적요
Powdery mildew (Blumeria graminis f. sp. tritici) significantly affects wheat yield and flour quality. Plant resistance to powdery mildew has been investigated for decades, and numerous resistance genes and quantitative trait loci (QTLs) for molecular markers have been discovered. In Korea, powdery mildew occurs initially in spring, due to frequent rain and low temperatures, becoming severe during the harvest season. In Korea, systematic monitoring and quantitative and qualitative impact assessments of powdery mildew outbreaks have never been conducted properly. Herein, the lifecycle of powdery mildew, resistance genes, QTLs, and selection markers in wheat were examined to elucidate powdery mildew resistance, develop resistant varieties, and genetic markers suitable for the domestic environment.
서 언
평균적으로 매년 전 세계 밀 생산량의 약 10-20%가 질병과 해충으로 인해 손실되는 것으로 추정되며(
Bowen et al. 1991,
Conner et al. 2003,
Oerke 2006) 예상 수확량 손실의 약 10%가 녹병(
Puccinia ssp.), Septoria 잎무늬병(
Septoria spp.), 흰가루병(
Blumeria graminis) 및
Fusarium 종과 같은 곰팡이 병원균 때문인 것으로 보고되었다(
Aboukhaddour et al. 2020,
Oerke 2006,
Simoon et al. 2021).
Blumeria graminis f. sp.
tritici (
Bgt) 에 의해 발생하는 흰가루병은 전세계에서 주로 발생하는 상위 10대 곰팡이병 중에 과학적으로나 경제적 중요성에서 6번째 순위로 꼽히며 병해충으로 인한 수량 감소에서는 8번째로 많은 피해를 일으킨다고 보고되었다(
Dean et al. 2012,
Jankovics et al. 2015,
Savary et al. 2019). 피해정도를 정확한 수치로 나타내기는 어렵지만 일부 연구에서는 수확량 감소의 10-40%, 극단적인 경우 최대 50%의 수확량 손실을 초래할 수 있다고 보고하였다(
Parlange et al. 2015,
Zhang et al. 2017b,
Zhu et al. 2015).
흰가루병은 습도가 높고 온난한 지역에서 질소시비를 많이 하면 쉽게 발생하고, 단작 중심의 과량 시비를 하는 현재 밀 재배 방식에서 발병이 잦을 수 있다. 특히 겨울밀 재배를 주로 하는 우리나라, 일본과 중국을 포함한 아시아, 북⋅동아프리카, 북유럽과 북미지역에서 흔하게 발병하며, 밀 재배에 관수와 질소 시비가 늘어남에 따라 중국에서는 흰가루병은 줄 녹병을 넘어서 가장 피해가 심각한 질병이 되었다(
Austin et al. 2011,
Chen et al. 2007,
Gao et al. 2018,
He et al. 2011,
Li et al. 2014,
Miedaner & Flath 2007,
Mwale et al. 2014,
Shah et al. 2018,
Tang et al. 2018,
Wang et al. 2012). 흰가루병에 의한 피해는 품종의 저항성 정도에 따라 10-34%정도 수량 감소가 나타났으며, 심한 경우에는 50-100%의 피해를 받기 때문에 2005년에서 2008년까지 평균 약 7백만 ha가 흰가루병 피해를 입는 것으로 보고되었다(
Alam et al. 2013,
Gao et al. 2014a,
Gao et al. 2018,
He et al. 2011,
Li et al. 2014,
Luo et al. 2009). 미국 근동질 밀 계통을 이용한 수량 검정에서는 흰가루병에 의한 수량 감소는 34%에 이르렀으며, 흰가루병 감수성인 캐나다 밀 품종의 수량 감소는 최대 19.9%로 나타났고, 러시아, 중국과 브라질에서 흰가루병에 의한 수량 감소는 각각 35%, 40%와 60%에 이른다(
Conner et al. 2003,
Gao et al. 2018,
Johnson et al. 1979).
흰가루병 병원균은 생육 초기에는 뿌리 생장을 지연시키고 분얼을 감소시키며, 출수기와 개화기에는 1수립수를 감소시키고 종실의 충실도와 무게를 급격히 떨어뜨리는데, 곰팡이로 인해 광합성 부위가 줄어 심한 경우 이삭이 맺히지 않거나 이삭이 차지 않아 수량이 급격히 줄어들게 된다. 흰가루병이 일찍 발생하고 병징이 식물체 상단부까지 올라가게 되면 수량 감소는 더욱 심각 해진다(
Gao et al. 2018,
Kang et al. 2020,
Li et al. 2014,
Mwale et al. 2014,
Serrago et al. 2011,
Shah et al. 2018,
Tang et al. 2018). 흰가루병은 전분 및 단백질 함량에 영향을 주기 때문에 종실의 충실도, 밀가루의 반죽 점도 및 가공제품의 품질에도 영향을 미치는 것으로 알려져 있다(
Gao et al. 2018,
Mwale et al. 2014,
Shah et al. 2018,
Tang et al. 2018).
본 론
흰가루병 병원균 생활사
밀의 흰가루병을 일으키는 병원균은
Blumeria graminis f. sp.
tritici (
Bgt)로 활물기생성(biotrophic)이면서 매우 기주 특이적(host specificity)이며 무성생식 단계에서는 분생포자(conidia, conidiospores)를 생성하고, 유성생식 단계에서는 자낭포자(ascospores)를 생성한다. 분생포자와 자낭포자 모두 표면에 수분이 있거나 습한 조건에서 발아하며 95-100% 습도, 10-22°C 온도와 적은 직사광선이 적합한 조건이다. 하지만 25°C 이상의 고온이 지속되면 생장을 멈춰 오히려 발생이 줄어든다(
Beest et al. 2008,
Jankovics et al. 2015). 분생포자는 온화한 겨울에도 번식이 가능하고, 무색의 단세포로 크기는 32-44×12-15 μm이다. 자낭구(cleistothecia)는 검은 갈색으로 거의 구형이며 크기는 130-280 μm으로 자낭은 장타원형으로 이른 봄에 자낭구 안에 무색의 단세포인 20-23×10-13 μm의 크기의 자낭포자를 만드는데, 이러한 분생포자와 자낭포자를 일차 접종원(primary inoculum)이라고 한다(
Glawe 2008). 흰가루병 병원균은 생육 주기가 짧고 공기중에 퍼져 쉽게 전파되며 병원성이 높은 새로운 레이스(race) 발생도 쉽다. 포자가 잎에 떨어지면 2시간 안에 특수한 배아관(germ tube)이 신장해서 부착기(appressorium)를 만들어 잎, 엽초, 줄기와 이삭 등의 표피세포(epidermal cell)에 침투해서 영양분을 흡수하기 위한 원반형의 흡기(haustorium)를 만들고, 5일에서 8일이 지나서 분생포자가 만들어지는데 이를 2차 접종원(secondary inoculum)이라고 한다. 이러한 분생포자는 밀 재배기간 동안에 반복적으로 생장하고 번식하며 밀 생육에 피해를 준다(
Chen et al. 2007,
Kuckenberg et al. 2009,
Walters et al. 2008,
Wright et al. 1995). 무성생식이 끝나고 식물체 표면에 흰색의 고름주머니의 반점(pustules)이 생긴 후 점점 확대되어 원형 또는 타원형이 되고, 표면이 밀가루 뿌린 것과 비슷하게 된 후에 병반은 회색에서 담갈색으로 변하여 흑색의 작은 자낭구가 만들어진다(
Fig. 1) (
Acevedo-Garcia et al. 2017).
Bgt는 감염후에 기주 식물의 양분을 흡수해서 이용하는데, 감염된 조직에서 모든 종류의 당분해효소인 invertase 활성이 증가해서 당을
Bgt에 공급하기 때문에 감염된 잎에서 종자로 전류 될 당이 줄어들고 뿌리의 생육이 억제되는 등 여러 생리적 활성이 억제된다(
Deng et al. 2010,
Sutton et al. 2007).
Bgt는 감염 식물의 전분합성효소인 ADP-glucose pyrophosphorylase, α-1,4-phosphorylase나 sucrose synthase의 발현을 억제하기 때문에 종실의 전분합성이 줄고 이로 인해 수량 감소가 일어난다. 그리고 phosphoglycerate mutase, 3-phosphoglycerate kinase와 UTP-glucose-1-phosphate uridylyltransferase의 발현이 증가해서 해당작용(glycolysis)이 활발해지면서 전분 분해 및 소비를 촉진하게 된다(
Gao et al. 2014b,
Li et al. 2018). 단백질 생합성에 관련된 peptidylprolyl isomerase, cyclophilin A-2와 GTPase ObgE의 발현이 새롭게 나타나는데, 이런 현상은 종실 등숙 과정에서 단백질 함량이 증가하는 것과 유사해서 밀 품질과 밀접한 관련이 있는 triticin, serpin과 high molecular weight-glutenin subunit (HMW-GS)를 증가시키는 걸로 나타난다(
Gao et al. 2014b,
Li et al. 2018). 하지만
Bgt 감염에 의해 전분 및 단백질 합성 관련 효소의 활성 변화가 종실의 단백질 함량 및 질적 특성에 어떻게 얼마나 어떤 영향을 주는지에 대한 관련 연구는 진행중에 있다.
