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Research Article

Comparative SNP Analysis of Chloroplast Genomes and 45S nrDNAs Reveals Genetic Diversity of Perilla Species

Plant Breeding and Biotechnology 2018;6(2):125-139.
Published online: June 1, 2018

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*Corresponding author: Tae-Ho Kim, thkim@rda.go.kr, Tel: +82-63-238-4563, Fax: +82-63-238-4554
• Received: March 20, 2018   • Revised: April 10, 2018   • Accepted: April 10, 2018

Copyright © 2018 The Korean Society of Breeding Science

This is an Open-Access article distributed under the terms of the Creative Commons Attribution Non-Commercial License (http://creativecommons.org/licenses/by-nc/4.0) which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original work is properly cited.

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Comparative SNP Analysis of Chloroplast Genomes and 45S nrDNAs Reveals Genetic Diversity of Perilla Species
Plant Breed. Biotech.. 2018;6(2):125-139.   Published online June 1, 2018
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Comparative SNP Analysis of Chloroplast Genomes and 45S nrDNAs Reveals Genetic Diversity of Perilla Species
Plant Breed. Biotech.. 2018;6(2):125-139.   Published online June 1, 2018
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Comparative SNP Analysis of Chloroplast Genomes and 45S nrDNAs Reveals Genetic Diversity of Perilla Species
Image Image Image Image Image
Fig. 1 Chloroplast genome map of the nine Perilla species. Genes labeled inside the circle are transcribed clockwise and genes outside are transcribed counter-clockwise. Genes belonging to different functional groups are color coded. The dark gray ring corresponds to the GC content, while the light gray ring corresponds to the AT content.
Fig. 2 Assembly of the complete 45S nrDNA unit. The completed 45S nrDNA unit of P. citriodora, consisting of 18S rRNA, an internal transcribed spacer (ITS) 1, 5.8S rRNA, ITS 2, 26S rRNA, and the intergenic spacer region. The blue shading indicates the mapping depth of raw reads on the 45S nrDNA sequence of P. citriodora.
Fig. 3 Number of SNPs for different genes and the frequency of non-synonymous (ns) vs. synonymous (sy) SNPs. Interactive comparative analysis of the sequences of nine Perilla species showed that 33 genes contained 81 SNPs, including 44 sySNPs in 27 genes and 37 nsSNPs, which modify amino acid residues, in 15 genes. All genes of the transfer RNA and ribosomal RNA were conserved.
Fig. 4 Validation of SNP and InDel polymorphic sites using 4–8 individuals of each Perilla species. (A-C) SNP analysis of SNP positions in ndhF (A) using primer set CAPS-1 (SacII), in ycf1 (B) using primer set CAPS-2 (VspI), in ycf2 (C) using primer set CAPS-3 (AluI), and in ITS1 (D) using primer set CAPS-4 (BsmAI). (E) PCR analysis of InDel regions in trnKUUU-matK using primer set InDel-1. Abbreviated names on the gels indicate 100-bp marker (M), ‘Chajogi’ (CH), ‘Apureunchajogi’ (AC), ‘PureunJureumchajogi’ (PJC), ‘Jureumchajogi’ (JC), ‘Deulkkae’ (DK), ‘Pureunchajogi’ (PC), and negative control (N). Horizontal arrows represent 300 bp of the 100-bp marker.
Fig. 5 Phylogenomic analysis of chloroplast genomes of Perillas and representative species in the Lamiaceae family. The Olea europaea chloroplast genome was used as outgroup. GenBank accession numbers are in parentheses, and the number at an internal node represents the bootstrap value.
Comparative SNP Analysis of Chloroplast Genomes and 45S nrDNAs Reveals Genetic Diversity of Perilla Species

List of cultivated and wild Perilla species used in this study.

