中国水稻科学 ›› 2021, Vol. 35 ›› Issue (2): 121-129.DOI: 10.16819/j.1001-7216.2021.0805
潘阳阳, 陈宜波, 王重荣, 李宏, 黄道强, 周德贵, 王志东, 赵雷, 龚蓉, 周少川*()
收稿日期:
2020-08-05
修回日期:
2020-09-16
出版日期:
2021-03-10
发布日期:
2021-03-10
通讯作者:
周少川
Yangyang PAN, Yibo CHEN, Chongrong WANG, Hong LI, Daoqiang HUANG, Degui ZHOU, Zhidong WANG, Lei ZHAO, Rong GONG, Shaochuan ZHOU*()
Received:
2020-08-05
Revised:
2020-09-16
Online:
2021-03-10
Published:
2021-03-10
Contact:
Shaochuan ZHOU
摘要:
【目的】结合代谢组和转录组技术,分析香稻和非香稻不同发育时期籽粒中γ-氨基丁酸(GABA)和2-乙酰-1-吡咯啉(2AP)代谢通路主要化合物和基因的动态变化,为高含量GABA和2AP的水稻育种提供理论依据。【方法】本研究收集优质香籼稻美香占2号和优质籼稻黄华占花后8 d(乳熟期)、花后15 d(蜡熟期)、30 d(完熟期)的脱壳籽粒和花后40 d收获的籽粒的精米,对各样本采用LC-MS/MS和RNA-seq技术进行代谢产物检测和转录表达分析,分析两个水稻品种GABA和2AP代谢通路相关化合物的空间分布以及含量变化特征。【结果】代谢组数据PCA分析表明两个水稻品种乳熟期籽粒和花后40 d收获的籽粒的精米明显分离,蜡熟期和完熟期籽粒分离不明显。代谢组共鉴定出623种代谢物,四个时期差异代谢物合计161个。GABA和2AP通路中9个代谢物被检出,两个水稻品种的相关代谢物变化趋势相似,其中腐胺主要定位于籽粒的糊粉层,而亚精胺在内胚乳特异累积。转录组共检出3.3万个左右基因,其中检出GABA通路18个酶中的14个酶对应的基因有较高的表达水平,除Badh2外其他基因均不是差异表达基因,同一基因在两份水稻材料中的表达水平和变化趋势相似,GAD1、GABA-T1、DAO4和PAO4在相应同源基因中表达水平较高。黄华占籽粒中L-精氨酸和腐胺的代谢在蜡熟期已基本结束,美香占2号在籽粒整个发育期具有持续较强的多胺降解水平。美香占2号籽粒具有合成2AP的物质基础,多胺降解途径和谷氨酸-脯氨酸转化通路同时有助于2AP的积累。【结论】水稻籽粒GABA代谢以降解为主,GABA-T1和GABA-T2共同调控GABA的降解。GABA相关代谢物主要集中在糊粉层,黄华占籽粒中代谢物分布和基因表达特征与美香占2号相似,美香占2号籽粒成熟过程中具有持续较高的线粒体活性。
潘阳阳, 陈宜波, 王重荣, 李宏, 黄道强, 周德贵, 王志东, 赵雷, 龚蓉, 周少川. γ-氨基丁酸和2-乙酰-1-吡咯啉代谢通路在水稻籽粒发育过程中的变化分析[J]. 中国水稻科学, 2021, 35(2): 121-129.
Yangyang PAN, Yibo CHEN, Chongrong WANG, Hong LI, Daoqiang HUANG, Degui ZHOU, Zhidong WANG, Lei ZHAO, Rong GONG, Shaochuan ZHOU. Metabolism of γ-aminobutyrate and 2-acetyl-1-pyrroline Analyses at Various Grain Developmental Stages in Rice (Oryza sativa L.)[J]. Chinese Journal OF Rice Science, 2021, 35(2): 121-129.
