Guoqiang GUO,Jianying YIN,Wuqiang GUO,Lianggui ZENG,Yiguo HUANG,Jing GAO, Xiaohu YANG,Mingqi GUO

1.Hengyang Institute of Agricultural Science,Hengyang 421101,China;

2.College of Agronomy,Hunan Agricultural University,Changsha 410125,China;

3.China National Hybrid Rice R&D Center,Changsha 410125,China;

4.Hunan Longping Seed Industry Co.,Ltd.,Changsha 410125,China

Inheritance of Aroma of Good-quality Indica Type Rice PTGMS Line GHS and Its Application in Hybrid Rice Breeding

Guoqiang GUO1,2,Jianying YIN1,Wuqiang GUO3,4,Lianggui ZENG1,Yiguo HUANG1,Jing GAO1, Xiaohu YANG1,Mingqi GUO3*

1.Hengyang Institute of Agricultural Science,Hengyang 421101,China;

2.College of Agronomy,Hunan Agricultural University,Changsha 410125,China;

3.China National Hybrid Rice R&D Center,Changsha 410125,China;

4.Hunan Longping Seed Industry Co.,Ltd.,Changsha 410125,China

The inheritance of aroma of GHS was studied by crossing GHS with nonaromatic lines 9311,Xianhui 207,Yuefengzhan,Taihuangzhan,Yuexiangzhan and aromatic lines Yuxiangyouzhan,Xiangwanxian 17,Tianlongxiang 103,Xinwanxiang and Chuangxiang 5.For the grains of F1generation,the performance of aroma was tested by chewing method,while for the plants of generations of F1,F2and BC1F1, the performance of aroma was tested by KOH solution method.The results showed that the non-aroma of restorer line is a dominant trait,but the aroma of GHS line is a recessive trait;the aroma genes of GHS,Yuexiangzhan and Xiangwanxian 17 were non-allelic;the aroma genes of GHS,Yuxiangyouzhan,Tianlongxiang 103,Xinwanxiang and Chuangxiang 5 may have the same allelic loci.Among the combinations of GHS and non-aromatic restorer lines,the non-aromatic to aromatic ratios of the F2plants and the BC1F1plants were 3∶1 and 1∶1,respectively,and the inheritance of aroma of GHS was controlled by a pair of recessive nuclear genes.In hybrid rice breeding,using GHS,Qiongxiang S has been bred out,and a series of aromatic hybrid rice combinations have also been selected.Among the hybrid combinations,Qiongxiangliangyou 1,Qiongxiangliangyou 2 and other high-yielding and high-quality combinations have been approved by provincial government,and they have good application prospects.

Rice;Aroma;Inheritance;Breeding

A romatic rice is popular in consumers and world rice market for its fragrance and delicious taste.The aroma of rice has been treated internationally as an important indicator for evaluating quality.In the international market,the brand-name high-quality rice is generally mixed with aromatic rice,so the price of aromatic rice is 2-3 times higher than that of ordinary rice.The Thai rice has become synonymous with high-grade and high-quality rice.Its international trade always ranks first among the world’s rice trade[1].With the continuous improvement of people’s living standard and adjustment of cropplanting structure,development opportunity and motivation have been provided for the breeding and largescale production of aromatic rice. Therefore,conducting studies on inheritance of aroma and breeding of aromatic rice,by fully exploiting and utilizing the precious resources of aromatic rice,plays an important role in improving China’s agricultural planting structure,increasing farmers’income, promoting rice industrialization,developing brand-name rice and increasing the competitiveness of China’s rice in the international market.

