Development of a New High-Quality and Multi-Resistant japonica Rice Variety, Huxianggeng 216, Through Molecular Pyramiding Breeding

doi:10.16819/j.1001-7216.2025.240204

Abstract

Abstract:

【Objective】 Good eating quality and comprehensive resistance have always been important directions in the breeding of high-quality japonica varieties. To meet the demand for high-quality and multi-resistant rice varieties in production, it is urgent to enhance the synergistic improvement of quality and resistance and correct unfavorable traits in the breeding process. 【Method】Nangeng 46, a high-quality japonica rice variety harboring the fragrance gene badh2-E2 and the low-amylose-content gene Wx^mp, was employed as the female parent, while Wuxianggeng 1, a Luzon-flavor high-quality japonica variety, served as the male parent for cross-breeding. During the self-breeding process, molecular marker-assisted selection was carried out using badh2-E2 and Wx^mp, and seed selection was conducted based on field performance. The line with the most favorable comprehensive traits was chosen from the stable lines of F₈ and designated as Hu 12-30. Subsequently, Xiushui 114, a blast-resistant variety suitable for direct seeding and carrying the Pita and Pik-m genes, was used as the female parent, and Hu 12-30 as the male parent for cross-breeding. In the course of backcrossing and self-crossing, molecular marker-assisted selection was performed using badh2-E2, Wxmp, Pita, and Pik-m. By integrating quality determination, blast resistance identification, and identification of suitability for direct seeding, a new high-quality and resistant japonica rice variety, Huxianggeng 216, was developed. 【Result】Huxianggeng 216 contains the fragrance gene badh2-E2, the low-amylose-content gene Wx^mp, and the blast-resistance genes Pita and Pik-m. This variety has a fragrant aroma, a soft and elastic texture, with the amylose content ranging from 9.2% to 10.4%. The comprehensive resistance index for rice blast is 2.5, and the disease grade is 3, indicating moderate resistance. The resistance grade for rice stripe leaf blight is 3, showing resistance. Huxianggeng 216 has a high direct-seeding rate, with a plant height of 87 cm. It has strong lodging resistance and a yield of 650 kg per 667 m².【Conclusion】Molecular marker-assisted selection combined with phenotypic identification of target traits is an important means to achieve multi-target trait aggregation breeding, which can accurately and effectively develop high-quality multi-resistant japonica rice varieties.

Key words: rice, direct sowing, molecular pyramiding breeding, resistance to rice blast, taste quality

摘要：

【目的】食味品质优良、综合抗性好一直是优质粳稻育种的重要方向。为满足生产上对水稻品种优质多抗的需求，迫切需要在育种中加强对优质与抗病、抗逆性状的协同改良。【方法】以携带香味基因型badh2-E2和低直链淀粉含量基因型Wx^mp的优质粳稻品种南粳46为母本，浓香型优质粳稻品种武香粳1号为父本进行杂交配组。在连续多代的自交过程中，利用基因badh2-E2和Wx^mp进行分子标记辅助选择，结合田间表现进行选种，在F₈稳定株系中选择综合性状最优株系，命名为沪12-30。随后，以携带抗稻瘟病基因Pita和Pik-m的宜直播、抗稻瘟病品种秀水114为母本，沪12-30为父本进行杂交，回交一代后连续自交选种。在回交和自交过程中，对badh2-E2、Wx^mp、Pita和Pik-m进行分子标记辅助选择，结合品质测定、抗稻瘟病鉴定和宜直播性状鉴定筛选育成优质多抗粳稻新品种沪香粳216。【结果】沪香粳216含有香味基因型badh2-E2、低直链淀粉含量基因型Wx^mp、抗稻瘟病基因型Pita和Pik-m。该品种具有香味，口感柔软有弹性，直链淀粉含量为9.2%~10.4%；稻瘟病综合抗病指数为2.5，病级3级，表现为中抗；水稻条纹叶枯病抗病等级为3级，表现为抗病；直播成苗率高，株高87 cm，抗倒性强，每667 m²产量达650 kg。【结论】利用分子标记辅助选择结合目标性状表型鉴定是实现多目标性状聚合育种的重要手段，能够准确、有效地实现优质多抗粳稻品种选育。

