中国水稻科学 ›› 2019, Vol. 33 ›› Issue (6): 513-522.DOI: 10.16819/j.1001-7216.2019. 9090
陈专专1,2, 李先锋1, 仲敏1, 葛家奇1, 范晓磊1,2, 张昌泉1,2, 刘巧泉1,2,*()
收稿日期:
2019-08-07
修回日期:
2019-09-13
出版日期:
2019-11-10
发布日期:
2019-11-10
通讯作者:
刘巧泉
基金资助:
Zhuanzhuan CHEN1,2, Xianfeng LI1, Min ZHONG1, Jiaqi GE1, Xiaolei FAN1,2, Changquan ZHANG1,2, Qiaoquan LIU1,2,*()
Received:
2019-08-07
Revised:
2019-09-13
Online:
2019-11-10
Published:
2019-11-10
Contact:
Qiaoquan LIU
摘要:
目的 稻米糊化温度是影响稻米品质的重要指标,该性状受主效基因ALK/SSII-3调控,ALK基因具有多个复等位基因,本研究旨在通过RNAi技术明确籼稻亚种中两个不同ALK等位基因的效应。方法 以分别含有ALKc和ALKb等位基因的高糊化温度品种珍汕97B和低糊化温度品种龙特甫B为试验材料,使用RNAi技术构建ALK表达下调的转基因株系,通过对其稻米理化品质的测定来明确不同等位基因表达下调对稻米品质的影响。结果 对不同转基因水稻目的基因的表达分析显示本研究中转基因株系的ALK基因受到了不同程度的干扰。重点分析了不同RNAi株系稻米的糊化温度,结果表明珍汕97B的RNAi转基因稻米的糊化温度极显著降低,而在低糊化温度品种龙特甫B背景中下调表达ALK基因后对糊化温度的影响较小;转基因株系与未转化亲本相比,米粉的起始糊化温度都显著降低,表现为提前糊化;在珍汕97B背景下干扰系的峰值温度与未转化亲本相比极显著降低,而在龙特甫背景下米粉的峰值温度与未转化对照相比显著降低。对不同转基因系的理化品质分析表明,ALK下调表达植株稻米的表观直链淀粉含量显著增加,下调表达ALK后会引起米粉峰值黏度和崩解值的改变。高糊化温度品种珍汕97B干扰系与未转化对照相比胶稠度呈现极显著性差异,而低糊化温度品种龙特甫干扰系的胶稠度与未转化对照相比没有差异。结论 下调表达ALK等位基因对稻米理化品质产生显著影响,并且干扰不同等位基因的效应存在明显差异,即籼稻中的两个ALK等位基因的效应存在显著差异。
中图分类号:
陈专专, 李先锋, 仲敏, 葛家奇, 范晓磊, 张昌泉, 刘巧泉. 籼稻背景下抑制不同ALK等位基因表达对稻米品质的影响[J]. 中国水稻科学, 2019, 33(6): 513-522.
Zhuanzhuan CHEN, Xianfeng LI, Min ZHONG, Jiaqi GE, Xiaolei FAN, Changquan ZHANG, Qiaoquan LIU. Grain Quality as Affected by Down-regulation of Expression of Different ALK Alleles in indica Rice (Oryza sativa L.)[J]. Chinese Journal OF Rice Science, 2019, 33(6): 513-522.
