Identification of quantitative trait loci under drought conditions in tropical maize. 1. Flowering parameters and the anthesis-silking interval

Drought is an important climatic phenomenon which, after soil infertility, ranks as the second most severe limitation to maize production in developing countries. When drought stress occurs just before or during the flowering period, a delay in silking is observed, resulting in an increase in the length of the anthesis-silking interval (ASI) and in a decrease in grain yield. Selection for reduced ASI in tropical open-pollinated varieties has been shown to be correlated with improved yields under drought stress. Since efficient selection for drought tolerance requires carefully managed experimental conditions, molecular markers were used to identify the genomic segments responsible for the expression of ASI, with the final aim of developing marker-assisted selection (MAS) strategies. An F₂population of 234 individuals was genotyped at 142 loci and F₃ families were evaluated in the field under several water regimes for male flowering (MFLW), male sterility (STER), female flowering (FFLW) and ASI. The genetic variance of ASI increased as a function of the stress intensity, and the broad-sense heritabilites of MFLW, FFLW and ASI were high under stress conditions, being 86%, 82% and 78%, respectively. Putative quantitative trait loci (QTLs) involved in the expression of MFLW and/or FFLW under drought were detected on chromosomes 1, 2, 4, 5, 8, 9 and 10, accounting for around 48% of the phenotypic variance for both traits. For ASI, six putative QTLs were identified under drought on chromosomes 1, 2, 5, 6, 8 and 10, and together accounted for approximately 47% of the phenotypic variance. Under water stress conditions, four QTLs were common for the expression of MFLW and FFLW, one for the expression of ASI and MFLW, and four for the expression of ASI and FFLW. The number of common QTLs for two traits was related to the level of linear correlation between these two traits. Segregation for ASI was found to be transgressive with the drought-susceptible parent contributing alleles for reduced ASI (4 days) at two QTL positions. Alleles contributed by the resistant line at the other four QTLs were responsible for a 7-day reduction of ASI. These four QTLs represented around 9% of the linkage map, and were stable over years and stress levels. It is argued that MAS based on ASI QTLs should be a powerful tool for improving drought tolerance of tropical maize inbred lines.     

玉米开花期相关性状的QTL分析

利用玉米强优势组合(Mo17×黄早四)自交衍生的191个F_2单株构建了由SSR和AFLP标记组成的分子连锁图谱,用F2进一步自交产生的184个F_(2:3)家系调查散粉期、吐丝期和开花-吐丝间隔期(ASI)的表型值,采用基于混合线性模型的复合区间作图法和相应的作图软件QTLmapper/V2.0,在两个生长环境下定位了与散粉期、吐丝期和ASI相关的QTL数目分别为13、7和5个,检测到3对控制散粉期、17对控制吐丝期和5对控制ASI的上位性效应位点;同时发现了与环境存在显著互作的3个散粉期、3个吐丝期和2个ASI单位点标记区域以及1对散粉期、3对吐丝期和2对ASI上位性效应区域.对玉米散粉期、吐丝期和ASI遗传基础中遗传因素相对作用大小分析表明,加性效应、部分显性效应和上位性效应是玉米开花期相关性状的重要遗传基础.     

镉胁迫下甜、糯玉米开花期性状的全基因组关联分析

以183份甜、糯玉米自交系构成的关联群体为材料,在2种Cd污染水平下对甜、糯玉米开花期(DA,days to anthesis)、吐丝期(Ds,days to silking)和开花吐丝间隔(ASI,anthesis silking interval)进行全基因组关联分析,以揭示重金属Cd胁迫对甜、糯玉米开花期的影响及...     