关键词: 水稻, 直播, 分子聚合育种, 抗稻瘟病, 食味品质

YAN Ying, WANG Kai, ZHANG Lixia, HU Zejun, YE Junhua, YANG Hang, GU Chunjun, WU Shujun. Development of a New High-Quality and Multi-Resistant japonica Rice Variety, Huxianggeng 216, Through Molecular Pyramiding Breeding[J]. Chinese Journal OF Rice Science, 2025, 39(2): 209-219.

闫影, 王凯, 张丽霞, 胡泽军, 叶俊华, 杨航, 顾春军, 吴书俊. 利用分子聚合育种培育优质多抗粳稻新品种沪香粳216[J]. 中国水稻科学, 2025, 39(2): 209-219.

Figures/Tables 11

References 35

[1]	国家统计局. 关于2021年粮食产量数据的公告[EB/OL]. (2021-12-06) [2023-12-16]. http://www.stats.gov.cn/tjsj/zxfb/202112/t20211206_1825053.html.
	National Bureau of Statistics. Announcement on grain production data for 2021[EB/OL]. (2021-12-06) [2023-12-16]. http://www.stats.gov.cn/tjsj/zxfb/202112/t20211206_1825053.html. (in Chinese)
[2]	Qiao W H, Chen Y T, Wang R S, Wei X, Cao L R, Zhang W X, Yang Q W. Nucleotide diversity in waxy gene and validation of single nucleotide polymorphism in relation to amylose content in Chinese microcore rice germplasm[J]. Crop Science, 2012, 52(4): 1689-1697.
[3]	张昌泉, 冯琳皓, 顾铭洪, 刘巧泉. 江苏省水稻品质性状遗传和重要基因克隆研究进展[J]. 遗传, 2021, 43(5): 425-441.
	Zhang C Q, Feng L H, Gu M H, Liu Q Q. Progress on inheritance and gene cloning for rice grain quality in Jiangsu Province[J]. Hereditas(Beijing), 2021, 43(5): 425-441. (in Chinese with English abstract)
[4]	Verma D K, Srivastav P P. Introduction to rice aroma, flavor, and fragrance//Verma D K, Srivastav P P. Science and Technology of Aroma, Flavour and Fragrance in Rice[M]. USA: Apple Academic Press, 2019: 3-34.
[5]	He Q, Park Y J. Discovery of a novel fragrant allele and development of functional markers for fragrance in rice[J]. Molecular Breeding, 2015, 35(11): 217.
[6]	Bradbury L M T, Fitzgerald T L, Henry R J, Jin Q S, Waters D L E. The gene for fragrance in rice[J]. Plant Biotechnology Journal, 2005, 3(3): 363-370.
[7]	闫影, 诸光明, 张丽霞, 万常照, 曹黎明, 赵志鹏, 吴书俊. 水稻香味基因分子标记的开发及应用[J]. 西北植物学报, 2015, 35(2): 269-274.
	Yan Y, Zhu G M, Zhang L X, Wan C Z, Cao L M, Zhao Z P, Wu S J. Development of molecular markers for fragrant gene and its application[J]. Acta Botanica Boreali-Occidentalia Sinica, 2015, 35(2): 269-274. (in Chinese with English abstract)
[8]	Dean R, van Kan J A L, Pretorius Z A, Hammond- Kosack K E, Di Pietro A, Spanu P D, Rudd J J, Dickman M, Kahmann R, Ellis J, Foster G D. The top 10 fungal pathogens in molecular plant pathology[J]. Molecular Plant Pathology, 2012, 13(4): 414-430.
[9]	Zhou B, Qu S H, Liu G F, Dolan M, Sakai H, Lu G D, Bellizzi M, Wang G L. The eight amino-acid differences within three leucine-rich repeats between Pi2 and Piz-t resistance proteins determine the resistance specificity to Magnaporthe grisea[J]. Molecular Plant-Microbe Interactions, 2006, 19: 1216-1228.
[10]	Deng Y W, Zhai K R, Xie Z, Yang D Y, Zhu X D, Liu J Z, Wang X, Qin P, Yang Y Z, Zhang G M, Li Q, Zhang J F, Wu S Q, Milazzo J, Mao B Z, Wang E T, Xie H A, Tharreau D, He Z H. Epigenetic regulation of antagonistic receptors confers rice blast resistance with yield balance[J]. Science, 2017, 355: 962-965.
[11]	Lee S K, Song M Y, Seo Y S, Kim H K, Ko S, Cao P J, Suh J P, Yi G, Roh J H, Lee S, An G, Hahn T R, Wang G L, Ronald P, Jeon J S. Rice Pi5-mediated resistance to Magnaporthe oryzae requires the presence of two coiled- coil-nucleotide-binding-leucine-rich repeat genes[J]. Genetics, 2009, 181: 1627-1638.
[12]	Ashikawa I, Hayashi N, Yamane H, Kanamori H, Wu J Z, Matsumoto T, Ono K, Yano M. Two adjacent nucleotide- binding site-leucine-rich repeat class genes are required to confer Pikm-specific rice blast resistance[J]. Genetics, 2008, 180: 2267-2276.
