Research Advances in Wheat Breeding and Genetics for Fusarium Head Blight Resistance

Myoung-Hui Lee; Sumin Hong; Kyeong-Min Kim; Sun-Hwa Kwak; Changhyun Choi; Chon-Sik Kang; Chul Soo Park; Youngjun Mo; Kyeong-Hoon Kim

doi:10.9787/KJBS.2023.55.3.195

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밀 붉은곰팡이병 저항성 유전 육종 주요 연구동향

이명희^1,†, 홍수민^2,†, 김경민¹, 곽순화², 최창현¹, 강천식¹, 박철수², 모영준², 김경훈^1,*

Research Advances in Wheat Breeding and Genetics for Fusarium Head Blight Resistance

Myoung-Hui Lee^1,†, Sumin Hong^2,†, Kyeong-Min Kim¹, Sun-Hwa Kwak², Changhyun Choi¹, Chon-Sik Kang¹, Chul Soo Park², Youngjun Mo², Kyeong-Hoon Kim^1,*

Korean Journal of Breeding Science 2023;55(3):195-217.
Published online: September 1, 2023

DOI: https://doi.org/10.9787/KJBS.2023.55.3.195

¹국립식량과학원

²전북대학교 작물생명과학과

¹National Institute of Crop Science, Rural Development Administration, Wanju, 55365, Republic of Korea

²Department of Crop Science and Biotechnology, Jeonbuk National University, Jeonju, 54896, Republic of Korea

*Corresponding Author (E-mail: k2h0331@korea.kr, Tel: +82-63-238-5457, Fax: +82-63-238-5463)

†

Author Contributions: Myoung-Hui Lee and Sumin Hong contributed equally.

• Received: April 14, 2023 • Revised: May 1, 2023 • Accepted: June 25, 2023

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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적요

Fusarium head blight (FHB) is a severe disease of wheat, mainly caused by Fusarium graminearum, which greatly reduces wheat production and directly affects human and animal health due to the mycotoxins produced in wheat grains. To develop high-quality, stable yields, and mycotoxin-free crop, it is essential to first understand the genetic basis of wheat FHB-resistance, and to design molecular markers facilitating the selection of FHB-resistant varieties. However, despite extensive global research efforts, genetic research and marker development for the selection of FHB-resistant varieties, in Korea, are insufficient. Here, we summarize recent studies on FHB-resistance genes, resistance resources, quantitative trait locus analysis, and genome-wide association studies to enhance our understanding of FHB and the breeding of FHB-resistant domestic wheat cultivars.

서 론

밀의 붉은곰팡이병(Fusarium head blight, FHB)은 주로 Fusarium graminearum (F. graminearum)에 의해 겨울철 고온과 출수기에 잦은 강우로 인해 상대습도가 80% 이상인 날이 3일이상 지속되면 발생하며 수량 감소와 품질 저하를 일으킨다(Azimi et al. 2020, Beak et al. 2020, Ryu et al. 2011, Shin et al. 2014, Shin et al. 2018). 그리고 FHB 병원균에 감염된 곡물에서 생산되는 트리코테신(trichothecenes, TCT)에 속하는 독소는 가축은 물론 사람에게도 중독증 피해를 일으켜 우리나라를 포함한 아시아는 물론 유럽, 북⋅남미 등 전세계적으로 관심을 갖는 주요 관리 병해이다(Buerstmayr et al. 2020, Ghimire et al. 2020, Khan et al. 2020, Ma et al. 2020). 최근 들어 기후변화로 인한 잦은 이상 기후 발생으로 밀에서 발생 빈도와 심각성이 증가되고 있다. Fusarium은 고온 다습한 조건에서 토양에 잔류하고 있는 포자 형태로 공기중으로 방출되어 식물체의 꽃가루에 붙어 식물체를 감염시키며, 살아있는 식물체는 물론 죽은 식물 조직에서도 생존하기 때문에 경운이 충분하지 않은 최소 경운이나 무경운 재배로 윤작하는 경우는 피해가 최대 10-70%까지 증가된다(Matthies & Buchenauer 2000, Shah et al. 2017).

밀에서 FHB 발생으로 인한 경제적 손실은 매우 심각하다. “wheat scab”으로 알려진 최초 보고는 1884년 영국의 대규모 감염(epidemic)이며, 미국은 1891년에 일리노이주에서 대규모 감염이 처음 보고되었다. 1990년대에는 미국, 캐나다, 중국, 유럽과 남미에서 대규모 피해가 발생하였다(Buerstmayr et al. 2020, Ma et al. 2020, Zhu et al. 2019). 미국은 1991년부터 1997년까지 FHB에 의한 수량 감소로 인한 직접적인 손해액은 약 13억달러였으며, 부가적인 산업피해를 합치면 약 48억달러에 달한다. 그리고 1998년에서 2007년까지 미국 대평원 지역과 캐나다 맥류 재배지역의 직접적인 피해 금액은 27억 달러로 추정된다(Bai & Shaner 2004, Gorash et al. 2021). 중국은 1950년 이후 전체 수확량 손실이 10-50%에 달하는 대규모 감염이 30회 이상 보고되었으며, 2012년에는 200만톤 이상의 피해를 입었다(Bai et al. 2018, Ma et al. 2020, Wilson et al. 2017). 우리나라는 1963년에 남부지역에서 대발생한 맥류 FHB으로 평균 50%, 심한 지역은 80% 이상 수량이 감소하였고, 식량 및 사료 부족과 이병곡물 섭취로 인한 인축의 심각한 중독증이 발생하였다. 2012년 호남 지역 맥류 FHB 발생 조사 결과, 전남지역에서는 해남군이 25.7%으로 가장 심했으며, 전북지역에서는 순창군이 14% 발생으로 가장 높았다. FHB 방제를 위한 다양한 약제가 사용되고 있고, 국내에는 캡탄(Captan), 메트코나졸(metconazole), 플루디옥소닐(fludioxonil) 수화제와 디페노코나졸⋅프로피코나졸(difenoconazole⋅propiconazole) 유제가 등록되어 있다(http://pis.rda.go.kr). 윤작, 경운 등 경작 방법에 의해서도 FHB 발생을 줄일 수 있지만 FHB의 방제에 가장 경제적이며 효과적인 방법은 저항성 품종을 육종하는 것이다(Boutigny et al. 2008, Hao et al. 2020, Jones 2000, Nakajima 2010, Park et al. 2012, Paul et al. 2008). 그러나 복잡한 밀 게놈, 양적 유전, 표현형의 가변성 및 숙주-병원체 상호 작용의 불분명한 저항성 메커니즘과 효과적인 저항성 유전자의 부족으로 인해 밀 품종 육종이 쉽지 않다(Bai & Shaner 1996, Son et al. 2013). 현재 주요 밀 생산국들은 FHB 저항성 증진을 위한 유용자원 탐색 및 저항성 품종 육성과 육종 효율성 증진을 위한 표지인자 개발에 많은 노력을 기울이고 있다(Buerstmayr et al. 2020, Ghimire et al. 2020, Gorash et al. 2021, Hao et al. 2020, Khan et al. 2020, Li et al. 2020, Ma et al. 2020, Zhu et al. 2019). FHB는 국내에서도 발생이 높아지고 있고 국산밀 안정적 공급을 위해서 반드시 관리해야 하는 병해이다(Beak et al. 2020, Chung 1975, Ryu et al. 2011, Shin et al. 2014, Shin et al. 2018). 국내에서도 FHB 심도와 mycotoxin 축적 평가를 통해 FHB평가가 이루어지고 있으며 FHB 중도저항성 밀 품종들이 개발되었다(Baek et al. 2022, Jung et al. 2011, Kang et al. 2014, Kim et al. 2019, Kim et al. 2021, Shin et al. 2014). 그리고 기존에 보고된 SSR marker와 STS marker를 이용한 저항성 품종 선발 결과가 보고되었지만 FHB 관련 유용유전자 탐색 및 선발 마커 개발 연구는 매우 미흡한 실정이다. 본 연구는 이러한 연구 동향 정리를 통하여 국산밀의 FHB 저항성 증진에 기여하고자 한다.

본 론

붉은곰팡이병 병원균 생활사

우리나라에서는 FHB가 전국에 걸쳐 발병하지만 주로 포자를 퍼트리기 좋은 온난 다습한 이모작 지대에서 많이 발생한다. 특히 4월 하순과 5월 상순의 밀 출수기와 유숙기 기간의 잦은 강우로 습도가 높아지면 이삭의 일부나 전체가 갈색으로 변하고 성숙이 되면 붉은색의 분생포자층(acervulus)이 생긴다. 심한 경우에는 출수가 안되거나 지연되고, 기형 이삭이 발생하기도 하며 양분이 이삭으로 가는 통로가 막혀 종실이 제대로 채워지지 않고 붉은색이나 검은색으로 변한다(Shah et al. 2017). Fusarium은 FHB외에도 파종 직후 종자를 썩게 하거나 유묘의 근부에 침입해서 뿌리를 썩게 하고, 잎이 오그라들며 작아지는 현상을 나타나기도 한다(Karlsson et al. 2021).

