Wx与ALK主要等位基因不同组合对稻米品质的影响

doi:10.16819/j.1001-7216.2020.9140

中国水稻科学 ›› 2020, Vol. 34 ›› Issue (3): 228-236.DOI: 10.16819/j.1001-7216.2020.9140

Wx与ALK主要等位基因不同组合对稻米品质的影响

陈专专¹, 杨勇¹, 冯琳皓¹, 孙晔¹, 张昌泉^1,², 范晓磊^1,², 李钱峰^1,², 刘巧泉^1,^2,^*()

¹扬州大学农学院植物功能基因组学教育部重点实验室/江苏省作物基因组学和分子育种重点实验室, 江苏扬州 225009
²扬州大学江苏省作物遗传生理重点实验室/江苏省粮食作物现代产业技术协同创新中心, 江苏扬州225009

收稿日期:2019-12-29 修回日期:2020-01-13 出版日期:2020-05-15 发布日期:2020-05-10
通讯作者: 刘巧泉
基金资助:
国家自然科学基金资助项目(31825019, U19A2032);国家重点研发计划资助项目(2016YFD0100501);江苏省科技计划资助项目[BE2018357, CX(18)1001和19KJA560006];扬州大学农学院本科生创新训练计划资助项目

Effects of Different Combinations of Wx and ALK Main Alleles on Rice Grain Quality

Zhuanzhuan CHEN¹, Yong YANG¹, Linhao FENG¹, Ye SUN¹, Changquan ZHANG^1,², Xiaolei FAN^1,², Qianfeng LI^1,², Qiaoquan LIU^1,^2,^*()

¹Key Laboratory of Plant Functional Genomics of Ministry of Education / Jiangsu Key Laboratory of Crop Genomics and Molecular Breeding, College of Agriculture, Yangzhou University, Yangzhou 225009, China
²Jiangsu Key Laboratory of Crop Genetics and Physiology / Jiangsu Collaborative Innovation Center for Modern Industrial Technology of Grain Crops, Yangzhou University, Yangzhou 225009, China

Received:2019-12-29 Revised:2020-01-13 Online:2020-05-15 Published:2020-05-10
Contact: Qiaoquan LIU

摘要/Abstract

摘要：

【目的】稻米蒸煮和食味品质主要受Wx和ALK两个主效基因的调控。这两个基因在栽培稻中存在多个复等位变异,本研究旨在明确Wx和ALK基因的主要等位变异及其组合对稻米品质的影响。【方法】将Wx的3个主要等位基因（Wx^a、Wx^b和wx）与ALK的2个主要等位基因（ALK^a和ALK^c）进行不同的组合,在粳稻品种日本晴（Nip）背景下创建6个含有Wx和ALK基因不同等位变异组合的近等基因系。通过测定近等基因系的稻米理化品质明确不同等位基因及其组合对稻米品质的效应。【结果】在相同的ALK基因型背景下,含Wx^a等位基因稻米的糊化温度极显著低于wx型和Wx^b型的近等基因系。稻米黏滞性受Wx和ALK等位基因组合影响较大,Wx等位变异主要影响稻米的峰值黏度、峰值时间和冷胶黏度;在相同Wx背景下,ALK等位基因主要影响稻米RVA谱的起浆温度,对稻米崩解值和消减值的影响不显著。总体来看,Nip-Wx^a/ALK^a和Nip-Wx^a/ALK^c型稻米具有较高直链淀粉含量和崩解值,较低的胶稠度,因而食味值较差;Nip-wx/ALK^a和Nip-Wx^b/ALK^a型水稻具有适中的直链淀粉含量、较软的胶稠度以及较低的糊化温度,因此这两个近等基因系热饭和冷饭的食味值较好;虽然Nip-Wx^b/ALK^c的糊化温度较高,但因其适中的直链淀粉含量和较高的胶稠度,其冷饭食味值降幅较小;Nip-wx/ALK^c稻米的热饭食味值与Nip-wx/ALK^a、Nip-Wx^b/ALK^a和Nip-Wx^b/ALK^c无差异,但因其较高的糊化温度,其冷饭食味值显著下降。【结论】Wx基因变异主要影响稻米直链淀粉含量和胶稠度,对糊化温度有一定的影响;ALK基因变异主要影响糊化温度,对直链淀粉含量无显著影响,对胶稠度存在微效影响。