[13]	Bryan G T, Wu K S, Farrall L, Jia Y, Hershey H P, McAdams S A, Faulk K N, Donaldson G K, Tarchini R, Valent B. A single amino acid difference distinguishes resistant and susceptible alleles of the rice blast resistance gene Pi-ta[J]. The Plant Cell, 2000, 12: 2033-2046.
[14]	李莹. 抗稻瘟病早粳稻空育131(Pid2/Pid3)的培育[D]. 哈尔滨: 黑龙江大学, 2015.
	Li Y. Cultivation of blast resistant early japonica rice Kongyu 131 (Pid2/Pid3)[D]. Harbin: Heilongjiang University, 2015. (in Chinese with English abstract)
[15]	国家水稻数据中心. 中国水稻品种及其系谱数据库[DB/OL]. http://www.ricedata.cn/variety/.
	National Rice Data Center. China Rice Varieties and Pedigree Database[DB/OL]. http://www.ricedata.cn/gene/. (in Chinese)
[16]	陈涛, 骆名瑞, 张亚东, 朱镇, 赵凌, 赵庆勇, 周丽慧, 姚姝, 于新, 王才林. 利用四引物扩增受阻突变体系PCR技术检测水稻低直链淀粉含量基因Wx-mq[J]. 中国水稻科学, 2013, 27(5): 529-534.
	Chen T, Luo M R, Zhang Y D, Zhu Z, Zhao L, Zhao Q Y, Zhou L H, Yao S, Yu X, Wang C L. Detection of Wx-mq gene for low-amylose content tetra-primer amplification refractory mutation system PCR in rice[J]. Chinese Journal Rice Science, 2013, 27(5): 529-534. (in Chinese with English abstract)
[17]	姚姝, 张亚东, 刘燕清, 赵春芳, 周丽慧, 陈涛, 赵庆勇, 朱镇, Pillay B, 王才林. Wx^mp基因背景下可溶性淀粉合成酶基因SSⅡa和去分支酶基因PUL对水稻蒸煮食味品质的影响[J]. 中国水稻科学, 2020, 34(3): 217-227.
	Yao S, Zhang Y D, Liu Y Q, Zhao C F, Zhou L H, Chen T, Zhao Q Y, Zhu Z, Pillay B, Wang C L. Allelic effects on eating and cooking quality of SSⅡa and PUL genes under Wx^mp background in rice[J]. Chinese Journal of Rice Science, 2020, 34(3): 217-227. (in Chinese with English abstract)
[18]	王军, 杨杰, 陈志德, 仲维功. 水稻香米基因标记的开发与应用[J]. 分子植物育种, 2008, 6(6): 1209-1212.
	Wang J, Yang J, Chen Z D, Zhong W G. Development and application of fragrance gene markers in rice[J]. Molecular Plant Breeding, 2008, 6(6): 1209-1212. (in Chinese with English abstract)
[19]	王忠华. 水稻抗稻瘟病基因Pi-ta分子标记的建立及其应用[D]. 杭州: 浙江大学, 2003.
	Wang Z H. Development of molecular markers of the rice blast resistance gene Pi-ta and its application[D]. Hangzhou: Zhejiang University, 2003. (in Chinese with English abstract)
[20]	范方军, 王芳权, 刘永峰, 王军, 朱金燕, 李文奇, 仲维功, 杨杰. Pi-b、Pi-ta、Pikm和Pi54对水稻穗颈瘟的抗性评价[J]. 华北农学报, 2014, 29(3): 221-226.
	Fan F J, Wang F Q, Liu Y F, Wang J, Zhu J Y, Li W Q, Zhong W G, Yang J. Evaluation of resistance to rice panicle blast with resistant genes Pi-b, Pi-ta, Pikm and Pi54[J]. Acta Agriculturae Boreali-sinica, 2014, 29(3): 221-226. (in Chinese with English abstract)
[21]	吴书俊, 闫影, 张丽霞, 万常照, 龚凯, 顾永平. 分子标记辅助选择培育抗条纹叶枯病香粳品种沪香粳106[J]. 上海农业学报, 2017, 33(3): 25-30.
	Wu S J, Yan Y, Zhang L X, Wan C Z, Gong K, Gu Y P. Breeding of RSD-resistant and fragrant japonica variety ‘Huxiangjing106’ by marker assisted selection in rice(Oryza sativa)[J]. Acta Agriculturae Shanghai, 2017, 33(3): 25-30. (in Chinese with English abstract)
[22]	陈桂华, 邓化冰, 张桂莲, 唐文帮, 黄璜. 水稻茎秆性状与抗倒性的关系及配合力分析[J]. 中国农业科学, 2016, 49(3): 407-417.
	Chen G H, Deng H B, Zhang G L, Tang W B, Huang H. The correlation of stem characters and lodging resistance and combining ability analysis in rice[J]. Scientia Agricultura Sinica, 2016, 49(3): 407-417. (in Chinese with English abstract)
[23]	陈涛, 赵庆勇, 朱镇, 赵凌, 姚姝, 周丽慧, 赵春芳, 张亚东, 王才林. 利用分子标记辅助选择培育优良食味、低谷蛋白香粳稻新品系[J]. 中国水稻科学, 2023, 37(1): 55-65.
	Chen T, Zhao Q Y, Zhu Z, Zhao L, Yao S, Zhou L H, Zhao C F, Zhang Y D, Wang C L. Development of new low glutelin content japonica rice lines with good eating quality and fragrance by molecular marker-assisted selection[J]. Chinese Journal of Rice Science, 2023, 37(1): 55-65. (in Chinese with English abstract)
