Quantitative Trait Loci Mapping of Maize Yield and Its Components Under Different Water Treatments at Flowering Time

Drought or water stress is a serious agronomic problem resulting in maize (Zea mays L.) yield loss throughout the world. Breeding hybrids with drought tolerance is one important approach for solving this problem. However, lower efficiency and a longer period of breeding hybrids are disadvantages of traditional breeding programs. It is generally recognized that applying molecular marker techniques to traditional breeding programs could improve the efficiency of the breeding of drought‐tolerant maize. To provide useful information for use in studies of maize drought tolerance, the mapping and tagging of quantitative trait loci (QTL) for yield and its components were performed in the present study on the basis of the principle of a mixed linear model. Two hundred and twenty‐one recombinant inbred lines (RIL) of Yuyu 22 were grown under both well‐watered and water‐stressed conditions. In the former treatment group, plants were well irrigated, whereas those in the latter treatment group were stressed at flowering time. Ten plants of each genotype were grown in a row that was 3.00 m × 0.67 m (length × width). The results show that a few of the QTL were the same (one additive QTL for ear length, two additive QTL and one pair of epistatic QTL for kernel number per row, one additive QTL for kernel weight per plant), whereas most of other QTL were different between the two different water treatment groups. It may be that genetic expression differs under the two different water conditions. Furthermore, differences in the additive and epistatic QTL among the traits under water‐stressed conditions indicate that genetic expression also differs from trait to trait. Major and minor QTL were detected for the traits, except for kernel number per row, underwater‐stressed conditions. Thus, the genetic mechanism of drought tolerance in maize is complex because the additive and epistatic QTL exist at the same time and the major and minor QTL all contribute to phenotype under water‐stressed conditions. In particular, epidemic QTL under water‐stressed conditions suggest that it is important to investigate the drought tolerance of maize from a genetic viewpoint. (Managing editor: Wei Wang)     

Characterizing drought stress and trait influence on maize yield under current and future conditions

Global climate change is predicted to increase temperatures, alter geographical patterns of rainfall and increase the frequency of extreme climatic events. Such changes are likely to alter the timing and magnitude of drought stresses experienced by crops. This study used new developments in the classification of crop water stress to first characterize the typology and frequency of drought‐stress patterns experienced by European maize crops and their associated distributions of grain yield, and second determine the influence of the breeding traits anthesis‐silking synchrony, maturity and kernel number on yield in different drought‐stress scenarios, under current and future climates. Under historical conditions, a low‐stress scenario occurred most frequently (ca. 40%), and three other stress types exposing crops to late‐season stresses each occurred in ca. 20% of cases. A key revelation shown was that the four patterns will also be the most dominant stress patterns under 2050 conditions. Future frequencies of low drought stress were reduced by ca. 15%, and those of severe water deficit during grain filling increased from 18% to 25%. Despite this, effects of elevated CO₂ on crop growth moderated detrimental effects of climate change on yield. Increasing anthesis‐silking synchrony had the greatest effect on yield in low drought‐stress seasonal patterns, whereas earlier maturity had the greatest effect in crops exposed to severe early‐terminal drought stress. Segregating drought‐stress patterns into key groups allowed greater insight into the effects of trait perturbation on crop yield under different weather conditions. We demonstrate that for crops exposed to the same drought‐stress pattern, trait perturbation under current climates will have a similar impact on yield as that expected in future, even though the frequencies of severe drought stress will increase in future. These results have important ramifications for breeding of maize and have implications for studies examining genetic and physiological crop responses to environmental stresses.     

