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作物学报 ›› 2020, Vol. 46 ›› Issue (5): 700-711.doi: 10.3724/SP.J.1006.2020.94107

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

谷子PEPC基因的鉴定及其对非生物逆境的响应特性

赵晋锋,杜艳伟,王高鸿,李颜方,赵根有,王振华,王玉文,余爱丽()   

  1. 山西省农业科学院谷子研究所 / 特色杂粮种质资源发掘与育种山西省重点实验室, 山西长治 046011
  • 收稿日期:2019-07-24 接受日期:2020-01-15 出版日期:2020-05-12 网络出版日期:2020-02-14
  • 通讯作者: 余爱丽
  • 作者简介:赵晋锋, E-mail:zhaojfmail@126.com, Tel: 0355-2204195.
  • 基金资助:
    本研究由山西省农业科学院项目(YGJPY2009);本研究由山西省农业科学院项目(YGG17021);本研究由山西省农业科学院项目(YCX2019T05);国家现代农业产业技术体系建设专项(CARS-06-13.5-A23);国家农业环境数据中心观测检测任务项目资助(ZX03S0410)

Identification of PEPC genes from foxtail millet and its response to abiotic stress

Jin-Feng ZHAO,Yan-Wei DU,Gao-Hong WANG,Yan-Fang LI,Gen-You ZHAO,Zhen-Hua WANG,Yu-Wen WANG,Ai-Li YU()   

  1. Millet Research Institute, Shanxi Academy of Agricultural Sciences / Shanxi Key Laboratory of Genetic Resources and Breeding in Minor Crops, Changzhi 046011, Shanxi, China
  • Received:2019-07-24 Accepted:2020-01-15 Published:2020-05-12 Published online:2020-02-14
  • Contact: Ai-Li YU
  • Supported by:
    This study was supported by the Project Plan of Shanxi Academy of Agricultural Sciences(YGJPY2009);This study was supported by the Project Plan of Shanxi Academy of Agricultural Sciences(YGG17021);This study was supported by the Project Plan of Shanxi Academy of Agricultural Sciences(YCX2019T05);the China Agricultural Research System(CARS-06-13.5-A23);the National Agricultural Environmental Data Center Observation and Detection Mission(ZX03S0410)

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摘要：

磷酸烯醇式丙酮酸羧化酶(phosphoenolpyruvate carboxylase, PEPC)是C4植物光合作用关键酶, 并在植物多种代谢途径及逆境信号应答过程中起重要作用。本研究通过序列比对, 从谷子基因组中筛选出6个SiPEPC候选基因。SiPEPC蛋白特征参数相似度很高, 序列非常保守, 都含有PEPC基因特征功能域PEPcase Motif。SiPEPC成员主要被定位在细胞质、细胞核和线粒体。在SiPEPC成员启动子序列中含大量有光、激素、逆境以及其他生长调控相关的顺式应答元件。苗期逆境qRT-PCR表达谱分析表明, 5个SiPEPC基因(SiPEPC1、SiPEPC2、SiPEPC3、SiPEPC5、SiPEPC6)不同程度受ABA、PEG、高盐和低温诱导表达, 表明其参与了苗期对非生物逆境的响应。5个SiPEPC基因表达量在正常生长条件下随着谷子的生长呈增强趋势, 且在不同生育时期干旱胁迫下明显增加, 表明其参与了拔节、抽穗、灌浆期的干旱胁迫应答。拔节期弱光可诱导5个SiPEPC基因的表达, 而在拔节期中等强度光照以及抽穗期和灌浆期的中等光照和弱光照下表达量均急剧降低, 揭示光照强度严重影响SiPEPC基因的表达。

关键词: 谷子, 磷酸烯醇式丙酮酸羧化酶, 非生物逆境, 表达分析

Abstract:

Phosphoenolpyruvate carboxylase (PEPC) is a key enzyme in photosynthesis of C4 plants and plays an important role in a variety of metabolic and stress pathways. In this study, we identified six candidate PEPC genes from foxtail millet genome via sequence alignment. The characteristic parameters of all SiPEPC protein were very similar and the sequences were very conservative. All SiPEPC genes contained the PEPcase motif, which is the characteristic domain of PEPC gene. SiPEPCs were localized in cytoplasm, nucleus and mitochondrion. Promoter analysis identified a variety of light, hormonal, stress, and other growth-related cis-elements in the promoter sequences of SiPEPC members. The qRT-PCR expression profiles showed that the five SiPEPC genes (SiPEPC1, SiPEPC2, SiPEPC3, SiPEPC5, SiPEPC6) were induced by ABA, PEG, high salt and low temperature at seedling stage, indicating that five SiPEPC genes ate involved in abiotic signaling pathway at the seedling stage. The expression of five SiPEPC genes increased with the growth of foxtail millet under normal growth conditions, and increased significantly under drought stress at different growth stages, indicating that five SiPEPCs are involved in drought stress response at jointing, heading and filling stages. The weak light at jointing stage could induce the expression of five SiPEPC genes, while the expression level decreased sharply under moderate light intensity at jointing stage, moderate and weak light intensity at heading and filling stages, showing that light intensity seriously affects the expression of SiPEPC genes.

Key words: foxtail millet, phosphoenolpyruvate carboxylase, abiotic stress, expression analysis

表1

本试验所用引物"

基因
Gene		 上游引物
Forward primer (5'-3')		 下游引物
Reverse primer (5'-3')
SiPEPC1 TGCGTGCTGGAATGAGTTACT CCAATAAGCATACATCCCGTG
SiPEPC2 CAAGAGCCAAACTATTGACCTG TCATACCAGCACGCATTTCA
SiPEPC3 ATGGAAGGGTGTCCCAAAGTT CCAAGAAGAGAACTGAATGAGAGG
SiPEPC5 GCTACTTCGACGACACCATC AGGAGGAGAACTGGATGAGC
SiPEPC6 GAGGTGGAACTGTGGGAAGA GTAAAACGTTGCAGGGTTCTAA
β-Actin CAGTGGACGCACAACAGGTAT AGCAAGGTCAAGACGGAGAAT

表2

预测谷子PEPC基因参数"

基因
Gene		 基因ID
Gene ID		 染色体位置
Chr. location		 基因长度
Genome
length (bp)		 氨基酸数目
No. of amino acid		 等电点
pI		 分子量
Molecular weight (kD)		 内含子数
Intron number		 不稳定指数
Instability index		 脂肪系数
Aliphatic index		 平均疏水指数
Grand average of hydropathicity		 PEPcse保守域位置
PEPcse domain location
SiPEPC1 Seita.1G020700 1784608-1791259 6652 965 5.83 109.82 9 48.48 87.36 -0.392 163-965
SiPEPC2 Seita.2G173700 26198697-26204510 5814 967 5.77 110.20 9 44.74 92.41 -0.351 164-967
SiPEPC3 Seita.4G175200 28034663-28043265 8603 964 5.95 109.98 8 45.83 85.81 -0.418 162-964
SiPEPC4 Seita.5G074200 6390259-6398514 8256 1032 7.14 114.89 20 52.09 93.87 -0.277 144-348, 414-1032
SiPEPC5 Seita.5G147000 13061164-13065839 4676 1015 5.89 113.53 6 49.05 87.69 -0.319 213-1015
SiPEPC6 Seita.5G324500 37380598-37387338 6741 969 5.70 110.44 9 44.10 89.79 -0.375 165-969

图1

SiPEPC成员的基因结构"

表3

SiPEPC蛋白亚细胞定位预测"

基因
Gene		 预测位置
Localization		 可信度
Reliability
SiPEPC1 Cytoplasmic; nuclear; mitochondrial 3.395; 0.685; 0.414
SiPEPC2 Cytoplasmic; nuclear; mitochondrial 3.398; 0.831; 0.317
SiPEPC3 Cytoplasmic; nuclear; mitochondrial 3.679; 0.620; 0.321
SiPEPC4 Cytoplasmic; mitochondrial; nuclear 1.838; 1.290; 1.230
SiPEPC5 Cytoplasmic; mitochondrial; nuclear 2.258; 1.092; 0.472
SiPEPC6 Cytoplasmic; nuclear; mitochondrial 3.428; 0.758; 0.338

表4

SiPEPC基因启动子区域顺式元件预测"

