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밀 줄녹병 저항성 유전 육종 주요 연구 동향

김경민1, 조성우2, 강천식1, 김경훈1, 최창현1, 손재한1, 박철수3, 모영준1,*

Research Advances in Wheat Breeding and Genetics for Stripe Rust Resistance

Kyeong-Min Kim1, Seong-Woo Cho2, Chon-Sik Kang1, Kyeong-Hoon Kim1, Chang-Hyun Choi1, Jae-Han Son1, Chul Soo Park3, Youngjun Mo1,*
Korean Journal of Breeding Science 2020;52(2):93-103.
Published online: June 1, 2020

1농촌진흥청 국립식량과학원

2경남과학기술대학교 농학한약자원학부

3전북대학교 작물생명과학과

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

2Department of Agronomy and Medicinal Plant Resources, Gyeongnam National University of Science and Technology, Jinju 575, Republic of Korea

3Department of Crop Science and Biotechnology, Chonbuk National University, Jeonju 54896, Republic of Korea

* Corresponding Author (E-mail: moyj82@korea.kr, Tel: +82-63-238-5232, Fax:+82-63-238-5205)
• Received: May 6, 2020   • Revised: May 6, 2020   • Accepted: May 12, 2020

Copyright © 2020 by the Korean Society of Breeding Science

This is an open-access article distributed under the terms of the Creative Commons Attribution Non-Commercial License (http://creativecommons.org/licenses/by-nc/3.0) which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original work is properly cited.