Effect of Drought and Weed Management on Maize Genotypes and the Tensiometric Soil Water Content of an Eutric Nitisol in South Western Nigeria

In the dry savannas of West and Central Africa, where low soil fertility, unpredictable rainfall, weed competition and recurrent drought are major constraints to maize production, the development of tropical maize genotypes with high and stable yields under drought and low-nitrogen condition is very important since access to these improved genotypes may be the only affordable alternative to many small scale farmers. Field trials were conducted in 2002 and 2003 at Ikenne southwestern Nigeria to investigate the effect of weed pressures and drought stress on 2 maize (Zea mays L.) hybrids (9134-14, 9803-9) and 2 open-pollinated varieties (STREVIWD, IYFDCO1). Irrigation was withdrawn 4 weeks after planting (about four weeks to mid-flowering) in the drought stress while the adjacent watered treatment had irrigation throughout the growing period. The weed pressures were the completely weeded plots (hand weeding every week) and weedy plots (weeded once, 2 weeks after planting). The experiment was a split plot in a randomized complete block design with four replicates. Drought stress reduced the stover weight and grain yield of the maize cultivars by 6% and 34% respectively. Weed-free plots had maize with higher agronomic traits than unweeded treatments. Hybrid 9803-9 was more susceptible to drought and weed stress as indicated in the stover weight and grain yield. STREVIWD an open-pollinated variety (OPV) and Hybrid 9134-14 had superior performances in terms of grain yield and shorter anthesis silking interval. Soil moisture content was higher in the unweeded plots while the uptake of moisture was highest in drought susceptible hybrid 9803-9. Irrespective of the genotypes, maize (hybrid and OPV) was more tolerant to drought in a weed-free environment than in unweeded conditions. There existed a negative but significant correlation between weed biomass and chlorophyll content (−0.29, P < 0.01), grain yield (−0.45, P < 0.05), ear plant⁻¹ (−0.27, P < 0.05) and kernel-number (−0.366 P < 0.01).     

鱼和熊掌的选择:反向重复序列变异介导的玉米环境适应与产量平衡

作物育种的目标是找到产量和抗性的最佳平衡点,其中涉及“鱼和熊掌”二者兼得的选择策略。哪些逆境负调控位点影响产量性状,以及如何调控等是突破育种瓶颈的重要科学问题。近百年来,高产玉米(Zeamays)育种使玉米单产不断提高,同时现代玉米品种对干旱的敏感性也呈现出增强趋势,故而存在高产稳产的潜在风险。可对于这一现象背后确切的遗传机制却知之甚少,从而限制了既高产又高抗玉米新品种的培育。玉米的非生物胁迫抗性与产量性状均为多基因控制的复杂数量性状,涉及全基因组范围内大量基因的表达与调控。玉米基因组内存在大量的小RNA (sRNA),其对基因表达起精细调控作用,但人们对sRNA调控作物环境胁迫应答与产量性状机制的理解仍然有限。近日,华中农业大学代明球课题组与李林和李峰两个课题组合作,基于对338份玉米关联群体在不同环境下的sRNA表达组分析,鉴定到大量干旱应答的sRNA,以及调控这些sRNA表达的遗传位点(eQTL);并克隆了8号染色体上1个干旱特异性eQTL热点DRESH8。生物信息学分析显示, DRESH8是1个由转座子组成的长度约为21.4 kb的反向重复序列(TE-IR)。DRESH8通过产...     

Effects of drought stress on phosphorus and potassium uptake dynamics in summer maize (Zea mays) throughout the growth cycle

The effects of drought stress on the phosphorus (P) and potassium (K) uptake dynamics of summer maize (Zea mays L.) throughout the growth cycle were studied. Field trials were conducted under a completely randomized design with three field water capacity (FC) regimes: 75?% FC was well watered and considered to be the control, 55?% FC represented moderate stress (MS), and 35?% FC represented severe stress (SS). The water regimes were applied from the third leaf stage until maturity. Drought stress induced sharp decreases in total K and P uptake of maize organs at different developmental stages and, in particular, detrimentally affected the nutrient uptake capability of roots. SS caused more deleterious effect than MS on both total K and P uptake by plant organs. The results suggested maize plants differ in their ability to maintain nutrient uptake under drought stress, and it is highly dependent on the intensity and duration of drought stress and the developmental stage. The decrease in total K and P uptake caused by both MS and SS was accompanied by reduction in biomass production in drought-stressed tissues. The biomass allocation patterns in response to drought stress fluctuated strong mostly because of competitive changes in the shoot and roots at different stages, thus the root:shoot ratio increased at some stages and decreased at other stages. SS induced a dramatic reduction in the harvest index (HI), whereas MS slightly decreased HI. Thus, water limitation caused lower K and P uptake and HI.     