흰가루병 방제
Bgt 방제에 사용되는 살균제는 트리아졸론(triazolone), 플푸트리아폴(flutriafol), 벤지미다졸(benzimidazole), 디니코나졸(diniconazole)과 티오카바메이트(thiocarbamate)가 사용되며 메톡시 아크릴레이트(methoxy acrylate)가 효과적이다. 그러나 같은 메톡시 아크릴레이트계인 메토미노스트로빈(metominostrobin)이나 피라메토스트로빈(pyrametostrobin)과 아족시스트로빈(azoxystrobin)의 동시 사용은 효과를 반감시키고, 프로피코나졸(propiconazole)과 플루시나졸(flusilazole)을 트리아졸론(triazolone)과 교대로 사용하면 밀 수량 감소를 막을 수 있다(
Yang et al. 2013). 살균제를 종자에 처리하거나 엽면 살포를 하면 흰가루병을 감소시킬 수 있으며 종자소독과 엽면 살포를 같이 하면 방제 효율이 더 높아 진다(
Christ & Frank 1989,
Da Luz & Bergstrom 1986,
Hardwick et al. 1994). 살균제를 과도하게 사용하거나 반복적인 사용은 종실의 단백질과 황(sulfur) 함량을 낮추어 빵 품질을 떨어드리고, 식품 안전성에도 치명적이기 때문에 적량을 처리해야 한다(
Gao et al. 2018,
Ruske et al. 2004).
우리나라에서는 4-5월경 비가 잦고 온도가 낮으면 발생하기 시작해서 수확기에 심하게 발생하는데, Bgt방제를 위해서는 적기 파종, 적당한 재식 밀도와 질소 시비, 윤작과 수확 후 Bgt에 감염된 밀 잔재물의 확실한 제거가 도움이 된다. 하지만 병을 일으키는 포자가 공기로 비산되기 때문에 Bgt방제에는 한계가 있다. 국내 Bgt 발생은 강수량과 포장의 과습 상태에 따른 발생이 연차간 지역 간에 차이가 있을 것으로 추정되지만 체계적인 예찰과 병 발생에 따른 수량 및 품질에 미치는 영향에 대한 평가는 이루어지지 않았다. 국내에서는 밀과 보리에 사용하는 흰가루병 살균제로 펜티오피라드(Penthiopyrad) 유제, 마이클로뷰타닐(Myclobutanil) 수화제와 아족시스트로빈 디메토모르프(azoxystrobin difenoconazole) 입상 수화제가 등록되어 있다.
흰가루병 병원균에 대한 면역 반응
최근 흰가루병 병원균 감염 경로에 대한 식물체의 면역 반응 및 기작에 대한 연구가 활발하게 진행되고 있다(
Fig. 2). 식물은 세포 표면의 수용체를 통해 외부 신호를 감지하는데 세포 표면의 pattern recognition receptor (PRR)는 pathogen-associated molecular patterns (PAMP)를 인식하여 pattern-triggered immunity (PTI)를 활성화한다(
Zhao et al. 2022). PTI 활성화는 이온 흐름변화, 활성산소종(reactive oxygen species, ROS) 생성, mitogen-activated protein kinase (MAPK) cascades 같은 하류(downstream) 신호전달로 이어진다(
DeFalco & Zipfel 2021,
Yu et al. 2017). 식물에서 밝혀진 PRR은 대부분이 원형질막에 위치한 receptor-like kinase (RLK)나 receptor-like protein (RLP) 계열에 속하며, RLK와 RLP는 ligand가 결합하는 N-말단 영역에 위치하는 extracellular domain (ectodomain, ECD)의 형태에 따라 leucine-rich repeat (LRR), lectin-like domain, cysteine-rich domain, lysine motif (LysM)와 epidermal growth factor (EGF)-like domain으로 분류된다(
Lee et al. 2021,
Sinha et al. 2022,
Tang et al. 2017, Wu et al. 2013). RLK와 RLP의 ECD는 그에 맞는 ligand와 결합하고, 세포질 kinase domain은 인산화를 통해 세포 내로 신호를 전달하여 특정 리간드에 반응한다(
Couto & Zipfel 2016,
He et al. 2018b,
Hohmann et al. 2017). RLP는 세포질의 kinase domain이 없기 때문에 외부 신호를 세포 내 신호로 변환하기 위해서 RLK와 상호작용을 한다(
Fritz-Laylin et al. 2005,
Mapuranga et al. 2022).
면역수용체(Immune receptor)는 병원체의 침입 시 면역 반응을 일으키는 신호를 감지하여 빠르게 활성상태로 전환된다. PRR이 수용체, 공수용체(co-receptor)와 BIK1 같은 receptor-like cytoplasmic kinases와 복합체를 형성하여 세포질 kinase domain이 인산화 되고, receptor-like cytoplasmic kinases (RLCKs)가 활성화된다(
DeFalco & Zipfel 2021,
Köster et al. 2022,
Zhao et al. 2022). RLCK는 기질 단백질의 인산화로 다양한 하류 신호 전달 경로를 활성화하는데, 수용복합체의 활성화로 인해 막의 이온 채널이 열리면서 Ca
2+이 세포 내로 유입되고 ROS가 생성되며 세포 내 ROS 증가에 따라 살리실산(salicylic acid, SA)과 같은 호르몬 합성 및 전사 재조절 등을 통해 저항성 반응이 나타난다(
DeFalco & Zipfel 2021,
Denance et al. 2013,
He et al. 2018b, Herrera-Vásquez et al. 2015,
Koster et al. 2022,
Liang & Zhou 2018,
Malik et al. 2020).
밀의 leucine-rich repeat (LRR)-RLK인
TaRLK1와
TaRLK2, 이배체 밀
Haynaldia villosa의 lectin type RLK (
LecRK-V)와 malectin-like/LRR-RLK (
RLK-V) 유전자는 곰팡이 결정인자(fungal determinants)를 인식하여 세포 내부로 신호를 전달하는데 관여하는 막단백질 조절을 통하여
Bgt 저항성을 조절하는 것으로 알려져 있다(
Chen et al. 2016,
Hu et al. 2018,
Wang et al. 2018).
TaRLK1와
TaRLK2를 과발현 시킨 계통은 감염 부위에 ROS를 증가시켜
Bgt에 대한 저항성이 증가하였다(
Chen et al. 2016). LecRK-V매개 저항성은 ROS 및 SA 신호전달과 관련된 저항성 기작을 매개하는 것으로 알려져 있으며 LecRK-V 과발현체는
Bgt 흡기 형성 감소로 흰가루병 저항성이 증가하였다(
Hu et al. 2018,
Wang et al. 2018).
RLK-V는 기본 저항성(basal defense)과
Pm21 매개 저항성을 조절하며 선택적 스플라이싱(alternative splicing)에 의해 온전한 형태의
RLK-V1.1과 잘린형태(truncated form)의
RLK-V1.2 두 가지 형태의 전사체를 가지며
RLK-V1.1이 ROS 축적 및 세포사멸을 유도하여 저항성 반응을 일으키는 것으로 알려져 있다(
Hu et al. 2018,
Wang et al. 2018).
식물호르몬인 자스몬산(Jasmonic acid, JA)과 에틸렌(ethylene, ET)의 신호 전달 경로는 병 관련 유전자들의 발현과 ROS 조절을 통해
Bgt에 대한 저항성에 관여한다(
Cao et al. 2014,
Jing et al. 2019,
Li et al. 2017,
Xu et al. 2016,
Zhao et al. 2014). JA에 의해 유도된 basic helix-loop-helix (bHLH) 전사 인자인 TaMYC4는 Jasmonate-ZIM domain (JAZ) 유전자인 TaJAZ1 과 직접적인 상호작용을 통해 전사활성이 억제된다. TaJAZ1의 ZIM 도메인은 NOVEL INTERACTOR OF JAZ (MINJA)의 C-말단과 상호 작용하고 TaNINJA의 N-말단 EAR 모티프는 TOPLESS (TPL)와 상호작용하여 TaJAZ1-TaNINJA-TaTPL 전사공동억제자 복합체를 형성한다.
Bgt 감염에 의해 JA가 축적되면 TaJAZ1을 분해하고 전사공동억제자 복합체 형성을 방해하여
TaPR유전자들의 발현과 ROS 축적을 억제한다(
Jing et al. 2019). 그리고 RNA Polymerase Ⅱ (RNA Pol Ⅱ)는 TaMED25와 ETHYLENE INSENSITIVE3-LIKE1 (TaEIL1) 사이의 상호작용에 의해 ETHYLENE RESPONSE FACTOR 1 (
TaERF1)의 전사를 상향조절하여
PR 유전자의 발현과 ROS의 축적을 억제한다(
Liu et al. 2016).
다양한 탈아세틸화효소(histone deacetylase, HDAC)도 병원균 감염에 대한 식물 저항성에 양성 및 음성 조절자 역할을 한다(
Ding & Wang 2015,
Kong et al. 2020,
Ramirez-Prado et al. 2018).
Bgt 감염에 의해 히스톤 아세틸화효소(Histone acetyltransferase, HAT)가 증가되고, 히스톤 아세틸화효소가 히스톤의 라이신(Lys) 잔기에 아세틸기를 증가시키면 염색질(chromatin) 구조가 느슨해저 방어 관련 유전자의 발현이 촉진된다. 반면에 탈아세틸화효소에 의해 아세틸 그룹이 제거가 되면 염색질 구조가 긴밀해지면서 전사가 억제된다. 밀의 RPD3타입 탈아세틸화효소인 TaHDA6는 WD repeats (WDR) 단백질인 TaHOS15와 상호작용하여 히스톤의 탈아세틸화를 통해 방어 관련 유전자의 발현을 억제한다(
Liu et al. 2019). 탈아세틸화효소 2 (HD2)의 구성원인 TaHDT701은 TaHDA6 및 TaHOS15와 상호작용하여 HDAC복합체를 형성하는데 이 HDAC복합체는
TaPR1,
TaPR2,
TaPR5 및
TaWRKY45와 같은 밀 방어 관련 유전자에서 히스톤 탈아세틸화를 매개하여
Bgt에 대한 방어 관련 유전자 전사 및 방어 반응을 억제한다.
