高粱与苏丹草的遗传及其杂种优势利用的研究
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摘要
高粱[Sorghum bicolor(L.)Moench和苏丹草[Sorghum sudanense(Piper)Stapf]均属于禾本科高粱属,普遍被认为起源于热带非洲,二者在亲缘关系上有一定距离,但无生殖隔离。高粱的分类问题,在几百年来一直被众多植物学家探索着。目前,苏丹草与高粱是否属于同一个种至今存在争议,亟待澄清两物种分类学上混乱。
高粱与苏丹草没有生殖隔离,这就使利用高粱与苏丹草的杂种优势成为可能。开展高粱-苏丹草杂种优势利用及其指纹图谱和分子标记研究,对饲用高粱和苏丹草遗传育种具有重要的理论和实际意义。
本研究旨在通过数值分类学、细胞遗传学、生理生物化学和分子生物学等方法,探讨苏丹草与高粱形态学与分子性状的遗传亲缘关系,开展高粱与苏丹草杂交优势利用研究,建立高粱与苏丹草及其杂交种DNA指纹图谱,进行高粱/苏丹草褐色中脉基因的遗传分析和SSR标记定位,为进一步选育更加高产优质品种创造前提和条件。本研究获得的主要结论如下:
1.高粱与苏丹草生物学特性的数值分析
选取高梁属不同来源和类型高粱品种6个,苏丹草品种4个,另加1份与苏丹草和高粱染色体数相同的饲用墨西哥玉米(Zea Mexicana)作为参照物种开展数值分析。在植物营养生长阶段和生殖生长阶段观测高粱、苏丹草生物学性状共64个。对苏丹草与高粱10个品种共300个个体的两组判别分析表明,苏丹草与高粱品种间F测验达极显著水平。因此,苏丹草与高粱类间差异明显。至于苏丹草类内和高粱类内部分品种间F测验显著,说明苏丹草与高粱类内又存在分化现象。对苏丹草与高粱的10个品种各30株共300个个体进行判别分析,结果每品种的30个个体都能回判到各自品种中,表明参试的30个个体是合理的。用观测的64个性状指标对苏丹草和高粱共10个品种进行分层聚类,结果苏丹草和高粱明显被分成两大类。
与此同时,本研究还系统分析了苏丹草和高粱64个生物学性状的分类学意义;利用主成分分析法确定出6个主分量,在特定生态条件和栽培模式下,主成分Ⅰ、Ⅱ值的大小决定着苏丹草和高粱共同祖先向各自发展方向进行演化,以6个主成分的回归因子得分值进行10个品种的快速聚类,结果苏丹草和高粱被分成2大类。进一步扩大参试品种的数量,对48个苏丹草和高粱及其近缘种进行数量分类研究,结果基本支持原10个品种的结论,但全部苏丹草和高梁两类群不能被完全分离开来。
2.高粱与苏丹草及其杂种的细胞遗传学研究
采用去壁低渗火焰干燥法对4个苏丹草和6个高粱品种的核型进行比较研究,结果表明苏丹草和高粱体细胞染色体数均为20(2n=20)。苏丹草品种均为1A核型,并具有1对随体染色体,其中1、2号品种的第2对染色体为近中间着丝点染色体,3、4号品种为中间着丝点染色体;高粱品种除5号为1A核型、中间着丝点染色体外,其余的5个品种为2A或2B核型,且都有1或2对近中间着丝点染色体,9号品种还出现1对近端着丝点染色体;高粱8、9号品种各观察到1对随体染色体。染色体数量分析表明,第1到第10对染色体长短臂的绝对长度、相对长度以及绝对全长在苏丹草和高粱两类间的差异均不显著(P＞0.05)。因此,苏丹草和高粱的遗传差异不在染色体长度上。
进一步观察了3个杂种F_1减数分裂染色体行为和2个杂种F_2体细胞中染色体数目。结果杂种F_1花粉母细胞减数分裂过程中,中期Ⅰ染色体核型和数目清晰可见(2n=2x=20),配对行为规则;棒状和环状二价体的频率因组合不同而有差异,Tx623A×S722 F_1、3042A×Sa F_1和Tx623A×Sa F_1棒状二价体频率分别为4.887,5.710和5.126,环状二价体频率分别为5.113,4.290和4.874;在后期Ⅰ,配对的染色体能够正常分离。杂种F_2体细胞中染色体数目为20(2n=20)。说明苏丹草与高粱染色体具有很高的同源性,能够正常配对并稳定传递,二者亲缘关系非常近。
3.高粱与苏丹草同工酶以及光合特性的比较研究
采用聚丙烯酰胺电泳法,对4个苏丹草和6个高粱品种进行过氧化物酶同工酶的分析比较,结果表明苏丹草和高粱各品种的酶带不完全相同,但却未能找到苏丹草和高粱两类的特异带谱。进一步对以上材料进行酯酶同工酶分析,结果苏丹草被聚为一类,高粱被聚为另一类,因此,酯酶同工酶谱可以作为鉴别苏丹草和高粱的有效生化标记。
采用CI-310便携式光合测定系统,对32份高粱和10份苏丹草品种进行强弱光下光合特性的比较研究。结果表明:高粱和苏丹草的各项光合指标除后者的胞间CO_2浓度外,在强弱两种光照条件下均有显著或极显著差异。在弱光条件下,各项光合指标在高粱和苏丹草间均没有显著差异;但在强光条件下,高粱的蒸腾速率和胞间CO_2浓度极显著或显著超过苏丹草。在弱光条件下,无论高粱或苏丹草,其净光合速率、蒸腾速率和气孔导度三者间均呈显著或极显著正相关;但在强光条件下,苏丹草的各项指标无显著相关,而高粱大部分指标间相关极显著。
4.应用RAPD和SSR分子标记对高粱和苏丹草遗传多态性的研究