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  • Stripe rust (or yellow rust) caused by Puccinia striiformis f. sp. tritici is the major wheat disease responsible for deteriorating global wheat yield and quality. Although stripe rust appears to be prevalent mainly in temperate areas, new races adapting to high temperature have recently appeared in warmer areas, such as Australia, the Middle East, and Africa, increasing the threat to global food security. Among the approximately 80 reported genes for stripe rust resistance, six (Yr5, Yr7, Yr15, Yr18, Yr36, Yr46) have been cloned. Stripe rust resistance genes are generally classified into race-specific (or all-stage) and non-race-specific (or high temperature adult plant) resistance genes. While resistance conferred by most race-specific genes are overcome by the appearance of new pathogen races within few years, non-race-specific genes, such as Yr18, Yr36, and Yr46, provide more durable resistance and are often stable for several decades. Yr18/Lr34/Sr57/Pm38 and Yr46/Lr67/Sr55/Pm46 are especially useful in breeding as they confer resistance to leaf rust, stem rust, and powdery mildew as well as stripe rust. Unlike the extensive global research efforts, few studies have been conducted in Korea regarding breeding and genetics for stripe rust resistance. To prevent damage by stripe rust in advance, it is important to monitor the changes in major pathogen races in Korea, evaluate major wheat breeding lines and landraces for stripe rust resistance by establishing an efficient screening system, and introduce new germplasm with various resistance genes. Reinforcing wheat molecular genetics and genomics capacity is also important to enable identification of new stripe rust resistance genes and efficient transfer of the novel genes into elite wheat cultivars using molecular markers.
  • Keywords: wheat; stripe rust; yellow rust; resistance; breeding
녹병은 전세계적으로 밀의 수량과 품질을 위협하는 대표적인 질병으로 잎녹병(leaf rust), 줄기녹병(stem rust 또는 black rust), 줄녹병(stripe rust 또는 yellow rust)으로 구분된다(Cummins & Hiratsuka 2004, Duplessis et al. 2011, Figueroa et al. 2018). 밀 줄녹병균인 Puccinia striiformis Westend. f. sp. tritici (Pst)는 서늘하고 습한 온대 지역에서 흔히 발생해서 지난 50년 동안 60개국 이상에서 보고되었지만, 2000년 이후에는 고온환경에 적응한 전염력이 강한 Pst 병원균 발생으로 점차 발생 지역과 빈도가 증가하는 추세이다(Chen 2005, Ali et al. 2014, Beddow et al. 2015). 유럽, 호주 및 북미에서 발생하는 Pst는 대부분 다양성이 낮으나 중국 서부, 중앙아시아, 히말라야 근처에서 발병하는 Pst는 유전적 다양성이 상대적으로 높다고 보고되었다(Hovmoller et al. 2011, Ali et al. 2014, Chen et al. 2014). 2011~2013년과 2015년에 히말라야 근처에서 기원하는 Pst 병원균의 변이체는 유럽 전역에 발병하여 큰 피해를 초래하였다(Hovmoller et al. 2015, Hubbard et al. 2015).