土壤水分胁迫对玉米形态发育及产量的影响

未来气候变化可能加剧的干旱化将对我国主要粮食和最重要的饲料作物玉米产生严重影响。为增进玉米对干旱化响应与适应的理解及制定应对策略 ,利用大型活动遮雨棚及池栽对玉米进行了全程水分控制试验研究。对不同土壤水分胁迫下的玉米形态表征、生长发育及产量的分析表明 ,玉米受干旱胁迫的影响程度因受旱轻重、持续时间以及生育进程的不同而不同 ,受旱越重 ,持续时间越长 ,影响越甚。大喇叭口期前 ,玉米株高和生物产量受有限供水或轻度干旱影响不算很大 ,但从大喇叭口期后直至抽雄和灌浆期 ,轻度干旱胁迫持续久了也会对株高和生物产量产生较大不良影响。严重干旱胁迫则从拔节始至灌浆期均对株高和生物产量影响更为不利。进而引起果穗性状恶化 ,穗粒数和百粒重减小 ,最终导致经济产量大幅下降。说明玉米生育前期(大喇叭口期前 )进行有限的控水可行。而玉米生育前期干旱胁迫将使生育进程明显延缓 ,严重干旱胁迫可使抽雄、吐丝期较水分充足滞后 4 d左右 ,并引起成熟期推迟     

Genetic Architecture of Delayed Senescence,Biomass, and Grain Yield under Drought Stress in Cowpea

The stay-green phenomenon is a key plant trait with wide usage in managing crop production under limited water conditions. This trait enhances delayed senescence, biomass, and grain yield under drought stress. In this study we sought to identify QTLs in cowpea (Vigna unguiculata) consistent across experiments conducted in Burkina Faso, Nigeria, Senegal, and the United States of America under limited water conditions. A panel of 383 diverse cowpea accessions and a recombinant inbred line population (RIL) were SNP genotyped using an Illumina 1536 GoldenGate assay. Phenotypic data from thirteen experiments conducted across the four countries were used to identify SNP-trait associations based on linkage disequilibrium association mapping, with bi-parental QTL mapping as a complementary strategy. We identified seven loci, five of which exhibited evidence suggesting pleiotropic effects (stay-green) between delayed senescence, biomass, and grain yield. Further, we provide evidence suggesting the existence of positive pleiotropy in cowpea based on positively correlated mean phenotypic values (0.34< r <0.87) and allele effects (0.07< r <0.86) for delayed senescence and grain yield across three African environments. Three of the five putative stay-green QTLs, Dro-1, 3, and 7 were identified in both RILs and diverse germplasm with resolutions of 3.2 cM or less for each of the three loci, suggesting that these may be valuable targets for marker-assisted breeding in cowpea. Also, the co-location of early vegetative delayed senescence with biomass and grain yield QTLs suggests the possibility of using delayed senescence at the seedling stage as a rapid screening tool for post-flowering drought tolerance in cowpea breeding. BLAST analysis using EST sequences harboring SNPs with the highest associations provided a genomic context for loci identified in this study in closely related common bean (Phaseolus vulgaris) and soybean (Glycine max) reference genomes.     

A unifying theory for general multigenic heterosis: energy efficiency, protein metabolism, and implications for molecular breeding

Hybrids between genetically diverse varieties display enhanced growth, and increased total biomass, stress resistance and grain yield. Gene expression and metabolic studies in maize, rice and other species suggest that protein metabolism plays a role in the growth differences between hybrids and inbreds. Single trait heterosis can be explained by the existing theories of dominance, overdominance and epistasis. General multigenic heterosis is observed in a wide variety of different species and is likely to share a common underlying biological mechanism. This review presents a model to explain differences in growth and yield caused by general multigenic heterosis. The model describes multigenic heterosis in terms of energy-use efficiency and faster cell cycle progression where hybrids have more efficient growth than inbreds because of differences in protein metabolism. The proposed model is consistent with the observed variation of gene expression in different pairs of inbred lines and hybrid offspring as well as growth differences in polyploids and aneuploids. It also suggests an approach to enhance yield gains in both hybrid and inbred crops via the creation of an appropriate computational analysis pipeline coupled to an efficient molecular breeding program.     

