Rice Science ›› 2021, Vol. 28 ›› Issue (4): 358-367.DOI: 10.1016/j.rsci.2021.05.006
收稿日期:
2020-11-12
接受日期:
2021-01-28
出版日期:
2021-07-28
发布日期:
2021-07-28
. [J]. Rice Science, 2021, 28(4): 358-367.
Fig. 1. Knock-out of OsbZIP09 alleviates rice pre-harvest sprouting (PHS).A, Schematic diagram of the two targets in OsbZIP09 gene and the mutation information of the related mutants. The protospacer adjacent motif (PAM) is highlighted in blue, insert nucleotide is highlighted in red, and deleted nucleotides are shown with minus sign ‘-’. B, Germination phenotype of mature rice panicles after 6 d imbibition in water. Scale bar, 2 cm. ZH11, Zhonghua 11.C, Time-course analysis of germination rates of seeds in the panicles. Data are Mean ± SD (n = 30).
Fig. 2. Agronomic traits and normal seed germination assay of osbzip09 mutants.A, Morphology of osbzip09 mutants and its wild type Zhonghua 11 (ZH11). Scale bar, 10 cm. B?J, Plant height (B), heading date (C), tiller number per plant (D), panicle length (E), grain number per panicle (F), 1000-grain weight (G), grain shape (scale bar, 1 cm) (H), grain length (I) and grain width (J) of osbzip09 mutants and wild type. Data are Mean ± SD (n = 10 in B?G, and n = 30 in I and J). **, P < 0.01 (t-test).
Fig. 3. Expression and sub-cellular localization analysis of OsbZIP09.A, Expression profiling of OsbZIP09 transcript abundance in developing seeds of rice. B, Time-course expression analysis of OsbZIP09 after imbibition. C, Expression analysis of OsbZIP09 in response to abscisic acid (ABA, 5 μmol/L) and gibberellins (GA, 10 μmol/L) treatments. D, Sub-cellular localization of OsbZIP09 (35S::OsbZIP09-GFP) in tobacco epidermal cells. Scale bars, 20 μm. Actin served as an internal reference gene for normalization in A?C. Data are Mean ± SD (n = 3). **, P < 0.01 (t-test).
Fig. S2. Expression analysis of ABA or GA responsive marker genes.A, Expression analysis of RAB16A, ABA8ox1 and ABA8ox2 in response to ABA (5 μmol/L) treatment.B, Expression analysis of GA20ox1 and GA2ox1 in response to GA (10 μmol/L). Actin served as an internal reference gene for normalisation in A and B. Data are Mean ± SD (n = 3). ** indicates significant difference (P < 0.01, t-test).
Fig. 4. Identification of OsbZIP09 binding motif and its targets involved in regulation of rice seed germination.A, Conserved motif identified from the DNA affinity purification sequencing (DAP-seq) data. B, Distribution of OsbZIP09 binding motifs in the 2-kb region preceding the initial codon (ATG) of ABA8ox1 and LEA3. C and D, Time-course expression analysis of ABA8ox1 and LEA3 after imbibition. Data are Mean ± SD (n = 3).E, Schematic program of the reporter and effector constructs used in the dual-luciferase reporter assay. F and G, OsbZIP09 activates the promoter activity of ABA8ox1 but suppresses the promoter activity of LEA3. Data are Mean ± SD (n = 5). ** indicates significant difference (P < 0.01, t-test).
Chromosome | Start | End | -log10(P-value)' | Strand | Distance_ to_TSS | Category | Gene ID | Gene name |
---|---|---|---|---|---|---|---|---|
5 | 26927280 | 26927615 | 7.60524 | + | 3 | Promoter | Os05g0542500 | OsLEA3; Similar to isoform 2 of late embryogenesis abundant protein 2C group 3. |
2 | 28994350 | 28994666 | 8.95194 | + | 1052 | Promoter | Os02g0703600 | OsABA8ox1; Similar to abscisic acid 8'-hydroxylase 1. (Os02t0703600-01) |
5 | 4663664 | 4663921 | 6.50438 | - | -1100 | Promoter | Os05g0178100 | OsGA3ox1;3beta-hydroxylase GA metabolism (Os05t0178100-01) |
Table S1. DAP-seq data related to ABA8ox1, LEA3 and GA3ox1.