图1 GABA和2AP代谢通路 ADC—精氨酸脱羧酶;AgmAH—鲱精胺酶;ARG—精氨酸酶;BADH2—甜菜碱醛脱氢酶2;DAO—二胺氧化酶;GABA-T—氨基丁酸转氨酶;GAD—谷氨酸脱羧酶;GDH—谷氨酸脱氢酶;OAT—鸟氨酸转氨酶;ODC—鸟氨酸脱羧酶;P5CDH—吡咯啉-5-羧酸脱氢酶;P5CR—吡咯啉-5-羧酸还原酶;P5CS—吡咯啉-5-羧酸合成酶;PAO—多胺氧化酶;ProDH—脯氨酸脱氢酶;SpdS—亚精胺合成酶;SpmS—精胺合成酶;SSADH—琥珀酸半醛脱氢酶。
Fig. 1. Interconnectivity of GABA and 2AP metabolism. ADC, Arginine decarboxylase; AgmAH, Agmatine amidinohydrolase; ARG, Arginase; BADH2, Betaine aldehyde dehydrogenase 2; DAO, Diamine oxidase; GABA-T, GABA transaminase; GAD, Glutamate decarboxylase; GDH, Glutamate dehydrogenase; OAT, Ornithine aminotransferase; ODC, Ornithine decarboxylase; P5CDH, Pyrroline-5-carboxylate dehydrogenase; P5CR, Pyrrolidine -5- carboxylate reductase; P5CS, Pyrrolidine -5- carboxylate synthetase; PAO, Polyamine oxidase; ProDH, Proline dehydrogenase; SpdS, Spermidine synthase; SpmS, Spermine synthase; SSADH, Succinic semialdehyde dehydrogenase.
图2 黄华占(HHZ)和美香占2号(MXZ)四个时期籽粒代谢物的主成分分析 PC1:主成分1解释总变异的42.9%;PC2:主成分2解释总变异的21.9%。8D, 15D,30D和40D分别代表花后8 d(乳熟期)、15 d(蜡熟期)、30 d(完熟期,糙米)脱壳籽粒,花后40 d籽粒碾成的精米。
Fig. 2. PCA of metabolites at four grain developmental stages of Huanghuazhan(HHZ) and Meixiangzhan 2(MXZ). The principal component 1 accounted for 42.9% of the total variation, and the component 2 accounted for 21.9% of the total variation by PCA (principal components analyses). 8D, 15D, 30D and 40D, indicated milky grains(8 days after flowering), dough grains(15 days after flowering), mature grains(30 days after flowering) and milled rice(40 days after flowering), respectively.
图4 黄华占(HHZ)和美香占2号(MXZ)籽粒四个发育时期GABA通路相关代谢物的含量*表示差异代谢物。
Fig. 4. Contents of metabolites involved in GABA metabolism in Huanghuazhan(HHZ) and Meixiangzhan 2(MXZ) grains. * Represents differential metabolite.
