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들깨 종간 교잡( × ) 집단의 양적 형질 유전자좌 분석

Detection of QTLs in an Interspecific Cross between Perilla citriodora × P. hirtella Mapping Population

Myoung Hee Lee1, Ki Won Oh1,*, Myung Sik Kim2, Sung Up Kim1, Jung In Kim1, Eun Young Oh1, Suk Bok Pae1, Un Sang Yeo1, Tae-Ho Kim3, Jeong Hee Lee4, Chan Sik Jung1, Do Yeon Kwak1, Yong Chul Kim1
Korean Journal of Breeding Science 2018;50(1):13-20.
Published online: February 28, 2018

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*Corresponding Author (ohkw1004@korea.kr, +82-55-350-1212, +82-55-353-3050)
• Received: September 19, 2017   • Accepted: November 24, 2017

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  • An interspecific cross between P. citriodora and P. hirtella constitutes ideal material for a linkage map construction in genome project of Perilla. The chromosomes of the species are same with n=10 and progenies of the cross are normal in growth and seed set. The phenotype of F2 population of the cross are normally distributed and this is a proof of high affinity of the chromosomes during their sexual reproduction. Rosmarinic acid, Luteolin and Apigenin contents of F2 plants were distributed in similar range with tetraploid perilla cultivars. Luteolin and Apigenin are positively correlated with correlation coefficient of 0.762. 21 QTLs were detected in agronomic traits and Rosmarinic acid, Luteolin and Apigenin contents. Even though Luteolin and Apigenin are positively correlated, QTLs were located in different position. Purple leaf color related QTL was mapped in Chromosome 3 with LOD of 14.3, PVE of 50.4%. Three anthocyanin biosynthesis transcription factor like sequences, WD40 repeat-like superfamily protein, myb domain protein 43 and basic helix-loop-helix (bHLH) DNA-binding superfamily protein, were detected near from the QTL.
들깨(Perilla frutescens Britt.)는 우리나라의 전통적인 유지작 물로 재배되어 오고 있으며 한국을 포함한 중국, 네팔, 베트남, 일본, 대만, 라오스 등 다양한 지역에서 재배종 및 야생종 자원이 보고되어 있다(Hu et al. 2010, Ito et al. 2008, Jung et al. 2005, Lee & Ohnishi 2001, Nitta et al. 2005, Nitta et al. 2003, Pandey & Bhatt 2008, Woo et al. 2016). 들깨 속 식물에 는 우리가 일반적으로 들깨로 부르고 있는 하나의 재배종 4배체 (P. frutescens, 2n=40)와 3종의 2배체 야생종인 P. citriodora (2n=20), P. setoyensis(2n=20), P. hirtella(2n=20)가 보고되어 있다(Nitta et al. 2005).
