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장수형 밀의 계통 특성 분석 I. 농업 형질과 관련 유전자 특성

Characterization of Korean Wheat Line with Long Spike. I. Agronomic Traits and Genetic Variations

Jae-Han Son1, Chon-Sik Kang1, Young-Keun Cheong1, Kyeong-Hoon Kim1, Hag-Sin Kim1, Jong-Chul Park1, Kyoung-Ho Kim1, Bo-Kyoung Kim1, Chul Soo Park2,*
Korean Journal of Breeding Science 2015;47(3):219-228.
Published online: August 31, 2015

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*Corresponding author (pcs89@jbnu.ac.kr, Tel: +82-63-270-2533, Fax: +82-63-270-2640)
• Received: July 27, 2015   • Revised: September 15, 2015   • Accepted: September 15, 2015

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  • One hundred sixty four wheat (Triticum aestivum L.) lines expressing the long spike were developed by the National Institute of Crop Science, Rural Development Administration. Six of 11 genes showed allelic variation among 164 lines and they were highly related to yield traits containing spike and kernel number, kernel length, 1000-kernel weight and grain yield. Analysis of allelic variation of each gene showed high correlation between allelic variation and agricultural characteristics. Coefficient of correlation between culm length and spike length was the highest 0.436. The heading date of the lines expressed Vrn-D1 allele was late and the lines expressed Vrn-D1 allele had longer spike length and higher number of kernel per spike than other lines. There was no significant difference in agronomic characteristics of analysis of Vrn-B1. Ppd-D1b was related to heading date and higher number of spike/m2. Ppd-D1a was corresponded to number of kernel per spike and 1000-kernel weight. Three haplotypes, Hap-L, Hap-6A-G and TaCwi-A1a, were related to 1000-kernel weight.
기상 이변과 병충해로 인해 밀 수량 안정성은 그 어느 때 보다 불안한 실정이기 때문에 수량 증대 및 안정성 증진은 매 우 중요한 육종목표라고 할 수 있다(Miralles & Slafer 2007, Rosegrant & Cline 2003). 밀 수량은 단위면적당 이삭수, 1 수립수, 종자 무게 영향을 가장 많이 받는데, 개별적인 종자의 무게는 차이는 있지만 이삭의 개수와 1수립수는 단위면적당 이삭수의 증가와 밀접한 관련이 있기 때문에 이삭 길이와 1수 립수의 증가는 밀 수량 증대에 직접적인 원인이 된다(Brancourt-Hulmel et al. 2003, Ledent & Stoy 1988, Ma et al. 2007, Perry & d’Antuono 1989). 그러므로 밀 육종 프로그램에 서 수량 증진을 위한 이삭에 달린 종자 개수 증가를 유전적으 로 조절하는 것이 중요하다고 할 수 있다(Reynolds et al. 2005, Fischer 2007, Foulkes et al. 2011, Reynolds et al. 2012).