[24]	降好宇, 曾盖, 郝明, 黄湘桂, 肖应辉. 广谱抗稻瘟病种质75-1-127的褐飞虱抗性基因鉴定及分子标记辅助选择育种[J]. 中国水稻科学, 2019, 33(2): 227-234.
	Jiang H Y, Zeng G, Hao M, Huang X G, Xiao Y H. Identification of brown planthopper resistance genes in broad-spectrum blast resistant rice germplasm 75-1-127 and its molecular marker-assisted selection breeding[J]. Chinese Journal of Rice Science, 2019, 33(2): 227-234. (in Chinese with English abstract)
[25]	马作斌, 全东兴, 时羽, 王昌华, 周广春, 郑文静. 分子标记辅助选育聚合抗稻瘟病基因Pi5及Pita的水稻新品系[J]. 分子植物育种, 2021(1): 173-179.
	Ma Z B, Quan D X, Shi Y, Wang C H, Zhou G C, Zheng W J. Molecular markers-assisted selection of new rice lines with blast resistance gene Pi5 and Pita[J]. Molecular Plant Breeding, 2021(1): 173-179. (in Chinese with English abstract)
[26]	上海市农业农村委员会. 关于2021年水稻机械化种植情况的通报[EB/OL]. (2021-08-25) [2023-12-25]. http://nyncw.sh.gov.cn/xxgk/20210825/1234567890abcdef.html.
	Shanghai Agriculture and Rural Affairs Committee. Bulletin on Mechanized Rice Planting in 2021[EB/OL]. (2021-08-25) [2023-12-25]. http://nyncw.sh.gov.cn/xxgk/20210825/1234567890abcdef.html. (in Chinese)
[27]	熊怀阳, 李阳生. 水稻的耐淹性状及其Sub1基因[J]. 遗传, 2010, 32(9): 886-892.
	Xiong H Y, Li Y S. Submergence tolerance and Sub1 locus in rice[J]. Hereditas(Beijing), 2010, 32(9): 886-892. (in Chinese with English abstract)
[28]	Kretzschmar T, Pelayo M A F, Trijatmiko K R, Gabunada L F M, Alam R, Jimenez R, Mendioro M S, Slamet-Loedin I H, Sreenivasulu N, Bailey-Serres J, Ismail A M, Mackill D J. A trehalose-6-phosphate phosphatase enhances anaerobic germination tolerance in rice[J]. Nature Plants, 2015, 1(9): 15124.
[29]	Ye N H, Wang F Z, Shi L, Chen M X, Cao Y Y, Zhu F Y, Wu Y Z, Xie L J, Liu T Y, Su Z Z, Xiao S, Zhang H, Yang J, Gu H Y, Hou X X, Hu Q J, Yi H J, Zhu C X, Zhang J, Liu Y G. Natural variation in the promoter of rice calcineurin B-like protein 10(OsCBL10) affects flooding tolerance during seed germination among rice subspecies[J]. The Plant Journal, 2018, 94(4): 612-625.
[30]	Sun J, Zhang G C, Cui Z B, Kong X M, Yu X Y, Gui R, Han Y Q, Li Z, Lang H, Hua Y C, Zhang X M, Xu Q, Tang L, Xu Z J, Ma D R, Chen W F. Regain flood adaptation in rice through a 14-3-3 protein OsGF14h[J]. Nature Communications, 2022, 13: 5664.
[31]	He Y Q, Sun S, Zhao J, Huang Z B, Peng L L, Huang C W, Tang Z B, Huang Q Q, Wang Z F. UDP- glucosyltransferase OsUGT75A promotes submergence tolerance during rice seed germination[J]. Nature Communications, 2023, 14: 2296.
[32]	Lü Y S, Shao G N, Jiao G A, Sheng Z H, Xie L H, Hu S K, Tang S Q, Wei X J, Hu P S. Targeted mutagenesis of POLYAMINE OXIDASE 5 that negatively regulates mesocotyl elongation enables the generation of direct-seeding rice with improved grain yield[J]. Molecular Plant, 2021, 14(2): 344-351.
[33]	Xiong Q, Ma B, Lu X, Huang Y H, He S J, Yang C, Yin C C, Zhao H, Zhou Y, Zhang W K, Wang W S, Li Z K, Chen S Y, Zhang J S. Ethylene-inhibited jasmonic acid biosynthesis promotes mesocotyl/coleoptile elongation of etiolated rice seedlings[J]. The Plant Cell, 2017, 29: 1053-1072.
[34]	Sun S Y, Wang T, Wang L L, Li X M, Jia Y C, Liu C, Huang X H, Xie W B, Wang X L. Natural selection of a GSK3 determines rice mesocotyl domestication by coordinating strigolactone and brassinosteroid signaling[J]. Nature Communications, 2018, 9(1): 2523.
[35]	李先锋. 水稻热精胺合成酶基因OsACL5的功能研究[D]. 扬州: 扬州大学, 2022.
	Li X F. Functional analysis of OsACL5, a gene putatively encoding thermospermine synthase[D]. Yangzhou: Yangzhou University, 2022. (in Chinese with English abstract)