GHS is a PTGMS line with aroma of jasmine that was found by Guo in Myanmar in November of 1997 when he collected upland rice resources.Its sterility gene is not allelic with those of Nongken 58S and Annong S-1.So GHS is a new photoperiod-and thermo-sensitive genic male sterile material[2].Under the long-day(＞13.5 h)condition,the sterility of GHS will not be changed although it encountered with abnormal low temperature(daily average temperature of 23℃for 3 consecutive days).GHS is typical photoperiod-thermo interactive and dualusing genic sterile line that has lower sterile threshold temperature and shorter sterile threshold day length[3]. The growth period of GHS lasts for 95 d in Changsha.It is sowed on May 20th,and its heading is started on September 1st;while in Hainan,the growth period of GHS lasts for 75-80 d.GHS has plant height of about 90 cm,moderate plant type,strong tillering ability,stiff stems and developed roots.The leaves and sheathes of GHS are all gray-green.There are about 14.5-16.5 leaves on the main stem with photosensitivity.The apiculus are white,the panicles,in length of 23.0 cm,are large,and the longshaped grains are more.There are 140 grains in average in each panicle. The kilo-grain weight of GHS is 24.0 g. The grain quality of GHS line has been analyzed by the Rice c sterile line and Products Quality Supervision and Testing Center of the Ministry of Agriculture.The analysis results show that the GHS line has grain length of 6.6 mm,length to width ratio of grains of 3.2,chalky rice rate of 2%,chalkiness of 0.2%,alkali spreading value of 7.0, gel consistency of 78 mm,protein content of 16.1%,milled rice rate of 71.0%,amylose content of 9.6%and transparency of 2 grade.Both the appearance qualities and tastes of GHS/ Haixiang 1 and GHS/296 hybrid rice are higher than those of their male parents[4].In this study,the inheritance law of aroma trait of GHS and its application in aromatic hybrid rice breeding were discussed,enriching the breeding theory and practice of aromatic hybrid rice.

Inheritance of Aroma Trait of GHS

Materials and Methods

MaterialsThe tested materials included aromatic sterile line GHS,nonaromatic restorer lines Xianhui 207, 9311,Yuefengzhan and Taihuangzhan,and aromatic rice varieties Yuexiangzhan,Xiangwanxian 17, Yuxiangyouzhan,Tianlongxiang 103, Xinwanxiang and Chuangxiang 5.The Xianhui 207 and 9311 were introduced from the Hunan Hybrid Rice Research Center;the Xiangwanxian 17,Tianlongxiang 103 and Chuangxiang 5 were introduced from the Hunan Rice Research Institute;the Yuefengzhan, Taihuangzhan,Yuexiangzhan,Yuxiangyouzhan and Xinwanxiang were introduced from the Rice Research Institute of Guangdong Province.

Combination methodsThe GHS was used as female parent,and it was crossed with the non-aromatic restorer lines Xianhui 207,9311,Yuefengzhan and Taihuangzhan and the aromatic rice varieties Yuexiangzhan,Xiangwanxian 17,Yuxiangyouzhan,Tianlongxiang 103,Xinwanxiang and Chuangxiang 5.Their F1hybrid rice seeds were obtained.Through the self-crossing of F1hybrid rice seeds, the F2rice seeds were obtained.The F1rice seeds were back-crossed with the GHS,and then the BC1F1rice seeds were prepared.At final,the F1, F2and BC1F1rice seeds were cultivated to obtain their corresponding plants.

Identification method of aroma

The presence or absence of aroma and its degree in F1brown rice were identified by chewing method grain by grain.The aroma in plant leaves was identified using the KOH solution method[5].At the heading stage,certain amount(about 2 g)of leaves was sampled from each of the samples.The fresh leaves were cut into pieces indoors and then placed into tubes.Certain amount(10 ml)of 1.7%KOH solution was added to each of the tubes. After covering the lids,the leaves and KOH solution were fully mixed.After stood for 20 min at room temperature, the samples were identified.Due to the differences in people’s sense of smell, the samples were all identified by three staffs respectively.If the identification result of a certain sample is controversial,it will be identified again.For the two identification method,the combination of corresponding restorer line and GHS was treated as the control. The aromatic to non-aromatic ratios of F1,F2and BC1F1plants were calculated,and the values were tested by χ2test.

Results and analysis

Appearance and allelism of aroma trait of rice grains of parents and F1generationsThe pollens of non-aromatic restorer lines Xianhui 207,9311, Yuefengzhan and Taihuangzhan and aromatic rice varieties Yuexiangzhan, Xiangwanxian 17,Yuxiangyouzhan, Tianlongxiang 103,Xinwanxiang and Chuangxiang 5 were pollinated to the aromatic sterile line GHS,respectively. The F1hybrid rice seeds were collected from the GHS 25 d later.After the shelling,brown rice was obtained.The presence or absence of aroma in brown rice was identified grain by grain.The results showed that the F1brown rice of GHS/Xianhui 207,GHS/ 9311,GHS/Yuefengzhan and GHS/Taihuangzhan all had no aroma.The F1brown rice of GHS/Yuexiangzhan and GHS/Xiangwanxian also had no aroma.The F1brown rice of GHS/ Yuxiangyouzhan,GHS/Tianlongxiang 103,GHS/Xinwanxiang and GHS/ Chuangxiang 5 had aroma,but the degree of aroma is different among different combinations.The F1brown rice of GHS/Yuxiangyouzhan has only slight aromatic smell,the F1brown rice of GHS/Tianlongxiang 103 had light aromatic smell,but the F1brown rice of GHS/Xinwanxiang and GHS/Chuangxiang 5 had thick aromatic smell (Table 1).All the results above suggested that the non-aroma of restorer lines is a dominant trait,but the aroma of GHS is a recessive trait.The aroma genes of GHS,Yuexiangzhan and Xiangwanxian 17 are not allelic.But the aroma genes of GHS,Yuxiangyouzhan,Tianlongxiang 103,Xinwanxiang and Chuangxiang 5 may have the alleles.