국내 밀 생산량은 년간 44,000 톤 정도 생산되고 있지만, 자급률은 2.2% 정도로 매우 낮아 대부분 수입에 의존하고 있 는 실정이지만, 최근 국가 정책과 소비자 의식 변화 등에 의 해 밀 재배 면적과 생산량이 증가하고 있는 추세이다 (MAFRA 2014). 국내 밀 육종프로그램은 수량성 증진, 숙기 단축과 밀가루 품질 향상을 주요 목표로 품종 및 계통 선발과 육성이 이루어지고 있는데, 1970년 이후 35개의 품종이 육성 되었다(Kang et al. 2014). 이모작 남부지역에서 주로 재배되 고 있는 금강밀과 조경밀은 제분율이 높고 제빵 적성이 양호 한 백립계 경질밀이지만, 20년 전에 육성되어 현재 재배 농가 에서 요구하는 수량성에 미치지 못하여 재배면적이 점차 감 소하고 있다(Kang et al. 2014). 재배 농가의 요구에 부응하 는 수량성 높은 밀 품종 개발을 위하여 금강밀과 중국밀 자원 인 Xian83의 교배를 통해 이삭 길이와 1수립수가 많은 장수 형 계통을 개발하였다(Kang et al. 2013).
Atsmon과 Jacobs (1977)가 개발한 “Gigas”는 최대 30개 의 이삭을 갖는 장수형 밀로 종자 1개의 무게는 63 mg이고 한 소수당 9개의 종자가 열린다. 그러나 Gigas는 분얼이 왕성 하지 않고 억제되는 경향을 나타냈는데, 이것은 염색체 1AS 에 위치하고 있는 tin 열성유전자가 원인으로 알려져 있다 (Richards 1988, Spielmeyer & Richards 2004). tin은 수량 에 영향을 미치는 유전자로서 1수립수는 9% 정도 증가시키 지만, 수수와 립중을 각각 11%와 2% 감소시킨다고 보고되어 있다(Duggan et al. 2005). 장수형 밀은 국제밀옥수수연구소 (CIMMYT)의 밀과 비교했을 때, 1수영화수, 1수립수, 개개 의 립중이 각각 4%, 5%, 10%가 증가하는 것으로 보고되었 지만, 대비품종인 Bacanora에 비해 수량이 8% 감소하고, 수 수가 26% 감소하는 것으로 확인 되었다(Gaju et al. 2009, 2014).
수수 및 1수립수 등 수량성과 관련된 유전자 및 다양한 분 자마커 개발은 밀의 육종에 매우 유용하게 사용될 수 있다 (Bernardo 2008). 종실의 무게와 크기는 수량에 직접적으로 관련되어 있기 때문에 매우 중요하며 이와 관련된 유전자로 Sucrose synthase 2 (Sus2), grain width (GW2)와 cell-wall invertase (CWI) 등 3개가 보고되어 있다(Liu et al. 2012). 각각의 유전자형에 따라 TaSus2-2B의 Hap-H과 TaGW2-6A 의 Hap-6A-A는 천립중 증가와 매우 밀접하게 연관되어 있는 것으로 보고되어 있다(Jiang et al. 2011, Su et al. 2011). 또 한 Ma et al. (2012)은 TaCwi-A1의 분석을 통해 TaCwi-A1a 대립유전자의 천립중은 TaCwi-A1b 보다 많은 것을 확인하였 다. 반왜성 유전자(Rht)는 녹색혁명을 이끈 중요한 유전자로 서 서로 다른 대립유전자는 식물의 길이 생장에 서로 다른 영 향을 미친다(Börner et al. 1996). 특히 밀의 염색체 4B와 4D 에 위치하고 있는 Rht-B1b과 Rht-D1b는 DELLA 단백질을 암호화하고 있고 지베렐린산(GA3)에 둔감한 유전자이다 (Tosovic-Maric et al. 2008). 이들 반왜성 유전자는 간장을 감소시키고 고온 등 기후 변화 아래에서도 밀의 수량을 증대 를 기대할 수 있게 하였다(Flintham et al. 1997). 파성 (vernalization)과 일장(photoperiod)은 밀의 개화를 조절하는 주요한 인자이다. 이들은 생장온도와 반응을 통해 개화를 시 작하는 것과 다른 환경적인 자극을 통해 개화의 시기를 앞당 기거나 늦출 수 있다. Vrn-1 과 Ppd-1 등은 각각 파성과 일장 과 관련되어 있는 유전자로서 각각 3개의 상동유전자를 포함 하고, 5번과 2번 염색체에 위치하고 있다(Kamran et al. 2014). Vrn-1은 개화 시기와 식물의 키 등에 영향을 미치고 (Stelmakh 1998), Ppd-1는 일장 둔감형 유전자로서 일장에 민감한 유전자보다 개화 시기를 앞당기기 때문에 여름의 고 온과 건조 조건에서도 수량을 증대시킬 수 있는 중요한 유전 자이다(Snape et al. 2001).
국내 품종들에 대한 품질 관련 유전자와 같이 농업형질과 관련된 TaCwi-A1, TaGW2, TaSux2-2B, Rht, Ppd-1, Vrn-1 등 유전자의 연구가 이루어졌지만, 새로 개발된 장수형 밀 164계통에 대한 연구는 아직 없다. 따라서 다수성 밀 품종 개 발을 위한 장수형 밀의 농업적인 분석과 유전적인 구성요소 를 함께 분석하여 한국의 밀 육종에서 다수성 밀 품종개발을 위한 중요한 자원으로 활용하고자 하였다.
공시재료
2005년도에 숙기는 늦지만 이삭이 길고 립수가 많은 CIMMYT의 수집종인 Xian83을 모본으로 하고, 숙기가 빠르 고 다목적용 밀 품종인 금강밀을 부본으로 인공교배하여 IW20050259 조합을 육성하였다. 국립식량과학원 시험포장 (전북 익산시)에서 교잡종(F1)의 분리세대(F2) 종자를 국립식 량과학원 운봉출장소(전북 남원시)에서 일반적인 집단육종법 에 의해 내한성이 양호한 계통을 선발하였다. 잡종 제3세대 (F3)이후에는 국립식량과학원 시험포장에 집단 재배하여 이 삭이 길고 립수가 많은 계통을 선발하였다. 본 연구에는 잡종 제 8세대(F8) 164계통을 공시하였다. 부본으로 이용된 Xian83 은 CIMMYT에서 중국자원에 대한 붉은곰팡이병 평가를 위 해 도입되었지만, 평가 이후 폐기되어 농업형질 및 관련 표지 인자와의 비교는 부본인 금강밀과 비교했다.
