干旱胁迫下表观遗传机制对转C4型PEPC基因水稻种子萌发的影响

doi:10.11983/CBB20048

植物学报 ›› 2020, Vol. 55 ›› Issue (6): 677-692.DOI: 10.11983/CBB20048

干旱胁迫下表观遗传机制对转C₄型PEPC基因水稻种子萌发的影响

宋凝曦¹^,², 谢寅峰², 李霞¹^,²^,³^,^*

¹江苏省农业科学院粮食作物研究所/江苏省优质水稻工程技术研究中心/国家水稻改良中心南京分中心, 南京 210014
²南京林业大学生物与环境学院, 南京 210037
³扬州大学农学院, 江苏省粮食作物现代产业技术协同创新中心, 扬州 225009

收稿日期:2020-03-21 接受日期:2020-08-26 出版日期:2020-11-01 发布日期:2020-11-11
通讯作者: 李霞
作者简介:*E-mail: jspplx@jaas.ac.cn
基金资助:
国家自然科学基金(31571585);国家重点研发计划(2016YFD0300501-03)

Effects of Epigenetic Mechanisms on C₄ Phosphoenolpyruvate Carboxylase Transgenic Rice (Oryza sativa) Seed Germination Under Drought Stress

Ningxi Song¹^,², Yingfeng Xie², Xia Li¹^,²^,³^,^*

¹Nanjing Branch of China National Center for Rice Improvement/Jiangsu High Quality Rice Engineering Technology Research Center/Institute of Food Crops, Jiangsu Academy of Agricultural Sciences, Nanjing 210014, China
²College of Biology and Environment, Nanjing Forestry University, Nanjing 210037, China
³Collaborative Innovation Center for Modern Production Technology of Grain Crops of Jiangsu Province, Agricultural College, Yangzhou University, Yangzhou 225009, China

Received:2020-03-21 Accepted:2020-08-26 Online:2020-11-01 Published:2020-11-11
Contact: Xia Li

摘要/Abstract

摘要： 为探究干旱胁迫下表观遗传机制对高表达玉米(Zea mays) C₄型PEPC转基因水稻(Oryza sativa)种子萌发的影响, 以转C₄型PEPC水稻(PC)和野生型水稻Kitaake (WT)为试材, 采用10% (m/v)聚乙二醇6000 (PEG6000)模拟干旱条件, 通过单独和联合施用PEG6000、DNA甲基化抑制剂5-氮杂胞苷(5azaC)和可变剪接抑制剂大环内酯类(PB)进行种子发芽实验, 测定种子活力、萌发过程中可溶性糖和可溶性蛋白含量、α-淀粉酶活性以及PEPC、糖信号相关基因和部分剪接因子基因的表达。结果表明, 0.25 µmol·L^-1PB处理对2种供试水稻在干旱条件下种子萌发均表现出显著抑制作用, 使干旱条件下种子萌发过程中可溶性总糖、蔗糖、葡萄糖和果糖含量以及可溶性蛋白含量均有所下降, PB也抑制糖信号-蔗糖非发酵1 (SNF1)相关蛋白激酶(SnRKs)家族和剪接因子丝氨酸/精氨酸富集蛋白家族(SR proteins)相关基因的表达以及α-淀粉酶的活性, 但对PC的抑制作用小于WT。5 µmol·L^-15azaC处理对干旱条件下种子萌发的效果与可变剪接抑制剂相反。5 µmol·L ^-1 5azaC联合PEG6000干旱处理部分减缓了干旱对水稻种子发芽率的抑制作用, 使供试材料发芽率升高, 表明DNA甲基化和可变剪接机制参与了水稻芽期干旱耐性, 其中对PC的作用更大。