Table 1Performance of aroma trait in grains of F1generation and parents

Appearance of aroma trait in leaves of F1plantsAt the tillering stage,the aroma in F1plants of the hybrid combinations(GHS/Xianhui 207,GHS/9311, GHS/Yuefengzhan,GHS/Taihuangzhan,GHS/Yuexiangzhan,GHS/Xiangwanxian 17,GHS/Yuxiangyouzhan, GHS/Tianlongxiang 103,GHS/Xinwanxiang and GHS/Chuang-xiang 5) was identified.The results showed that there was no aromatic smell in the leaves of the F1plants of GHS/9311, GHS/Xianhui 207,GHS/Yuefengzhan or GHS/Taihuangzhan,and also no aromatic smell in the leaves of the F1plants of GHS/Yuexiangzhan or GHS/ Xiangwanxian 17.But the leaves of the F1plants of GHS/Yuxiangyouzhan, GHS/Tianlongxiang 103,GHS/Xinwanxiang and GHS/Chuangxiang 5 had aromatic smell.Therefore,it can be concluded that the aroma trait of GHS is controlled by recessive gene.Segregation of aroma trait in leaves among F2plantsThe appearance of aroma trait in the F2plants of the combinations was further analyzed(Table 2).There was no aroma in the F1plants of GHS/Yuexiangzhan or GHS/ Xiangwanxian 17,but aromatic plants were obtained from their F2plants.It indicated that the aroma gene of GHS was different from those of Yuexiangzhan and Xiangwanxian 17.The F1plants of GHS/Yuxiangyouzhan,GHS/ Tianlongxiang 103,GHS/Xinwanxiang and GHS/Chuangxiang 5 were all aromatic,and the aroma trait was not segregated among their F2plants,so their F2plants were also all aromatic.It suggested that the GHS,Yuxiangyouzhan,Tianlongxiang 103,Xinwanxiang and Chuangxiang 5 had the same aroma gene.The χ2test of the aromatic plant to non-aromatic plant ratios among the F2populations of GHS/Xianhui 207,GHS/9311,GHS/Yuefengzhan and GHS/Taihuangzhan showed that aromatic plant to nonaromatic plant ratio met the segregation ratio of 1:3,indicating that the aroma trait of GHS is controlled by one pair of recessive genes.

Segregation of aroma trait in leaves among BC1F1plantsAs shown in Table 3,among the four BC1F1populations,there were all aromatic and non-aromatic plants.The aromatic plant to non-aromatic plant ratios among the BC1F1plants of GHS//GHS/ 9311,GHS//GHS/Xianhui 207,GHS// GHS/Yuefengzhan and GHS//GHS/ Taihuangzhan were 1∶1.24,1∶0.89, 1∶0.80 and 1∶0.84,respectively.The χ2test showed that the aromatic plant to non-aromatic plant ratios among the four BC1F1populations was all 1∶1(χ2values ranged from 0.08 to 0.34,which were much lower than the critical value of χ20.05),meeting the segregation pattern of one pair of genes.So it was further proved that the aroma trait of GHS is controlled by one pair of recessive nuclear genes.

Table 2Segregation performance of aroma trait in leaves of F2plants of all the combinations

Table 3Segregation performance of aroma trait in leaves of BC1F1plants of the combinations between GHS and non-aromatic male parents

Discussion

Allelism of aroma geneThis study showed that the aroma gene of GHS is not allelic with those of Yuexiangzhan and Xiangwanxian 17.Therefore, there are at least two kinds of aroma gene in rice.It is generally considered that the main component of the aroma of aromatic rice is 2-acetyl-1-pyrroline (AP)[6].But some scholars believe that the main component of the aroma of aromatic rice is A-pyrrolidinone[7],and some others consider the main components are propionaldehyde,valeraldehyde and acetaldehyde[8].The difference in chemical composition may also be one of the causes of non-allelic aroma gene.Pinson speculates that there may be a kind of denaturase that promotes the biosynthesis of 2-acetyl-1-pyrroline in aromatic rice,and the multiple alleles of the single aroma genes can slightly change the denaturase,resulting in the generation of rice varieties with different-degree aromas.The changed metabolic pathways and the other additional steps are caused by compound aroma genes.

