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Korean. J. Breed. Sci. : Korean Journal of Breeding Science

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나도풍란과 하이그로칠러스 교잡종의 꽃 무늬 발현 양상 분석

Analysis of the Floral Pattern of Sedirea japonica and Its Hybrids with Hygrochilus parishii

The Korean Journal of Breeding Science 2016;48(4):399-405.
Published online: November 30, 2016

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*Corresponding author (jungheon1@korea.kr, +82-63-238-4651, +82-238-4654)
• Received: August 5, 2016   • Accepted: November 21, 2016

© 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/3.0) which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original work is properly cited.

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  • Floral pattern plays a vital role in flower structure. The most basic factors that make up the floral pattern are spots and lines in Sedirea japonica. These factors were expressed in a various pattern according to their location and the type of the flower. Frequent expression pattern is higher in the left side of lateral sepal and labellum than petal and dorsal sepal. The expression type in one plant is usually very different according to right or left of sepals, flowering orders, and flowering times and in some individuals, expression pattern remained constant in all flowers. Two hybrids from the cross between Hygrochilus parishii and H. pasishii var. purpureus hybrid (Hongjadan) have shown similar flower characteristics (flower size, flower background color, and pattern distribution) to those of H. pasishii with the exception of the pattern, size, and expression of the number of spots, and pattern shape of labellum including morphology. The flower of an offspring from Hongjadan x S. japonicum (K10) showed petals and sepals without pattern, red flecks, red spot, and red sectors. The shape and back ground color of labellum were similar to those of Hongjadan and S. japonica. The results of this study will be helpful for breeding new cultivars of Sedirea japonica.
난초과의 다년생 초본류인 나도풍란[Sedirea japonica (Linden & Rchb.f.) Garay & H. R. Sweet]은 나무나 바위에 붙어 자라는 착생식물로 제주한란, 죽백란 등과 함께 환경부 지정 멸종위기 식물종(1급)이다(Kim et al. 2014). 무분별한 남획으로 인해 자생지에서 나도풍란을 관찰하기란 매우 힘든 것으로 알려져 있지만 다행스럽게도 자생지 복원을 위한 움직임 이 최근까지도 계속되고 있다. 자생지에서의 사정과는 다르게 나도풍란은 지난 20여 년 동안 원예작물로서 많은 인기를 누려 왔는데, 이는 꽃 향기와 자태가 뛰어나고 무균배양법 연구 (Chung et al. 1984)를 통한 대량 번식이 가능한대서 기인한 결과로 판단된다.
자연상태에서 나도풍란은 꽃대 한 개에 4~10개의 자색줄무늬 가 있는 연한 녹백색 꽃이 총상으로 달리어 6~8월에 개화한다. 소비자들이 선호하는 나도풍란의 주요 원예형질은 꽃의 경우 향기와 줄무늬 유무, 잎은 작고 둥근 모양, 노란색과 줄무늬 유무 등인데, 꽃에 줄무늬가 없어 단아하게 보이는 개체(소심)와 잎에 무늬가 있는 개체는 대량번식이 쉽지 않다. 반면에 청옥환으 로 대표되는 잎이 작고 둥근 환엽형 개체는 열성형질로 유전되어 무균파종을 통한 대량 번식이 가능하지만, 균일한 개체 생산이 어려운 것으로 알려져 있다(Hur 2006, Park et al. 2014). 이러한 문제점 때문에 환엽형 나도풍란 생산을 위한 생장억제제 처리연 구(Chung et al. 2004)와 독농가들의 부단한 노력으로 환엽 개체 생산이 가능해졌지만, 농약사용으로 인한 인체유해성과 장기간 재배시 원래 형질인 장엽으로 복원하는 문제는 여전히 풀어야 할 숙제로 남아 있다.