들깨는 다양한 지역에 분포하고 있으나 이용 방법에는 나라별 로 차이가 있는데 우리나라에서는 들기름과 깻잎으로 이용되고 있으며 농업적 가치가 매우 큰 반면 일본에서는 차조기를 많이 이용하고 있지만 들기름 소비는 많지 않다. 중국에서는 우리나라 로 수출을 목적으로 들깨를 재배하고 있으며 배트남에서는 깻잎을 차로 이용하고 있다고 알려져 있다. 국내의 들깨 재배면적은 최근 지속적으로 증가하여 2010년 33,376ha에서 2016년 45,474ha 까지 증가하여 재배면적 기준으로 국내에서 3번째로 많이 재배 되었다(2017, KOSIS). 이와 같이 농업적 중요성이 커짐에 따라 농촌진흥청 국립식량과학원에서는 착유 용으로 이용되는 종실 용 들깨 품종과 깻잎으로 이용되는 잎들깨 품종을 지속적으로 개발하고 있으며 들깨와 관련된 다양한 분야의 연구를 지속하고 있다(Jung et al. 2006, Lee et al. 2014, Lee et al. 2011).
최근 들깨 유전체 정보 해독을 위해 연구가 진행중에 있으며 2배체 야생종인 P. citriodora를 해독 재료로 이용하고 있다. 이 연구 과정에서 P. citriodora의 엽록체 정보가 해독되어, 총 길이가 152,588bp로 보고되었고(Mo et al. 2016), 발현체 정보 로부터 신규 유전자를 분석한 결과 기존에 알려진 유전자와 유사성이 높아 P. citriodora유전체 해독에 유용한 정보가 보고된 바 있다(Kim et al. 2016b). 4배체 들깨는 식물성 오메가-3 지방 산인 Linolenic acid 비율이 60% 이상으로 높아 기름의 기능과 이용에 대한 관심이 높아 종실의 발달 과정에서 지방산 생합성 관련 유전자 발현(Kim et al. 2016)과 더불어 cDNA Library 작성 등 발현체 중심으로 연구가 진행되고 있다(Fukushima et al. 2015, Seong et al. 2015).
본 연구에서는 P. citriodora 유전체 해독에 필요한 매핑 집단 을 육성하기 위해 종간 교잡을 실시하고 이로부터 분리 세대인 F2 집단을 육성하여 이를 이용하여 주요한 농업 형질에 대한 양적형질유전자좌를 분석한 결과에 대하여 보고하고자 한다.
들깨 신규 유전체 해독을 위한 매핑 집단 육성을 위해 모본은 레몬들깨(P. citriodora)를, 부본은 범꼬리소엽(P. hirtella)을 이 용하였다. P. citriodora종내 교잡을 통해 집단을 육성하는 것이 일반적인 방법이지만 국내에서 수집된 유전자원이 1점밖에 보고 되어 있지 않아 부득이 하게 종간 교잡을 통해 매핑 집단을 육성하였다. 종간교잡에 사용된 레몬들깨는 강원대학교 응용식 물과학과 이주경 교수로부터 분양받은 재료를 사용하였고, 범꼬 리소엽은 제주도 비자림에서 수집된 제주-17(Jung et al. 2005) 을 사용하였다. 2015년 6월 파종하여 재배 한 후 9월 27일 개화 하여 인공 교배를 실시하였다. 인공 교배 결과 30개의 꽃에서 4개의 교배립을 얻었다. 교배립은 11월에 온실에 파종하였으며 이듬해 1월까지는 밤 10시부터 4시간 동안 조명하여 충분한 영양생장이 되도록 하였으며 2월부터 야간조명을 중단하여 개화 를 유도하여 3월에 개화하였고 4월 중순에 F2 종자를 채종하였 다. 채종된 종자는 2016년 6월에 상순에 육묘 포트에 파종하여 육묘하고 6월 25일에 이랑 넓이 70cm, 포기간격 25cm로 250개 체를 이식하였다. 경장, 마디수, 분지수, 화방군수, 화방군장, 화방군당 삭수 등 주요 농업 형질은 농촌진흥청 조사 기준에 따라 조사하였으며, 엽색은 잎 뒷면이 자색, 녹색, 연한 자색으로 구분하여 조사하였다.
Rosmarinic acid, Luteolin, Apigenin 분석을 위해 종자 시료 1g을 분쇄하여 80% methanol 30 ml에 상온에서 3시간동안 추출한 후 0.45μm 필터로(Millipore, MSI, Westborom MA, US) 필터링한 후 HPLC로 분석하였다(Lee et al. 2009). HPLC 분석 조건은 Dionex Ultimate 3000 series(Dionex Softron GmbH, Germering, Germany), 역상 C-18 컬럼(150 mm × 4.6 mm, Eclipse XDB-C18, 5μm, Agilent, USA), 이동상 (A) 0.1% acetic acid in water, 이동상 (B) 0.1% acetic acid in acetonitrile 으로 하여 변화 조건은 0 min, 5% B; 5 min, 10% B; 10 min, 20% B 15 min, 25% B; 20 min, 30% B; 30 min, 50% B; 40 min, 65% B로 하였다. 기타 분석 조건은 유속 1.0 mL/min; 컬럼 온도 30℃; 검출 파장 330nm; 시료 주입량 20μl로 하였다.