选用了32份高粱品种、10份苏丹草品种及2份高粱近缘种拟高粱和假高粱进行了RAPD分析。结果表明:12对RAPD引物产生的68条DNA扩增片段中,52条(76.5%)具有多态性。高粱之间的相似系数从55%到95%;苏丹草之间的相似系数从52%到84%,因此选择的品种之间的多态性高,具有代表意义。高粱不育系和保持系之间的相似度很大为89%以上。以0.66为阈值将44个品种分为十个类群。第1类群为4份苏丹草品种;在2、3、4群中既有苏丹草也有高粱;5、6、7、8群中则全部为高粱。因此,从RAPD分子标记聚类结果看,高粱和苏丹草遗传差异较小,难以完全分开。
用SSR标记分析了48份高粱、苏丹草及其近缘植物拟高粱、假高粱、墨西哥玉米、玉米的多态性及其亲缘关系。91对SSR引物在48个品种中扩增得到723个等位基因,平均每个位点有7.945个等位基因,多态性率达98.480%,多态性信息含量PIC为0.783。其中,在高粱品种内扩增出669个等位基因,平均每个位点有7.352个,多态性率为84.530%,多态性信息含量PIC为0.774;在苏丹草品种内扩增出649个等位基因,平均每个位点有7.132个,多态性率为78.483%,多态性信息含量PIC为0.770。通过对高粱与苏丹草多态性信息含量PIC值差异性t检验表明,二类群间无显著差异(p＞0.05)。高粱和苏丹草两类间平均遗传距离仅0.035。全部48份材料间的遗传相似系数最小为0.217,最大为0.975,据此进行聚类分析,结果显示:以相似系数0.766作为分类临界值,可以将48份材料分成5类,即全部高粱和苏丹草品种聚为1类,拟高梁(S.propinquum)1类,假高粱(S.halepense)1类,墨西哥玉米(Zeamexicana)1类,玉米(Zea mays)1类。因此,将苏丹草作为1个独立的植物分类种并不合适,建议将其划归双色高粱种(S.bicolor)。
5.高粱与苏丹草杂种优势利用的研究
苏丹草(S.sudanense)与高粱(S.bicolor)均为禾本科高粱属植物,二者的杂种优势明显。20世纪80年代,作者首次在我国开展高粱与苏丹草杂种优势利用的研究,研究结果表明:F_1单株生物产量性状高于双亲平均值,甚至超过或接近高亲。影响高粱-苏丹草杂交种单株茎叶鲜重的主要因素是主茎粗、叶长;决定单位面积鲜草产量的主要是密度、叶长和单株茎叶鲜重,其中在一定范围内密度对单位面积产量的直接通径系数最大,为1.0299~(**)。多年随机区组试验结果表明,杂交种单位面积鲜草产量显著超过苏丹草,并以30万株/hm~2为最适种植密度。杂交种营养品质与苏丹草相近,氰化物含量低,在孕穗后期含糖量达到最大值,为10.0%(Bx)。
6.高粱与苏丹草及杂交种DNA指纹图谱构建和褐色中脉基因的定位
从100个RAPD引物中筛选到9个多态性高、重复性好的引物,多态性条带比率为64.06%,利用4个核心RAPD引物可以为42份高粱与苏丹草品种资源中的每份品种构建一张特定的数字指纹,并通过其中1个引物F-01构建了一张能鉴别2份国审品种皖草2号和皖草3号高粱-苏丹草杂交种的RAPD指纹图谱,不过该图谱不能区别皖草3号与其父本Sa。从95对SSR引物中筛选出多态性丰富的引物73对,多态性条带比率为86.06%,通过3对核心SSR引物就可以构建42份高粱和苏丹草的SSR数字指纹,同时利用其中1对SSR引物txp18,寻找到两个杂交种的互补带,从而构建了两个高粱-苏丹草杂交种的SSR指纹图谱,这张SSR指纹图谱不仅能鉴别皖草2号和皖草3号,还可以把杂交种与其亲本区别开来。
高粱/苏丹草褐色中脉品种的叶片中脉和茎杆髓部是褐色的,茎杆的表皮呈深褐色。此类品种能使难以消化的木质素含量降低40%-60%,从而大幅度地提高消化率。利用高粱经典材料Tx623A的褐色中脉突变体(bmr-6类型)与白色中脉苏丹草(Sa-line)进行杂交,构建F_2分离群体,F_1表现为白色中脉,F_2白褐分离比符合3:1。这说明褐色中脉基因受一对等位基因控制,表现为隐性遗传。用已知定位到连锁群上的SSR标记和分离群体分析法,对褐色中脉bmr-6基因进行了连锁分析,发现1个与褐色中脉bmr-6基因连锁的微卫星标记,连锁距离为4.2cM,并将该基因初步定位于第7连锁群上。
Sorghum[Sorghum bicolor(L) Moench]and sudangrass[S.sudanense(piper) stapf] are two members of the agriculturally-important genus Sorghum(Gramineal),but some disagreement exists as to whether they actually belong to the same species.Hybrids of these two species have exhibited favorable forage yields and overall improved quality and disease resistance.They are widely used in aquaculture,production of livestock food and environmental protection.It is urgent to study the morphological characteristics,heterosis, forage value and molecular genetic for hybrid.