Fusarium 종은 분생포자를 생산하며 몇몇 종은(예, F. graminearum, F. avenaceum) 유성생식으로 자낭포자를 가지고 있다. 밀의 대표적인 FHB 병원균은 F. graminearum Schwabe (teleomorph: Gibberella zeae)인데, F. graminearum은 무성생식 단계의 포자에 대한 학명이고, Gibberella zeae (teleomorph)는 유성생식으로 자낭포자를 만드는 세대에 대한 학명이다(Geiser et al. 2013, Leslie & Summerell 2006). 무성생식 포자인 분생포자(macroconidia)와 유성생식 포자인 자낭포자(ascospore)가 주요 감염원이다(Shah et al. 2018). 분생포자는 퍼지는 거리가 짧고, 자낭포자는 바람이나 곤충에 의해 널리 퍼지며, FHB에 감염된 작물 부스러기나 토양 잔류물 등도 감염원으로 알려져 있다(Dweba et al. 2017). FHB 병원균의 발육온도는 10-30℃이며 최적온도는 25℃이고, 발육에 필요한 pH는 2.5-8.5이며 최적 pH는 4.0-4.5 이다. 월동이 가능하기 때문에 토양이나 잔류물에 부착되었다가 자낭포자를 방출시켜 포자를 공기중으로 퍼트려 어린 이삭이나 수술에 붙어 감염을 일으킨다(Beyer et al. 2005, Dweba et al. 2017, Fulcher et al. 2019, Osborne & Stein 2007) (Fig. 1). 대규모 단작 재배 및 옥수수 후작 밀 무경운 재배 확대와 최근 기후변화로 인해서 FHB 감염은 점점 확대될 것으로 전망하고 있다(Fulcher et al. 2019, Soresi et al. 2017, Zhang et al. 2014).

붉은곰팡이병 병원균의 독성

FHB 병원균에 의한 독성은 체중감소, 구토, 복통과 설사를 일으키고 심한 경우에는 사망에 이르게 되는데, 트리코테신(trichothecenes, TCT)의 B 타입에 속하는 밀 FHB 병원균 독소는 데옥시니발레놀(deoxynivalenol, DON), 니발레놀(nivalenol, NIV), 제랄레논(zearalenone, ZEN), 모닐리포민(moniliformin, MON)과 다양한 아세틸 유도체들이 있다(Osborne & Stein 2007, Soresi et al. 2017, Ward et al. 2008). 대표적 독소는 DON과 NIV이며 독성은 NIV가 DON보다 강하지만 주로 발생되는 밀 FHB 병원균 독소에 의한 독성은 주로 DON이기 때문에 DON에 의한 피해가 많으며, TCT B-타입 독소는 열 변성에 안정적이어서 식품 제조시 가열에 의해서도 독소가 제거되지 않기 때문에 세계 각국은 인축 피해를 최소화하기 위해 일일 섭취 한계량 등을 규정하고 있다(Ji et al. 2019, Torres et al. 2019). 이들 독소 피해를 줄이기 위한 DON과 NIV의 다양한 유도체들 생합성에 관련된 유전자 조절 연구와 카로테노이드, 항진균 펩타이드, 과산화 수소 및 지방 산화물이 해독(detoxification)에 미치는 영향에 대한 연구가 수행되고 있으며, 최근에는 감염된 식물 대사에서 생성되는 접합된(conjugated) 독소 형태인 가려진 곰팡이 독소(masked mycotoxins)도 보고되었다(Boutigny et al. 2008, Villafana et al. 2019, Xu & Nicholson 2009, Zhang et al. 2020).

붉은곰팡이병 병원균의 감염 저항성유형

밀에서 FHB은 다양한 양상으로 발현되므로 감염과 저항성에 대한 정확한 이해가 매우 중요하다. 밀의 FHB 병원균 감염에 대한 저항성 유형은 5가지로 나뉘는데, 초기 감염(initial infection)에만 관련된 저항성(Type I), 감염된 FHB 병원균의 이삭내 확산(spread)에 대한 저항성(Type Ⅱ), DON과 같은 곰팡이 독소 축적(accumulation)에 대한 저항성(Type Ⅲ), 곰팡이의 종자 감염(Fusarium-damaged kernels, FDK)에 대한 저항성(Type Ⅳ)과 수량 감소에 대한 저항성(Type Ⅴ)이다(Mesterházy 1995, Miller et al. 1985, Miller & Arnison 1986, Schroeder & Christensen 1963) (Fig. 2).

Type I 저항성은 FHB 병원균의 포자 현탁액(spore suspension)을 개화기 이삭에 분사해서 발병된 이삭의 비율을 측정한 FHB 발병율(FHB incidence)로 저항성 정도를 평가하고, Type Ⅱ 저항성은 개화기 이삭의 중앙에 있는 소수(spikelet)에 병원균을 접종(inoculation)한 후 3주후 이삭 내 감염이 확산된 종실 비율을 측정하여 FHB 심도(FHB severity)로 저항성 정도를 평가한다(Bai et al. 2003, Bai & Shaner 2004, Mesterházy 1995). Type Ⅰ과 Ⅱ 저항성이 가장 많이 사용되는 방법이다. Type Ⅰ은 Type Ⅱ에 비해 노력이 적게 들지만, 전장, 망 유무, 수축 길이와 개화기 등 형태적 특성에 관련된 농업형질이 Type Ⅰ의 저항성에 영향을 미치기 때문에 같은 품종이라도 환경에 따라 저항성 정도가 차이가 난다. 그래서 Type Ⅱ의 저항성 평가를 포장이나 실내 검정으로 가장 많이 이용한다(Bai et al. 2003, Bai & Shaner 2004, Bushnell et al. 2003, Mesterházy 1995). 대부분 연구는 분사나 접종을 통해서 FHB 발병률과 심도를 동시에 평가하는데, 이를 통해서 Type I 저항성과 Type I 과 Ⅱ 저항성을 동시에 평가한다(Buerstmayr et al. 2020). 최근에는 FHB 발병률과 심도를 FHB 등급지수(FHB visual rating index, VRI; VRI=(FHB incidence×FHB severity)/100)로 전환하여 평가하기도 한다(McCartney et al. 2016). DON 함량 및 전사체 분석과 Sumai 3와 감수성 품종의 병징 특성 평가를 통해서 Type Ⅱ 저항성은 수축 절간(rachis internode) 길이가 중요한 것이 밝혀졌다(Hofstad et al. 2016, Wang et al. 2018).

최근 들어 식품의 안전성 때문에 Type Ⅲ에 대한 저항성에 대한 관심도 높아지고 있는데, Type Ⅲ 저항성 품종이 감수성 품종에 비해 DON 축적량이 상대적으로 적다(Miller et al. 1985). Type Ⅳ는 종자 내 병원균 침입 정도나 배유 전분의 구조 변형과 손상정도가 적을수록 저항성을 나타내며, Type Ⅴ는 FHB 병원균에 감염되어도 수량 감소가 적은 것이 저항성이기 때문에 저항성 보다는 내성(tolerance)에 가까운 유형으로 볼 수 있으며, 최근에는 Type Ⅳ와 Ⅴ의 발생 원인과 결과가 유사하기 때문에 단일 유형으로 보기도 한다(Gong et al. 2020, Jackowiak et al. 2002). Type Ⅲ, Ⅳ와 Ⅴ 저항성과 관련된 독소 축적과 수량 감소는 Type I 과 Ⅱ 저항성을 지닌 경우 쉽게 극복이 가능하기 때문에 밀 FHB 저항성 연구는 주로 Type I 과 Ⅱ에 많은 연구가 이루어 졌다. 하지만 최근에는 미국 남서부 지역의 연질밀 재배 지역은 Type Ⅲ과 Ⅳ 저항성 증진에 많은 노력을 기울이고 있다(Wu et al. 2022). 근동질계통(near isogenic lines, NIL)을 이용한 양적유전자좌(quantitative trait loci, QTL) 분석한 결과, Type Ⅰ 저항성 QTL의 일부는 Type Ⅱ 저항성을 나타내지 않았고, Type Ⅱ 저항성 QTL의 일부는 Type I 저항성을 나타내지 않았다(Xue et al. 2010a). 그리고 Type Ⅱ, Ⅲ과 Ⅳ 저항성 QTL은 중복되지 않는 것으로 나타났으며, Type Ⅳ 저항성 QTL과 수량구성요소 관련 QTL은 중복되는 걸로 나타났다(Abate et al. 2008, Bonin & Kolb 2009, Draeger et al. 2007, Jiang et al. 2007b, Li et al. 2008, Szabó-Hevér et al. 2012, Szabó-Hevér et al. 2014, Wiśniewska et al. 2016, Yang et al. 2005).