图1 ALK-RNAi载体结构及转基因水稻的鉴定 A-用引物4342(GC/TT)鉴定亲本对照龙特甫B(LTFB)和珍汕97B(ZS97B)的ALK基因型。B-含ALK-RNAi 结构双元载体的T-DNA区结构。P35S和Tnos, 分别表示CaMV35S基因的启动子和终止子区; NOS-农杆菌胭脂碱合成酶基因终止子; Hyg-潮霉素抗性基因; LB和RB分别表示T-DNA区的左右边界序列; Anti和Sense分别表示目的基因片段的反向和正向结构;图中箭头表示设计引物方向。C-ALK-RNAi转基因水稻植株的PCR鉴定。D-ALK-RNAi转基因水稻植株胚乳中ALK基因表达量分析。L-ALKb-RNAi-1、L-ALKb-RNAi-2和L-ALKb-RNAi-3为龙特甫背景下的ALKb-RNAi转基因纯合系;Z-ALKc-RNAi-2-1、Z-ALKc-RNAi-2-2和Z-ALKc-RNAi-2-3为同时携带有ALKc等位基因和ALKb-RNAi干扰结构的转基因材料纯合系。对不同背景下的转基因系和未转化对照的表达量进行方差分析,并用新复极差法进行多重比较。**表示转基因系与未转化对照之间的差异达0.01显著水平,*表示转基因系与未转化对照之间的差异达0.05显著水平(n=3)。
Fig. 1. ALK-RNAi construction and idendification of transgenic rice. A, ALK genotype in Longtefu B(LTFB) and Zhenshan 97B(ZS97B) on sites 4342 and 4343 based on primer 4342(GC/TT). B, The T-DNA structure of ALK-RNAi construct. P35S and Tnos represent the promoter and terminator region of the CaMV 35S gene respectively; NOS represents agrobacterium nopaline synthetase gene terminator; Hyg represents hygromycin resistance gene; LB and RB represent the left and right boundary sequences of the T-DNA region respectively; the anti-sense and the sense represent the reverse and forward structure of the target gene segment respectively; and the arrow in the figure indicates the direction of the designed primer. C, PCR analysis of ALK-RNAi transgenic rice plants. D, Transcription level expression of ALK Gene in RNAi Line. L-ALKb-RNAi-1, L-ALKb-RNAi-2 and L-ALKb-RNAi-3 are ALKb-RNAi transgenic homozygous lines under the background of Longtefu and Z-ALKc-RNAi-2-1, Z-ALKc-RNAi-2-2 and Z-ALKc-RNAi-2-3 are homozygous transgenic lines carrying both ALKb-RNAi interference structure and ALKc allele. Data were subjected to one-way analysis of variance (ANOVA) depending on the experiment, followed by a comparison of the means according to a Duncan’s multiple range test at P< 0.05 or P < 0.01. Double asterisks denote a highly significant difference between transgenic line and its wild type (P<0.01), single asterisk denotes a significant difference between transgenic line and it’s wild type (0.01≤ P<0.05). (n=3).
正向引物 Forward primer | 引物序列 Primer sequence(5′-3′) | 反向引物 Reverse primer | 引物序列 Primer sequence(5′-3′) |
---|---|---|---|
ALK-5 | GTCCATGGTCGACCTCAAGTAAGAACGGAG | ALK-3 | GTACTAGTCGCCTTTGGCTTC |
4342(GC/TT)-F1 | CAAGGAGAGCTGGAGGGGGC | 4342(GC/TT)-R1 | CATGCCGCGCACCTGGAAA |
4342(GC/TT)-F | TCGGCGGGCTGAGGGACAC | 4342(GC/TT)-R | TCGCATCAATGGACATAACAAACAC |
INT-F | CCTCGTAATCAATTTTAGAT | NOS-R | GACCGCACAGGATTCAAT |
ALK-RT-F | TGATCTGAACGAACCGGACG | ALK-RT-R | TGGGTAAAGCACCTGCAACA |
表1 本研究所用的PCR引物
Table 1 Primers used in this study.
正向引物 Forward primer | 引物序列 Primer sequence(5′-3′) | 反向引物 Reverse primer | 引物序列 Primer sequence(5′-3′) |
---|---|---|---|
ALK-5 | GTCCATGGTCGACCTCAAGTAAGAACGGAG | ALK-3 | GTACTAGTCGCCTTTGGCTTC |
4342(GC/TT)-F1 | CAAGGAGAGCTGGAGGGGGC | 4342(GC/TT)-R1 | CATGCCGCGCACCTGGAAA |
4342(GC/TT)-F | TCGGCGGGCTGAGGGACAC | 4342(GC/TT)-R | TCGCATCAATGGACATAACAAACAC |
INT-F | CCTCGTAATCAATTTTAGAT | NOS-R | GACCGCACAGGATTCAAT |
ALK-RT-F | TGATCTGAACGAACCGGACG | ALK-RT-R | TGGGTAAAGCACCTGCAACA |
图2 ALK-RNAi转基因稻米的糊化温度 A-ALK-RNAi水稻株系精米在1.7% KOH中的碱消值; B-ALK-RNAi 水稻株系米粉的DSC曲线。
Fig 2. Gelatinization properties of ALK-RNAi transgenic rice. A, Alkali spreading value of milled rice in 1.7% KOH; B, DSC curve of rice flours of ALK-RNAi transgenic rice.
