Combining Ability Analysis for Yield and Yield Components of Yinshui-type Japonica Hybrid Rice

Cheng ZHANG，Yajun CHEN，Xianju WANG，Xiu PANG，Wei JlANG，Guojun SHAOLiaoning Rice Research Institute，Shenyang 110101，China

Combining Ability Analysis for Yield and Yield Components of Yinshui-type Japonica Hybrid Rice

Cheng ZHANG，Yajun CHEN，Xianju WANG，Xiu PANG，Wei JlANG，Guojun SHAO*
Liaoning Rice Research Institute，Shenyang 110101，China

［Objective］This study aimed to analyze the yield potential of newly developed Yinshui-type japonica hybrid rice parents and provide the theoretical basis for breeding of Yinshui-type japonica hybrid rice.［Method］An incomplete diallel cross of four cytoplasmic male sterile（CMS）lines and four restorer lines was designed to investigate the combining ability for yield and yield components of Yinshui-type japonica hybrid rice. ［Result］The variances of general combining ability （GCA）for yield traits were extremely significantly different.GCA variances of male parents were higher than that of female parents except for seed-setting rate.The variances of specific combining ability （SCA）for grain yield per plant，number of effective spikes and seed-setting rate were significantly or extremely significantly different.139A and Wanlun 422 exhibited higher GCA effects.Most yield traits of these two parents belonged to typeⅡ.Combinations 139A×Wanlun 422 and Liaojing10A×Wanlun 422 had relatively high grain yield per plant and SCA effects.GCA effects of restorer lines and SCA effects of combinations greatly affected yield traits，while GCA effects of CMS lines slightly affected yield traits. ［Conclusion］In this study，restorer lines exhibited greater genetic differences and impact on yield traits than CMS lines；139A and Wanlun 422 possessed higher application potential.

Yinshui-type japonica hybrid rice；Yield and yield components；Combining ability

Materials and Methods

Materials

Four cytoplasmic male sterile（CMS）lines and four restorer lines of Yinshui-type japonica hybrid rice were used as experimental materials.CMS lines:Liaojing 6A，Liaojing 10A，65A and 139A；restorer lines:R382，Wanlun 422，C2106 and Yejinghui（introduced by Guangde Institute of Agricultural Sciences）.

Methods

Hybrid rice seed production was carried out in the experimental field of Liaoning Rice Research Institute and Hainan experimental base in August 2011 and February 2012，respectively. Using four CMS lines as female parents and four restorer lines as male parents，cross combinations were set according to NC II genetic mating design.In April 2012，F(xiàn)1hybrids were planted in the experimental field of Liaoning Rice Research Institute. Randomized block design was employed with three replications.There were three rows in each plot and each row was 2 m in length；the spacing in the rows and spacing between rows were 30 cm×13 cm.Single-plant cultivation pattern was adopted and other management measures were in accordance with field practice.

After maturity，five representative plants in the middle row of each plot were selected for indoor test.The combining ability for yield and yield components was analyzed according to NC Ⅱ genetic mating design method by using SPSS and Excel software［19-20］.

Table 1 Variances analysis for yield and yield components of 16 combinations

Table 2 GCA effects on yield and yield components of parents

Results and Analysis

Variance Analysis of combining ability

According to variance analysis results of yield and yield components of 16 combinations（Table 1），yield and yield components varied extremely significantly among various combinations.To be specific，among six traits，GCA variances of female parents for grain number per spike，seed-setting rate，number of filled grains per spike and 1 000-grain weight were significantly or extremely significantly different；GCA variances of female parents for grain yield per plant and number of effective spikes were significantly different at P＜0.08；GCA variances of male parents for six traits were extremely significantly different；SCAvariances of combinations for grain yield per plant，number of effective spikes and seed-setting rate were significantly or extremely significantly different；among various yield traits，GCA variances of male parents were higher compared with female parents except for seed-setting rate，which indicated that the parents involved in this study exhibited significant genetic differences in yield traits；the combining ability of traits with significant differences in GCA variances and SCA variances could be further analyzed.

