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

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‘Tano Red’와 ‘Ruby Seedless’ 교배 후대에서의 주요 과실 형질에 대한 평가

Evaluation of Major Fruit Traits in Progenies Derived from a Cross between ‘Tano Red’ and ‘Ruby Seedless’ Grapevines (Vitis spp.)

Korean Journal of Breeding Science 2017;49(4):324-333.
Published online: November 30, 2017

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*Corresponding author : (khc7027@korea.kr), +82-63-238-6712, +82-63-238-6705
• Received: July 31, 2017   • Accepted: October 11, 2017

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  • In this study, phenotypic distribution of 15 major fruit quality traits were analyzed using 252 progenies derived from a cross between ‘Tano Red’ (seed) and ‘Ruby Seedless’ (seedless), to obtain basic data for improving the breeding efficiency of grapevine cultivars. Berry skin color was dark red-violet in 46.4% of the progenies, which is the color of ‘Tano Red’ and ‘Ruby Seedless’, and berry shape was elliptic in 48.4%. Most of the progenies were very juicy with soft flesh, and closely related to the characteristics of ‘Tano Red’. Seeds were well developed in 67.1% of the progenies, rudimentary in 30.1%, and 2.8% were seedless, with seed weight being less than 0.15 g in 84.9% of the progenies. Among the 15 fruit quality traits assessed, bunch density, ease of berry detachment from pedicel, berry weight, berry seed number, berry longitudinal diameter, berry transverse diameter, berry soluble solids, and berry acidity showed normal distributions. Heritability of berry weight, berry longitudinal diameter, berry transverse diameter, berry soluble solids, and berry acidity was 0.89, 0.82, 0.78, 0.86, and 0.93, respectively. Berry weight was positively correlated with seed weight (r = 0.486**), presence of seeds (r = 0.483**), and seed number (r = 0.211**). Seed weight significantly increased with presence of seeds (r = 0.607**) and seed number (r = 0.725**). In addition, presence of seeds was positively correlated with seed number (r = 0.319**). These results could be useful for the identification of quantitative trait loci associated with fruit quality to assist in grapevine breeding.
포도(Vitis spp.)는 포도과(Vitaceae) 포도속(Vitis)에 속하는 식물로 약 60종이 존재하는 것으로 알려져 있다(Olmo 1976). 포도는 약 5000년 동안 재배되어 왔으며 영양번식을 통해 많은 품종들이 널리 퍼져 있다(Moreno-Sanz et al. 2008). 전 세계에 서 주로 재배되고 있는 유럽종(Vitis vinifera L.) 포도 외에 미국종 (Vitis labrusca L.) 등 100종의 야생종 포도를 품종 육종에 이용 하고 있으며 교배 육종으로 창출되는 변이의 폭을 넓히고 있다 (Ledbetter & Ramming 1989). 2017년 현재 우리나라의 포도 재배면적은 전년보다 11% 감소한 13,383 ha로 예측되는데, 이는 2016년 FTA 폐업면적이 2015년에 비해 10% 증가한 것으 로 추정되기 때문이다(KREI 2017).
최근 껍질째 먹을 수 있는 고품질 무핵 포도에 대한 관심이 증가하면서 국내 소비자 및 생산자의 기호 변화에 따른 다양한 포도 품종과 국내 포도 재배면적의 약 70%를 차지하는 ‘캠벨얼 리’를 대체할 새로운 품종 육성이 필요하다. 그러나, 포도 품종 육종은 장기간의 육종 기간(15년 이상)과 교배실생 유지에 대면 적의 포장과 많은 비용이 요구되는 과수 특성상 국가기관에서만 수행되고 있고, 유전조성이 이형접합(heterozygous) 상태이므 로 유전육종을 수행하는 데도 어려움이 있다. 또한 특정형질을 목표로 하는 포도 육종 프로그램의 역사는 매우 짧다(Hur et al. 2014). 우리나라에서 교배를 통한 포도 품종 육성은 1993년 ‘청수’ 품종을 시작으로 현재까지 30종의 품종이 육성되었다. 최근에 육성된 품종들은 기존 품종들을 모본으로 사용하는 경우 가 많아 유전적 유사도가 높은 것으로 추정되고 있어 변이의 폭이 좁다(Cho et al. 2011). 또한 포도 유전자원의 유전체 및 유전양상의 체계적인 정보의 부족으로 인하여 육종 소재로서의 유전자원의 활용성은 미흡하다. 따라서 다양한 유전자원의 도입 과 평가를 통해 유용형질을 보유한 유전자원을 육종에 활용하는 것이 필요하다. 선발 시 형질의 유전력과 형질들 간의 상관 분석 과 같은 정보는 육종 프로그램에서 유전적 과정을 예측하고 효율적인 육종 전략을 수립하는데 있어서 매우 유용하다 (Falconer & Mackay 1996).