현재까지 일반 밀 및 야생 근연종에서 63개의 유전자좌(
Pm1-Pm66)에서 100개 이상의 흰가루병 저항성 유전자가 확인되었는데(
Mapuranga et al. 2022,
Zhang et al. 2019)
Pm1 (
a-e),
Pm2 (
a-c),
Pm3 (
a-g,
k-t),
Pm4 (
a-d),
Pm5 (
a-e)와
Pm24 (
a-b)는 다중대립유전자(multiple alleles)를 지니고 있다(
Hsam et al. 1998,
Hsam et al. 2001,
Huang et al. 2003,
Huang et al. 2000b,
Ma et al. 2015,
Schmolke et al. 2012,
Singrun et al. 2003,
Xu et al. 2015,
Xue et al. 2012). 그 중에 몇 개의 유전자
Pm1a,
Pm2a,
Pm2b,
Pm3b,
Pm5e,
Pm8,
Pm17,
Pm21,
Pm24,
Pm38,
Pm41,
Pm46과
Pm60 만이 클로닝 되었다(
He et al. 2018a,
Hewitt et al. 2021,
Hurni et al. 2013,
Jin et al. 2022,
Krattinger et al. 2009,
Li et al. 2020,
Lu et al. 2020,
Moore et al. 2015,
Sanchez-Martin et al. 2016,
Singh et al. 2018,
Xie et al. 2020,
Xing et al. 2018,
Yahiaoui et al. 2004,
Zou et al. 2018). 그러나 이들 저항성 유전자들의 대부분은 레이스 특이적인 유전자로 새로운 레이스에 의해 쉽게 대체된다. 그리고 현재까지
Bgt 저항성 유전자로 명명되지 않은 임시유전자(temporary gene)는
MlBrock,
MlIW170,
Pm5055,
PmAF7DS,
PmD57,
PmFG,
PmG3M,
PmHNK54,
PmLX66,
PmW14,
PmZ155,
Pm-M53,
PmTb7A.1,
PmTb7A.2,
PmV,
PmX3986-2와
PmYm66 등이 보고되었다(
Huang et al. 2012,
Li et al. 2009,
Li et al. 2014,
Li et al. 2017,
Ma et al. 2011,
Ma et al. 2014,
Qi et al. 1996,
Xie et al. 2012). 2018년까지 보고된 54개 유전자좌 중에서 27개가 B게놈에 위치하며 13개가 D 게놈에 위치한다. A와 D 게놈에서 매핑 된
Pm 유전자는 각 42개와 17개이며, 유전자좌당 평균
Pm 대립유전자 개수는 A게놈이 3.00개로 B게놈(1.19)이나 D게놈(1.31)보다 많았다(
Tang et al. 2018). 다중대립유전자인
Pm1,
Pm2,
Pm3,
Pm4,
Pm5와
Pm24는 각각 7AL, 5DS, 1AS, 2AL, 7BL과 1DS에 위치하고 있으며(
Ma et al. 2016), 임시유전자를 제외한 상태에서는 2B (
Pm6,
Pm26,
Pm33,
Pm42,
Pm49), 5D (
Pm2,
Pm34,
Pm35,
Pm46), 6B (
Pm11,
Pm12,
Pm14,
Pm27)와 7D (
Pm15,
Pm19,
Pm29,
Pm39)에 많은 유전자가 위치하고 있다(
Gao et al. 2018,
Kang et al. 2020,
Li et al. 2014,
Mwale et al. 2014,
Qin et al. 2011,
Shah et al. 2018,
Tang et al. 2018).
Pm 저항성 유전자는 레이스 특이적(race-specific) 저항성과 레이스 비특이적(race-non-specific) 저항성 타입으로 나뉜다. 첫번째 레이스 특이적 저항성은 질적 저항성(qualitative resistance), 모든 식물 성장 단계 저항성(all stage resistance, ASR) 또는 단일유전자 저항성이라고도 부른다. 레이스 특이적 저항성은
Bgt에 대한 유전자 대 유전자 대응 방식(gene-for-gene manner)으로 관련된 단일 주동 유전자(major gene, R gene)에 의해서 저항성을 가지게 된다(
Mapuranga et al. 2022,
Maurya et al. 2021).
Pm4a (MCTP-kinase),
Pm24 (tandem kinase gene WTK3),
Pm38 (ATP-binding cassette (ABC) transporter)과
Pm46 (hexose transporter)을 제외한 대부분의 흰가루병 저항성 유전자는 주로 Nucleotide binding site-Leucine rich Repeat (NLR) 형태의 수용체를 암호한다(
Koller et al. 2018,
Krattinger et al. 2009,
Lu et al. 2020,
Moore et al. 2015,
Sanchez-Martín et al. 2021). 그러나 대부분의 레이스 특이적 저항성은
Bgt에 대한 내구성(durability)이 낮아 3-5년 지나면 새로운 레이스 출현으로 쉽게 저항성을 잃게 된다(
Lillemo et al. 2010,
Shamanin et al. 2019,
Wang et al. 2005). 1980년대에 중국에서는
Pm8저항성 유전자를 포함하는 품종이 70%에 달하였으나
Pm8저항성이 무너졌고(
Sheng et al. 1995), 현재 중국 일부지역에서는 단일
Pm21 유전자의 대규모 빈번한 사용으로 저항성이 무너질 위기에 놓여 있다. 미중부지역에서는
Pm3a (
Niewoehner & Leath 1998)와
Pm4a (
Parks et al. 2008)가 몇 년을 넘기지 못했고,
Pm17도 5년을 넘기지 못했다(
Cowger et al. 2009,
Cowger et al. 2018). 호주에서는
Pm4b,
Pm7,
Pm24와
Pm28이 4년을 넘기지 못했다(
Golzar et al. 2016). 따라서 흰가루병에 대한 넓은 범위의 병원체 저항성을 가진 새로운 저항성 유전자의 탐색 및 품종개발이 필요하다.
Bgt 유전자 중에서 1AS에 위치한
Pm3에 대한 연구가 많이 이루어졌는데, 현재까지 20개(
Pm3a-
Pm3t)의 대립유전자가 보고되었으며 이중에 17개(
Pm3a-
Pm3g,
Pm3k-
Pm3t) 유전자가 클로닝 되었다(
Bhullar et al. 2010a,
Yahiaoui et al. 2006). 이들 유전자는 단백질 수준에서 97% 유사성을 보였으며
Pm3h,
Pm3i와
Pm3j는 각각
Pm3d,
Pm3c,
Pm3b와 일치하였다(
Bhullar et al. 2009,
Bhullar et al. 2010b,
Bougot et al. 2002,
Huang et al. 2004,
Yahiaoui et al. 2006). 특히
Pm3 유전자좌를 특이적으로 검출하는 184 bp의 단편인 UP3 서열이
Pm3 대립유전자 뿐만 아니라 호밀에서 유래한 유전자인
Pm8과
Pm17에서도 발견되었다(
Hurni et al. 2013,
Singh et al. 2018),
Pm3는 감수성 자원인
Pm3CS에서 진화한 것으로 보이며(
Yahiaoui et al. 2004)
Pm3 형질전환체 연구를 통해서 하나의 대립유전자만 있는 모계보다 특정 조합에서 저항성이 증가하는 것으로 나타났다(
Koller et al. 2018). 그러나 같은 품종에
Pm3가 있는 경우에는
Pm3가 번역 후기 단계(post-translational stage)에서
Pm8유도 신호전달을 방해해서 오히려 저항성을 억제하는 것으로 나타났으며 (
Hurni et al. 2014) 이러한 상호억제는
Pm3b와
Pm3f 에서도 보였다(
Stirnweis et al. 2014). 그리고
Pm3 NLR receptor는 숙주 특이성과 흰가루병 레이스 저항성을 결정하는 것으로 밝혀졌다(
Bourras et al. 2015,
Bourras et al. 2019).
두번째로 레이스 비특이적 저항성은 양적 저항성(quantitative resistance)이라고 부르며 성체 식물 단계에서 저항성의 징후와 관련이 있어 성체 식물 저항성(adult plant resistance, APR)이라고도 하는데 성체 식물 단계에서 활성화되며 종종 부분저항성 표현형으로 나타난다(
Mapuranga et al. 2022,
Maurya et al. 2021). 레이스 비특이적 저항성은 소수의 유전자가 저항성을 부여하는 역할을 해 미동 유전자 (minor gene) 저항성이라고도 한다(
Maurya et al. 2021). 레이스 비특이적 저항성 유전자는 R 유전자 보다 특정 유전자에 대한 저항성은 낮지만 거의 모든 레이스에 효과적이기 때문에 특정
Bgt대발생으로 인해 피해가 커지는 것을 막을 수 있는 내구성은 레이스 특이 저항성보다 높다(
Burdon et al. 2014,
Li et al. 2014,
Liu et al. 2001). 레이스 비특이적 저항성을 조절하는 유전자의 수나 효과는 레이스 특이적인 저항성과 다르지만 여러 조절 메커니즘 중에 레이스 특이 저항성과 중복되기도 하며(
Poland et al. 2009) 미국에서 널리 재배되는 Genesee 및 Knox 같은 일부 APR 품종은 조절된 조건(controlled condition)에서 단일 우성 유전자에 의해 조절되는 질적 패턴을 보이기도 했다(
Bennett 1984,
Ellingboe 1976). APR 유전자가 R 유전자와 달리 대부분의
Bgt에 부분저항성을 보이는 것은 질적 유전자에 해당하는 주동유전자, APR 유전자와 미미한 영향을 미치는 저항성 유전자들의 불완전 발현으로 보고 있다(
Burdon et al. 2014,
Niks et al. 2015).