转基因系 | 起始温度 | 峰值温度 | 终止温度 | 热焓值 | 直链淀粉含量 | 胶稠度 | 粗蛋白含量 | |
---|---|---|---|---|---|---|---|---|
Transgenic line | To /℃ | Tp /℃ | Tc /℃ | △H /(J∙g-1) | AAC/% | GC/mm | Protein content/% | |
龙特甫B LTFB | 63.6±0.1 | 69.5±0.1 | 77.2±0.3 | 6.30±0.41 | 23.16±1.55 | 19.0±0.0 | 10.47±0.12 | |
L-ALKb-RNAi-1 | 61.8±0.4* | 68.2±0.0** | 76.2±0.5* | 6.40±0.49 | 24.42±0.13 | 18.0±0.0 | 9.80±0.17 | |
L-ALKb-RNAi-2 | 62.2±0.3* | 68.3±0.1** | 76.0±0.5* | 6.30±0.82 | 24.01±0.15 | 18.0±0.1 | 9.73±0.19 | |
L-ALKb-RNAi-3 | 62.2±0.5* | 68.6±0.1** | 76.0±0.0* | 6.00±0.31 | 23.66±0.26 | 17.7±0.1 | 10.11±0.49 | |
珍汕97B ZS97B | 74.6±0.2 | 79.2±0.2 | 85.2±0.1 | 9.43±0.16 | 25.11±0.04 | 58.5±0.4 | 10.61±0.04 | |
Z-ALKc-RNAi-2-1 | 67.1±0.3** | 73.4±0.1** | 80.2±0.1** | 8.20±0.08** | 27.73±0.24** | 25.5±0.1** | 9.21±0.05* | |
Z-ALKc-RNAi-2-2 | 66.3±0.0** | 72.9±0.1** | 79.4±0.2** | 7.99±0.12** | 27.85±0.13** | 28.0±0.3** | 9.65±0.42* | |
Z-ALKc-RNAi-2-3 | 65.6±0.1** | 72.4±0.1** | 79.0±0.0** | 7.29±0.07** | 26.69±0.01** | 45.0±0.0** | 9.03±0.46* |
表2 ALK-RNAi转基因水稻株系米粉的热力学特性和理化特性
Table 2 Gelatinization properties and physical-chemical qualities of ALK-RNAi transgenic rice.
