基于配合力和遗传距离的甜高粱杂种优势预测

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王黎明^,, 严洪冬^,, 焦少杰, 姜艳喜, 苏德峰, 孙广全黑龙江省农业科学院作物资源研究所,哈尔滨 150086

Heterosis Prediction of Sweet Sorghum Based on Combining Ability and Genetic Distance

WANG LiMing^,, YAN HongDong^,, JIAO ShaoJie, JIANG YanXi, SU DeFeng, SUN GuangQuanCrop Resource Institute of Heilongjiang Academy of Agricultural Sciences, Harbin 150086

王黎明和严洪冬为同等贡献作者
责任编辑: 李莉
收稿日期:2019-07-9接受日期:2019-09-20网络出版日期:2020-07-16

基金资助:

国家现代农业产业技术体系.CARS-06-13.5-A1
国家国际科技合作项目.2011DFR30840-10

Received:2019-07-9Accepted:2019-09-20Online:2020-07-16
作者简介 About authors
王黎明,Tel：0451-86668645;E-mail：dawnw@126.com。

严洪冬,E-mail：hljcrop@163.com。

摘要
【目的】对甜高粱主要农艺性状进行杂种优势、一般配合力及特殊配合力分析,同时,分析配合力、表型遗传距离以及分子遗传距离用于杂种优势预测的可行性,为甜高粱的种质创新和杂交种选育提供理论参考。【方法】采用不完全双列杂交设计,以8个甜高粱不育系为母本及8个甜高粱恢复系为父本配制64个杂交组合。对亲本及杂交后代进行2年的性状调查,包括：出苗至开花日数、生育期、株高、穗长、茎粗、分蘖、单穗粒重、千粒重、籽粒产量、单株重、生物产量和含糖量。分析不同性状的杂种优势、一般配合力、特殊配合力、表型遗传距离、分子遗传距离以及配合力、遗传距离与杂种优势的相关性。【结果】各性状的中亲优势由强到弱分别为单株重、籽粒产量、单穗粒重、生物产量、株高、穗长、千粒重、茎粗、生育期、至开花日数、分蘖和含糖量,其中,生育期、至开花日数、分蘖和含糖量等性状为负优势。不同性状的中亲优势和超亲优势由强到弱的顺序基本相同。配合力分析表明,每个性状中,不同亲本的一般配合力相差较大,且不同组合的特殊配合力也有很大差异。大多数特殊配合力高的组合,其亲本的一般配合力也较高。杂种优势与配合力和遗传距离的相关性为单株重、籽粒产量、单穗粒重、生物产量、穗长、千粒重、分蘖以及含糖量等性状的杂种优势与其亲本的一般配合力和特殊配合力均为极显著正相关。生育期的杂种优势与特殊配合力为极显著正相关,至开花日数与特殊配合力为显著正相关。亲本间的表型遗传距离为2.86—6.82,分子遗传距离为0.50—0.96。单株重、籽粒产量、单穗粒重、生物产量、株高、穗长、茎粗及含糖量等性状的杂种优势与分子遗传距离的相关性大于表型遗传距离,其中,生物产量、单株重、穗长和茎粗的杂种优势与分子遗传距离为极显著正相关。【结论】所有性状中,与产量相关性状的杂种优势较高,而含糖量和分蘖的杂种优势较低。在杂种优势预测上,利用亲本的配合力可有效预测杂种优势,预测效果优于遗传距离。与表型遗传距离相比,分子遗传距离对杂种优势的预测更有效。
关键词： 甜高粱;杂种优势;配合力;遗传距离;预测

Abstract

【Objective】 The heterosis, general combining ability (GCA) and special combining ability (SCA) of main agronomical traits of sweet sorghum were analyzed. Meanwhile, the feasibility of heterosis prediction based on combining ability, phenotypic genetic distance (PGD) and molecular genetic distance (MGD) was estimated. The result will provide theoretical reference for germplasm improvement and hybrid breeding of sweet sorghum.【Method】 Sixty-four hybridized combinations were generated with 8 sterile lines as females and 8 restorer lines as males in accordance with a North Carolina Design II mating scheme. Twelve agronomical traits including days to flowering, growth duration, plant height, panicle length, stem diameter, tillers, panicle weight, 1000-grain weight, grain yield per plot, biomass per plant, biomass per plot and sugar content of hybrids and their parent lines were investigated in two years. The heterosis, GCA, SCA, PGD and MGD were analyzed, as well as the correlation between combining ability, genetic distance and heterosis. 【Result】 Mid-parent heterosis (MPH) of traits from high to low was: biomass per plant, grain yield per plot, panicle weight, biomass per plot, plant height, panicle length, 1000-grain weight, stem diameter, growth duration, days to flowering, tillers and sugar content. Among which, heterosis of growth duration, days to flowering, tillers and sugar content was negative. The sequence of MPH for all traits was almost the same as that of better parent heterosis (BPH). Combining ability analysis showed that the GCA of different parents varied greatly in each trait. Meanwhile, the SCA of different combinations was also significantly different. Most combinations with high SCA also showed high GCA in their parent lines. Heterosis of biomass per plant, grain yield per plot, panicle weight, biomass per plot, panicle length, 1000-grain weight, tillers and sugar content was positively and highly significantly correlated with GCA and SCA of their parent lines simultaneously. Heterosis of growth duration was positively and highly significantly correlated with SCA. Meanwhile, heterosis of days to flowering was positively and significantly correlated with SCA. The PGD of parent lines was 2.86-6.82, and MGD was 0.50-0.96. The correlation between heterosis and MGD was greater than that of PGD in biomass per plant, grain yield per plot, panicle weight, biomass per plot, plant height, panicle length, stem diameter and sugar content. Among which, heterosis of biomass per plot, biomass per plant, panicle length and stem diameter was positively and highly significantly correlated with MGD. 【Conclusion】 Heterosis of yield related traits was higher, and that of sugar content and tillers was lower among all traits. In the prediction of heterosis, combining ability was more effective than genetic distance, therefore, can be used to predict heterosis. MGD was more effective in heterosis prediction compared with PGD.

Keywords：sweet sorghum;heterosis;combining ability;genetic distance;prediction

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本文引用格式
王黎明, 严洪冬, 焦少杰, 姜艳喜, 苏德峰, 孙广全. 基于配合力和遗传距离的甜高粱杂种优势预测[J]. 中国农业科学, 2020, 53(14): 2786-2794 doi:10.3864/j.issn.0578-1752.2020.14.003
WANG LiMing, YAN HongDong, JIAO ShaoJie, JIANG YanXi, SU DeFeng, SUN GuangQuan. Heterosis Prediction of Sweet Sorghum Based on Combining Ability and Genetic Distance[J]. Scientia Acricultura Sinica, 2020, 53(14): 2786-2794 doi:10.3864/j.issn.0578-1752.2020.14.003