농업형질 조사
농업 형질 조사는 농촌진흥청 농업 과학 기술 조사 분석 기 준(RDA 2012)에 준하여 실시하였다. 출수기는 전체 경수 중 에 40%가 출수한 날로 하였으며, 성숙기는 대부분의 이삭이 황화한 날로 조사하였다. 간장은 지면에서 이삭 목까지의 길 이로 나타내었고, 수장은 이삭 목에서 이삭 끝까지의 길이를 측정하였고, 단위 면적당 수수는 생육이 고른 3개소에서 25 cm × 50 cm에 있는 총 수수를 측정하여 1 m 2당 경수로 환산 하였다. 1수립수는 생육이 균일한 장소에서 무작위로 20수를 채취하여 탈곡 후 조사하였다. 리터중은 탈곡한 종자를 2.0 mm 종목체를 통과하지 않은 종자를 리터중 측정기를 이용하 여 측정하였으며, 천립중도 탈곡한 종자를 2.0 mm 종목체를 통과하지 않은 종자를 1,000립을 측정하여 무게를 조사하였 다. 수량은 수확 탈곡 건조 후(수분 14% 기준) 설립을 제외하 고 종실을 평량하여 10a당으로 환산하여 조사하였다.
Genomic DNA 추출
금강밀과 장수형 계통을 온실에 파종하여 3주 간 생육시켰 다. 유식물체의 엽을 채취하여 액체질소로 급속냉동 시킨 후 유발을 이용하여 미세분말로 분쇄한 뒤 -70°C에 보관하였다. Genomic DNA는 잎 분말(100 mg)로부터 식물용 Genomic DNA 분리시약(Solgent, Korea)을 이용하여 추출하였다.
PCR 분석
Rht-B1와 Rht-D1 유전자를 이용한 Polymerase chain reaction (PCR)은 Ellis et al. (2002)의 방법으로 수행되었다. Vrn-1 유전자 PCR 분석은 Yan et al. (2004), Fu et al. (2005), Santra et al. (2009), Zhang et al. (2012), Chen et al. (2013)의 연구방법에 따라 수행되었고, Ppd-1는 Beales et al. (2007)과 Bentley et al. (2011)의 방법을 이용하였다. PCR 생산물은 1.5% 아가로스젤에 전기영동하여 확인하였고, TaSus2-2B, TaGW2-6A, TaCwi-A1 등의 대립유전자 다양성 은 Jiang et al. (2011), Su et al. (2011)과 Ma et al. (2012) 의 방법에 의해 분석되었다.
통계 분석
모든 시험은 최소 3회 이상 반복을 두어 실시하였으며, 시 험성적은 SAS 프로그램(SAS Institute, Cary, NC)을 이용하 여 분산분석을 실시하고 LSD로 유의성을 검정하였으며, 품종 과 연차간 변이의 상호작용을 보기 위한 ANOVA (Analysis of variance) 분석은 GLM을 이용하였으며, 품종과 연차 변 이의 상호작용 성분은 오차항을 이용하였다. 상관관계는 피어 슨의 상관 계수를 이용하였으며, 통계적 유의 수준은 P<0.05 으로 실시하였다.
장수형 계통의 농업 형질
장수형 계통의 잎색은 녹색이며 폭은 금강밀에 비해 넓고 긴 편이다. 줄기 굵기는 중 정도이며 이삭은 방추형으로 까락 은 중간 정도이고, 종실 크기는 중 정도에서 대립크기의 분포 를 보였다(Fig. 1). 장수형 계통은 출수까지 평균 200.8일이 걸렸고, 금강밀(193일)보다 약 7일 정도 많은 일수가 요구되 었다. 장수형의 간장은 88.3 cm로 금강밀보다 약 16 cm 더 길었다(Table 1). 밀의 수량과 관련된 특징 중 이삭길이, 단위 면적 당 수수, 1수립수, 리터중, 천립중과 생산량 등은 각각 11.7 cm, 650.7개, 43.4개, 776.8 g, 44.6 g, 480.4 kg/10a으 로 나타났다. 비교 품종인 금강밀은 7.0 cm, 711.1개, 27.5개, 843.3 g, 45.1 g, 470.4 kg/10a였는데, 장수형 계통은 단위면 적당 수수, 리터중은 금강밀보다 낮았지만 이삭의 길이가 길 고, 1수립수가 많아 전체적인 수량은 금강밀보다 약 10 kg/10a 증수되는 것으로 나타났다(Table 1).
Table 1.
Agronomic traits in Korean wheat line with long spike and Keumkang.
Table 1.
Agronomic traits Korean wheat population with long spike lengthz (n = 164) Keumkang
Days to heading date 200.8 ± 1.3 (197.3 – 204.0) 193.0
Culm length (cm) 88.3 ± 6.8 (66.1 – 102.3) 72.1
Spike length (cm) 11.7 ± 1.1 (9.1 – 14.4) 7.0
Number of spike per m2 650.7 ± 117.0 (389.0 – 950.0) 711.1
Number of kernel per spike 43.4 ± 4.3 (30.8 – 54.7) 27.5
Test weight (g) 776.8 ± 16.9 (713.0 – 815.0) 843.3
1000-kernel weight (g) 44.6 ± 5.7 (31.6 – 55.9) 45.1
Grain yield (kg/10a) 480.4 ± 74.7 (243.2 – 654.1) 470.4