性状 Trait	等位基因 Allele	分子标记 Molecular marker	引物序列（5’-3’） Primer sequence(5’-3’)	多态性 Polymorphism
香味Fragrance	badh2-E2	InDel-E2	F：GGGAGGCGCTGAAGAGGA	100 bp/107 bp
香味Fragrance	badh2-E2	InDel-E2	R：GGGTAGTCACCACCCTACCTTG	100 bp/107 bp
低直链淀粉含量 Low amylose content	Wx^mp	Wx-mp-o	F：ATGTTGTGTTCTTGTGTTCTTTGCAGGC	A/G
		Wx-mp-o	R：GTAGATCTTCTCACCGGTCTTTCCCCAA	A/G
		KASP-WXMP	F1：GAAGGTGACCAAGTTCATGCTGATGAA CACACGGTCGACTCCAT	A/G
			F2：GAAGGTCGGAGTCAACGGATTATGAAC ACACGGTCGACTCCAC
			R：GAGAGGGTGAGGTTTTTCCATTGCTA
抗稻瘟病 Resistance to rice blast	Pita	YL155/YL87	F：AGCAGGTTATAAGCTAGGCC	1042 bp/-
抗稻瘟病 Resistance to rice blast	Pita	YL155/YL87	R：CTACCAACAAGTTCATCAAA	1042 bp/-
感稻瘟病 Blast susceptibility	pita	YL183/YL87	F：AGCAGGTTATAAGCTAGCTAT	-/1042 bp
感稻瘟病 Blast susceptibility		YL183/YL87	R：CTACCAACAAGTTCATCAAA	-/1042 bp
抗稻瘟病 Resistance to rice blast	Pikm	Pikm-1	1F：TGAGCTCAAGGCAAGAGTTGAGGA	174 bp / 213 bp
		Pikm-1	1R：TGTTCCAGCAACTCGATGAG	174 bp / 213 bp
		Pikm-2	2F：CAGTAGCTGTGTCTCAGAACTATG	290 bp / 332 bp
		Pikm-2	2R：AAGGTACCTCTTTTCGGCCAG	290 bp / 332 bp