0 引言

【研究意义】栽培高粱根据其农艺类型分为粒用高粱、甜高粱、帚高粱、苏丹草及其与高粱的杂交种^[1]。甜高粱（Sorghum bicolor (L.) Moench）植株高大、茎秆粗壮多汁且富含糖分^[2],籽粒和茎秆同时具有贮存生物量的功能,既可用作粮食^[3],又可用作饲料^[4]和能源作物^[5],在国内外广泛种植^[6,7,8]。利用三系配套的杂种优势进行品种选育已在粒用高粱中普遍应用^[9,10]。与粒用高粱相同,杂种优势利用在甜高粱的品种选育中同样具有极其重要的作用,因而,一直是甜高粱育种的主要研究内容^[11,12]。在杂种优势利用中,亲本的一般配合力和特殊配合力对后代的性状表现具有重要作用,是评价亲本利用价值的重要指标和组配强优势杂交后代的基础^[13,14]。同时,亲本间的遗传距离可数量化地体现亲本的遗传差异,在配制优异杂交组合时对亲本的选配原则具有指导意义^[15,16]。因此,研究亲本的配合力和遗传距离在杂种优势预测中的作用将有效提高甜高粱的杂种优势利用效率。【前人研究进展】在甜高粱的杂种优势研究方面,韩东倩等^[17]研究的9个性状中,生物量的杂种优势最高,而籽粒产量的杂种优势最低,且只有生物量的杂种优势为正值。但BUNPHAN等^[18]通过对15个杂交后代7个性状的杂种优势研究,发现籽粒产量的杂种优势最高,而生物产量的杂种优势中等。在配合力研究方面,UMAKANTH等^[19]研究表明生物产量和籽粒产量的特殊配合力高于一般配合力,茎秆重和含糖量的一般配合力高于特殊配合力。而MAKANDA等^[20,21]认为一般配合力和特殊配合力对含糖量、生物产量、株高及生育期等性状同样重要。在高粱的遗传距离与杂种优势的相关性方面,只对粒用高粱进行了少量研究。其中,侯荷亭等^[22]采用最短距离法分析表型遗传距离与杂种优势和配合力的相关性表明,杂种优势在一定范围内随表型遗传距离的增大而增强,但遗传距离太大或太小都会使特殊配合力降低。而AMELEWORK等^[23]对粒用高粱的研究表明,表型遗传距离和分子遗传距离与籽粒产量杂种优势的相关性均不显著。但王瑞等^[24]认为高粱亲本间的分子遗传距离与杂种优势水平关系密切。【本研究切入点】甜高粱的杂种优势、配合力及杂种优势预测等方面的研究结果差异较大。主要原因可能是由于试验材料的选择差异大,有些研究的亲本材料遗传背景差异小,且用于分析的杂交组合数量有限,缺乏代表性。同时,多数研究只针对少数几个性状进行研究,缺乏系统性。尤其对甜高粱的遗传距离与杂种优势的相关性方面缺少研究。【拟解决的关键问题】本研究利用8个甜高粱不育系和8个恢复系进行不完全双列杂交,配制64个杂交组合。对杂交后代的含糖量、产量、生育期及株形等主要农艺性状进行系统研究,分析性状间的杂种优势、一般配合力及特殊配合力,同时,分析配合力、表型遗传距离以及分子遗传距离用于杂种优势预测的可行性。为甜高粱的种质创新和杂交种选育奠定基础。

1 材料与方法

1.1 试验材料

分别利用在农艺性状上差异较大的8个甜高粱不育系为母本及8个甜高粱恢复系为父本配制杂交组合,其中,307A和8431为对照,454A、4190A和314A为生产上常用的亲本材料（电子附表1）。

1.2 性状测定方法

采用不完全双列杂交设计（NCⅡ设计）,以8个不育系和8个恢复系为亲本共配制64个杂交组合。于2012年和2013年种植在黑龙江省农业科学院（哈尔滨）试验基地,随机区组排列,行长5 m,5行区,垄距65 cm,3次重复。对亲本及杂交组合进行2年的主要农艺性状调查,调查项目包括出苗至开花日数、生育期、株高、穗长、茎粗、分蘖、单穗粒重、千粒重、籽粒产量、单株重、生物产量和含糖量等性状。所有性状的调查标准参照陆平^[25]的方法。其中,含糖量的测定为田间采样后,整株榨汁,用测糖仪进行测定,以锤度计算。选择中间3行测定籽粒产量和生物产量。

1.3 分子遗传距离测定

利用筛选出的41对多态性高、扩增稳定且分布于所有连锁群上的SSR引物用于PCR扩增（电子附表2）。DNA提取及PCR扩增的具体方法见文献[26]。

1.4 数据分析

运用SAS软件对杂种优势、一般配合力、特殊配合力、表型遗传距离和分子遗传距离进行分析。其中,杂种优势分析包括中亲优势和超亲优势,计算公式为：

中亲优势（%）=[hF₁-h(p₁+p₂)/2]/h(p₁+p₂)×100;

超亲优势（%）=(hF₁-hHp)/hHp×100;

式中,hF₁为杂交后代值,hp₁为母本值,hp₂为父本值,hHp为高亲值。

参照孔繁玲等^[27]方法计算一般配合力、特殊配合力和表型遗传距离。其中,表型遗传距离采用欧氏距离。

按照NEI等^[28]公式计算亲本间的遗传相似系数GS=2Nxy/(Nx+Ny),其中,GS为亲本间的相似系数,Nxy代表2个亲本共有的等位基因,Nx和Ny分别代表亲本x和y的等位基因数,遗传距离GD=1-GS。

Table 2
表2
表2主要农艺性状的超亲优势分析
Table 2Better parent heterosis analysis of main agronomical traits

性状 Trait	单株重 BP	籽粒产量 GY	单穗粒重 PW	生物产量 BM	株高 PH	穗长 PL	千粒重 TGW	茎粗 SD	生育期 GD	至开花日数 FD	含糖量 SC	分蘖 TL
最大Maximum	158.30	281.97	257.78	108.49	63.23	53.33	50.00	44.44	6.77	10.16	49.56	333.33
最小Minimum	-8.55	-36.87	-29.47	-24.11	-22.27	-7.41	-23.23	-21.67	-14.18	-19.30	-62.97	-100.00
平均Mean	61.70	50.31	48.12	36.99	13.74	13.00	7.23	-2.72	-3.89	-6.95	-25.67	-41.00

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2 结果

2.1 主要农艺性状的杂种优势分析

2.1.1 中亲优势分析各性状的中亲优势分析结果（表1）表明,不同性状的中亲优势差别很大,由强到弱分别为单株重、籽粒产量、单穗粒重、生物产量、株高、穗长、千粒重、茎粗、生育期、至开花日数、分蘖和含糖量。产量性状的中亲优势强于其他性状,其中,单株重的中亲优势最高,平均为89.17,且所有组合均为正值。与产量性状相比,株高的中亲优势略低,均为正优势,且最高值较高,为633.33。另外,生育期、至开花日数、分蘖和含糖量的平均中亲优势均为负值,其中,含糖量的中亲优势最低。至开花日数和生育期的中亲优势组合间最高值和最低值相差较小。分蘖的中亲优势平均值虽然为负值,但后代的杂种优势差异很大,最高值达766.67,最低值为-100.00,可利用组合间的差异性选择目标性状。各性状中,至开花日数、生育期、茎粗、分蘖、单穗粒重和含糖量等性状中,超过半数杂交组合的中亲优势高于对照组合,而株高、穗长、千粒重、籽粒产量、生物产量和单株重等性状中,只有少数组合的中亲优势高于对照组合。

Table 1
表1
表1主要农艺性状的中亲优势分析
Table 1Mid-parent heterosis analysis of main agronomical traits