In this research,the genetics of sorghum(S.bicolor) and sudangrass(S.sudanense) was studied by numerical taxonomy,cytogenetics,physiology/biochemistry and molecular biology.Heterosis of hybrid between sorghum and sudangrass was utilized for the first time in China.DNA fingerprints for 42 sorghum and sudangrass varieties and 2 sorghum-sudangrass hybrids were constructed.The brown midrib gene(bmr-6) of sorghum-sudangrass hybrids was mapped.The results were listed as the following:
1.Numerical Analysis of Genetic Variation in Sorghum[S.bicolor(L.) Moench]and Sudangrass[S.sudanense(Piper) Stapf]
Sixty-four biological characteristics were evaluated for a comparative study between sorghum(S.bicolor) and sudangrass(S.sudanense) by means of variance,discrimination and cluster analyses.The significant differences in biological characteristics between sorghum and sudangrass were found.The F-test divergence values between the varieties of sorghum and sudangrass were highly significant.Discrimination analysis of all plants suggested that 100%of originally grouped cases were classified correctly.Cluster analysis showed that the 10 varieties of sorghum and sudangrass in the study fell into 2 groups:four sudangrass varieties making up one kind,and six sorghum varieties making up the other. Analysis of variance showed the characteristics chosen for cluster analysis were reasonable.
The characters of significant positive correlation and significant negative correlation had been found.In terms of a combined line,64 characters were clustered to 5 groups.The taxonomic significance of these characters had also been analyzed.6 principal branch quantitative characters were determined using the principal component analysis.Principal component-Ⅰwas named for principal sudangrass factor and principal component-Ⅱwas named for sorghum principal factor.10 varieties could be divided into 2 main groups depending on the result of cluster analysis by principal component regression values. Sudangrass was clustered to 1 group,and sorghum was clustered to the other group.
The results of quantity classification for 48 varieties,i.e.32 sorghum varieties and 10 sudangrass varieties and 4 relatives,supported basically above viewpoint.
2.Cytogenetical Studies of Relationship on Sudangrass(S.sudanense) and Sorghum (S.bicolor)
The Karyotypes of 4 sudangrass varieties and 6 sorghum varieties during mitosis were investigated using the method of wall degradation and hypotonic treatment of making chromosomal preparations.The results showed that the numbers of chromosome are 20 in sudangrass(S.sudanense) and sorghum(S.bicolor).The karyotypes of sudangrass varieties were 1A,and their karyotype formulas were shown as follows:No1 and No2 2n=18m+2sm(2SAT),No3 2n=18m+2sm(2SAT),No4 2n=20m(2SAT).Their karyotypes were not all same among sorghum varieties.Only No5 was 1A,and all the other varieties were 2A or 2B.The karyotype formulas of sorghum varieties were:No5 2n=20m,No6 2n=16m+4sm,No7 2n=18m+2sm,No8 2n=18m(2SAT)+2sm,No9 2n=16m+2sm(2SAT)+2st,No10 2n=16m+4sm.There were not different significantly between sudangrass and sorghum in short arm,long arm and absolute total length(P＞0.05).
The Karyotypes of F_1 hybrids and their parents were analyzed,and the meiosis behavior of 3 F_1 hybrids and the chromosome numbers of 2 F_2 hybrids were observed The result showed that the karyotypic types of sorghum and sudangrass were 1A,and the karyotype formula of Sa was 2n=18m+2sm(sat);the karyotype formulas of 3042A and 3042A×Sa F_1 were 2n=20m,but the others were 2n=20m(sat).The difference among 3 groups(i.e.sudangrass,sorghum and sorghum-sudangrass hybrid F_1) weren't significant in absolute long arm,absolute short arm,absolute total length,arm ratio(l/s),relative total length from 1st to 10th chromosome pair(P＞0.05);This result supported above viewpoint. The paired chromosome configuration of sorghum-sudangrass hybrid F_1 in pollen mother cells at metaphase I was 2n=2x=20(10Ⅱ),and the chromosome pairing behavior was regular,but the frequencies of rod bivalent of Tx623A×S722 F_1,3042A×Sa F_1 and Tx623A×Sa F_1 were different,which were 4.887,5.710 and 5.126,respectively;that of ring bivalent were 5.113,4.290 and 4.874,respectively.At anaphase I,the paired chromosomes of sorghum-sudangrass hybrid F_1 could separate from each other.The chromosome numbers of F_2 hybrid were 20(2n=20).Therefore,the sudangrass/sorghum relationship is sufficiently close.
3.Photosynthetic Characteristics,Peroxidase(POD) and Esterase(EST) Isozyme Analysis in Sudangrass and Sorghum
Six sorghums and four sudangrasses varieties was assayed by preoxidase(POD) and EST isozyme based on Polyacryamide gel electropheresis(PAGE),respectively.The results showed that preoxidase isozyme could determine different characteristics of different varieties with enzyme patterns difference,but could not distinguish sorghum with sudangrass.EST isozyme could determine characteristic of different varieties with enzyme pattern difference,and could distinguish sudangrass and sorghum.