나도풍란은 재배가 쉽고 향기는 청아하며 나무나 석부, 목부작 의 소재로 활용도가 높지만 대중적 품목인 호접란에 비해 꽃무늬 가 단조로운 편이다. 이러한 단점을 개선하고자 이종간 교배육종 이 꾸준히 시도되었고, 최근에는 Hygrochilus parishii 또는 유사 종과의 교배를 통해 노란색 바탕에 갈색 줄무늬 꽃이 피는 프린스 와 붉은색 꽃이 피는 홍자단 품종이 개발되기도 하였다. 일부에서 는 도리티스 또는 호접란과 교잡한 F1 개체를 육성한 경우도 알려져 있지만(Kim et al. 2015) 현재까지 품종으로 보고된 예는 없다.
오늘날 국제식물신품종보호연맹(UPOV)이나 세계지적재산 권기구(WIPO) 등은 유전자원의 지적 재산권 보호를 주장하고 있고, 이런 상황에서 자국의 유전자원을 활용하는 것은 자원 주권 확보와 지식 재산권 강화에 매우 중요하다. 나도풍란은 국내 고유 생물종 가운데 자원 주권을 확보하는 측면에서 고부가 신품종 육종을 위한 소재로서 중요한 의미를 가진다. 아울러 지난 20년간의 재배법 확립과 향기분석(Been & Kim 2013) 및 우수개체 증식 연구 등도 신품종 나도풍란 육성에 매우 고무적 인 여건으로 판단된다. 그럼에도 불구하고 나도풍란을 이용한 고부가 신품종 육종에 필요한 계통도 분석을 통한 근연 관계 구명, 우수 원예형질 발굴, 우수개체 최적 대량 증식법, 주요 형질의 유전양상 연구 등이 미미한 실정이다.
본 연구에서는 우선적으로 나도풍란 신품종 육성에 필요한 기초자료를 얻고자 나도풍란의 꽃무늬 변화, 나도풍란과 Hygrochilusi parishii 유래 교배종인 홍자단 교잡종의 꽃 형질 발현 양상을 조사하였다.
식물재료 및 꽃 사진 분석
시중에서 구입한 나도풍란 수집종 80개체 가운데 생장 상태가 양호한 30개체(T3, T11, T12, T14, T15, T16, T19, T21, T22, T23, T25, T26, T27, T28, T29, T30, T31, T32, T33, T34, T35, T37, T39, T46, T47, T49, T58, T59, T67, k10, T55)를 실험재료로 사용하였다. 실험재료는 과천시 소재 비닐온실에서 재배하면서, 2011년 2월부터 2015년 6월까지 개화된 꽃을 디지 털 카메라(Nikon D90, Japan)로 촬영하여 총 536 장의 꽃 사진을 획득하였다. 이들 가운데 꽃잎(petal) 536개, 주판(dorsal sepal) 536개, 왼쪽 부판(lateral sepal) 512개, 설판(labellum) 431개를 알씨 V7.64(엔씨소프트, 한국) 프로그램을 사용하여 발췌하고 각 부위에서 무늬 발현 양상을 육안으로 분석하였다. 개화 년도에 따른 꽃 잎과 주판의 무늬 발현 빈도 변화는 3년간 (2011, 2012, 2013) 연속적으로 개화한 5개 개체(T15, T25, T27, T29, T35)를 대상으로 Excel 프로그램을 이용해 분석하였다.
나도풍란과 Hygrochilus parishii 교잡종 생산 및 특성 분석
2010년 3월에 나도풍란(K10), Hygrochilus parishii(K11), H. parishii var. purpureus와 나도풍란 교잡종으로 알려진 홍자 단(K8)간에 교배를 수행하고 수분 후 170일 경에 꼬투리를 수확 하였다. 수확한 꼬투리를 2% 락스에 5분간 침지하고 멸균수로 2회 세척한 다음 1회 화염 살균하였으며, 하이포넥스 배지에 증식하고(Chung et al. 1984) 배양 9개월 후 F1(K11 x K8) 과 F1(K11 x K10) 각각 30개체와 F1(K8 x K10) 150개체를 온실에서 순화하고 재배하였다. 이후 2012년에서 2015년 4월까 지 개화된 12개체의 꽃 특성을 분석하였다.