유전지도 작성에는 F2 집단 250개체 중 96개체를 이용하였다. 유전지도 작성은 농촌진흥청 포스트게놈 다부처유전체사업의 ‘들깨 유전체 해독 소재 구축 및 유전체 해독’ 과제를 통해 ㈜씨더 스에서 수행 하였으며, GBS Library를 구축한 후 이로부터 분석 된 2518개의 SNP 마커를 이용하여 유전지도를 작성하였고 10개 의 염색체에서 총 1309cM의 영역이 분석되었다(결과 미발표).
조사자료는 SAS 9.2를 이용하여 상관분석과 기본통계 분석을 실시하였다. 엽색의 경우 적합도검정을 통해 분리비 검정을 실시 하였고 SNP마커 유전지도 상의 엽색의 유전지도 작성을 위해서 QTL IciMapping Ver. 4.0(http://www.isbreeding.net)을 이용하 여 LOD 3.0 조건에서 유전자지도 분석을 수행하였다. QTL 분석 은 QTL IciMapping Ver. 4.0을 이용하여 LOD 3.0의 조건에서 inclusive composite interval mapping을 수행하였다. 분석된 QTL 중 엽색과 관련된 QTL 주변의 유전자 분석은 ㈜씨더스에서 수행하였다. P. citriodora의 영양생장기 잎에서 추출한 RNA로부 터 얻어진 발현체 분석 정보(Kim et al. 2016)를 P. citriodora 염기서열 분석 정보와 비교하여 QTL 주변 SNP 마커가 위치하는 Scaffold에 자리하는 발현체 목록을 만든 후 P. citriodora 전사체 정보를 Phytozome(https://phytozome.jgi.doe.gov)에 등록되어 있는 다른 식물 유전자 정보와 비교하여 분석하였다.
주요 농업 형질 및 성분 12개 형질에 대한 분포도를 비교하였 다(Fig. 1). 경장, 마디수, 분지수, 화방군장, 화방군당 삭수, 종실 중, Rosmarinic acid 함량, Luteolin 함량, Apigenin 함량 등 대부분의 양적 형질은 평균을 중심으로 좌우로 고르게 분포되었 고, 모본(P. citriodora)과 부본(P. hirtella)의 값 보다 넓은 범위에 서 조사되었다. 개체당 화방군수는 중위수가 평균보다 작아 왜도 가 4.81로 양의 수를 보여 분포의 꼬리가 오른쪽으로 길게 나타난 반면 나머지 형질들은 왜도가 작고 첨도가 크지 않아 정규분포에 가까운 모양으로 분포하였다(Table 1). 이와 같은 형질의 분포는 실험재료로 사용된 F2 집단의 개체들이 종간 교잡 후 세대를 거치는 유성생식 과정에서 염색체 재조환이 정상적으로 진행되 었음을 알 수 있다. Perilla속 식물의 종간 교잡에 대한 선행연구 결과에 따르면 4배체로 알려진 재배종 들깨 P. frutescens(2n=40) 와 야생종 2배체 P. citriodora(2n=-20)는 종간 교잡을 통해 3배 체(2n=30)인 F1 식물체가 만들어 지지만 감수분열 과정에서 10개의 2가염색체와 10개의 1가염색체가 형성되어 정상적인 감수분열을 하지 못해 불임이 된다(Honda et al. 1994). 본 실험 에서 수행한 종간 교잡 에서 F1 식물체 생존율은 12.5%(종간 교잡 종자 30개 중 4개체 생존)로서 F1 개체 생존율이 12.6%라 고 보고한 선행연구결과(Ito and Honda 1999)와 비슷하였다. 들깨 속의 2배체간 종간 교잡 F1으로부터 세대를 진전시킨 선행 연구결과는 없으나 콜히친 처리를 통해 P. citriodora × P. hirtella 종간 교잡 F1 식물체의 염색체를 배가하여 육성한 복 이배체의 결실율은 0.9~41.4% 까지 다양하였다(Ito and Honda 1999).
F2 집단, 모본 및 부분의 종자에서 Rosmarinic acid, Luteolin, Apigenin 등 3종의 성분을 분석한 결과 Fig. 1의 (h), (i), (j)에서 보는 바와 같이 모본(P1)인 P. citriodora의 함량이 부본(P2)인 P. hirtella 보다 상당히 높았다. F2 집단에서는 Rosmarinic acid 는 886~2,503ug/g, Luteolin은 13~406 ug/g, 그리고 Apigenin은 61~534 ug/g 사이에 분포하였다(Table 1). 재배종 4배체인 들깨 유전자원의 함량은 Rosmarinic acid는 16~2,717ug/g, Luteolin 은 2~582 ug/g, 그리고 Apigenin은 6~476 ug/g 사이에 분포하는 것으로 보고되어(Lee et al. 2009) 본 실험의 결과와 유사한 경향이었다.