Pm 유전자 중에서
Pm38 (
Lagudah et al. 2009),
Pm39 (
Lillemo et al. 2008)와
Pm46 (
Moore et al. 2015)이 대표적인 APR유전자이다(
He et al. 2018a). 일부 APR유전자는 동일한 유전자좌에서 잎녹병(
P. triticina, Lr), 줄무늬 녹병(
P. striiformis f. sp.
tritici, Yr), 줄기 녹병(
graminis f. sp.
tritici, Sr) 과 흰가루병 등 하나이상의 광범위한 병원체 종에 대한 저항성을 제공하고 경우에 따라 복합병원체 종에 대한 저항성을 제공하는데
Lr34/Yr18/Sr57/Pm38/Ltn1 (
Krattinger et al. 2009),
Lr46/Yr29/Pm39/Sr58/Lnt2 (
Singh et al. 2013)와
Lr67/Yr46/Pm46/Sr55/Lnt3 (
Moore et al. 2015)가 그 예이다. 잎녹병, 줄무늬 녹병, 줄기 녹병과 흰가루병의 대한 저항성과, 잎 끝 괴사 표현형(leaf tip necrosis) 관련 유전자(
Ltn)와도 연관되어 있다(
Dyck 1991,
Hulbert et al. 2007,
Lagudah et al. 2006,
Risk et al. 2012,
Shah et al. 2011,
Kim et al. 2020a,
Kim et al. 2020b). 이들 유전자가 abscisic acid (ABA)와 당(sugar) 신호전달과 상관 있기 때문에 방어활성화(defense activation) 기작과 상관이 있으며(
Krattinger et al. 2009,
Moore et al. 2015) 병원균에 의해 유도되는 PTI와 ETI와도 상관이 있다(
Andolfo & Ercolano 2015,
Bolouri Moghaddam & Van den Ende 2012,
Eveland & Jackson 2012,
Robert-Seilaniantz et al. 2011,
Ton et al. 2009,
Valluru & Van den Ende 2011). 이들 유전자는 멘델의 유전법칙을 따르고,
Lr34/Yr18/Pm38/ Sr57의 경우 병저항성 효과를 크게 나타내는 것이
Pm38이기 때문에 주동유전자인
Pm38로 표기하기도 한다(
Lillemo et al. 2008).
Pm3a,
Pm5,
Pm6,
Pm12,
Pm16,
Pm20,
Pm35와
Pm43도 APR 유전자로 보고되었으며
Pm1,
Pm2,
Pm3,
Pm4,
Pm12,
Pm13,
Pm21과
Pm25는 레이스 특이적 저항성이면서 APR인것으로 보고되었다(
Cenci et al. 1999,
Hartl et al. 1999,
Hsam et al. 2003,
Hu et al. 1997,
Jia et al. 1996,
Lan et al. 2009,
Liang et al. 2006,
Ma et al. 1994,
Muranty et al. 2009,
Purnhauser et al. 2011,
Qi et al. 1996,
Shi et al. 1998,
Zhao et al. 2013). APR 유전자는 생육기간 전반에 걸쳐 중간이상의 저항성을 나타내며 이와 관련된 유전자는 4B, 5A, 5D와 7번 염색체에 위치하는 것으로 알려져 있다(
Burdon et al. 2014,
Li et al. 2014,
Poland et al. 2009).
감수성유전자(susceptibility genes, S gene)도 흰가루병 병원균 감염에 관여를 하는데, S gene를 비활성화하거나 knock-down 시키면 저항성이 생긴다(
Appiano et al. 2015,
Consonni et al. 2006,
Pessina et al. 2016,
Wang et al. 2014). 대표적인 S gene은
MLO (Mildew-Locus-O)이며, 많은 식물종에서
MLO의 기능상실 돌연변이(
mlo)가 흰가루병에 대한 저항성을 유발한다(Büschges et al. 1997).
MLO는
R 유전자와 달리 열성인
mlo가 저항성을 부여하지만(
Jorgensen 1992) 밀에서는 자연 발생적인
mlo 유전자 돌연변이가 발견되지 않았다(
Acevedo-Garcia et al. 2017). 3개의 보리 MLO 유사체
TaMLO-A1,
TaMLO-B1과
TaMLO-D1이 각 5AL, 4BL과4DL 염색체에서 보고되었으며(
Devoto et al. 2003,
Gao et al. 2018,
Varallyay et al. 2012) nucleotide와 단백질 수준에서 각 98%와 99%의 유사도를 보였고(
Wang et al. 2014), 보리
MLO와는 88% 유사도를 보였다(
Elliott et al. 2002).
TaMLO-B1의 과발현으로 흰가루병에 대한 감수성이 높아질 뿐만 아니라 표면의 원반형의 흡기(haustorium)도 비정상적으로 증가하였다(
Elliott et al. 2002).
mlo 돌연변이는 의도치 않은 다면발현 효과(pleiotropic effects)로 인한 생장저하(growth penalty)를 유발하는데, 보리와 애기장대에서
mlo 돌연변이는 캘러스 축적(callose deposition) 및 세포사(cell death), 노화(senescence) 유발 등이 보고되었다(
Consonni et al. 2006,
Humphry et al. 2006,
Piffanelli et al. 2004). 하지만 targeting induced local lesions in genomes (TILLING) 기술을 이용한 밀의
TaMLO 돌연변이체에서
Bgt 저항성을 보였으나 식물성장에 영향을 나타내지는 않았고 (
Acevedo-Garcia et al. 2017), 밀의 A, B와 D 게놈의
TaMLO을 모두 knock-out 시킨 돌연변이체(
Tamlo-aabbdd) 중에
Tamlo-R32는 흰가루병에 강한 저항성을 가지면서도 성장저하 및 생산량 감소를 나타내지 않았다(
Wang et al. 2014).
Tamlo-R32는 4B 염색체에서
MLO 유전자를 포함하여 앞쪽으로 304 kb 결실이 발생하면서 앞쪽에 있던 Tonoplast monosaccharide transporter 3 (
TaTMT3B) 유전자의 발현이 증가하여 성장 및 생산량 저하를 상쇄시킨 것으로 보인다(
Wang et al. 2014,
Li et al. 2022). 이와 유사한 음성적 조절자(negative regulator)로 enhanced disease resistance 1 (
EDR1)의 돌연변이는 흰가루병 뿐만 아니라 일반적인 저항성도 나타냈다(
Huckelhoven 2005).
캐나다 품종인 6배체 밀 Axminster (
Pm1a) (
Sears & Briggle 1969)에서 처음으로 보고된 이후에 많은 6배체 밀 저항성 자원들이 보고되었으며 현재까지 보고된 자원 중에 중국 자원이 가장 많다. 중국자원으로는 Ulka (
Pm2a), KM2939 (
Pm2b), Xiaobaidong (
Pm5a), CI 10904 (
Pm5d), Fuzhuang 30 (
Pm5e), Chinese Spring (
Pm11), 82-7241 (
Pm23), Chiyacao (
Pm24a), Baihulu (
Pm24b), D57 (
Pm45)과 Hongyanglazi (
Pm47), 일본 자원으로는 Asosan (
Pm3a), Norin 10 (
Pm14)과 Norin 26 (
Pm10,
Pm15), 독일 자원으로는 Kolibri (
Pm3d), Ibis (
Pm5c), N14 (
Pm9)과 Tabasco (
Pm46), 러시아 자원으로는 Chul (
Pm3b)과 GUS 122 (
Pm3j), 호주 계통 W150 (
Pm3e)과 RL6058 (
Pm38)이 보고되었다. 그리고 멕시코 Sonora (
Pm3c)와 Amber (
Pm3f), 프랑스 Aristide (
Pm3g), 네팔 N324 (
Pm3i), 이탈리아 Virest (
Pm1e), 에스토니아 Meri (
Pm28), 노르웨이 Saar (
Pm39)와 방글라데시 Hombar (
Pm44) 자원도 보고 되었다(
Alam et al. 2011,
Briggle 1969,
Gao et al. 2012,
Hsam et al. 1998,
Huang et al. 2000b,
Huang et al. 2004,
Li et al. 2014,
Lillemo et al. 2008,
Peusha et al. 2000,
Schneider et al. 1991,
Sears & Briggle 1969,
Singrün et al. 2003,
Sourdille et al. 1999,
Spielmeyer et al. 2005,
Tosa et al. 1987,
Tosa et al. 1988,
Tosa & Sakai 1990,
Xiao et al. 2013,
Xue et al. 2012). 그리고 국제 밀-옥수수 연구소(CIMMYT)에서 육성된 Francolin#1과 Quaiu#3 계통은 중국에서 분리한
Bgt에 APR저항성을 지닌 것으로 보고되었고, 미국 추파밀인 Knox 62, Massey와 Redcoat는 중도이상의 APR 유전자를 2-3개 지닌 것으로 알려져 있다. 특히 Knox 62는 20년 이상 APR효과를 지닌 품종으로 유명하다(
Liu et al. 2001). 그리고 프랑스 추파밀인 RE714는 또다른
Bgt 저항성 유전자 powdery mildew resistance gene (
MlRE)을 가지고 있는 것으로 보고되었다(
Chantret et al. 1999,
Das & Griffey 1994,
Ren et al. 2017). 3개의 APR유전자
Pm38 (
Lr34/Yr18/Pm38/Sr57),
Pm39 (
Lr46/ Yr29/Pm39/Sr58)와
Pm46 (
Lr67/Yr46/Pm46/Sr55)를 지닌 계통이나 자원은 전세계적으로 유용하게 이용되고 있으며, 중국의 경우 CIMMYT의 유용자원을 활용한 shuttle breeding을 통해서 흰가루병과 녹병에 저항성인 08RC2525 (Chuanmai 32*2/Chapio), 07RC3941 (SW119*2/Tukuru)과 07RC3929 (SW119*2/Tukuru)를 육성하였다(
Li et al. 2014).