Four photosynthetic indices of sorghum and sudangrass were measured under strong and weak light conditions by CI-310 Portable Photosynthesis System.The results showed that the differences of net photosynthetic rate,transpiration rate,stomatal conductance of sorghum and sudangrass were significant between weak(PAR:353.07μmol·m~(-2)s~(-1),in the overcast day) and strong(PAR:1331.86μmol·m~(-2)s~(-1),in clear-cloudy day,) radiation,but internal CO_2 of sudangrass was not changed significantly.Under weak radiation,the difference of every index between sorghum and sudangrass was not significant.But transpiration rate and internal CO_2 were significant between sorghum and sudangrass under strong radiation.Under weak radiation,there were significantly positive correlated among net photosynthetic rate,transpiration rate and stomatal conductance in sorghum or sudangrass.Under strong radiation,there were not significant correlated among all indices of sudangrass;but there were significantly positive correlated among most indices in sorghum.
4.Diversity Comparison and Phylogenetic Relationships of S.bicolor and S.sudanense as Revealed by RAPD and SSR markers
Thirty-two accession of S.bicolor,ten varieties of S.sudanense and two related species of S.bicolor were analyzed using random amplified polymorphic DNA(RAPD) markers.The results were as follows:(1)Atotal of 68 bands were amplified by 12 primers, among which 52(76.5%) bands were found to be polymorphic.(2)The similarity coefficient of S.bicolor and S.sudanense were 55%to 95%and 52%to 84%,respectively.The polymorphy is high in both.Restorer and maintainer lines of Sorghum bicolor can be differentiated by RAPD although the similarity of them was more than 89%.(3)Cluster analysis showed the 44 varieties fell into 10 groups by the threshold value of 0.66.The whole accession of the first group were S.sudanense and the whole accession of the 5,6,7 and 8 groups were S.bicolor.But there were S.sudanense and S.bicolor in the 2,3 and 4 groups.It suggested that S.bicolor and S.sudanense can not be distinguished by RAPD.
Phylogenetic relationships among 48 accessions of sorghum(S.bicolor),sudangrass (S.sudanense) and their relatives(S.propinquum,S.halepense,Zea mexicana,Z.mays) were investigated using SSR markers.The 91 SSR primer pairs generated a total of 723 polymorphic alleles,with an average of 7.945 alleles per locus,and a range of 2 to 19 alleles.The average genetic diversity,as measured by the polymorphic information content (PIC),was 0.783.The average polymorphic rates were 84.530%and 78.483%within sorghum and sudangrass,respectively.The PIC values were 0.774 and 0.770,respectively, and there was no significant difference(P＞0.05) between sorghum and sudangrass. Additionally,the genetic distance(GD_(sor-su)) between sorghum and sudangrass was only 0.035,suggesting a high degree of genetic homogeneity.Genetic similarity(GS) values between all varieties ranged from 0.217 to 0.975 and were used to produce a dendrogram. The 48 accessions were clustered into five groups(GS=0.766),specifically,groupⅠ(consisting of sorghum,sudangrass and sorghum-sudangrass hybrids),groupⅡ(S. propinquum),groupⅢ(S.halepense),groupⅣ(Z.mexicana) and group V(Z.mays). Results of our analyses suggest the sudangrass/sorghum relationship is sufficiently close to place them both within the same species-sorghum(S.bicolor).
5.Heterosis Utilization of Hybrid between Sorghum[S.bicolor(L.) Moench]and Sudangrass[S.sudanense(Piper) Stapf]
The morphological characteristics,heterosis and the forage value of hybrid between sorghum(S.bicolor) and sudangrass(S.sudanense) were studied.The results showed that the hybrid vigor was obvious.F_1 was higher than mid-parent,even higher than high-parent in leaf width,leaf length,plant height,tiller and fresh weight per plant.The forage yield of hybrid was higher significantly than that of sudangrass.Main stem diameter and leaf length had strong direct effects on fresh weight per plant,and the same was true for density,leaf length and weight per plant on yield per unit area.The path coefficient of density with yield per unit area(p=1.0299~(**)) was significant at 0.01 level.The best density suitable for the hybrid was 30×10~4 plants/hm~2.The nutrient level of the hybrid was close to that of sudangrass.CN~- content in the hybrid was lower.Sugar content was 10.0%(Bx) at later stage of booting,and it was higher than other developmental stages.
6.Establishment of DNA Fingerprinting and Mapping of the Brown Midrid Gene (bmr-6) for Sorghum/Sudangrass and Hybrids
Nine RAPD primers with high polymorphism and repeatability were screened from 100 RAPD primers,and the polymorphism rate was 64.06%.Every accession could construct a special data fingerprint by 4 RAPD core primers in 42 sorghum and sudangrass accessions.One RAPD fingerprint was constructed by primers F-01,and could differentiate two sorghum-sudangrass hybrids,but didn't distinguish Wancao No 3 and Sa.73 SSR primer pairs were screened from 95 SSR primer pairs,and the polymorphism rate was 86.06%.Each of 42 sorghum and sudangrass accessions could construct a special data fingerprint by 3 SSR core primer pairs.Primer txp18 showed obvious complementary band between the two sorghum-sudangrass hybrids.One SSR fingerprint was constructed by primer txp18,which could distinguish Wancao No 2,Wancao No 3 and their parents.
The midrib and stem marrow were brown in the brown midrib sorghum/sudangrass varieties.The scarfskin was dark brown.The content of lignin reduced 40%to 60%than normal varieties.So the digestibility increased effectively.F_1 and F_2 population were constructed by the hybrid of Tx623A(brown midrib,bmr-6) and Sa-line(white midrib).F_1 generation showed white midrib.The ration of white and brown midrib was suitable to 3:1 in F_2 generation.The results showed that the brown midrib gene(bmr-6) was a single recessive gene.Using microsatellite(SSR) markers and F_2 segregation population,one SSR marker was linkage with the brown midrib gene.The linkage map distance was 4.2 cM.The SSR marker was in LG-07.Therefore,the brown midrib gene(bmr-6) was mapped in LG-07.