Physiological Responses of Contrasting Maize (Zea mays L.) Hybrids to Repeated Drought

Maize is the most important crop worldwide in terms of production and yield, but every year a considerable amount of yield is lost due to drought. The foreseen increase in the number of drought spells due to climate change raises the question whether the ability to recover quickly after a water pulse may be a relevant trait for overall drought resistance. We here address the following hypotheses: (i) different maize hybrids exhibit distinct physiological adaptive responses to drought stress and (ii) these responses affect the ability to recover from the stress. (iii) The relative biomass production of maize hybrids, which show severe drought symptoms but are able to recover quickly after a water pulse, is comparable to those hybrids, which invest more energy into tolerance mechanisms. The physiological responses of eight maize hybrids to repeated drought were elucidated employing physiological parameters such as electrolyte leakage, osmolality, relative water content, growth rate and gas-exchange measurements. Only one hybrid was able to maintain biomass production under drought conditions. Amongst the others, two hybrids with similar growth inhibition but contrasting physiological responses were identified by a PCA analysis. Both strategies, i.e. stabilization of leaf water content via resistance mechanisms versus high recovery potential were equally effective in maintaining aboveground biomass production in the scenario of a long drought intermitted by a water-pulse. However, each strategy might be advantageous under different drought stress scenarios. Overall, the recovery potential is underestimated in drought resistance under natural conditions, which includes periodic cycles of drought and rewatering, and should be considered in screening trials.

     

The TaDREB3 transgene transferred by conventional crossings to different genetic backgrounds of bread wheat improves drought tolerance

Drought tolerance of the wheat cultivar Bobwhite was previously enhanced by transformation with a construct containing the wheat DREB3 gene driven by the stress‐inducible maize Rab17 promoter. Progeny of a single T₂ transgenic line were used as pollinators in crosses with four elite bread wheat cultivars from Western Australia: Bonnie Rock, IGW‐2971, Magenta and Wyalkatchem, with the aim of evaluating transgene performance in different genetic backgrounds. The selected pollinator line, BW8‐9‐10‐3, contained multiple transgene copies, had significantly improved drought tolerance compared with wild‐type plants and showed no growth and development penalties or abnormalities. A single hybrid plant was selected from each cross‐combination for three rounds of backcrossing with the corresponding maternal wheat cultivar. The transgene was detected in all four F₁BC₃ combinations, but stress‐inducible transgene expression was found in only three of the four combinations. Under well‐watered conditions, the phenotypes and grain yield components of the F₂BC₃ transgene‐expressing lines were similar to those of corresponding recurrent parents and null‐segregants. Under severe drought conditions, the backcross lines demonstrated 12–18% higher survival rates than the corresponding control plants. Two from four F₃BC₃ transgenic lines showed significantly higher yield (18.9% and 21.5%) than control plants under limited water conditions. There was no induction of transgene expression under cold stress, and therefore, no improvement of frost tolerance observed in the progenies of drought‐tolerant F₃BC₃ lines.     

Maize leaf temperature responses to drought: Thermal imaging and quantitative trait loci (QTL) mapping

Leaf temperature has been shown to vary when plants are subjected to water stress conditions. Recent advances in infrared thermography have increased the probability of recording drought tolerant responses more accurately. The aims of this study were to identify the effects of drought on leaf temperature using infrared thermography. Furthermore, the genomic regions responsible for the expression of leaf temperature variation in maize seedlings (Zea mays L.) were explored. The maize inbred lines Zong3 and 87-1 were evaluated using infrared thermography and exhibited notable differences in leaf temperature response to water stress. Correlation analysis indicated that leaf temperature response to water stress played an integral role in maize biomass accumulation. Additionally, a mapping population of 187 recombinant inbred lines (RILs) derived from a cross between Zong3 and 87-1 was constructed to identify quantitative trait loci (QTL) responsible for physiological traits associated with seedling water stress. Leaf temperature differences (LTD) and the drought tolerance index (DTI) of shoot fresh weight (SFW) and shoot dry weight (SDW) were the traits evaluated for QTL analysis in maize seedlings. A total of nine QTL were detected by composite interval mapping (CIM) for the three traits (LTD, RSFW and RSDW). Two co-locations responsible for both RSFW and RSDW were detected on chromosomes 1 and 2, respectively, which showed common signs with their trait correlations. Another co-location was detected on chromosome 9 between LTD and shoot biomass, which provided genetic evidence that leaf temperature affects biomass accumulation. Additionally, the utility of a thermography system for drought tolerance breeding in maize was discussed.     