性状 Trait	单株重 BP	籽粒产量 GY	单穗粒重 PW	生物产量 BM	株高 PH	穗长 PL	千粒重 TGW	茎粗 SD	生育期 GD	至开花日数 FD	分蘖 TL	含糖量 SC
最大Maximum	190.30	302.08	276.61	141.40	633.33	53.33	57.93	44.45	6.56	14.34	766.67	54.06
最小Minimum	10.59	-23.30	-23.86	-5.38	14.41	-7.40	-11.47	-17.46	-9.88	-14.29	-100.00	-59.82
平均Mean	89.17	75.39	73.69	59.49	53.64	20.94	17.24	6.51	-0.86	-3.31	-9.45	-17.40

FD: Days to flowering; GD: Growth duration; SD: Stem diameter; TL: Tillers; PH: Plant height; PL: Panicle length; BM: Biomass; BP: Biomass/plant; GY: Grain yield; PW: Panicle weight; TGW: 1000-grain weight; SC: Sugar content. The same as below
FD：至开花日数;GD：生育期;SD：茎粗;TL：分蘖;PH：株高;PL：穗长;BM：生物产量;BP：单株重;GY：籽粒产量;PW：单穗粒重;TGW：千粒重;SC：含糖量。下同

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2.1.2 超亲优势分析从各性状的超亲优势结果（表2）可以看出,杂交后代在各性状中的超亲优势差别很大。超亲优势由强到弱分别为单株重、籽粒产量、单穗粒重、生物产量、株高、穗长、千粒重、茎粗、生育期、至开花日数、含糖量和分蘖。各性状中,产量性状的超亲优势强于其他性状,其中单株重的超亲优势最高,平均为61.70。超亲优势略低的性状为株高、穗长和千粒重。而茎粗、生育期、至开花日数、含糖量和分蘖的超亲优势均为负值。其中,分蘖的超亲优势最低,但后代中的超亲优势差异很大。各性状中,至开花日数、生育期、茎粗、分蘖、单穗粒重和含糖量等性状中,超过半数杂交组合的超亲优势高于对照组合,而株高、穗长、千粒重、籽粒产量、生物产量和单株重等性状中,只有少数组合的超亲优势高于对照组合。

通过对比各性状的中亲优势和超亲优势可以看出,各性状的中亲优势和超亲优势由强到弱的顺序基本相同,只有含糖量和分蘖略有不同。

2.2 杂种优势与配合力的相关性分析

2.2.1 一般配合力分析通过对不育系和恢复系在所有性状中的一般配合力效应进行分析（表3）,结果显示,每个性状中,不同亲本的一般配合力效应相差较大。同时,同一亲本在不同性状的配合力表现上也有很大差异。各性状中,一般配合力最高的亲本分别为至开花日数、籽粒产量、生物产量以及单株重为A3;生育期为R6;株高为R2;穗长以及单穗粒重为R4;茎粗为R5;分蘖为A5;千粒重为A8以及含糖量为R1。以上亲本在多个性状中的一般配合力高于对照。

Table 3
表3
表3亲本主要农艺性状的一般配合力效应
Table 3General combining ability effect of parent lines in main agronomical traits

亲本 Parent lines	至开花日数 FD	生育期 GD	株高 PH	穗长 PL	茎粗 SD	分蘖 TL	籽粒产量 GY	单穗粒重 PW	千粒重 TGW	生物产量 BM	单株重 BP	含糖量 SC
A1	0.46	-0.30	29.12	-0.16	-0.02	0.14	3.22	-0.03	0.70	4.63	30.37	0.31
A2	-1.46	-2.00	13.85	-0.14	-0.01	0.03	-1.25	3.73	0.75	-0.43	6.74	0.11
A3	5.06	4.50	20.37	1.09	-0.02	-0.01	5.93	-3.30	-2.72	7.63	46.36	1.15
A4	-2.40	-2.90	-6.65	0.52	-0.07	-0.09	-8.11	1.60	1.68	-9.12	-93.45	-0.94
A5	2.48	3.90	-12.15	1.19	-0.02	0.17	1.46	-6.61	-3.19	-0.58	11.61	-0.86
A6	1.46	3.70	-12.96	1.17	0.09	-0.09	4.21	4.46	-0.76	5.17	35.83	0.18
A7	-4.46	-4.40	-19.67	-0.89	-0.05	-0.11	-8.11	-0.27	0.82	-10.18	-81.35	0.20
A8	-1.15	-2.50	-11.92	-2.77	0.10	-0.04	2.65	0.41	2.73	2.87	43.89	-0.16
R1	0.79	0.70	18.98	0.54	0.00	-0.12	2.70	0.25	0.62	1.55	9.84	1.55
R2	-0.19	-1.10	75.33	-2.25	-0.04	-0.11	1.14	4.98	-0.84	-0.33	3.69	1.26
R3	-0.65	-1.30	4.41	0.59	-0.09	0.01	-0.16	-5.30	-0.15	-2.02	9.37	-0.34
R4	0.13	-0.60	0.29	1.71	0.04	-0.07	4.26	14.65	2.24	4.09	30.98	1.01
R5	-0.65	0.60	-49.46	-0.33	0.11	-0.02	-4.42	-1.08	0.11	-4.16	-37.20	-0.07
R6	2.94	5.20	-40.40	0.32	0.05	0.13	-0.88	-7.45	-0.17	0.95	2.38	0.27
R7	-1.04	-0.70	-34.11	-0.98	0.01	0.03	-2.03	-2.77	-0.62	-1.64	-20.21	-2.64
R8	-1.33	-2.90	24.96	0.40	-0.08	0.15	-0.57	-3.29	-1.19	1.55	1.15	-1.03

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2.2.2 特殊配合力分析在所测定的杂交组合中,不同亲本间各性状的特殊配合力效应差异较大（表4）。不同组合在同一性状间以及同一组合在不同性状上的特殊配合力表现均有较大差异。其中,大多数性状中特殊配合力最高的组合中至少有1个亲本的一般配合力最高或较高,如株高A3/R2、穗长A3/R4、分蘖A5/R8、籽粒产量A3/R1、单穗粒重A4/R4、千粒重A8/R5、单株重A2/R4、含糖量A1/R4以及至开花日数A5/R6。而生育期、茎粗和生物产量中特殊配合力最高的组合,其亲本的一般配合力均不高,但这几个性状中特殊配合力较高的组合中,其亲本的一般配合力也较高。与对照相比,至开花日数、生育期、茎粗和分蘖等性状中一半以上组合的特殊配合力高于对照,但其他性状中高于对照的杂交组合较少。

Table 4
表4
表4亲本主要农艺性状的特殊配合力效应
Table 4Special combining ability effect of parent lines in main agronomical traits

极值 Extremum	至开花日数 FD	生育期 GD	株高 PH	穗长 PL	茎粗 SD	分蘖 TL	籽粒产量 GY	单穗粒重 PW	千粒重 TGW	生物产量 BM	单株重 BP	含糖量 SC
最大值 Maximum	9.56	7.80	102.16	3.07	0.15	0.61	12.22	25.75	6.06	12.41	269.50	3.81
最小值 Minimum	-7.94	-6.90	-70.67	-4.93	-0.20	-0.31	-16.69	-32.75	-6.84	-14.53	-218.40	-4.93