6배체 재배밀 이외에 2배체와 4배체 밀
T. durum (
Pm3h),
T. spelta (
Pm1d),
T. monococcum (
Pm1b,
Pm4d),
T. dicoccoides (
Pm3k,
Pm16,
Pm26,
Pm30,
Pm31 (=
Pm21),
Pm36,
Pm41,
Pm42),
T. dicoccum (
Pm4a,
Pm5a,
Pm49,
Pm50),
T. carthlicum (
Pm4b,
Pm33),
T. sphaerococcum (
Pm5c)과
T. timopheevii (
Pm6,
Pm27,
Pm37)에서도 저항성 유전자가 도입되었으며, 야생밀
Ae. tauschii (
Pm2,
Pm19,
Pm34,
Pm35),
Ae. speltoides (
Pm12,
Pm32),
Ae. longissimi (
Pm13)와
Ae. ovata (
Pm29), 그리고 밀 이외에 호밀(
Pm7,
Pm8,
Pm17,
Pm20),
Haynaldia vilosa (
Pm21)와
Elytrigia intermedium (
Pm40,
Pm43)에서도 저항성 유전자가 도입되었다(
Alam et al. 2011,
Bhullar et al. 2009,
Bhullar et al. 2010a,
Bhullar et al. 2010b,
Briggle 1969,
Chen et al. 1995,
He et al. 2009,
Hsam et al. 1998,
Hu et al. 2008,
Hua et al. 2009,
Huang et al. 2000a,
Huang et al. 2004,
Huang & Roder 2011,
Luo et al. 2009,
Miranda et al. 2007,
Mohler et al. 2013,
Nematollahi et al. 2008,
Peusha et al. 1996,
Piarulli et al. 2012,
Rong et al. 2000,
Schmolke et al. 2012,
Singrun et al. 2003,
Sourdille et al. 1999,
Srnic et al. 2005,
Tao et al. 2000,
The et al. 1979,
Xiao et al. 2013,
Xie et al. 2004,
Xue et al. 2012,
Yahiaoui et al. 2004,
Yahiaoui et al. 2009,
Zhong et al. 2016).
염색체별로 보면 염색체 1A에는
T. militinae으로부터 유래한 유전자와 QTL 영역이 포함되어 있으며 염색체 2A의 QTL 부위는 다양한 야생밀 자원(
T. monococcum,
T. dicoccum,
T. timopheevii,
T. carthilicum)의 유전자와 QTL이 도입된 영역을 포함한다. 이러한 도입은 염색체 2B (
T. carthlicum,
T. dicoccoides), 5A (
T. monococcum,
T. militinae), 5B (
T. dicoccoides)와 6B (
T. timopheevi,
T. dicoccoides,
Ae. speltoides)에서도 나타난다(
Jakobson et al. 2012,
Marone et al. 2013).
저항성 유전자
Pm1a,
Pm1c,
Pm2,
Pm3a,
Pm3b,
Pm3c,
Pm3g,
Pm4a,
Pm6,
Pm12,
Pm13,
Pm17,
Pm18,
Pm21,
Pm26,
Pm27과
Pm29와 연관된 restriction fragment length polymorphisms (RFLP) 마커가 개발되었다(
Cenci et al. 1999,
Hartl et al. 1995,
Hartl et al. 1999,
Hsam et al. 2000,
Jarve et al. 2000,
Jia et al. 1996,
Liu et al. 1999,
Liu et al. 2000,
Ma et al. 1994,
Mohler & Jahoor 1996,
Rong et al. 2000,
Sourdille et al. 1999,
Tao et al. 2000,
Zeller et al. 2002). 그리고
Pm1,
Pm2,
Pm3,
Pm3a,
Pm3b,
Pm3c,
Pm4a,
Pm12,
Pm13,
Pm18,
Pm21과
Pm25에서 random amplified polymorphisms DNA (RAPD) 마커가 개발되었으며
Pm21에서는 sequence characterized amplified region (SCAR) 마커가 개발되었다(
Cenci et al. 1999,
Hartl et al. 1995,
Hu et al. 1997,
Liu et al. 1996,
Liu et al. 1999,
Ma et al. 1994,
Qi et al. 1996,
Shi et al. 1998).
Pm1c,
Pm4a,
Pm17,
Pm24,
Pm29와
Pm42에서는 amplified fragment length polymorphisms (AFLP) 마커가 개발되었고(
Hsam et al. 2000,
Hua et al. 2009,
Huang et al. 2000b,
Zeller et al. 2002),
Pm1e,
Pm5e,
Pm24a,
Pm24a,
Pm24b,
Pm27,
Pm30,
Pm33,
Pm34,
Pm35,
Pm36,
Pm40,
Pm41,
Pm42,
Pm43과
Pm45에서 simple sequence repeats (SSR) 마커가 개발되었으며(
Table 1,
Fig. 3),
Pm1a,
Pm2,
Pm4a,
Pm6,
Pm13,
Pm41,
Pm42와
Pm45에서 sequence tagged site (STS) 마커가 개발되었다(
Blanco et al. 2008,
Cenci et al. 1999,
He et al. 2009,
Hu et al. 1997,
Hua et al. 2009,
Huang et al. 2000b,
Jarve et al. 2000,
Li et al. 2009,
Liu et al. 2002,
Luo et al. 2009,
Ma et al. 2011,
Miranda et al. 2006,
Miranda et al. 2007,
Mohler & Jahoor 1996). 현재 SSR와 STS 마커가 가장 많이 이용되고 있다.
현재까지 100여개가 넘는 APR QTL이 보고되었는데 APR QTL은 완전히 감염이 없는 저항성이 아니라 생육 기간에
Bgt 감염의 최소화를 의미하며 레이스 특이적 저항성을 포함하여 여러가지 유전자가 관여하는 양적인 의미의 저항성으로 이해해야 한다(
Burdon et al. 2014,
Li et al. 2014,
Poland et al. 2009). APR QTL은 R 유전자 보다
Bgt 저항성에 내구성이 높지만 감염력이 높은 경우에는 QTL의 효과는 낮아진다. 따라서 흰가루병 발병이 심한 경우에는 R 유전자가 효과적이고 중간 이하의 발병에서는 APR QTL이 효과적이다(
Gao et al. 2018,
Kang et al. 2020,
Li et al. 2014,
Liu et al. 2001,
Mundt 2014,
Mwale et al. 2014,
Shah et al. 2018,
Tang et al. 2018). CIMMYT의 Francolin#1과 Quaiu#3의 recombinant inbred line (RIL) 집단에서 6개의 QTL (
QPm.heau-1BL (
Pm39),
QPm.heau-1DL,
QPm.heau- 2DL,
QPm.heau-4BL,
QPm.heau-5BL,
QPm.heau-6BS)이 보고되었는데
QPm.heau-1DL은 새롭게 보고된 것이고,
QPm.heau- 1BL (
Pm39)과
QPm.heau-2DL은
Yr54과 같은 위치에 있으며 다면발현 저항성 유전자인 APR QTL이다(
Ren et al. 2017). 중국 계통인 Zhou8425B에서 4개의
Bgt 저항성 QTL로 전체
Bgt 저항성의 27%를 설명할 수 있었고, 이들 4개 QTLs중에서
QPm.caas-3BS는 녹병 유전자인
QLr.hebau-3BS와 같은 위치에 있는 새로운 APR QTL이다(
Jia et al. 2018,
Zhang et al. 2017a). APR QTL는 1AS, 1AL, 2AS, 2AL, 3AS, 4AL, 5AL와 5BS에 각각 8개씩으로 많았고, 16개 APR QTL는 잎 녹병과 흰가루병에 동시에 저항성이었으며 11개 유전자좌는 잎 녹병, 줄 녹병과 흰가루병에 저항성이었다(
Table 2,
Fig. 3). 중국 계통 Bainong 64에서 보고된
QPm.caas-1AL과
QPm.caas-4DL은 흰가루병의 발병 심도를 낮추는 것과 상관이 있었으며
QPm.caas-3BS는 녹병 QTL인
QLr.hebau-3BS과 같은 위치인 것으로 보아 흰가루병과 녹병에 동시에 저항성 인 것으로 추정된다(
Bai et al. 2012,
Zhang et al. 2017a). Genome-wide association mapping (GWAS)나 연관 맵핑(linkage mapping)을 통해서 중간 정도의 발병에서는 4-5 QTL를 가진 품종이나 계통이 흰가루병에 저항성을 가지는 것으로 나타났다(
Asad et al. 2012,
Jia et al. 2018). 보리에서는 과산화효소 유전자군(peroxidase gene family)이
Bgt와 녹병에 저항성인 것으로 알려져 있는데, 밀에서도 이와 유사할 가능성이 높기 때문에 관련 유전자에 대한 연구가 이루어지고 있다(
Gonzalez et al. 2010).