Inheritance pattern of aroma trait

The inheritance pattern of aroma trait of aromatic rice is till controversial. Most scholars have reported that the aroma of aromatic rice is a single recessive inheritance[9,15].Zhou et al.[16]and Ren et al.[17]consider that the aroma is a trait controlled by two recessive genes.There are also studies reporting that the aroma is a trait controlled by one pair of aroma genes and one pair of suppressor genes[18],and it is affected by the complementary effects among the three dominant genes [1 9].The differences above may be caused by the differences in tested aromatic rice materials.In different aromatic rice materials,the origins and components of aroma are different, and the genes that control the aroma trait and their action patterns are also different.In addition,the difference in concluded inheritance pattern of aroma trait in aromatic rice may also be caused by different aroma identification methods.There is aroma in F1brown rice of GHS/Yuxiangyouzhan, GHS/Tianlongxiang 103,GHS/Xinwanxiang and GHS/Chuangxiang 5, but the degree of aroma differs among the four populations,which may be due to the presence of multiple aroma alleles or multiple minor genes.

Breeding of aromatic riceThe aroma of rice is restricted by the genotype of endosperm(3n).The sterile line provides the endosperm of hybrid rice with 2n(egg nucleus and polar nuclei), and the restorer line only provides with 1n(sperm nucleus).In terms of breeding naturally-combined aromatic hybrid rice,the less the number of genes that control the aroma of sterile line is,the larger the percentage of aromatic plants in F2generation is[20].This study preliminarily indicates that the aroma trait of GHS is controlled by one pair of recessive major genes,and its hereditary behavior is simple,and it is easy to be transferred and stabilized in breeding.So GHS is an ideal parent material for breeding aromatic hybrid rice.In hybrid rice breeding,if GHS is used as the aroma donor and it is crossed with a parent with excellent comprehensive traits,new excellent two-line sterile lines can be bred out from the offspring.If GHS is crossed with a non-aromatic restorer line,there are about 1/4 of the F2hybrid rice grains with aromatic flavor.If GHS is crossed with a restorer line with different allelic aroma gene(controlled by a single recessive gene),one half of the F2hybrid rice grains will have aromatic flavor.So the F2generation is also semi-aroma typed.If GHS is crossed with a restorer line with same allelic aroma gene,the F2hybrid rice grains will all have aromatic flavor.So the F2generation is whole-aroma typed.

Breeding of Aromatic Hybrid Rice

Breeding of photoperiod-and thermo-sensitive genic male sterile line Qiongxiang S

The sterile line was bred systematically from the GHS system by Guo using seven years.It passed the science and technology achievement testimonial of Hainan Province in 2004[21].The successful breeding of GHS makes it possible to bred two-line high-y-qng and high-quality aromatic hybrid rice.Utilizing the heterosis between upland rice(Qiongxiang S-1) and general rice,several high-quality hybrid rice combinations have been bred and approved by the Hainan Government.They have been also started to be promoted in production, combining perfectly high quality and high yielding,thereby making important contributions to the quality improvement of rice grains in Hainan Province.

Breeding of hybrid rice combinations

Qiongxiangliangyou 1The Qiongxiangliangyou 1 is bred by crossing the dual-using genic male sterile line Qiongxiang S and the high-yielding and high-quality conventional rice variety Yuefengzhan,and it is a highquality two-line hybrid rice combination[22].The Qiongxiangliangyou 1 has the entire growth period of 135.8 d in average,plant height of 112.5 cm,effective panicle number per hectare of 3 015 000,panicle length of 23.9 cm,total grain number per panicle of 144.7,kilo-grain weight of 24.1 g and yield of 7 050-7 800 kg/hm2.The Qiongxiangliangyou 1 has been tested by the Rice c sterile line and Products Quality Supervision and Testing Center of the Ministry of Agriculture.The results showed that the Qiongxiangliangyou 1 has the brown rice rate of 80.7%,milled rice rate of 73.6%,head rice rate of 69.1%,grain length of 6.7 mm,length to width ratio of grain of 3.2,chalky rice rate of 8%,chalkiness of 2.0,transparency of 1 grade,alkali spreading value of 7.0,gel consistency of 46 mm,amylose content of 24.4% and protein content of 7.9%.Most indicators of Qiongxiangliangyou 1 meet the national second grade high-quality rice standards.In fields,Qiongxiangliangyou 1 shows resistance against rice blast and bacterial leaf blight.The successful breeding of Qiongxiangliangyou 1 sets a precedent for the high-quality and high-yielding aromatic rice breeding using two-line method in Hainan Province.