高粱与苏丹草没有生殖隔离,这就使利用高粱与苏丹草的杂种优势成为可能。开展高粱-苏丹草杂种优势利用及其指纹图谱和分子标记研究,对饲用高粱和苏丹草遗传育种具有重要的理论和实际意义。
本研究旨在通过数值分类学、细胞遗传学、生理生物化学和分子生物学等方法,探讨苏丹草与高粱形态学与分子性状的遗传亲缘关系,开展高粱与苏丹草杂交优势利用研究,建立高粱与苏丹草及其杂交种DNA指纹图谱,进行高粱/苏丹草褐色中脉基因的遗传分析和SSR标记定位,为进一步选育更加高产优质品种创造前提和条件。本研究获得的主要结论如下:
1.高粱与苏丹草生物学特性的数值分析
选取高梁属不同来源和类型高粱品种6个,苏丹草品种4个,另加1份与苏丹草和高粱染色体数相同的饲用墨西哥玉米(Zea Mexicana)作为参照物种开展数值分析。在植物营养生长阶段和生殖生长阶段观测高粱、苏丹草生物学性状共64个。对苏丹草与高粱10个品种共300个个体的两组判别分析表明,苏丹草与高粱品种间F测验达极显著水平。因此,苏丹草与高粱类间差异明显。至于苏丹草类内和高粱类内部分品种间F测验显著,说明苏丹草与高粱类内又存在分化现象。对苏丹草与高粱的10个品种各30株共300个个体进行判别分析,结果每品种的30个个体都能回判到各自品种中,表明参试的30个个体是合理的。用观测的64个性状指标对苏丹草和高粱共10个品种进行分层聚类,结果苏丹草和高粱明显被分成两大类。
与此同时,本研究还系统分析了苏丹草和高粱64个生物学性状的分类学意义;利用主成分分析法确定出6个主分量,在特定生态条件和栽培模式下,主成分Ⅰ、Ⅱ值的大小决定着苏丹草和高粱共同祖先向各自发展方向进行演化,以6个主成分的回归因子得分值进行10个品种的快速聚类,结果苏丹草和高粱被分成2大类。进一步扩大参试品种的数量,对48个苏丹草和高粱及其近缘种进行数量分类研究,结果基本支持原10个品种的结论,但全部苏丹草和高梁两类群不能被完全分离开来。
2.高粱与苏丹草及其杂种的细胞遗传学研究
采用去壁低渗火焰干燥法对4个苏丹草和6个高粱品种的核型进行比较研究,结果表明苏丹草和高粱体细胞染色体数均为20(2n=20)。苏丹草品种均为1A核型,并具有1对随体染色体,其中1、2号品种的第2对染色体为近中间着丝点染色体,3、4号品种为中间着丝点染色体;高粱品种除5号为1A核型、中间着丝点染色体外,其余的5个品种为2A或2B核型,且都有1或2对近中间着丝点染色体,9号品种还出现1对近端着丝点染色体;高粱8、9号品种各观察到1对随体染色体。染色体数量分析表明,第1到第10对染色体长短臂的绝对长度、相对长度以及绝对全长在苏丹草和高粱两类间的差异均不显著(P＞0.05)。因此,苏丹草和高粱的遗传差异不在染色体长度上。
进一步观察了3个杂种F_1减数分裂染色体行为和2个杂种F_2体细胞中染色体数目。结果杂种F_1花粉母细胞减数分裂过程中,中期Ⅰ染色体核型和数目清晰可见(2n=2x=20),配对行为规则;棒状和环状二价体的频率因组合不同而有差异,Tx623A×S722 F_1、3042A×Sa F_1和Tx623A×Sa F_1棒状二价体频率分别为4.887,5.710和5.126,环状二价体频率分别为5.113,4.290和4.874;在后期Ⅰ,配对的染色体能够正常分离。杂种F_2体细胞中染色体数目为20(2n=20)。说明苏丹草与高粱染色体具有很高的同源性,能够正常配对并稳定传递,二者亲缘关系非常近。
3.高粱与苏丹草同工酶以及光合特性的比较研究
采用聚丙烯酰胺电泳法,对4个苏丹草和6个高粱品种进行过氧化物酶同工酶的分析比较,结果表明苏丹草和高粱各品种的酶带不完全相同,但却未能找到苏丹草和高粱两类的特异带谱。进一步对以上材料进行酯酶同工酶分析,结果苏丹草被聚为一类,高粱被聚为另一类,因此,酯酶同工酶谱可以作为鉴别苏丹草和高粱的有效生化标记。
采用CI-310便携式光合测定系统,对32份高粱和10份苏丹草品种进行强弱光下光合特性的比较研究。结果表明:高粱和苏丹草的各项光合指标除后者的胞间CO_2浓度外,在强弱两种光照条件下均有显著或极显著差异。在弱光条件下,各项光合指标在高粱和苏丹草间均没有显著差异;但在强光条件下,高粱的蒸腾速率和胞间CO_2浓度极显著或显著超过苏丹草。在弱光条件下,无论高粱或苏丹草,其净光合速率、蒸腾速率和气孔导度三者间均呈显著或极显著正相关;但在强光条件下,苏丹草的各项指标无显著相关,而高粱大部分指标间相关极显著。
4.应用RAPD和SSR分子标记对高粱和苏丹草遗传多态性的研究
选用了32份高粱品种、10份苏丹草品种及2份高粱近缘种拟高粱和假高粱进行了RAPD分析。结果表明:12对RAPD引物产生的68条DNA扩增片段中,52条(76.5%)具有多态性。高粱之间的相似系数从55%到95%;苏丹草之间的相似系数从52%到84%,因此选择的品种之间的多态性高,具有代表意义。高粱不育系和保持系之间的相似度很大为89%以上。以0.66为阈值将44个品种分为十个类群。第1类群为4份苏丹草品种;在2、3、4群中既有苏丹草也有高粱;5、6、7、8群中则全部为高粱。因此,从RAPD分子标记聚类结果看,高粱和苏丹草遗传差异较小,难以完全分开。