붉은곰팡이병 병원균에 대한 방어기작

FHB 병원균에 대한 저항성은 능동적 방어와 수동적 방어로 나뉘고, 수동적 방어는 식물체의 형태나 생리적인 특성에 의해 저항성을 가지게 되는데, 수축 길이와 관련 있는 수밀도(kernel density), 출수기(heading stage), 개화기간 및 전장(height) 등이 해당된다(Kusch & Panstruga 2017, Kushalappa et al. 2016,). 이삭의 미세환경이 FHB 병원균에 저항성을 지니게도 하는데, 소수의 밀납(wax)이 있으면 FHB 병원균의 침투가 어렵지만 약 추출(anther extrusion)이 클 경우에는 침투가 용이하게 된다(Buerstmayr et al. 2020). 균사가 식물세포 내부층을 침투하는 감염 초기 단계(36 hpi)를 Type Ⅰ 저항성, 36 hpi 이후를 Type Ⅱ 저항성으로 구분한다(Fig. 3) (Kang & Buchenauer 2000, Wu et al. 2022). Type Ⅰ 저항성 단계에서 glycolysis pathway (EMP), tricarboxylic acid cycle (TCA), reactive oxygen species (ROS)생산 및 ascorbate peroxidase (APX), polyphenol oxidase (PPO)와 곰팡이 flavin oxidoreductase (FR)의 활성이 유도되며, 식물호르몬(salicylic acid, SA; jasmonic acid, JA; ethylene, ET; abscisic acid, ABA) 관련 경로가 조절된다(Buhrow et al. 2021, Dong 1998, Fonseca et al. 2009, Li & Yen 2008, Spanic et al. 2017, Wang et al. 2002, Wang et al. 2005, Zhou et al. 2006, Zhou et al. 2009). 병원체가 기주식물에 침투하기 위해 기주식물의 생리 대사와 PAMP-triggered Immunity (PTI)를 극복하고 효과인자를 분비하면 기주식물은 세포내 수용기나 식물저항성 단백질들(R proteins)이 발현을 촉진시켜 effector-triggered-immunity (ETI)가 활성화된다(Selin et al. 2016). 병원체는 ETI를 무력화시키기 위해 변형 또는 돌연변이를 통해서 새로운 효과인자를 생산해서 PTI를 억제하고 R proteins의 공격을 피하게 되는데, FHB 병원균이 침입하면 기주 식물 세포를 괴사하게 하여 곰팡이독소(mycotoxins)를 만들고, 식물체는 RR 저항성 관련 단백질(resistance-related proteins) 및 대사 산물의 생산으로 병원체를 막아낸다(Bönnighausen et al. 2019, Gunupuru et al. 2017, Karre et al. 2019). FHB 병원균에 의해 유도되는 유전자는 163개이고 단백질은 37개로 알려져 있다(Ding et al. 2011). FHB 병원균에 감염이 되면 6시간 이내에 SA와 포스파티드산(phosphatidic acid)의 신호전달과 활성산소(reactive oxygen species, ROS) 생성 및 제거되는 과정을 통해 식물체의 과민반응(hyper-sensitive responses)과 식물세포 사멸(plant cell death)를 통한 병원체 성장의 억제가 일어나기 때문에 이들 신호전달 물질이 FHB 병원균 감염의 초기에 있어서 매우 중요하다. 이후 24시간이내에 JA와 ET 신호전달 활성화를 통해 FHB 병원균의 괴사가 이루어진다(Ding et al. 2011, Ma et al. 2020). 감염초기단계(36 hpi) 이후에 Type Ⅱ 저항성이 유도되는데 hemicellulose를 구성하는 조성물, 세포벽 단백질 hydroxyproline-rich glycoproteins (HRGPs)이 증가하고 phenylpropanoid pathway가 촉진된다(Deepak et al. 2010, Dong & Lin 2021, Lionetti et al. 2015). Type Ⅲ 저항성에는 mycotoxin 해독을 위해 UDP-glucuronosyltransferases (UGTs), ATP-binding cassette transporters (ABC transporters), glutathione S-transferases (GSTs), cytochromes P450s (CYP450s) 발현이 유도되어진다(Bernardo et al. 2007, Gunnaiah et al. 2012, Jia et al. 2009, Kong et al. 2005, Nilsen et al. 2020). FHB 병원균 감염에 신호 전달 물질의 역할이 밝혀지고 있고, 이와 더불어 감염 식물체의 독소 제거 능력 또한 FHB 저항성에 있어서 중요하기 때문에 최근 들어서 관련 연구가 수행되고 있다(Gatti et al. 2019, Lemmens et al. 2005, Wang et al. 2020).

붉은곰팡이병에 대한 저항성 유전자원

중국과 일본을 포함한 아시아와 캐나다, 미국, 멕시코를 포함한 라틴아메리카에서는 봄밀 저항성 자원이 보고되었고, 북미와 유럽 자원에서는 겨울밀 저항성 자원이 보고되었다(Bai & Shaner 2004, Buerstmayr et al. 2009, Dill-Macky & Jones 2000). Sumai 3는 가장 많이 알려진 중국 봄밀 중 Type Ⅱ에 저항성인 자원으로, 여러 다양한 환경에서도 저항성이 유지되어 각국에서 저항성 증진을 위한 교배본으로 많이 활용되고 있다. 이외에도, Baishanyuehuang, Haiyanzhong, Huangcandou, Huangfangzhu, Ning 7840, Ning 8331, Qing Shoumai와 Wangshuibai가 저항성 자원으로 알려져 있다(Anderson et al. 2001, Bai & Shanger 1996, Bai & Shaner 2004, Buechley & Shaner 1999, Buerstmayr et al. 2009, Jia et al. 2005, Lin et al. 2006, Rudd et al. 2001, Somers et al. 2003, Waldron et al. 1999, Yu et al. 2008, Zhang et al. 2004, Zhu et al. 2019). 중국은 일본계통의 Norin 129를 자국의 FHB 저항성 증진을 위한 자원으로 이용하였다(Zhu et al. 2019). 일본 자원으로는 Nobeokabozu-komugi, Nyubai와 Shinchunaga가 있으며(Gocho 1985, Nakagawa 1955) 국내 품종 중에는 조광이 저항성으로 알려져 있다(Yang et al. 2006). 유럽의 저항성 자원은 스위스 품종인 Arina (Chrpova et al. 2012, Mesterházy et al. 1999, Paillard et al. 2004, Saur 1991), 프랑스 품종인 Renan (Gervais et al. 2003)과 Type Ⅱ 저항성 선발에 이용되는 마커인 Fhb1를 지닌 Jaceo (Steiner et al. 2017), 체코 품종 Praag 8 (Snijders 1990), 루마니아 품종인 Fundulea 201R (Shen et al. 2003a)와 오스트리아 계통인 CM82036이 있다(Buerstmayr et al. 2003). 브라질의 저항성 자원으로는 Encruzilhada, Frontana, Polyssu와 Toropi가 있으며(Kohli & de Ackermann 2013, Steiner et al. 2004), 미국의 저항성 자원으로는 AGS 3015, Becker, Ernie, Freedom, Jamestown, KY06C-11-3-10, Massey, MO 94-317, NC-Neuse, Roane, Truman, VA00W-38, VA04W-433과 VA04W-474가 있다(Chen et al. 2012, Clark et al. 2014, Griffey et. 2010, Jin et al. 2013, Murphy et al. 2004). 2007년에 버지니아주에서 육성된 Jamestown은 1BS에 FHB 저항성 관련 인자가 위치한 것으로 알려져 있어서 육종프로그램에서 많이 이용되고 있다(Griffey et al. 2010, Zhu et al. 2019). 국제 밀-옥수수연구소(International Maize and Wheat Improvement Centre, CIMMYT)에서는 Catbird 및 파생 계통과 합성품종(synthetic bread wheat)을 육성하였다(Gilchrist et al. 1997, Singh & Van Ginkel 1997). 아르헨티나자원인 MS INTA 416는 잎 녹병(leaf rust) 저항성 Lr47 유전자와 Fhb1을 지녔고(Bainotti et al. 2017), Barletta, Klein Sin Rival과 Klein Vencedor도 저항성 품종으로 알려져 있으나 이들 자원은 Sumai 3와 저항성에 대한 기원이 전혀 다른 것으로 알려져 있다(Bai et al. 1999, Bainotti et al. 2017, Buerstmayr et al. 2003, Gilchrist et al. 1997, He et al. 2013a, He et al. 2013b, Kohli & de Ackermann 2013, Rudd et al. 2001). Sumai 3를 포함한 아시아 저항성 자원들은 대부분 농업형질이 좋지 않아서 우수한 농업형질을 지닌 중도저항성 자원들이 육종에 이용되는데, 대표적인 자원들은 미국의 Alsen, Bacup, Everest, Glenn, ND2710, Overland, Sabin과 Steele 자원들이 있으며, 이들은 FHB 저항성 관련 유전 분석을 위해서도 많이 이용되고 있다(Mergoum et al. 2007, Waldron et al. 1999). 중국은 FHB 저항성 증진을 위해 Norin 129를 Fhb1의 주요 저항성 제공자(major donor)로 이용해서 Jimai 22, Zhoumai 16 및 여러 우량 계통에 도입하였다. Wangshuibai는 Type Ⅰ 저항성 증진을 위한 자원으로 이용하고 있으며, Type Ⅱ 저항성 증진을 위해서는 Baishanyuehuang, Haiyanzhong, Huangfangzhu, Huangcandou과 PI672538를 활용하고 있다. 이들 자원이외에도 Fanshanxiaomai, Frontana, Hongheshang, Ning 7840, Nobeokabozu-komugi, Saikai-165, Shinchunaga Sumai 3와 Wenzhou도 이용되고 있다(Bai et al. 1989, Ban & Suenaga 2000, Cai et al. 2016, Cai & Bai 2014, Li et al. 2011, Li et al. 2012, Li et al. 2017, Lin et al. 1992, Lin et al. 2006, Lu et al. 2013, Singh et al. 1995, Van Ginkel et al. 1996, Wang et al. 1992, Zhang et al. 2012, Zhu et al. 2019). CIMMYT는 Fhb1과 매우 근접해 있는 줄기 녹병(stem rust) 유전자중에 하나인 Sr2를 저항성 자원 선발에 이용을 하고 있으며, Fhb1-Sr2재조합 계통인 Hartog/Sumai 3로부터 Fhb1 저항성인자를 Shuttle breeding과 marker-associated selection (MAS) 시스템을 활용하여 Quaiu, Munal와 Super 152와 같은 우량계통에 도입하는데 성공하였다(Hao et al. 2020).