포도 과실 형질에 관한 유전 연구는 과피색, 과립 크기, 무핵, 종자 무게, 종자 개수 등 일부 형질에 대해서만 보고되었다 (Bešlić et al. 2005; Bouquet & Danglot 1996; Cabezas et al. 2006; Ebadi et al. 2009; Fisher et al. 2004; Mejia et al. 2007; Park et al. 2003; Salmaso et al. 2008). 포도의 과피색은 과피에 축적되는 안토시아닌 색소의 함량과 종류에 의해 결정되 는데(Azuma et al. 2009), 품종들 간의 과피색이 황록색에서부 터 흑청색까지 다양하여 유전적 변이가 넓은 것으로 알려져 있다(Lijavetzky et al. 2006). Bešlić et al. (2005)은 ‘Drenak crveni’와 ‘Afuz-ali’의 교배조합 후대 실생 130개체를 대상으로 과실 특성을 조사한 결과, 과립 크기, 과립 모양 등의 형질과 달리 과립의 과피색은 χ2-검정을 통해서 단일 인자에 의해 발현하 는 것으로 추정하였다. 또한 양적형질유전자좌(quantitative trait loci: QTL) 분석을 통해서 과립의 과피색을 결정하는 단일 유전 자좌가 linkage group (LG) 2번에서 동정되었다(Doligez et al. 2002; Lijavetzky et al. 2006; Salmaso et al. 2008). 다른 과립 형질에 대한 유전 분석은 생식용 포도의 육종 프로그램 목표인 무핵과 과립 크기에 중점을 두고 연구되었다(Cabezas et al. 2006; Doligez et al. 2002; Mejia et al. 2007). Cabezas et al. (2006)은 ‘Dominga’와 ‘Autumn Seedless’의 교배조합 후대 실생 118개체를 대상으로 과립중, 종자 무게와 종자 개수를 조사한 결과 모두 정규분포를 나타내어 양적형질이 관여하는 것으로 판단하였다. 수정은 정상적으로 이루어지나 배주가 발육 도중 퇴화하여 종자를 형성하지 못하는 위단위결과성 (stenospermic) 무핵 형질은 무핵 품종과 후대에서 종자 발달의 정도를 모두 관찰할 수 있어 양적형질로 여겨져 왔다(Cabezas et al. 2006). 또한 무핵 형질과 관련 있는 종자 개수 및 종자 무게 등과 연관된 다수의 QTL이 동정되었다(Cabezas et al. 2006; Mejía et al. 2007, 2011). 현재 국내에서도 주요 품종의 형질을 대상으로 포도 육종 효율을 증진할 수 있는 분자 유전 연구가 진행되고 있다(Hur et al. 2015).
본 연구는 포도 유핵과 무핵의 교배조합 후대 실생 집단에서 주요 과실 형질에 대해 평가하여 포도 유전 육종 연구의 기반을 조성하고 품종의 육종 효율을 증진하기 위한 기초자료를 확보하 고자 하였다.
시험재료
포도 과실 형질의 평가를 위하여 종자친인 ‘Tano Red’와 화분친인 ‘Ruby Seedless’를 2001년에 교배하여 국립원예특작 과학원에 재식 되어 보존 중인 4년생 교배실생 252주를 사용하였 다. 양친인 ‘Tano Red’는 유핵이고, ‘Ruby Seedless’는 무핵 품종이다. 교배실생의 재배는 개량일자형의 노지 비가림 재배로 재식거리는 5 m × 5 m 간격으로 하였다. 시비 및 병해충 방제 등 재배관리는 농촌진흥청의 ‘포도 재배기술’ (RDA 1997)에 준하였다.
과실형질 조사
양친과 교배실생의 과실 형질 조사는 각 실생의 한 나무에서 중간 크기의 과방 3개를 채취하여 수행하였다. 수확기는 2016년 8월 하순부터 9월 중순까지였다. 조사항목은 과방의 착립 밀도 (bunch density: BND), 과립 모양(berry shape: BS), 과립중 (berry weight: BW), 과립의 과피색(과분 제외, berry skin color: BSC), 과립경에서의 과립 분리 정도(berry ease of detachment from pedicel: BED), 과육에서의 종자 분리 정도(berry ease of seeds separating from flesh: BES), 과즙량(berry juiciness of flesh: BJF), 과육 경도(berry firmness of flesh: BFF), 종자 무게(berry seed weight: BSW), 종자 형태(berry presence of seeds: BPS), 과립당 종자 개수(berry seed number: BSN), 과립 종경(berry longitudinal diameter: BLD), 과립 횡경(berry transverse diameter: BTD), 당도(berry soluble solids: BSS), 산 함량(berry acidity: BA)의 총 15개였다. 각 형질에 따른 조사 방법은 국제식물유전자원연구소(International Plant Genetic Resources Institute)에서 정한 ‘Descriptors for grapevine’ (1997)을 참고하였다(Table 1). 포도의 과립중, 과립 종경, 과립 횡경, 과립당 종자 개수 등은 과방당 과립 15개를 조사하여 평균 값을 이용하였다.
Table 1
Fruit characteristics measured on 252 progenies derived from a cross between ‘Tano Red’ and ‘Ruby Seedless’.
Table 1
Trait Abbreviation Unit Remark