用SSR标记分析了48份高粱、苏丹草及其近缘植物拟高粱、假高粱、墨西哥玉米、玉米的多态性及其亲缘关系。91对SSR引物在48个品种中扩增得到723个等位基因,平均每个位点有7.945个等位基因,多态性率达98.480%,多态性信息含量PIC为0.783。其中,在高粱品种内扩增出669个等位基因,平均每个位点有7.352个,多态性率为84.530%,多态性信息含量PIC为0.774;在苏丹草品种内扩增出649个等位基因,平均每个位点有7.132个,多态性率为78.483%,多态性信息含量PIC为0.770。通过对高粱与苏丹草多态性信息含量PIC值差异性t检验表明,二类群间无显著差异(p＞0.05)。高粱和苏丹草两类间平均遗传距离仅0.035。全部48份材料间的遗传相似系数最小为0.217,最大为0.975,据此进行聚类分析,结果显示:以相似系数0.766作为分类临界值,可以将48份材料分成5类,即全部高粱和苏丹草品种聚为1类,拟高梁(S.propinquum)1类,假高粱(S.halepense)1类,墨西哥玉米(Zeamexicana)1类,玉米(Zea mays)1类。因此,将苏丹草作为1个独立的植物分类种并不合适,建议将其划归双色高粱种(S.bicolor)。
5.高粱与苏丹草杂种优势利用的研究
苏丹草(S.sudanense)与高粱(S.bicolor)均为禾本科高粱属植物,二者的杂种优势明显。20世纪80年代,作者首次在我国开展高粱与苏丹草杂种优势利用的研究,研究结果表明:F_1单株生物产量性状高于双亲平均值,甚至超过或接近高亲。影响高粱-苏丹草杂交种单株茎叶鲜重的主要因素是主茎粗、叶长;决定单位面积鲜草产量的主要是密度、叶长和单株茎叶鲜重,其中在一定范围内密度对单位面积产量的直接通径系数最大,为1.0299~(**)。多年随机区组试验结果表明,杂交种单位面积鲜草产量显著超过苏丹草,并以30万株/hm~2为最适种植密度。杂交种营养品质与苏丹草相近,氰化物含量低,在孕穗后期含糖量达到最大值,为10.0%(Bx)。
6.高粱与苏丹草及杂交种DNA指纹图谱构建和褐色中脉基因的定位
从100个RAPD引物中筛选到9个多态性高、重复性好的引物,多态性条带比率为64.06%,利用4个核心RAPD引物可以为42份高粱与苏丹草品种资源中的每份品种构建一张特定的数字指纹,并通过其中1个引物F-01构建了一张能鉴别2份国审品种皖草2号和皖草3号高粱-苏丹草杂交种的RAPD指纹图谱,不过该图谱不能区别皖草3号与其父本Sa。从95对SSR引物中筛选出多态性丰富的引物73对,多态性条带比率为86.06%,通过3对核心SSR引物就可以构建42份高粱和苏丹草的SSR数字指纹,同时利用其中1对SSR引物txp18,寻找到两个杂交种的互补带,从而构建了两个高粱-苏丹草杂交种的SSR指纹图谱,这张SSR指纹图谱不仅能鉴别皖草2号和皖草3号,还可以把杂交种与其亲本区别开来。
高粱/苏丹草褐色中脉品种的叶片中脉和茎杆髓部是褐色的,茎杆的表皮呈深褐色。此类品种能使难以消化的木质素含量降低40%-60%,从而大幅度地提高消化率。利用高粱经典材料Tx623A的褐色中脉突变体(bmr-6类型)与白色中脉苏丹草(Sa-line)进行杂交,构建F_2分离群体,F_1表现为白色中脉,F_2白褐分离比符合3:1。这说明褐色中脉基因受一对等位基因控制,表现为隐性遗传。用已知定位到连锁群上的SSR标记和分离群体分析法,对褐色中脉bmr-6基因进行了连锁分析,发现1个与褐色中脉bmr-6基因连锁的微卫星标记,连锁距离为4.2cM,并将该基因初步定位于第7连锁群上。
Sorghum[Sorghum bicolor(L) Moench]and sudangrass[S.sudanense(piper) stapf] are two members of the agriculturally-important genus Sorghum(Gramineal),but some disagreement exists as to whether they actually belong to the same species.Hybrids of these two species have exhibited favorable forage yields and overall improved quality and disease resistance.They are widely used in aquaculture,production of livestock food and environmental protection.It is urgent to study the morphological characteristics,heterosis, forage value and molecular genetic for hybrid.
