o2突变引起糯玉米籽粒淀粉积累差异研究

doi:10.3724/SP.J.1006.2024.33046

Abstract

Abstract:

The primary variety of fresh maize known as waxy maize undergoes a transformation with the introduction of the opaque2 (o2) mutant gene, resulting in an increased lysine content, thus improving the grain's nutritional composition. Yet, the o2 mutation brings about desirable agronomic traits such as wrinkle formation and a decrease in starch content, which restrict its use in breeding applications. To explore the high performing waxy maize o2 receptors, we capitalized on the use of o2 near-isogenic line (o2-NIL), specifically the plump and round grain type Nuo 2/wx1wx1o2o2 and its wrinkled counterpart, Huangnuo2/wx1wx1o2o2. Measurements of 100-grain weight and grain composition at the fresh ear and mature stages showed that there was difference in starch and soluble sugar content, which might be the primary cause of kernel phenotype variation between the two waxy maize o2-NILs. Genetic analysis of starch synthesis in the two o2-NILs was performed using qRT-PCR technique revealed that six gene-regulated trends fluctuated between 10 and 24 days after pollination, among which Sh1, Sh2, SSIIIa, and SBEIIb genes were significant differences. Endosperm transcriptomes indicated that 24 genes encoding trehalose and glycosyl hydrolases and 48 genes involved in endosperm modification exhibited distinct changes between the two o2-NILs. There was no detectable alteration in the 100-grain weight or the starch content of Nuo 2/wx1wx1o2o2, which may well be tied to the early high-level expression of the primary starch synthesis gene, leaving later stages unchanged compared with the control. Furthermore, the shifts in the expression of sugar metabolism genes was beneficial to starch synthesis, which may be an important reason why starch content and 100-kernel weight of Nuo 2/wx1wx1o2o2 were unaffected by the o2 mutation, and grain traits were significantly better than the superior grain traits compared with Huangnuo 2/wx1wx1o2o2. These results may be directly related to the differential expression of multiple endosperm modifying genes. The results of this study can provide important reference for the future utilization of o2 mutants in maize breeding.

Key words: waxy maize, Nuo 2/wx1wx1o2o2, Huangnuo 2/wx1wx1o2o2, kernel fullness degree, starch, sucrose metabolism, different expression genes

HAN Jie-Nan, ZHANG Ze, LIU Xiao-Li, LI Ran, SHANG-GUAN Xiao-Chuan, ZHOU Ting-Fang, PAN Yue, HAO Zhuan-Fang, WENG Jian-Feng, YONG Hong-Jun, ZHOU Zhi-Qiang, XU Jing-Yu, LI Xin-Hai, LI Ming-Shun. Analysis of differential accumulation of starch in waxy maize grain caused by the o2 mutation gene[J].Acta Agronomica Sinica, 2024, 50(5): 1207-1222.

Figures/Tables 11

Table S1

Fig. 1

Table 1

Fig. 2

Fig. 3

Fig. 4

Fig. 5

Fig. 6

Table S2

DEGs related to zein protein in waxy maize o2-NILs"