Pst 병원균의 발병지역은 크게 10개의 역학 지역(epidemiological zones)으로 구별되는데, 최대 발병지역은 80% 이상의 수량 감소가 일어나고, 발병이 적은 지역은 5~10% 정도 수량이 감소하며, 평균적으로는 35% 이상의 수량 감소가 발생한다(Ali et al. 2014). 전세계 밀 품종의 88%는 Pst에 저항성이 없기 때문에 이로 인해 매년 약 10억 달러 정도의 경제적 손실이 일어나는데, 호주에서만 약 9천만 달러의 손실이 보고되고 있다(Murray & Brennan 2009, Beddow et al. 2015, Wellings 2011, Chen et al. 2014).
녹병균을 제어하는 방법은 화학적인 방법과 유전적인 방법이 있다. 병원균 매개체가 될 수 있는 녹색교량(green bridges)를 제거하는 경작적인 방법이 우선적인 방법이 될 수 있고, 다음으로는 살균제를 이용한 화학적 방법인데, 현재 승인된 녹병 살균제로는 quinone outside inhibitors (QoIs), 14α-demethylation inhibitors (DMIs), succinate dehydrogenase inhibitors (SDHI) 등이 이용되고 있다(Oliver 2014). 그러나 많은 국가에서 적절한 살균제를 찾기가 쉽지 않고 부적절한 사용으로 인하여 인체와 환경에 심각한 피해가 발생하기도 하여 이로 인한 비용 증가가 문제점으로 대두되고 있다(Kolmer et al. 2007, Arduim et al. 2012, Kumar et al. 2013, Oliver 2014). 많은 지역에서는 녹병 방제를 위해서 초기 생육시기의 살균제를 통한 방제와 지속적인 포장 관리와 모니터링을 기본으로 하는 병해충 종합관리(integrated pest management, IPM)가 실질적인 방안으로 제시되고 있다(Jorgensen et al. 2008).
보다 효과적이고 경제적이면서 환경친화적인 방법은 저항성 품종 개발이지만 이를 위해서는 시간과 노력이 많이 요구된다(Singh et al. 2000, Todorovska et al. 2009, Oliver 2014). 줄녹병은 현재까지 80여개의 저항성 유전자가 알려져 있는데(McIntosh et al. 1995, 2014, Chen 2005, Dakouri et al. 2013, Gessese et al. 2019), 이를 이용하여 저항성 품종을 효과적으로 육성하기 위해서는 목표 지역의 우점 균계를 파악하고 해당 균계에 효과적인 저항성 유전자를 우량 품종에 신속히 이전시켜야 하며, 다수의 유전자를 한 품종에 집적시킴으로써 균계 변화에도 대응해야 한다(Periyannan et al. 2017, Mundt 2018). 본 논문은 국내 환경에 적합한 줄녹병 저항성 밀 품종 육성에 필요한 정보를 정리하였다.
줄녹병 생활사
녹병균은 숙주식물에 기생하여 성장과 번식을 하는데, 밀에서는 여름포자(urediniospores), 겨울포자(teliospores), 담자포자(basidiospores)를 형성하고 밀이 아닌 다른 대체숙주에서는 병포자(pycniospores)와 녹포자(aeciospores)를 형성한다(Cummins & Hiratsuka 2004, Bolton et al. 2008, McIntosh 2009, Jin et al. 2010, Duplessis et al. 2011). 줄녹병은 P. striiformis f. sp. tritici (Pst)에 의하여 발생하며 생활사는 Fig. 1과 같다(Zheng et al. 2013). 매자나무속(barberry)인 Mahonia spp.와 Berberis spp.이 줄녹병의 대체숙주로 알려져 있지만 잎녹병과는 달리 자연상태에서는 이들이 밀에 녹병을 퍼뜨리는 역할을 하지는 않는다(Chen et al. 2014).
줄녹병의 감염은 4°C에서 시작되어 7°C가 적정온도이며 잎녹병이나 줄기녹병과 달리 밀에 침투하기 위한 부착기(appressoria)를 형성하지 않고, 생식관이 잎의 기공을 직접 관통하여 흡입 모세포(haustorial mother cells)를 형성한 후에 긴 균사(hyphae)가 잎의 길이 방향으로 자라게 되어 잎에 노랑색 내지 주황색의 포자(uredospore pustules)가 직선의 줄무늬 모양으로 나타나게 된다(Moldenhauer et al. 2006, Chen et al. 2014, Fig. 2). 줄녹병 감염은 건물중, 뿌리 생장, 간장, 이삭 및 종실의 개수와 크기에 영향을 주며, 이러한 영향은 유묘기 감염이 가장 크지만 개화⋅수정기에 시작된 감염도 수량에 영향을 준다(Wellings 2011).
줄녹병 저항성 기작
줄녹병 저항성은 두 가지 형태가 존재한다. 첫번째 형태는 전단계 저항성(ASR, all-stage resistance)이라고 하는데, 이들은 잎녹병의 유묘기 저항성(SR, seedling resistance) 유전자처럼 생육 초기에 저항성을 나타내며, 같은 병원균에 대해서는 성체에서도 저항성을 유지하기 때문에 줄녹병에서는 ASR로 구분한다. 줄녹병의 ASR은 잎녹병의 유묘기 저항성과 마찬가지로 주동유전자(major gene)가 주로 관여한다(Chen & Line 1995a, 1995b).