Analysis of combining ability effects

GCA analysis results of eight parents were shown in Table 2.139A exhibited high GCA effects on grain yield per plant，grain number per spike，seed-setting rate and number of filled grains per spike，which was identified as a good female parent of japonica hybrid rice；Wanlun 422 exhibited high GCA effects on grain yield per plant，number of effective spikes，grain number per spike，seed-setting rate and number of filled grains per spike，which was identified as a good male parent of japonica hybrid rice.In addition， Liaojing 6A exhibited high GCA effects on seed-setting rate； 65A and Yejinghui exhibited high GCA effects on 1 000-grain weight；R382 and C2106 had high GCA effects on grain number per spike and number of filled grains per spike，which indicated that these parents had important application significance.

According to SCA analysis results（Table 3），combinations with the highest SCA effects did not have the highest yield.To be specific，combinations 139A×Wanlun 422 and Liaojing10A× Wanlun 422exhibited the highest grain yield per plant of 87.56 g and 79.67 g，respectively；SCA effects of these two combinations on grain yield per plant were 4.08 and 5.24，respectively（differences reached the 0.2 significant level compared with Liaojing 10A×R382 and 65A×C2106），indicating that 139A×Wanlun 422 and Liaojing10A ×Wanlun 422were good japonica hybrid rice combinations. High-yield combinations exhibited relatively high SCA effects on grain yield per plant and number of effective spikes，with the variation ranges of 4.08-9.89 and 0.87-2.00，respectively.Moreover，SCA effects of a combination exhibited a correlation with GCA effects of its male parent.For instance，male parents were classified into high-type （a），moderate-type（b）and low-type（c）according to GCA effects on grain yield per plant.Male parents of five combinations with the highest SCA effects were mostly hightype and moderate-type；male parents of five combinations with the lowest SCA effects were mostly moderatetype and low-type.

Table 3 SCA effects on yield and yield components of ten combinations with the maximum and minimum yield

Table 4 Classification of parents based on utilization value of yield and yield components

Analysis of utilization value of parents

According to GCA effects and SCA variances，eight parents were classified based on their utilization value in breeding according to Mo［20］（Table 4）.On the basis of grain yield per plant，139A and Wanlun 422 were good parents belonging to typeⅡ，while R382 belonged to typeⅢ.Thus，SCA of these parents could be used to strengthen test-crossing.139A was conducive to improving number of effective spikes，grain number per spike，seed-setting rate and number of filled grains per spike of hybrid combinations，while Wanlun 422 was con-ducive to improving number of effective spikes，grain number per spike，seed-setting rate，number of filled grains per spike and1000-grain weight of hybrid combinations，which indicated that these two parents were good Yinshui-type japonica hybrid rice parents.

The utilization value of ten hybrid combinations with the highest and lowest grain yield per plant was analyzed （Table 5）.According to the results，male parents of five combinations with the highest yield mostly belonged to typeⅡ and typeⅢ；male parents of five combinations with the lowest yield mostly belonged to typeⅢand typeⅣ.GCA played a dominant role in the classification of utilization value of parents.Therefore，high-GCA male parents were very important to breeding of Yinshui-type japonica hybrid rice.Grain yield per plant was influenced by GCA of parents and SCA of combinations.Correlation analysis showed that the correlation coefficient between GCA of male parents and grain yield per plant was 0.709，which reached an extremely significant level；the correlation coefficient between SCA of combinations and grain yield per plant was 0.603，which reached a significant level；the correlation coefficient between GCA of female parents and grain yield per plant was 0.366，which reached 1% significant level.Therefore，two combinations with the highest grain yield per plant were attributed to the combined action of GCA of male parents and SCA of combinations.Hybrid combinations with significantly improved yield can be obtained when GCA of male parents and SCA of combinations were both high.GCA of female parents exerted a certain impact on yield when GCA of male parents and SCA of combinations exhibited no significant differences.