붉은곰팡이병 병원균에 대한 저항성 유전자

밀의 FHB 저항성은 양적유전의 양상을 보이는 것으로 나타났으며, Type Ⅰ-Ⅳ의 FHB 유전력은 각각 0.70, 0.73, 0.54과 0.68로 낮지 않지만, 저항성 평가와 선발에 있어서 온도나 습도 등 환경에 영향을 많이 받는 걸로 알려져 있다(Draeger et al. 2007, Gu 1983, Liu et al. 1992, Petersen et al. 2017, Singh et al. 1995, Steiner et al. 2019, Tessmann et al. 2019, Zhang & Pan 1982). 현재까지 400개 이상의 QTL이 보고되었고, 다양한 유전적 배경이나 환경조건에서 저항성이 검증된 Fhb 유전자는 7개(Fhb1-Fhb7)이다. 유전자 수준에서 동정(cloning)된 것은 Fhb1과 Fhb7이고, 현재까지 가장 많이 이용되고 있는 것은 Fhb1와 Fhb2이다(Table 1) (Bai & Shaner 2004, Ma et al. 2020). Fhb1은 Sumai 3에서 유래한 Type Ⅱ 저항성 유전자로 현재 육종 사업에서 가장 광범위하게 이용되고 있는데, 염색체 3BS 말단의 Qfhs.ndsu-3BS로 보고되었던 QTL을 Sumai 3와 Nyu Bai를 이용하여 육성된 여교배 및 F₂ 집단을 이용한 정밀 유전자 지도 작성(fine-mapping)으로 밝혀졌다(Bai et al. 2018, Bernardo et al. 2012, Gunnaiah et al. 2012, Jia et al. 2018, Liu et al. 2006). Fhb1은 유전자 기능검정을 통해서 PFT (pore forming toxin-like)로 밝혀졌지만, FHB 감수성 품종도 PFT 유전자를 지닌 것으로 밝혀졌고, 후속 연구를 통해서 PFT로부터 약 50 kb 떨어진 위치에 TaHRC (histidine-rich calcium-binding protein, His)가 Fhb1 해당 유전자로 동정 되었다(Jia et al. 2018, Li et al. 2019, Rawat et al. 2016, Su et al. 2019). TaHRC의 FHB 저항성 기작에는 기능획득(gain of function)과 기능상실(loss of function)의 2가지 가설이 제기되고 있다(Li et al. 2019, Su et al. 2019). 기능획득 가설은 저항성인 His^R은 감수성인 His^S에 비해 14개 아미노산이 많은 데 이러한 이유는 상류영역(upstream)에 위치한 개시코돈에서 TaHRC 유전자의 기능획득으로 일어난 현상이라는 것이다. 기능상실 가설은 TaHRC를 과발현 해도 FHB 저항성이 증가하지 않고, 오히려 TaHRC를 knockout 또는 Knockdown 시켰을 때 저항성이 향상된 것으로 보아 FHB 저항성은 TaHRC 유전자의 두번째 인트론(intron) 내 결실(deletion)이 하류영역(downstream)에 위치한 개시코돈을 제거한 기능상실로 발생한다는 것이다. 아직까지 명확히 밝혀지지는 않았지만 TaHRC나 His가 밀의 FHB 저항성에 매우 중요한 역할을 하며 이러한 특성은 다른 작물에도 적용이 된다(Bai et al. 2018, Hao et al. 2020, Li et al. 2019).

현재 Fhb1을 통한 FHB 저항성은 아직 밝혀지지 않은 복합적인 유전자 상호작용이나 PFT와 TaHCR를 포함한 다수의 유전자로 구성된 오페론(operon) 형태의 유전자복합체(gene cluster)에 의하여 조절될 가능성이 있다고 보고 있다(Hao et al. 2020, Lagudah & Krattinger 2019). 현재까지 Sumai 3에서 유래된 Fhb1은 다양한 환경과 자원에서 안정적으로 Type Ⅱ에 저항성을 지니는 것으로 알려져 있다(Anderson et al. 2001, Bai et al. 1999, Buerstmayr et al. 2003, Chen et al. 2006, Cuthbert et al. 2006, Jayatilake et al. 2011, Jiang et al. 2007a, Jiang et al. 2007b, Lemmens et al. 2005, Shen et al. 2003b, Somers et al. 2003, Yu et al. 2008). 미국 북서부 지역 밀 자원은 Fhb1 유전자를 지닌 자원이 적어서 FHB 저항성 선발에서 의미가 없었지만, 중서부와 동부 밀에서는 Fhb1 유전자가 FHB 저항성에서 중요한 역할을 하고 있으며, South Dakota주립대의 봄밀 육종 프로그램은 Fhb1을 활용해서 FHB 저항성 증진시키고 있다(Arruda et al. 2016, Eckard et al. 2015a, Wang et al. 2017). Fhb2는 Sumai 3파생 계통의 재조합 근교계집단(recombinant inbred lines, RILs)을 이용한 유전분석으로 6BS 염색체의 Xgwm133과 Xgwm644 사이에 위치함이 확인되었고, Wangshuibai에서도 저항성 QTL인 Qfhs.nau-6B로 탐지되어서 Fhb1과 함께 Type Ⅱ 저항성 선발 마커로 이용되고 있다(Anderson et al. 2001, Buerstmayr et al. 2002, Cuthbert et al. 2007, Jia et al. 2018, Waldron et al. 1999, Xue et al. 2010b, Zhou et al. 2002). Fhb2 유전자좌 근처에는 FHB 저항성 관련되어 DON 독소 제거, 세포벽 강화 및 화기(floret)나 수축에서 병원균 확산 억제에 관련된 6개 정도의 유전자가 있을 것으로 추정하고 있다(Dhokane et al. 2016). Fhb3는 4배체인 Leymus racemosus의 7Lr#1염색체가 밀의 7A 염색체에 전위된 계통에서 유래되었고 Sumai 3와 유사한 수준의 Type Ⅱ 저항성을 지니고 있는 것으로 밝혀졌다(Qi et al. 2008). Fhb4는 Wangshuibai와 Nanda2419의 RIL과 BC₃F₂ 유전 분석을 통해서 4BL 염색체의 Xhbg226와 Xgwm149 사이에 존재하는 마커로 Type Ⅰ 저항성 자원 선발에 이용되고 있다(Jia et al. 2018, Xue et al. 2010b). Fhb5 (Qfhs.ifa-5A)는 5A 염색체에 존재하며 Type Ⅰ 저항성 선발 마커로 이용된다. Qfhs.ifa-5A는 두개의 독립된 QTL인 Qfhs.ifa-5Ac와 Qfhs.ifa-5AS로 분리되며 각각 Xgwm304‒Xgwm415과 Xwgrb0222-Xwgrb1621사이에서 매핑 되었다. 그리고 Fhb1 보다는 적지만 Type Ⅱ 저항성 선발에도 이용이 가능하다(Buerstmayr et al. 2018, Jia et al. 2018, Lin et al. 2006, Xue et al. 2011). Fhb3와 유사하게 Fhb6는 Elymus tsukushiensis의 1Ets#1S 영역이 밀의 1AS에 삽입된 계통의 QTL Mapping에서 이들 계통이 Type Ⅱ에 저항성을 나타내어 Fhb6으로 명명되었다(Cainong et al. 2015). Fhb7은 밀의 근교계인 Thinopyrum elongatum과의 교배를 통하여 도입된 저항성 유전자로 글루타치온(glutathione)합성을 촉매해서 DON 독성을 해독하는 GST를 암호화하며 Type Ⅱ와 Type Ⅲ 저항성을 지닌 것으로 알려져 있다(Fu et al. 2012, Wang et al. 2020, Zheng et al. 2020). 중국 품종인 Nanda 2419에서 보고된 QTL인 Qfhb. nau-2B은 Type Ⅰ과 Ⅱ에 저항성으로 알려졌다(Li et al. 2019, Lin et al. 2006). 그리고 Glenn, Wanshuibai, Frontana와 L699 품종은 Fhb1 이외에 Fhb2, Fhb4와 Fhb5 유전자의 존재로 인해 Type Ⅰ과 Type Ⅱ에 저항성을 나타낸다(Bokore et al. 2017, Cai et al. 2016, Lin et al. 2006, Mergoum et al. 2006, Steiner et al. 2004, Yang et al. 2016, Zhang et al. 2011). FHB 저항성은 세포질 효과(cytoplasmic effect)는 크게 나타나지 않는 것으로 알려져 있지만, Ning 7840과 CNO79조합 및 Funo과 Taiwanxiaomai 조합 같은 특정 조합에서는 모계효과(maternal effects)가 나타나는 것으로 알려져 있다(Van Ginkel et al. 1996, Zhang & Pan 1982).