두번째 형태는 고온성체저항성(HTAP, high temperature adult plant resistance)으로 구분되는데, HTAP는 생육 후기의 온도가 21°C 이상이 되면서 병원균이 쉽게 퍼지는 시기에 저항성을 보인다(Qayoum & Line 1985). 줄녹병의 HTAP 저항성 유전자는 유묘기에는 감수성을 나타내지만, 잎녹병의 성체 저항성(APR, adult plant resistance)처럼 성체가 되면 줄녹병에 대한 저항성을 나타낸다(Chen & Line 1995a, 1995b, Line & Chen 1995). 이러한 형태의 저항성 유전자로 대표되는 것들은 Yr18, Yr36과 Yr46이 있지만 이들의 정확한 저항성 기작에 대해서는 알려진 것이 적다(Ellis et al. 2014). 유전자 동정을 통하여 Yr36이 WKS (Wheat Kinase-START) 단백질을 코딩하는 것이 밝혀졌고, Yr18과 Yr46이 각각 ABC (ATP-binding cassette) transporter와 hexose transporter를 만든다는 정도만 밝혀졌다(Krattinger et al. 2009, Moore et al. 2015, Dodds & Lagudah 2016).
줄녹병 저항성 유전자
줄녹병 저항성 유전자는 Yr로 표기하며 주요 저항성 유전자의 정보는 Table 1에 정리하였다. 줄녹병균인 Pst의 높은 다양성과 빠른 변이로 인하여 새로운 병원균이 지속적으로 발생하는데, 이로 인해 기존 유전자가 무력화 되는 경우가 많이 발생한다. 1990년대에 Yr2와 Yr9 유전자가 무력화 되었고, Yr17과 Yr27이 각각 유럽과 인도에서 무력화 되면서 줄녹병이 대발생하였고, 2011년에도 유럽에서 병원성이 강한 새로운 균계의 발생으로 큰 피해가 초래되었다(Bahri et al. 2011, Hovmoller et al. 2015). 그러므로 균계 분화에 대응하기 위하여 종특이적 유전자와 종비특이적 유전자의 다양한 조합으로 줄녹병 저항성을 유지하는 것이 효과적이다(Singh et al. 2000, Herrera-Foessel et al. 2011). 예를 들어 종특이적 유전자인 Yr5이나 Yr15에 종비특이적 유전자인 Yr18, Yr29과 Yr46를 조합하는 것이 줄녹병 저항성 증진에 효과적임이 보고된 바 있다(Chen et al. 2014).
대부분의 줄녹병 저항성 유전자는 ASR에 속하며 주로 육배체 일반밀에서 유래하지만, 호밀(Yr9), 야생밀(Yr5, Yr8, Yr15, Yr24, Yr26, Yr28, Yr35, Yr42)에서 유래하기도 한다(Table 1). ASR의 경우 NBS-LRR 형태의 저항성 유전자에 의해 주로 조절될 것으로 추정되는 반면 HTAP 저항성은 보다 다양한 유전자나 환경인자에 영향을 받을 것으로 예측되고 있다(Hulbert et al. 2007, Chen 2013).
줄녹병의 HTAP 저항성 유전자는 대부분 다른 녹병이나 흰가루병 저항성과 연관이 되어있는데, Yr17 (Lr37/Sr38), Yr18 (Lr34/Sr57/Pm38), Yr29 (Lr46/Sr55/Pm46), Yr30 (Lr27/Sr2), Yr46 (Lr67/Sr55/Pm46)과 Yr47 (Lr52)이 그 예로, Yr18, Yr29과 Yr36는 미국에서 HTAP 저항성 품종 육성에 많이 사용되고 있다(Table 1). 특히 6BS 염색체에 위치한 Yr36은 종실 단백질 함량에 영향하는 Gpc-B1 유전자에 근접해 있어 빵 품질향상을 위한 분자표지와 병행하여 이용되고 있다(Chen 2013). 위에서 언급한 Yr5, Yr15, Yr17, Yr18, Yr29, Yr36, Yr46를 비롯하여 주요 줄녹병 저항성 유전자의 도입을 위한 분자표지 관련 정보는 MAS Wheat 홈페이지(https://maswheat.ucdavis.edu/)에 잘 정리되어 있으므로 육종사업에 유용하게 활용할 수 있다.
줄녹병은 온대지방에서 주로 발생하지만 최근 광범위한 기후에 적응하는 균계가 발생하면서 전세계적으로 밀의 생산성과 품질에 큰 위협이 되고 있다. 줄녹병 저항성 유전자는 약 80여개가 보고되었고 동정이 이루어진 것은 6개로 Yr5, Yr7, Yr15, Yr18/Lr34/Sr57/Pm38, Yr36, Yr46/Lr67/Sr55/Pm46이다(Table 1). 새로운 줄녹병 저항성 유전자원을 발굴하고 다수의 저항성 유전자를 우수 품종에 집적시키기 위한 연구가 국제적으로 활발하게 이루어지고 있으나 우리나라의 줄녹병 저항성 유전육종 연구는 아직 초기단계이다. 국내 육성 밀 계통인 수원11호와 수원92호가 줄녹병 저항성 유전자인 YrSu, YrSo를 각각 보유하였다는 외국의 보고 사례가 있었으나(Wan et al. 2004, Mamun et al. 2018) 이들 계통이 줄녹병 저항성 품종 개발을 위하여 국내에서 활용된 사례는 찾아보기 어렵다.
줄녹병은 저항성이 없는 품종에서 발병하는 경우 적게는 5~10%에서 많게는 80% 이상 밀 수량을 감소시키며 지속적인 균계 분화로 병원성 양상과 환경적응성이 빠르게 변화하므로 우리나라도 저항성 품종 육성을 통하여 발병피해를 선제적으로 예방해야 한다. 이를 위해서는 대량⋅정밀 검정체계를 구축하여 주요 줄녹병 균계에 대한 국내 밀 품종 및 육성계통, 재래종의 저항성을 판정해야 하며, 다양한 저항성 유전자를 보유한 유전자원을 적극적으로 도입하여 육종에 활용해야 한다.