Bunch density BND 1-9 1. Very loose (berries in grouped formation, many visible pedicels)
3. Loose (single berries with some visible pedicels)
5. Medium (densely distributed berries, pedicels not visible)
7. Dense (berries not readily movable)
9. Very dense (berries pressed out of shape)
Berry shape BS 1-9
2. Narrow elliptic
3. Elliptic
4. Round
5. Oblate
6.Ovate
7. Obtuse-ovate
8. Obovate
9. Arched
Berry weight BW g Weight of berry
Berry skin color (without bloom) BSC 1-6 1. Green-yellow
2. Rose
3. Red
4. Red-grey
5. Dark red-violet
6. Blue-black
Berry ease of detachment from BED 1-3 1. Difficult
pedicel 2. Slightly easy
3. Very easy
Berry ease of seeds separating BES 1-3 1. Difficult
from flesh 2. Slightly easy
3. Very easy
Berry juiciness of flesh BJF 1-3 1. Very slightly juicy
2. Slightly juicy
3. Very juicy
Berry firmness of flesh BFF 1-3 1. Soft
2. Medium
3. Firm
Berry seed weight BSW g Weight of seed
Berry presence of seeds BPS 1-3 1. Seedless (absent)
2. Rudimentary
3. Well developed
Berry seed number BSN 1-4 Seed number of berry
Berry longitudinal diameter BLD mm Longitudinal diameter of berry
Berry transverse diameter BTD mm Transverse diameter of berry
Berry soluble solids BSS oBrix Sugar content of berry
Berry acidity BA % Acid content of berry
통계분석
교배실생에 대한 과실특성의 실생 분포도는 Excel 프로그램 을 이용하여 작성하였고, 과립중, 과립 종경 등의 양적형질의 유전력 분석은 양친 품종(P1, P2)의 분산평균과 교배실생의 분산 을 산출하여 다음과 같은 공식으로 계산하였다(Hong 1991).
유전력(h2)=VGVG+VE유전분산(VG)=VF1VE환경분산(VE)=VP1+VP22
또한 형질 중에서 과립중과 종자 형태, 종자 무게, 종자 개수의 형질 간의 상관분석도 Excel 프로그램을 이용하여 수행하였다.
양친의 과실 형질 특성
양친인 ‘Tano Red’와 ‘Ruby Seedless’의 수확 시기는 각각 2016년 9월 26일과 9월 6일이었다. Fig 1AB는 양친의 과실 모습이고 과실 형질의 특성은 Table 2와 같다. ‘Tano Red’는 일본에서 개인 육종가가 육성한 종간 교잡 품종으로 양친은 알려져 있지 않다. 과실 형질 특성 조사 결과, 과방의 착립 밀도는 매우 빽빽하고(very dense) 과립 모양이 도란형(obovate)으로 과 립중은 4.11 g이었다. 과립의 과피색은 암자적색(dark red-violet) 으로 과립경에서의 과립 분리와 과육에서의 종자 분리는 매우 쉬운 특성을 가지고 있다. 또한 과즙량은 많은(very juicy) 반면에 과육 경도는 무르며(soft) 평균 종자 개수는 3.2개, 종자 무게는 0.16 g으로 완전한(well developed) 종자 형태를 가지고 있다. 과립 종경은 21.73 mm이고, 횡경은 18.08 mm, 당도는 11.80 °Brix, 산 함량은 0.76%였다.
Fig. 1
Fruits of ‘Tano Red’ (A), ‘Ruby Seedless’ (B), and progenies derived from a cross between ‘Tano Red’ and ‘Ruby Seedless’ (C).
KJBS-49-324_F1.gif
Table 2
Comparison of the fruit characteristics of the parents.
Table 2
Parent Traitz

BND (1-9) BS (1-9) BW (g) BSC (1-6) BED (1-3) BES (1-3) BJF (1-3) BFF (1-3) BSW (g) BPS (1-3) BSN (0-4) BLD (mm) BTD (mm) BSS (°Brix) BA (%)

Tano Red 9 (very dense) 8 (obovate) 4.11 5 (dark red-violet) 3 (very easy) 3 (very easy) 3 (very juicy) 1 (soft) 0.16 3 (well developed) 3.2 21.73 18.08 11.80 0.76
Ruby Seedless 1 (very loose) 2 (narrow elliptic) 3.58 5 (dark red-violet) 2 (slightly easy) - 2 (slightly juicy) 2 (medium) - 2 (rudimentary) 0 20.85 16.85 19.56 0.64

zBND, bunch density; BS, berry shape; BW, berry weight; BSC, berry skin color; BED, berry ease of detachment from pedicel; BES, berry ease of seeds separating from flesh; BJF, berry juiciness of flesh; BFF, berry firmness of flesh; BSW, berry seed weight; BPS, berry presence of seeds; BSN, berry seed number; BLD, berry longitudinal diameter; BTD, berry transverse diameter; BSS, berry soluble solids; BA, berry acidity.