결 론
밀은 흰가루병에 의해 수량 감소뿐만 아니라 전분 및 단백질 함량과 밀가루 품질에도 영향을 받는데 품종이나 발병시기 및 평가 방법에 따른 차이가 크다. Bgt방제를 위해서는 적기 파종, 적당한 재식 밀도와 질소 시비, 윤작과 수확 후 Bgt에 감염된 밀 잔재물의 확실한 제거와 적정한 살균제 처리 등이 도움이 된다. 하지만 병을 일으키는 포자가 공기로 비산되기 때문에 Bgt방제에 한계가 있고 병원성이 높은 새로운 레이스 발생도 쉽다. 현재까지 많은 저항성 유전자(Pm gene)가 알려져 있지만 클로닝된 유전자는 몇 개 되지 않으며 이들 저항성 유전자들의 대부분은 레이스 특이적인 유전자로 Bgt에 대한 내구성이 낮아 3-5년 지나면 새로운 레이스 출현으로 쉽게 저항성을 잃게 된다. 따라서 흰가루병에 대한 넓은 범위의 병원체 저항성을 가진 새로운 저항성 유전자의 지속적인 탐색 및 품종개발이 필요하다. 우리나라에서는 4-5월경 비가 잦고 온도가 낮으면 발생하기 시작해서 수확기에 심하게 발생하는데 흰가루병 발생에 대한 체계적인 예찰이나 흰가루병 발생에 따른 수량 및 품질의 영향에 대한 평가는 제대로 이루어진 적이 없는 실정이다. 따라서 국산 밀 안정적 공급과 국내환경에 맞는 저항성 품종 육성을 위해 먼저 정밀 검정체계 구축을 통한 표현형 데이터 획득이 필요하다. 그리고 효율적인 분자 마커 개발을 위해 genome-wide association mapping (GWAS)와 Genomic selection (GS) 분석으로 QTL 및 유전자 선별 등 marker-associated selection (MAS) 연구를 통해 국내환경에 맞는 품종개발 및 선발 마커 개발이 이루어져야 한다. 또한 면역체계의 신호전달 연구와 수량이나 농업형질의 피해 없이 저항성을 증진한 유전자의 유전자 편집을 통한 저항성 증진 연구도 함께 이루어져야 한다.
적 요
흰가루병(Blumeria graminis f. sp. tritici)은 밀 생산에 상당한 경제적 손실을 초래한다. 수십 년 동안 밀 흰가루병에 대한 식물 저항성이 조사되었으며 많은 저항성 유전자 및 양적 형질 유전자좌(QTL)를 발견하였다. 국내에서는 4-5월경 비가 잦고 온도가 낮으면 발생하기 시작해서 수확기에 심하게 발생하며 수확량, 단백질 함량 및 밀가루 품질에도 영향을 준다. 그러나 국내에서는 흰가루병 발생에 대한 체계적인 예찰이나 흰가루병 발생에 따른 수량 및 품질의 영향에 대한 평가도 제대로 이루어진 적이 없다. 본 연구에서는 흰가루병에 대한 이해와 보고된 저항성 유전자, QTL 및 유전자 마커에 대한 최근 연구 동향을 정리하여 국내에 맞는 저항성 품종 개발과 선별 마커 개발에 도움이 되고자 하였다.
사 사
본 연구는 2023년도 농촌진흥청 국립식량과학원 전문연구원 과정 지원사업과 농촌진흥청 연구사업(과제명: 가공 용도별 밀 품종개발 및 육종효율 증진연구(4단계), 과제번호: PJ 016771012023)에 의해 이루어진 것임.
Fig. 1Wheat powdery mildew (Blumeria graminis f. sp. tritici) life cycle.
Fig. 2Schematic representation of wheat defense responses to
Bgt infection from
Mapuranga et al. (2022). ABA, abscisic acid; ET, ethylene; ETI, effector-triggered immunity; ETS, effector-triggered susceptibility; HAT, histone acetyltransferase; JA, Jasmonate; PR, pathogenesis-related; PRR, pattern recognition receptor; PTI, PAMP-triggered immunity; ROS, reactive oxygen species.
Fig. 3Projection of wheat
Pm genes (in bold) and QTL (blue bars) onto wheat chromosomes, grey section on each chromosome represents centromere. The positions of these genes and QTL are estimated from their flanking molecular markers from previous publications in integrated map (
Maccaferri et al. 2015) and from
Kang et al. (2020). For those with either flanking marker or close marker not available, are excluded from map. Some Pm genes which are placed at the same position, only one gene randomly chosen to be present, the number of other genes in this case is given in bracket after.
Table 1Genes associated with powdery mildew resistance, their chromosomal location, cultivar/line, source and molecular markers modified from
Kang et al. (2020).
Table 1
|
Gene |
Location |
Type of marker |
Closest/flanking marker |
Cultivar/line |
Source |
|
Pm1a
|
7AL |
RFLP, RAPD, STS |
UBC320420, UBC638550, CDO347
WHS178, Xmwg2062, Xcdo347, Xpsr121, Xpsr148, Xpsr680, Xpsr687, Xwir148, XC607, STS638542, Xksuh9
|
Axminster |
T. aestivum
|
|
Pm1b
|
|
|
MocZlatka |
T. monococcum
|
|
Pm1c (Pm18) |
RFLP, AFLP, RAPD |
Xwhs178, OPH-111900, S19M22, S14M20
|
Weihestephan M1N |
T. aestivum
|
|
Pm1d
|
|
|
Duhamelianum |
T. spelta
|
|
Pm1e (Pm22) |
SSR, AFLP |
Xgwm344, S13M26
|
Virest |
T. aestivum
|
|
Pm2a
|
5DS |
RFLP, STS |
Xwhs350, Xwhs295
Xbcd1871, STSwhs350
|
Ulka/XX 194 |
T. aestivum
/Ae. tauschii
|
|
Pm2b
|
SSR |
Xcfd81, Xbwm25, Xbwm21, Xbwm20
|
KM2939 |
A. cristatum
|
|
Pm2c
|
SSR |
Xcfd81 - Xcfd78
|
Niaomai |
T. aestivum
|
|
Pm3a
|
5AS |
RFLP |
Xwhs179
|
Asosan |
T. aestivum
|
|
Pm3b
|
RFLP |
Xbcd1434
|
Chul |
T. aestivum
|
|
Pm3c
|
|
|
Sonora |
T. aestivum
|
|
Pm3d
|
|
|
Kolibri |
T. aestivum
|
|
Pm3e
|
|
|
W150 |
T. aestivum
|
|
Pm3f
|
|
|
Michigan Amber |
T. aestivum
|
|
Pm3g
|
RFLP |
Gli-A5
|
Aristide |
T. aestivum
|
|
Pm3h
|
|
|
Abessi |
T. durum
|
|
Pm3i
|
|
|
N324 |
T. aestivum
|
|
Pm3j
|
|
|
GUS 122 |
T. aestivum
|
|
Pm3k
|
|
|
IG46439 |
T. dicoccoides
|
|
Pm4a
|
2AL |
RFLP, AFLP, STS |
Xbcd1231, Xcdo678, 4aM1
STSbcd1231
|
Khapli |
T. dicoccum
|
|
Pm4b
|
|
|
Armada |
T. carthlicum
|
|
Pm4c
(Pm23)
|
|
|
81-7241 |
T. aestivum
|
|
Pm4d
|
SRS |
Xbarc122, Xgwm526
|
Tm27d2 |
T. monococcum
|
|
Pm4e
|
SRS |
Xwgrc763, Xwgrc982
|
D29 |
T. aestivum
|
|
Pm5
|
7BL |
|
|
Xiaobaidong |
T. aestivum
|
|
Pm5a
|
|
|
Hope |
T. dicoccum
|
|
Pm5b
|
|
|
Ibis |
T. aestivum
|
|
Pm5c
|
|
|
Kolandi |
T. aestivum spp. sphaerococcum |
|
Pm5d
|
|
|
IGV 1-455 |
T. aestivum
|
|
Pm5e
|
SSR |
Xgwm1267-136
|
Fuzhuang 30 |
T. aestivum
|
|
Pm6
|
2BL |
RFLP |
Xbcd135
|
TP 114 |
T. timopheevii
|
|
Pm7
|
4BS.4BL-2RL |
|
|
Transec |
S. cereal
|
|
Pm8
|
1RS.1BL |
RFLP, RAPD, STS |
IAG95, OPJ07-1200, OPR19-1350
SEC-1b, STSiag95
|
Disponent |
S. cereal
|
|
Pm9
|
7AL |
|
|
N14 |
T. aestivum
|
|
Pm10
|
1D |
|
|
Norin 26 |
T. aestivum
|
|
Pm11
|
6BS |
|
|
Chinese Spring |
T. aestivum
|
|
Pm12
|
6BS-6SS.6SL |
RFLP |
Xpsr10, Xpsr106, Xnor-2, Xpsr141, Xpsr113, Xpsr142, Xpsr149, Xpsr2
|
Trans. Line 31 |
Ae. speltoides
|
|
Pm13
|
3BL.3SS-3S
3DL.3SS-3S |
RFLP
RFLP, RAPD, STS |
Xpsr305, Xpsr1196
Xcdo460, Xutv135, OPV13800,
UTV13, OPX12570, UTV14
|
C strans. Line |
Ae. longissima
|
|
Pm14
|
6BS |
|
|
Norin 10 |
T. aestivum
|
|
Pm15
|
7DS |
|
|
Norin 26 |
T. aestivum
|
|
Pm16
|
4A |
|
|
Norman rec. line |
T. dicoccoides
|
|
Gene
|
Location |
Type of marker |
Closest/flanking marker
|
Cultivar/line |
Source
|
|
Pm17
|
1RS.1AL |
RFLP, AFLP |
IAG95-CA/CT-355
|
Amigo |
S. cereal
|
|
Pm19
|
7D |
|
|
XX 186 |
Ae. tauschii
|
|
Pm20
|
6BS.6RL |
|
|