玉米品种耐阴性指标的筛选与评价

采用大田试验,研究了24个玉米品种在50%遮光处理下的形态、生理与产量性状的变化.结果表明：与自然光照相比,遮光处理后,玉米的株高降低,茎粗减小,雌雄间隔期延长,净光合速率减小,比叶重减小,地上部干物质量减少,果穗缩短变细,穗轴直径减小,行粒数减少,籽粒产量显著降低,其中雌雄间隔期、净光合速率、比叶重和行粒数变化的百分率与地上部干物质量和籽粒产量减少的百分率之间呈显著或极显著相关,可作为田间鉴定玉米耐阴性的有效指标.采用综合耐阴性状作为评价参数,经聚类分析表明,郑单958、浚单20、登海602等14个品种属耐阴型品种,安玉12、豫玉22等10个品种属非耐阴型品种.表明以形态、生理指标结合产量性状来评价玉米耐阴性较为客观,且简单易行.     

Identification of quantitative trait loci for leaf area and chlorophyll content in maize (Zea mays) under low nitrogen and low phosphorus supply

To investigate responses to nitrogen and phosphorus stress, 218 recombinant inbred maize (Zea mays L.) lines were grown under low nitrogen, low phosphorus, and control (i.e., nitrogen and phosphorus sufficient) conditions and evaluated at the silking stage for various traits, including leaf area, leaf chlorophyll content, flowering time, the interval between anthesis and silking, and grain yield. Among the 83 quantitative trait loci (QTL) detected, 29 were for controls, another 29 were for low nitrogen, and 25 were low phosphorus. These loci indicate that there were both common and specific genetic mechanisms underlying the investigated traits. Overlapping QTL for leaf size (area, length, and width) leaf chlorophyll level, flowering time, anthesis?Csilking interval, and grain yield were located at chromosome bin 2.03/2.04, bin 2.06/2.07/2.08, bin 4.01/4.02, bin 5.03/5.04, bin 6.07, bin 9.03, and bin 10.03/10.04. Many of these loci overlapped with previously reported loci controlling root growth as well as tolerance or response to nutrient deficiency. These QTL identify chromosome regions as targets for genetic improvement of low nitrogen and low phosphorus tolerance.     

Genetic analysis of the related traits of flowering and silk for hybrid seed production in maize

In the field, asynchrony of flowering and silk for male and female plants always occurs under different abiotic stresses, and reduces yield in the seed industry. Under stress conditions, a female parent with a short duration of silk emergence and a long time of silk receptivity can decrease the risk of the asynchrony for flowering and silking in the process of hybrid seed production. For dissecting the genetic basis of the related traits of flowering and silk in maize, a set of recombinant inbred lines (RIL) was evaluated at three different environments. Correlation analysis showed that anthesis silk interval (ASI) had a significant positive relationship with day to silk (DTS) and duration of silk emergence (DSE). Silk receptivity (SR) had a significant negative correlation to DTS, DSE, and ASI. This implied that the female parent with a long DTS, DSE, and ASI could easy lose its SR, and required synchrony of pollen shedding for the male parent in the field. A total of sixteen different QTLs were identified for the seven traits of flowering and silking traits, including three QTLs for day to tassel (DTT), two for day to pollen (DTP), two for duration of pollen shedding (DPS), three for SR, two for DTS, two for DSE, and two for ASI. The QTLs detected for the related traits of flowering and silk could select ideal traits for male and female parents to raise the yield in the seed industry under certain stress conditions.     

Two consensus quantitative trait loci clusters controlling anthesis–silking interval, ear setting and grain yield might be related with drought tolerance in maize

Unravelling the molecular basis of drought tolerance will provide novel opportunities for improving crop yield under water-limited conditions. The present study was conducted to identify quantitative trait loci (QTLs) controlling anthesis–silking interval (ASI), ear setting percentage (ESP) and grain yield (GY). The mapping population included 234 F₂ plants derived from the cross X178 (drought tolerant) × B73 (drought susceptible). The corresponding F_2:3 progenies, along with their parents, were evaluated for the above-mentioned traits under both well-watered and water-stressed field conditions in three different trials carried out in central and southern China. Interval mapping and composite interval mapping identified 45 and 65 QTLs for the investigated traits, respectively. Two QTL clusters influencing ASI and ESP on chromosomes 1 (bin 1.03) and 9 (bins 9.03–9.05) were identified in more than two environments, showing sizeable additive effects and contribution to phenotypic variance; these two QTL clusters influenced GY only in one environment. No significant interaction was detected between the two genomic regions. A comparative analysis of these two QTL clusters with the QTLs controlling maize drought tolerance previously described in three mapping populations confirmed and extended their relevance for marker-assisted breeding to improve maize production under water-limited conditions.     

