摘要
光合作用是“地球上最重要的化学反应”,与农业生产有着十分密切的关系,提高光合效率是提高作物产量的重要途径,其目的是选择高光效品种。这是因为高光效品种的净光合速率与单位叶面积内光合单位数呈正相关。然而,大量研究表明,在过剩强光甚至中等光强逆境条件下,作物光合器官就会受到伤害,会产生光氧化现象。这种生理病害,影响作物的光合生产和产量,愈来愈受到注意。
利用溧水中子黄豆(P,)×南农493-1(P2)大豆杂交组合的F2、F2：3和F2：43个群体两个试验点(南京农业大学江浦试验站和山东临沂农科院试验站)的试验资料研究大豆光氧化等有关数量性状的遗传规律。首先,构建大豆遗传图谱；然后,利用该连锁图,定位大豆光氧化等的数量性状基因座(quantitative trait loci, QTL),获得不同群体、不同作图方法下都稳定存在的QTL。这些QTL定位结果可为大豆高光效育种和分子标记辅助育种的提供有用的参考信息。
1大豆遗传图谱的构建
采用972对SSR引物扫描亲本P1和P2间的遗传差异,发现150对引物呈现多态性。利用这150对多态性的SSR引物扫描244株F2个体间的遗传差异,获得多态性F2植株分子数据资料,构建了包含91个SSR分子标记的28个连锁群的遗传图谱。连锁群全长1356.42cM,平均间距14.91cM,单个连锁群长度在6.9-108.3cM之间,每个连锁群的标记数为2-9个,平均间距最大的连锁群是Ⅰ-2,为51.3cM,最小的连锁群是L,为6.9cM,有28个间距大于20cM的标记区间,59个SSR标记不属于任何连锁群。与大豆“公共遗传图谱”比较,91个标记中86个标记所在连锁群及其顺序与公共遗传图谱一致,satt077与sat_110(E)、satt266和sat_2542(Dlb)排序与公共图谱相反,只有标记satt669所在的连锁群与公共图谱不一致。
2大豆光氧化相关性状QTL定位
光氧化试验包括2007年江浦试验站F2：3群体试验与2008年江浦试验站和临沂农科院试验站F2：4群体试验,共获得三组试验资料。每组资料用复合区间作图(composite interval mapping:CIM)和多区间作图(multiple interval mapping:MIM)分析,三组资料联合用多标记联合分析(multi-marker joint analysis:MJA),共7次分析,检测了光氧化前叶绿素含量(Normal chlorophyll concentration before photooxidation:NCC)、光氧化后叶绿素含量(chlorophyll concentration after photooxidation:PCC)、光氧化率(photooxidation rate:POR)和黄叶率(yellow leaf rate:YLR)的主效QTL共181个、上位性QTL共37个、环境效应共4个和环境互作共29个,其中,7次检测PCC的52个QTL中共同检测到2～5次的有10个,NCC的41个QTL中共同检测到2-3次的有8个,POR的29个QTL中共同检测到2～4次的有5个和YLR的59个QTL中共同检测到2～4次的有7个。这些QTL有聚集成簇的现象,在C2连锁群中的satt640-satt42区间、D1b连锁群的sat_160-satt147区间、D2连锁群的satt413-satt256区间、E连锁群的satt263-satt045区间、G连锁群的sat_418-sat_419区间及M连锁群的sat_391-satt150区间成簇出现,推测这些染色体区域可能是控制光氧化相关性状的基因。方法间以CIM和MIM的共检测率较高,约50%。年份间的共检测率不高,而年份内不同地点和同一地点不同的年份间共检测率较高。4性状间的QTL检测结果也可解释性状间的相关性。
3大豆叶绿素含量动态表达的QTL定位
在大豆生育期内不同时间点,测定2007年江浦试验站244个F2：3家系8次叶绿素含量(资料Ⅰ)、2008年江浦试验站244个F2：4家系1次叶绿素含量(资料Ⅱ)和2008年临沂农科院试验站244个F2：4家系4次叶绿素含量(资料Ⅲ)。在Ⅰ～Ⅲ资料中,求每组资料不同时间点的平均叶绿素含量,得到各家系平均叶绿素含量(资料Ⅳ)。用复合区间作图法分析资料Ⅰ～Ⅲ,用多标记联合分析方法分析资料Ⅳ,共检测到45个QTL。
2007年江浦试验站的8次测量中共检测到18个QTL,第1～8时间点分别有3、2、2、3、3、2、3和0个。4个时间点检测到N连锁群上的satt234-satt022标记区间存在QTL,2～3个时间点检测到D1a、D1b、D2和F连锁群上存在QTL。从贡献率上看,大于10%的有11个,占总个数的61%；最大贡献率达57%,该QTL也正是2007年江浦8次检测中重复检测次数最多的。
2008年临沂试验站的4次测量中共检测到11个QTL,第1～4时间点分别检测到2、3、4和2个QTL,只有2个时间点检测到D2和D1a连锁群上存在QTL,但QTL位置均不同。从贡献率上看,大于10%的有2个,占总个数的18%；最大贡献率14%,该QTL也是2008年临沂试验站重复检测次数最多的QTL之一。
同一地点的不同年份间只检测到1个共同的QTL,即位于sat_160-satt147的qchl-D1a-1。江浦和临沂不同地点间共同检测到的QTL有3个,分别是位于D1a连锁群的gchl-D1a-1;位于D2连锁群的qchl-D2-1和位于K连锁群上的qchl-K.