‘Ruby Seedless’는 ‘Emperor’와 ‘Pirovano 75’ (‘Muscat of Alexandria’ × ‘Tomson Seedless’)를 교배하여 육성된 품종 으로 무핵 형질은 ‘Tomson Seedless’에서 유래하였다(Viticultu ral information, http://iv.ucdavis.edu/Viticultural_Informatio n). 과방의 착립 밀도는 매우 성기고(very loose) 과립 모양이 장타원형(narrow elliptic)이었으며, 과립중은 3.58 g으로 ‘Tano Red’보다 작았다. 과립의 과피색은 암자적색이었고, 과립경에서 의 과립 분리는 약간 쉽고(slightly easy), 과즙량은 적으며(slight ly juicy), 과육 경도는 약간 단단(medium)하였다. 종자는 흔적 만 있는 상태(rudimentary)였으며, 과립 종경은 20.85 mm, 횡경 은 16.85 mm이었다. 당도는 19.56 °Brix로 ‘Tano Red’보다 높았고 산 함량은 0.64%로 ‘Tano Red’보다 낮았다.
교배실생의 과실 형질 특성
포도 분자 육종 연구가 진행되면서 과실 형질 중에서 과피색, 무핵(종자 무게, 종자 개수), 과립중, 과육 경도, 당도 및 산함량에 대한 관련 유전자좌가 동정되었지만(Cabezas et al. 2006; Chen et al. 2015; Correa et al. 2015; Doligez et al. 2002) 다른 형질에 대해서는 보고된 바 없다. 본 연구에서는 ‘Tano Red’ × ‘Ruby Seedless’조합의 교배실생 252주에 대한 주요 과실 형질의 분포를 분석하였다(Fig. 2). 교배실생 과방의 착립 밀도는 중간 정도가 51.6%였고, 빽빽한 것이 23.8%, 성긴 것이 17.4% 로 정규분포를 나타내었다. ‘Tano Red’의 특성인 매우 빽빽한 착립 밀도를 나타낸 교배실생은 2.8%였고, ‘Ruby Seedless’의 특성인 매우 성긴 착립 밀도를 나타낸 교배실생은 4.4%였다. 착립 밀도가 빽빽한 것은 과립경의 부착뿐만 아니라 과립 크기를 감소시키기 때문에 생식용 포도에서는 바람직하지 않은 형질로 알려져 있다(Ebadi et al. 2009). 또한 착립 밀도는 수확기를 제외하고 품질 관련 특성과는 거의 연관이 없어 일반적으로 수확기가 늦은 포도나무에서 착립 밀도가 더 빽빽한 과방을 가진 과실이 생산된다고 보고되었다(Wei et al. 2002). 과립 모양은 단타원형이 48.4%로 가장 많았고, 그 다음은 장타원형 (24.6%), 원형(20.6%), 도란형(5.6%) 순이었다.
Fig. 2
Phenotypic distribution of the 15 fruit quality traits of 252 progenies derived from a cross between ‘Tano Red’ and ‘Ruby Seedless’. Filled arrows represent ‘Tano Red’ values, while empty arrows represent ‘Ruby Seedless’ values.
KJBS-49-324_F2.gif
과립중은 전체적으로 1.25-11.0 g의 범위로서 정규분포를 나타내어 Cabezas et al. (2006)의 결과와 일치하였다. ‘Tano Red’가 속하는 4.1~5.0 g의 범위에 포함되는 교배실생은 30.9% 였고, 3.1-4.0 g은 23.0%, 5.1-6.0 g은 17.5%, 6.1-7.0 g은 10.7% 순 등이었다. 교배실생 과립의 과피색은 Fig. 1C와 같이 다양하 게 분포하였는데, ‘Tano Red’와 ‘Ruby Seedless’의 과피색인 암자적색이 46.4%로 가장 많았고 그 다음은 황녹색이 23.4%, 적색이 15.1%, 그리고 흑청색이 14.7%였다.
과립경에서의 과립 분리 정도는 ‘Ruby Seedless’의 특성과 같이 교배실생의 63.5%가 약간 쉬운 것으로 조사되었고, 매우 쉬운 것이 25.4%, 어려운 것이 11.1%였다. 과립경에서의 과립 분리 정도의 특성은 생식용 포도에 있어서 중요한 형질 중의 하나로 완전한 종자를 가진 과립이 과립경에 더 강하게 붙어 있어 있는 것으로 알려져 있는데(Ebadi et al. 2009), 금후 정밀하 게 조사하여 종자 유무와의 상관을 검토할 예정이다. 과육에서의 종자 분리 정도는 어려운 것이 36.5%, 약간 쉬운 것이 44.4%, ‘Tano Red’의 특성인 매우 쉬운 것이 19.1%로 교배실생의 63.5%에서 종자 분리가 용이하였다. 과즙량이 많은 교배실생이 76.6%로 대부분이었고 적은 것은 22.6%, 매우 적은 것은 0.8% 였으며. 과육 경도는 무른 것이 59.5%였고, 약간 단단한 것이 23.8%, 매우 단단한 것은 16.7%였다. 따라서 대부분의 교배실생 은 과즙이 많고 과육이 부드러운 ‘Tano Red’의 특성과 가깝게 분포되었다.