Mapping quantitative trait loci associated with southern leaf blight resistance in maize (Zea mays L.)

Southern leaf blight (SLB) caused by the fungus Cochliobolus heterostrophus (Drechs.) Drechs. is a major foliar disease of maize worldwide. Our objectives were to identify quantitative trait loci (QTL) for resistance to SLB and flowering traits in recombinant inbred line (RIL) population derived from the cross of inbred lines LM5 (resistant) and CM140 (susceptible). A set of 207 RILs were phenotyped for resistance to SLB at three time intervals for two consecutive years. Four putative QTL for SLB resistance were detected on chromosomes 3, 8 and 9 that accounted for 54% of the total phenotypic variation. Days to silking and anthesis–silking interval (ASI) QTL were located on chromosomes 6, 7 and 9. A comparison of the obtained results with the published SLB resistance QTL studies suggested that the detected bins 9.03/02 and 8.03/8.02 are the hot spots for SLB resistance whereas novel QTL were identified in bins 3.08 and 8.01/8.04. The linked markers are being utilized for marker‐assisted mobilization of QTL conferring resistance to SLB in elite maize backgrounds. Fine mapping of identified QTL will facilitate identification of candidate genes underlying SLB resistance.     

弱光胁迫对不同耐荫型玉米果穗发育及内源激素含量的影响

2010—2011年以耐荫性较弱的玉米杂交种豫玉22和耐荫性较强的玉米杂交种郑单958为材料,在抽雄前3 d开始进行弱光胁迫处理,吐丝后10 d恢复自然光照,研究弱光胁迫及光恢复对不同耐荫型玉米果穗生长发育及其内源激素含量的影响。结果表明:弱光胁迫下,玉米果穗生长发育明显减缓,穗长、穗粗和果穗干物质积累显著减小,秃尖度变大;穗行数、穗粒数和籽粒库容显著降低;吐丝期果穗顶部小穗子房发育停滞,已有败育迹象的籽粒在恢复自然光照后无明显改善;豫玉22果穗和籽粒性状在处理间的差异程度均大于郑单958。弱光胁迫下,两玉米杂交种果穗的ABA和ZR含量均升高,而GA含量和GA/ABA比值均降低;IAA含量和IAA/ABA比值在郑单958果穗中表现为升高,而豫玉22则表现为下降。     

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).     

Apoplastic infusion of sucrose into stem internodes during female flowering does not increase grain yield in maize plants grown under nitrogen‐limiting conditions

Nitrogen (N) limitation reduces leaf growth and photosynthetic rates of maize (Zea mays), and constrains photosynthate translocation to developing ears. Additionally, the period from about 1 week before to 2 weeks after silking is critical for establishing the reproductive sink capacity necessary to attain maximum yield. To investigate the influence of carbohydrate availability in plants of differing N status, a greenhouse study was performed in which exogenous sucrose (Suc) was infused around the time of silking into maize stems grown under different N regimes. N deficiency significantly reduced leaf area, leaf longevity, leaf chlorophyll content and photosynthetic rate. High N‐delayed leaf senescence, particularly of the six uppermost leaves, compared to the other two N treatments. While N application increased ear leaf soluble protein concentration, it did not influence glucose and suc concentrations. Interestingly, ear leaf starch concentration decreased with increasing N application. Infusion of exogenous suc tended to increase non‐structural carbohydrate concentrations in the developing ears of all N treatments at silking and 6 days after silking. However, leaf photosynthetic rates were not affected by suc infusion, and suc infusion failed to increase grain yield in any N treatment. The lack of an effect of suc infusion on ear growth and the high ear leaf starch concentration of N‐deficient maize, suggest that yield reduction under N deficiency may not be due to insufficient photosynthate availability to the developing ear during silking, and that yield reduction under N deficiency may be determined at an earlier growth stage.     