Fig. 1.
Phenotypic variation in F2 population and their parents. P1 and P2 represent maternal and paternal parent respectively.
KJBS-50-13_F1.jpg
Table 1
Phenotypic variation in F2 population of a cross P. citriodora × P. hirtella.
Table 1
Trait Mean SE Min Max Skewness Kurtosis
SL(Stem Length, cm) 91 19.3 35 135 0.09 0.09
NN(Node Number) 12.1 1.9 7 18 0.31 1.03
BN(Branch Number) 8.5 3.3 1 18 0.41 0.64
FCN(Flower Cluster Number) 234 225.7 33 512 4.81 26.88
FCL(Flower Cluster Length, cm) 13.95 3.47 5.6 24.1 -0.01 0.15
FN(Flower No./Flower Cluster) 51 9.8 28 73 -0.32 -0.58
SW(Seed Weight) 8.3 5.02 0.1 22.0 0.52 -0.41
RA(Rosemarinic Acid, ug/g) 1431 294 886 2503 0.70 0.94
LU(Luteolin, ug/g) 176 81 13 406 0.43 -0.18
AP(Apigenin, ug/g) 250 96 61 534 0.55 -0.05
주요 농업 형질 및 기능성 성분 간의 상관관계 분석결과 농업형 질들은 대부분 상호간에 정의 상관이 인정되었다(Table 2). 경장 은 마디수, 분지수, 화방군수, 화방군장, 화방군당 삭수 및 종실중 등 모든 형질과 유의한 정의 상관관계를 보였고, 마디수는 분지수 및 화방군수와 정의 상관관계를, 분지수는 화방군수, 화방군당 삭수 및 종실중과 정의 상관관계를, 화방군수는 화방군장, 화방 군당 삭수 및 종실중과 정의 상관관계를, 화방군장은 화방군당 삭수와 정의 상관관계를 보였는데 이는 생육이 강한 개체가 키가 크고 분지가 많으며 이에 따라 화방군이 많이 형성되고 종실중이 많아진 때문이었다. 이차대사산물인 Rosmarinic acid, Apigenin, Luteolin의 상관관계를 분석한 결과 Apigenin과 Luteolin 간에 0.762의 높은 정의 상관이 있었다. 이는 들깨 유전자원에서 이 두 물질 간에 0.81의 높은 정의 상관이 있었다는 선행 연구결과와 일치하였다(Lee et al. 2009). Flavone 생합성 대사과정에서 두 물질의 전구 물질인 Naringenin에 Flavone synthase II (FNSII)가 이중결합을 하나 추가하면 Apigenin이 합성되고, Flavonoid 3’-hydroxylase (F3’H)가 작용하여 3’의 H를 OH로 치환하여 Eriodictyol을 합성한 후 FNSII가 작용하여 이중결합을 추가하면 Luteolin이 합성된다(Mizuno et al. 2016). 이와 같이 Luteolin과 Apigenin은 3’ 탄소에 결합된 OH또는 H간의 차이만 있어 분자구조가 유사하고, 대사과정도 아주 밀접 하게 연관되어 있기 때문에 두 물질의 함량 간에 높은 상관관계가 있는 것으로 보인다.
Table 2
Phenotypic Pearson correlations among traits in F2 populationz.
Table 2
Trait SL NN BN FCN FCL FN SW RA LU
NN 0.505**
BN 0.622** 0.622**
FCN 0.432** 0.343** 0.57**
FCL 0.3** 0.014 0.119 0.304**
FN 0.323** 0.114 0.224* 0.428** 0.747**
SW 0.298** 0.076 0.293** 0.558** 0.204 0.135
RA -0.082 -0.092 -0.234* -0.057 0.262* 0.125 0.029
LU 0.066 -0.022 0.070 -0.071 -0.257* -0.311** 0.107 -0.084
AP -0.087 0.026 -0.024 -0.243* -0.27* -0.35** -0.025 0.052 0.762**