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2.2.3 杂种优势与配合力的相关性通过对配合力和中亲优势的相关性进行研究（表5）,大多数性状的杂种优势与其亲本的一般配合力和特殊配合力相关显著或极显著。其中,单株重、籽粒产量、单穗粒重、生物产量、穗长、千粒重、分蘖以及含糖量的杂种优势与其亲本的一般配合力和特殊配合力均为极显著正相关。并且,含糖量的杂种优势与一般配合力和特殊配合力的相关程度较高,相关系数均在70以上,同时,籽粒产量与一般配合力的相关系数也达到70以上。在所有相关系数达极显著的性状中,除籽粒产量的杂种优势与一般配合力的相关性显著高于特殊配合力外,其余性状的一般配合力和特殊配合力之间的相关系数差异较小。生育期的杂种优势与特殊配合力为极显著正相关,至开花日数与特殊配合力为显著正相关。而株高和茎粗的杂种优势与各配合力相关均不显著。

Table 5
表5
表5杂种优势与配合力的相关性
Table 5Correlation between heterosis and combining ability

配合力 Combining ability	单株重 BP	籽粒产量 GY	单穗粒重 PW	生物产量 BM	株高 PH	穗长 PL	千粒重 TGW	茎粗 SD	生育期 GD	至开花日数 FD	分蘖 TL	含糖量 SC
一般配合力 GCA	55.79**	71.73**	49.99**	48.36**	3.10	34.19**	32.65**	4.23	8.78	2.66	42.71**	70.02**
特殊配合力 SCA	61.83**	39.66**	49.94**	59.16**	6.82	44.46**	46.26**	8.78	52.44**	30.56*	54.57**	76.43**

GCA: General combining ability; SCA: Special combining ability; *: Significance at P<0.05; **: Significance at P<0.01. The same as below
GCA：一般配合力;SCA：特殊配合力;*：P<0.05水平相关性差异显著;**：P<0.01水平相关性差异显著。下同

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2.3 杂种优势与遗传距离的相关性分析

2.3.1 表型遗传距离分析亲本间的表型遗传距离平均为4.73（表6）,其中,最高值为6.82（A7和R6之间）,最低值为2.86（A3的R2之间）,不同亲本的表型性状差异较大。

Table 6
表6
表6亲本间的表型遗传距离
Table 6Phenotypic genetic distance (PGD) between parent lines

亲本Parent lines	A1	A2	A3	A4	A5	A6	A7	A8
R1	4.49	3.44	4.86	3.46	5.97	5.35	5.90	5.04
R2	5.03	3.41	2.86	4.14	5.71	5.15	4.64	5.68
R3	4.61	3.30	4.37	3.78	4.89	4.76	4.39	4.40
R4	5.20	4.13	3.91	4.09	6.10	5.50	6.32	6.49
R5	3.97	3.51	2.95	3.86	6.44	5.57	5.84	5.25
R6	4.74	5.05	4.48	5.58	5.92	5.91	6.82	6.45
R7	5.10	4.11	4.16	4.03	3.75	3.83	3.96	3.48
R8	3.95	4.46	4.98	4.46	5.08	4.67	4.44	4.29

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2.3.2 分子遗传距离分析利用SSR引物扩增后,每个引物检测到的等位基因为2—11个,平均4.6个。引物的多样性指数介于0.094—0.870,平均为0.586。

分子遗传距离结果（表7）表明,亲本间的分子遗传距离平均为0.74,其中,最高值为0.96（A3和R6之间）,最低值为0.50（A5和R5之间）,说明不同亲本在遗传水平上差异较大。

Table 7
表7
表7亲本间的分子遗传距离
Table 7Molecular genetic distance (MGD) between parent lines

亲本Parent lines	A1	A2	A3	A4	A5	A6	A7	A8
R1	0.70	0.75	0.78	0.76	0.70	0.71	0.79	0.73
R2	0.69	0.73	0.80	0.75	0.63	0.70	0.81	0.75
R3	0.67	0.68	0.71	0.73	0.65	0.68	0.64	0.70
R4	0.65	0.70	0.74	0.72	0.63	0.70	0.70	0.72
R5	0.84	0.92	0.95	0.89	0.50	0.90	0.90	0.93
R6	0.88	0.93	0.96	0.95	0.60	0.91	0.94	0.94
R7	0.53	0.59	0.63	0.56	0.56	0.59	0.67	0.58
R8	0.72	0.72	0.76	0.78	0.67	0.73	0.76	0.71

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2.3.3 杂种优势与遗传距离的相关性分别对表型遗传距离和分子遗传距离与中亲优势的相关性进行分析（表8）。结果表明,单株重、籽粒产量、单穗粒重、生物产量、株高、穗长、茎粗及含糖量等性状的杂种优势与分子遗传距离的相关性大于表型遗传距离。所有性状的杂种优势与表型遗传距离相关性均不显著,但单株重、生物产量、穗长和茎粗的杂种优势与分子遗传距离为极显著正相关。而籽粒产量、单穗粒重、株高和含糖量等性状的杂种优势与其分子遗传距离虽然相关系数较高,但没达到显著水平。另外,籽粒产量、单穗粒重、株高、千粒重、茎粗、生育期、至开花日数和分蘖的杂种优势与表型遗传距离的相关性较高,也没有达到显著水平。以上结果表明,可利用分子遗传距离对甜高粱部分性状的杂种优势进行预测。

Table 8
表8
表8杂种优势和遗传距离的相关性
Table 8Correlation between heterosis and genetic distance

遗传距离 Genetic distance	单株重 BP	籽粒产量 GY	单穗粒重 PW	生物产量 BM	株高 PH	穗长 PL	千粒重 TGW	茎粗 SD	生育期 GD	至开花日数 FD	分蘖 TL	含糖量 SC
表型遗传距离 PGD	3.84	16.28	15.77	5.52	-16.07	9.35	15.09	20.18	-15.82	-23.93	-20.02	6.54
分子遗传距离 MGD	39.31**	21.23	21.44	42.93**	-21.96	37.89**	5.27	46.45**	-5.79	-0.59	-1.14	17.67

PGD: Phenotypic genetic distance; MGD: Molecular genetic distance
PGD：表型遗传距离;MGD：分子遗传距离

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3 讨论

3.1 甜高粱杂种优势的利用

杂交后代各性状的中亲优势和超亲优势由强到弱的顺序基本相同。在所有性状中,单株重、籽粒产量、单穗粒重及生物产量等产量性状的杂种优势高,其次为株高,说明利用杂种优势易获得植株高大且籽粒产量和生物产量均高的杂交后代。穗长、千粒重和茎粗的杂种优势为中等,杂交后代的优势一般。至开花日数、生育期、分蘖和含糖量的杂种优势均为负优势,说明杂交后代的生育期容易变早,且不易获得分蘖多以及含糖量高的杂交后代。BUNPHAN等^[18]研究也发现甜高粱的生物产量和籽粒产量的杂种优势较高,但含糖量的杂种优势低,研究结果与本研究相近。

从配合力分析结果来看,亲本在不同性状中的一般配合力和特殊配合力效应相差较大。因此,根据不同育种目标,对亲本的一般配合力和特殊配合力可有针对地利用。同时,特殊配合力较高的组合,其亲本的一般配合力往往也较高。以往研究也表明,虽然在一些性状上特殊配合力比一般配合力重要^[12,18-19],但在亲本选择时不能只依靠特殊配合力效应,应将特殊配合力和一般配合力相结合,在一般配合力高的基础上,再选择特殊配合力高的亲本^[20,21],以增加创造优异组合的机率。