Chromosome | Start | End | -log10(P-value)' | Strand | Distance_ to_TSS | Category | Gene ID | Gene name |
---|---|---|---|---|---|---|---|---|
5 | 26927280 | 26927615 | 7.60524 | + | 3 | Promoter | Os05g0542500 | OsLEA3; Similar to isoform 2 of late embryogenesis abundant protein 2C group 3. |
2 | 28994350 | 28994666 | 8.95194 | + | 1052 | Promoter | Os02g0703600 | OsABA8ox1; Similar to abscisic acid 8'-hydroxylase 1. (Os02t0703600-01) |
5 | 4663664 | 4663921 | 6.50438 | - | -1100 | Promoter | Os05g0178100 | OsGA3ox1;3beta-hydroxylase GA metabolism (Os05t0178100-01) |
Fig. 5. Expression analysis of ABA8ox1 and LEA3 in rice seeds before and after imbibition of abscisic acid (ABA).A and B, Expression analysis of ABA8ox1 (A) and LEA3 (B) in osbzip09 mutants and wild type Zhonghua 11 (ZH11) with or without ABA treatment. Actin served as an internal reference gene for normalization. Different lowercase letters indicate significant differences at P < 0.05 level based on one-way analysis of variance. C, ABA content in osbzip09 mutants and ZH11 before or at 12 h after imbibition (HAI).
Fig. S3. Expression analysis of GA3ox1 and quantification of GA3 content in rice seeds before and after imbibition. A, Expression analysis of GA3ox1 in osbzip09 mutants and ZH11. Actin served as an internal reference gene for normalisation. B, GA3 content in osbzip09 mutants and ZH11 before or at 12 h after imbibition (HAI). Data are Mean ± SD (n = 3). ** indicates significant difference (P < 0.01, t-test). NS, Not significant.
Fig. 6. Simplified model showing OsbZIP09 promotes rather than suppresses seed germination via reducing abscisic acid (ABA) accumulation and attenuating ABA-induction of downstream responsive genes in rice.
Primer name | Sequence ( 5' → 3' ) | Usage |
---|---|---|
OsbZIP09a-SG-seq | GTACAGCCTCACCCTCGACG | SG sequence-1 for editing OsbZIP09 SG sequence-2 for editing OsbZIP09 |
OsbZIP09b-SG-seq | AAGAGGAATGCCGAAACGGG | |
OsbZIP09seq-F | ATCTCGTCTCGATCATGGGG | Primers for sequencing and confirming mutation of OsbZIP09 |
OsbZIP09seq-R | CCCCAGCTTTGACCAGGAAA | |
bZIP09qRT-F | GAGGCATGTCAGGGGATGTG | For qRT-PCR analysis of OsbZIP09 |
bZIP09qRT-R | ACTTGGGTTCTGGTACAGGC | |
RAB16AqRT-F | ATGAGGGAGGAGCACAAGAC | For qRT-PCR analysis of RAB16A |
RAB16AqRT-R | TTGATCCCCTTCTTCCTCCT | |
ABA8ox1qRT-F | AAGCTGGCAAAACCAACATC | For qRT-PCR analysis of ABA8ox1 |
ABA8ox1qRT-R | TGGATTCCGTATTAGCACGG | |
ABA8ox2qRT-F | GCGAGACGCTCCAGCTCT | For qRT-PCR analysis of ABA8ox2 |
ABA8ox2qRT-R | GGGCACCCCAGCAGATT | |
GA20ox1qRT-F | GCCACTACAGGGCCGACAT | For qRT-PCR analysis of GA20ox1 |
GA20ox1qRT-R | TGGTTGCAGGTGACGATGAT | |
GA2ox1qRT-F | TGACGATGATGACAGCGACAA | For qRT-PCR analysis of GA2ox1 |
GA2ox1qRT-R | CCATAGGCATCGTCTGCAATT | |
LEA3qRT-F | TCACTTCAAATTCGGTGCAA | For qRT-PCR analysis of LEA3 |
LEA3qRT-R | CACACCCGTCAGAAATCCTC | |
Actin1qRT-F | CCAAGGCCAATCGTGAGAAGA | Reference gene for qRT-PCR assay |
Actin1qRT-R | AATCAGTGAGATCACGCCCAG | |
pZIP09-Smal-F | TCCCCCGGGATGGCGTCGAAGGCCGGAG | For subcellular assay of OsbZIP09 |
pZIP09-XbaI-R | GCTCTAGAGAAATCTGCGGAGCTTGTTC | |
pZIP09-EcoRI-F | CGGAATTCATGGCGTCGAAGGCCGGAG | For construction of OsbZIP09-62-SK |
pZIP09-KpnI-R | GGGGTACCTCAGAAATCTGCGGAGCTTG | |
proLEA3Kpn-F | GGGGTACCGTCGGCCACCCTCCGCAATA | For construction of LEA3pro-pGreenII |
proLEA3PstI-R | AACTGCAGACCTGCCCCGCCTTCTCCTG | |
proABA8ox1Kpn-F | GGGGTACCGTAGCCTGTTGGTAGCTTGG | For construction of ABA8ox1pro-pGreenII |
proABA8ox1PstI-R | AACTGCAGCGCCTGCTCACTTCACTTAT | |
DAP-seq Adapter A | CACGACGCTCTTCCGATCT | For DAP-seq sampling |
DAP-seq Adapter B | GATCGGAAGAGCACACGTCTG |
Table S2. Oligonucleotide primer sequences used.