꽃 무늬 유형
나도풍란의 꽃은 바탕색이 연한 녹백색이고 부판(later sepal) 과 설판(labellum)에 적자색 무늬가 분포하는 것이 일반적인 특징이다(Fig. 1-A). 본 실험 사용한 개체에서는 일반적인 특징 이외에도 꽃의 부위에 따라 무늬 발현 양상이 다양하였다. 꽃에서 발현되는 무늬는 점과 선으로 크게 구별하여 나눌 수 있는데, 점 무늬는 지름에 따라 작은 점(2mm 이하)과 큰 점(2mm 이상) 으로 세분화 할 수 있었다(Fig. 2-A2 & 3). 선 무늬는 꽃 받침 (sepal)의 횡단면을 기준으로 끝까지 이어지지 못한 짧은 것(단 선)(Fig. 2-A4)과 완전하게 이어진 것(온선)(Fig. 2)으로 대별할 수 있었고, 발현 수에도 차이를 보였으며(Fig. 2) 점, 선, 또는 둘 다가 합쳐지거나 퍼진 무늬(Fig. 2)로 발현되었다.
Fig. 1.
Flowers of Sedirea japonica, Hygrochilus parishii, and H. pasishii var. purpureus hybrid (Hongjadan). The Arabic number in circle indicates petal (➀), dorsal sepal (➁), lateral sepals (➂), labellum (➃), column (➄), side lobe (➅), and middle lobe(➆), respectively.
KJBS-48-4-399_F1.gif
꽃잎, 꽃받침, 설판의 무늬 발현 양상
꽃잎(petal)의 무늬는 점, 단선, 온선 또는 이들의 복합 형태로 꽃잎의 왼쪽, 오른쪽 또는 양쪽에 나타났는데(Fig. 2) 무늬가 없는 것(Fig. 2-A1)은 446개, 있는 것은 90개였다. 주판(dorsal sepal)의 무늬 형태는 기본적으로 꽃잎과 비슷하였으나 좀 더 복잡하고, 무늬가 없는 것(Fig. 2-B1)과 있는 것은 각각 327개와 209개였다. 주판을 위에서 아래로 10등분 했을 때 이들은 주로 9, 6, 3 등분 지점에 분포하였고(Fig. 2-B2~14), 3등분 아래 쪽 지점에서는 점선이 여러 개 나타나는 경우도 관찰할 수 있었다 (Fig. 2-B10~12). 왼쪽 꽃받침에서는 조사한 모든 시료(총 512 개)에서 온선(Fig. 2-C1 & 2)과 단선(Fig. 2-C 3), 이들의 복합체 (Fig. 2-C4 & 5), 선의 두께가 다른 형태(Fig. 2-C6 & 7)로 나타났다. 꽃받침을 긴 쪽으로 기준으로 횡단했을 때, 꽃술대 (column)의 기저부까지 절반 이하에는(Fig. 2-C1) 모든 시료에 서 무늬가 나타났으나, 절반 이상(Fig. 2-C2)에서는 386개에서 만 관찰되었다. 드물기는 하지만 꽃 받침을 긴 쪽으로 종단했을 때 아래쪽에만 무늬가 나타나는 경우(Fig. 2-C3)도 한 개체(T25) 에서 관찰되었는데 9개 꽃 가운데 7개에서 관찰되었다. 한편, 같은 시기에 동일개체에서 개화한 꽃의 무늬 발현 정도를 조사한 결과 꽃 마다 무늬 발현 정도가 다르게 나타났고(Fig. 3), 개화 년도에 따라 꽃 무늬 발현은 꽃 받침과 주판 둘 다에서 무늬 발현 빈도가 높고 표준편차가 낮은 것, 빈도와 표준편차가 둘 다가 낮은 것, 표준편차 값의 오차가 심한 것으로 나눌 수 있었다 (Fig. 4).