基因名称 Gene ID	糯2/wx1wx1o2o2 Nuo 2/wx1wx1o2o2 log₂FC	黄糯2/wx1wx1o2o2 Huangnuo 2/wx1wx1o2o2 log₂FC	注释 Annotation
Zm00001d005793	ns	-1.21	Prolamin 16 kD gamma zein precursor
Zm00001d048847	-1.80	-5.05	Prolamin 19 kD alpha zein z1A1_2 precursor
Zm00001d048848	ns	-3.56	Prolamin 19 kD alpha zein z1A1_3 precursor
Zm00001d048849	ns	-3.63	Prolamin 19 kD alpha zein z1A1_4 precursor
Zm00001d048850	-1.34	-3.67	Prolamin 19 kD alpha zein z1A1_5 precursor
Zm00001d048851	ns	-3.90	Prolamin 19 kD alpha zein z1A1_6 precursor
Zm00001d048852	ns	-3.76	Prolamin 19 kD alpha zein z1A1_7 precursor
Zm00001d019155	-1.36	-3.70	Prolamin 19 kD alpha zein z1B_4 precursor
Zm00001d030855	-1.33	-2.18	Prolamin 19 kD alpha zein z1D_4 precursor
newGene_32946	-1.67	-2.30	Prolamin 19 kD alpha zein z1D_2 precursor
newGene_32956	-2.52	-2.73	Prolamin 19 kD alpha zein z1D_2 precursor
newGene_17461	ns	-3.84	Prolamin 19 kD alpha zein z1A2_2 precursor
newGene_33790	-2.19	-3.66	Prolamin 19 kD alpha zein z1B_1 precursor
Zm00001d048816	-3.78	-4.24	Prolamin 22 kD alpha zein z1C1_10 precursor
Zm00001d048806	-2.78	-6.63	Prolamin 22 kD alpha zein z1C1_12 precursor
Zm00001d048817	-5.20	-5.80	Prolamin 22 kD alpha zein z1C1_19 precursor
Zm00001d048812	-5.50	-5.71	Prolamin 22 kD alpha zein z1C1_7 precursor
Zm00001d048813	-2.75	-3.58	Prolamin 22 kD alpha zein z1C1_8 precursor
Zm00001d049243	-4.90	-5.63	Prolamin 22 kD alpha zein z1C2 precursor
Zm00001d048810	-3.69	-4.77	Prolamin 22 kD alpha-zein 14
Zm00001d048809	-4.49	-4.61	Prolamin 22 kD alpha-zein 4
Zm00001d048818	-4.61	-4.30	Prolamin 22 kD alpha-zein 8
Zm00001d020591	ns	-1.87	Prolamin 50 kD gamma zein
Zm00001d035760	-1.70	ns	Prolamin PPROL 17 precursor
Zm00001d049476	-1.36	-2.90	Z1A alpha zein protein
Zm00001d045937	ns	-3.18	Zein protein
Zm00001d048807	-4.37	-4.79	Zein seed storage protein, hypothetical protein
Zm00001d019160	-2.11	-4.64	Zein seed storage protein, hypothetical protein
Zm00001d019162	-1.49	-5.11	Zein seed storage protein, hypothetical protein
Zm00001d019156	-1.22	-3.64	Zein seed storage protein, hypothetical protein
Zm00001d013100	-1.20	ns	Zein-binding
Zm00001d048808	-4.10	-4.96	Kafirin PSKR2 Precursor

Table S2

Table S3

Table S4

DEGs related to endosperm modification in waxy maize o2-NILs"