KS93WGRC28 |
S. cereal
|
|
Pm21
(Pm31)
|
6VS.6AL |
RAPD, SCAR |
OPH171900, OPH171400, SCAR1265, SCAR1400
|
Yangmai 5 line |
Haynaldia villosa
|
|
Pm23
(Pm4c)
|
2AL |
|
|
82-7241 |
T. aestivum
|
|
Pm24a
|
1DS |
AFLP, SSR |
E34/M51, Xgwm337, Xgwm1291
|
Chiyacao |
T. aestivum
|
|
Pm24b
(mlbhl)
|
SSR |
Xgwm603/Xgwm789, Xbarc229
|
Baihulu |
T. aestivum
|
|
Pm25
|
1A |
RAPD |
OPA04950
|
NC96BGTA5 |
T. boeoticum
|
|
Pm26
|
2BS |
RFLP |
wg516
|
TTD140 |
T. dicoccoides
|
|
Pm27
|
6B-6G |
RFLP, SSR |
psp3131
|
146-155-T |
T. timopheevii
|
|
Pm28
|
1B |
|
|
Meri |
T. aestivum
|
|
Pm29
|
7DL |
RFLP, AFLP |
S24M13, S19M23, S22M26, S25M15, S13M23, S22M21, S17M25 |
Pova |
A. ovate
|
|
Pm30
|
5BS |
SSR |
Xpsp3029
|
C20 |
T. dicoccoides
|
|
Pm31
(Pm21)
|
6AL |
SSR |
BJ261635
|
G-305-M/781 |
T. dicoccoides
|
|
Pm32
|
1BL.1SS |
|
|
L501 |
Ae. speltoides
|
|
Pm33
|
2BL |
|
|
PS5 |
T. carthlicum
|
|
Pm34
|
5DL |
|
|
NC97BGTD7 |
Ae. tauschii
|
|
Pm35
|
5DL |
|
|
NC96BGTD3 |
Ae. tauschii
|
|
Pm36
|
5BL |
SSR |
Xgwm297
|
MG29896 |
T. dicoccoides
|
|
Pm37
|
7AL |
|
|
NC99BGTAG11 |
T. timopheevii
|
|
Pm38
|
7DS |
|
|
RL6058 |
T. aestivum
|
|
Pm39
|
1BL |
|
|
Saar |
T. aestivum
|
|
Pm40
|
7BS |
|
|
GRY19 |
E. intermedium
|
|
Pm41
|
3BL |
SSR, ISBP, STS |
BE489472
|
IW2 |
T. dicoccoides
|
|
Pm42
|
2BS |
SSR, AFLP-SCAR, EST-STS, RFLP-STS |
BF146221
|
G-303-1M |
T. dicoccoides
|
|
Pm43
|
2DL |
SSR |
Xwmc41
|
CH5025 |
T. intermedium
|
|
Pm44
|
3AS |
|
|
Hombar |
T. aestivum
|
|
Pm45
|
6DS |
SSR, STS |
Xmag6176
|
D57 |
T. aestivum
|
|
Pm46
|
5DS |
SSR |
Xgwm205, Xcf81
|
Tabasco |
T. aestivum
|
|
Pm47
|
7BS |
SSR, EST |
Xgwm46, BE606897
|
Hongyanglazi |
T. aestivum
|
|
Pm49
(MI533)
|
2BS |
EST-SSR |
CA695634
|
MG5323 |
T. dicoccum
|
|
Pm50
|
2AL |
SSR |
Xgwm294
|
K2 |
T. dicoccum
|
|
Pm51
|
2BL |
SSR |
Xwmc332 - Xwmc317
|
CH7086 |
Th. ponticum
|
|
Pm52
(MlLX99)
|
2BL |
SSR |
Xcfd73, Xwmc441, XBE604758, Xgwm120
|
Liangxing 99 |
T. aestivum
|
|
Pm53
|
5BL |
SSR |
Xwmc759, Xgwm499, IWA6024, IWA2454, Xgwm408
|
NC09BGTS16 |
Ae. Speltoides
|
|
Pm54
|
6BL |
SSR, SNP |
Xgpw2344, wPt-9256, Xbarc134
|
AGS 2000 |
T. aestivum
|
|
Pm55
|
5AL/5DL |
EST |
5EST-237 |
NAU421 |
D. villosum
|
|
Pm56
|
6AS |
|
|
LM47-6 |
S.cereale |
|
Pm57
|
2BL |
RFLP |
X2L4g9p4/HaeIII |
Line 89-346(TA5108)/
Line 89(5)69 (TA5109) |
Ae. searsi
|
|
Pm58
|
2DS |
KASP |
K-TP338253-K-TP159900 |
TA1662 |
Ae. tauschii
|
|
Pm59
|
7AL |
SSR |
Xmag1759 - Xmag1714
|
PI 181356 |
T. aestivum
|
|
Pm60
|
7AL |
SSR |
Xwmc273.3
|
PI 428309 |
T. urartu
|
|
Pm61
|
4AL |
SSR |
Xgwm160-Xicsx79
|
Xuxusanyuehuang |
T. aestivum
|
|
Pm62
|
T2BS.2VL#5 |
|
|
NAU1823 |
D. villosum
|
Table 2Summary of quantitative trait loci (QTLs) for adult-plant resistance to powdery mildew in wheat (
Triticum aestivum L.) modified from
Li et al. (2014) and
Kang et al. (2020).
Table 2
|
QTLs |
Chromosome |
Marker interval |
R2z
|
|
QPm.caas-1AL
|
1AL |
Xbarc148-Xwmc550
|
7.4-9.9% |
|
QPm.crag-1A
|
1AL |
Xcdo572-Xbad442
|
39.3-43.0% |
|
QPm.sfr-1A
|
1AL |
Xpsr1201b-Xpsr941
|
7.7% |
|
QPm.caas-1AS
|
1AS |
Xgdm33-Xpsp2999
|
19.9-26.6% |
|
QPm.osu-1A
|
1AS |
Pm3a
|
63.0% |
|
Qaprpm.cgb-1B
|
1B |
WMC269.2-CWM90
|
4.8-20.3% |
|
Qaprpm.cgb-1B
|
1B |
P4133-170-Xgwm582
|
4.8% |
|
QPm.caas-1BL.1
|
1BL |
IWB72835-IWB18787
|
7.2% |
|
QPm.osu-1B
|
1BL |
WMC134
|
14.0% |
|
QPm.vt-1B
|
1BL |
WG241
|
17.0% |
|
QPm.vt-1BL
|
1BL |
Xgwm259-Xbarc80
|
15.0%-17.0% |
|
QPm.sfr-1B
|
1BS |
CD9b-Xpsr593a
|
11.6% |
|
QPm.ttu-1B
|
1BS |
Xgwm3000
|
4.0-5.0% |
|
QPm.heau-1DL
|
1DL |
wPt-5721-wPt-1865
|
6.1-8.5% |
|
QPm.sfr-1D
|
1DL |
Xpsr168-Xglk558b
|
9.5% |
|
QPm.inra-1D.1
|
1DS |
Xgwm106
|
12.6% |
|
Qpm.sdau-2A
|
2A |
cfa2263-D-1395795
|
8.6-9% |
|
QPm.crag-2A
|
2AL |
Pm4b-gbxG303
|
22.7-39.2% |
|
QPm.ttu-2A
|
2AL |
Xgwm311-Xgwm382
|
5.0% |
|
QPm.vt-2A
|
2AL |
Xgwm304-Xgwm294
|
26.0-29.0% |
|
QPm.vt-2AL
|
2AL |
Xgwm304a-Xgwm312
|
29.0% |
|
QPm.inra-2A
|
2AS |
Xgwm275
|
7.4% |
|
QPm.sfr-2A
|
2AS |
Xpsr380-Xglk293b
|
7.7% |
|
Qaprpm.cgb-2B
|
2B |
Xwmc477-Xwmc272
|
5.4% |
|
QPm.sdau-2B
|
2B |
D-1054006—D-1114401 |
12.6-13.4% |
|
QPm.caas-2B
|
2BL |
Xgwm877.1-Xgwm435.1
|
5.7-8.0% |
|
QPm.caas-2BL
|
2BL |
Xbarc1139-Xgwm47
|
5.2-10.1% |
|
QPm.inra-2B
|
2BL |
Xrtp114R-Xcfd267b
|
10.3-36.3% |
|
QPm.vt-2B
|
2BL |
WG338-Xgwm526a
|
11.0% |
|
QPm.vt-2BL
|
2BL |
Xgwm501-Xgwm191
|
11.0-15.0% |
|
QPm.caas-2BS
|
2BS |
Xbarc98-Xbarc1147
|
10.6-20.6% |
|
QPm.crag-2B
|
2BS |
Xgwm148-gbxG553
|
23.6-71.5% |
|
QPm.umb-2BS
|
2BS |
wPt-9402
|
3.9-13.0% |
|
QPm.umb-2BS
|
2BS |
Xgwm410b-Xgwm148
|
8.0-10.2% |
|
QPm.caas-2DL
|
2DL |
Xwmc18-Xcfd233
|
5.7-11.6% |
|
QPm.ipk-2D
|
2DL |
Xglk558-XksuD23
|
unknown |
|
QPm.sfr-2D
|
2DL |
Xpsr932-Xpsr331a
|
10.0% |
|
QPm.umb-2DL
|
2DL |
Xwmc167-Xgwm301
|
4.3-9.5% |
|
QPm.caas-2DS
|
2DS |
Xcfd51-Xcfd56
|
2.3-3.4% |
|
QPm.inra-2D-a
|
2DS |
Xgwm102
|
19.0% |
|
QPm.inra-2D-b
|
2DS |
Xcfd2e
|
16.5% |
|
Qaprpm.cgb-3A
|
3A |
Xwmc21-Xwmc505.2
|
9.8% |
|
QPm.crag-3A
|
3AS |
Xpsr598-Xgwm5
|
21.4-25.9% |
|
QPm.nuls-3AS
|
3AS |
Xstm844tcac-Xbarc310
|
8.1-20.7% |
|
QPm.sfr-3A
|
3AS |
Xpsr598-Xpsr570
|
10.4% |
|
Qaprpm.cgb-3BL
|
3B |
Xgwm181-Xgwm340
|
13.3% |
|
QPm.caas-3B
|
3BS |
XksuG53-Xfba190
|
7.3% |
|
QPm.caas-3BS
|
3BS |
IWB21064-IWB64002
|
7.1% |
|
QPm.inra-3B
|
3BS |
Xgwm389
|
22.7% |
|
QPm.osu-3B
|
3BS |
WMS533
|
10.0% |
|
QPm.inra-3D
|
3DS |
Xcfd152, Xgwm707
|
9.3-15.2% |
|
QPm.sfr-3D
|
3DS |
Xpsr1196a-Lrk10-6
|
15.7% |
|
QPm.tut-4A
|
4A |
Xwmc232-Xrga3.1
|
24.0-46.0% |
|
QPm.crag-4A.
