摘要
小麦的生育期、株高、产量、品质、光合功能、抗病和抗逆性等大多数性状都是由多基因控制的数量性状。随着分子生物学的发展,QTL的定位、克隆和分子标记辅助选育将成为作物产量、品质和抗性改良的重要途径和手段。本研究收集了近年来发表的与小麦株高、产量、品质、赤霉病抗性、穗发芽抗性、种子休眠性相关的QTL定位数据,并对来自不同试验的数据进行整合、比对,进行QTL元分析。同时,利用重组自交系对小麦不同生育阶段的光合功能、农艺性状以及芽期和苗期的耐湿性进行QTL定位,主要结果如下：
1.收集了近年来发表的56篇有关小麦株高(PH)和赤霉病抗性(FHB) QTL的文献,元分析后获得了27个控制株高的一致性QTL(MQTL)和30个赤霉病抗性一致性QTL(MQTL),比较分析后发现：位于2D、3A、4B、4D和7A上的株高MQTL"Rht-B1"、"Rht-Dl"、"Rht8" "PT"和“P26”均与相应染色体上的赤霉病抗性QTL完全或部分重叠,这些一致性区段或许能从分子水平上解释小麦株高和赤霉病抗性之间的相关性。对于Rht-B1、Rht-D1、Rht8和赤霉病抗性的负相关性,前人已有相关的田间实验报道；但是有关3A和7A上株高和赤霉病抗性QTL的一致性,尚未见相关的报道,其相关性还有待于进一步的田间试验来验证。同时,还发现来源于不同抗源的FHB抗性基因在染色体上聚集在相同的区段；部分染色体区段与两种或三种类型的赤霉病抗性均相关。对这些多效性区段的研究和应用将有助于赤霉病抗性机理的深入了解和多抗型小麦品种的选育。
2.整合分析了26篇有关小麦穗发芽抗性和种子休眠性QTL的文献数据,在3A、3B、3D和4A上得到了9个主效MQTL。其中,位于3AS上的"PHS2"与4A染色体上的4个主效MQTL均与种皮颜色无关,可用于选育白皮抗穗发芽小麦；对MQTL的染色体区段进行分析,发现小麦3B染色体上穗发芽抗性MQTL"PHS3"和控制休眠的MQTL"SD2"在49.45cM-49.74cM的置信区间内重叠；3D染色体上"PHS6"和“SDY”在69.35 cM-73.15 cM区间重叠,元分析结果进一步证实了种子休眠性是影响穗发芽的一个重要因素。
3.用元分析软件整合了299个与小麦产量相关的QTL、154个与籽粒硬度和蛋白含量相关的QTL,共得到26个与产量相关的MQTL、14个与品质相关的MQTL。在4A的55.58cM的MY7为主效热点MQTL,能解释22.75%的产量变异；2DS的21.67cM-28.47cM区段与产量和穗粒数均相关,变异解释度分别为20.93%和13.68%,是影响产量的主效区段；4A的54.27cM-56.89cM、4D的27.63cM-41.09cM和7DS上的33.46cM-43.09cM均为与产量相关的主效区段；在5D和1AS上各发现了1个与籽粒硬度相关的主效热点MQTL,且5D上的MQTL与“Ha”基因位置一致；此外,在6B的117.22cM处还发现了一个高蛋白含量MQTL,,贡献率为64.03%,该位点对小麦籽粒蛋白含量改良具有重要的应用价值。
4.用Opata85×W7984构建的重组自交系对小麦苗期、抽穗期和灌浆期的叶绿素含量(Chl)、叶片净光合速率(Pn)、气孔导度(Cond)、胞间C02浓度(Ci)和蒸腾速率(Trmmol)进行QTL定位,未检测到在三个生长发育时期都表达的QTL,仅发现部分加性QTL在两个生育阶段都有表达,大多数QTL表达均具时空特异性；对两年的QTL数据进行比对,在3A、4D和6A上发现了4个相对稳定的与叶绿素含量相关的QTL位点。其中,位于4D的102.3cM处的QTL与抽穗期Chl相关,两年的变异解释度均超过10%,为主效QTL；在3A上分别检测到了1个与Pn和Cond相关的稳定QTL；在2A上82.9cM也检测到了一个与Cond相关的稳定QTL；在2D上检测到1个控制千粒重和1个控制穗粒数的稳定QTL,在4D上和7B上分别检测到了1个与株高和经济学产量相关的稳定QTL。此外,对控制各性状的QTL进行上位性分析,发现大多数性状都受上位性效应影响,且上位性QTL大多与环境之间存在互作效应,而与环境存在互作效应的加性QTL则相对较少。同时,还在1D、2D和3A上发现了4个与光合功能、产量等多个性状均相关的多效性区段,这些重要区段对于聚合高光效和高产基因,培育高产和超高产小麦有着一定的应用价值。
5.用两套重组自交系对小麦芽期和苗期的耐湿性进行QTL定位,在芽期检测到了6个耐湿性QTL,其中,用ITMI群体定位的位于7A上57.9cM处的耐湿性QTL能解释23.92%的芽期耐湿性变异；在苗期,用ITMI群体在16条染色体上检测到了28个耐湿性QTL；在SHW-L1×SW8188群体内共检测到11个苗期耐湿性QTL,分别位于1A、1B、5D、6A和7D上；两套重组自交系均未检测到与耐湿性相关的上位性QTL,表明耐湿性主要受加性效应控制；对逆境和非逆境条件下的QTL进行比较,发现仅少量QTL在两个环境中同时表达,大多数QTL均受环境诱导；此外,还在6A、6β、7B和7D上发现了几个与多项耐湿性指标相关的多效性区段,其中7D上的289.4cM处,加性效应来自于SW8188的耐湿性QTL同时与叶绿素含量、根长和株高耐湿性指数相关,能解释30%以上的苗期耐湿性,对该位点的深入研究将有助于了解小麦苗期的耐湿性机制。
Most important traits of wheat, such as earliness, plant height, grain yield, quality, biotic stress and abiotic stress tolerance are generally reflecting quantitative inheritance. With the development of molecular biology, QTL location, isolation and MAS (molecular marker-assisted selection) are playing a much important role in crop genetic improvement on yield, quality and stress resistance. In this study, QTLs on plant height, garin yield, quality, Fusarium head blight, pre-harvesting sprouting and seed dormancy were collected for meta-anlysis. Meanwhile, QTL analysis for photosynthesis, agronomic traits and waterlogging tolerance were performed. The results were described as follows:
1. QTLs on plant height (PH) and Fusarium head blight (FHB) were investigated by QTL meta-analysis from fifty six experiments. Twenty seven PH meta-QTLs (MQTLs) and thirty FHB MQTLs were predicted by mete-analysis. Coincident MQTLs for PH and FHB were found on chromosomes 2D,3A,4B,4D and 7A. Rht-Bl, Rht-DI, Rht8, MQTLs P7 and P26 were consistent with FHB MQTLs. The meta-analysis results confirmed the negative associations of Rht-Bl, Rht-Dl, and Rht8 with FHB resistance. The associations of PH and FHB resistance on chromosomes 3A and 7A have not been reported and need further investigation. These regions should be given attention in breeding programs. MQTLs derived from several resistance sources were also observed. Some FHB MQTLs for different types of resistance overlapped. These findings could be useful for improving wheat varieties with resistance to FHB.