关键词: 蜡质基因, 糊化温度基因, 等位基因, 近等基因系, 稻米品质

Abstract:

【Objective】Rice eating and cooking qualities are mainly controlled by two key genes, Wx and ALK. There are multiple alleles of both genes among rice cultivars. In this study, our aim is to clarify the effects of the main allelic variation combinations of Wx and ALK genes on rice grain quality.【Method】Three main alleles of Wx (Wx^a, Wx^b and wx) and two of ALK (ALK^a and ALK^c) were combined within a series of near-isogenic lines (NILs) in the japonica Nipponbare (Nip) background. The effects of different allelic combinations on grain quality were determined by measuring the physical and chemical characteristics of NILs.【Result】Gelatinization temperature (GT) was regulated by the major gene ALK and the minor gene Wx. Under the same ALK allele background, there was no significant difference in GT between rice lines carrying wx and Wx^b alleles, but the GT of Nip-Wx^a rice lines was significantly lower than that of Nip-Wx^b and Nip-wx. The starch viscosity was greatly affected by the combination of Wx and ALK alleles. The breakdown value of Nip-Wx^a rice line was lower and the setback value was higher (lower tasting value) than those of Nip-Wx^b and Nip-wx. Under the same Wx allele background, ALK allelic variation mainly affected the pasting temperature of RVA curve, but had little effect on breakdown value and setback value. Nip-Wx^a/ALK^a and Nip-Wx^a/ALK^c rice grains had poor tasting value due to the high AC and breakdown value and low GC. But the protein content in above two NILs was higher than other rice lines. Nip-wx/ALK^a and Nip-Wx^b/ALk^a rice grains had better tasting value of hot cooked and cooled cooked rice due to the moderate AC, GC and protein content, as well as lower GT. Although the GT of Nip-Wx^b/ALK^c grains was higher, the moderate AC and high GC of this line resulted in a slight decrease in the tasting value of cooled cooked rice compared with hot cooked rice. The tasting value of hot cooked rice of Nip-wx/ALK^c rice remained the same as those of Nip-wx/ALK^a, Nip-Wx^b/ALK^a and Nip-Wx^b/ALK^c, but the tasting value of cooled rice decreased dramatically because of its high GT. 【Conclusion】Amylose content (AC) and gel consistency (GC) were mainly controlled by Wx gene; while ALK gene had no significant effect on AC, but a slight effect on GC.

Key words: waxy gene, alkali digestion gene, allelic variation, near isogenic lines, grain quality

中图分类号:

陈专专, 杨勇, 冯琳皓, 孙晔, 张昌泉, 范晓磊, 李钱峰, 刘巧泉. Wx与ALK主要等位基因不同组合对稻米品质的影响[J]. 中国水稻科学, 2020, 34(3): 228-236.

Zhuanzhuan CHEN, Yong YANG, Linhao FENG, Ye SUN, Changquan ZHANG, Xiaolei FAN, Qianfeng LI, Qiaoquan LIU. Effects of Different Combinations of Wx and ALK Main Alleles on Rice Grain Quality[J]. Chinese Journal OF Rice Science, 2020, 34(3): 228-236.

图/表 8

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近等基因系 Near-isogenic line	Wx基因型 Wx allele	ALK基因型 ALK allele	Wx基因供体 Donor of Wx gene	ALK基因供体 Donor of ALK gene	世代 Generation
Nip-wx/ALK^a	wx	ALK^a	苏御糯 Suyunuo	日本晴 Nipponbare	BC₉F₈
Nip-wx/ALK^c	wx	ALK^c	苏御糯 Suyunuo	中恢9308 Zhonghui 9308	BC₆F₁₀
Nip-Wx^b/ALK^a	Wx^b	ALK^a	日本晴 Nipponbare	日本晴 Nipponbare
Nip-Wx^b/ALK^c	Wx^b	ALK^c	日本晴 Nipponbare	中恢9308 Zhonghui 9308	BC₈F₁₀
Nip-Wx^a/ALK^a	Wx^a	ALK^a	龙特甫 Longtefu	日本晴 Nipponbare	BC₈F₇
Nip-Wx^a/ALK^c	Wx^a	ALK^c	龙特甫 Longtefu	中恢9308 Zhonghui 9308	BC₆F₃