*Correlation is significant at 0.05 and 0.01 probability levels, respectively.

**Correlation is significant at 0.05 and 0.01 probability levels, respectively.

zAbbreviations of traits are same with Table 1.

주요 농업형질 및 기능 성분과 관련된 양적 형질 유전자좌를 분석한 결과 21개의 QTL을 찾았다. 잎 뒷면 엽색(LC)은 3번 염색체에서 하나의 QTL이 존재하였고, 모용(PB)의 밀도는 2번 과 5번 염색체에서 각각 2개, 경장(SL)은 9번 염색체에서 1개, 화방군수(FCN)는 3번 염색체에서 1개, 화방군장(FCL)은 1번과 3번 염색체에서 각각 1개, 종실중(SW)은 10번염색체에서 1개, Rosmarinic acid는 1, 5, 6번 염색체에 각각 1개, 7번 염색체에 2개, Luteolin은 1번과 10번 염색체에 각각 1개, Apigenin은 1번과 4번 염색체에 각각 1개의 QTL이 분석되었다(Table 3).
Table 3
Results of composite interval mapping for agronomic traits in F2.
Table 3
Traitu Chr. Left Marker Right Marker Position (cM) LODz PVE(%)y Addx Domw |d/a|v
LC 3 scaffold8_17030380 scaffold8_16101189 95.8 14.3 50.4 0.66 0.15 PD
PB 2 scaffold25_4598423 scaffold25_3912693 57.9 10.0 25.4 0.54 0.06 A
PB 2 scaffold25_1773512 scaffold25_744098 76.2 7.8 19.1 0.05 -0.64 OD
PB 5 scaffold3_9875772 scaffold3_11292324 38.0 5.4 12.2 0.39 0.12 PD
PB 5 scaffold64_1191653 scaffold22_5567527 100.4 3.3 7.1 -0.28 -0.03 A
SL 9 scaffold35_5711921 scaffold134_133385 13.9 3.8 20.1 9.79 -14.67 OD
FCN 3 scaffold31_6872402 scaffold31_6250465 40.9 3.8 19.0 -54.43 -3.61 A
FCL 1 scaffold32_5830494 scaffold29_6732047 13.0 3.2 15.2 -1.64 1.04 PD
FCL 3 scaffold49_35668 scaffold6_4743200 63.4 4.0 17.8 -1.92 0.21 A
FN 3 scaffold6_9130731 scaffold21_10557309 58.2 5.1 23.8 -5.00 2.14 PD
SW 10 scaffold28_5887083 scaffold28_6525916 24.2 3.7 19.6 3.13 -1.60 PD
RA 1 scaffold16_7569624 scaffold16_7019064 150.9 4.2 8.3 -115.90 0.47 A
RA 5 scaffold10_10061999 scaffold10_9181391 4.0 7.1 14.8 171.59 47.89 PD
RA 6 scaffold43_1927956 scaffold43_2688122 34.6 3.7 7.2 103.81 27.49 PD
RA 7 scaffold59_753444 scaffold59_2135864 83.7 4.0 7.8 -27.91 -134.19 OD
RA 7 scaffold34_6133629 scaffold34_6235270 156.9 3.4 6.3 108.83 -31.67 PD
RA 10 scaffold12_7503648 scaffold12_9560702 2.1 12.8 32.1 -229.27 11.30 A
LU 1 scaffold46_600664 scaffold46_534033 191.8 5.2 22.4 59.15 -10.69 A
LU 10 scaffold32_1386668 scaffold32_1454570 21.8 3.3 13.6 44.55 -6.67 A
AP 1 scaffold2_12959065 scaffold2_13146855 89.8 3.8 12.6 46.97 26.22 PD
AP 4 scaffold24_857542 scaffold24_1017077 77.3 5.8 21.0 -6.65 90.29 OD

zLOD score for the detected QTLs (LOD threshold=3.0).

yproportion of variance explained by a QTL at the test site under H0:H3.

xAdditive effect under H1.

wDominant effect under H2.

vestimation of gene action, A (additive effect) 0-0.20, PD (partial dominance) 0.21-0.80, D (dominance) 0.81-1.20, OD (over dominance)>1.20.

uAbbreviations ; LC (Leaf Color), PB (Pubescence), others are same with Table 1.