关键词: 可变剪接, DNA甲基化, 干旱逆境, 萌发, PEPC基因, 水稻

Abstract: In order to reveal the effect of epigenetic mechanism under drought stress toward seed germination of transgenic rice within high maize C₄-type PEPC gene expressing, C₄-PEPC transgenic rice (PC) and wild type rice Kitaake (WT) were used in this study. By introducing DNA methylation inhibitor (5-azacytidine, 5azaC) and alternative splicing inhibitors (macrolides pladienolide B, PB), the drought simulation treatments with 10% (m/v) polyethylene glycol-6000 (PEG6000) alone or combining with the inhibitors were used for seed germination experiments. Seed vigor, soluble sugar and soluble protein content, α-amylase activity and the expression levels of related genes, PEPC-related genes, sugar signal-related genes, and some splicing factor genes during germination were measured. In the results, when treated with 0.25 µmol·L^-1, PB had showed a significant inhibitory effect on the seed germination of the two tested rice lines under drought conditions. The content of total soluble sugar, sucrose, glucose, fructose and soluble protein during seed germination after PB addition was reduced to a certain extent under drought conditions. PB treatment also inhibited the gene expression of sucrose nonfermenting-1 (SNF1)-related protein kinase SnRKs family and the splicing factor arginine/serine-rich proteins (SR proteins), and the activity of α-amylase as well, but the inhibitory effect on PC is less than those on WT. 5 µmol·L ^-1 5azaC treatment had an opposite effect with alternative splicing inhibitors. The combination treatment with 5azaC and PEG6000 partially alleviated the inhibitory effect of drought stress on rice seed germination, and the germination rate of the tested materials increased. It can be seen that the mechanism of DNA methylation and alternative splicing are involved in drought tolerance at the bud stage of rice lines, with a larger effect on PC.

Key words: alternative splicing, DNA methylation, drought stress, germination, PEPC gene, rice (Oryza sativa)

宋凝曦, 谢寅峰, 李霞. 干旱胁迫下表观遗传机制对转C₄型PEPC基因水稻种子萌发的影响. 植物学报, 2020, 55(6): 677-692.

Ningxi Song, Yingfeng Xie, Xia Li. Effects of Epigenetic Mechanisms on C₄ Phosphoenolpyruvate Carboxylase Transgenic Rice (Oryza sativa) Seed Germination Under Drought Stress. Chinese Bulletin of Botany, 2020, 55(6): 677-692.

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https://www.chinbullbotany.com/CN/Y2020/V55/I6/677

图/表 10

表1 引物序列

Table 1 Primers used in this study

Gene	Forward primer (5'-3')	Reverse primer (5'-3')
Actin	CCCTCAAACATCGGTATGGA	TTGATCTTCATGCTGCTTGG
OsK1a	AACCAGAGGTAACAGGCAGG	AACCAGAGGTAACAGGCAGG
OsK24	CGTGTTGGCTTCAGTGAAT	CCTTCTCTATCTAAGGGCCG
OsK35	TTGTGTTGGCTTCAGTGAAA	CCTTCGCTGTCTAAGGACTG
C₄-PEPC	CCCACTATCCTTCGCAAGAC	CTAGCCAGTGTTCTGCATGCCGG
Osppc2a	CTGGTTGAGATGGTTTTCGC	GGTGTGAATTCAGGCACTTC
SAPK8	ATAGATGATAATGTCCAGCG	GTTCCTACAGTGGATTTTGG
SAPK9	CACAGCAACGCCGTCTCC	CACACTTCCACCGCTACCAA
SAPK10	TGCTGATGTGTGGTCGTGTG	TGCTGGTATGGTCGCCTCT
SR40	CAATCTGGGGACTGCTTTC	TCCTGCTTGGGCTTTTACT
SR33	ATATTGCCTGCTACCCGAAAG	CAGAGCAGCACCCAGTTTATTAC
OsAmy1A	TTTCGGTCCTCATCGTCCTCC	TCCACGACTCCCAGTTGAATC
OsAmy1C	TGGTATCGATCAGAAACCGGC	GTCCGACCTTCGTGATGACC
OsAmy3C	AAGCATTCCACCACAATGAGC	AGGAAGTTGTACCACCCACC
OsAmy3E	TCACCCTGTGTTGTGTCGTT	AAAGTTGTACCACCCGCCTT

图1 不同处理对转基因水稻(PC)和野生型水稻(WT)种子活力的影响 (A) 发芽势; (B) 发芽率; (C) 发芽指数; (D) 活力指数; (E) 种子发芽图片。不同小写字母表示差异显著(P<0.05)。Bars=0.5 cm