转基因系 | 起始温度 | 峰值温度 | 终止温度 | 热焓值 | 直链淀粉含量 | 胶稠度 | 粗蛋白含量 | |
---|---|---|---|---|---|---|---|---|
Transgenic line | To /℃ | Tp /℃ | Tc /℃ | △H /(J∙g-1) | AAC/% | GC/mm | Protein content/% | |
龙特甫B LTFB | 63.6±0.1 | 69.5±0.1 | 77.2±0.3 | 6.30±0.41 | 23.16±1.55 | 19.0±0.0 | 10.47±0.12 | |
L-ALKb-RNAi-1 | 61.8±0.4* | 68.2±0.0** | 76.2±0.5* | 6.40±0.49 | 24.42±0.13 | 18.0±0.0 | 9.80±0.17 | |
L-ALKb-RNAi-2 | 62.2±0.3* | 68.3±0.1** | 76.0±0.5* | 6.30±0.82 | 24.01±0.15 | 18.0±0.1 | 9.73±0.19 | |
L-ALKb-RNAi-3 | 62.2±0.5* | 68.6±0.1** | 76.0±0.0* | 6.00±0.31 | 23.66±0.26 | 17.7±0.1 | 10.11±0.49 | |
珍汕97B ZS97B | 74.6±0.2 | 79.2±0.2 | 85.2±0.1 | 9.43±0.16 | 25.11±0.04 | 58.5±0.4 | 10.61±0.04 | |
Z-ALKc-RNAi-2-1 | 67.1±0.3** | 73.4±0.1** | 80.2±0.1** | 8.20±0.08** | 27.73±0.24** | 25.5±0.1** | 9.21±0.05* | |
Z-ALKc-RNAi-2-2 | 66.3±0.0** | 72.9±0.1** | 79.4±0.2** | 7.99±0.12** | 27.85±0.13** | 28.0±0.3** | 9.65±0.42* | |
Z-ALKc-RNAi-2-3 | 65.6±0.1** | 72.4±0.1** | 79.0±0.0** | 7.29±0.07** | 26.69±0.01** | 45.0±0.0** | 9.03±0.46* |
转基因系 Transgenic line | 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 /℃ |
---|---|---|---|---|---|---|---|
龙特甫B LTFB | 3699±11 | 3217±18 | 482±7 | 6385±18 | 2686±5 | 6.5 | 87.9 |
L-ALKb-RNAi-1 | 3923±7** | 3390±10** | 533±13** | 6390±11 | 2467±8** | 6.5 | 87.0* |
L-ALKb-RNAi-2 | 3810±13** | 3312±25** | 498±8* | 6055±7** | 2245±6** | 6.5 | 86.9* |
L-ALKb-RNAi-3 | 3843±8** | 3137±17* | 706±10** | 6345±14* | 2502±7** | 6.5 | 87.8* |
珍汕97B ZS97B | 2885±12 | 2510±16 | 375±9 | 4974±15 | 2089±9 | 6.5 | 88.6 |
Z-ALKc-RNAi-2-1 | 3608±9** | 2945±17** | 663±12** | 4749±17** | 1141±11** | 6.5 | 76.5** |
Z-ALKc-RNAi-2-2 | 3930±11** | 3249±13** | 681±11** | 4590±16** | 660±12** | 6.6 | 77.4** |
Z-ALKc-RNAi-2-3 | 3976±6** | 3337±14** | 639±8** | 4956±19 | 980±14** | 6.5 | 76.5** |
表3 ALK-RNAi 转基因水稻株系米粉的RVA谱特征值
Table 3 RVA parameters of mature rice flour in ALK-RNAi transgenic rice lines.
转基因系 Transgenic line | 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 /℃ |
---|---|---|---|---|---|---|---|
龙特甫B LTFB | 3699±11 | 3217±18 | 482±7 | 6385±18 | 2686±5 | 6.5 | 87.9 |
L-ALKb-RNAi-1 | 3923±7** | 3390±10** | 533±13** | 6390±11 | 2467±8** | 6.5 | 87.0* |
L-ALKb-RNAi-2 | 3810±13** | 3312±25** | 498±8* | 6055±7** | 2245±6** | 6.5 | 86.9* |
L-ALKb-RNAi-3 | 3843±8** | 3137±17* | 706±10** | 6345±14* | 2502±7** | 6.5 | 87.8* |
珍汕97B ZS97B | 2885±12 | 2510±16 | 375±9 | 4974±15 | 2089±9 | 6.5 | 88.6 |