丛枝菌根真菌对杨树生长、气孔和木质部微观结构的影响

植物气孔与木质部导管及纤维的功能直接关系着植物的水分利用, 进而影响植物的生长。为研究丛枝菌根真菌(AMF)对杨树抗旱性的影响, 采用温室盆栽的方法, 研究两种水分条件下, 接种根内球囊霉(Rhizophagus irregularis)对速生杨107 Populus × canadensis (P. nigra × P. deltoides) ‘Neva’气孔及木质部微观结构的影响。结果表明: AMF的侵染显著提高了杨树幼苗地上和地下部分生物量, 对叶片气孔长度、茎部导管细胞直径和纤维细胞长度也有促进作用。AMF对生物量和导管细胞直径的增加幅度表现出干旱条件下>正常水分条件下, 而对气孔长度的提高幅度表现出干旱条件下<正常水分条件下。正常水分条件下, AMF增加了杨树叶片的气孔密度, 减小了纤维细胞直径, 对相对水分饱和亏缺无影响; 干旱条件下, AMF增加了纤维细胞直径, 降低了相对水分饱和亏缺, 对气孔密度无影响。综上所述, 干旱条件下, AMF对导管水分传输能力的促进作用明显增加, 而对气孔蒸腾能力的促进作用有所减少, 从而更利于杨树在遭遇干旱时保持水分, 减少干旱对菌根杨树造成的水分亏缺, 提高菌根杨树对干旱的耐受性。     

Engineering of enhanced glycine betaine synthesis improves drought tolerance in maize

Glycine betaine plays an important role in some plants, including maize, in conditions of abiotic stress, but different maize varieties vary in their capacity to accumulate glycine betaine. An elite maize inbred line DH4866 was transformed with the betA gene from Escherichia coli encoding choline dehydrogenase (EC 1.1.99.1), a key enzyme in the biosynthesis of glycine betaine from choline. The transgenic maize plants accumulated higher levels of glycine betaine and were more tolerant to drought stress than wild-type plants (non-transgenic) at germination and the young seedling stage. Most importantly, the grain yield of transgenic plants was significantly higher than that of wild-type plants after drought treatment. The enhanced glycine betaine accumulation in transgenic maize provides greater protection of the integrity of the cell membrane and greater activity of enzymes compared with wild-type plants in conditions of drought stress.     

Response of wheat canopy CO2 and water gas-exchange to soil water content under ambient and elevated CO2

The nature of the interaction between drought and elevated CO₂ partial pressure (pC_a) is critically important for the effects of global change on crops. Some crop models assume that the relative responses of transpiration and photosynthesis to soil water deficit are unaltered by elevated pC_a, while others predict decreased sensitivity to drought at elevated pC_a. These assumptions were tested by measuring canopy photosynthesis and transpiration in spring wheat (cv. Minaret) stands grown in boxes with 100 L rooting volume. Plants were grown under controlled environments with constant light (300 µmol m^?2 s^?1) at ambient (36 Pa) or elevated (68 Pa) pC_a and were well watered throughout growth or had a controlled decline in soil water starting at ear emergence. Drought decreased final aboveground biomass (?15%) and grain yield (?19%) while elevated pC_a increased biomass (+24%) and grain yield (+29%) and there was no significant interaction. Elevated pC_a increased canopy photosynthesis by 15% on average for both water regimes and increased dark respiration per unit ground area in well‐watered plants, but not drought‐grown ones. Canopy transpiration and photosynthesis were decreased in drought‐grown plants relative to well‐watered plants after about 20–25 days from the start of the drought. Elevated pC_a decreased transpiration only slightly during drought, but canopy photosynthesis continued to be stimulated so that net growth per unit water transpired increased by 21%. The effect of drought on canopy photosynthesis was not the consequence of a loss of photosynthetic capacity initially, as photosynthesis continued to be stimulated proportionately by a fixed increase in irradiance. Drought began to decrease canopy transpiration below a relative plant‐available soil water content of 0.6 and canopy photosynthesis and growth below 0.4. The shape of these responses were unaffected by pC_a, supporting the simple assumption used in some models that they are independent of pC_a.     