近等基因系 Near-isogenic line	Wx基因型 Wx allele	ALK基因型 ALK allele	Wx基因供体 Donor of Wx gene	ALK基因供体 Donor of ALK gene	世代 Generation
Nip-wx/ALK^a	wx	ALK^a	苏御糯 Suyunuo	日本晴 Nipponbare	BC₉F₈
Nip-wx/ALK^c	wx	ALK^c	苏御糯 Suyunuo	中恢9308 Zhonghui 9308	BC₆F₁₀
Nip-Wx^b/ALK^a	Wx^b	ALK^a	日本晴 Nipponbare	日本晴 Nipponbare
Nip-Wx^b/ALK^c	Wx^b	ALK^c	日本晴 Nipponbare	中恢9308 Zhonghui 9308	BC₈F₁₀
Nip-Wx^a/ALK^a	Wx^a	ALK^a	龙特甫 Longtefu	日本晴 Nipponbare	BC₈F₇
Nip-Wx^a/ALK^c	Wx^a	ALK^c	龙特甫 Longtefu	中恢9308 Zhonghui 9308	BC₆F₃

鉴定基因 Target gene	引物名称 Primer	引物序列 Primer sequence (5'-3')	区分基因型 Genotype	检测突变位置 Mutation site
Wx	QRM190-F	ATTCCTTCAGTTCTTTGTCTATCTCA	Wx^a/Wx^b	第1内含子CT序列多态性 (CT)₇ in Intron 1
	QRM190-R	TCCTGATGAACAACAGAACAACAC	Wx^a/Wx^b	第1内含子CT序列多态性 (CT)₇ in Intron 1
	ND-F	CACAGCAACAGCTAGACAACCAC	Wx/wx	第2外显子插入缺失23 bp 23 bp InDel in Exon 2
	ND-R	CACGACGACGGAGGGGAAC	Wx/wx	第2外显子插入缺失23 bp 23 bp InDel in Exon 2
ALK	4211(A/G)-F₁	GCGGGCTGAGGGACACCG	ALK^a/ALK^c	第8外显子单核苷酸多态性(A/G) A/G SNP in Exon 8
	4211(A/G)-R₁	AACGGGTCGAACGCCGACGT
	4211(A/G)-F	AACAGCAAGGTGCGCGGGTG
	4211(A/G)-R	CCACCGAGTCGCACCTCCACA

鉴定基因 Target gene	引物名称 Primer	引物序列 Primer sequence (5'-3')	区分基因型 Genotype	检测突变位置 Mutation site
Wx	QRM190-F	ATTCCTTCAGTTCTTTGTCTATCTCA	Wx^a/Wx^b	第1内含子CT序列多态性 (CT)₇ in Intron 1
	QRM190-R	TCCTGATGAACAACAGAACAACAC	Wx^a/Wx^b	第1内含子CT序列多态性 (CT)₇ in Intron 1
	ND-F	CACAGCAACAGCTAGACAACCAC	Wx/wx	第2外显子插入缺失23 bp 23 bp InDel in Exon 2
	ND-R	CACGACGACGGAGGGGAAC	Wx/wx	第2外显子插入缺失23 bp 23 bp InDel in Exon 2
ALK	4211(A/G)-F₁	GCGGGCTGAGGGACACCG	ALK^a/ALK^c	第8外显子单核苷酸多态性(A/G) A/G SNP in Exon 8
	4211(A/G)-R₁	AACGGGTCGAACGCCGACGT
	4211(A/G)-F	AACAGCAAGGTGCGCGGGTG
	4211(A/G)-R	CCACCGAGTCGCACCTCCACA