no Species Sample code Status GenBank Accession number

Chloroplast 45S nrDNA
1 Perilla citriodora P. citriodora Wild KT220690 KT220699
2 Perilla frutescens Britt. var. acuta Kudo ‘Chajogi’ Cultivated KT220685 KT220694
3 Perilla frutescens Britt. var. japonica Hara. for. discolor Makino ‘Apureunchajogi’ Cultivated KT220686 KT220695
4 Perilla frutescens Britt. var. crispa Hand.-Mazz. ‘PureunJureumchajogi’ Cultivated KT220687 KT220696
5 Perilla frutescens Britt. Var. crispa Hand.-Mazz. frutescens atropurpurea ‘Jureumchajogi’ Cultivated KT220688 KT220697
6 Perilla frutescens Britt. var. japonica Hara ‘Deulkkae’ Cultivated KT220689 KT220698
7 Perilla frutescens Britt. var. viridis Makino ‘Pureunchajogi’ Cultivated KT220684 KT220693
8 Perilla setoyensis P. setoyensis Wild KT220692 KT220701
9 Perilla frutescens var. hirtella P. hirtella Wild KT220691 KT220700

Chloroplast genome sizes of the nine Perilla species based on de novo assembly.

Sample Total Length (bp) LSCz) (bp) SSCy) (bp) IRax) (bp) IRbw) (bp)
P. citriodora 152,602 83,705 17,545 25,676 25,676
‘Chajogi’ 152,588 83,699 17,537 25,676 25,676
‘Apureunchajogi’ 152,598 83,701 17,545 25,676 25,676
‘PureunJureumchajogi’ 152,598 83,701 17,545 25,676 25,676
‘Jureumchajogi’ 152,598 83,701 17,545 25,676 25,676
‘Deulkkae’ 152,598 83,701 17,545 25,676 25,676
‘Pureunchajogi’ 152,588 83,699 17,537 25,676 25,676
P. setoyensis 152,607 83,710 17,547 25,675 25,675
P. hirtella 152,656 83,757 17,547 25,676 25,676

z)Large single copy,

y)Small single copy,

x)Inverted repeat region a,

w)Inverted repeat region b.

Chloroplast gene products of the nine Perilla species.

Group of genes Name of genes
Photosystem I psaA, B, C, I, J, ycf3y), ycf4
Photosystem II psbA, B, C, D, E, F, H, I, J, K, L, M, N, T, Z
Cytochrome b6/f petA, Bz), Dz), G, L, N
ATP synthase atpA, B, E, Fz), H, I
Rubisco rbcL
NADH oxidoreductase ndhAz), Bz,x), C, D, E, F, G, H, I, J, K
Large subunit ribosomal proteins rpl2z,x), 14, 16z), 20, 22, 23x), 32, 33, 36
Small subunit ribosomal proteins rps2, 3, 4, 7x), 8, 11, 12y,x,w), 14, 15, 16z), 18, 19
RNA polymerase rpoA, B, C1z), C2
Unknown function protein coding gene ycf1x), 2x), 15x)
Other genes accD, ccsA, cemA, clpPy) matK, InfA
Ribosomal RNAs rrn16x), 23x), 4.5x), 5x)
Transfer RNAs trnA-UGCz,x), trnC-GCA, trnD-GUC, trnE-UUC, trnF-GAA, trnG-UCCz), trnG-GCC, trnH-GUG, trnI-CAUx), trnI-GAUz,x), trnK-UUUz), trnL-UAAz), trnL-UAG, trnL-CAAx), trnfM-CAU, trnM-CAU, trnN-GUUx), trnP-UGG, trnQ-UUG, trnR-ACGx), trnR-UCU, trnS-GCU, trnS-GGA, trnS-UGA, trnT-GGU, trnT-UGU, trnV-UACz), trnV-GACx), trnW-CCA, trnY-GUA

z)Gene containing a single intron,

y)Gene containing two introns,

x)Two gene copies in IRs,

w)Trans-splicing gene.

Non-synonymous substitutions that modify amino acid residues among the nine Perilla species.