利用多标记联合分析方法检测到资料Ⅳ的分别位于9个连锁群上的7个主效QTL、4个环境与标记互作QTL和1个环境效应。MJA方法与CIM方法共同检测到的QTL共有5个,分别位于D1a、D2、M和N连锁群上。环境效应的存在说明环境对叶绿素合成影响较大。此外,还检测到6个QTL和4个环境与标记互作。
在不同时间点共检测到45个QTL,但除N连锁群外不同时间点上共同的QTL不多,揭示了叶绿素表达时空表达差异；CIM和MJA两种方法共同检测到的QTL共有5个；MJA方法新检测到的主效QTL6个和环境互作QTL4个,存在环境效应。这些结果为叶绿素性状的遗传剖析和标记辅助育种提供理论依据。
4大豆对豆卷叶螟抗性的遗传分析
在田间自然虫源条件下,利用P1、P2、F1和F2共4个世代进行了遗传分离分析,表明对豆卷叶螟抗性符合2对主基因+多基因遗传模型。利用F2单株叶片损失率数据和已构建的连锁遗传图谱,定位大豆对豆卷叶螟抗性的QTL。复合区间作图法检测到位于D1b和K连锁群上的2个QTL；多区间作图法检测到位于A2、D1b、K和N连锁群上的4个QTL和6个互作QTL；其中有两个共同的QTL,至少解释表型变异的19.2%。
Photosynthesis is the most important chemical reaction on the earth and very closely related to agricultural production. Improving photosynthetic efficiency is an important way to increase crop yield and to select the cultivar with high photosynthetic efficiency, because net photosynthetic rate is significantly related to the number of photosynthetic units per area. However, a number of studies have suggested that the photosynthetic apparatus is hurt and photo-oxidation phenomenon often happens under the situation of strong or middle strong light in plants. Therefore, more and more attention was paid to the effect of this physiological disease on photosynthetic production.
The F2, F2:3and F2:4populations derived from the soybean cross of Lishuizhongzi-huang and Nannong493-1at Jiangpu experimental station of Nanjing Agricultural University or/and at Linyi experimental station of Linyi Agricultural Academy were used to study the inheritance of quantitative traits related to photooxidation. First, genetic linkage maps in soybean were constructed. Then, mapping quantitative trait loci (QTL) responsible for the above traits was carried out in order to obtain stable QTL across various populations and mapping approaches, based on the genetic linkage maps above. The results presented will provide some useful information for molecular design breeding and high photosynthetic efficiency in soybean.
1. Construction of genetic linkage maps in soybean
The genetic linkage map of soybean using244F2plants derived from the soybean cross above was constructed in this paper. Before the construction of a linkage map, the two parents were screened for polymorphism with972SSR primer combinations. In total,150of972primers (15.4%) could produce polymorphic loci. All these polymorphic markers were performed in the F2population. In the construction of a genetic linkage map, there are three steps. The first step is to cluster markers into linkage groups. Ninety-one molecular markers were assigned to28linkage groups with Mapmaker3.0software. The second step is to estimate genetic distances in each of the linkage groups. The results show that the length of each linkage group ranges from6.9to108.3cM and all these molecular markers covered1356.42cM for the whole genome. The minimum, maximum and average of marker spacing are6.9(linkage group L),51.3(linkage group1-2) and14.91cM, respectively. Number of markers included in each linkage group varies from2to9. There are twenty-eight marker intervals with marker spacing longer than20cM. Fifty-nine markers are not placed on any linkage groups. The last step is to optimize the orders of all mapped markers in all linkage groups. The presented maps were the optimized linkage groups. Among ninety-one markers, the linkage gropup and order for eighty-six marker are consistent with the public genetic maps in soybean while different order between satt077and sat_110, and between satt266and sat_2542, and different linkage group for satt669are found in this paper.