In this research,the genetics of sorghum(S.bicolor) and sudangrass(S.sudanense) was studied by numerical taxonomy,cytogenetics,physiology/biochemistry and molecular biology.Heterosis of hybrid between sorghum and sudangrass was utilized for the first time in China.DNA fingerprints for 42 sorghum and sudangrass varieties and 2 sorghum-sudangrass hybrids were constructed.The brown midrib gene(bmr-6) of sorghum-sudangrass hybrids was mapped.The results were listed as the following:
1.Numerical Analysis of Genetic Variation in Sorghum[S.bicolor(L.) Moench]and Sudangrass[S.sudanense(Piper) Stapf]
Sixty-four biological characteristics were evaluated for a comparative study between sorghum(S.bicolor) and sudangrass(S.sudanense) by means of variance,discrimination and cluster analyses.The significant differences in biological characteristics between sorghum and sudangrass were found.The F-test divergence values between the varieties of sorghum and sudangrass were highly significant.Discrimination analysis of all plants suggested that 100%of originally grouped cases were classified correctly.Cluster analysis showed that the 10 varieties of sorghum and sudangrass in the study fell into 2 groups:four sudangrass varieties making up one kind,and six sorghum varieties making up the other. Analysis of variance showed the characteristics chosen for cluster analysis were reasonable.
The characters of significant positive correlation and significant negative correlation had been found.In terms of a combined line,64 characters were clustered to 5 groups.The taxonomic significance of these characters had also been analyzed.6 principal branch quantitative characters were determined using the principal component analysis.Principal component-Ⅰwas named for principal sudangrass factor and principal component-Ⅱwas named for sorghum principal factor.10 varieties could be divided into 2 main groups depending on the result of cluster analysis by principal component regression values. Sudangrass was clustered to 1 group,and sorghum was clustered to the other group.
The results of quantity classification for 48 varieties,i.e.32 sorghum varieties and 10 sudangrass varieties and 4 relatives,supported basically above viewpoint.
2.Cytogenetical Studies of Relationship on Sudangrass(S.sudanense) and Sorghum (S.bicolor)
The Karyotypes of 4 sudangrass varieties and 6 sorghum varieties during mitosis were investigated using the method of wall degradation and hypotonic treatment of making chromosomal preparations.The results showed that the numbers of chromosome are 20 in sudangrass(S.sudanense) and sorghum(S.bicolor).The karyotypes of sudangrass varieties were 1A,and their karyotype formulas were shown as follows:No1 and No2 2n=18m+2sm(2SAT),No3 2n=18m+2sm(2SAT),No4 2n=20m(2SAT).Their karyotypes were not all same among sorghum varieties.Only No5 was 1A,and all the other varieties were 2A or 2B.The karyotype formulas of sorghum varieties were:No5 2n=20m,No6 2n=16m+4sm,No7 2n=18m+2sm,No8 2n=18m(2SAT)+2sm,No9 2n=16m+2sm(2SAT)+2st,No10 2n=16m+4sm.There were not different significantly between sudangrass and sorghum in short arm,long arm and absolute total length(P＞0.05).
The Karyotypes of F_1 hybrids and their parents were analyzed,and the meiosis behavior of 3 F_1 hybrids and the chromosome numbers of 2 F_2 hybrids were observed The result showed that the karyotypic types of sorghum and sudangrass were 1A,and the karyotype formula of Sa was 2n=18m+2sm(sat);the karyotype formulas of 3042A and 3042A×Sa F_1 were 2n=20m,but the others were 2n=20m(sat).The difference among 3 groups(i.e.sudangrass,sorghum and sorghum-sudangrass hybrid F_1) weren't significant in absolute long arm,absolute short arm,absolute total length,arm ratio(l/s),relative total length from 1st to 10th chromosome pair(P＞0.05);This result supported above viewpoint. The paired chromosome configuration of sorghum-sudangrass hybrid F_1 in pollen mother cells at metaphase I was 2n=2x=20(10Ⅱ),and the chromosome pairing behavior was regular,but the frequencies of rod bivalent of Tx623A×S722 F_1,3042A×Sa F_1 and Tx623A×Sa F_1 were different,which were 4.887,5.710 and 5.126,respectively;that of ring bivalent were 5.113,4.290 and 4.874,respectively.At anaphase I,the paired chromosomes of sorghum-sudangrass hybrid F_1 could separate from each other.The chromosome numbers of F_2 hybrid were 20(2n=20).Therefore,the sudangrass/sorghum relationship is sufficiently close.
3.Photosynthetic Characteristics,Peroxidase(POD) and Esterase(EST) Isozyme Analysis in Sudangrass and Sorghum
Six sorghums and four sudangrasses varieties was assayed by preoxidase(POD) and EST isozyme based on Polyacryamide gel electropheresis(PAGE),respectively.The results showed that preoxidase isozyme could determine different characteristics of different varieties with enzyme patterns difference,but could not distinguish sorghum with sudangrass.EST isozyme could determine characteristic of different varieties with enzyme pattern difference,and could distinguish sudangrass and sorghum.
Four photosynthetic indices of sorghum and sudangrass were measured under strong and weak light conditions by CI-310 Portable Photosynthesis System.The results showed that the differences of net photosynthetic rate,transpiration rate,stomatal conductance of sorghum and sudangrass were significant between weak(PAR:353.07μmol·m~(-2)s~(-1),in the overcast day) and strong(PAR:1331.86μmol·m~(-2)s~(-1),in clear-cloudy day,) radiation,but internal CO_2 of sudangrass was not changed significantly.Under weak radiation,the difference of every index between sorghum and sudangrass was not significant.But transpiration rate and internal CO_2 were significant between sorghum and sudangrass under strong radiation.Under weak radiation,there were significantly positive correlated among net photosynthetic rate,transpiration rate and stomatal conductance in sorghum or sudangrass.Under strong radiation,there were not significant correlated among all indices of sudangrass;but there were significantly positive correlated among most indices in sorghum.