zValue indicates average and standard deviation of Korean wheat line with long spike and parenthesis includes ranges of Korean wheat line with long spike

장수형 계통의 농업적 형질 간의 상관관계 분석을 통해 간 장은 출수까지의 일수와 수장과 매우 높은 상관을 갖는 것으 로 나타났다. 이와 같은 결과는 장수형 계통이 금강보다 긴 간장과 수장을 갖고 있기 때문에 그 만큼 이들이 자랄 수 있 는 충분한 일장이 요구되기 때문인 것으로 생각된다. 단위면 적당 수수는 금강에 비해 50개가 적었지만, 1수립수와 전체 적인 수량은 금강보다 많았다. 장수형 계통의 종자 크기는 금 강밀과 비슷하고, 긴 이삭에 종자가 많이 형성되기 때문에 수 량이 많게 나온 것으로 생각된다(Fig. 1). 이와 같이 수장과 1수립수의 상관도 매우 높게 나타났다. 리터중은 천립중과 생 산량과 매우 높은 정의상관을 나타냈지만, 출수까지의 일수와 는 부의상관을 나타냈다. 뿐만 아니라 천립중과 생산량도 출 수까지 일수와 고도의 부의상관을 나타냈다(Table 2).
Fig. 1.
Image of spike and spikelet (A), heading stage (B), maturing stage (C) and grain (D) of Korean wheat 164 line with long spike and Keumkang.
KJBS-47-219_F1.gif
Table 2.
Correlation coefficients among agronomic traits of Korean wheat line with long spike.
Table 2.
Parametersz DHD CL SL NSP NKPS TW TKW
CL 0.30***
SL 0.13 0.44***
NSP 0.21** 0.20** –0.10
NKPS 0.03 0.08 0.37*** –0.01
TW –0.50*** –0.17* –0.16* –0.17* –0.15
TKW –0.36*** 0.16* 0.20** –0.48*** –0.13 0.40***
GY –0.38*** –0.11 0.06 –0.09 0.14 0.29*** 0.26***

zDHD = days to heading date, CL= culm length, SL= spike length, NSP= number of spike per m2, NKPS= number of kernel per spike, TW= test weight, TKW=1000-kernel weight, GY= grain yield

y* indicates significance at 0.05 level, ** at 0.01 level, and *** at 0.001 level