|
4AL |
XgbxG036-XgbxG542
|
22.3% |
|
QPm.inra-4A
|
4AL |
XgbxG036
|
4.9-6.9% |
|
QPm.osu-4A
|
4AL |
WMS160
|
12.0% |
|
QPm.sfr-4A.1
|
4AL |
Xgwm111c-Xpsr934a
|
14.7% |
|
QPm.sfr-4A.2
|
4AL |
Xmwg710b-Xglk128
|
14.3% |
|
QPm.ttu-4A
|
4AL |
Xwmc232-Xgwm160
|
35.0-54.0% |
|
QPm.caas-4BL
|
4BL |
Xgwm375-Xgwm251
|
5.9% |
|
QPm.Caas-4BL.1
|
4BL |
Xgwm149-Xgwm495
|
9.1-14.7% |
|
QPm.caas-4BL.2
|
4BL |
IWB35851-IWB60096
|
8.8% |
|
QPm.ipk-4B
|
4BL |
Xcdo795-Xbcd1262
|
Unknown |
|
QPm.nuls-4BL
|
4BL |
XwPt1505-Xgwm149
|
21.0-40.2% |
|
QPm.sfr-4B
|
4BL |
Xpsr593b-Xpsr1112
|
7.5% |
|
QPm.caas-4DL
|
4DL |
Xbarc200-Xwmc33
|
15.2-22.7% |
|
QPm.sfr-4D
|
4DL |
Xglk302b-Xpsr1101a
|
14.4% |
|
Qaprpm.cgb-5A
|
5A |
P3616-185-P3616-195
|
13.2% |
|
QPm.tut-5A
|
5A |
Xgwm666-Xcfd30-Xbarc319
|
14.0-16.0% |
|
QPm.nau-5AL
|
5AL |
Xcfd39/Xmag1491-Xmag1493
|
59.0% |
|
QPm.nuls-5A
|
5AL |
Xgwm617b-Xwmc327
|
4.2-15.2% |
|
QPm.sfr-5A.2
|
5AL |
Xpsr1194-Xpsr918b
|
16.6% |
|
QPm.sfr-5A.3
|
5AL |
Xpsr911-Xpsr120a
|
10.5% |
|
QPm.umb-5AL
|
5AL |
wPt-2426
|
4.0-9.7% |
|
QPm.sfr-5A.1
|
5AS |
Xpsr644a-Xpsr945a
|
22.9% |
|
QPm.ttu-5A
|
5AS |
Xgwm186-Xgwm415
|
4.0-6.0% |
|
Qaprpm.cgb-5B
|
5B |
Xgwm213-Xgwm499
|
19.8% |
|
QPm.inra-5B.2
|
5BL |
Xgwm790b
|
11.1% |
|
QPm.sfr-5B
|
5BL |
Xpsr580b-Xpsr143
|
12.6% |
|
QPm.nuls-5B
|
5BS |
Xbarc4-Xgwm274b
|
9.7% |
|
QPm.ttu-5B
|
5BS |
Xgwm133.mi6-Xgwm205.mi1
|
4.0-6.0% |
|
QPm.umb-5BS
|
5BS |
wPt-1261
|
3.1% |
|
QPm.umb-5BS
|
5BS |
Xbarc128a-Xgwm213
|
8.1-12.9% |
|
QPm.caas-5D
|
5D |
Xmwg922-Xbcd1103
|
5.9% |
|
QPm.crag-5D.1
|
5DL |
Xgwm639a-Xgwm174
|
30.2-38.9% |
|
QPm.crag-5D.2
|
5DL |
Xcfd8B9-Xcfd4A6
|
24.0-37.8% |
|
QPm.inra-5D.1
|
5DL |
Xcfd26
|
28.1-37.7% |
|
QPm.inra-5D.2
|
5DL |
XgbxG083c
|
37.7% |
|
QPmVpn.inra-5D
|
5DL |
Xcfd8
|
11.0% |
|
QPm.inra-5D
|
5DS |
cfd189
|
9.0% |
|
QPm.crag-6A
|
6AL |
MlRE
|
19.8-53.9% |
|
QPm.inra-6A
|
6AL |
MIRE(Xgwm427)
|
8.8-13.4% |
|
Qaprpm.cgb-6B
|
6B |
Xgwm193-P3470-210
|
21.0% |
|
QPm.caas-6BL.1
|
6BL |
Xgwm219-Xbarc24
|
2.5-5.2% |
|
QPm.caas-6BL.2
|
6BL |
Xbarc24-Xbarc345
|
0.5-1.9% |
|
QPm.caas-6BS
|
6BS |
Xbarc79-Xgwm518
|
10.3-16.0% |
|
QPm.sfr-6B
|
6BS |
Xpsr167b-Xpsr964
|
8.7% |
|
QPm.umb-6BS
|
6BS |
wPt-6437-Xwmc494
|
6.5-10.3% |
|
QPm.osu-6D
|
6DS |
BARC196
|
5.0% |
|
Qaprpm.cgb-7A
|
7A |
CWM462.2-Xgwm635.2
|
8.0% |
|
Qaprpm.cgb-7A
|
7A |
Xgwm282-P1111-202
|
15.7% |
|
QPm.tut-7A
|
7A |
Xgwm635-Xbarc70-Waxy
|
9.0-28.0% |
|
QPm.umb-7AL
|
7AL |
Xgwm428-Xcfa2040
|
6.4-13% |
|
QPm.caas-7A
|
7AS |
Xbarc127-Xbarc174
|
6.3-7.1% |
|
QPm.inra-7A
|
7AS |
Xfba069-Xgwm344
|
2.9-6.4% |
|
Qaprpm.cgb-7B
|
7B |
Xwmc273-Xwmc276
|
12.6% |
|
QPm.crag-7B
|
7BL |
XpdaC01-XgbxR035b
|
22.8-33.5% |
|
QPm.inra-7B
|
7BL |
Xgwm577
|
1.7% |
|
QPm.nuls-7BL
|
7BL |
Xwmc581-XwPt8007
|
4.90% |
|
QPm.sfr-7B.1
|
7BL |
Xpsr593c-Xpsr129c
|
11.3% |
|
QPm.sfr-7B.2
|
7BL |
Xglk750-Xmwg710a
|
31.8% |
|
Qaprpm.cgb-7D
|
7D |
Xwmc436-Xgwm44
|
3.8-4.6% |
|
Qaprpm.cgb-7D
|
7D |
Xgdm88-WMC463
|
14.20% |
|
QPm.caas-7D
|
7D |
Xwg834-Xbcd1438
|
29.6% |
|
QPm.caas-7DS
|
7DS |
XcsLV34-Xgwm295
|
7.6-13.8% |
|
QPm.caas-7DS
|
7DS |
IWB41108-IWB53819
|
4.21% |
|
QPm.caas-7DS
|
7DS |
Ltn-Xgwm295.1
|
12.0% |
|
QPm.inra-7D.1
|
7DS |
Xgpw1106
|
10.6% |
|
QPm.ipk-7D
|
7DS |
Xwg834-Xbcd1872
|
Unknown |
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