4.Diversity Comparison and Phylogenetic Relationships of S.bicolor and S.sudanense as Revealed by RAPD and SSR markers
Thirty-two accession of S.bicolor,ten varieties of S.sudanense and two related species of S.bicolor were analyzed using random amplified polymorphic DNA(RAPD) markers.The results were as follows:(1)Atotal of 68 bands were amplified by 12 primers, among which 52(76.5%) bands were found to be polymorphic.(2)The similarity coefficient of S.bicolor and S.sudanense were 55%to 95%and 52%to 84%,respectively.The polymorphy is high in both.Restorer and maintainer lines of Sorghum bicolor can be differentiated by RAPD although the similarity of them was more than 89%.(3)Cluster analysis showed the 44 varieties fell into 10 groups by the threshold value of 0.66.The whole accession of the first group were S.sudanense and the whole accession of the 5,6,7 and 8 groups were S.bicolor.But there were S.sudanense and S.bicolor in the 2,3 and 4 groups.It suggested that S.bicolor and S.sudanense can not be distinguished by RAPD.
Phylogenetic relationships among 48 accessions of sorghum(S.bicolor),sudangrass (S.sudanense) and their relatives(S.propinquum,S.halepense,Zea mexicana,Z.mays) were investigated using SSR markers.The 91 SSR primer pairs generated a total of 723 polymorphic alleles,with an average of 7.945 alleles per locus,and a range of 2 to 19 alleles.The average genetic diversity,as measured by the polymorphic information content (PIC),was 0.783.The average polymorphic rates were 84.530%and 78.483%within sorghum and sudangrass,respectively.The PIC values were 0.774 and 0.770,respectively, and there was no significant difference(P＞0.05) between sorghum and sudangrass. Additionally,the genetic distance(GD_(sor-su)) between sorghum and sudangrass was only 0.035,suggesting a high degree of genetic homogeneity.Genetic similarity(GS) values between all varieties ranged from 0.217 to 0.975 and were used to produce a dendrogram. The 48 accessions were clustered into five groups(GS=0.766),specifically,groupⅠ(consisting of sorghum,sudangrass and sorghum-sudangrass hybrids),groupⅡ(S. propinquum),groupⅢ(S.halepense),groupⅣ(Z.mexicana) and group V(Z.mays). Results of our analyses suggest the sudangrass/sorghum relationship is sufficiently close to place them both within the same species-sorghum(S.bicolor).
5.Heterosis Utilization of Hybrid between Sorghum[S.bicolor(L.) Moench]and Sudangrass[S.sudanense(Piper) Stapf]
The morphological characteristics,heterosis and the forage value of hybrid between sorghum(S.bicolor) and sudangrass(S.sudanense) were studied.The results showed that the hybrid vigor was obvious.F_1 was higher than mid-parent,even higher than high-parent in leaf width,leaf length,plant height,tiller and fresh weight per plant.The forage yield of hybrid was higher significantly than that of sudangrass.Main stem diameter and leaf length had strong direct effects on fresh weight per plant,and the same was true for density,leaf length and weight per plant on yield per unit area.The path coefficient of density with yield per unit area(p=1.0299~(**)) was significant at 0.01 level.The best density suitable for the hybrid was 30×10~4 plants/hm~2.The nutrient level of the hybrid was close to that of sudangrass.CN~- content in the hybrid was lower.Sugar content was 10.0%(Bx) at later stage of booting,and it was higher than other developmental stages.
6.Establishment of DNA Fingerprinting and Mapping of the Brown Midrid Gene (bmr-6) for Sorghum/Sudangrass and Hybrids
Nine RAPD primers with high polymorphism and repeatability were screened from 100 RAPD primers,and the polymorphism rate was 64.06%.Every accession could construct a special data fingerprint by 4 RAPD core primers in 42 sorghum and sudangrass accessions.One RAPD fingerprint was constructed by primers F-01,and could differentiate two sorghum-sudangrass hybrids,but didn't distinguish Wancao No 3 and Sa.73 SSR primer pairs were screened from 95 SSR primer pairs,and the polymorphism rate was 86.06%.Each of 42 sorghum and sudangrass accessions could construct a special data fingerprint by 3 SSR core primer pairs.Primer txp18 showed obvious complementary band between the two sorghum-sudangrass hybrids.One SSR fingerprint was constructed by primer txp18,which could distinguish Wancao No 2,Wancao No 3 and their parents.
The midrib and stem marrow were brown in the brown midrib sorghum/sudangrass varieties.The scarfskin was dark brown.The content of lignin reduced 40%to 60%than normal varieties.So the digestibility increased effectively.F_1 and F_2 population were constructed by the hybrid of Tx623A(brown midrib,bmr-6) and Sa-line(white midrib).F_1 generation showed white midrib.The ration of white and brown midrib was suitable to 3:1 in F_2 generation.The results showed that the brown midrib gene(bmr-6) was a single recessive gene.Using microsatellite(SSR) markers and F_2 segregation population,one SSR marker was linkage with the brown midrib gene.The linkage map distance was 4.2 cM.The SSR marker was in LG-07.Therefore,the brown midrib gene(bmr-6) was mapped in LG-07.
引文
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