Gene Positionz) Samples

P. citriodora CHy) ACy) PJCy) JCy) DKy) PCy) P. setoyensis P. hirtella
accD 58337–58339 Phe (TTT)x) Phe (TTT) Phe (TTT) Phe (TTT) Phe (TTT) Phe (TTT) Phe (TTT) Ile (ATT) Phe (TTT)
ccsA 113262–113264 Ile (ATT) Ile (ATT) Ile (ATT) Ile (ATT) Ile (ATT) Ile (ATT) Ile (ATT) Ile (ATT) Phe (TTT)
clpP 70863–70865 Val (CTC) Val (CTC) Val (CTC) Val (CTC) Val (CTC) Val (CTC) Val (CTC) Ile (TTC) Val (CTC)
matK 3342–3344 Ser (GAA) Ser (GAA) Ser (GAA) Ser (GAA) Ser (GAA) Ser (GAA) Ser (GAA) Ser (GAA) Leu (AAA)
2130–2132 Phe (AAA) Phe (AAA) Phe (AAA) Phe (AAA) Phe (AAA) Phe (AAA) Phe (AAA) Phe (AAA) Ser (GAA)
2094–2096 Glu (TCC) Glu (TCC) Glu (TCC) Glu (TCC) Glu (TCC) Glu (TCC) Glu (TCC) Gly (CCC) Gly (CCC)
ndhF 110721–110723 Met (CAT) Met (CAT) Met (CAT) Met (CAT) Met (CAT) Met (CAT) Met (CAT) Ile (TAT) Met (CAT)
110195–110197 Arg (GAA) Arg (GAA) Arg (GAA) Arg (GAA) Arg (GAA) Arg (GAA) Arg (GAA) Arg (GAA) Cys (AAA)
110180–110182 Ser (TGT) Ser (TGT) Ser (TGT) Ser (TGT) Ser (TGT) Ser (TGT) Ser (TGT) Arg (GGT) Arg (GGT)
110068–110070 Tyr (TAA) Tyr (TAA) Tyr (TAA) Tyr (TAA) Tyr (TAA) Tyr (TAA) Tyr (TAA) Ser (GAA) Ser (GAA)
110021–110023 Lys (TTT) Lys (TTT) Lys (TTT) Lys (TTT) Lys (TTT) Lys (TTT) Lys (TTT) Gly (CCT) Lys (TTT)
110020–110022 Lys (TTT) Lys (TTT) Lys (TTT) Lys (TTT) Lys (TTT) Lys (TTT) Lys (TTT) Gly (CTT) Lys (TTT)
109895–109897 Val (CAA) Val (CAA) Val (CAA) Val (CAA) Val (CAA) Val (CAA) Val (CAA) Ile (TAA) Val (CAA)
109888–109890 Ser (GAT) Ser (GAT) Ser (GAT) Ser (GAT) Ser (GAT) Ser (GAT) Ser (GAT) Phe (AAT) Ser (GAT)
petB 75774–75776 Ile (ATT) Ile (ATT) Ile (ATT) Ile (ATT) Ile (ATT) Ile (ATT) Ile (ATT) Val (GTT) Val (GTT)
rpl22 83142–83144 Phe (GAA) Phe (GAA) Phe (GAA) Phe (GAA) Phe (GAA) Phe (GAA) Phe (GAA) Leu (TAA) Phe (GAA)
83135–83137 Gln (GAG) Lys (TAG) Lys (TAG) Lys (TAG) Lys (TAG) Lys (TAG) Lys (TAG) Gln (GAG) Gln (GAG)
rpoA 78559–78561 Glu (CTC) Glu (CTC) Glu (CTC) Glu (CTC) Glu (CTC) Glu (CTC) Glu (CTC) Asp (GTC) Asp (GTC)