잎의 크기는 생산량과 바이오매스 생산에 매우 중요한 요 소이다(Gonzalez et al. 2009). 잎의 크기와 관련된 생장은 세 포증식과 세포신장의 중첩된 두 단계로 나뉘어져 있고, 큰 잎은 세포의 크기와 관련이 있는 것으로 보고되어 있다(Anastasiou & Lenhard 2007). Shin et al. (2015)은 금강과 비교한 결과 세포의 크기는 장수형 계통이 금강보다 30% 증가했지만 세 포의 수나 광합성 효율은 차이가 없었고, 장수형 계통의 수장 은 금강보다 6.0 cm 이상 긴 것으로 보고하였다. 본 연구에서 수장과 1수립수와 고도의 정의상관을 나타내는 것을 알 수 있 다. 수장이 길수록 립수가 많고, 줄기는 길었지만 도복에 강한 특성을 나타냈다. 장수형 계통의 줄기 두께는 5.02 cm로서 금강 4.35 cm보다 약 0.8 cm가 더 두껍기 때문에 많은 립수 에도 불구하고 줄기가 수를 지탱하는 것으로 생각된다(Son et al. 2014). 장수형 계통의 출수기는 금강보다 8일에서 10일 정도 늦고, 개화기는 5일에서 6일 정도 늦다고 보고되어 있다 (Son et al. 2014). 리터중, 천립중과 생산량 등은 출수까지의 일수와 고도의 부의 상관을 나타내는 것은 출수기와 개화가 늦기 때문에 수확기까지 요구되는 충분한 시간이 우리나라 기후 여건에서 충족될 수 없는 환경적 영향에 의한 것으로 생 각되고, 이들 장수형 계통의 품종과 유전자원으로서 활용을 위해서는 숙기 단축이 매우 중요하게 요구될 것으로 판단된다.
밀의 농업적 특성과 관련된 유전자 분석
TaCwi-A1, TaGW2, TaSux2-2B, Rht-B1, Rht-D1, Vrn-A1, Vrn-B1, Vrn-D1, Ppd-A1, Ppd-B1, Ppd-D1등 밀의 농업특 성과 관련이 있는 유전자 11개의 다양성을 분석하여 금강밀 과 비교하였다. PCR을 통해 증폭된 밴드는 167에서 1,671 bp 범위 안에서 확인되었다(Fig. 2). Rht-B1, Rht-D1, Vrn-B1, Ppd-A1, Ppd-B1 등 5개 유전자는 164개 장수형 계통에서 차 이가 없고 모두 같은 유전자형으로 나타났다. Rht-B1과 Rht-D1 의 유전자형은 두 가지(237, 264bp)로서 장수형 계통은 Rht- B1b와 Rht-D1b (null-type) 금강밀은 Rht-B1a와 Rht-D1a로 서로 다른 것을 확인하였다. 장수형 164 계통과 금강밀은 모 두 Vrn-B1b, Ppd-A1b, Ppd-B1b를 갖고 있었으며, 각각 1,149 bp, 453 bp, 1,292 bp의 크기의 밴드로 증폭되었다 (Fig. 2).
Fig. 2.
Agarose gel electrophoresis of PCR amplified Rht-B1 and Rht-D1 (A), Vrn-1 (B), Ppd-1 (C) and TaSus2-2B, TaGW2 and TaCwi-A1 alleles (D). M, molecular size marker; 1-12, Korean wheat line with long spike (GHW001~GHW012).
KJBS-47-219_F2.gif
Vrn-A1과 Vrn-D1 유전자는 각각 Vrn-A1a와 Vrn-A1b, Vrn-D1a와 Vrn-D1b의 유전자 형을 갖고 있었으며 1,068bp 와 522 bp의 크기로 증폭되었다(Fig. 2). Vrn-A1a는 금강을 포함하여 156 계통에서 확인되었고, Vrn-A1b는 8개 계통에 서 확인되었다. Vrn-D1은 금강과 157 계통에서 Vrn-D1b (997 bp), 7개 계통에서 Vrn-D1a (1,671 bp)가 확인되었다. Ppd-D1 또한 Ppd-D1a (288 bp)과 Ppd-D1b (414 bp) 두 가 지 유전자형을 갖고 있었고, 금강과 130개 계통이 Ppd-D1a, 34 계통이 Ppd-D1b를 갖고 있었다. 164 계통의 농업적 특성 과 비교해 보았을 때, Vrn-A1a와 Vrn-A1b는 유전자형에 따 른 특성차이에 유의성이 없었고, Vrn-D1은 출수기까지 일수, 수장, 1수립수에서 유의성의 차이가 나타났다. Vrn-D1이 Vrn-D1a를 갖는 계통이 출수까지 일수와 수장, 1수립수가 많 게 나타났다. 일장과 관련이 있는 Ppd-D1 유전자 분석에서 Ppd-D1a와 Ppd-D1b에 따라 출수까지 일수, 단위면적당 수 수, 1수립수, 리터중을 제외한 다른 특성에서 Ppd-D1a를 갖 는 계통이 많게 나타났다(Table 3). 파성과 일장 관련 유전자 는 earliness per se (Eps) 유전자와 함께 밀의 환경에 대한 적 응을 조절하는 한 요인이다(Kato & Yamagata 1988). 이들 유전자는 밀의 개화를 조절하는 것으로서 특정 환경에서 유 전자형에 따라 개화 시기를 앞당기거나 늦추는 것을 조절한 다(Worland 1996). 특히 파성은 보통밀에서 Vrn-A1, Vrn-B1, Vrn-D1, Vrn4, Vrn-B3 등 적어도 5개 대립유전자에 의해 조 절되는 것으로 알려져 있고(Goncharov 2003, McIntosh et al. 1998, Pugsley 1971), 본 연구에서 Vrn-D1은 7개 계통에서 확인되었고, 수장과 1수립수와 매우 깊게 연관되어 있는 것으 로 확인되었다. Ppd-D1b를 가지는 34 계통은 출수기까지 일 수와 수수가 많았지만 전체적인 수량은 Ppd-D1a보다 적었 다. Ppd-D1a는 1수립수, 리터중, 천립중, 생산량에서 모두 많 은 것으로 확인되었기 때문에 Ppd-D1a는 수량에 직접적인 연관이 있는 것으로 생각된다.
Table 3.
Difference in agronomic traits of Korean wheat line with long spike carrying different vernalization and photoperiod genotypes using pair-wise t-test.
Table 3.
Agronomic Traitsz Vernalization Photoperiod
Vrn-A1 Vrn-D1 Ppd-D1
vrn-A1 Vrn-A1b vrn-D1 Vrn-D1a Ppd-D1a Ppd-D1b
No. Line 156 8 7 157 130 34
DHD 200.8ay 200.4a 200.7b 201.3a 200.5b 201.8a
CL (cm) 88.3a 90.0a 88.5a 85.7a 88.0a 89.5a
SL (cm) 11.7a 11.8a 11.8a 10.8b 11.8a 11.5a
NSP 651.0a 645.8a 673.8a 649.7a 640.3b 690.5a
NKPS 43.4a 42.6a 43.6a 39.3b 43.7a 41.9b
TW (g) 777.2a 769.7a 776.3a 787.8a 778.5a 770.2b
TKW (g) 44.7a 42.4a 44.7a 42.7a 45.6a 40.5b
GY (kg/10a) 479.9a 490.1a 481.3a 460.0a 501.1a 401.1b