性状 Trait	等位基因 Allele	分子标记 Molecular marker	引物序列（5’-3’） Primer sequence(5’-3’)	多态性 Polymorphism
香味Fragrance	badh2-E2	InDel-E2	F：GGGAGGCGCTGAAGAGGA	100 bp/107 bp
香味Fragrance	badh2-E2	InDel-E2	R：GGGTAGTCACCACCCTACCTTG	100 bp/107 bp
低直链淀粉含量 Low amylose content	Wx^mp	Wx-mp-o	F：ATGTTGTGTTCTTGTGTTCTTTGCAGGC	A/G
		Wx-mp-o	R：GTAGATCTTCTCACCGGTCTTTCCCCAA	A/G
		KASP-WXMP	F1：GAAGGTGACCAAGTTCATGCTGATGAA CACACGGTCGACTCCAT	A/G
			F2：GAAGGTCGGAGTCAACGGATTATGAAC ACACGGTCGACTCCAC
			R：GAGAGGGTGAGGTTTTTCCATTGCTA
抗稻瘟病 Resistance to rice blast	Pita	YL155/YL87	F：AGCAGGTTATAAGCTAGGCC	1042 bp/-
抗稻瘟病 Resistance to rice blast	Pita	YL155/YL87	R：CTACCAACAAGTTCATCAAA	1042 bp/-
感稻瘟病 Blast susceptibility	pita	YL183/YL87	F：AGCAGGTTATAAGCTAGCTAT	-/1042 bp
感稻瘟病 Blast susceptibility		YL183/YL87	R：CTACCAACAAGTTCATCAAA	-/1042 bp
抗稻瘟病 Resistance to rice blast	Pikm	Pikm-1	1F：TGAGCTCAAGGCAAGAGTTGAGGA	174 bp / 213 bp
		Pikm-1	1R：TGTTCCAGCAACTCGATGAG	174 bp / 213 bp
		Pikm-2	2F：CAGTAGCTGTGTCTCAGAACTATG	290 bp / 332 bp
		Pikm-2	2R：AAGGTACCTCTTTTCGGCCAG	290 bp / 332 bp