생육과 관련된 특성 중 마디수와 분지수는 유의한 수준의 QTL이 나타나지 않았고 경장의 경우 1개의 QTL이 분석되었다. Luteolin과 Apigenin의 함량 간에는 높은 정의 상관관계가 있어 QTL의 위치 또한 가까이 위치할 것으로 예상되었으나 두 성분의 함량과 관련된 QTL이 같이 위치한 1번 염색체 내에서도 Apigenin 의 QTL은 89.8cM에 있는 반면 Luteolin의 QTL은 191.8cM에 자리하고 있어 상당한 거리가 있는 것으로 분석되었다. 엽색(LC) 의 분리비를 조사한 결과 자색, 헤테로(연한자색), 녹색이 각각 14, 62, 20개체로 조사되어(Fig. 1) 적합도검정 결과 1:2:1의 분리비에 적합하였다. 엽색의 유전지도상의 위치를 조사한 결과 3번 염색체의 111.4cM에 위치하였다(자료 미표시). 반면 엽색 을 양적형질로 가정하여 녹색, 헤테로(연한자색), 자색을 각각 0, 1, 2로 값을 준 후 분석한 경우 QTL은 3번 염색체의 95.8cM에 위치하여 질적형질의 위치와 큰 차이를 보이지 않았다. 엽색(LC) 을 양적형질로 분석하였을 때 QTL은 3번 염색체에 하나만 존재 하였는데 LOD가 14.3으으로 매우 높고 표현형 변이에 대한 QTL의 설명 비율이 50.4%로 높은 부분 우성 형질이었다(Table 3). 엽색 유전자 주변에 위치한 P. citriodora 전사체 정보를 분석한 결과 안토시아닌 생합성 조절인자로 알려진 basic helix-loop-helix (bHLH) DNA-binding superfamily protein, myb domain protein 43, Transducin/WD40 repeat-like superfamily protein이 확인 되었으며 이들 유전자는 비교대상 유전자와 염기서열 유사성이 각각 72.25, 73.06, 93.39%였다(Table 4). 식물의 안토시아닌 생합성 과정의 후기 생합성 관련 유전자의 발현은 MYB, Basic helix-loop-helix(bHLH) 그리고 WD40 전사인자 등 3가지 요소 로 구성된 WD40-bHLH-MYB(WBM) 복합체에 의해 조절된다 (Shi and Xie 2014). PAP1/MYB75, PAP2/MYB90, MYB113 그리고 MYB114 등 염기서열이 서로 유사한 MYB 전사인자는 식물 조직의 안토시아닌 생합성을 조절하는 것으로 밝혀져 있으 며, 이들 유전자는 R2R3-MYB type 단백질로서 두개의 불완전 한 반복 서열을 MYB domain에 가지고 있다(Dubos et al. 2010, Stracke et al. 2001). PAP1또는 PAP2 도메인의 MYB113, MYB114를 과다 발현 시키면 식물체의 안토시아닌 함량이 증가 하는 반면, 이들 유전자 발현을 억제하면 안토시아닌이 결핍된다 (Borevitz et al. 2000, Gonzalez et al. 2008). bHLH 전사인자는 GL3 (Glabra 3), EGL3 (Enhancer of Glabra 3) 그리고 TT8 (Transparent testa 8) 등 3가지 유전자로 구성되어 있으며 이들 유전자는 안토시아닌 생합성을 촉진시키는 방향으로 조절하는 반면 WD40 도메인에는 TTG1 유전자만이 알려져 있고 이 유전 자가 결핍되면 식물체의 안토시아닌 결핍이 발생한다(Shi & Xie 2014). 이상에서 설명한 WBM 복합체는 구성요소에 따라 다른 형태로 존재하며 이에 따라 안토시아닌 생합성을 활성화시 키는 반면, 1R-MYB type 단백질인 CPC와 MYBL2 등은 WBM 복합체의 활동을 억제하여 안토시아닌 생합성을 감소시킨다. 즉 촉진 조절 인자인 PAP1/PAP2/MYB113/MYB114와 감소 조절 인자인 CPC/MYBL2 간의 양적 경쟁이 안토시아닌 생합성 경로의 유전자 활성화 또는 억제에 영향을 미치는 것으로 보인다 (Shi & Xie 2014). 엽색은 색의 발현 유무에 따라 질적형질로 볼 수도 있으나 온도, 광도 등 환경에 따라 발현 강도가 달라지기 도 하여 양적형질로 판단할 수도 있을 것으로 판단되며 들깨의 경우 잎의 앞면, 뒷면, 줄기, 꽃 등 다양한 식물체 부위에 안토시아 닌이 발현되고 각 부위의 유전자 발현은 독립적으로 조절되기 때문에 엽색의 유전과 관련하여 더 많은 연구가 필요하다.
Table 4
Anthocyanin biosynthesis transcription factor like sequences located near from Leaf Color (LC) QTL of Chromosome 3.
Table 4
Transcript No. Speciesz Sequence Identity TAIRy TAIR description
1SL023156t001 Mimulus guttatus v1.1 72.25 AT1G72210.1 basic helix-loop-helix (bHLH) DNA-binding superfamily protein
1SL003771t001 Vitis vinifera 73.06 AT5G16600.1 myb domain protein 43
1SL006007t004 Mimulus guttatus v2.0 93.39 AT5G66240.2 Transducin/WD40 repeat-like superfamily protein

zMost similar sequence DB with P. citriodora transcript sequence.

yLocus number in TAIR(The Arabidopsis Information Resource, http://www.arabidopsis.org).