붉은곰팡이병 병원균 저항성 QTL Mapping

1999년에 RILs (Sumai 3/Stoa, Ning 7840/Clark)을 이용한 Type Ⅱ 저항성 관련 QTL이 보고된 이후 다양한 유전 집단에서 많은 QTL의 보고가 이루어지고 있으며, 이를 기반으로 한 관련된 마커와 MAS 활용이 이루어졌다. 최근에는 단일염기형성(single nucleotide polymorphism, SNP) array와 전장유전체 연관분석(genome-wide association study, GWAS)으로 고밀도 유전지도(high resolution genetic map) 작성을 통한 선발 효율을 높이는 노력이 이루어지고 있다(Anderson 2012, Bai et al. 1999, Buerstmayr et al. 2020, Del Blanco et al. 2003, Ghimire et al. 2020, Hao et al. 2020, Khan et al. 2020, Mesterházy 2020, Rawat et al. 2016, Rutkoski et al. 2012, Waldron et al. 1999, Yang et al. 2003, Zhou et al. 2003) (Fig. 2). FHB 저항성 QTL mapping의 정확성에는 다른 형질과 마찬가지로 집단의 크기, 마커 밀도(marker density), 표현형 평가와 유전형의 정확성이 중요하며, minor QTL 이기는 하지만 환경적인 영향이 작용하기 때문에 일반적으로 다양한 환경에서 다년간 실험이 행해진다(Collard et al. 2005, Haley & Knott 1992, Kolb et al. 2001). 2020년까지 FHB 저항성 관련 QTL은 432개가 보고되었고, 이중에서 40개 정도는 검증이 되었다(Table 2) (Ma et al. 2020). 메타 분석을 통해서 최소 10개의 마커 연계된 특성(marker-trait associations)관련 65개의 meta-QTL이 보고되었다(Ma et al. 2020, Venske et al. 2019). Arina, CJ 9306, DH181, Frontana, NC-Neuse, Sumai 3, Truman와 Wangshuibai 등의 품종에서는 3개이상의 염색체 부분에서 QTL이 보고되었으며, 우리나라 재래품종인 조광에서도 3BS, 4BL과 5DL에서 QTL이 보고되었는데, 이러한 결과는 FHB 병원균 저항성에 많은 유전자가 상당히 복잡하게 연결되어 있음을 의미한다(Table 3) (Bai et al. 2018, Bonin & Kolb 2009, Brown-Guedira et al. 2008, Eckard et al. 2015b, Gilsinger et al. 2005, Islam et al. 2016, Kang et al. 2011, Van Sanford et al. 1997, Zhu et al. 2019).

FHB 저항성 관련 QTL은 다른 농업형질과 매우 밀접하게 관련이 있는데, 2009년에서 2019년까지 연구를 정리한 결과, 전장, 약 추출과 개화기 관련 QTL과 FHB 병원균 저항성 관련 QTL의 중복 정도는 각각 44%, 60%와 25%로 나타났다(Buerstmayr et al. 2020). 1960년대 녹색혁명으로 수량 증대가 이루어 졌지만 Rht-B1b와 Rht-D1b 유전자 도입으로 인한 중단간화(Semi-dwarf) 밀육종으로 전장은 41% 감소하였으며(Flintham et al. 1997, Jobson et al. 2019), FHB의 Type Ⅰ 감수성은 41%까지 증가했고, 약 추출은 13-23% 감소한 걸로 알려져 있다(He et al. 2016, Miedaner & Voss 2008). 토양에 있는 곰팡이 병원균이 60 cm 이상에서는 전파 가능성이 적기 때문에 키가 클수록 병원균 접촉 가능성이 적어지고, 까락이 없을수록 병원균 접촉면이 적기 때문에 무망(awnless) 품종이 유리하며, 화기 구조 측면에서는 개화시 영이 열리는 정도가 적을수록 유리한 것으로 알려져 있다(Buerstmayr et al. 2009, Buerstmayr & Buerstmayr 2016, Gilsinger et al. 2005, Mesterhazy 1995, Steiner et al. 2017). 다양한 높이의 밀 이삭은 FHB 심도에 영향을 미치는 다양한 온도와 습도 같은 다양한 미기상(microclimate)에 노출되므로 전장은 FHB 저항성과 관련이 깊다. 수밀도(spike compactness)도 병원균 접종(point inoculation)과 상관이 있으며, 전장과 수밀도는 병원균 분사(spray inoculation)에 따른 FHB 심도와 상관이 있다(Buerstmayr & Buerstmayr 2016, Yi et al. 2018). 장간 품종이 Type Ⅱ에서는 단간(dwarf) 품종 보다 저항성을 나타내지는 않았지만, NIL을 통한 연구에서는 Rht-B1b 지닌 단간 계통이 Type Ⅰ, Ⅱ 또는 III에서 저항성을 보인 것으로 나타났다(Buerstmayr et al. 2013, He et al. 2016, Kollers et al. 2013, Miedaner et al. 2011, Srinivasachary et al. 2009, Yan et al. 2011). Rht-1 유전자는 세포 신장(cell elongation)을 저해하기 때문에 수밀도를 높이고, 수술 길이와 약 추출을 감소시켜 FHB 감수성을 높이는데, Rht-D1b 유전자의 영향력이 Rht-B1b에 비해 큰 것으로 알려져 있다(Boeven et al. 2016, Buerstmayr & Buerstmayr 2016, Cheng et al. 2004, Miralles et al. 1998, Miedaner & Voss 2008, Okada et al. 2019, Srinivasachary et al. 2009).

약의 특성은 유전력은 높지만 많은 유전자가 관련되어 있는 것으로 알려져 있고, 현재까지 FHB의 QTL과 관계된 연구는 9개 유전 집단에서 7개 연구만이 이루어졌는데, 15개 염색체의 23개 영역에서 약 보유(anther retention)와 관련 있는 QTL이 보고되었다(Buerstmayr et al. 2020, Buerstmayr & Buerstmayr 2015, He et al. 2016, Lu et al. 2013, Steiner et al. 2019). 약 추출에 이삭 형태 및 수술 관련 특성에 관련된 다양한 유전자가 영향을 미치기 때문에 Rht1유전자가 약 추출을 억제하기도 하고 그렇지 않기도 하다. 약 추출에 Rht-B1은 영향을 주지만 Rht-D1은 상관이 없다는 상반된 보고가 있지만, 모든 결과에서 약 보유 비율이 높으면 FHB 심도는 커지는 것으로 나타났다(Boeven et al. 2016, Buerstmayr & Buerstmayr 2016, He et al. 2016, Lu et al. 2013, Muqaddasi et al. 2017, Okada et al. 2019, Wuerschum et al. 2018). 그러므로 약 추출이 높은 자원을 선발하는 것이 Rht1 유전자에 의해서 단간화로 인해 발생하는 FHB 피해를 줄이는 것으로 제안되고 있다. 약 추출과 상관 있는 QTL중에 Qfhs.ifa-5A과 Rht1만이 전장을 줄이고 약 보유 비율을 증가시키는데, Qfhs.ifa-5A에 의해서 약 추출이 증가하여 FHB 저항성이 증진되는 효과는 Rht-D1b에 의해서 전장 감소로 일어나는 FHB 감수성 증가 정도와 유사한 걸로 알려져 있다(Miedaner et al. 2019, Steiner et al. 2019).