基因名称 Gene_ID	糯2/wx1wx1o2o2 Nuo 2/wx1wx1o2o2 log₂FC	黄糯2/wx1wx1o2o2 Huangnuo 2/wx1wx1o2o2 log₂FC	注释 Annotation
Zm00001d015504	1.70	1.70	Protein phosphatase 2C isoform gamma
Zm00001d039942	1.12	-2.28	16.9 kD class I heat shock protein 3
Zm00001d037717	2.25	-1.70	Heat shock 70 kD protein 14
Zm00001d011241	-1.58	ns	15.7 kD heat shock protein
Zm00001d039566	1.20	ns	17.5 kD class II heat shock protein
Zm00001d008841	1.61	ns	17.8 kD class II heat shock protein
Zm00001d031329	1.14	ns	Catalytic/protein phosphatase type 2C
Zm00001d039933	ns	-1.02	16.9 kD class I heat shock protein 1
Zm00001d039936	ns	-1.54	16.9 kD class I heat shock protein 1
Zm00001d017813	ns	-2.42	17.8 kD heat shock protein isoform X4
Zm00001d028557	ns	-1.47	17.9 kD class I heat shock protein
Zm00001d028561	ns	-2.11	17.9 kD class I heat shock protein
Zm00001d047841	ns	-2.32	17.9 kD class I heat shock protein
Zm00001d028555	ns	-1.64	18.1 kD class I heat shock protein
Zm00001d003554	ns	-1.24	22.0 kD class IV heat shock protein precursor
Zm00001d025508	ns	-1.46	22.0 kD class IV heat shock protein precursor
Zm00001d010693	ns	1.21	22.3 kD class VI heat shock protein
Zm00001d050119	ns	-1.44	Activator of 90 kDa heat shock protein ATPase
Zm00001d015227	ns	-2.02	Activator of Hsp90 ATPase
Zm00001d031740	ns	-1.38	Activator of Hsp90 ATPase
Zm00001d015777	ns	-1.90	Chloroplast small heat shock protein
Zm00001d018298	ns	-1.59	Class II heat shock protein
Zm00001d039259	ns	-1.30	DNAJ heat shock family protein
Zm00001d047726	ns	-1.71	DnaJ protein ERDJ3A
Zea_mays_new Gene_4468	ns	-1.13	Heat shock 70 kD protein
Zm00001d048073	ns	-1.88	Heat shock 70 kD protein 1
Zm00001d042922	ns	-1.17	Heat shock 70 kD protein-like
Zm00001d047799	ns	-1.81	Heat shock cognate 70 kD protein
Zm00001d028630	ns	-1.75	Heat shock cognate 70 kD protein 2
Zm00001d012420	ns	-2.06	Heat shock protein 1
Zm00001d024903	ns	-1.71	Heat shock protein 82
Zm00001d020898	ns	-1.33	Heat shock protein 90-2
Zm00001d031332	ns	-1.32	Heat shock protein 90-2
Zm00001d039935	ns	-1.34	Heat shock protein 17-2
Zm00001d028408	ns	-2.16	Heat shock protein26
Zm00001d030346	ns	2.95	Hsp20/alpha crystallin family protein
Zm00001d002542	ns	1.32	Probable protein phosphatase 2C
Zea_mays_new Gene_647	1.62	ns	Probable protein phosphatase 2C
Zm00001d021817	-1.01	ns	Probable protein phosphatase 2C
Zm00001d017643	-1.48	ns	Probable protein phosphatase 2C 25
Zm00001d047807	-2.29	ns	probable protein phosphatase 2C 31
Zm00001d020100	1.23	ns	Probable protein phosphatase 2C 68
Zm00001d045919	-1.88	ns	Pyrophosphate-fructose 6-phosphate-Phosphotransferase subunit alpha 2
Zm00001d028615	ns	-1.39	Probable protein phosphatase 2C 31
Zm00001d025055	ns	-1.88	Probable protein phosphatase 2C 37
Zm00001d011195	ns	-1.35	Probable protein phosphatase 2C 38
Zm00001d047847	ns	-1.19	Serine/arginine-rich splicing factor SR45a
Zm00001d031325	ns	-2.33	Small heat shock protein
Zm00001d032893	ns	-1.78	Small heat shock-like protein