종자 무게는 0.1 g 미만인 교배실생이 52.4%였고, 그 다음은 0.11-0.15 g (32.5%)>0.16-2.0 g (11.9%)>0.21-0.25 g (2.8%)>0.26-0.30 g (0.4%) 순이었다. 종자 무게가 0.15 g 미만 인 교배실생이 84.9%로 대부분을 차지하였다. 종자 형태는 완전 한 것이 67.1%였고, 흔적만 있는 것이 30.1%, 종자가 없는 것이 2.8%였다. 이는 후대에서 무핵의 출현 비율은 교배에 이용 한 양친에 따라 다양하지만, 전반적으로 50% 미만이었다는 Lahogue et al.(1998)Spiegel-Roy et al.(1985)의 연구 결과 와 일치하였다. Ebadi et al. (2009)도 무핵과 유핵 품종을 교배하 여 얻은 22개 집단의 후대 총 381개체에서 유핵과 무핵의 출현 비율을 4 그룹으로 나누어 조사한 결과 완전한 종자를 가진 개체의 비율이 51.2%이었음을 보고하였다. 과립당 종자 개수는 교배실생의 48.4%가 1.1-2.0개였으며 2.1-3.0개가 30.9%, 1.0 개 미만인 교배실생은 17.9%였다. 또한 종자 개수가 3.1-4.0개인 교배실생은 2.8%로 정규분포를 나타냈는데 이는 Cabezas et al. (2006)의 결과와 일치하였다. 포도의 유핵과 무핵 형질을 구분하는 기준은 종자 흔적의 검출 유무 및 풍미(Loomis & Weinberger 1979), 과립의 폴리페놀(polyphenol) 함량(Merin et al. 1983), 종자 무게와 개수(Cabezas et al. 2006) 등에 따라 분류하였다. 본 연구에서는 종자 형태 조사에 있어서 육안으로 정상적으로 보이는 종자를 완전한 형태로 구분하였고 압착 종자 등 비정상의 종자를 종자 흔적으로 분류하였다. 이와 같이 무핵과 유핵 형태의 구분은 시험방법과 연구자에 따라 다양하며, 서로 다른 결과를 도출하는 문제가 있어 먼저 무핵에 대한 정확한 검정 기준이 정립되어야 한다(Park et al. 2003). Mejía et al. (2007)은 무핵 형질과 관련된 주동 유전자와 과립 크기, 과립중과 수확기에 대한 QTL들이 동일한 위치에 존재함을 보고하였는데, 이는 하나의 유전자가 두 개 이상의 형질에 관여하는 다면발현의 효과로 추정하였다.
교배실생의 과립 종경은 양친의 크기인 20.1-22.0 mm (32.1%) 를 중심으로 정규분포하였다. 종경이 22.1-24.0 mm에 속하는 교배 실생이 25.0%였고, 그 다음은 18.1-20.0 mm (20.2%)>24.1-26.0 mm (13.5%)>16.1-18.0 mm (6.4%)>14.0-16.0 mm(2.8%) 순이 었다. 과립 횡경은 ‘Tano Red’가 속하는 18.1-20.0 mm (40.9%) 를 중심으로 정규분포하였으며, 14.0-16.0 mm와 20.1-22.0 mm 범위에 속하는 교배실생은 각각 25.8%와 21.8%였다. 당도에 있어서 교배실생의 분포는 ‘Ruby Seedless’가 속하는 18.1-20.0 °Brix는 전체의 31.8%였고, 16.1-18.0 °Brix와 20.1-22.0 °Brix 에 속하는 교배실생은 각각 28.2%, 20.6%로, 당도가 16.1~22.0 °Brix의 범위에 속하는 교배실생은 전체의 80.6%였다. 산 함량 은 ‘Ruby Seedless’가 속하는 0.61-0.70% (37.7%)를 중심으로 정규분포하였으며, 0.71-0.80%와 0.51-0.60% 범위에 속하는 교배실생은 각각 25.8%와 21.8%였다. 포도의 당도와 산 함량은 일반적으로 환경적 요인과 수량, 수확시기에 큰 영향을 받는다고 알려져 있는데(Eibach 1990), Chen et al. (2015)은 당도와 산 함량의 간의 상관관계는 환경적 요인에 의해 영향을 받기는 하지만 연차 간 비교적 안정적이었다고 보고하였다. 이 결과를 이용하여 포도에서 당도와 산 함량과 관련된 QTL 분석을 수행하 였고 각각 10개와 3개의 QTL이 동정되었다(Chen et al. 2015). 이와 같이 당도와 산 함량의 형질은 복잡한 대사 경로를 조절하는 여러 개의 유전자에 의해 영향을 받는 것으로 추정할 수 있다.
유전력 분석
본 연구에서는 교배실생을 대상으로 양적형질의 특성을 나타 내는 과립중, 과립 횡경, 과립 종경, 당도, 산함량에 대한 유전력을 분석하였다. 일반적으로 유전력이 높은 형질들은 환경적 요인에 의한 영향을 덜 받는다(Ebadi et al. 2009). ‘Tano Red’와 ‘Ruby Seedless’의 교배실생에서 과립중과 과립 횡경, 과립 종경의 유전력은 각각 0.89, 0.82, 0.78으로 조사되었다(Table 3). Wei et al. (2002)은 이들 형질에 대한 유전력을 각각 0.63, 0.69, 0.68로 보고하였고, Ebadi et al. (2009)은 0.65, 0.54, 0.67으로 보고하였다. Firoozabady & Olmo (1987)Singh & Jalikop (1986) 등의 연구자에 의해 과립중의 유전력은 0.49-0.92의 범위 로 보고되었다. 본 연구에서 당도와 산 함량의 유전력은 각각 0.86과 0.93으로 Wei et al. (2002)이 보고한 0.48, 0.36과 Ebadi et al. (2009)이 보고한 0.29, 0.37보다 높았다. 이는 유전력 분석에 사용한 교배실생의 양친의 특성이 매우 유사하여 다른 보고에서와 달리 이들 형질에 대한 유전력이 높았던 것으로 추정되었다. 따라서 양친의 형태적 특성이 상이한 교배조합을 이용하여 주요 형질들에 대한 정밀한 추가적인 유전력의 비교 분석이 필요할 것으로 생각되었다.
Table 3
Estimate of heritability (h2) for five fruit quality traits in 252 progenies derived from a cross between ‘Tano Red’ and ‘Ruby Seedless’.
Table 3
Trait Heritability (h2)

