,生物量增加,从而提高单产.但气候变暖的同时,气候变干会限制热量资源的利用,将缩短玉米灌浆时间,降低灌浆速率,使千粒重下降,从而造成明显减产,而且减产幅度明显大于温度升高的增产幅度.在水分条件基本得到满足的条件下,未来夏半年气候变暖对东北地区玉米生产是有利的,偏晚熟玉米品种比例可以适当扩大,东北玉米带可以向北部和东部扩展,单产和总产都会增加;但如果水分得不到满足,气候的暖干化趋势会使东北地区的中、西部玉米主产区的农业干旱变得更加严重且频繁,造成产量下降和不稳定,给玉米生产带来严重威胁,因而更应加强农业干旱的综合防御工作.