zDHD = days to heading date, CL= culm length, SL= spike length, NSP= number of spike per m2, NKPS= number of kernel per spike, TW= test weight, TKW=1000-kernel weight, GY= grain yield

yValues followed by same letters within same characteristics at the same locus are not significantly different at P<0.05

TaCwi-A1 유전자 또한 두 가지 유전자형을 가지고 있다. 장수형 164계통 중 75 계통은 a형(402 bp), 89 계통은 b형 (404 bp)을 갖고 있었다. TaGW2 유전자는 Hap-6A-A와 Hap-6A-G의 두 대립유전자를 갖고 있으며, Hap-6A-A는 167 bp, Hap-6A-G는 218 bp의 크기로 각각 확인되었다. 24 계통에서 Hap-6A-A가 나타났고 나머지 계통에서 Hap-6A-G 로 나타났다. TsSus2-2B 유전자는 Hap-L과 Hap-H형의 유전 자형을 갖고 있고, 각각 318 bp와 423 bp의 밴드로 증폭되었 다. Hap-L과 Hap-H의 두 유전자형은 164계통에서 각각 134 계통과 30 계통에서 확인되었다. Hap-L, TaCwi-A1b, Hap-6A-G 등 3개의 유전자형을 갖는 계통의 천립중은 다른 유전자형에 비해 현저하게 높게 나타났다(Fig. 3). 본 연구를 통해 TaCwi-A1, TaGW2, TaSus-2B, Vrn-D1, Ppd-D1 등 5개 유전자는 밀의 농업형질 확인을 위한 분자마커로 매우 유용하게 사용될 수 있을 것으로 생각된다. TaCwi-A1 유전자는 저장조직 발달, 탄소 구배, 종자무게와 깊게 관련되어 있는 효소로 알려져 있 는 cell wall invertase (CWI)를 암호화 하고 있다(Miller & Chourey 1992, Sturm 1999, Tang et al. 1999). TaCwi-A1 은 밀의 염색체 2번에 위치하고 있으며 open reading frame (ORF)는 총 1,767 bp로서 7개의 엑손과 6개의 인트론으로 구 성되어 있다(Ma et al. 2012). TaSus2 유전자는 염색체 2A, 2B, 2D에 위치하고 있으며 sucrose synthase 2 (Sus2)를 암 호화하고 있다(Jiang et al. 2011). TaSus2는 내배유에서 발 현되는 유전자로서 유전형에 따라 농업적 형질이 다르게 나 타나는 것으로 알려져 있다. 이 유전자는 중국 밀 품종 육종 단계에서 분자표지이용선발(MAS)을 위해 평가되었다(Jiang et al. 2011). TaGW2는 종자무게와 연관되어 있는 것으로 보 고되었고, 벼의 OsGw2와 상동성이 높은 것으로 발견되었다.
Fig. 3.
Difference in 1,000-kernel weight according to the genotype of TaSus2-2B (A), TaGW2 (B) and TaCwi-A1 alleles (C) in Korean wheat line with long spike. Each bar represents the mean ± standard error. Bars with different letters above are significantly different at p<0.05.
KJBS-47-219_F3.gif
TaGW2-6A의 유전자 코딩 지역의 염기서열 내에서는 변이가 발견되지 않았고, 프로모터 지역에서 2개의 single nucleotide polymorphisms (SNPs)가 발견되어 두 가지 유전자형을 갖게 된 것으로 보고되었다(Su et al. 2011). 본 연구 결과를 통해 TaSus2는 천립중과 관련이 있는 것을 알 수 있었다. 이들 유 전자형과 농업형질이 깊은 연관이 있음이 확인되었지만 생산 량과 관련된 여러 형질은 환경적인 영향도 많이 받기 때문에 장수형 계통에 대한 연구가 더욱 필요할 것으로 생각된다.