2. Mapping QTL for photooxidation-related traits in soybean
Two-hundred and forty-four F2:3families derived from the soybean cross above at Jiangpu experimental station in2007and the corresponding F2:4families at the Jiangpu experimental station and at Linyi experimental station in2008were measured for four photooxidation-related traits, including normal chlorophyll concentration before photo-oxidation (NCC), chlorophyll concentration after photo-oxidation (PCC), photo-oxidation rate (POR) and yellow leaf rate (YLR). Its purposes is to perform the inheritance analysis of the above traits by using the CIM, MIM and multi-marker joint analysis (MJA). Each dataset was analyzed respectively by the first two approaches and all data was jointly analyzed by the MJA. The results showed that181main-effect QTL along37epistatic QTL,4environmental effects and29QTL-by-environment interactions were detected in the seven genetic analyses. Of52QTL for PCC,10QTL were identified repeatedly2to5times. Of41QTL for NCC,8QTL were mapped repeatedly2to3times. Of29QTL for POR,5QTL were identified repeatedly2to4times. Of59QTL for YLR,7QTL were found repeatedly2to4times. The phenomenon of QTL cluster was present in our results. Several QTL that influence multiple traits were detected in the same genomic regions. A total of six intervals were found to be involved in the control of two or more traits and located on marker intervals satt640-satt42, sat_160-satt147, satt413-satt256, satt263-satt045, sat_418-sat_419and sat_391-satt150in the linkage groups C2, Dlb, D2, E, G and M, respectively. We deduce that there are some genes responsible for the traits related to photooxidation in these regions above. In addition, results showed that there are high consistency between the CIM and the MIM, between different places in the same year, and between different years in the same place, and low replication between different years. Results from QTL mapping may explain the correlation between photooxidation-related traits.
3. Mapping QTL for dynamic chlorophyll content in soybean
Two-hundred and forty-four F2:3families derived from the soybean cross above at Jiangpu experimental station in2007and the corresponding F2:4families at the Jiangpu experimental station and at Linyi experimental station in2008were measured for chlorophyll content at eight, one and four different developmental stages, respectively. Therefore, the average for each family across different developmental stages could be calculated at fixed year or experimental station. The average data was analyzed by using the MJA while the data at different developmental stages was analyzed by using the CIM and MIM. The results showed that45QTL were detected.
At the measured times1to8at Jiangpu experimental station in2007,3,2,2,3,3,2,3and0QTL for chlorophyll content were detected, respectively. Of these QTL, one QTL located on the marker interval satt234-satt022in the linkage group N, with maximum percentage57.0%of the total phenotypic variance explained, was repeatedly identified across four times, some QTL placed on the linkage groups D1a, D1b, D2and F were repeatedly confirmed across2to3times, and there were11QTL with more than10.0%of the total phenotypic variance explained.
At the measured stages1to4at Linyi experimental station in2008,2,3,4and2QTL for chlorophyll content were detected, respectively. Of these QTL, the QTL located on different positions of the linkage groups D2and D1a were identified across two times, there were2QTL with more than10.0%of the total phenotypic variance explained, and maximum percentage of the total phenotypic variance explained was14.0%.
One common QTL, qchl-D1a-1, located on the marker interval sat_160-sattl47in the linkage group Dla, was mapped across different years at Jiangpu experimental station. Three common QTL, qchl-D1a-1, qchl-D2-1and qchl-K, located on the linkage groups D1a, D2and K, respectively, was mapped between Jiangpu and Linyi experimental stations.
Using the multi-marker joint analysis for the average data above,7main-effect QTL,4environmental interactions QTL and1environmental effect were identified on nine linkage groups. Five common QTL placed on the linkage groups D1a, D2, M and N were detected between the CIM and MJA approaches. Significant environmental effect indicates that environment have a large effect on chlorophyll synthesis. In addition,6new QTL and four environmental interactions were found by the MJA.
4. Genetic analyses for resistance of soybean to bean pyralid(Lamprosema indicata Fabricius)
Two-hundred and forty-four F2plants derived from the soybean cross between Lishui-zhongzihuang (resistance) and Nannong493-1(susceptible) were measured for resistance in soybean to natural population of bean pyralid in the field according to its defoliation percentage in2006. QTL detection for the resistance was carried out using the F2population. The results showed that two QTL were mapped on the linkage groups Dlb and K using composite interval mapping (CIM). Four QTLs along with six interactions were identified in the linkage groups A2, Dlb, K and N using multiple intervals mapping (MIM). Among these QTL, there were two common QTL with more than19.2%of total phenotypic variance explained.
引文
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