此外,在选育甜高粱杂交种时,应根据杂交种的具体用途对不同性状重点选育。当用于提取乙醇或制糖时,要重点对生物产量和含糖量进行选育,而籽粒产量及其他性状则不是选育的重点^[19,29]。当用于青贮饲料时,则要求杂交种的生物产量和籽粒产量都高,但含糖量可以不必太高^[30]。此外,研究表明,黑龙江、吉林、内蒙古以及辽宁等省/自治区是中国利用甜高粱提取乙醇最具生产潜力的省份^[31],为适应这些地区机械化水平相对较高的特点,可通过降低株高、增加种植密度来增加群体生物产量和糖产量的方式,选育适宜机械化种植的甜高粱品种^[6,32],以有效降低生产成本,大力发展甜高粱种植。

3.2 甜高粱的杂种优势预测

在甜高粱的杂交育种中,含糖量及产量性状很难根据亲本的表型性状来预测杂交种的性状表现^[33]。为此,育种者也在探讨更有效的预测方法。随着分子标记的不断发展,利用分子标记进行杂种优势的预测已在多种作物上进行了研究^[34,35,36]。

本研究中,通过对比甜高粱亲本的表型遗传距离和分子遗传距离与杂种优势的相关性发现,大多数性状的杂种优势与分子遗传距离的相关性大于表型遗传距离,其中,单株重、生物产量、穗长和茎粗的杂种优势与亲本的分子遗传距离为极显著正相关。由此可见,与表型遗传距离相比,分子遗传距离对杂种优势的预测更有效。王瑞等^[24]分析了粒用高粱的杂种优势与亲本遗传距离的相关性,也认为高粱亲本的选配应充分考虑遗传距离,为获得杂种优势强的后代,应选择遗传距离大的亲本杂交。同时,WEGARY等^[15]采用欧氏距离的表型遗传距离和SSR分子遗传距离分析与玉米杂种优势的相关性,结果表明,分子遗传距离和籽粒产量为极显著正相关,和至开花期和抽丝期为极显著负相关,认为分子遗传距离对杂交种产量及一些农艺性状的预测比表型遗传距离更有效,研究结果与本研究相近。在水稻^[37]、棉花^[16]等作物中也有相似的研究结果。另外,本研究对甜高粱配合力与杂种优势的相关性研究显示,包括单株重、籽粒产量、单穗粒重、生物产量、穗长、千粒重、分蘖以及含糖量等大多数性状的杂种优势与其亲本的一般配合力和特殊配合力均为极显著正相关,亲本的配合力在甜高粱杂种优势的预测上与遗传距离相比,可对更多性状进行有效预测。AMELEWORK等^[23]和JORDAN等^[38]对粒用高粱的研究表明,表型遗传距离和分子遗传距离与籽粒产量杂种优势的相关性虽然不显著,但和一般配合力效应有关,认为在杂种优势的预测上,基于一般配合力的预测方法比基于分子标记的预测方法更有效。

在不同种质和不同物种上遗传距离用于杂种优势的预测效果不同,可能有以下几种原因。首先,研究结果因所选择材料群体的数量、亲本的多样性以及所研究的性状而不同,且不同类型的农艺性状受环境影响程度也不同。另外,缺乏和测定性状连锁的控制基因,所选择的分子标记和数量具有随机性,使标记在染色体上的分布不均衡以及显性基因的多样性效应等^[39,40]。利用分子多样性来预测杂交种表现的一个基础假设是由于在性状QTL和标记等位基因间存在高水平的配子阶段连锁不平衡^[34,38],因此,当用于计算遗传距离的所选标记比例超过50%以上与目标性状和杂种优势的QTL相关时,则利用分子标记预测杂种优势是可行的^[39,40]。

4 结论

在甜高粱杂种优势预测上,利用亲本的一般配合力和特殊配合力可对单株重、籽粒产量、单穗粒重、生物产量、穗长、千粒重、分蘖、含糖量、至开花日数及生育期等性状的杂种优势进行预测,预测效果优于表型遗传距离和分子遗传距离。与表型遗传距离相比,分子遗传距离对杂种优势的预测更有效,尤其在单株重、生物产量、穗长和茎粗等性状的杂种优势预测上。

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Biomass sorghums are being developed as a dedicated energy crop. These sorghums are managed to maximize biomass yields and are tall (3.5-5 m) with dry, pithy stems. To facilitate seed production, they are produced as hybrids using grain sorghum seed parents. Besides facilitating seed production, these hybrids may provide high-parent heterosis for improved biomass yields. To measure potential heterosis, biomass hybrids derived from 13 different biomass pollinators and 4 grain sorghum seed parents were evaluated for biomass yield in 4 environments. High-parent heterosis in these hybrids averaged 24.8% across all the environments and was available in many hybrid combinations. Hybrids out-yielded the pollinators in each environment. Across environments, the hybrids yielded 32.4 Mg/ha compared to the 27.0 Mg/ha of the pollinators. While heterosis was available in many hybrids, specific combining abilities and environmental interactions combine to maximize heterosis and biomass yields. Heterosis can serve as an effective tool for improving biomass sorghum yields. (C) 2014 Elsevier B.V.

[13]

BIRCHLER

J A

, YAO

, CHUDALAYANDI

, VAIMAN

, VEITIA

R A

. Heterosis
The Plant Cell, 2010,22:2105-2112.

URLPMID:20622146 [本文引用: 1]

[14]

REDDY

B V S

, RAMESH

, REDDY

P S

, RAMAIAH

. Combining ability and heterosis as influenced by male-sterility inducing cytoplasms in sorghum [Sorghum bicolor(L.) Moench]
Euphytica, 2007,154:153-164.

DOI:10.1007/s10681-006-9281-6 URL [本文引用: 1]

An investigation was carried out to assess the efficiency of A₂ cytoplasmic-nuclear male-sterility (CMS) system in comparison to the widely used A₁ cytoplasm in terms of general combining ability (gca) effects of male-sterile (A-) lines and mean performance, specific combining ability (sca) effects and mid-parent heterosis of hybrids for days to 50% flowering, plant height and grain yield at International Crops Research Institute for the Semi-Arid Tropics, Patancheru, Andhra Pradesh, India in 2001 and 2002 rainy seasons. The material for the study consisted of six pairs of iso-nuclear, allo-plasmic (A₁ and A₂) A-lines and 36 iso-nuclear hybrids produced by crossing these A-lines with three dual restorer (R-) lines. The results revealed that cytoplasm and its first-order interaction with year, R- and A-lines did not appear to contribute to variation in iso-nuclear hybrids for plant height and grain yield. Cytoplasm had limited effect on gca effects of A-lines and on sca effects and mid-parent heterosis of iso-nuclear hybrids for days to 50% flowering, plant height and grain yield. The mean days to 50% flowering, plant height and grain yield of A₂ cytoplasm-based hybrids were comparable with those of widely used A₁ cytoplasm-based hybrids. The relative frequency of the occurrence of the A₁- and A₂-based hybrids with significant sca effects and mid-parent heterosis indicated that A₂ CMS system is as efficient as A₁ with a slight edge over A₁ for commercial exploitation. The implications of these results are discussed in relation to opportunities for broadening not only cytoplasm base but also nuclear genetic base of both the hybrid parents.