Conclusion and Discussion

In this study，SCA variances of combinations for grain number per spike，number of filled grains per spike and 1 000-grain weight did not reach the 5%significant level，suggesting that these traits were controlled by additive effects；GCA variances and SCA variances of combinations for grain yield per plant，number of effective spikes and seed-setting rate reached the 5%or 1%significant levels，indicating that these traits were controlled by additive effects and non-additive effects.Except for seed-setting rate，GCA variances of male parents for yield traits were higher compared with female parents，which indicated that genetic differences of male parents were greater than female parents. Male parents of five combinations with the highest SCA effects mostly belonged to high-type and moderatetype；SCA effects of combinations and GCA effects of male parents exhibited a certain correlation.GCA effects of 139A and Wanlun 422 on grain yield per plant were 6.30 and 11.97，which ranked first among CMS lines and restorer lines，respectively；furthermore，the utilization value of yield and yield components of 139A and Wanlun 422 mainly belonged to typeⅡ，indicating that these two lines were good Yinshui-type japonica hybrid rice parents. Grain yield per plant of 139A×Wanlun 422 and Liaojing 10A×Wanlun 422 was relatively high（87.56 and 79.67 g）among various combinations，with SCA of 4.08 and 5.24，indicating that they were good Yinshui-type japonica hybrid rice combinations.The correlation coefficients between GCA of male parents，SCA of combinations，GCA of female parents and grain yield per plant were 0.709，0.603 and 0.366，respectively.Therefore，GCA effects of male parents and SCA effects of combinations greatly affected yield traits；GCA effects of female parents slightly affected yield traits.

Table 5 Comparative analysis of combination yield and parental type

Yinshui-type japonica hybrid rice parents were used as experimental materials in this study.According to the results，grain number per spike，number of filled grains per spike，and 1 000-grain weight were mainly controlled by additive effects，while grain yield per plant，number of effective spikes and seed-setting rate were controlled by additive effects and nonadditive effects，which were inconsistent with previous conclusions［21-22］. This might be due to the different genetic backgrounds of experimental materials.In this study，female parents were bred by conventional japonica rice in Liaoning Province，exhibiting relatively narrow genetic backgrounds；male parents were bred by indicajaponica hybrid breeding，exhibiting extensive genetic backgrounds.Specific affinity genes can overcome infertility of indica-japonica hybrids［23］.Male parents used in this study had certain genetic relationship with indica rice，and differences in the specific affinity among female parents might result inhigher GCA variances of female parents for seed-setting rate than male parents.Results indicate that SCA effects of combinations exhibit a certain correlation with GCA effects of their male parents，which is inconsistent with previous conclusions［24］and this may be related with the slight genetic differences among female parents.

Among the existing parental materials，male parents have abundant genetic basis，while female parents have relatively narrow genetic basis. Therefore，test-crossing and screening of existing restorer lines should be strengthened to construct high SCA combinations and fully utilize germplasm resources；new germplasm resources should be introduced into maintainer lines to improve the genetic diversity of female parents and breed high GCA female parents.139A and Wanlun 422 are excellent Yinshui-type japonica hybrid rice parents，which can be used as major parents to improve maintainer lines and restorer lines on the basis of test-crossing，thus contributing to breeding of Yinshui-type japonica hybrid rice.In the breeding process of new combinations of Yinshui-type japonica hybrid rice，parents with high GCA effects on yield traits should be selected for extensive testcrossing and screening to breed specific combinations with high SCA effects on number of effective spikes，seed-setting rate and grain yield per plant.

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印水型雜交粳稻產(chǎn)量及產(chǎn)量構(gòu)成因素的配合力分析