Transgenic alteration of ethylene biosynthesis increases grain yield in maize under field drought‐stress conditions

A transgenic gene‐silencing approach was used to modulate the levels of ethylene biosynthesis in maize (Zea mays L.) and determine its effect on grain yield under drought stress in a comprehensive set of field trials. Commercially relevant transgenic events were created with down‐regulated ACC synthases (ACSs), enzymes that catalyse the rate‐limiting step in ethylene biosynthesis. These events had ethylene emission levels reduced approximately 50% compared with nontransgenic nulls. Multiple, independent transgenic hybrids and controls were tested in field trials at managed drought‐stress and rain‐fed locations throughout the US. Analysis of yield data indicated that transgenic events had significantly increased grain yield over the null comparators, with the best event having a 0.58 Mg/ha (9.3 bushel/acre) increase after a flowering period drought stress. A (genotype × transgene) × environment interaction existed among the events, highlighting the need to better understand the context in which the down‐regulation of ACSs functions in maize. Analysis of secondary traits showed that there was a consistent decrease in the anthesis‐silking interval and a concomitant increase in kernel number/ear in transgene‐positive events versus nulls. Selected events were also field tested under a low‐nitrogen treatment, and the best event was found to have a significant 0.44 Mg/ha (7.1 bushel/acre) yield increase. This set of extensive field evaluations demonstrated that down‐regulating the ethylene biosynthetic pathway can improve the grain yield of maize under abiotic stress conditions.     

Drought affects the rate and duration of organ growth but not inter-organ growth coordination

Drought at flowering and grain filling greatly reduces maize (Zea mays) yield. Climate change is causing earlier and longer-lasting periods of drought, which affect the growth of multiple maize organs throughout development. To study how long periods of water deficit impact the dynamic nature of growth, and to determine how these relate to reproductive drought, we employed a high-throughput phenotyping platform featuring precise irrigation, imaging systems, and image-based biomass estimations. Prolonged drought resulted in a reduction of growth rate of individual organs—though an extension of growth duration partially compensated for this—culminating in lower biomass and delayed flowering. However, long periods of drought did not affect the highly organized succession of maximal growth rates of the distinct organs, i.e. leaves, stems, and ears. Two drought treatments negatively affected distinct seed yield components: Prolonged drought mainly reduced the number of spikelets, and drought during the reproductive period increased the anthesis-silking interval. The identification of these divergent biomass and yield components, which were affected by the shift in duration and intensity of drought, will facilitate trait-specific breeding toward future climate-resilient crops.

When affected by drought, the plant responds by adjusting the growth of individual organs but not the coordination between the organs.     

玉米雌雄开花间隔天数、结穗率与产量的数量性状位点(QTL)分析

采用SSR标记连锁图谱和复合区间作图法在山西灌溉和干旱胁迫条件下,对玉米(Zea mays L.)自交系黄早四×掖107组合的F3群体雌雄开花间隔天数(ASI)、结穗率和籽粒产量进行了数量性状位点(QTL)定位及基因效应分析.结果表明,在两种水分处理下,ASI、结穗率与籽粒产量的相关性均达到显著水平(P＜0.05).在灌溉和干旱胁迫下,分别检测到3个和2个控制ASI的QTL,位于第1、2、3和第2、5染色体上.在灌溉条件下,在第3和第6染色体上各检测到1个控制结穗率的QTL,基因作用方式呈加性或部分显性,可解释19.9%的表型变异;在干旱条件下,在第3、 7、10染色体上共检测到4个控制结穗率的QTL,基因作用方式为显性或部分显性,可解释60.4%的表型变异.在灌溉和干旱胁迫下,控制产量的QTL分别定位在第3、6、7和第1、2、4、8染色体上,基因作用方式均以加性或部分显性为主,可解释的表型变异为7.3%～22.0%.在干旱条件下,借助连锁分子标记和基因效应分析,可构建包含ASI、结穗率和产量QTL的选择指数,用于分子标记辅助育种.     