   Berry weight 0.89
   Berry longitudinal diameter 0.82
   Berry transverse diameter 0.78
   Berry soluble solids 0.86
   Berry acidity 0.93
과립중과 종자 형태, 종자 무게, 종자 개수 형질 간의 상관분석
Table 4와 Fig. 3은 과립중과 종자 무게, 종자 형태, 종자 개수의 형질 간의 상관을 분석한 결과이다. 과립중은 종자 무게 (r = 0.486**)와 종자 형태(r = 0.483**) 간에 고도의 정의 상관이 인정되었으며 종자 개수(r = 0.211**)와도 정의 상관이 확인되었 다. 이는 과립중과 종자 무게, 종자 유무와 고도의 정의 상관이 있었다는 Ebadi et al. (2009)의 보고와 일치하였다. Ledbetter & Ramming (1989)은 과립에 있는 지베렐린 유사 물질 (gibberellin-like substance)의 농도와 종자 개수 간의 정의 상관 이 있음을 보고하였고, 유핵 품종에서는 종자에서 지베렐린 유사 물질이 생성되어 과립을 발달시키는데 대부분의 유핵 품종이 위단위결과성인 품종보다 큰 과립을 가지고 있다고 보고되었다 (Ebadi et al. 2009). 종자 무게는 예상대로 종자 형태와는 r = 0.607**, 종자 개수와는 r = 0.725**의 고도로 유의한 상관이 인정되었다. 또한 종자 형태와 종자 개수 간에는 r = 0.319**로 유의한 정의 상관이 확인되었다. 두 형질 간의 유전적 상관이 높고 특히 두 형질 간의 정의 상관이 존재할 때 한 형질에 대한 선발은 결국 두 번째 형질을 증가시키는 결과가 된다(Ebadi et al. 2009). 따라서 포도 과립중과 종자 형태와 무게 사이에 밀접한 상관이 존재함으로 과립중의 선발에 의해 종자형태와 종자 무게에 관한 간접 선발이 가능할 것으로 판단되었다.
Table 4
Correlation coefficient (r) between berry weight and berry seed weight, berry presence of seeds, and berry seed number in 252 progenies derived from a cross between ‘Tano Red’ and ‘Ruby Seedless’ (n = 252).
Table 4
Trait Berry weight Berry seed weight Berry presence of seeds

