Chinese Bulletin of Botany ›› 2021, Vol. 56 ›› Issue (5): 520-532.DOI: 10.11983/CBB21119
• EXPERIMENTAL COMMUNICATIONS • Previous Articles Next Articles
Jiangyuan Shang, Yan Chun, Xueyong Li*()
Received:
2021-07-21
Accepted:
2021-09-16
Online:
2021-09-01
Published:
2021-09-16
Contact:
Xueyong Li
Jiangyuan Shang, Yan Chun, Xueyong Li. Map-based Cloning and Natural Variation Analysis of the PAL3 Gene Controlling Panicle Length in Rice[J]. Chinese Bulletin of Botany, 2021, 56(5): 520-532.
Primer name | Forward primer (5′→3′) | Reverse primer (5′→3′) |
---|---|---|
R11-3 | AGAGAGACATCCGGAGACAA | TAAGACGAAAGGTCAAACGT |
R11-5 | GTGCTAACGTTTCGTCTAAC | AATAGCCTTCGGTGGTCTCA |
R11-10 | GTTCGTAATGTGGGCGTCTT | TGGGCACTCTTCTCACACTG |
R11-12 | CAATCTTGCTCTACTAGCTAGTG | GTGGCAACTAACAGATTAGATG |
M1 | GCAGTATATATTCGGCGGCG | GCCGTCGCCATATAGCTG |
M2 | GAGCCTCTCCTACTGTGCTA | AGAGCCCTCAGTTCCTCAAT |
M3 | GCTGACTACAGTAAGATCATGC | AGACAAACGGTCAAACATGT |
M4 | AAGGATCCAAGCTAGCCTCC | CCTGACAGCAAGCGAGAGAT |
M5 | CTTCAGCAAGTGAACTACGA | CCTAAACTAGCACGGATCATAGC |
M6 | TGTGAGGTTTAGGTTCTCGGA | TGAATAGAGATGCGGTCCAAC |
M7 | GGATTCGGCCACTGGTTGTT | GAATGTACTCGGATAAACCC |
Table 1 Primers used for mapping
Primer name | Forward primer (5′→3′) | Reverse primer (5′→3′) |
---|---|---|
R11-3 | AGAGAGACATCCGGAGACAA | TAAGACGAAAGGTCAAACGT |
R11-5 | GTGCTAACGTTTCGTCTAAC | AATAGCCTTCGGTGGTCTCA |
R11-10 | GTTCGTAATGTGGGCGTCTT | TGGGCACTCTTCTCACACTG |
R11-12 | CAATCTTGCTCTACTAGCTAGTG | GTGGCAACTAACAGATTAGATG |
M1 | GCAGTATATATTCGGCGGCG | GCCGTCGCCATATAGCTG |
M2 | GAGCCTCTCCTACTGTGCTA | AGAGCCCTCAGTTCCTCAAT |
M3 | GCTGACTACAGTAAGATCATGC | AGACAAACGGTCAAACATGT |
M4 | AAGGATCCAAGCTAGCCTCC | CCTGACAGCAAGCGAGAGAT |
M5 | CTTCAGCAAGTGAACTACGA | CCTAAACTAGCACGGATCATAGC |
M6 | TGTGAGGTTTAGGTTCTCGGA | TGAATAGAGATGCGGTCCAAC |
M7 | GGATTCGGCCACTGGTTGTT | GAATGTACTCGGATAAACCC |
Figure 1 Comparison of the rice phenotype between wild type and the pal3 mutants (A) Gross morphology of wild type and the pal3 mutants at mature stage (Bar=20 cm); (B) Closed panicle of wild type and the pal3 mutants (Bar=2 cm); (C) Spread panicles of wild type and the pal3 mutants (right) (Bar=2 cm); (D)-(G) Statistic analysis of panicle length (D), primary branch number (E), secondary branch number (F) and grain number per panicle (G) of wild type and the pal3 mutants. WT: Wild type. Data in figure are means±SD (n=20). ** indicate the significant differences at P<0.01 level by Students’t test.