基因名称 Gene name | 基因缩写 Gene symbol | 基因登录号 Locus ID | HHZ8D | HHZ15D | HHZ30D | MXZ8D | MXZ15D | MXZ30D |
---|---|---|---|---|---|---|---|---|
谷氨酸脱羧酶 | GAD1 | LOC_Os08g36320 | 10.65±0.67 | 64.70 ±9.21 | 60.76±3.22 | 10.41±0.96 | 48.13±6.90 | 70.77±18.90 |
Glutamate decarboxylase | GAD2 | LOC_Os04g37500 | 1.92±0.20 | 7.98±1.07 | 6.11±0.67 | 1.30±0.38 | 5.86±1.68 | 7.06±2.38 |
GAD3 | LOC_Os03g13300 | 0.29±0.11 | 0.71±0.04 | 0.55±0.15 | 0.16±0.03 | 0.98±0.92 | 0.81±0.22 | |
GAD4 | LOC_Os03g51080 | 3.88±0.41 | 5.55±1.07 | 5.06±0.74 | 5.43±1.03 | 3.77±0.05 | 4.50±0.36 | |
GAD5 | LOC_Os04g37460 | 0.46±0.28 | 0.12±0.07 | 0.16±0.13 | 0.32±0.16 | 0.25±0.03 | 0.37±0.30 | |
γ-氨基丁酸转氨酶 | GABA-T1 | LOC_Os04g52450 | 44.71±2.12 | 79.40±4.99 | 79.46±1.66 | 31.79±3.80 | 65.98±6.65 | 69.10±20.38 |
γ-aminobutyrate | GABA-T2 | LOC_Os08g10510 | 28.11±0.90 | 9.01±1.69* | 7.83±1.83 | 25.10±2.85 | 21.29±2.84 | 12.73±1.14 |
transaminase | GABA-T3 | LOC_Os04g52440 | 0.83±0.20 | 1.42±0.77 | 0.41±0.10 | 1.47±0.34 | 1.11±0.48 | 0.59±0.07 |
GABA-T4 | LOC_Os02g02210 | N/A | N/A | N/A | N/A | N/A | N/A | |
谷氨酸脱氢酶 | GDH1 | LOC_Os03g58040 | 17.13±0.80 | 6.87±0.35 | 6.22±0.84 | 13.65±0.85 | 8.15±1.12 | 9.07±0.94 |
Glutamate dehydrogenase | GDH2 | LOC_Os04g45970 | 58.97±2.54 | 17.06±2.58 | 20.12±1.04 | 43.85±1.93 | 19.51±5.50 | 13.24±1.93 |
琥珀酸半醛脱氢酶 Succinic semialdehyde dehydrogenase | SSADH | LOC_Os02g07760 | 92.65±8.12 | 105.31±16.3 | 100.52±3.45 | 96.75±9.28 | 102.63±6.72 | 167.60±46.58 |
甜菜碱醛脱氢酶2 Betaine aldehyde dehydrogenase 2 | BADH2 | LOC_Os08g32870 | 77.12±3.91* | 64.64±3.54* | 62.18±4.62* | 10.32±0.84 | 9.21±2.19 | 8.41±1.17 |
二胺氧化酶 | DAO1 | LOC_Os06g23114 | 0.22±0.10 | 0.17±0.05 | 0.26±0.09 | 0.11±0.04 | 0.10±0.04 | 0.12±0.07 |
Diamine oxidase | DAO2 | LOC_Os04g20164 | 3.99±0.75 | 27.83±4.11 | 31.22±0.37 | 2.21±0.38 | 19.15±0.56 | 23.76±3.57 |
DAO3 | LOC_Os04g04950 | N/A | N/A | N/A | N/A | N/A | N/A | |
DAO4 | LOC_Os04g40040 | 71.02±9.02 | 159.52±13.5 | 200.30±5.59 | 57.67±6.26 | 116.76±13.03 | 124.73±13.74 | |
DAO5 | LOC_Os02g37990 | 0.00 | N/A | 0.00 | 0.21±0.12 | N/A | 0.20±0.26 | |