   Berry seed weight 0.486**z
   Berry presence of seeds 0.483** 0.607**
   Berry seed number 0.211** 0.725** 0.319**

z*, **Significant correlation coefficient at p=0.05, 0.01, respectively.

Fig. 3
Relationships between berry weight and berry seed weight (A), berry presence of seeds (B), and berry seed number (C) of 252 progenies derived from a cross between ‘Tano Red’ and ‘Ruby Seedless’.
KJBS-49-324_F3.gif
본 연구에서는 포도 교배실생을 이용하여 주요 과실 형질에 관한 조사를 통하여 대략적인 형질의 분포도를 파악하였다. 금후 에는 더 정밀한 분석을 통해 경도 등 일부 형질에 대한 객관적인 수치 제시와 연차간 변이 및 다른 교배조합들의 과실 특성 평가 결과와도 비교 분석할 예정이다. 또한 포도 육종에 있어서의 실질적인 적용 대상인 양적 형질에 대한 구명과 이에 대한 선발 지표를 개발하여 원하는 특성을 조기에 선발할 수 있는 시스템을 구축하고자 한다.
본 연구는 포도 품종의 육종 효율 증진을 위한 기초자료를 확보하기 위하여 ‘Tano Red’(유핵)와 ‘Ruby Seedless’(무핵)의 교배조합으로부터 얻은 실생 252주를 이용하여 주요 과실 형질 15항목에 대한 후대 실생의 분포 특성을 분석하였다. 교배실생의 과립의 과피색은 ‘Tano Red’와 ‘Ruby Seedless’의 과피색인 암자적색이 46.4%였고, 과립 모양은 단타원형이 48.4%였다. 대부분의 교배실생은 과즙이 많고 과육이 부드러운 ‘Tano Red’ 의 특성과 가깝게 분포되었다. 종자 형태는 완전한 것이 67.1%, 흔적만 있는 것이 30.1%, 종자가 없는 것이 2.8%를 차지하였고, 종자 무게가 0.15 g 미만인 교배실생이 84.9%로 대부분이었다. 15가지 과실 형질 중에서 과방의 착립 밀도, 과립경에서의 과립 분리 정도, 과립중, 종자 개수, 과립 종경, 과립 횡경, 당도, 산 함량은 정규분포를 나타내었다. 과립중과 과립 횡경, 과립 종경, 당도, 산 함량의 유전력은 각각 0.89, 0.82, 0.78, 0.86, 0.93이었 다. 과립중과 종자와 관련 있는 형질 간의 상관을 분석한 결과, 과립중은 종자 무게(r = 0.486**), 종자 형태(r = 0.483**), 종자 개수(r = 0.211**)와 정의 상관이 확인되었다. 종자 무게는 종자 형태(r = 0.607**), 종자 개수(r = 0.725**)와 고도로 유의한 상관이 인정되었다. 또한 종자 형태와 종자 개수(r = 0.319**) 간에는 유의한 정의 상관이 확인되었다. 본 결과는 포도 육종을 지원하기 위한 과실 품질과 관련된 양적형질유전자좌를 동정하 는데 이용될 수 있을 것으로 기대한다.
본 논문은 차세대바이오그린21사업(농생물게놈활용연구사 업단, 세부과제번호: PJ01185601)의 지원에 의해 이루어진 것임
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Evaluation of Major Fruit Traits in Progenies Derived from a Cross between ‘Tano Red’ and ‘Ruby Seedless’ Grapevines (Vitis spp.)
Korean. J. Breed. Sci.. 2017;49(4):324-333.   Published online December 1, 2017
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Evaluation of Major Fruit Traits in Progenies Derived from a Cross between ‘Tano Red’ and ‘Ruby Seedless’ Grapevines (Vitis spp.)
Korean. J. Breed. Sci.. 2017;49(4):324-333.   Published online December 1, 2017
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Evaluation of Major Fruit Traits in Progenies Derived from a Cross between ‘Tano Red’ and ‘Ruby Seedless’ Grapevines (Vitis spp.)
Image Image Image
Fig. 1 Fruits of ‘Tano Red’ (A), ‘Ruby Seedless’ (B), and progenies derived from a cross between ‘Tano Red’ and ‘Ruby Seedless’ (C).
Fig. 2 Phenotypic distribution of the 15 fruit quality traits of 252 progenies derived from a cross between ‘Tano Red’ and ‘Ruby Seedless’. Filled arrows represent ‘Tano Red’ values, while empty arrows represent ‘Ruby Seedless’ values.
Fig. 3 Relationships between berry weight and berry seed weight (A), berry presence of seeds (B), and berry seed number (C) of 252 progenies derived from a cross between ‘Tano Red’ and ‘Ruby Seedless’.
Evaluation of Major Fruit Traits in Progenies Derived from a Cross between ‘Tano Red’ and ‘Ruby Seedless’ Grapevines (Vitis spp.)

Fruit characteristics measured on 252 progenies derived from a cross between ‘Tano Red’ and ‘Ruby Seedless’.

