玉米种胚HSP20基因对人工老化处理的响应

doi:10.3724/SP.J.1006.2018.01733

Abstract

Abstract:

Maize (Zea mays L.) cultivar ‘Zhengdan 958’ seeds were treated by artificial aging (45°C, 100% relative humidity) and RNA-seq was used to study the response of HSP20 genes to artificial aging treatment of seeds, so as to provide evidences for uncovering the molecular mechanism of seed aging. In the study, the seed vigor decreased dramatically with increasing aging time. The activity of catalase in seed embryo showed a decreasing trend. The content of hydrogen peroxide increased to the maximum at the third day of aging and then decreased. The content of malonaldehyde increased, which indicated that the damage to the membrane system increased. Twenty-five HSP20 genes were identified. These genes encoded HSP20 and mainly distributed in nuclei, mitochondria and chloroplasts. There was a conserved ACD sequence (RVDWRETPDAHEIVVDVPGMRREDLRIEVE DNRVLRVSGERRRAEERKGDHWHREERSYGRFW RRFRLPENADLDSVAASLDSGVLTVRFRK) in HSP20 protein which contains more amino acids, such as Arg (11.2%), Lys (7.2%), Pro (4.2%), and Thr (3.9%). In the aging process these amino acids might be oxidized by ROS accumulated in the embryo, leading to protein structure damaged and loss of functions. The expression patterns of HSP20 genes in cytoplasm, chloroplasts and mitochondria were validated by qRT-PCR, showing that the expressions of cytoplasmic HSP20 genes were up-regulated and those of chloroplast and mitochondrial HSP20 genes reached peak at the third day of aging treatment, and then declined. These suggested that the HSP20 genes play an important role in seeds aging, and the targeted oxidation of Arg, Lys and other amino acids in the ACD structural domain may be an essential cause leading to seed deterioration.

Key words: maize, seed embryo, seed aging, differential expressed genes, HSP20

Lu-Man XING, Wei-Zeng LYU, Wei LEI, Yu-Huan LIANG, Yang LU, Jun-Ying CHEN. Response of HSP20 Genes to Artificial Aging Treatment in Maize Embryo[J].Acta Agronomica Sinica, 2018, 44(11): 1733-1742.

Figures/Tables 9

Supplementary table 1

Primer for expression analysis of HAP20s genes"

基因ID号 Gene ID	引物序列 Primer sequence (5'-3')		退火温度 Annealing temperature (℃)	产物长度 Product length (bp)
GRMZM2G049767	F	GCTGTACTACAGGTAGTATAAC	59	105
	R	GCACAACACTTCCATACA	59
GRMZM2G306679	F	TAGTGATTATCCGTTATGTACTC	59	141
	R	ACAGAACAGATTGTATTAGGTT	59
GRMZM2G479260	F	GTCTTCGTCTTAGTGTTGAT	58.9	109
	R	TACTCAAGACAGCCATGA	59
GRMZM2G375517	F	AGATGTGAATGTGATCTGAC	59	116
	R	AGTACATATTAACTCACGCAA	58.9
GRMZM5G858128	F	TCTAACAGAACTCGCTGA	59	179
	R	CATCATCATACCACCGATC	59.1
GRMZM2G346839	F	CATCATCATACCACCGATC	59.1	179
	R	TCTAACAGAACTCGCTGA	59
GRMZM2G335242	F	GTACGTCTGAATTCTGGTC	59.1	98
	R	CTCACTCGGATTACTTGC	58.9
GRMZM2G080724	F	GATTCATCAGGACGATGAG	59	101
	R	AAGAAGAGTGGTAGCTAGAT	58.9
GRMZM2G012455	F	AGACCATCGAGATTAAGGT	59	179
	R	ACACATCAGGTAGGATGAT	59
GRMZM2G311710	F	ACACATCAGGTAGGATGAT	59	179
	R	AGACCATCGAGATTAAGGT	59
GRMZM2G333635	F	GAGGACAAGAACGACAAG	59.1	154
	R	TCTTCACCTCCGTCTTAG	59.1
GRMZM2G158232	F	TGAGGTGAAGGCTATTGA	59	89
	R	ATATCGTAGAAGACACAGGA	59
GRMZM2G331701	F	ATGACTCAATGACGGAGA	59.1	90
	R	GCATATACGCTGTCACAA	58.9
GRMZM5G849535	F	ATATCGTAGAAGACACAGGA	59	110
	R	GATATCTGGTTGAGCATCC	59
GRMZM2G034157	F	GGTATCACCAATCATCCTATC	59.2	83
	R	GTTACAAGAGACATGTAGAGAT	58.9
GRMZM2G083810	F	ACCAATTCCTATCTGATGTATC	59	112
	R	GTCAATGAAGACTACGGATTA	59
GRMZM2G332512	F	GCTTGTTGTGTTGGTTTC	58.9	117
	R	CCTTCCAGAAGAACAGAAG	59
GRMZM2G404274	F	ACACATCAGGTAGGATGAT	59	179
	R	AGACCATCGAGATTAAGGT	59
GRMZM2G085934	F	CGATGGACATAGTGGAGA	59.2	113
	R	CTTCATCACTAGCATCCG	58.7
GRMZM2G149647	F	GGTCTTTGTGTAGCACTAG	59.1	109
	R	ATAGCAGAATTGAAGCGTT	59
GRMZM2G098167	F	AGAGTAGCAGCAGTAGAAT	59	154
	R	CAGAGACACGATTGAATAATTC	59
GRMZM2G046382	F	GTCGTAATGTCGTAGGATG	59	85
	R	ATCCATACATAGATTCAGACAC	59
AC208204.3_FGT006	F	CAACGTCCTTCAGATCAG	58.9	175
	R	CACAGTGACTGTAAGCAC	59.4
NM-001155179.1(Actin1)	F	GTATGAGCAAGGAGATCAC	58.9	194
	R	TTAGAAGCACTTCATGTGG	59.0