보다 장기적으로는 밀 분자유전학 및 유전체 연구 강화를 통하여 국내에서도 새로운 녹병 저항성 유전자를 자체적으로 발굴해야 한다. 밀은 배수체이며 유전체의 크기가 벼의 40배 이상으로 매우 방대하여 유전연구가 까다로운 편이다. 그러나 최근 밀 표준유전체(IWGSC 2018)가 확립됨에 따라 전통적인 유전지도기반 유전자분리(map-based cloning) 이외에도 다양한 유전분석 기술이 활용되고 있다. 목표 염색체만 선별(flow cytometry)하여 차세대염기서열분석(NGS)을 진행하는 MutChromSeq (Sanchez-Martin et al. 2016), 병 저항성 관련 부위만 선별하는 RenSeq (Steuernagel et al. 2016), 유전자 부위를 선별하는 ExomeSeq (Mo et al. 2018), 전유전체연관분석(GWAS)을 병 저항성 관련 부위에 집중하여 진행하는 AgRenSeq (Arora et al. 2019) 등이 그 사례이다. 따라서 신규 유전자원 탐색과 함께 유전체 기반 기술을 활용한 유전분석을 병행한다면 국내 고유의 녹병 저항성 유전자 및 분자표지를 개발하고 이를 녹병 저항성 밀 육종에 활용하는 것도 가능해질 것으로 기대한다.
Puccinia striiformis f. sp. tritici에 의하여 발생하는 줄녹병은 잎녹병, 줄기녹병과 함께 밀의 생산성과 품질을 떨어뜨리는 주요 질병이다. 줄녹병은 과거에 온대지역에 주로 발생하여 잎녹병에 비하여 피해가 적은 것으로 인식되어 왔으나, 최근 균계 분화로 호주, 중동, 아프리카를 포함한 고온지역으로도 피해가 확대되면서 위협이 커지고 있다. 줄녹병 저항성 유전자는 현재까지 80여개가 보고되었고 그 중 6개(Yr5, Yr7, Yr15, Yr18, Yr36, Yr46)가 동정되었다. 줄녹병 저항성은 크게 종특이적 저항성(전단계저항성)과 종비특이적 저항성(고온성체저항성)으로 구분되는데, 종특이적 저항성 유전자는 균계 분화로 인하여 저항성이 수년 안에 쉽게 무너지는 반면 Yr18, Yr36, Yr46을 비롯한 종비특이적 저항성 유전자는 저항성이 수십년 이상 유지되기도 한다. 특히 Yr18/Lr34/Sr57/Pm38과 Yr46/Lr67/Sr55/Pm46 유전자는 줄녹병뿐아니라 잎녹병, 줄기녹병, 흰가루병에도 저항성을 나타내므로 육종적으로 매우 유용하다. 우리나라는 줄녹병 저항성 유전육종 연구가 아직 초기단계이므로 한반도 내 주요 균계의 분포와 변화, 저항성 검정체계 구축, 주요 밀 품종과 육성계통의 저항성 검정, 유망 유전자원 확보 및 활용 연구를 활성화하여 발병으로 인한 피해를 사전에 예방하는 것이 시급하다. 또한 장기적으로 분자유전학 및 유전체 분야의 연구를 강화하여 국내 고유의 줄녹병 저항성 유전자를 발굴하고 이를 밀 분자육종에 활용하기 위한 역량을 갖추어야 한다.
본 연구는 농촌진흥청 국립식량과학원 농업과학기술연구개발사업(과제명: 조기 육성 기술을 활용한 고품질 밀 우량 계통 육성 및 유전 분석, 과제번호: PJ014989032020)의 지원에 의해 이루어진 것으로 이에 감사드립니다.
Fig. 1
Life cycle of Puccinia striiformis (Zheng et al. 2013). This figure was imported from the open access article licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License (http://creativecommons.org/licenses/by-nc-nd/3.0/).
KJBS-52-093-f1.gif
Fig. 2
Symptoms of wheat stripe rust disease caused by Puccinia striiformis f. sp. tritici. Photo courtesy: Yue Jin, USDA Agricultural Research Service, Image Number D517-1 (http://www.ars.usda.gov/is/graphics/photos/jun06/d517-1.htm). As indicated in the copyright policy (https://www.ars.usda.gov/oc/images/copyright/), photos in the USDA-ARS Image Gallery are available free of charge and are copyright-free, public domain, images unless otherwise indicated.
KJBS-52-093-f2.gif
Table 1
Summary of major stripe rust (Yr) resistance genes.
Table 1
Locus Acc.z Source Chr.y Typex Referencesw
Yr1 - T. aestivum 2AL ASR Bansal et al. 2009, Hasancebi et al. 2014
Yr4 - T. aestivum 3BS ASR Bansal et al. 2010
Yr5 (YrSP) JN631792 T. spelta album 2BL ASR Chen et al. 2003, Yan et al. 2003, Smith et al. 2007, Murphy et al. 2009, Zhang et al. 2009, Naruoka et al. 2016, Marchal et al. 2018
Yr6 - T. aestivum 7B ASR Li & Niu 2007
Yr7 MN273771 T. aestivum 2B ASR Yao et al. 2006, Marchal et al. 2018
Yr8 - T. comosa 2D ASR Niu et al. 2004