개화와 관련된 특성이 FHB와 상관이 높기 때문에 관련된 유전자인 광주기성 및 파성에 관련된 유전자인 Ppd-D1a (photoperiod insensitivity gene)와 Vrn-A1 및 Vrn-B1 (vernalization requirement genes)이 FHB QTL과 관련이 있으며, Vrn-1이 Ppd-1보다 영향이 큰 것으로 알려 있다. 하지만 유럽밀에서는 개화기와 FHB 심도는 상관이 없었으며, 출수기와 전장으로 정확한 보정을 했을 때에도 Vrn-A1이 FHB 심도에 미치는 영향은 없었다고 보고하였다(Buerstmayr et al. 2008). 순화(domestication)와 관련이 있는 유전자인 Q (major wheat domestication gene)는 탈립성 뿐만 아니라 여러가지 이삭의 형태 및 특성에 영향을 미치기 때문에 FHB와 상관이 있는 것으로 알려져 있다(Buerstmayr et al. 2020, Faris & Gill 2002, He et al. 2016, Islam et al. 2016, Liu et al. 2013, Miedaner & Voss 2008, Petersen et al. 2017, Simons et al. 2006, Yi et al. 2018). 전장, 출수기 및 화기 구조가 FHB 저항성에 영향을 미치는 것은 일종의 다면 발현현상(pleiotropic effects)으로도 볼 수 있기 때문에 FHB 저항성 품종 선발 효율을 높이기 위해서는 육종가가 여러가지 농업형질을 동시에 고려해야만 한다.

GWAS를 통한 밀의 FHB 저항성 관련 연구로 보다 정밀하게 잘 정리된 QTL 뿐만 아니라 FHB 저항성에 관련된 많은 유전자가 밝혀지고 있으며, 선발 효율성을 높이기 위한 Genomic selection (GS)에 대한 관심도 높아지고 있다. 455개 유럽 적립계 연질 추파밀의 GWAS 분석에서는 1D와 3A 염색체에서 9개의 QTL이 보고되었고, 최근 육성된 358개 유럽 밀 품종에 대한 GWAS 분석에서는 6B 염색체를 제외하고는 기존에 알려진 QTL과 일치하였다(Kollers et al. 2013, Miedaner et al. 2011). 교배본이 다양한 F₁ 800계통에 대한 분석을 통해 FHB 저항성에 관련된 15개 QTL이 염색체 2A, 2B, 3B와 7B에 위치하고, 기존에 알려진 Fhb1, Fhb5, Rht-D1와 Rht-B1 포함하고 있음이 밝혀졌다(Eckard et al. 2015a, Eckard et al. 2015b). 273개 미국 중부 추파 밀 계통에서 GWAS 분석 결과, FHB 병원균저항성과 관련된 QTL은 주로 1D, 3B, 4A, 4D, 6A, 7A와 7D에 위치하고, 3B에서 발견된 몇몇 SNP는 Fhb1과 관련 있는 것으로 나타났으며, 일본과 중국 밀 자원에 대한 GWAS 분석으로 11개 QTL을 찾았지만 10개는 기존에 알려진 QTL이었으며, 새롭게 밝혀진 QTL은 5D 염색체에 존재하는 것으로 나타났다(Arruda et al. 2016, Li et al. 2016). 두개의 서로 다른 환경에서 재배된 238개 연질 추파밀에서 29개의 SNP가 보고되었으며, 4A, 5B와 6B에 위치한 QTL은 Type Ⅲ과 IV 저항성과 상관있는 것으로 나타났고, 354개 연질밀 계통에 대한 GWAS 분석으로 Type Ⅰ, Ⅱ, Ⅲ과 IV에 관련 있는 41개의 SNP가 보고되었다(Larkin et al. 2020, Tessmann et al. 2019, Tessmann & Van Sanford 2018). 2,500개 유럽밀에 대해서 GWAS와 GS를 통한 선발을 하면 단순한 FHB 저항성 선발 보다 Type Ⅱ 선발 효율이 10.62% 높아졌고, 2,325개 유럽밀에서도 GWAS+GS을 통한 Type Ⅰ의 예측 확률이 0.6으로 매우 높았다. 미국 태평양 북서부 지역의 170개 품종에 대한 Type Ⅲ에 대한 선발 효율이 60%에 달했으며, 273개 미국 중부 추파 밀 계통에서도 GWAS+GS 모델이 FHB 저항성을 예측에 효과적인 것으로 나타났다(Arruda et al. 2016, Dong et al. 2018, Herter et al. 2019, Mirdita et al. 2015a, Mirdita et al. 2015b).

결 론

최근 들어 기후변화로 인한 잦은 이상 기후 발생으로 붉은곰팡이병(FHB)의 빈도와 심각성이 증가되고 있어 주요 밀 생산국들은 FHB 저항성 증진을 위한 유용자원 탐색 및 저항성 품종 육성과 육종 효율성 증진을 위한 표지인자 개발에 많은 노력을 기울이고 있다. 다양한 유전 집단에서 많은 QTL의 보고가 이루어지고 있으며 이를 기반으로 한 관련된 마커와 MAS 활용이 이루어졌는데 현재까지 Fhb (Fhb1-Fhb7) 병저항성 검정 유전자를 가장 많이 활용하고 있다. GWAS와 GS를 통해 FHB 저항성 예측과 선발 효율이 높아졌고, SNP array와 GWAS로 고밀도 유전자지도작성을 통한 선발 율을 높이는 노력이 이루어지고 있다. 국내에도 기후변화로 인한 이상기온으로 FHB 발생이 높아지고 있고, 국산 밀 안정적 공급을 위해서 국내환경에 맞는 저항성 품종을 육성해야 한다. 이를 위해서 정밀 검정체계 구축, 표현형 데이터 획득 및 효율적인 분자 마커 개발, GWAS와 GS 분석으로 QTL 및 유전자 선별 등 MAS 연구가 활발히 수행되어 저항성 품종 개발과 육종에 활용하여야 한다.

적 요

붉은곰팡이병(FHB)은 밀의 수량 감소와 품질 저하를 일으키며, FHB 병원균 감염에 의해 곡물에서 생산되는 독소는 가축은 물론 사람에게도 중독증 피해를 일으킨다. 최근 들어 기후변화로 인한 잦은 이상 기후 발생으로 국내에도 발생이 높아지고 있어 국산 밀 안정적 공급을 위해서 반드시 관리해야 하는 병해이지만 국내에서는 저항성 품종 선발 마커 개발과 품종 육성이 미흡한 실정이다. 본 연구에서는 FHB 저항성에 대한 이해와 보고된 저항성 자원, 저항성 유전자 및 유전자 마커에 대한 최근 연구 동향을 정리하여 국내에 맞는 마커 개발과 저항성 품종 개발에 도움이 되고자 하였다.

사 사

본 연구는 2023년도 농촌진흥청 국립식량과학원 전문연구원 과정 지원사업과 농촌진흥청 연구사업(과제명: 가공 용도별 밀 품종개량 및 육종효율 증진연구(4단계), 과제번호: PJ 016771022023)에 의해 이루어진 것임.

Fig. 1

Fusarium head blight disease cycle.

Fig. 2

Schematic flow diagram showing overall breeding effort for assessing FHB resistance types, QTL detection and validation, and development of FHB resistant wheat cultivars (Source from Ghimire et al. 2020).

Fig. 3

A model illustrating pathways and different types of FHB resistance from Wu et al. (2022). The dotted arrows represent indirect interactions; the vertical upward arrows represent a positive correlation with resistance and the vertical downward arrows represent a negative correlation with resistance. EMP, glycolysis pathway; TCA, tricarboxylic acid cycle; APX, ascorbate peroxidase; PPO, polyphenol oxidase; HRGPs, hydroxyproline-rich glycoproteins; HCAA, Hydroxycinnamic acid amide; UGT, UDP-glucuronosyltransferase; ABC transporter, ATP-binding cassette transporter; GST, glutathione S-transferase; CYP450, cytochromes P450; hpi, hour post-infection; hpi, h post inoculation.