顺式元件
cis-element		 典型序列
Typical sequence		 特性
Characteristic		 基因
Gene
Box 4 ATTAAT Light responsive element SiPEPC1, SiPEPC2, SiPEPC3
G-Box CACGTT Light responsive element SiPEPC1, SiPEPC2, SiPEPC3, SiPEPC4, SiPEPC5, SiPEPC6
GATA-motif AAGGATAAGG Light responsive element SiPEPC1, SiPEPC6
AE-box AGAAACAA Light responsive element SiPEPC2, SiPEPC4, SiPEPC6
GT1-motif GGTTAAT Light responsive element SiPEPC2, SiPEPC4, SiPEPC5
ACE GACACGTATG Light responsive element SiPEPC3, SiPEPC5
Sp1 GGGCGG Light responsive element SiPEPC3, SiPEPC4
ABRE ACGTG Abscisic acid responsiveness SiPEPC1, SiPEPC2, SiPEPC3, SiPEPC4, SiPEPC5
P-box CCTTTTG Gibberellin-responsive SiPEPC1, SiPEPC2
TATC-box TATCCCA Gibberellin-responsiveness SiPEPC2, SiPEPC4
GARE-motif TCTGTTG Gibberellin-responsive element SiPEPC6
TCA-element CCATCTTTTT Salicylic acid responsiveness SiPEPC1, SiPEPC2, SiPEPC5, SiPEPC6
ERE ATTTCATA Ethylene-responsive element SiPEPC2, SiPEPC4
CGTCA-motif CGTCA MeJA-responsiveness SiPEPC3, SiPEPC6
TGACG-motif TGACG MeJA-responsiveness SiPEPC3, SiPEPC6
TGA-element AACGAC Auxin-responsive element SiPEPC1, SiPEPC4
TC-rich repeats GTTTTCTTAC Defense and stress responsiveness SiPEPC1, SiPEPC3
LTR CCGAAA Low-temperature responsiveness SiPEPC2, SiPEPC3, SiPEPC4, SiPEPC6
WUN-motif TTATTACAT Wound-responsive element SiPEPC3
MBS CAACTG MYB binding site involved in drought-inducibility SiPEPC5
ARE AAACCA Anaerobic induction SiPEPC1, 3, 4, 6
CCGTCC-box CCGTCC Meristem specific activation SiPEPC1, 2, 4, 6
GC-motif CCCCCG Anoxic specific inducibility SiPEPC1, 2, 6
O2-site GATGACATGG Zein metabolism regulation SiPEPC2, 4, 5
RY-element CATGCATG Seed-specific regulation SiPEPC3
circadian CAAAGATATC Circadian control SiPEPC5
CAT-box GCCACT Meristem expression SiPEPC6

图2

PEPC家族蛋白序列的氨基酸序列多重比对 AtPEPC1、AtPEPC2、AtPEPC3: 拟南芥AtPEPC1、AtPEPC2、AtPEPC3; SbPEPC1、SbPEPC2、SbPEPC3: 高粱SbPEPC1、SbPEPC2、SbPEPC3; GmPEPC: 大豆GmPEPC; ZmPEPC1、ZmPEPC2: 玉米ZmPEPC1、ZmPEPC2; StPEPC: 马铃薯StPEPC; AhPEPC: 花生AhPEPC; RcPEPC: 蓖麻RcPEPC; NtPEPC: 烟草 NtPEPC。对应GenBank蛋白序列号分别为Q9MAH0.1、Q5GM68.2、Q84VW9.2、P29195.1、P29194.1、P15804.2、P51061.1、P04711.2、P51059.1、P29196.2、ABY87944.1、ABR29878.1、P27154.1。相同氨基酸残基用黑色表示, 相似氨基酸残基用灰色表示(相似性 ≥ 60%)。"

图3

不同物种PEPC 蛋白的进化关系"

图4

SiPEPC基因苗期逆境表达谱 图中浅绿、深绿、黑色、浅红、深红5种颜色代表基因表达水平。绿色表示基因表达弱, 红色表示基因表达强。"

图5

正常和干旱条件下SiPEPC基因在不同发育阶段的相对表达量 *表示在0.05水平上显著; **表示在0.01水平上显著。"

图6

光照和干旱条件下SiPEPC 的相对表达量 图中浅绿、深绿、黑色、浅红、深红五色代表基因表达水平。绿色表示基因表达弱, 红色表示基因表达强。J-1: 拔节期对照; J-2: 拔节期干旱; J-3: 拔节期中等光照; J-4: 拔节期弱光照; H-1: 抽穗期对照; H-2: 抽穗期干旱; H-3: 抽穗期中等光照; H-4: 抽穗期弱光照; F-1: 灌浆期对照; F-2: 灌浆期干旱; F-3: 灌浆期中等光照; F-4: 灌浆期弱光照。"

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