Supplementary table 1

Table 1

Fig. 1

Table 2

Table 3

HSP20 related genes and their expression in artificial aged maize seed embryo"

基因ID Gene ID	log₂比值 log₂Ratio	P值 P-value	错误发现率 False discovery rate	基因注释 Gene annotation	亚细胞定位 Subcellular localization
GRMZM2G481605	8.84	1.89E-259	5.63E-257	17.9 kD class I heat shock protein	Nuclear
GRMZM2G049767	8.19	1.07E-164	2.04E-162	17.9 kD class I heat shock protein	Nuclear
GRMZM2G306679	7.37	0	0	17.9 kD class I heat shock protein	Nuclear
GRMZM2G479260	6.71	0	0	17.9 kD class I heat shock protein	Nuclear
GRMZM2G375517	6.70	0	0	17.8 kD class I heat shock protein	Cytoplasmic
GRMZM5G858128	6.53	6.93E-222	1.83E-219	22.0 kD class IV heat shock protein	Mitochondria
GRMZM2G306714	6.28	0	0	Class I heat shock protein 3	Nuclear
GRMZM2G346839	5.97	7.04E-66	5.89E-64	18.3 kD class I heat shock protein	Cytoplasmic
GRMZM2G335242	5.96	0	0	15.7 kD class I heat shock protein	Cytoplasmic
GRMZM2G080724	5.61	0	0	Retrotransposon protein [Zea mays]	Mitochondrial
GRMZM2G012455	5.19	8.52E-135	1.34E-132	17.6 kD class II heat shock protein	Cytoplasmic
GRMZM2G311710	4.98	0	0	17.0 kD class II heat shock protein	Nuclear
GRMZM2G333635	4.48	0	0	17.9 kD class I heat shock protein	Nuclear
GRMZM2G158232	4.33	0	0	17.9 kD class I heat shock protein	Nuclear
GRMZM2G331701	4.31	0	0	18.3 kD class I heat shock	Cytoplasmic
GRMZM5G849535	4.30	0	0	16.9 kD class I heat shock protein 1-like	Mitochondrial
GRMZM2G034157	4.27	2.84E-22	8.22E-21	17.8 kD class II heat shock protein	Cytoplasmic
GRMZM2G083810	4.18	0	0	17.8 kD class II heat shock protein	Cytoplasmic
GRMZM2G332512	3.94	1.36E-70	1.22E-68	Heat shock protein18c (hsp18c)	Nuclear
GRMZM2G085934	3.88	3.28E-10	4.36E-09	18.6 kD class III heat shock protein	Cytoplasmic
GRMZM2G404274	3.88	1.34E-197	3.16E-195	18.0 kD heat shock protein 18a	Nuclear
GRMZM2G149647	3.52	0	0	Hsp26 - heat shock protein 26	Chloroplast
GRMZM2G098167	3.49	0	0	17.0 kD class II heat shock protein	Cytoplasmic
GRMZM2G046382	3.22	0	0	17.9 kD class I heat shock protein	Nuclear
AC208204.3_FG006	3.10	0	0	17.4 kD class I heat shock protein	Nuclear