Genetic analysis of drought tolerance in maize by molecular markers. II. Plant height and flowering

Drought is a serious agronomic problem, and one of the most important factors contributing to crop yield loss. In maize grown in temperate areas, drought stress occurs just before and during the flowering period; consequently, tolerance to water stress in this species is largely determined by events that occur at or shortly after flowering. The purposes of our investigation were: (1)?to identify the chromosomal regions where factors conferring drought tolerance for traits related to plant development and flowering are located and (2)?to compare these regions with those carrying QTLs controlling these traits, in order to get indirect information on the genetic and physiological basis of maize response to water stress. To this aim, we performed a linkage analysis between the expression of male and female flowering time, anthesis-silking interval (ASI), plant height and molecular markers. The experiment was carried out under two environmental conditions, well-watered and water-stressed, on a maize population of 142 recombinant inbred lines obtained by selfing the F₁ between lines B73 and H99 and genotyped by RFLP, microsatellites (SSR) and AFLP markers, for a total of 153 loci. Linkage analysis revealed that, for male flowering time and plant height, most of the QTLs detected were the same under control and stress conditions. In contrast, with respect to female flowering time and ASI diverse QTLs appeared to be expressed either under control conditions or under stress. All of the QTLs conferring tolerance to drought were located in a different chromosome position as compared to the map position of the factors controlling the trait per se. This suggests that plant tolerance, in its different components, is not attributable to the presence of favourable allelic combinations controlling the trait but is based on physiological characteristics not directly associated with the control of the character.     

Carbohydrate, Nitrogen and Dry Matter Accumulation and Partitioning of Maize Hybrids Under Drought Stress

Maize (Zea mays L.) productivity under drought stress dependsto some extent upon a hybrid's capacity to produce and translocateassimilate to its developing kernels during the stress periodand/or after the stress is relieved. The objective of this studywas to evaluate differences in carbon and nitrogen accumulationand partitioning under drought stress among maize hybrids thatdiffer in yield potential and/or physiological metabolism duringreproductive development. The hybrids B73 x LH38, FS854, B73xMol7and US13 were subjected to drought stress from the 7th leafstage until pollination was completed, at which time the soilof the stressed plots was replenished with water. For d. wtand chemical constituent determinations, plants of each hybridwere harvested from the irrigated and drought stressed plotsat silking, mid-grain fill, and physiological maturity. Averagedover hybrids, vegetative biomass at silking was reduced 25%as a result of the drought stress treatment, with B73 x LH38and FS854 accumulating more total biomass during the later portionof grain fill than the other two hybrids under both soil moisturetreatments. At silking, the total non-structural carbohydratecontent of the hybrids' vegetative tissue was not changed asa result of drought stress, whereas their reduced nitrogen (N)contents were decreased by an average of 33%. B73 x LH38 andFS854 had greater grain carbohydrate and reduced N contentsunder irrigation and smaller decreases in those variables asa result of soil moisture deficit than did the other two hybrids.These results indicate that the greater drought tolerance ofB73 x LH38 and FS854 to stress imposed during vegetative andearly reproductive development resulted from their more activeN uptake and assimilation and sugar production during the laterportion of grain fill and from their more efficient partitioningof assimilate to the developing kernels. Zea mays L., maize, drought stress, nitrogen, carbohydrates, hybrids, partitioning