Trait Abbreviation Unit Remark

Bunch density BND 1-9 1. Very loose (berries in grouped formation, many visible pedicels)
3. Loose (single berries with some visible pedicels)
5. Medium (densely distributed berries, pedicels not visible)
7. Dense (berries not readily movable)
9. Very dense (berries pressed out of shape)
Berry shape BS 1-9
2. Narrow elliptic
3. Elliptic
4. Round
5. Oblate
6.Ovate
7. Obtuse-ovate
8. Obovate
9. Arched
Berry weight BW g Weight of berry
Berry skin color (without bloom) BSC 1-6 1. Green-yellow
2. Rose
3. Red
4. Red-grey
5. Dark red-violet
6. Blue-black
Berry ease of detachment from BED 1-3 1. Difficult
pedicel 2. Slightly easy
3. Very easy
Berry ease of seeds separating BES 1-3 1. Difficult
from flesh 2. Slightly easy
3. Very easy
Berry juiciness of flesh BJF 1-3 1. Very slightly juicy
2. Slightly juicy
3. Very juicy
Berry firmness of flesh BFF 1-3 1. Soft
2. Medium
3. Firm
Berry seed weight BSW g Weight of seed
Berry presence of seeds BPS 1-3 1. Seedless (absent)
2. Rudimentary
3. Well developed
Berry seed number BSN 1-4 Seed number of berry
Berry longitudinal diameter BLD mm Longitudinal diameter of berry
Berry transverse diameter BTD mm Transverse diameter of berry
Berry soluble solids BSS oBrix Sugar content of berry
Berry acidity BA % Acid content of berry

Comparison of the fruit characteristics of the parents.

Parent Traitz

BND (1-9) BS (1-9) BW (g) BSC (1-6) BED (1-3) BES (1-3) BJF (1-3) BFF (1-3) BSW (g) BPS (1-3) BSN (0-4) BLD (mm) BTD (mm) BSS (°Brix) BA (%)

Tano Red 9 (very dense) 8 (obovate) 4.11 5 (dark red-violet) 3 (very easy) 3 (very easy) 3 (very juicy) 1 (soft) 0.16 3 (well developed) 3.2 21.73 18.08 11.80 0.76
Ruby Seedless 1 (very loose) 2 (narrow elliptic) 3.58 5 (dark red-violet) 2 (slightly easy) - 2 (slightly juicy) 2 (medium) - 2 (rudimentary) 0 20.85 16.85 19.56 0.64

zBND, bunch density; BS, berry shape; BW, berry weight; BSC, berry skin color; BED, berry ease of detachment from pedicel; BES, berry ease of seeds separating from flesh; BJF, berry juiciness of flesh; BFF, berry firmness of flesh; BSW, berry seed weight; BPS, berry presence of seeds; BSN, berry seed number; BLD, berry longitudinal diameter; BTD, berry transverse diameter; BSS, berry soluble solids; BA, berry acidity.

Estimate of heritability (h2) for five fruit quality traits in 252 progenies derived from a cross between ‘Tano Red’ and ‘Ruby Seedless’.

Trait Heritability (h2)

   Berry weight 0.89
   Berry longitudinal diameter 0.82
   Berry transverse diameter 0.78
   Berry soluble solids 0.86
   Berry acidity 0.93

Correlation coefficient (r) between berry weight and berry seed weight, berry presence of seeds, and berry seed number in 252 progenies derived from a cross between ‘Tano Red’ and ‘Ruby Seedless’ (n = 252).

Trait Berry weight Berry seed weight Berry presence of seeds

   Berry seed weight 0.486**z
   Berry presence of seeds 0.483** 0.607**
   Berry seed number 0.211** 0.725** 0.319**

z*, **Significant correlation coefficient at p=0.05, 0.01, respectively.

Table 1 Fruit characteristics measured on 252 progenies derived from a cross between ‘Tano Red’ and ‘Ruby Seedless’.
Table 2 Comparison of the fruit characteristics of the parents.

BND, bunch density; BS, berry shape; BW, berry weight; BSC, berry skin color; BED, berry ease of detachment from pedicel; BES, berry ease of seeds separating from flesh; BJF, berry juiciness of flesh; BFF, berry firmness of flesh; BSW, berry seed weight; BPS, berry presence of seeds; BSN, berry seed number; BLD, berry longitudinal diameter; BTD, berry transverse diameter; BSS, berry soluble solids; BA, berry acidity.

Table 3 Estimate of heritability (h2) for five fruit quality traits in 252 progenies derived from a cross between ‘Tano Red’ and ‘Ruby Seedless’.
Table 4 Correlation coefficient (r) between berry weight and berry seed weight, berry presence of seeds, and berry seed number in 252 progenies derived from a cross between ‘Tano Red’ and ‘Ruby Seedless’ (n = 252).

Significant correlation coefficient at p=0.05, 0.01, respectively.