Fig. 2.
Representative patterns and its location on the petal (A), and dorsal sepal (B), and the left side of lateral sepal (C) in Sedirea japonica.
KJBS-48-4-399_F2.gif
Fig. 3.
Patterns and its location on the entire inflorescence of an accession T29 of Sedirea japonica. Photographs (A~L) were taken in order of flowering in March 2013.
KJBS-48-4-399_F3.gif
Fig. 4.
Means of pattern frequency on the petal or the left side of lateral sepal in five representatives of Sedirea japonica. The frequency count was calculated over the entire inflorescence of each plant at a flowering time in 2011, 2012, and 2013 years.
KJBS-48-4-399_F4.gif
Fig. 5.
Representative patterns on the labellum of Sedirea japonica (A) and pattern changes depending on flowering time in T15 (B), T27 (C), T29 (D), T32 (E), T35 (F), and T46 (G) plants. Photographs were taken in the year shown.
KJBS-48-4-399_F5.gif
순판에서는 주로 점과 이들의 복합체 형태로 무늬가 나타났는 데, 모양과 분포 위치에 따라 6가지 정도로 대별할 수 있었으며, 동일 개체에서도 개화 년도에 따라 다르게 발현되는 경향을 보였다(Fig. 5).
Hygrochilus parishii 와 홍자단 교배종의 꽃 특성 비교
Hygrochilus parishii(K11)에 나도풍란(K10)를 교배한 조합 에서는 30개체 중 1개체도 개화하지 않았고, K11과 홍자단(K8) 교배조합에서는 30개체 중 2개체만이 개화하여 2개체의 꽃 특성 을 조사하였다. 꽃의 크기와 모양은 전체적으로 H. parishii와 유사하였고 바탕색은 노랑, 점무늬 크기는 H. parishii에 비해 조금 작아졌고 갯수는 늘어 났으며, 향기는 양친의 어느 쪽과도 같지 않았다. 순판 모양은 양친의 중간 형태였고 무늬는 K10과 유사하였으며, side lobe와 middle lobe의 형태는 H. parishii와 유사하였다(Fig. 6-A & B). 홍자단(K8)에 나도풍란(K10)을 교배했을 때, 후대의 꽃은 다양한 특성을 보였다. 120개체 중 10개체가 개화하였는데, 6개체는 꽃 모양이 K10과 유사하였고, 나머지는 4개체는 K8과 유사하였다. 꽃잎의 무늬는 바탕색이 붉은 한 개체(Fig. 6-K)를 포함해 무늬가 없는 것은 7개체, 붉은 색 얼룩(Fig. 6-C), 붉은색 점무늬(Fig. 6-E), 붉은색 점(Fig. 6-L)의 형태로 나타났다. 주판과 꽃 잎은 동일한 유형을 보였으나 부판에서는 꽃잎의 절반 이하(Fig. 6-F, H, J)와 전체(Fig. 6-E & L)에 줄무늬가 있는 것, 꽃잎의 절반 이하(Fig. 6-D)와 전체 (Fig. 6-C)에 얼룩무늬 퍼져있는 것, 무늬가 없는 것(Fig. 6-G, I, & K)을 관찰할 수 있었다. 순판 모양은 K8과 K10형이 각각 4개와 6개였고 줄무늬가 있는 것은 5개였으나 점 무늬의 분포 유형이 모두 달랐다. 꽃잎의 바탕색은 붉은색(Fig. 6-K), 노랑색 (Fig. 6-L), 연한 녹백색이었고, 순판의 바탕색은 붉은색(Fig. 6-C, D, I, K, & L)과 연한 녹백색이였다(Fig. 6).
Fig. 6.
Photographs of hybrids from Hygrochilus parishii x Honjadan (A and B) and Honjadan x Sedirea japonica (C~L) crosses.