Z-ALKc-RNAi-2-1 | 3608±9** | 2945±17** | 663±12** | 4749±17** | 1141±11** | 6.5 | 76.5** |
Z-ALKc-RNAi-2-2 | 3930±11** | 3249±13** | 681±11** | 4590±16** | 660±12** | 6.6 | 77.4** |
Z-ALKc-RNAi-2-3 | 3976±6** | 3337±14** | 639±8** | 4956±19 | 980±14** | 6.5 | 76.5** |
样品 Sample | 转基因系 Transgenic line | 综合 Integrated score | 外观 Appearance | 口感 Taste |
---|---|---|---|---|
热饭 Cooked rice | 龙特甫B LTFB | 53.3±1.3 | 5.0±0.1 | 4.5±0.2 |
L-ALKb-RNAi-1 | 48.3±1.3** | 4.3±0.1** | 3.9±0.2* | |
L-ALKb-RNAi-2 | 51.0±0.8* | 4.7±0.1* | 4.2±0.1* | |
L-ALKb-RNAi-3 | 50.5±1.7** | 4.6±0.2* | 4.1±0.2* | |
珍汕97B ZS97B | 55.0±2.6 | 5.1±0.3 | 4.8±0.3 | |
Z-ALKc-RNAi-2-1 | 43.8±1.9** | 3.6±0.3** | 3.3±0.3** | |
Z-ALKc-RNAi-2-2 | 42.0±0.8** | 3.5±0.0** | 3.1±0.1** | |
Z-ALKc-RNAi-2-3 | 44.5±0.6** | 3.8±0.0** | 3.4±0.0** | |
冷饭 Cooled rice | 龙特甫B LTFB | 49.8±0.5 | 4.6±0.0 | 3.9±0.1 |
L-ALKb-RNAi-1 | 44.8±0.5** | 4.1±0.1** | 3.6±0.1** | |
L-ALKb-RNAi-2 | 48.5±1.0* | 4.3±0.1* | 3.7±0.1* | |
L-ALKb-RNAi-3 | 46.5±0.6** | 4.3±0.1* | 3.7±0.1* | |
珍汕97B ZS97B | 54.0±0.8 | 5.1±0.0 | 4.7±0.1 | |
Z-ALKc-RNAi-2-1 | 45.8±0.5** | 3.0±1.4** | 3.4±0.0** | |
Z-ALKc-RNAi-2-2 | 44.8±1.7** | 3.4±0.2* | 3.1±0.4** | |
Z-ALKc-RNAi-2-3 | 47.8±2.1** | 4.0±0.2 | 3.6±0.3** |
表4 ALK-RNAi转基因水稻株系稻米食味值
Table 4 Sensory evaluation of milled rice in ALK-RNAi transgenic rice lines.
样品 Sample | 转基因系 Transgenic line | 综合 Integrated score | 外观 Appearance | 口感 Taste |
---|---|---|---|---|
热饭 Cooked rice | 龙特甫B LTFB | 53.3±1.3 | 5.0±0.1 | 4.5±0.2 |
L-ALKb-RNAi-1 | 48.3±1.3** | 4.3±0.1** | 3.9±0.2* | |
L-ALKb-RNAi-2 | 51.0±0.8* | 4.7±0.1* | 4.2±0.1* | |
L-ALKb-RNAi-3 | 50.5±1.7** | 4.6±0.2* | 4.1±0.2* | |
珍汕97B ZS97B | 55.0±2.6 | 5.1±0.3 | 4.8±0.3 | |
Z-ALKc-RNAi-2-1 | 43.8±1.9** | 3.6±0.3** | 3.3±0.3** | |
Z-ALKc-RNAi-2-2 | 42.0±0.8** | 3.5±0.0** | 3.1±0.1** | |
Z-ALKc-RNAi-2-3 | 44.5±0.6** | 3.8±0.0** | 3.4±0.0** | |
冷饭 Cooled rice | 龙特甫B LTFB | 49.8±0.5 | 4.6±0.0 | 3.9±0.1 |
L-ALKb-RNAi-1 | 44.8±0.5** | 4.1±0.1** | 3.6±0.1** | |
L-ALKb-RNAi-2 | 48.5±1.0* | 4.3±0.1* | 3.7±0.1* | |
L-ALKb-RNAi-3 | 46.5±0.6** | 4.3±0.1* | 3.7±0.1* | |
珍汕97B ZS97B | 54.0±0.8 | 5.1±0.0 | 4.7±0.1 | |
Z-ALKc-RNAi-2-1 | 45.8±0.5** | 3.0±1.4** | 3.4±0.0** | |
Z-ALKc-RNAi-2-2 | 44.8±1.7** | 3.4±0.2* | 3.1±0.4** | |
Z-ALKc-RNAi-2-3 | 47.8±2.1** | 4.0±0.2 | 3.6±0.3** |
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