Table 3

Fig. 2

Table 4

Supplementary fig. 1

Fig. 3

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处理时间 Treatment time	发芽率 Germination rate (%)	发芽指数 Germination index	活力指数 Vigor index	平均发芽日数 Mean days of germination (d)
0 d	100±0.00 a	9.44±0.25 a	124.14±5.57 a	2.17±0.08 b
1 d	100±0.00 a	8.22±0.54 b	100.07±4.93 b	2.53±0.16 b
2 d	100±0.00 a	8.11±0.25 b	101.32±1.17 b	2.56±0.08 b
3 d	86.67±0.06 b	6.29±0.32 c	75.73±10.45 c	2.86±0.03 b
4 d	66.67±0.08 c	3.96±0.50 d	41.63±5.39 d	3.48±0.11 ab
5 d	21.67±0.10 d	1.11±0.42 e	8.95±1.84 e	4.11±1.02 a

处理时间 Treatment time	过氧化氢酶活性 CAT activity (U g^-1 FW min^-1)	过氧化氢含量 H₂O₂content (μmol g^-1 FW)	丙二醛含量 MDA content (μmol g^-1 FW)
0 d	0.39±0.11 a	0.38±0.03 b	0.16±0.11 c
1 d	0.26±0.13 a	0.40±0.02 b	0.18±0.03 c
2 d	0.18±0.12 ab	0.42±0.02 b	0.33±0.10 c
3 d	0.14±0.07 b	0.51±0.06 a	0.43±0.19 c
4 d	0.14±0.03 b	0.43±0.02 ab	0.87±0.26 b
5 d	0.06±0.02 b	0.29±0.03 c	1.955±0.21 a

基因ID Gene ID	ACD氨基酸数 Amino acid number	个数/百分比Number/percentage (%)
基因ID Gene ID	ACD氨基酸数 Amino acid number	精氨酸 Arg (R)	赖氨酸 Lys (K)	脯氨酸 Pro (P)	苏氨酸 Thr (T)
GRMZM2G481605	109	12/11.0	3/2.7	3/2.7	4/3.7
GRMZM2G049767	96	14/14.6	5/5.2	4/4.2	3/3.1
GRMZM2G306679	92	9/9.8	10/10.9	4/4.3	3/3.3
GRMZM2G479260	92	8/8.7	9/9.8	3/3.3	8/8.7
GRMZM2G375517	87	16/18.4	4/4.6	4/4.6	3/3.4
GRMZM2G346839	93	17/18.3	2/2.3	3/3.2	2/2.2
GRMZM2G335242	97	6/6.2	8/8.2	7/7.2	4/4.1
GRMZM2G080724	97	10/10.3	4/4.1	6/6.2	3/3.1
GRMZM2G012455	97	11/11.3	5/5.2	2/2.1	3/3.1
GRMZM2G333635	92	8/8.7	11/12	4/4.3	3/3.3
GRMZM2G158232	92	8/8.7	11/12	4/4.3	4/4.3
GRMZM2G331701	92	19/20.7	2/2.2	3/3.3	2/2.3
GRMZM2G034157	98	10/10.2	5/5.1	3/3.1	2/2.0
GRMZM2G083810	87	10/11.5	6/6.9	3/3.4	2/2.3
GRMZM2G085934	78	9/11.5	5/6.4	4/5.1	1/1.3
GRMZM2G149647	107	6/5.6	11/10.3	3/2.8	3/2.8
GRMZM2G098167	113	11/9.7	10/8.8	7/6.2	6/5.3
GRMZM2G046382	92	8/8.7	9/9.8	4/4.3	7/7.6
AC208204.3_FG006	92	8/8.7	9/9.8	4/4.3	7/7.6
平均值Mean	95	11/11.2	7/7.2	4 /4.2	4/3.9

Response of HSP20 Genes to Artificial Aging Treatment in Maize Embryo

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