Yr9 DQ167397 S. cereale 1BL ASR Mago et al. 2005
Yr10 AF149112 T. aestivum 1BS ASR Wang et al. 2002, Singh et al. 2009, Yuan et al. 2012, 2018, Liu et al. 2014
Yr11 - T. aestivum 2D HTAP McIntosh et al. 2003, Nazari & Wellings 2008
Yr12 - T. aestivum 2D HTAP McIntosh et al. 2003, Nazari & Wellings 2008
Yr13 - T. aestivum 2D HTAP McIntosh et al. 2003, Nazari & Wellings 2008
Yr14 - T. aestivum 2D HTAP McIntosh et al. 2003, Nazari & Wellings 2008
Yr15 (Wtk1) MG649384 T. dicoccoides G25 1BS ASR Peng et al. 2000, Murphy et al. 2009, Yaniv et al. 2015, Klymiuk et al. 2018
Yr16 - T. aestivum 2DL HTAP Agenbag et al. 2012
Yr17 GQ850586 T. ventricosa 2AS HTAP Helguera et al. 2003, Jia et al. 2011
Yr18/Lr34/Sr57 FJ436983 T. aestivum 7DS HTAP Lagudah et al. 2006, 2009, Krattinger et al. 2009
Yr24/Yr26 JX270677 D. villosum 1BS ASR Ma et al. 2001, Zhang et al. 2013, Zeng et al. 2014
Yr28 KX181569 T. tauschii W-219 4DS ASR Zhang et al. 2019
Yr29 - T. aestivum 1BL HTAP Rosewarne et al. 2006
Yr30 - T. aestivum 3BS HTAP Spielmeyer et al. 2003, Hayden et al. 2004
Yr33 - T. aestivum 7D - Zahravi et al. 2003
Yr34 - T. aestivum 5AL HTAP Qureshi et al. 2018
Yr35 - T. dicoccoides 6BS ASR Dadkhodaie et al. 2011
Yr36 (WKS1) EU835199 T. dicoccoides 6BS HTAP Fu et al. 2009
Yr37 A. kotschyi 2DL - Marais et al. 2005
Yr38 A. sharonensis 6AL - Marais et al. 2010
Yr39 - T. aestivum 1BS HTAP Lin & Chen 2007
Yr40 - A. geniculata 5DS - Kuraparthy et al. 2009
Yr41 - T. aestivum 2BS - Luo et al. 2008
Yr42 - A. neglecta 6AS ASR Marais et al. 2009
Yr43/Yr44 - T. aestivum 2BL - Sui et al. 2009
Yr45 - T. aestivum 3DL ASR Li et al. 2011
Yr46/Lr67/Sr55 KR604818 T. aestivum 4DS HTAP Herrera-Foessel et al. 2011, Abdelbacki et al. 2014, Moore et al. 2015
Yr47 - T. aestivum 5BS HTAP Obert et al. 2005, Bansal et al. 2011, Qureshi et al. 2017
Yr48 - T. aestivum 5AL HTAP Lowe et al. 2011
Yr50 - T. intermedium 4BL HTAP Liu et al. 2013
Yr51 - T. aestivum 4AL ASR Randhawa et al. 2014
Yr52 - T. aestivum 7BL HTAP Ren et al. 2012
Yr53 - T. aestivum 2BL ASR Xu et al. 2013
Yr57 - T. aestivum 3BS ASR Randhawa et al. 2015
Yr58 - T. aestivum 3BS ASR Chhetri et al. 2016
Yr59 - T. aestivum 7BL HTAP Zhou et al. 2014a
Yr60 - T. aestivum 4AL ASR, HTAP Herrera-Foessel et al. 2015
Yr61 - T. aestivum 7AS ASR Zhou et al. 2014b
Yr62 - T. aestivum 4BL HTAP Lu et al. 2014
Yr64 - T. durum 1BS - Cheng et al. 2014, Qie et al. 2019
Yr65 - T. durum 1BS ASR Cheng et al. 2014
Yr67 - T. aestivum 7BL - Li et al. 2009
Yr69 - T. aestivum 2AS HTAP Hou et al. 2016
Yr71 - T. aestivum 3D HTAP Bariana et al. 2016
Yr76 - T. compactum 3AS - Xiang et al. 2016
Yr81 - T. aestivum 6A HTAP Gessese et al. 2019