77863–77865 Asp (ATC) Asp (ATC) Asp (ATC) Asp (ATC) Asp (ATC) Asp (ATC) Asp (ATC) Glu (CTC) Asp (ATC)
rpoC2 17958–17960 Leu (GGA) Leu (GGA) Leu (GGA) Leu (GGA) Leu (GGA) Leu (GGA) Leu (GGA) Leu (GGA) Phe (AGA)
17565–17567 Ile (TAA) Ile (TAA) Ile (TAA) Ile (TAA) Ile (TAA) Ile (TAA) Ile (TAA) Phe (AAA) Ile (TAA)
rps3 82916–82918 Ser (CTT) Ser (CTT) Ser (CTT) Ser (CTT) Ser (CTT) Ser (CTT) Ser (CTT) Ser (CTT) Asn (TTT)
rps8 79996–79998 Pro (GTC) Ala (CTC) Ala (CTC) Ala (CTC) Ala (CTC) Ala (CTC) Ala (CTC) Pro (GTC) Pro (GTC)
79982–79984 Leu (CAA) Leu (CAA) Leu (CAA) Leu (CAA) Leu (CAA) Leu (CAA) Leu (CAA) Phe (AAA) Leu (CAA)
rps16 4943–4945 Val (CAG) Val (CAG) Val (CAG) Val (CAG) Val (CAG) Val (CAG) Val (CAG) Leu (GAG) Val (CAG)
rps19 83719–83721 Asn (ATT) Asn (ATT) Asn (ATT) Asn (ATT) Asn (ATT) Asn (ATT) Asn (ATT) Lys (TTT) Asn (ATT)
109217–109219 Glu (GAA) Glu (GAA) Glu (GAA) Glu (GAA) Glu (GAA) Glu (GAA) Glu (GAA) Glu (GAA) Lys (AAA)
127091–127093 Glu (CTT) Glu (CTT) Glu (CTT) Glu (CTT) Glu (CTT) Glu (CTT) Glu (CTT) Glu (CTT) Lys (TTT)
126527–126529 Lys (TTT) Lys (TTT) Lys (TTT) Lys (TTT) Lys (TTT) Lys (TTT) Lys (TTT) Glu (CTT) Glu (CTT)
ycf1 125843–125845 Ala (CTC) Ala (CTC) Ala (CTC) Ala (CTC) Ala (CTC) Ala (CTC) Ala (CTC) Thr (TTC) Ala (CTC)
123676–123678 Thr (GTA) Thr (GTA) Thr (GTA) Thr (GTA) Thr (GTA) Thr (GTA) Thr (GTA) Thr (GTA) Ile (ATA)
123615–123617 Glu (CTC) Glu (CTC) Glu (CTC) Glu (CTC) Glu (CTC) Glu (CTC) Glu (CTC) Asp (ATC) Glu (CTC)
122632–122634 Ile (ATG) Ile (ATG) Ile (ATG) Ile (ATG) Ile (ATG) Ile (ATG) Ile (ATG) Ser (CTG) Ser (CTG)
ycf2 88843–88845 Met (TGA) Met (TGA) Met (TGA) Met (TGA) Met (TGA) Met (TGA) Met (TGA) Thr (CGA) Met (TGA)
147465–147467 Met (ATC) Met (ATC) Met (ATC) Met (ATC) Met (ATC) Met (ATC) Met (ATC) Thr (GTC) Met (ATC)
90151–90153 Val (TTG) Val (TTG) Val (TTG) Val (TTG) Val (TTG) Val (TTG) Val (TTG) Ala (CTG) Val (TTG)
146157–146159 Val (ACT) Val (ACT) Val (ACT) Val (ACT) Val (ACT) Val (ACT) Val (ACT) Ala (GCT) Val (ACT)