Table 1

Fusarium head blight (FHB) genes with closely linked markers summarized by Ma et al. (2020).

Table 1
	Chr^zGene	Closely linked markersy	Genetic distance (cM)	FHB resistance typex	Donor parents
Fhb1	3BS	PFT TaHRC Xwgrb619	0	Ⅱ	Wangshuibai Chokwang Sumai 3
Fhb2	6BS	Qfhs.nau-6B Xwgrb682-Xmag3017	2.2	Ⅱ	Wangshuibai Sumai 3 Frontana
Fhb3	7AS	7Lr#1S BE586744-STS, BE404728-STS, BE586111-STS	-	Ⅱ
Fhb4	4BL	Qfhi.nau-4B Xmag8990-Xmag8894	0.14	Ⅰ	Wangshuibai Chokwang
Fhb5	5AS	Qfhs.ifa-5Ac Qfhs.ifa-5AS Xwgrb0222-Xwgrb1621	0.09	Ⅰ	Wangshuibai Frontana
Fhb6	1AS	1Ets#1S tplb0017E15 tplb0029J02 AK357509	-	Ⅰ, Ⅱ
Fhb7	7DL	GST XsdauK66-Xcfa2240	1.7	Ⅱ, Ⅲ
Qfhb. nau-2B		Xwgrb1561-Xwgrb1410	0.9	Ⅰ, Ⅱ	Nanda2419

^zChr=chromosome.

^yPFT=pore-forming toxins, HRC=histidine-rich calcium-binding protein, GST=Glutathione S-transferase.

^xType I=FHB incidence, Type II=FHB spread, Type III=DON content.

Table 2

Quantitative trait loci (QTLs) validated and used in marker-assisted selection (MAS) associated with Fusarium head blight resistance in wheat summarized from Ma et al. (2020).

Table 2
Chrz	Markers	FHB resistance typey	Commentx
2B	Xgwm47	II	QTLs
2B	Xwmc474-Xwmc499	I, I/II	MAS
2D	Xgwm539, Xgwm608	II, III, IV	MAS
	Xcfd233	III, IV	QTLs
	Xcfd233, Xgwm539	II, III	QTLs
3A	Xbarc19, Xbarc67	II, III, IV	QTLs
3A	Xgwm720	II, III	MAS
3B	Xgwm533-Xgwm274	I/II	QTLs
	Xgwm533-Xbarc147	I/II	MAS
	Xgwm533-Xgwm493	I/II, II, III	MAS
	Xgwm389-Xgwm493	I/II	MAS
	Xgwm533, Xgwm493	II, III, IV	QTLs
	Xgwm533-Xgwm493	I/II	MAS
	Xgwm389-Xgwm533-Xbarc133	II, III	MAS
	Xbarc133-Xgwm533-Xgwm493	II, IV	QTLs
	Xgwm389-Xgwm533-Xgwm493	I/II	MAS
	Xgwm533, Xcfd79, Xumn10	II, III, IV	MAS
	Xumn10	II, III	QTLs, MAS
	Xgwm533	III, IV	QTLs
	Xumn10, Xgwm533	II, III	QTLs
	TaHRC-GSM	II	MAS
	Xgwm566, Xwmc231, Xwmc625, Xwmc693, Xwmc307, Xwmc418	II, III	MAS
4B	Xwmc238, Xgwm149	II, III	MAS
	Xwmc245, Xgwm608	II, III	MAS
	Xbarc20-Xwmc349	I	MAS
5A	Xgwm291	I/II	MAS
	Xwmc705, Xgwm304, Xgwm154	II, III	MAS
	Xgwm156, Xgwm304	II, III	MAS
	Xbarc180, Xmag3794	I	MAS
	Xgwm293-Xgwm1057	II	QTLs
	Xgwm304, Xbarc186	II, III, IV	MAS
	Xgwm304, Xbarc117, Xwmc705, Xgwm293	II, III	MAS
6A	Xgwm82	II	QTLs
6B	Xgwm644	I/II	MAS
	Xwmc398-Xgwm644	I/II	MAS
	Xgwm133, Xgwm644	II, III	MAS
7B	Xgwm46	II	QTLs

^zChr=chromosome.

^yFHB Type I=FHB incidence, FHB Type II=FHB severity, FHB Type III=DON content, FHB Type IV=Fusarium-damaged kernels, FHB Type I/II=FHB spread.

^xQTLs=QTLs validation in breeding lines.

Table 3

Sources of FHB resistance and the location of the involved quantitative trait loci (QTLs) in wheat summarized from Khan et al. (2020).

Table 3
Chromosome	Source
1A	Bess (1A), CJ 9306 (1AS), Ernie (1A), NC-Neuse (1A), NK93604 (1AL)
1B	Arina (1BL), CM-82036 (1B), INW0411 (1B), Seri82 (1B), Remus (1B), Wangshuibai (1B)
1D	DH181 (1DL), Sincron (1DS)
2A	Ning 7840 (2AS), Freedom (2AS), Remus (2A), Renan (2A), Stoa (2AL), Truman (2ASc), Wangshuibai (2A)
2B	Dream (2BL), Ernie (2B), Frontana (2B), Goldfield (2B), IL94-1653 (2B), INW0304 (2B), Ning 7840 (2BL), Renan (2BS)
2D	Alondra (2DS), CJ 9306 (2DL), CS-SM3-7ADS (2D), DH181 (2DS), Gamenya (2DS), McCormick (2D), Nuy Bai (2D), Truman (2DS), Wangshuibai (2D/2DL), Wuhan 1 (2DL)
3A	Forno (3AL), Frontana (3A), Heyne (3AS), Huapei 57-2 (3AS), INW0411 (3A), ND2603 (3AL), Wangshuibai (3AS),
3B	Arina (3BL), Bess (3B), Chokwang (3BS), CJ 9306 (3BS), CM-82036 (3BS), COKER 9474 (3BS), CS-SM3-7ADS (3BS), DH181 (3BS), Ernie (3B/3BL), Huapei 57-2 (3BS/3BL), INW0411 (3BS), Massey (3BL), ND2603 (3BS), Ning 7840 (3BS), Ning 894037 (3BS), Nuy Bai (3BS), McCormick (3BS), Patton (3B), Sumai 3(3BS), Wangshuibai (3BS), Wesley (3BS), Wu 14 (3BS)
3D	Forno (3DS), Patterson (3D), Truman (3DS), Wangshuibai (3DL),
4A	Arina (4AL), Ernie (4A), Heyne (4AL), NC-Neuse (4A)
4B	Chokwang (4BL), Ernie (4BL), IL94-1653 (4B), Stoa (4BS), Wangshuibai (4B), Wuhan 1 (4BS)
4D	Arina (4DS), CS-SM3-7ADS (4D), DH181 (4DL), Heyne (4DL)
5A	Arina (5AL), CJ 9306 (5AS), CM-82036 (5A), DH181 (5AS), Ernie (5A), Frontana (5A), McCormick (5A), Nuy Bai (5AS), Wangshuibai (5A), Wu 14 (5A), Renan (5AL)
5B	Forno (5BL), Patterson (5BL), Wangshuibai (5B)
5D	Chokwang (5DL), Wangshuibai (5DL)
6A	Dream (6AL), Ernie (6A), NC-Neuse (6A), ND2603 (6AS)
6B	Arina (6BL), DH181 (6BS), Frontana (6B), Sumai 3 (6BS), Patton (6B), Wangshuibai (6B)
6D	Arina (6DL)
7A	Frontana (7AS), NK93604 (7AL), Sumai 3 (7A), Wangshuibai (7A)
7B	CJ 9306 (7BS), DH181(7BL), Dream (7BS), Goldfield (7B)

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Research Advances in Wheat Breeding and Genetics for Fusarium Head Blight Resistance

Korean. J. Breed. Sci.. 2023;55(3):195-217. Published online September 1, 2023

DOI: https://doi.org/10.9787/KJBS.2023.55.3.195

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Research Advances in Wheat Breeding and Genetics for Fusarium Head Blight Resistance

Korean. J. Breed. Sci.. 2023;55(3):195-217. Published online September 1, 2023

DOI: https://doi.org/10.9787/KJBS.2023.55.3.195

Figure

Research Advances in Wheat Breeding and Genetics for Fusarium Head Blight Resistance

Fig. 1 Fusarium head blight disease cycle.

Fig. 2 Schematic flow diagram showing overall breeding effort for assessing FHB resistance types, QTL detection and validation, and development of FHB resistant wheat cultivars (Source from Ghimire et al. 2020).