Qiongxiangliangyou 2The Qiongxiangliangyou 2 is bred by crossing the dual-using genic male sterile line Qiongxiang S and the high-yielding and high-quality conventional rice variety Texianzhan 25,and it is a two-line hybrid indica rice combination[23].The Qiongxiangliangyou 2 has the entire growth period of 114-136 d(which islonger than that of Texianzhan 25 by 1 d),effective panicle number of 3 060 000/hm2,plant height of 90.7-123.5 cm,panicle length of 23.6 cm, total grain number per panicle of 150.2,fruiting rate of 83.4%,kilo-grain weight of 25.1 g and yield of 7 050-7 800 kg/hm2.After tested by the Food Quality Supervision and Testing Center(Wuhan)of Ministry of Agriculture, the Qiongxiangliangyou 2 has the brown rice rate of 80.9%,milled rice rate of 71.3%,head rice rate of 57.0%, grain length of 7.0 mm,length to width ratio of grain of 3.0,chalky rice rate of 18%,chalkiness of 2.7,transparency of 2 grade,alkali spreading value of 7.0,gel consistency of 78 mm,amylose content of 18.65%.The grain quality of Qiongxiangliangyou 2 reaches the national second grade highquality rice standards by GB/T17891-1999.The average resistance of Qiongxiangliangyou 2 to rice blast is of 4th grade,and to bacterial leaf blight is of 8th grade.In fields,the resistances of Qiongxiangliangyou 2 to spike neck blast and bacterial leaf blight are equivalent to those of II-You 128 and Texianzhan 25.

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Responsible editor:Tingting XU

Responsible proofreader:Xiaoyan WU

香稻不育系GHS香味遗传及其杂交育种研究

郭国强1,2，尹建英1，郭武强3,4，曾良贵1，黄益国1，高静1，阳小虎1，郭名奇3*
（1.衡阳市农科所，湖南衡阳421000；2.湖南农业大学农学院，湖南长沙410125；3.国家杂交水稻工程技术研究中心，湖南长沙410125；4.湖南隆平种业有限公司，湖南长沙410125）

以香稻不育系GHS与无香味恢复系和香稻品种杂交，采用咀嚼法测定杂交组合F1米粒和KOH溶液法测定杂交组合F1、F2、BC1F1植株的香味遗传特点。结果表明，恢复系无香味为显性性状，GHS的香味为隐性性状；GHS的香味基因与粤香占、湘晚籼17号的香味基因不等位；GHS的香味基因与玉香油占、天龙香103、新万香、创香5号香味基因可能存有等位位点。与无香味恢复系杂交组合F2植株群体无香味与有香味之比符合3∶1的分离比，BC1F1植株群体符合1∶1的分离比，GHS的香味遗传受1对隐性核基因控制。在杂交水稻育种中，利用GHS系统选育出琼香S，并选配出一批香型杂交稻组合，其中琼香两优1号、琼香两优2号等高产优质组合已通过省级审定，具有较好的应用前景。

水稻；香味；遗传；育种

湖南省自然科学省市联合基金项目（14JJ5025）；衡阳市重点项目（2011KZ15, 2013KN01）；衡阳市一般项目（2010KN02）；国家水稻产业技术体系衡阳综合试验站项目（CARS-01-03A）。

郭国强（1970-），男，湖南隆回人，农艺师，博士研究生，主要从事杂交水稻遗传育种研究，E-mail：hnggq2008@163.com。*通讯作者，副研究员，从事杂交水稻遗传育种研究，E-mail：GGYYRRYY@163.com。

2015-01-29

修回日期 2015-05-09

Supported by Province-City Joint Funds of Natural Science of Hunan Province (14JJ5025);Key Program of Hengyang City(2011KZ15,2013KN01);General Program of Hengyang City(2010KN02);Hengyang Comprehensive Experimental Station of National Rice Industry Technology System(CARS-01-03A).

*Corresponding author.E-mail:GGYYRRYY@163.com

Received:January 29,2015 Accepted:May 9,2015