Hybrid combi- nations | Phenotype of F1 | F2 population | χ23:1 | ||
---|---|---|---|---|---|
Wild-type plant number | Mutant-phenotypic plant number | Total number | |||
pal3-1 × WT | WT | 190 | 60 | 250 | 0.13 |
pal3-2 × WT | WT | 196 | 68 | 264 | 0.08 |
pal3-3 × WT | WT | 180 | 63 | 243 | 0.11 |
pal3-4 × WT | WT | 201 | 69 | 270 | 0.04 |
Table 2 Genetic analysis of the pal3-1, pal3-2, pal3-3 and pal3-4 rice mutants
Hybrid combi- nations | Phenotype of F1 | F2 population | χ23:1 | ||
---|---|---|---|---|---|
Wild-type plant number | Mutant-phenotypic plant number | Total number | |||
pal3-1 × WT | WT | 190 | 60 | 250 | 0.13 |
pal3-2 × WT | WT | 196 | 68 | 264 | 0.08 |
pal3-3 × WT | WT | 180 | 63 | 243 | 0.11 |
pal3-4 × WT | WT | 201 | 69 | 270 | 0.04 |
Figure 2 Map-based cloning of the PAL3 gene (A) The PAL3 gene was mapped to a region between InDel markers R11-3 and R11-5 on chromosome 11; (B), (C) Fine-mapping of PAL3. The PAL3 gene was further delimited to the region between markers M3 and M5, which contains seven open reading frames (ORFs). The number of recombinants is indicated beneath the marker positions; (D) Schematic structure of the PAL3 gene and the mutation sites of pal3 mutants. WT: Wild type
Gene ID | Annotation |
---|---|
LOC_Os11g12680 | Expressed protein |
LOC_Os11g12690 | Retrotransposon protein, putative, unclassified, expressed |
LOC_Os11g12700 | Retrotransposon protein, putative, unclassified, expressed |
LOC_Os11g12710 | Retrotransposon protein, putative, unclassified, expressed |
LOC_Os11g12720 | Retrotransposon, putative, centromere-specific |
LOC_Os11g12730 | Transposon protein, putative, CACTA, En/Spm sub-class, expressed |
LOC_Os11g12740 | Peptide transporter PTR2, putative, expressed |
Table 3 Annotations of candidate genes in the fine-mapping region
Gene ID | Annotation |
---|---|
LOC_Os11g12680 | Expressed protein |
LOC_Os11g12690 | Retrotransposon protein, putative, unclassified, expressed |
LOC_Os11g12700 | Retrotransposon protein, putative, unclassified, expressed |
LOC_Os11g12710 | Retrotransposon protein, putative, unclassified, expressed |
LOC_Os11g12720 | Retrotransposon, putative, centromere-specific |
LOC_Os11g12730 | Transposon protein, putative, CACTA, En/Spm sub-class, expressed |
LOC_Os11g12740 | Peptide transporter PTR2, putative, expressed |
Figure 3 Transmembrane regions and phylogenetic analysis of PAL3 (A) 12 transmembrane domains of PAL3 and the mutation sites of the pal3-1, pal3-2 and pal3-4 mutants; (B) Phylogenetic analysis of PAL3 and its homologs. The phylogenetic analysis was carried out by MEGA X with 1 000 bootstrap replicates and was constructed using the distance method with maximum likelihood.
Species | Locus ID | Gene name |
---|---|---|
Rice | LOC_Os11g12740 | PAL3/SP1 |
LOC_Os07g01070 | OsPTR1 | |
LOC_Os12g44100 | OsPTR2 | |
LOC_Os10g33210 | OsPTR3 | |
LOC_Os07g41250 | OsPTR4 | |
LOC_Os04g50940 | OsPTR5 | |
LOC_Os04g50950 | OsPTR6 | |
LOC_Os01g04950 | OsPTR7 | |
LOC_Os03g51050 | OsPTR8 | |
LOC_Os06g49250 | OsPTR9 | |
LOC_Os03g13274 | OsNTR1.1 | |
LOC_Os10g40600 | OsNRT1.1B | |
LOC_Os01g54515 | OsNPF4.5 | |
Arabidopsis | At1g12110 | AtNRT1.1 |
At5g62680 | AtNRT1.10 | |
At1g52190 | AtNRT1.11 | |