2. QTL meta-analysis was performed on pre-harvest sprouting (PHS) and seed dormancy (SD) from twenty six experiments. Nine major MQTLs (R2>10%) were located on 3A,3B,3D and 4A. Furthermore, the MQTL PHS2 on 3AS and four MQTLs on 4A, independent of red kernel colour loci could be used to develop PHS tolerant white wheat varieties. PHS and SD MQTLs were co-located in 49.45cM-49.74cM on chromosome 3B, and 69.35 cM-73.15 cM on 3D. The co-location confirmed that SD was the majore inheritant component of PHS.
3. Two hundred and ninety nine QTLs for grain yield, one hundred and fifty four QTLs for grain hardness and grain protein content (GPC) were meta-analysed. Twenty six MQTLs on grain yield and fourteen MQTLs on quality were deteced. The major MQTL, located at 55.58cM on 4A could explain 22.75%of phenotypic variance to grain yield. The co-located MQTLs, integrated at 21.67cM-28.47cM on 2DS, could explain 20.93%and 13.68%of phenotypic variance to grain yield and kernels per spike (KPS), respectively. All the genomic regions of 54.27cM-56.89cM on 4A,27.63cM-41.09cM on 4D and 33.46cM-43.09cM on 7DS were major regions for grain yield. One major MQTL was located on 5D and 1AS, individually, and the MQTL on 5D was consistent with Ha gene. The major MQTL, detected at 117.22cM on 6B, was a high GPC locus for protein content improvement, explaining 64.03%of GPC variance.
4. The recombinant inbread lines (RILs), derived from the cross of Opata85 and W7984 were used for QTL analysis for phosynthesis associated traits. QTLs for chlorophyll content (Chl), net leaf photosynthetic rate (Pn), stomatal conductance (Cond), and internal CO2 concentration (Ci) at seedling stage, heading stage and grain filling stage were deteced. Wheareas, only a few QTLs were deteced at two different growth stages, non-comment QTLs were deteced at all three stages. Four stable QTLs for Chl were located on 3A,4D and 6A. Among them, the QTL, detected at 102.3cM on 4D was a major QTL, with a R2>10%. On chromosome 3A, two major QTLs for Pn and Cond were found. Another major QTL for Cond were located at 82.9cM on 2A. Two major QTLs for grain yield and KPS were deteced on 2D, and major QTLs for PH and economic yield were located on 4D and 7B, individually. Episistas QTLs were detected for most of the photosynthesis related traits and agronomic traits. The effect of Episistas QTLxEnvirnoment was much more important than that of Additive QTLxEnvirnoment. Four regions on 1D,2D and 3A, associated with both photosynthesis and grain yield were observed, which might be uself for developing wheat varieties with high or super high yield.
5. Two sets of RILs were used for QTL analysis on waterlogging tolerance at germination and seedling stages. Six resistance QTLs at germination stage were deteced. The major QTL, located at 57.9cM on 7A could explain 23.92%of tolerance variance at germination stage. Twenty eight QTLs, located on sixteen chromosomes were observed in ITMI population. In the RILs, derived from SHW-L1 xSW8188,11 QTLs were deteced on 1A, 1B,5D,6A and 7D. None of episistas QTLs were deteced in either population, which indicated that waterlogging tolerance was controlled mainly by additive QTLs. Only a few of QTLs on growth and germination ability were observed both in stress and normal conditions. Co-located regions for different waterlogging tolerancecoefficient (WTC) were found in the study. One of the regions at 289.4cM on 7D, associated with WTC for Chl, root length and PH, accounted for more than 30% waterlogging tolerance at seedling stage. For the QTL, allele from SW8188 contributed for an increase of the tolerance.
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