본 연구에 사용한 F2 집단은 들깨 야생종 2배체(P. citriodora) 표준유전체 분석을 위한 과정에서 사용된 시험재료로서 연구과 제의 목적 달성을 위해 유전지도작성이 필요하여 부득이하게 보통의 경우 양적형질 분석에 사용하지 않는 F2 세대에서 분석을 수행하였으나 현재 F5세대까지 RIL이 진전되어 있으며 향후 고세대까지 세대를 진전시킨 후 농업형질과 더불어 주요 형질에 대한 유전분석을 실시할 계획이다.
레몬들깨(P. citriodora)와 범꼬리소엽(P. hirtella) 등 2종의 2배체 야생 들깨 종간 교잡을 통해 들깨 유전체 사업의 연관 유전지도 작성에 필요한 집단을 육성하였다. 두 종의 염색체 수는 n=10으로 같고 종간 교잡 후대는 세대를 거치는 동안 정상 적인 생육과 결실을 보였다. F2 집단의 표현형은 정규분포를 보여 종간 교잡 식물체의 감수분열 과정에서 두 종의 염색체가 친화성이 높은 것으로 보였다. F2 집단으로부터 수확한 종자에서 Rosmarinic acid, Luteolin, Apigenin등 3종의 성분을 분석한 결과 4배체 재배종 들깨의 함량과 유사한 범위에 분포하였고 Apigenin과 Luteolin 간에 0.762의 높은 정의 상관이 있었다. 주요 농업형질 및 기능성분 분석 결과 21개의 QTL을 찾았다. Apigenin과 Luteolin 간에 높은 정상관이 있었으나 QTL 영역은 다르게 조사되었다. 엽색(LC)의 QTL은 3번 염색체에 하나만 존재하였는데 LOD가 14.3으으로 매우 높고 표현형 변이에 대한 QTL의 설명 비율이 50.4%로 높은 부분 우성 형질이었다. 또한 이 QTL 주변 영역에는 안토시아닌 생합성 조절인자로 알려진 basic helix-loop-helix(bHLH) DNA-binding superfamily protein, myb domain protein 43, Transducin/WD40 repeat-like superfamily protein이 확인되었다.
본 논문은 농촌진흥청 연구사업(세부과제명: 들깨 유전체 해 독소재 개발 및 매핑 집단 작성, 세부과제번호: PJ01040802)의 지원에 의해 이루어진 결과의 일부입니다.
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Detection of QTLs in an Interspecific Cross between Perilla citriodora × P. hirtella Mapping Population
Korean. J. Breed. Sci.. 2018;50(1):13-20.   Published online March 1, 2018
DOI: https://doi.org/10.9787/KJBS.2018.50.1.13
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Detection of QTLs in an Interspecific Cross between Perilla citriodora × P. hirtella Mapping Population
Korean. J. Breed. Sci.. 2018;50(1):13-20.   Published online March 1, 2018
DOI: https://doi.org/10.9787/KJBS.2018.50.1.13
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Detection of QTLs in an Interspecific Cross between Perilla citriodora × P. hirtella Mapping Population
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Fig. 1. Phenotypic variation in F2 population and their parents. P1 and P2 represent maternal and paternal parent respectively.

Fig. 1.

Detection of QTLs in an Interspecific Cross between Perilla citriodora × P. hirtella Mapping Population

Phenotypic variation in F2 population of a cross P. citriodora × P. hirtella.

Trait Mean SE Min Max Skewness Kurtosis
SL(Stem Length, cm) 91 19.3 35 135 0.09 0.09
NN(Node Number) 12.1 1.9 7 18 0.31 1.03
BN(Branch Number) 8.5 3.3 1 18 0.41 0.64
FCN(Flower Cluster Number) 234 225.7 33 512 4.81 26.88
FCL(Flower Cluster Length, cm) 13.95 3.47 5.6 24.1 -0.01 0.15
FN(Flower No./Flower Cluster) 51 9.8 28 73 -0.32 -0.58
SW(Seed Weight) 8.3 5.02 0.1 22.0 0.52 -0.41
RA(Rosemarinic Acid, ug/g) 1431 294 886 2503 0.70 0.94
LU(Luteolin, ug/g) 176 81 13 406 0.43 -0.18
AP(Apigenin, ug/g) 250 96 61 534 0.55 -0.05

Phenotypic Pearson correlations among traits in F2 populationz.

Trait SL NN BN FCN FCL FN SW RA LU
NN 0.505**
BN 0.622** 0.622**
FCN 0.432** 0.343** 0.57**
FCL 0.3** 0.014 0.119 0.304**
FN 0.323** 0.114 0.224* 0.428** 0.747**
SW 0.298** 0.076 0.293** 0.558** 0.204 0.135
RA -0.082 -0.092 -0.234* -0.057 0.262* 0.125 0.029
LU 0.066 -0.022 0.070 -0.071 -0.257* -0.311** 0.107 -0.084
AP -0.087 0.026 -0.024 -0.243* -0.27* -0.35** -0.025 0.052 0.762**

*Correlation is significant at 0.05 and 0.01 probability levels, respectively.

**Correlation is significant at 0.05 and 0.01 probability levels, respectively.

zAbbreviations of traits are same with Table 1.

Results of composite interval mapping for agronomic traits in F2.