At1g33440 | AtNRT1.13 | |
At1g59740 | AtNRT1.14 | |
At1g72120 | AtNRT1.15 | |
At1g72125 | AtNRT1.16 | |
At1g69850 | AtNRT1.2 | |
At3g21670 | AtNRT1.3 | |
At2g26690 | AtNRT1.4 | |
At1g32450 | AtNRT1.5 | |
At1g27080 | AtNRT1.6 | |
At1g69870 | AtNRT1.7 | |
At4g21680 | AtNRT1.8 | |
At1g18880 | AtNRT1.9 | |
At3g54140 | AtPTR1 | |
At2g02040 | AtPTR2/AtNTR1 | |
At5g46050 | AtPTR3 | |
At2g02020 | AtPTR4 | |
At5g01180 | AtPTR5 | |
At1g62200 | AtPTR6 | |
At1g27040 | AIT2 | |
At3g25260 | AIT3 | |
At3g25280 | AIT4 | |
At3g47960 | GTR1 | |
At3g45650 | NAXT1 | |
At1g68570 | Nitr |
Table 4 Locus ID and gene name of PAL3 and its homologs
Species | Locus ID | Gene name |
---|---|---|
Rice | LOC_Os11g12740 | PAL3/SP1 |
LOC_Os07g01070 | OsPTR1 | |
LOC_Os12g44100 | OsPTR2 | |
LOC_Os10g33210 | OsPTR3 | |
LOC_Os07g41250 | OsPTR4 | |
LOC_Os04g50940 | OsPTR5 | |
LOC_Os04g50950 | OsPTR6 | |
LOC_Os01g04950 | OsPTR7 | |
LOC_Os03g51050 | OsPTR8 | |
LOC_Os06g49250 | OsPTR9 | |
LOC_Os03g13274 | OsNTR1.1 | |
LOC_Os10g40600 | OsNRT1.1B | |
LOC_Os01g54515 | OsNPF4.5 | |
Arabidopsis | At1g12110 | AtNRT1.1 |
At5g62680 | AtNRT1.10 | |
At1g52190 | AtNRT1.11 | |
At1g33440 | AtNRT1.13 | |
At1g59740 | AtNRT1.14 | |
At1g72120 | AtNRT1.15 | |
At1g72125 | AtNRT1.16 | |
At1g69850 | AtNRT1.2 | |
At3g21670 | AtNRT1.3 | |
At2g26690 | AtNRT1.4 | |
At1g32450 | AtNRT1.5 | |
At1g27080 | AtNRT1.6 | |
At1g69870 | AtNRT1.7 | |
At4g21680 | AtNRT1.8 | |
At1g18880 | AtNRT1.9 | |
At3g54140 | AtPTR1 | |
At2g02040 | AtPTR2/AtNTR1 | |
At5g46050 | AtPTR3 | |
At2g02020 | AtPTR4 | |
At5g01180 | AtPTR5 | |
At1g62200 | AtPTR6 | |
At1g27040 | AIT2 | |
At3g25260 | AIT3 | |
At3g25280 | AIT4 | |
At3g47960 | GTR1 | |
At3g45650 | NAXT1 | |
At1g68570 | Nitr |
Figure 4 Sequences alignment of PAL3 and its homologs The black background represent that all amino acid sequences are homologous. The gray part represent that some of the amino acid sequences are homologous. The white part represent that the amino acid sequences are completely different. * indicate the mutation sites of the pal3-1, pal3-2 and pal3-4 mutants in turn; △: The variation sites between Hap1 and Hap3.
Figure 5 Haplotype analysis of PAL3 (A) Structure of PAL3 gene and the location of haplotypes; (B) Haplotypes of PAL3 in the cultivated rice. Haplotype data were analyzed by Rice Molecular Breeding Knowledgebase (MBKbase) (Peng et al., 2020), including 2 187 cultivated rice varieties. Variations on intron, heterozygous variations and synonymous variations were removed during analysis (-: The deletion of base pair); (C) Sequence alignment around the variation sites between Hap1 and hap3 (Δ: The variation sites between Hap1 and Hap3).
Figure 6 Haplotype of PAL3 in the wild rice Haplotype data of wild rice were from Gramene and Rice Relatives-GD (Mao et al., 2019). *: D475 and G476 deletion; ●: No datum
Figure 7 Subspecies composition and panicle phenotype analysis of Hap1, Hap2 and Hap3 of the PAL3 gene (A) Subspecies composition of Hap1, Hap2 and Hap3; (B) Comparison of panicle length among Hap1, Hap2 and Hap3; (C) Statistic of panicle length of Hap1, Hap2 and Hap3. Different lowercase letters indicate significant differences (P<0.05).
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