多胺氧化酶 | PAO1 | LOC_Os01g51320 | 1.35±0.26 | 0.22±0.01 | 0.20±0.16 | 1.03±0.38 | 0.24±0.08 | 0.66±0.43 |
Polyamine oxidase | PAO2 | LOC_Os03g09810 | 0.35±0.06 | 0.08±0.07 | 0.34±0.24 | 0.36±0.18 | 0.26±0.14 | 0.32±0.20 |
PAO3 | LOC_Os04g53190 | 11.90±1.27 | 28.45±1.63 | 26.83±1.56 | 11.26±0.69 | 22.94±0.81 | 19.37±4.34 | |
PAO4 | LOC_Os04g57550 | 60.57±3.81 | 96.12±6.23 | 106.53±6.95 | 55.90±2.42 | 55.71±3.12 | 89.17±6.59 | |
PAO5 | LOC_Os04g57560 | 29.95±0.50 | 24.23±1.45 | 26.50±1.55 | 28.07±0.50 | 22.78±1.35 | 25.90±1.94 | |
亚精胺合成酶 | SpdS1 | LOC_Os07g22600 | 65.33±7.15 | 57.42±4.88 | 41.89±3.78 | 77.27±11.83 | 44.77±3.59 | 45.24±1.23 |
Spermidine synthase | SpdS2 | LOC_Os06g33710 | 14.53±2.97 | 37.87±4.49 | 29.55±3.53 | 11.78±0.65 | 25.93±2.67 | 35.42±11.54 |
精胺合成酶 Spermine synthase | SpmS2 | LOC_Os02g15550 | 31.20±1.47 | 32.68±1.65 | 30.30±1.81 | 32.98±2.50 | 32.87±2.56 | 33.11±2.66 |
精氨酸酶 Arginase | ARG | LOC_Os04g01590 | 96.36±4.27 | 78.83±3.01 | 80.27±3.07 | 102.45±3.68 | 85.64±4.11 | 76.70±6.07 |
鸟氨酸脱羧酶 Ornithine decarboxylase | ODC | LOC_Os02g28110 | N/A | N/A | N/A | N/A | N/A | N/A |
脯氨酸脱氢酶 Proline dehydrogenase | ProDH | LOC_Os10g40360 | 23.89±5.83 | 66.88±7.54 | 78.81±7.47* | 15.76±4.19 | 49.54±8.94 | 25.74±4.81 |
1-吡咯啉-5-羧酸合成酶 1-pyrrolidine-5-carboxylate synthetase | P5CS1 | LOC_Os05g38150 | 43.21±2.08 | 31.28±1.56 | 25.46±2.29 | 41.61±2.96 | 42.14±2.06 | 39.42±3.01 |
1-吡咯啉-5-羧酸还原酶 1-pyrrolidine-5-carboxylic reductase | P5CR | LOC_Os01g71990 | 9.00±0.43 | 3.72±0.61 | 3.59±0.74 | 10.00±1.01 | 3.16±0.36 | 6.13±1.36 |
1-吡咯啉-5-羧酸脱氢酶 1-pyrrolidine-5-carboxylic dehydrogenase | P5CDH | Os05g0536400 | N/A | N/A | N/A | N/A | N/A | N/A |
鸟氨酸转氨酶 Ornithine aminotransferase | OAT | LOC_Os03g44150 | 52.54±0.78 | 32.10±7.61 | 25.48±4.34 | 57.42±4.49 | 33.33±1.53 | 38.72±6.02 |
精氨酸脱羧酶 Arginine decarboxylase | ADC1 | LOC_Os06g04070 | 13.55±1.72 | 15.43±1.21 | 18.36±1.78 | 15.41±3.11 | 14.40±0.84 | 19.42±6.76 |
鲱精胺酶 Agmatine amidinohydrolase | AgmAH | LOC_Os09g37200 | 1.78±0.11 | 5.52±1.75 | 5.33±0.98 | 3.17±0.32 | 8.65±0.21 | 3.78±0.15 |
表1 黄华占和美香占2号籽粒三个发育时期GABA通路相关基因表达量
Table 1 Gene expression levels involved in GABA metabolism in HHZ and MXZ rice grains.