[15]

WEGARY

, VIVEK

, LABUSCHAGNE

. Association of parental genetic distance with heterosis and specific combining ability in quality protein maize
Euphytica, 2013,191:205-216.

DOI:10.1007/s10681-012-0757-2 URL [本文引用: 2]

Genetic distance analysis among quality protein maize (QPM) inbred lines and the correlation of genetic distance with heterosis would help to design breeding strategy and predict hybrid performance. This study was carried out to determine the amount of genetic diversity among QPM inbred lines using SSR markers and morphological distances; to classify the inbred lines according to their relationships; and to estimate the correlations of SSR markers and morphological distances with hybrid performance, heterosis and specific combining ability (SCA). One-hundred and five hybrids generated by diallel crossing of 15 QPM inbred lines were evaluated with the 15 parents for 17 morphological traits at Harare, Zimbabwe and Bako, Ethiopia and also examined for DNA polymorphism using 40 SSR markers. SSR markers and morphological methods of genetic distance estimates showed moderately high genetic distance among the inbred lines studied. Cluster analysis based on the two distance measures grouped the 15 parental lines differently. The SSR marker-based genetic distance was positively and highly significantly correlated with grain yield (r = 0.37), and negatively and highly significantly with days to anthesis (r = -0.40) and days to silking (r = -0.42). These relationships suggest that high grain yield and earliness of QPM hybrids can be predicted from SSR marker determined distances of the parents, although the correlation values were not very high. The correlations of SSR marker distance with heterosis were too low to be of predictive value except for the case of plant height. Morphological distances were of less importance in predicting hybrid performance and SCA effects of hybrids.

[16]

曲玉杰, 孙君灵, 耿晓丽, 王骁, SARFRAZ

, 贾银华, 潘兆娥, 何守朴, 龚文芳, 王立如, 庞保印, 杜雄明. 陆地棉亲本间遗传距离与杂种优势的相关性研究
中国农业科学, 2019,52(9):1488-1500.

DOI:10.3864/j.issn.0578-1752.2019.09.002 URL [本文引用: 2]

【Objective】The correlation between heterosis and genetic distance (GD) of quantitative traits between parents was analyzed by 1500 hybrid combinations in upland cotton, and the possibility of using GD between parents of large-scale combinations to improve the efficiency of hybrid vigour prediction of upland cotton was discussed in order to provide theoretical guidance for cotton hybrid breeding and utilization of heterosis.【Method】305 upland cotton core collections from 15 countries and 23 provinces (municipalities) of China were selected as parents, and 1500 cross combinations were produced by L×T (Line×Tester) cross design. From 2012 to 2013, ten yield and fiber quality related traits, including plant height (PH), boll weight (BW), boll number per plant (BN), lint percentage (LP), fiber length (FL), fiber strength (FS), fiber elongation (FE), fiber length uniformity (FU), micronaire (MIC) and spinning consistent index (SCI), were investigated in 13 ecological conditions in north and south China. F₁ hybrids mid-parent heterosis (MPH), heterobeltiosis (HB), GD between parents and population structure were analyzed. The correlation between GD and hybrid vigour was calculated by four schemes (Cor1-Cor4). 【Result】The mean values of MPH of the ten traits ranged from 1.70% to 7.40%, with an average of 4.36%, and F₁ hybrids were divided into 5 groups (A-E) according to different male parents, the mean values of MPH: A>E>B>C>D. The mean values of HB ranged from -4.17% to 1.87%, with an average of -0.17%, and the average values of group A, B, and E were positive. In 5 groups, except for MIC of group D and E, other 9 traits had obvious MPH, among them, MPH of BW and FL were mainly positive (more than 80%) in the 5 groups, the maximum MPH values were 34.01% and 9.83% respectively, and the corresponding HB values were 24.25% and 5.80% respectively. The significant difference analysis between F₁ hybrids and their parents indicated that BW, PH, FL, FE, and FU showed some HB. The GDs between male parents (testers) and 300 female parents ranged from 2.280 to 61.430, with an average of 21.550. The mean GDs between 5 testers and female parents: D>C>E>A>B, in which the nearest value was 11.721, and the farthest value was 33.271. According to “Ward” clustering method, 305 upland cotton parents were divided into two groups, including five subgroups. The results of four correlation analysis methods between GD and heterosis showed that the consequences varied with the sample size, the range of GD, and the male parent, the correlation increased with the sample size. Cor1 was the overall embodiment of Cor2 results; compared with Cor1 and Cor2, Cor3 had different correlations between MPH and GD in some traits; Cor4 had the weakest correlations. To sum up, the genetic distance was positively correlated with the MPH of LP, FS, FU, and SCI, the correlation between GD and MPH of other traits was different due to the different analysis schemes. In the four schemes, except for FU, the relationship between GD and HB was negatively correlated on the whole, and there was a strong correlation between genetic distance and HB of MIC, FL and LP. 【Conclusion】There is a linear relationship between GD of quantitative traits and hybrid vigour in upland cotton. The correlations are positive or negative, strong or weak due to different traits, and the larger the sample size, the stronger the correlation. Thus, the large-scale hybrid combinations are used to well study the relationship between GD and heterosis in upland cotton.

Y J

, SUN

J L

, GENG

X L

, WANG

, SARFRAZ

, JIA

Y H

, PAN

Z E

, HE

S P

, GONG

W F

, WANG

L R

, PANG

B Y

, DU

X M

. Correlation between genetic distance of parents and heterosis in upland cotton
Scientia Agricultura Sinica, 2019,52(9):1488-1500. (in Chinese)

DOI:10.3864/j.issn.0578-1752.2019.09.002 URL [本文引用: 2]

韩东倩, 韩立朴, 薛帅, 尤明山, 谢光辉. 基于能源利用的高粱配合力和杂种优势分析
中国农业大学学报, 2012,17(1):26-32.

[本文引用: 1]

HAN

D Q

, HAN

L P

, XUE

, YOU

M S

, XIE

G H

. Combining ability and heterosis of sorghum for biomass energy
Journal of China Agricultural University, 2012,17(1):26-32. (in Chinese)

[本文引用: 1]

[18]

BUNPHAN

, JAISIL

, SANITCHON

, KNOLL

J E

, ANDERSON

W F

. Heterosis and combining ability of F₁ hybrid sweet sorghum in Thailand
Crop Science, 2015,55:178-187.

DOI:10.2135/cropsci2014.05.0363 URL [本文引用: 3]

[19]

UMAKANTH

A V

, PATIL

J V

, RANI

, GADAKH

S R

, KUMAR

S S

, RAO

S S

, KOTASTHANE

T V

. Combining ability and heterosis over environments for stalk and sugar related traits in sweet sorghum (Sorghum bicolor(L.) Moench)
Sugar Technology, 2012,14(3):237-246.

DOI:10.1007/s12355-012-0166-9 URL [本文引用: 3]

[20]

MAKANDA

, TONGOONA

, DERERA

, SIBIYA

, FATO

. Combining ability and cultivar superiority of sorghum germplasm for grain yield across tropical low- and mid-altitude environments
Field Crops Research, 2010,116:75-85.