Traitu Chr. Left Marker Right Marker Position (cM) LODz PVE(%)y Addx Domw |d/a|v
LC 3 scaffold8_17030380 scaffold8_16101189 95.8 14.3 50.4 0.66 0.15 PD
PB 2 scaffold25_4598423 scaffold25_3912693 57.9 10.0 25.4 0.54 0.06 A
PB 2 scaffold25_1773512 scaffold25_744098 76.2 7.8 19.1 0.05 -0.64 OD
PB 5 scaffold3_9875772 scaffold3_11292324 38.0 5.4 12.2 0.39 0.12 PD
PB 5 scaffold64_1191653 scaffold22_5567527 100.4 3.3 7.1 -0.28 -0.03 A
SL 9 scaffold35_5711921 scaffold134_133385 13.9 3.8 20.1 9.79 -14.67 OD
FCN 3 scaffold31_6872402 scaffold31_6250465 40.9 3.8 19.0 -54.43 -3.61 A
FCL 1 scaffold32_5830494 scaffold29_6732047 13.0 3.2 15.2 -1.64 1.04 PD
FCL 3 scaffold49_35668 scaffold6_4743200 63.4 4.0 17.8 -1.92 0.21 A
FN 3 scaffold6_9130731 scaffold21_10557309 58.2 5.1 23.8 -5.00 2.14 PD
SW 10 scaffold28_5887083 scaffold28_6525916 24.2 3.7 19.6 3.13 -1.60 PD
RA 1 scaffold16_7569624 scaffold16_7019064 150.9 4.2 8.3 -115.90 0.47 A
RA 5 scaffold10_10061999 scaffold10_9181391 4.0 7.1 14.8 171.59 47.89 PD
RA 6 scaffold43_1927956 scaffold43_2688122 34.6 3.7 7.2 103.81 27.49 PD
RA 7 scaffold59_753444 scaffold59_2135864 83.7 4.0 7.8 -27.91 -134.19 OD
RA 7 scaffold34_6133629 scaffold34_6235270 156.9 3.4 6.3 108.83 -31.67 PD
RA 10 scaffold12_7503648 scaffold12_9560702 2.1 12.8 32.1 -229.27 11.30 A
LU 1 scaffold46_600664 scaffold46_534033 191.8 5.2 22.4 59.15 -10.69 A
LU 10 scaffold32_1386668 scaffold32_1454570 21.8 3.3 13.6 44.55 -6.67 A
AP 1 scaffold2_12959065 scaffold2_13146855 89.8 3.8 12.6 46.97 26.22 PD
AP 4 scaffold24_857542 scaffold24_1017077 77.3 5.8 21.0 -6.65 90.29 OD

zLOD score for the detected QTLs (LOD threshold=3.0).

yproportion of variance explained by a QTL at the test site under H0:H3.

xAdditive effect under H1.

wDominant effect under H2.

vestimation of gene action, A (additive effect) 0-0.20, PD (partial dominance) 0.21-0.80, D (dominance) 0.81-1.20, OD (over dominance)>1.20.

uAbbreviations ; LC (Leaf Color), PB (Pubescence), others are same with Table 1.

Anthocyanin biosynthesis transcription factor like sequences located near from Leaf Color (LC) QTL of Chromosome 3.

Transcript No. Speciesz Sequence Identity TAIRy TAIR description
1SL023156t001 Mimulus guttatus v1.1 72.25 AT1G72210.1 basic helix-loop-helix (bHLH) DNA-binding superfamily protein
1SL003771t001 Vitis vinifera 73.06 AT5G16600.1 myb domain protein 43
1SL006007t004 Mimulus guttatus v2.0 93.39 AT5G66240.2 Transducin/WD40 repeat-like superfamily protein

zMost similar sequence DB with P. citriodora transcript sequence.

yLocus number in TAIR(The Arabidopsis Information Resource, http://www.arabidopsis.org).

Table 1 Phenotypic variation in F2 population of a cross P. citriodora × P. hirtella.
Table 2 Phenotypic Pearson correlations among traits in F2 populationz.

Correlation is significant at 0.05 and 0.01 probability levels, respectively.

Correlation is significant at 0.05 and 0.01 probability levels, respectively.

Abbreviations of traits are same with Table 1.

Table 3 Results of composite interval mapping for agronomic traits in F2.

LOD score for the detected QTLs (LOD threshold=3.0).

proportion of variance explained by a QTL at the test site under H0:H3.

Additive effect under H1.

Dominant effect under H2.

estimation of gene action, A (additive effect) 0-0.20, PD (partial dominance) 0.21-0.80, D (dominance) 0.81-1.20, OD (over dominance)>1.20.

Abbreviations ; LC (Leaf Color), PB (Pubescence), others are same with Table 1.

Table 4 Anthocyanin biosynthesis transcription factor like sequences located near from Leaf Color (LC) QTL of Chromosome 3.

Most similar sequence DB with P. citriodora transcript sequence.

Locus number in TAIR(The Arabidopsis Information Resource, http://www.arabidopsis.org).

Table 1

Table 2

Table 3

Table 4

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