張城，陳亞君，王先俱，龐秀，姜偉，邵國(guó)軍*
（遼寧省水稻研究所，遼寧沈陽 110101）

［目的］旨在為確定新育成印水型雜交粳稻親本的產(chǎn)量潛力，并為印水型雜交粳稻育種提供理論基礎(chǔ)。［方法］本研究以4個(gè)不育系和4個(gè)恢復(fù)系為親本進(jìn)行不完全雙列雜交，研究產(chǎn)量及其構(gòu)成因素的配合力。［結(jié)果］產(chǎn)量性狀一般配合力（GCA）方差均達(dá)極顯著水平，除結(jié)實(shí)率外，父本的一般配合力方差大于母本，單株產(chǎn)量、有效穗和結(jié)實(shí)率的特殊配合力（SCA）方差達(dá)顯著或極顯著水平。139A和晚輪422的GCA效應(yīng)較高，產(chǎn)量性狀以Ⅱ類為主，139A×晚輪422和遼粳10A×晚輪422的單株產(chǎn)量水平及組合SCA效應(yīng)較高。父本GCA效應(yīng)和組合SCA效應(yīng)對(duì)產(chǎn)量性狀的影響較大，母本GCA效應(yīng)對(duì)產(chǎn)量性狀的影響相對(duì)較小。［結(jié)論］本研究中，父本的遺傳差異大于母本，對(duì)產(chǎn)量性狀的影響也大于母本，139A和晚輪422有較大應(yīng)用潛力。

印水型雜交粳稻；產(chǎn)量及產(chǎn)量構(gòu)成因素；配合力

W ith the development of economy，the demand for highquality japonica rice is increasing continuously［1］.According to the historical evolution of japonica rice and indica rice in various countries，the trend"indica rice to japonica rice" has been observed in Taiwan and Jiangsu of China，South Korea and other regions［2］，indicating that japonica hybrid rice is developing towards the south.Yinshui-type sterile lines are widely used core parents of Chinese super hybrid rice［3］.Currently，Yinshuitype sterile cytoplasm has been introduced into japonica hybrid rice to overcome the shortcomings of BT sterile cytoplasm，breed Yinshui-type japonica sterile lines and increase genetic diversity of the cytoplasm，which is of great significance for overcoming the genetic vulnerability due to simple cytoplasm［4-6］.In addition，existing germplasm resources and segregation populations were screened with molecular marker-assisted selection method and field test and several restorer lines for Yinshui-type japonica CMS lines were obtained［7-8］，which constructed preliminarily three-ttline combinations of Yinshui-type japonica hybrid rice.

Combining ability plays an important role in parent breeding and combination selection of hybrid rice.Constructing high-SCA combinations on the basis of high-GCA parents is necessary for the improvement of hybrid rice breeding［9］；investigating the advantages and disadvantages of combi-nations is an important link in hybrid rice breeding［10］.A large number of studies have been conducted on combining ability of indica rice.At present，combining ability has become an important indicator for breeding highquality parents［11-13］.The combining ability of BT-type and Dian-type japonica hybrid rice in Jiangsu［14-16］and Yunnan［17-18］has been analyzed，and a number of major parents were selected，including Liuyan A，Dianjingyou 2，C418 and Nan 34.Currently，little information is available on the combining ability of northern japonica hybrid rice；no studies have been reported on the combining ability of Yinshui-type japonica hybrid rice，resulting in lack of in-depth understanding of genetic characteristics of northern Yinshuitype japonica hybrid rice parents.In this study，the combining ability for yield and yield components of northern Yinshui-type japonica hybrid rice was investigated to analyze the genetic characteristics of yield traits in parents，determine the combining ability of CMS lines and restorer lines and obtain parents and hybrid combinations with application potential，which provided a theoretical basis for breeding of northern Yinshui-type japonica hybrid rice.

遼寧省科技廳“粳稻新型不育胞質(zhì)在北方雜交粳稻育種中的研究與應(yīng)用”（20102105）。

張城（1979-），男，遼寧丹東人，助理研究員，碩士，主要從事粳稻常規(guī)品種和保持系選育，E-mail：zhch024@163.com。*通訊作者，陳亞君（1962-），女，遼寧蓋州人，副研究員，學(xué)士，主要從事北方雜交粳稻新組合選育，E-mail：lnsygjsh07@163.com。

2015-01-20

2015-02-15

Supported by Project of Liaoning Provincial Department of Science and Technology "Study and Application of A Novel Sterile Cytoplasm of Japonica Rice in Japonica Hybrid Rice Breeding in the North"（20102105）.

.E-mail:lnsygjsh07@163.com

January 20，2015Accepted:February 15，2015