KJBS-48-4-399_F6.gif
나도풍란(Sedirea japonica)의 꽃 무늬를 구성하는 기본적인 요인은 점과 선으로 비교적 단순하지만(Fig. 1), 각 구성요소가 연출하는 무늬 유형과 개화시간에 따른 분포양상 변화로 다양한 특성을 지닌 꽃 무늬가 관찰되었다. 꽃 무늬의 다양성은 수분 매개충(pollinator)의 방문 기회를 높임으로써 종 번식을 위한 기회 향상에 중요한 역할을 하는 것으로 알려져 있는데(Heather et al. 2009, Suetsugu & Tanaka 2013) 이와 관련해 연구가 가장 많이 된 난과 식물은 흑란(ophrys)이다. 흑란의 꽃 모양은 수분 매개충인 말벌과 부분적으로 닮아 있고, 꽃의 무늬 변화가 다양하여 수컷 말벌을 유혹하는 것으로 보고되어 있다(Heather et al. 2009).
나도풍란에서는 꽃의 색 변화가 수분 매개충인 어리뒤영벌 (Bombus diversus diversus)을 끌어들이는데 관계가 있을 것으로 추정한 보고도 있지만(Suetsugu & Tanaka 2013), 아직까지 무늬 발현이 어떤 기능을 하는지 알 수가 없다. 그러나 시계꽃 (Hibiscus trionum)에서 연구된 무늬 변화가 자외선 반사량을 증가시켜 곤충의 꽃 방문 횟수를 증가시키는데 기여한다는 보고 (Heather et al. 2009)에 비추어, 적어도 나도풍란의 무늬와 발현 유형은 수분 매개충을 유혹하는데 조금이나마 기여를 하지 않을 까 추정한다.
본 연구에서 나도풍란의 꽃 무늬 발현 변화에 영향을 주는 요인으로 가장 가능성이 높아 보이는 것은 DNA transposon의 활성 유무이다. DNA transposon은 나팔꽃이나 카네이션에서 부정형의 얼룩무늬 발현 변화에 주요한 역할을 하는 것으로 알려다(Inagaki et al. 1994, Iida et al. 1999, Itoh et al. 2002). 나도풍란의 경우 일부 개체의 부판(Fig. 2-C7)과 순판(Fig. 5-B~G)에서 얼룩무늬 변화가 관찰 되어 이와 유사한 특성을 보였고, 꽃 잎과 꽃 받침에서 점과 선의 개수나 분포 위치에 다양한 차이를 보였다(Fig. 2). 특히 같은 개체의 꽃과 개화 연도별로 다른 유형의 무늬가 발현되는 것을 볼 때 꽃 무늬 발현은 유전력이 매우 낮은 것으로 판단된다. 그러나 부판의 경우 무늬 발현 양상이 환경에 따라 다르게 나타나는 경향이 강하다 할지라도 조사한 모든 꽃에서 무늬가 발현되고 홍자단 x S. japonicum(K10) 교잡종 가운데 나도풍란과 유사한 개체의 부판에 줄무늬와 얼룩무늬가 발현되는 사실에 근거해 부판의 무늬 발현은 유전력이 높은 형질로 판단된다. 순판에서 발현되는 점무늬는 환경적 요인에 의해 많은 영향을 받지만 무늬 발현 유무는 유전력이 높을 것으로 판단된다(Fig. 6).
한편, 본 연구에서는 Hygrochilus parishii(K11)과 나도풍란 (K10) 교잡종의 꽃을 관찰할 수 없었고, 홍자단(K8)과 나도풍란 과 교잡 후대에서도 개화율 약 7% 정도로 낮게 나타났다. 이는 부분적으로 개화에 필요한 저온요구도가 두 종간에 다른 것에서 기인했다고 보여지며, Hygrochilus parishii나 홍자단을 이용한 나도풍란 신품종 육성시 꽃무늬 발현 변화와 함께 반드시 고려해 야 할 요인으로 판단된다. 아울러 나도풍란에 대한 보다 많은 이해를 위해 유전력이 강할 것으로 판단되는 꽃잎과 꽃 받침의 무늬, 꽃 배경색 유무, 부판의 무늬 발현 유무, 순판의 적색 형질 등과 같은 주요 원예형질에 대한 추가 연구가 수행되어야 할 것으로 판단된다.