品种 Variety	香味 Fragrance	直链淀粉含量 Amylose content(%)	穗颈瘟抗性评价 Evaluation of resistance to neck blast
南粳46 Nangeng 46	有 Yes	10.1±0.44 c	高感Highly susceptible(HS)
武香粳1号Wuxianggeng 1	有 Yes	16.0±0.17 b	高感Highly susceptible(HS)
秀水114 Xiushui 114	无 No	17.6±0.20 a	抗病 Resistant(R)

品种 Variety	香味 Fragrance	直链淀粉含量 Amylose content(%)	穗颈瘟抗性评价 Evaluation of resistance to neck blast
南粳46 Nangeng 46	有 Yes	10.1±0.44 c	高感Highly susceptible(HS)
武香粳1号Wuxianggeng 1	有 Yes	16.0±0.17 b	高感Highly susceptible(HS)
秀水114 Xiushui 114	无 No	17.6±0.20 a	抗病 Resistant(R)

品系 Line	直链淀粉 Amylose content(%)	胶稠度 Gel consistency (m)	食味值 Taste value	峰值黏度 Peak viscosity (RVU)	热浆黏度Hot paste viscosity (RVU)	崩解值 Breakdown (RVU)	终值黏度 Final viscosity (RVU)	消减值Setback (RVU)	糊化温度 Gelatinization temperature (℃)
ZX1	8.7±0.3 e	80.3±1.5 cd	74.5±1.0 d	315.6±6.4 ab	209.4±12.3 a	106.2±6.0 b	310.3±6.8 a	−5.3±0.4 b	73.1±1.2 a
ZX2	9.2±0.3 d	82.7±2.1 bc	81.8±0.8 c	235.3±7.2 c	135.1±6.3 b	100.1±0.8 b	238.7±7.4 c	3.4±0.3 a	70.5±1.1 ab
ZX3	11.6±0.2 ab	77.3±2.3 de	82.3±0.5 c	285.6±7.7 b	170.8±22.7 ab	114.8±15.0 b	263.7±13.2 bc	−22.0±5.5 c	71.1±0.5 ab
ZX4	11.2±0.3 b	79.3±1.2 d	84.8±0.4 b	290.4±7.1 ab	179.3±5.7 ab	111.1±12.8 b	245.6±5.5 c	−44.7±1.7 d	73.2±1.3 a
ZX5	9.8±0.4 c	79.7±1.5 d	84.3±0.5 b	303.0±41.9 ab	161.9±40.4 ab	141.1±1.6 a	219.8±41.9 c	−83.1±0.1 e	70.3±0.6 ab
ZX6	9.3±0.1 cd	85.3±0.6 ab	86.0±0.0 a	336.7±38.8 a	182.8±27.7 ab	153.9±11.2 a	236.7±38.1 c	−100.0±0.8 e	70.7±1.1 ab
ZX7	12.0±0.4 a	75.3±1.5 de	81.8±1.2 c	303.5±12.9 ab	199.9±15.2 a	103.5±2.3 b	305.7±13.1 ab	2.2±0.2 a	71.9±1.7 ab
ZX8	8.5±0.2 e	86.7±1.5 a	85.0±0.0 b	307.4±2.3 ab	168.2±6.5 ab	139.2±4.2 a	228.6±0.1 c	−78.8±2.4 e	70.0±0.0 b