Primer name | Sequence ( 5' → 3' ) | Usage |
---|---|---|
OsbZIP09a-SG-seq | GTACAGCCTCACCCTCGACG | SG sequence-1 for editing OsbZIP09 SG sequence-2 for editing OsbZIP09 |
OsbZIP09b-SG-seq | AAGAGGAATGCCGAAACGGG | |
OsbZIP09seq-F | ATCTCGTCTCGATCATGGGG | Primers for sequencing and confirming mutation of OsbZIP09 |
OsbZIP09seq-R | CCCCAGCTTTGACCAGGAAA | |
bZIP09qRT-F | GAGGCATGTCAGGGGATGTG | For qRT-PCR analysis of OsbZIP09 |
bZIP09qRT-R | ACTTGGGTTCTGGTACAGGC | |
RAB16AqRT-F | ATGAGGGAGGAGCACAAGAC | For qRT-PCR analysis of RAB16A |
RAB16AqRT-R | TTGATCCCCTTCTTCCTCCT | |
ABA8ox1qRT-F | AAGCTGGCAAAACCAACATC | For qRT-PCR analysis of ABA8ox1 |
ABA8ox1qRT-R | TGGATTCCGTATTAGCACGG | |
ABA8ox2qRT-F | GCGAGACGCTCCAGCTCT | For qRT-PCR analysis of ABA8ox2 |
ABA8ox2qRT-R | GGGCACCCCAGCAGATT | |
GA20ox1qRT-F | GCCACTACAGGGCCGACAT | For qRT-PCR analysis of GA20ox1 |
GA20ox1qRT-R | TGGTTGCAGGTGACGATGAT | |
GA2ox1qRT-F | TGACGATGATGACAGCGACAA | For qRT-PCR analysis of GA2ox1 |
GA2ox1qRT-R | CCATAGGCATCGTCTGCAATT | |
LEA3qRT-F | TCACTTCAAATTCGGTGCAA | For qRT-PCR analysis of LEA3 |
LEA3qRT-R | CACACCCGTCAGAAATCCTC | |
Actin1qRT-F | CCAAGGCCAATCGTGAGAAGA | Reference gene for qRT-PCR assay |
Actin1qRT-R | AATCAGTGAGATCACGCCCAG | |
pZIP09-Smal-F | TCCCCCGGGATGGCGTCGAAGGCCGGAG | For subcellular assay of OsbZIP09 |
pZIP09-XbaI-R | GCTCTAGAGAAATCTGCGGAGCTTGTTC | |
pZIP09-EcoRI-F | CGGAATTCATGGCGTCGAAGGCCGGAG | For construction of OsbZIP09-62-SK |
pZIP09-KpnI-R | GGGGTACCTCAGAAATCTGCGGAGCTTG | |
proLEA3Kpn-F | GGGGTACCGTCGGCCACCCTCCGCAATA | For construction of LEA3pro-pGreenII |
proLEA3PstI-R | AACTGCAGACCTGCCCCGCCTTCTCCTG | |
proABA8ox1Kpn-F | GGGGTACCGTAGCCTGTTGGTAGCTTGG | For construction of ABA8ox1pro-pGreenII |
proABA8ox1PstI-R | AACTGCAGCGCCTGCTCACTTCACTTAT | |
DAP-seq Adapter A | CACGACGCTCTTCCGATCT | For DAP-seq sampling |
DAP-seq Adapter B | GATCGGAAGAGCACACGTCTG |
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