Fig. 3 A model illustrating pathways and different types of FHB resistance from Wu et al. (2022). The dotted arrows represent indirect interactions; the vertical upward arrows represent a positive correlation with resistance and the vertical downward arrows represent a negative correlation with resistance. EMP, glycolysis pathway; TCA, tricarboxylic acid cycle; APX, ascorbate peroxidase; PPO, polyphenol oxidase; HRGPs, hydroxyproline-rich glycoproteins; HCAA, Hydroxycinnamic acid amide; UGT, UDP-glucuronosyltransferase; ABC transporter, ATP-binding cassette transporter; GST, glutathione S-transferase; CYP450, cytochromes P450; hpi, hour post-infection; hpi, h post inoculation.

Fig. 1

Fig. 2

Fig. 3

Research Advances in Wheat Breeding and Genetics for Fusarium Head Blight Resistance

Table 1

Fusarium head blight (FHB) genes with closely linked markers summarized by Ma et al. (2020).

	Chr^zGene	Closely linked markersy	Genetic distance (cM)	FHB resistance typex	Donor parents
Fhb1	3BS	PFT TaHRC Xwgrb619	0	Ⅱ	Wangshuibai Chokwang Sumai 3
Fhb2	6BS	Qfhs.nau-6B Xwgrb682-Xmag3017	2.2	Ⅱ	Wangshuibai Sumai 3 Frontana
Fhb3	7AS	7Lr#1S BE586744-STS, BE404728-STS, BE586111-STS	-	Ⅱ
Fhb4	4BL	Qfhi.nau-4B Xmag8990-Xmag8894	0.14	Ⅰ	Wangshuibai Chokwang
Fhb5	5AS	Qfhs.ifa-5Ac Qfhs.ifa-5AS Xwgrb0222-Xwgrb1621	0.09	Ⅰ	Wangshuibai Frontana
Fhb6	1AS	1Ets#1S tplb0017E15 tplb0029J02 AK357509	-	Ⅰ, Ⅱ
Fhb7	7DL	GST XsdauK66-Xcfa2240	1.7	Ⅱ, Ⅲ
Qfhb. nau-2B		Xwgrb1561-Xwgrb1410	0.9	Ⅰ, Ⅱ	Nanda2419

Quantitative trait loci (QTLs) validated and used in marker-assisted selection (MAS) associated with Fusarium head blight resistance in wheat summarized from Ma et al. (2020).

Chrz	Markers	FHB resistance typey	Commentx
2B	Xgwm47	II	QTLs
2B	Xwmc474-Xwmc499	I, I/II	MAS
2D	Xgwm539, Xgwm608	II, III, IV	MAS
	Xcfd233	III, IV	QTLs
	Xcfd233, Xgwm539	II, III	QTLs
3A	Xbarc19, Xbarc67	II, III, IV	QTLs
3A	Xgwm720	II, III	MAS
3B	Xgwm533-Xgwm274	I/II	QTLs
	Xgwm533-Xbarc147	I/II	MAS
	Xgwm533-Xgwm493	I/II, II, III	MAS
	Xgwm389-Xgwm493	I/II	MAS
	Xgwm533, Xgwm493	II, III, IV	QTLs
	Xgwm533-Xgwm493	I/II	MAS
	Xgwm389-Xgwm533-Xbarc133	II, III	MAS
	Xbarc133-Xgwm533-Xgwm493	II, IV	QTLs
	Xgwm389-Xgwm533-Xgwm493	I/II	MAS
	Xgwm533, Xcfd79, Xumn10	II, III, IV	MAS
	Xumn10	II, III	QTLs, MAS
	Xgwm533	III, IV	QTLs
	Xumn10, Xgwm533	II, III	QTLs
	TaHRC-GSM	II	MAS
	Xgwm566, Xwmc231, Xwmc625, Xwmc693, Xwmc307, Xwmc418	II, III	MAS
4B	Xwmc238, Xgwm149	II, III	MAS
	Xwmc245, Xgwm608	II, III	MAS
	Xbarc20-Xwmc349	I	MAS
5A	Xgwm291	I/II	MAS
	Xwmc705, Xgwm304, Xgwm154	II, III	MAS
	Xgwm156, Xgwm304	II, III	MAS
	Xbarc180, Xmag3794	I	MAS
	Xgwm293-Xgwm1057	II	QTLs
	Xgwm304, Xbarc186	II, III, IV	MAS
	Xgwm304, Xbarc117, Xwmc705, Xgwm293	II, III	MAS
6A	Xgwm82	II	QTLs
6B	Xgwm644	I/II	MAS
	Xwmc398-Xgwm644	I/II	MAS
	Xgwm133, Xgwm644	II, III	MAS
7B	Xgwm46	II	QTLs

Sources of FHB resistance and the location of the involved quantitative trait loci (QTLs) in wheat summarized from Khan et al. (2020).

Chromosome	Source
1A	Bess (1A), CJ 9306 (1AS), Ernie (1A), NC-Neuse (1A), NK93604 (1AL)
1B	Arina (1BL), CM-82036 (1B), INW0411 (1B), Seri82 (1B), Remus (1B), Wangshuibai (1B)
1D	DH181 (1DL), Sincron (1DS)
2A	Ning 7840 (2AS), Freedom (2AS), Remus (2A), Renan (2A), Stoa (2AL), Truman (2ASc), Wangshuibai (2A)
2B	Dream (2BL), Ernie (2B), Frontana (2B), Goldfield (2B), IL94-1653 (2B), INW0304 (2B), Ning 7840 (2BL), Renan (2BS)
2D	Alondra (2DS), CJ 9306 (2DL), CS-SM3-7ADS (2D), DH181 (2DS), Gamenya (2DS), McCormick (2D), Nuy Bai (2D), Truman (2DS), Wangshuibai (2D/2DL), Wuhan 1 (2DL)
3A	Forno (3AL), Frontana (3A), Heyne (3AS), Huapei 57-2 (3AS), INW0411 (3A), ND2603 (3AL), Wangshuibai (3AS),
3B	Arina (3BL), Bess (3B), Chokwang (3BS), CJ 9306 (3BS), CM-82036 (3BS), COKER 9474 (3BS), CS-SM3-7ADS (3BS), DH181 (3BS), Ernie (3B/3BL), Huapei 57-2 (3BS/3BL), INW0411 (3BS), Massey (3BL), ND2603 (3BS), Ning 7840 (3BS), Ning 894037 (3BS), Nuy Bai (3BS), McCormick (3BS), Patton (3B), Sumai 3(3BS), Wangshuibai (3BS), Wesley (3BS), Wu 14 (3BS)
3D	Forno (3DS), Patterson (3D), Truman (3DS), Wangshuibai (3DL),
4A	Arina (4AL), Ernie (4A), Heyne (4AL), NC-Neuse (4A)
4B	Chokwang (4BL), Ernie (4BL), IL94-1653 (4B), Stoa (4BS), Wangshuibai (4B), Wuhan 1 (4BS)
4D	Arina (4DS), CS-SM3-7ADS (4D), DH181 (4DL), Heyne (4DL)
5A	Arina (5AL), CJ 9306 (5AS), CM-82036 (5A), DH181 (5AS), Ernie (5A), Frontana (5A), McCormick (5A), Nuy Bai (5AS), Wangshuibai (5A), Wu 14 (5A), Renan (5AL)
5B	Forno (5BL), Patterson (5BL), Wangshuibai (5B)
5D	Chokwang (5DL), Wangshuibai (5DL)
6A	Dream (6AL), Ernie (6A), NC-Neuse (6A), ND2603 (6AS)
6B	Arina (6BL), DH181 (6BS), Frontana (6B), Sumai 3 (6BS), Patton (6B), Wangshuibai (6B)
6D	Arina (6DL)
7A	Frontana (7AS), NK93604 (7AL), Sumai 3 (7A), Wangshuibai (7A)
7B	CJ 9306 (7BS), DH181(7BL), Dream (7BS), Goldfield (7B)

Table 1 Fusarium head blight (FHB) genes with closely linked markers summarized by Ma et al. (2020).

^zChr=chromosome.

^yPFT=pore-forming toxins, HRC=histidine-rich calcium-binding protein, GST=Glutathione S-transferase.

^xType I=FHB incidence, Type II=FHB spread, Type III=DON content.

Table 2 Quantitative trait loci (QTLs) validated and used in marker-assisted selection (MAS) associated with Fusarium head blight resistance in wheat summarized from Ma et al. (2020).

^zChr=chromosome.

^yFHB Type I=FHB incidence, FHB Type II=FHB severity, FHB Type III=DON content, FHB Type IV=Fusarium-damaged kernels, FHB Type I/II=FHB spread.

^xQTLs=QTLs validation in breeding lines.

Table 3 Sources of FHB resistance and the location of the involved quantitative trait loci (QTLs) in wheat summarized from Khan et al. (2020).

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