基因名称 Gene name | 基因缩写 Gene symbol | 基因登录号 Locus ID | HHZ8D | HHZ15D | HHZ30D | MXZ8D | MXZ15D | MXZ30D |
---|---|---|---|---|---|---|---|---|
谷氨酸脱羧酶 | GAD1 | LOC_Os08g36320 | 10.65±0.67 | 64.70 ±9.21 | 60.76±3.22 | 10.41±0.96 | 48.13±6.90 | 70.77±18.90 |
Glutamate decarboxylase | GAD2 | LOC_Os04g37500 | 1.92±0.20 | 7.98±1.07 | 6.11±0.67 | 1.30±0.38 | 5.86±1.68 | 7.06±2.38 |
GAD3 | LOC_Os03g13300 | 0.29±0.11 | 0.71±0.04 | 0.55±0.15 | 0.16±0.03 | 0.98±0.92 | 0.81±0.22 | |
GAD4 | LOC_Os03g51080 | 3.88±0.41 | 5.55±1.07 | 5.06±0.74 | 5.43±1.03 | 3.77±0.05 | 4.50±0.36 | |
GAD5 | LOC_Os04g37460 | 0.46±0.28 | 0.12±0.07 | 0.16±0.13 | 0.32±0.16 | 0.25±0.03 | 0.37±0.30 | |
γ-氨基丁酸转氨酶 | GABA-T1 | LOC_Os04g52450 | 44.71±2.12 | 79.40±4.99 | 79.46±1.66 | 31.79±3.80 | 65.98±6.65 | 69.10±20.38 |
γ-aminobutyrate | GABA-T2 | LOC_Os08g10510 | 28.11±0.90 | 9.01±1.69* | 7.83±1.83 | 25.10±2.85 | 21.29±2.84 | 12.73±1.14 |
transaminase | GABA-T3 | LOC_Os04g52440 | 0.83±0.20 | 1.42±0.77 | 0.41±0.10 | 1.47±0.34 | 1.11±0.48 | 0.59±0.07 |
GABA-T4 | LOC_Os02g02210 | N/A | N/A | N/A | N/A | N/A | N/A | |
谷氨酸脱氢酶 | GDH1 | LOC_Os03g58040 | 17.13±0.80 | 6.87±0.35 | 6.22±0.84 | 13.65±0.85 | 8.15±1.12 | 9.07±0.94 |
Glutamate dehydrogenase | GDH2 | LOC_Os04g45970 | 58.97±2.54 | 17.06±2.58 | 20.12±1.04 | 43.85±1.93 | 19.51±5.50 | 13.24±1.93 |
琥珀酸半醛脱氢酶 Succinic semialdehyde dehydrogenase | SSADH | LOC_Os02g07760 | 92.65±8.12 | 105.31±16.3 | 100.52±3.45 | 96.75±9.28 | 102.63±6.72 | 167.60±46.58 |
甜菜碱醛脱氢酶2 Betaine aldehyde dehydrogenase 2 | BADH2 | LOC_Os08g32870 | 77.12±3.91* | 64.64±3.54* | 62.18±4.62* | 10.32±0.84 | 9.21±2.19 | 8.41±1.17 |
二胺氧化酶 | DAO1 | LOC_Os06g23114 | 0.22±0.10 | 0.17±0.05 | 0.26±0.09 | 0.11±0.04 | 0.10±0.04 | 0.12±0.07 |
Diamine oxidase | DAO2 | LOC_Os04g20164 | 3.99±0.75 | 27.83±4.11 | 31.22±0.37 | 2.21±0.38 | 19.15±0.56 | 23.76±3.57 |
DAO3 | LOC_Os04g04950 | N/A | N/A | N/A | N/A | N/A | N/A | |
DAO4 | LOC_Os04g40040 | 71.02±9.02 | 159.52±13.5 | 200.30±5.59 | 57.67±6.26 | 116.76±13.03 | 124.73±13.74 | |
DAO5 | LOC_Os02g37990 | 0.00 | N/A | 0.00 | 0.21±0.12 | N/A | 0.20±0.26 | |
多胺氧化酶 | PAO1 | LOC_Os01g51320 | 1.35±0.26 | 0.22±0.01 | 0.20±0.16 | 1.03±0.38 | 0.24±0.08 | 0.66±0.43 |