Co-expression of genes ApGSMT2 and ApDMT2 for glycinebetaine synthesis in maize enhances the drought tolerance of plants

Glycinebetaine plays an important role in the protection mechanism of many plants under various stress conditions. In this study, genetically engineered maize plants with an enhanced ability to synthesise glycinebetaine (GB) were produced by introducing two genes, glycine sarcosine methyltransferase gene (ApGSMT2) and dimethylglycine methyltransferase gene (ApDMT2), from the bacterium Aphanothece halophytica. Southern blotting and RT-PCR analysis demonstrated that the two genes were integrated into the maize genome and expressed. The increased expression levels of ApGSMT2 and ApDMT2 under drought conditions facilitated GB accumulation in the leaves of transgenic maize plants and conferred improved drought tolerance. Under drought conditions, the transgenic plants showed an increased accumulation of sugars and free amino acids, greater chlorophyll content, a higher photosynthesis rate and biomass, and lower malondialdehyde and electrolyte leakage compared to the wild-type; these results suggest that GB provides vital protection against drought stress. Under normal conditions, the transgenic plants did not show decreased biomass and productivity, which indicated that the co-expression of ApGSMT2 and ApDMT2 in maize plays an important role in its tolerance to drought stress and does not lead to detrimental effects. It was concluded that the co-expression of ApGSMT2 and ApDMT2 in maize is an effective approach to enhancing abiotic stress tolerance in maize breeding programmes.     

对生玉米产量性状的杂种优势和配合力分析

本文利用对生基因转育获得的对生与互生自交系及其杂交组合,研究了对生性状对产量性状杂种优势和配合力的遗传效应的影响。结果表明:对生玉米同互生玉米一样具有普遍的杂种优势,在产量性状上F1对生株优势互×互组合大于对×对组合和对×互(互×对)组合;对生性状的转育对产量性状的一般配合力有降低效应,而含有对生基因的互生玉米具有较高的一般配合力效应,因此,在对生玉米育种中可以利用含有不同对生基因的互生自交系作亲本来组配杂交组合(互×互),从而获得较高的制种产量和高产的对生杂交种。     

低氮诱导玉米幼苗叶片衰老过程中碳氮平衡的动态变化

叶片适时衰老对保证玉米产量有重要意义。本试验以玉米自交系PH6WC和PH4CV为研究对象,通过水培方法,设置低氮(0.04 mmol·L^-1,LN)和正常(4 mmol·L^-1,CK)氮素水平两种处理,在培养2、4、6和8 d后,对其幼苗第2和第3叶片表型、光合特性、叶片中氮素和糖分含量及碳氮比进行分析,旨在探究低氮胁迫下玉米幼苗叶片衰老过程中碳氮平衡的动态变化。结果表明: 与CK相比,LN造成两玉米自交系幼苗第2和第3叶片的面积、生物量、相对叶绿素含量、净光合速率、可溶性糖和淀粉含量均下降,而其氮物质生产能力均先后增加,但第2叶片的变化时间均早于第3叶片;在两叶片的各性状上,均为LN下PH6WC的变化幅度大于PH4CV,且仅幼苗叶片中的碳氮比在LN下显著提高;PH6WC的叶片衰老更快,PH4CV有更强的碳氮平衡能力,其叶片衰老相对滞后。综上,低氮会诱导玉米幼苗叶片衰老,高碳氮比具有促进叶片衰老的调控作用,低氮胁迫下幼苗叶片的碳氮平衡能力在两个玉米基因型间存在较大差异。