z)The positions of SNPs were based on the P. citriodora sequence.

y)CH, ‘Chajogi’; AC, ‘Apureunchajogi’; PJC, ‘PureunJureumchajogi’; JC, ‘Jureumchajogi’; DK, ‘Deulkkae’; PC, ‘Pureunchajogi’.

x)Amino acid (nucleotide).

Variable number of tandem repeats present in the nine Perilla species.

Region Location Size (bp) Copy number variation Consensus

P. citriodora, ACz), PJCz), JCz), DKz) CHz) PCz) P. setoyensis P. hirtella
LSCy) trnKUUU - matK intergenic 27 1 1 2 2 TTCAGAATAGAAATAGGGGAGATCCCA
LSC atpH - atpI intergenic 1 14 14 16 15 T
LSC petA - psbJ intergenic 7 1 1 1 2 ATTATAT
LSC petB intron 6 2 2 2 3 AAAGAA
SSCx) rpl33 - trnLUAG intergenic 8 2 1 2 2 CAAATTCC

z)CH, ‘Chajogi’; AC, ‘Apureunchajogi’; PJC, ‘PureunJureumchajogi’; JC, ‘Jureumchajogi’; DK, ‘Deulkkae’; PC, ‘Pureunchajogi’.

y)Large single copy.

x)Small single copy.

Summary of nucleotide polymorphisms in 45S nrDNA sequences of the nine Perilla species.

Type Locus Nucleotide Positionz) 9 perilla species

P. citriodora CHy) ACy) PJCy) JCy) DKy) PCy) P. setoyensis P. hirtella
SNP 18S rRNA 706 A G G G G G G G A
18S rRNA 1064 T A A A A A A A T
ITS1 1915 T G G G G G G G G
ITS1 1974 A C C C C C C C C
ITS1 1981 C T T T T T T T C
ITS2 2263 T C C C C C C C C
ITS2 2264 C T T T T T T C C
ITS2 2369 C T T T T T T T C
ITS2 2373 C A A A A A A A C
ITS2 2408 C C C C C C C C T
ITS2 2409 A C C C C C C C C
26S rRNA 2971 C T T T T T T T C
26S rRNA 2987 C T T T T T T T C
26S rRNA 4252 C C C C C T T C C
26S rRNA 5642 T T T T T T T T C
26S rRNA 5660 T A A A A A A A T
InDel ITS1 1864 -x) G G G G G G - G
ITS1 2006 G G G G G G G - G

z)The positions of SNPs were based on the P. citriodora sequence.

y)CH, ‘Chajogi’; AC, ‘Apureunchajogi’; PJC, ‘PureunJureumchajogi’; JC, ‘Jureumchajogi’; DK, ‘Deulkkae’; PC, ‘Pureunchajogi’.

x)No detection.

Table 1 List of cultivated and wild Perilla species used in this study.
Table 2 Chloroplast genome sizes of the nine Perilla species based on de novo assembly.

Large single copy,

Small single copy,

Inverted repeat region a,

Inverted repeat region b.

Table 3 Chloroplast gene products of the nine Perilla species.

Gene containing a single intron,

Gene containing two introns,

Two gene copies in IRs,

Trans-splicing gene.

Table 4 Non-synonymous substitutions that modify amino acid residues among the nine Perilla species.

The positions of SNPs were based on the P. citriodora sequence.

CH, ‘Chajogi’; AC, ‘Apureunchajogi’; PJC, ‘PureunJureumchajogi’; JC, ‘Jureumchajogi’; DK, ‘Deulkkae’; PC, ‘Pureunchajogi’.

Amino acid (nucleotide).

Table 5 Variable number of tandem repeats present in the nine Perilla species.

CH, ‘Chajogi’; AC, ‘Apureunchajogi’; PJC, ‘PureunJureumchajogi’; JC, ‘Jureumchajogi’; DK, ‘Deulkkae’; PC, ‘Pureunchajogi’.

Large single copy.

Small single copy.

Table 6 Summary of nucleotide polymorphisms in 45S nrDNA sequences of the nine Perilla species.

The positions of SNPs were based on the P. citriodora sequence.

CH, ‘Chajogi’; AC, ‘Apureunchajogi’; PJC, ‘PureunJureumchajogi’; JC, ‘Jureumchajogi’; DK, ‘Deulkkae’; PC, ‘Pureunchajogi’.

No detection.