zWheat nucleotide accession registered at NCBI (National Center for Biotechnology Information)

yChromosomal location

xASR: all-stage resistance, HTAP: high temperature adult plant resistance, -: uncertain

wReference for gene cloning is underlined

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Research Advances in Wheat Breeding and Genetics for Stripe Rust Resistance
Korean. J. Breed. Sci.. 2020;52(2):93-103.   Published online June 1, 2020
DOI: https://doi.org/10.9787/KJBS.2020.52.2.93
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Research Advances in Wheat Breeding and Genetics for Stripe Rust Resistance
Korean. J. Breed. Sci.. 2020;52(2):93-103.   Published online June 1, 2020
DOI: https://doi.org/10.9787/KJBS.2020.52.2.93
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Research Advances in Wheat Breeding and Genetics for Stripe Rust Resistance
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Fig. 1 Life cycle of Puccinia striiformis (Zheng et al. 2013). This figure was imported from the open access article licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License (http://creativecommons.org/licenses/by-nc-nd/3.0/).
Fig. 2 Symptoms of wheat stripe rust disease caused by Puccinia striiformis f. sp. tritici. Photo courtesy: Yue Jin, USDA Agricultural Research Service, Image Number D517-1 (http://www.ars.usda.gov/is/graphics/photos/jun06/d517-1.htm). As indicated in the copyright policy (https://www.ars.usda.gov/oc/images/copyright/), photos in the USDA-ARS Image Gallery are available free of charge and are copyright-free, public domain, images unless otherwise indicated.

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Research Advances in Wheat Breeding and Genetics for Stripe Rust Resistance

Summary of major stripe rust (Yr) resistance genes.