국내 밀의 수량 증진을 위해 이삭 길이가 길고 1수립수가 많은 장수형 164계통(F8)을 육성하였고, 본 연구에서는 이들 계통의 주요 농업형질과 이들 특성과 관련 있는 표지인자에 대한 평가를 수행하였다. 장수형 계통은 모본인 금강밀과 비 교했을 때, 단위면적당 수수와 리터중은 적고, 파종 후 출수까 지 일수가 길었고, 간장, 수장, 1수립수와 단위면적당 생산량 은 높게 나타났다. 농업 특성 관련 표지인자를 평가한 결과, 장수형 계통은 Rht-B1, Rht-D1, Vrn-B1, Ppd-A1, Ppd-B1 유전자 좌에서는 같은 유전자를 지닌 것으로 나타났다. 장수 형 계통 중 Vrn-D1를 지닌 계통은 Vrn-D1a를 지닌 계통에 비하여 출수에 걸리는 시간이 길었지만, 이삭길이와 1수립수 가 많은 것으로 나타났다. 하지만 Vrn-B1 유전자 변이는 장 수형 계통의 농업형질의 차이가 없는 것으로 나타났다. 장수 형은 Ppd-D1유전자 변이에 따른 간장이나 이삭 길이는 차이 가 없었지만, 출수 일수와 수수는 Ppd-D1b를 지닌 계통이 많 았지만, 1수립수, 천립중, 리터중과 수량은 Ppd-D1a를 지닌 계통이 높았다. 종실 특성을 나타내는 표지인자 변이는 출수 일수, 간장, 수장, 분얼이나 1수립수 및 수량에는 영향을 주지 않았지만 TaSus2-2B유전자의 Hap-L타입, TaGW2유전자의 Hap-6A-G타입과 TaCwi-A1a 가 각각의 대립 유전자형보다 천립중이 높게 나타났다. 향후 재배환경이 장수형의 농업형질 및 품질에 미치는 영향에 대한 평가가 필요하고, 만숙과 분얼 이 적은 장수형의 단점을 보완하기 위한 지속적인 연구가 필 요하다.
본 연구는 2015년도 농촌진흥청 국립식량과학원 박사후연 수과정지원사업(PJ008756012015)과 농촌진흥청 연구사업 (세부과제명: 용도별 초다수성 신초형 계통 육성, 세부과제번 호: PJ009230012015에 의해 이루어진 것임.
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Characterization of Korean Wheat Line with Long Spike. I. Agronomic Traits and Genetic Variations
Korean. J. Breed. Sci.. 2015;47(3):219-228.   Published online September 30, 2015
DOI: https://doi.org/10.9787/KJBS.2015.47.3.219
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Characterization of Korean Wheat Line with Long Spike. I. Agronomic Traits and Genetic Variations
Korean. J. Breed. Sci.. 2015;47(3):219-228.   Published online September 30, 2015
DOI: https://doi.org/10.9787/KJBS.2015.47.3.219
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Characterization of Korean Wheat Line with Long Spike. I. Agronomic Traits and Genetic Variations
Image Image Image
Fig. 1. Image of spike and spikelet (A), heading stage (B), maturing stage (C) and grain (D) of Korean wheat 164 line with long spike and Keumkang.
Fig. 2. Agarose gel electrophoresis of PCR amplified Rht-B1 and Rht-D1 (A), Vrn-1 (B), Ppd-1 (C) and TaSus2-2B, TaGW2 and TaCwi-A1 alleles (D). M, molecular size marker; 1-12, Korean wheat line with long spike (GHW001~GHW012).
Fig. 3. Difference in 1,000-kernel weight according to the genotype of TaSus2-2B (A), TaGW2 (B) and TaCwi-A1 alleles (C) in Korean wheat line with long spike. Each bar represents the mean ± standard error. Bars with different letters above are significantly different at p<0.05.