Table S4

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基因名称 Gene ID	引物序列 Primer sequence (5'-3')	片段长度 Production size (bp)	退火温度 Annealing temperature (℃)
Zm00001d045042 (Sh1)	TGTTTCACCGCAATTCGCA	190	60
Zm00001d045042 (Sh1)	AGACAGGTGAACGAGCAGGC	190	60
Zm00001d029091 (Sus1)	GAAGCGTTCGGTCTCACC	162	60
Zm00001d029091 (Sus1)	GAAGAAGTCGGCCATCAG	162	60
Zm00001d050032 (Bt2)	CTATGACCGTTTTGCTCCAAT	195	60
Zm00001d050032 (Bt2)	GCACCCATTAGTAAACTGTCCTC	195	60
Zm00001d044129 (Sh2)	CGTGTCAGCTCTGGATGTGAA	210	60
Zm00001d044129 (Sh2)	AGCCTCTTGGATGCCCTTA	210	60
Zm00001d045261 (SSI)	GACATTAATGATTGGAACCCTGCC	187	60
Zm00001d045261 (SSI)	GAATGAGATCAATGCCTTTCTG	187	60
Zm00001d037234 (SSIIa)	CTGCACTCCTGCCTGTTTAT	175	60
Zm00001d037234 (SSIIa)	GGATCGTACAGCTCGAAATGTT	175	60
Zm00001d000002 (SSIIIa)	CAAAAGGGGATCCACCTGATCA	208	60
Zm00001d000002 (SSIIIa)	CAGAGCCAGCGTATATCAGAT	208	60
Zm00001d016684 (SBEIIb)	TAGACTTATCACAATGGGTTTAGGA	227	60
Zm00001d016684 (SBEIIb)	GCATTGCCTGATCAAACTCTTG	227	60
Zm00001d017502	GTCGCAGGACTCGTGAAACA	206	60
Zm00001d017502	TTGAAGGCGTCTTCGTCTGT	206	60
Zm00001d032311	GTATGGCACTATTTGGACGCG	186	60
Zm00001d032311	GACCAGTGTGTGAATCAGCTTG	186	60
Zm00001d052060	TGAATCGTCTGTGCGAGGACC	164	60
Zm00001d052060	CCACCTTGTGAAGTAACCGTGCT	164	60
Zm00001d012748	GGTTACTTCTTGAGGTGGTCT	208	60
Zm00001d012748	GCATCTCCTTTGCCTGTGAG	208	60
Zm00001d029371	CCCGCCGAGGTCAAGGAGTT	160	60
Zm00001d029371	TGCCGGTGCAGCAGGTAGGT	160	60
Zm00001d012173	ATCACCGCCACGAAGAAGGG	210	60
Zm00001d012173	TAGGAACTAGGGCAAGCAAA	210	60
Zm00001d042536	GCCTTCCTTCAAAGTACAACA	220	60
Zm00001d042536	CTCTATTAGCTGCAAGACCTCC	220	60
Zm00001d036608	ACCTCCGCTACTCCATCAACACCA	234	60
Zm00001d036608	CCCTTGCGAACGGGTAGAACG	234	60
Zm00001d034017	TCTCCACCGTCATGATCTCCTA	200	60
Zm00001d034017	AATGCCAGCAAGAATCGAAGCC	200	60
Zm00001d033649	GGAGAAGTATGGGAATCCAACG	213	60
Zm00001d033649	TACCCTGCCAGCCACTCGAAG	213	60
Zm00001d028243	GCCTTCAACGCCTACTACCACG	144	60
Zm00001d028243	GGTGCCGTTCATCAACGACGTC	144	60
Zm00001d005546	CAGCACGAGTGTTCTTGGGATC	199	60
Zm00001d005546	GCGGTTGAGCGAAGCAGAGT	199	60
Zm00001d053960	GCTGGAGCTTTTGGTCAGTTTGC	190	60
Zm00001d053960	GGTCGCACCAGATACTGAAATCT	190	60
Zm00001d002256	CGACAGAGCCATAACCACAT	172	60
Zm00001d002256	AAACGAGCCTGATTTCCCTA	172	60
Zm00001d015327 (Ubiquitin)	TAAGCTGCCGATGTGCCTGCGTCG	206	60
Zm00001d015327 (Ubiquitin)	CTGAAAGACAGAACATAATGAGCACAG	206	60

时期 Stage	自交系 Inbred lines	基因型 Genotype	赖氨酸含量 Lysine content (%)	可溶性糖含量 Soluble sugar content (%)	总淀粉含量 Starch content (%)	百粒重 100-kernel weight (g)
鲜食期	黄糯2	O2O2	0.35	7.70	69.60	11.98
Fresh stage	Huangnuo 2	o2o2	0.54^**	7.35	65.34^**	10.86^*
	糯2	O2O2	0.42	6.76	64.11	16.06
	Nuo 2	o2o2	0.58^**	6.77	68.02^**	15.94
成熟期	黄糯2	O2O2	0.24	4.56	68.90	15.80
Mature stage	Huangnuo 2	o2o2	0.43^**	5.13^*	64.60^**	14.41^**
	糯2	O2O2	0.24	4.81	69.20	26.01
	Nuo 2	o2o2	0.38^**	5.09	69.10	25.02

基因名称 Gene_ID	糯2/wx1wx1o2o2 Nuo 2/wx1wx1o2o2 log₂FC	黄糯2/wx1wx1o2o2 Huangnuo 2/wx1wx1o2o2 log₂FC	注释 Annotation
Zm00001d020984	-2.43	-4.10	Probable sarcosine oxidase
Zm00001d003983	-2.48	ns	Aldehyde dehydrogenase family 7 member A1
Zm00001d008432	ns	-1.01	Putative acetyl-CoA acetyltransferase cytosolic 2
Zm00001d052079	ns	-2.65	Lysine-ketoglutarate reductase/saccharopine dehydrogenase1

Analysis of differential accumulation of starch in waxy maize grain caused by the o2 mutation gene

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