DOI:10.1016/j.fcr.2009.11.015 URL [本文引用: 2]

[21]

MAKANDA

, TONGOONA

, DERERA

. Combining ability and heterosis of sorghum germplasm for stem sugar traits under off-season conditions in tropical lowland environments
Field Crops Research, 2009,114:272-279.

DOI:10.1016/j.fcr.2009.08.009 URL [本文引用: 2]

[22]

侯荷亭, 杜志宏, 赵根弟. 高粱亲本遗传距离与杂种优势和特殊配合力的关系
遗传, 1995,17(1):30-33.

[本文引用: 1]

HOU

H T

, DU

Z H

, ZHAO

G D

. Studies on the relationships of genetic distance of sorghum parental lines with heterosis and specific combining ability
Hereditas, 1995,17(1):30-33. (in Chinese)

[本文引用: 1]

[23]

AMELEWORK

, SHIMELIS

, LAING

. Genetic variation in sorghum as revealed by phenotypic and SSR markers: Implications for combining ability and heterosis for grain yield
Plant Genetic Resources, 2016,3:1-13.

DOI:10.1079/PGR200566 URL [本文引用: 2]

[24]

王瑞, 王金胜, 张福耀, 程庆军, 田承华, 凌亮. 1970s—2000s中国高粱杂交种亲本遗传距离演变的 SSR 分析
中国农业科学, 2015,48(3):415-425.

DOI:10.3864/j.issn.0578-1752.2015.03.02 URL [本文引用: 2]

【Objective】To explore the genetic differences of Chinese sorghum varieties and evolution of the superior groups since heterosis utilization, SSR markers were used to identify the genetic distance of sorghum parental lines and F₁. Genetic background of Chinese sorghum main varieties in different periods were analyzed, and systematically understand of the genetic relationship, genetic diversity and genetic distance of sorghum hybrids and their parental lines can reduce the blindness of parental lines selection, effectively improve the predictability of cross breeding.【Method】SSR genetic distance of the representative hybrids and their parental lines grown in late-maturing area since 1970s were analyzed. Analysis of the superior groups of sorghum hybrids, research of the breeding progress and the evolution characteristics since heterosis utilization of sorghum were conducted at molecular level. 【Result】 A total 55 hybrids and their parental lines were evaluated by 109 pairs of SSR markers. The results revealed that 47 pairs of SSR markers were highly polymorphic. 373 varied alleles were amplified with an average of 7.5 alleles per marker, the ranges were 2 to 14. The ranges of PIC in 55 cultivars were 0.0351-0.8836, with an average of 0.6085. The genetic distance was averaged at 0.6011 with a range from 0.0889 to 0.9500. The sorghum materials were classified into 4 groups based on the SSR experiments. The clustering results were similar to those based on geographical origins and genetic backgrounds. The results showed that the average genetic distance between the two parental lines of hybrids from different eras showed a upward trend in 1970-1980, then a slight decline. According to the analysis of genetic distance between the two parental lines of hybrids from different heterosis models, Durra sterile lines×Kaoliang or trend Kaoliang restorer lines and Kafir-caudatum sterile lines×Kaoliang or trend kaoliang restorer lines have a far genetic distance, high yield, therefore, the heterosis model of Chinese sorghum hybrids should be Durra×kaoliang or trend kaoliang, Kafir-caudatum×kaoliang or trend kaoliang. 【Conclusion】In this study, it was found that the genetic distance between parental lines has a close relationship with heterosis level in sorghum, therefore, genetic distance should be fully considered in the matching of sorghum parental lines.

WANG

, WANG

J S

, ZHANG

F Y

, CHENG

Q J

, TIAN

C H

, LING

. Evolution of genetic distance between parental lines of Chinese sorghum hybrids from1970s-2000s based on SSR analysis
Scientia Agricultura Sinica, 2015,48(3):415-425. (in Chinese)

DOI:10.3864/j.issn.0578-1752.2015.03.02 URL [本文引用: 2]

陆平. 高粱种质资源描述规范和数据标准. 北京: 中国农业出版社, 2006:51-58.

[本文引用: 1]

. Descriptors and Data Standard for Sorghum [Sorghum bicolor(L.) Moench]. Beijing: China Agricultural Press, 2006:51-58. (in Chinese)

[本文引用: 1]

[26]

WANG

L M

, JIAO

S J

, JIANG

Y X

, YAN

H D

, SU

D F

, SUN

G Q

, YAN

X F

, SUN

L F

. Genetic diversity analysis in parent lines of sweet sorghum based on agronomical traits and SSR markers
Field Crops Research, 2013,149:11-19.

DOI:10.1016/j.fcr.2013.04.013 URL [本文引用: 1]

Agronomical traits and SSR markers were used to analyze genetic diversity of 142 parent lines of sweet sorghum. Parent lines were clustered into 5 groups based on agronomical traits and each group is characterized by morphological traits and main origins. 41 selected SSR markers were used to analyze the parent lines. The genetic distance (GD) of the parent lines ranged from 0.558 to 0.858, averaging 0.640. The average specific index of parent lines was 189.0, which ranged from 109.1 to 454.7. According to SSR markers, parent lines were clustered into 7 groups but the agronomical traits of which were not significantly different between each group, and clusters based on SSR markers did not coincide with the analysis results based on agronomical traits. Cluster analysis was failed to distinctly group restorer lines and maintainer lines by both agronomical traits and SSR markers. Parent lines should be selected both with different agronomical traits and distant genetic distance for higher heterosis. Groups clustered by agronomical traits and molecular markers should be taken into account simultaneously in hybrid breeding. Information of this study can be used to select parent lines for development of segregating populations and hybrid breeding of sweet sorghum. (C) 2013 Elsevier B.V.

[27]

孔繁玲. 植物数量遗传学. 北京: 中国农业大学出版社, 2006:403-412.

[本文引用: 1]

KONG

F L

. Quantitative Genetics in Plants. Beijing: China Agricultural University Press, 2006:403-412. (in Chinese)

[本文引用: 1]

[28]

NEI

, LI

. Mathematical model for studying genetic variation in terms of restriction endonucleases
Proceedings of the National Academy of Sciences of the USA, 1979,76(10):5269-5273.

URLPMID:291943 [本文引用: 1]

[29]

邹剑秋, 王艳秋, 张志鹏, 朱凯. A₃型细胞质能源用甜高粱生物产量、茎秆含糖锤度和出汁率研究
中国农业大学学报, 2011,16(2):8-13.

[本文引用: 1]

ZOU

J Q

, WANG

Y Q

, ZHANG

Z P

, ZHU

. Research on biomass, brix and juice extraction of A₃-type cytoplasmic sweet sorghum for energy use
Journal of China Agricultural University, 2011,16(2):8-13. (in Chinese)

[本文引用: 1]

[30]

高士杰, 刘晓辉, 李玉发, 李继洪. 中国甜高粱资源与利用
杂粮作物, 2006,26(4):273-274.

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GAO

S J

, LIU

X H

, LI

Y F

, LI

J H

. Sweet sorghum resources and its utilization in China
Rain Fed Crops, 2006,26(4):273-274. (in Chinese)

[本文引用: 1]

[31]

ZHANG

C X

, XIE

G D

, LI

S M

, GE

L Q

, HE

T T

. The productive potentials of sweet sorghum ethanol in China
Applied Energy, 2010,87:2360-2368.