Polyamine oxidase | PAO2 | LOC_Os03g09810 | 0.35±0.06 | 0.08±0.07 | 0.34±0.24 | 0.36±0.18 | 0.26±0.14 | 0.32±0.20 |
PAO3 | LOC_Os04g53190 | 11.90±1.27 | 28.45±1.63 | 26.83±1.56 | 11.26±0.69 | 22.94±0.81 | 19.37±4.34 | |
PAO4 | LOC_Os04g57550 | 60.57±3.81 | 96.12±6.23 | 106.53±6.95 | 55.90±2.42 | 55.71±3.12 | 89.17±6.59 | |
PAO5 | LOC_Os04g57560 | 29.95±0.50 | 24.23±1.45 | 26.50±1.55 | 28.07±0.50 | 22.78±1.35 | 25.90±1.94 | |
亚精胺合成酶 | SpdS1 | LOC_Os07g22600 | 65.33±7.15 | 57.42±4.88 | 41.89±3.78 | 77.27±11.83 | 44.77±3.59 | 45.24±1.23 |
Spermidine synthase | SpdS2 | LOC_Os06g33710 | 14.53±2.97 | 37.87±4.49 | 29.55±3.53 | 11.78±0.65 | 25.93±2.67 | 35.42±11.54 |
精胺合成酶 Spermine synthase | SpmS2 | LOC_Os02g15550 | 31.20±1.47 | 32.68±1.65 | 30.30±1.81 | 32.98±2.50 | 32.87±2.56 | 33.11±2.66 |
精氨酸酶 Arginase | ARG | LOC_Os04g01590 | 96.36±4.27 | 78.83±3.01 | 80.27±3.07 | 102.45±3.68 | 85.64±4.11 | 76.70±6.07 |
鸟氨酸脱羧酶 Ornithine decarboxylase | ODC | LOC_Os02g28110 | N/A | N/A | N/A | N/A | N/A | N/A |
脯氨酸脱氢酶 Proline dehydrogenase | ProDH | LOC_Os10g40360 | 23.89±5.83 | 66.88±7.54 | 78.81±7.47* | 15.76±4.19 | 49.54±8.94 | 25.74±4.81 |
1-吡咯啉-5-羧酸合成酶 1-pyrrolidine-5-carboxylate synthetase | P5CS1 | LOC_Os05g38150 | 43.21±2.08 | 31.28±1.56 | 25.46±2.29 | 41.61±2.96 | 42.14±2.06 | 39.42±3.01 |
1-吡咯啉-5-羧酸还原酶 1-pyrrolidine-5-carboxylic reductase | P5CR | LOC_Os01g71990 | 9.00±0.43 | 3.72±0.61 | 3.59±0.74 | 10.00±1.01 | 3.16±0.36 | 6.13±1.36 |
1-吡咯啉-5-羧酸脱氢酶 1-pyrrolidine-5-carboxylic dehydrogenase | P5CDH | Os05g0536400 | N/A | N/A | N/A | N/A | N/A | N/A |
鸟氨酸转氨酶 Ornithine aminotransferase | OAT | LOC_Os03g44150 | 52.54±0.78 | 32.10±7.61 | 25.48±4.34 | 57.42±4.49 | 33.33±1.53 | 38.72±6.02 |
精氨酸脱羧酶 Arginine decarboxylase | ADC1 | LOC_Os06g04070 | 13.55±1.72 | 15.43±1.21 | 18.36±1.78 | 15.41±3.11 | 14.40±0.84 | 19.42±6.76 |
鲱精胺酶 Agmatine amidinohydrolase | AgmAH | LOC_Os09g37200 | 1.78±0.11 | 5.52±1.75 | 5.33±0.98 | 3.17±0.32 | 8.65±0.21 | 3.78±0.15 |
图5 美香占2号水稻籽粒发育过程中6个基因表达水平动态变化*表示差异表达基因
Fig. 5. Dynamic changes of six genes expression levels at 3 developmental stages in MXZ. * Represents differentially expressed gene
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