基因型 Genotype	起始糊化温度 T_o /℃	峰值糊化温度 T_p /℃	终止糊化温度 T_c /℃	热焓值 △H / (J·g^-1)
Nip-wx/ALK^a	65.7±0.2 e	72.6±0.1 d	80.5±0.1 c	9.98±0.55 bc
Nip-Wx^b/ALK^a	67.5±0.1 d	72.6±0.1 d	79.7±0.4 c	9.45±0.31 c
Nip-Wx^a/ALK^a	63.6±0.3 f	69.4±0.1 e	76.6±0.6 d	6.80±0.33 d
Nip-wx/ALK^c	75.2±0.1 b	80.6±0.0 b	86.8±0.1 a	11.93±0.35 a
Nip-Wx^b/ALK^c	76.7±0.1 a	80.8±0.0 a	86.4±0.1 a	11.03±0.29 ab
Nip-Wx^a/ALK^c	73.7±0.0 c	78.2±0.0 c	84.2±0.0 b	9.19±0.46 c

Wx与ALK主要等位基因不同组合对稻米品质的影响

Effects of Different Combinations of Wx and ALK Main Alleles on Rice Grain Quality

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Metrics

基因型 Genotype	峰值黏度 Peak viscosity /cP	热浆黏度 Hot paste viscosity /cP	崩解值 Breakdown /cP	冷胶黏度Cool paste viscosity /cP	消减值 Setback /cP	峰值时间 Peak time /min	起浆温度 Pasting temperature /℃
Nip-wx/ALK^a	2251±7 e	1060±18 f	1191±7 d	1319±18 e	-932±5 d	3.7±0 f	74.1±0 e
Nip-wx/ALK^c	2876±11 d	1517±15 e	1359±9 c	1932±13 d	-944±7 d	4.4±0 e	81.3±0 b
Nip-Wx^b/ALK^a	4205±9 a	2392±22 c	1813±13 b	3619±11 b	-586±8 b	6.1±0 c	75.0±0 d
Nip-Wx^b/ALK^c	4164±20 b	2008±19 d	2156±15 a	3285±9 c	-879±9 c	5.7±0 d	82.1±0 a
Nip-Wx^a/ALK^a	3503±8 c	3145±23 a	358±8 f	4107±7 a	604±6 a	6.9±0 a	73.2±0 f
Nip-Wx^a/ALK^c	3533±15 c	2772±17 b	761±6 e	4169±12 a	636±5 a	6.5±0 b	80.5±0 c

样本 Sample	基因型 Genotype		外观 Appearance	硬度 Hardness
热饭 Hot cooked rice	Nip-wx/ALK^a	98.00±0.00 a	9.80±0.00 a	0.20±0.00 e	9.80±0.00 a	9.80±0.00 a
	Nip-wx/ALK^c	98.00±0.00 a	9.80±0.00 a	0.53±0.10 d	9.80±0.00 a	9.80±0.00 a
	Nip-Wx^b/ALK^a	97.00±0.00 a	9.80±0.00 a	2.03±0.05 c	9.43±0.05 b	9.80±0.00 a
	Nip-Wx^b/ALK^c	97.00±0.82 a	9.80±0.00 a	2.00±0.14 c	9.40±0.08 b	9.80±0.00 a
	Nip-Wx^a/ALK^a	81.25±0.50 c	8.50±0.08 c	4.18±0.10 a	6.40±0.08 c	8.50±0.08 c
	Nip-Wx^a/ALK^c	82.50±1.29 b	8.65±0.13 b	3.98±0.10 b	6.35±0.17 c	8.60±0.08 b
冷饭 Cooled cooked rice	Nip-wx/ALK^a	93.00±0.82 a	9.80±0.00 a	2.33±0.10 d	8.55±0.17 a	9.80±0.00 a
	Nip-wx/ALK^c	80.75±2.22 c	8.65±0.17 c	3.53±0.17 c	5.08±0.29 d	8.40±0.22 cd
	Nip-Wx^b/ALK^a	87.25±0.50 b	9.20±0.08 b	3.48±0.13 c	7.35±0.10 b	9.10±0.08 b
	Nip-Wx^b/ALK^c	87.00±0.82 b	9.15±0.17 b	3.35±0.17 c	7.15±0.24 b	9.15±0.17 b
	Nip-Wx^a/ALK^a	79.50±0.58 c	8.30±0.08 d	4.43±0.10 a	6.25±0.13 c	8.23±0.10 d
	Nip-Wx^a/ALK^c	81.25±0.96 c	8.53±0.10 c	4.03±0.15 b	6.08±0.05 c	8.50±0.14 c

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