Figure 1 Effects of different treatments on seed vigor of transgenic rice (PC) and wild type rice (WT) (A) Germination; (B) Germination rate; (C) Germination index; (D) Vigor index; (E) Images of germinating seeds at different time. Different lowercase letters indicate significant differences (P<0.05). Bars=0.5 cm

图2 不同处理下转基因水稻(PC)和野生型水稻(WT)萌发种子根长(A)、芽长(B)和根芽比(C)的变化不同小写字母表示差异显著(P<0.05)。

Figure 2 Effects of different treatments on root length (A), shoot length (B) and root/bud ratio (C) of transgenic rice (PC) and wild type rice (WT) germinating seeds Different lowercase letters indicate significant differences (P<0.05).

图3 不同处理对转基因水稻(PC)和野生型水稻(WT)种子萌发过程中可溶性总糖含量(A)和可溶性蛋白含量(B)的影响不同小写字母表示差异显著(P<0.05)。

Figure 3 Effects of different treatments on total soluble sugar content (A) and soluble protein content (B) of transgenic rice (PC) and wild type rice (WT) germinating seeds Different lowercase letters indicate significant differences (P<0.05).

图4 不同处理对转基因水稻(PC)和野生型水稻(WT)种子萌发过程中糖组分含量的影响 (A), (D) 分别为发芽24和48小时的蔗糖含量; (B), (E) 分别为发芽24和48小时的葡萄糖含量; (C), (F) 分别为发芽24和48小时的果糖含量。不同小写字母表示差异显著(P<0.05)。

Figure 4 Effects of different treatments on sugar content of transgenic rice (PC) and wild type rice (WT) germinating seeds (A), (D) Sucrose content at 24 h and 48 h after germination, respectively; (B), (E) Glucose content at 24 h and 48 h after germination, respectively; (C), (F) Fructose content at 24 h and 48 h after germination, respectively. Different lowercase letters indicate significant differences (P<0.05).

图5 不同处理对转基因水稻(PC)和野生型水稻(WT)种子萌发过程中SnRK相关基因表达的影响不同小写字母表示差异显著(P<0.05)。

Figure 5 Effects of different treatments on the expression of SnRK related genes in transgenic rice (PC) and wild type rice (WT) germinating seeds Different lowercase letters indicate significant differences (P<0.05).

图6 不同处理对转基因水稻(PC)和野生型水稻(WT)种子萌发过程中剪接因子相关基因表达的影响不同小写字母表示差异显著(P<0.05)。

Figure 6 Effects of different treatments on expression of splicing factor related genes in transgenic rice (PC) and wild type rice (WT) germinating seeds Different lowercase letters indicate significant differences (P<0.05).

图7 不同处理对转基因水稻(PC)和野生型水稻(WT)种子萌发过程中PEPC相关基因表达的影响不同小写字母表示差异显著(P<0.05)。

Figure 7 Effects of different treatments on expression of PEPC-related genes in transgenic rice (PC) and wild type rice (WT) germinating seeds Different lowercase letters indicate significant differences (P<0.05).

图8 不同处理对转基因水稻(PC)和野生型水稻(WT)种子萌发过程中α-淀粉酶相关基因表达的影响不同小写字母表示差异显著(P<0.05)。

Figure 8 Effects of different treatments on expression of α-amylase related genes in transgenic rice (PC) and type wild rice (WT) germinating seeds Different lowercase letters indicate significant differences (P<0.05).

图9 不同处理对转基因水稻(PC)和野生型水稻(WT)种子萌发过程中α-淀粉酶活性的影响不同小写字母表示差异显著(P<0.05)。

Figure 9 Effects of different treatments on α-amylase activity in transgenic rice (PC) and wild type rice (WT) germinating seeds Different lowercase letters indicate significant differences (P<0.05).

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干旱胁迫下表观遗传机制对转C₄型PEPC基因水稻种子萌发的影响

Effects of Epigenetic Mechanisms on C₄ Phosphoenolpyruvate Carboxylase Transgenic Rice (Oryza sativa) Seed Germination Under Drought Stress

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