Fig. 1.

Fig. 2.

Fig. 3.

Characterization of Korean Wheat Line with Long Spike. I. Agronomic Traits and Genetic Variations

Agronomic traits in Korean wheat line with long spike and Keumkang.

Agronomic traits Korean wheat population with long spike lengthz (n = 164) Keumkang
Days to heading date 200.8 ± 1.3 (197.3 – 204.0) 193.0
Culm length (cm) 88.3 ± 6.8 (66.1 – 102.3) 72.1
Spike length (cm) 11.7 ± 1.1 (9.1 – 14.4) 7.0
Number of spike per m2 650.7 ± 117.0 (389.0 – 950.0) 711.1
Number of kernel per spike 43.4 ± 4.3 (30.8 – 54.7) 27.5
Test weight (g) 776.8 ± 16.9 (713.0 – 815.0) 843.3
1000-kernel weight (g) 44.6 ± 5.7 (31.6 – 55.9) 45.1
Grain yield (kg/10a) 480.4 ± 74.7 (243.2 – 654.1) 470.4

zValue indicates average and standard deviation of Korean wheat line with long spike and parenthesis includes ranges of Korean wheat line with long spike

Correlation coefficients among agronomic traits of Korean wheat line with long spike.

Parametersz DHD CL SL NSP NKPS TW TKW
CL 0.30***
SL 0.13 0.44***
NSP 0.21** 0.20** –0.10
NKPS 0.03 0.08 0.37*** –0.01
TW –0.50*** –0.17* –0.16* –0.17* –0.15
TKW –0.36*** 0.16* 0.20** –0.48*** –0.13 0.40***
GY –0.38*** –0.11 0.06 –0.09 0.14 0.29*** 0.26***

zDHD = days to heading date, CL= culm length, SL= spike length, NSP= number of spike per m2, NKPS= number of kernel per spike, TW= test weight, TKW=1000-kernel weight, GY= grain yield

y* indicates significance at 0.05 level, ** at 0.01 level, and *** at 0.001 level

Difference in agronomic traits of Korean wheat line with long spike carrying different vernalization and photoperiod genotypes using pair-wise t-test.

Agronomic Traitsz Vernalization Photoperiod
Vrn-A1 Vrn-D1 Ppd-D1
vrn-A1 Vrn-A1b vrn-D1 Vrn-D1a Ppd-D1a Ppd-D1b
No. Line 156 8 7 157 130 34
DHD 200.8ay 200.4a 200.7b 201.3a 200.5b 201.8a
CL (cm) 88.3a 90.0a 88.5a 85.7a 88.0a 89.5a
SL (cm) 11.7a 11.8a 11.8a 10.8b 11.8a 11.5a
NSP 651.0a 645.8a 673.8a 649.7a 640.3b 690.5a
NKPS 43.4a 42.6a 43.6a 39.3b 43.7a 41.9b
TW (g) 777.2a 769.7a 776.3a 787.8a 778.5a 770.2b
TKW (g) 44.7a 42.4a 44.7a 42.7a 45.6a 40.5b
GY (kg/10a) 479.9a 490.1a 481.3a 460.0a 501.1a 401.1b

zDHD = days to heading date, CL= culm length, SL= spike length, NSP= number of spike per m2, NKPS= number of kernel per spike, TW= test weight, TKW=1000-kernel weight, GY= grain yield

yValues followed by same letters within same characteristics at the same locus are not significantly different at P<0.05

Table 1. Agronomic traits in Korean wheat line with long spike and Keumkang.

Value indicates average and standard deviation of Korean wheat line with long spike and parenthesis includes ranges of Korean wheat line with long spike

Table 2. Correlation coefficients among agronomic traits of Korean wheat line with long spike.

DHD = days to heading date, CL= culm length, SL= spike length, NSP= number of spike per m2, NKPS= number of kernel per spike, TW= test weight, TKW=1000-kernel weight, GY= grain yield

* indicates significance at 0.05 level, ** at 0.01 level, and *** at 0.001 level

Table 3. Difference in agronomic traits of Korean wheat line with long spike carrying different vernalization and photoperiod genotypes using pair-wise t-test.

DHD = days to heading date, CL= culm length, SL= spike length, NSP= number of spike per m2, NKPS= number of kernel per spike, TW= test weight, TKW=1000-kernel weight, GY= grain yield

Values followed by same letters within same characteristics at the same locus are not significantly different at P<0.05

Table 1.

Table 2.

Table 3.

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