DOI:10.1016/j.apenergy.2009.12.017 URL [本文引用: 1]

Abstract

As one of the important non-grain energy crops, sweet sorghum has attracted the attention of scientific community and decision makers of the world since decades. But insufficient study has been done about the spatial suitability distribution and ethanol potential of sweet sorghum in China. This paper attempts to probe into the spatial distribution and ethanol potential of sweet sorghum in China by ArcGIS methods. Data used for the analysis include the spatial data of climate, soil, topography and land use, and literatures relevant for sweet sorghum studies. The results show that although sweet sorghum can be planted in the majority of lands in China, the suitable unused lands for large-scale planting (unit area not less than 100 hm²) are only as much as 78.6 × 10⁴ hm²; and the productive potentials of ethanol from these lands are 157.1 × 10⁴–294.6 × 10⁴ t/year, which can only meet 24.8–46.4% of current demand for E10 (gasoline mixed with 10% ethanol) in China (assumption of the energy efficiency of E10 is equivalent to that of pure petroleum). If all the common grain sorghum at present were replaced by sweet sorghum, the average ethanol yield of 244.0 × 10⁴ t/year can be added, and thus the productive potentials of sweet sorghum ethanol can satisfy 63.2–84.9% of current demand for E10 of China. In general, Heilongjiang, Jilin, Inner Mongolia and Liaoning rank the highest in productive potentials of sweet sorghum ethanol, followed by Hebei, Shanxi, Sichuan, and some other provinces. It is suggested that these regions should be regarded as the priority development zones for sweet sorghum ethanol in China.

[32]

SHUKLA

, FELDERHOFF

T J

, SABALLOS

, VERMERRIS

. The relationship between plant height and sugar accumulation in the stems of sweet sorghum (Sorghum bicolor(L.) Moench)
Field Crops Research, 2017,203:181-191.

[本文引用: 1]

[33]

KNOLL

J E

, ANDERSON

W F

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K R

, NI

X Z

. The environment strongly affects estimates of heterosis in hybrid sweet sorghum
Sugar Technology, 2018,20(3):261-274.

DOI:10.1007/s12355-018-0596-0 URL [本文引用: 1]

[34]

JAIKISHAN

, RAJENDRAKUMAR

, HARIPRASANNA

, BHAT

B V

. Gene expression analysis in sorghum hybrids and their parental lines at critical developmental stages in relation to grain yield heterosis by exploiting heterosis-related genes from major cereals
Plant Molecular Biology Reporter, 2018,36:418-428.

DOI:10.1007/s11105-018-1079-x URL [本文引用: 2]

[35]

NDHLELA

, HERSELMAN

, SEMAGN

, MAGOROKOSHO

, MUTIMAAMBA

, LABUSCHAGNE

M T

. Relationships between heterosis, genetic distances and specific combining ability among CIMMYT and Zimbabwe developed maize inbred lines under stress and optimal conditions
Euphytica, 2015,204:635-647.

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[36]

PAVANI

, SUNDARAM

R M

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M S

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P B K

, KEMPARAJU

K B

. Prediction of heterosis in rice based on divergence of morphological and molecular markers
Journal of Genetics, 2018,97(5):1263-1279.

URLPMID:30555075 [本文引用: 1]

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王林友, 张礼霞, 勾晓霞, 范宏环, 金庆生, 王建军. 利用InDel标记鉴定浙优系列杂交稻籼粳属性和预测杂种优势
中国农业科学, 2014,47(7):1243-1255.

[本文引用: 1]

WANG

L Y

, ZHANG

L X

, GOU

X X

, FAN

H H

, JIN

Q S

, WANG

J J

. Identification of Indica-Japonica attribute and prediction of heterosis of Zheyou hybrids rice using InDel molecular markers
Scientia Agricultura Sinica, 2014,47(7):1243-1255. (in Chinese)

[本文引用: 1]

[38]

JORDAN

D R

, TAO

, GODWIN

I D

, HENZELL

R G

, COOPER

, MCINTYRE

C L

. Prediction of hybrid performance in grain sorghum using RFLP markers
Theoretical and Applied Genetics, 2003,106:559-567.

DOI:10.1007/s00122-002-1144-5 URLPMID:12589557 [本文引用: 2]

Heterosis is an important component of hybrid yield performance. Identifying high yielding hybrids is expensive and involves testing large numbers of hybrid combinations in multi-environment trials. Molecular marker diversity has been proposed as a more efficient method of selecting superior combinations. The aim of this study was to investigate the value of molecular marker-based distance information to identify high yielding grain sorghum hybrids in Australia. Data from 48 trials were used to produce hybrid performance-estimates for four traits (yield, height, maturity and stay green) for 162 hybrid combinations derived from 70 inbred parent lines. Each line was screened with 113 mapped RFLP markers. The Rogers distances between the parents of each hybrid were calculated from the marker information on a genome basis and individually for each of the ten linkage groups of sorghum. Some of the inbred parents were related so the hybrids were classified into 75 groups with each group containing individual hybrids that showed similar patterns of Rogers distances across linkage groups. Correlations between hybrid-group performance and hybrid-group Rogers distances were calculated. A significant correlation was observed between whole genome-based Rogers distance and yield ( r = 0.42). This association is too weak to be of value for identifying superior hybrid combinations. One reason for the generally poor association between parental genetic diversity and yield may be that important QTLs influencing heterosis are located in particular chromosome regions and not distributed evenly over the genome. Variation in the sign and magnitude of correlations between Rogers distance and hybrid-group performance for particular linkage groups observed in this study support this hypothesis. The concept of using diversity on individual linkage groups to predict performance was explored. Using data from just two linkage groups 38% of the variation in hybrid performance for grain yield could be explained. A model combining phenotypic trait data and parental diversity on particular linkage groups explained 71% of the variation in grain yield and has potential for use in the selection of heterotic hybrids.

[39]

MELCHINGER

A E

, LEE

, LAMKEY

K R

, WOODMAN

W L

. Genetic diversity for restriction fragment length polymorphisms: Relation to estimated genetic effects in maize inbreds
Crop Science, 1990,30:1033-1040.

DOI:10.2135/cropsci1990.0011183X003000050016x URL [本文引用: 2]

[40]

CHARCOSSET

, LEFORT-BUSEN

, GALLAIS

. Relationship between heterosis and heterozygosity at marker loci: A theoretical computation
Theoretical and Applied Genetics, 1991,81:571-575.

DOI:10.1007/BF00226720 URLPMID:24221369 [本文引用: 2]

In this paper we have studied the linear correlation between a genetic distance index between two parent lines (based on marker loci information) and the heterosis observed in the F1 hybrid from the two lines, for a quantitative character (determined by several loci, or QTL). Theoretical computations of the correlation coefficient (varrho) between the distance index and the heterosis were made, assuming the biallelic model (defined by Fisher). When the alleles at both marker loci and QTL are equally distributed among the whole population of considered lines, the coefficient varrho is a function of the squares of linkage disequilibria between alleles at marker loci and alleles at QTL. The QTL that are not marked by marker loci and marker loci that do not mark any QTL play symmetrical roles and can decrease varrho greatly. We conclude that the prediction of F1 hybrid heterosis based on marker loci would be more efficient if these markers were selected for their relationship to the alleles implicated in the heterotic traits considered.