Locus Acc.z Source Chr.y Typex Referencesw
Yr1 - T. aestivum 2AL ASR Bansal et al. 2009, Hasancebi et al. 2014
Yr4 - T. aestivum 3BS ASR Bansal et al. 2010
Yr5 (YrSP) JN631792 T. spelta album 2BL ASR Chen et al. 2003, Yan et al. 2003, Smith et al. 2007, Murphy et al. 2009, Zhang et al. 2009, Naruoka et al. 2016, Marchal et al. 2018
Yr6 - T. aestivum 7B ASR Li & Niu 2007
Yr7 MN273771 T. aestivum 2B ASR Yao et al. 2006, Marchal et al. 2018
Yr8 - T. comosa 2D ASR Niu et al. 2004
Yr9 DQ167397 S. cereale 1BL ASR Mago et al. 2005
Yr10 AF149112 T. aestivum 1BS ASR Wang et al. 2002, Singh et al. 2009, Yuan et al. 2012, 2018, Liu et al. 2014
Yr11 - T. aestivum 2D HTAP McIntosh et al. 2003, Nazari & Wellings 2008
Yr12 - T. aestivum 2D HTAP McIntosh et al. 2003, Nazari & Wellings 2008
Yr13 - T. aestivum 2D HTAP McIntosh et al. 2003, Nazari & Wellings 2008
Yr14 - T. aestivum 2D HTAP McIntosh et al. 2003, Nazari & Wellings 2008
Yr15 (Wtk1) MG649384 T. dicoccoides G25 1BS ASR Peng et al. 2000, Murphy et al. 2009, Yaniv et al. 2015, Klymiuk et al. 2018
Yr16 - T. aestivum 2DL HTAP Agenbag et al. 2012
Yr17 GQ850586 T. ventricosa 2AS HTAP Helguera et al. 2003, Jia et al. 2011
Yr18/Lr34/Sr57 FJ436983 T. aestivum 7DS HTAP Lagudah et al. 2006, 2009, Krattinger et al. 2009
Yr24/Yr26 JX270677 D. villosum 1BS ASR Ma et al. 2001, Zhang et al. 2013, Zeng et al. 2014
Yr28 KX181569 T. tauschii W-219 4DS ASR Zhang et al. 2019
Yr29 - T. aestivum 1BL HTAP Rosewarne et al. 2006
Yr30 - T. aestivum 3BS HTAP Spielmeyer et al. 2003, Hayden et al. 2004
Yr33 - T. aestivum 7D - Zahravi et al. 2003
Yr34 - T. aestivum 5AL HTAP Qureshi et al. 2018
Yr35 - T. dicoccoides 6BS ASR Dadkhodaie et al. 2011
Yr36 (WKS1) EU835199 T. dicoccoides 6BS HTAP Fu et al. 2009
Yr37 A. kotschyi 2DL - Marais et al. 2005
Yr38 A. sharonensis 6AL - Marais et al. 2010
Yr39 - T. aestivum 1BS HTAP Lin & Chen 2007
Yr40 - A. geniculata 5DS - Kuraparthy et al. 2009
Yr41 - T. aestivum 2BS - Luo et al. 2008
Yr42 - A. neglecta 6AS ASR Marais et al. 2009
Yr43/Yr44 - T. aestivum 2BL - Sui et al. 2009
Yr45 - T. aestivum 3DL ASR Li et al. 2011
Yr46/Lr67/Sr55 KR604818 T. aestivum 4DS HTAP Herrera-Foessel et al. 2011, Abdelbacki et al. 2014, Moore et al. 2015
Yr47 - T. aestivum 5BS HTAP Obert et al. 2005, Bansal et al. 2011, Qureshi et al. 2017
Yr48 - T. aestivum 5AL HTAP Lowe et al. 2011
Yr50 - T. intermedium 4BL HTAP Liu et al. 2013
Yr51 - T. aestivum 4AL ASR Randhawa et al. 2014
Yr52 - T. aestivum 7BL HTAP Ren et al. 2012
Yr53 - T. aestivum 2BL ASR Xu et al. 2013
Yr57 - T. aestivum 3BS ASR Randhawa et al. 2015
Yr58 - T. aestivum 3BS ASR Chhetri et al. 2016
Yr59 - T. aestivum 7BL HTAP Zhou et al. 2014a
Yr60 - T. aestivum 4AL ASR, HTAP Herrera-Foessel et al. 2015
Yr61 - T. aestivum 7AS ASR Zhou et al. 2014b
Yr62 - T. aestivum 4BL HTAP Lu et al. 2014
Yr64 - T. durum 1BS - Cheng et al. 2014, Qie et al. 2019
Yr65 - T. durum 1BS ASR Cheng et al. 2014
Yr67 - T. aestivum 7BL - Li et al. 2009
Yr69 - T. aestivum 2AS HTAP Hou et al. 2016
Yr71 - T. aestivum 3D HTAP Bariana et al. 2016
Yr76 - T. compactum 3AS - Xiang et al. 2016
Yr81 - T. aestivum 6A HTAP Gessese et al. 2019
Table 1 Summary of major stripe rust (Yr) resistance genes.

Wheat nucleotide accession registered at NCBI (National Center for Biotechnology Information)

Chromosomal location

ASR: all-stage resistance, HTAP: high temperature adult plant resistance, -: uncertain

Reference for gene cloning is underlined

Table 1

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