꽃 무늬는 꽃의 구조에서 필수적인 역할을 한다. 국내 자생종 인 나도풍란(Sedirea japonica)의 꽃 무늬를 구성하는 가장 기본 적인 요인은 점과 선이며, 꽃에서 발현되는 이들의 위치와 유형에 따라 다양한 꽃 무늬로 발현되었다. 무늬 발현되는 빈도는 꽃 잎과 주판보다는 부판과 설판에서 높게 나타났으며, 발현 유형은 오른쪽과 왼쪽 부판, 개화순서, 개화시기에 따라 동일 개체에서 도 다르게 나타나는 경향이 강했고, 일부 개체에서는 발현 빈도가 일정하게 유지되는 경우도 관찰되었다. Hygrochilus parishii x H. parishii 유래종(홍자단) 교잡종의 꽃 특성은 점 무늬 크기 및 발현 수, 순판의 형태 및 무늬 유형을 제외한 나머지 형질(꽃 크기, 바탕색, 무늬 분포 등)은 H. parishii와 유사하였다. 홍자단 x S. japonicum 후대에서는 꽃 잎과 꽃 받침에 무늬가 없는 것을 포함해 붉은 얼룩, 붉은 점무늬, 붉은색 퍼진 점이 특정 부분 또는 전체에 나타났고, 배경 색이 홍자단 처럼 붉은 꽃도 있었다. 순판 모양과 바탕색은 홍자단과 S. japonica형이 비슷한 비율로 나타났다. 이상의 결과는 향후 나도풍란을 이용한 신품종 착색란 육성에 도움이 될 것으로 판단되었다.
본 논문은 농촌진흥청 공동연구사업(PJ01183201)의 지원으 로 이루어진 것임.
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Analysis of the Floral Pattern of Sedirea japonica and Its Hybrids with Hygrochilus parishii
Korean. J. Breed. Sci.. 2016;48(4):399-405.   Published online December 31, 2016
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Analysis of the Floral Pattern of Sedirea japonica and Its Hybrids with Hygrochilus parishii
Korean. J. Breed. Sci.. 2016;48(4):399-405.   Published online December 31, 2016
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Analysis of the Floral Pattern of Sedirea japonica and Its Hybrids with Hygrochilus parishii
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Fig. 1. Flowers of Sedirea japonica, Hygrochilus parishii, and H. pasishii var. purpureus hybrid (Hongjadan). The Arabic number in circle indicates petal (➀), dorsal sepal (➁), lateral sepals (➂), labellum (➃), column (➄), side lobe (➅), and middle lobe(➆), respectively.
Fig. 2. Representative patterns and its location on the petal (A), and dorsal sepal (B), and the left side of lateral sepal (C) in Sedirea japonica.
Fig. 3. Patterns and its location on the entire inflorescence of an accession T29 of Sedirea japonica. Photographs (A~L) were taken in order of flowering in March 2013.
Fig. 4. Means of pattern frequency on the petal or the left side of lateral sepal in five representatives of Sedirea japonica. The frequency count was calculated over the entire inflorescence of each plant at a flowering time in 2011, 2012, and 2013 years.
Fig. 5. Representative patterns on the labellum of Sedirea japonica (A) and pattern changes depending on flowering time in T15 (B), T27 (C), T29 (D), T32 (E), T35 (F), and T46 (G) plants. Photographs were taken in the year shown.
Fig. 6. Photographs of hybrids from Hygrochilus parishii x Honjadan (A and B) and Honjadan x Sedirea japonica (C~L) crosses.
Analysis of the Floral Pattern of Sedirea japonica and Its Hybrids with Hygrochilus parishii