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밀 줄기녹병 저항성 유전·육종 주요 연구동향

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

Research Advances in Wheat Breeding and Genetics for Stem 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(3):225-234.
Published online: September 1, 2020

1국립식량과학원

2경남과학기술대학교 농학한약자원학부

3전북대학교 작물생명과학과

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*Corresponding Author (E-mail: moyj82@korea.kr, Tel: +82-63-238-5232, Fax: +82-63-238-5205)
• Received: May 8, 2020   • Revised: May 11, 2020   • Accepted: June 5, 2020

Copyright © 2020 by the Korean Society of Breeding Science

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  • AbstractStem rust is a major wheat disease caused by the fungus Puccinia graminis f. sp. tritici (Pgt). Occurring mainly in warm and humid climates, stem rust has generally been considered less prevalent than leaf rust (P. triticina) and stripe rust (P. striiformis f. sp. tritici). However, a highly virulent stem rust race, Ug99, appeared in Uganda in 1998 and has devastated wheat production throughout Africa and the Middle East. As damage caused by the Ug99 lineage and other newly diverging stem rust races continues to increase, extensive research on wheat breeding and genetics to enhance stem rust resistance has been conducted internationally. Among the 60 stem rust resistance genes reported thus far, 11 (Sr13, Sr21, Sr22, Sr33, Sr35, Sr45, Sr46, Sr50, Sr55, Sr57, and Sr60) have been cloned. New resistance sources have been sought by screening diverse wheat germplasms through international collaborations. Such efforts are urgently required in Korea to address the increasing threat of stem rust epidemics. Furthermore, major Pgt races in the Korean peninsula need to be pathotyped. This information can be used to screen major wheat cultivars, breeding lines, and landraces to identify effective resistance sources. Previously reported stem rust resistance genes should also be introduced and pyramided into the genetic background of Korean wheat breeding populations via available molecular markers. Finally, research capacity in molecular genetics and genomics needs to be strengthened to enable the identification of new stem rust resistance genes and the development of precise molecular markers.
  • Keywords: wheat; stem rust; resistance; breeding
녹병은 밀에서 발생하는 가장 심각한 질병 중 하나로 Puccinia triticina (Pt)에 의한 잎녹병(leaf rust), P. striiformis f. sp. tritici (Pst)에 의한 줄녹병(stripe rust, yellow rust), P. graminis f. sp. tritici (Pgt)에 의한 줄기녹병(stem rust, black rust)으로 구분된다(Cummins & Hiratsuka 2004, Duplessis et al. 2011, Figueroa et al. 2018). 줄기녹병은 잎녹병이나 줄녹병보다는 분포지역과 발생빈도가 낮은 편이지만 발병하였을 때의 피해는 밀 생산에 매우 큰 위협이 된다(Saari & Prescott 1985, Leonard & Szabo 2005). 줄기녹병은 일반적으로 따뜻하고 습한 조건이 우세한 지역에서 주로 발견되며 전형적인 감염 증상은 감수성 식물체의 잎집, 줄기, 포영과 까락에 적색 벽돌과 같은 포자 덩어리 형태로 나타난다(Kolmer 2005). 역사적으로 가장 공격적인 녹병으로 알려져 있으며 그 포자는 청동기 후기의 겉겨 조각에서 이미 발견되었다(Kislev 1982).
우간다에서 1998년 처음 보고된 치명적인 줄기녹병 균계인 Ug99은 아프리카 전역과 중동 및 남아시아로 퍼져서 30억 달러 이상의 경제적 손실을 야기했다(Huerta-Espino et al. 2011, Aktar-Uz-Zaman et al. 2017). 20세기 중반 유럽에서는 줄기녹병으로 30% 수준의 수량 감소가 있었고, 중국, 인도와 호주에서도 대규모 피해가 보고되었다(Zadoks 1963, Leonard & Szabo 2005, Draz et al. 2015). Ug99의 출현을 계기로 녹병 발병지역과 균계 분포에 대한 시공간적 데이터 수집을 위하여 국제 녹병 모니터링 시스템(global cereal rust monitoring system)이 개발되었는데, 이는 녹병 방제에 매우 효과적일 뿐 아니라 다른 병원균의 방제를 위한 모델로도 활용되고 있다(Hodson et al. 2009, Park et al. 2011). 이를 통하여 전세계 많은 지역에서 줄기녹병이 비교적 잘 관리되어 왔지만, 줄기녹병 저항성이 없다고 가정한 모델에서 밀의 수량 손실은 연간 평균 620만 톤 이상일 것으로 추정된다(Pardey et al. 2013).
현재까지 줄기녹병 저항성 유전자는 60여개가 보고되었으나(McIntosh et al. 1995, 2013, Pretorius et al. 2010, Olivera Firpo et al. 2015, Chen et al. 2020) 신균계 Ug99이 출현하고 빠르게 진화하면서 Sr21, Sr24, Sr31, Sr36, Sr38 및 SrTmp와 같은 육종적으로 중요한 저항성 유전자가 속수무책이 되었다(Singh et al. 2006, Jin et al. 2008, 2009, Pretorius et al. 2010, Olivera Firpo et al. 2015). 따라서 밀을 주식으로 하는 선진국을 중심으로 줄기녹병 저항성을 강화하기 위한 연구가 활발하게 진행되고 있다. 그러나 우리나라는 아직 관련 자원과 정보가 많이 부족한 실정이다. 본 논문은 국내 환경에 적합한 줄기녹병 저항성 밀 품종 육성에 필요한 정보를 정리하였다.
줄기녹병 생활사
줄기녹병을 일으키는 P. graminis f. sp. tritici (Pgt)의 생활사는 Fig. 1과 같다. 숙주의 종류에 따른 5가지 포자 형태 중 여름 포자(urediniospores)와 겨울 포자(teliospores), 담자 포자(basidiospores)는 밀에서 발생하고 병포자(pycniospores)와 녹포자(aeciospores)는 대체 숙주(alternative host)에서 발생한다(Bolton et al. 2008, McIntosh 2009, Jin et al. 2010, Chen et al. 2014). 줄녹병과마찬가지로매자나무속(barberry) Mahonia spp.와 Berberis spp.가 줄기녹병의 주요 대체숙주이다(Chen et al. 2014).
줄기녹병에 감염되면 잎녹병과 유사하게 주로 원형이나 타원형의 주황색 농포인 여름 포자가 나타나고 시간이 지나면서 점차 검정색의 겨울 포자로 발전한다(Fig. 2). 30°C 이상의 고온이 곰팡이 성장과 포자 형성의 최적 온도이며 포자의 발아 및 침투에 수분이 필수적인데, 줄기녹병 발생의 최적 조건에서는 잎녹병과 줄녹병 보다 훨씬 많은 수의 포자가 생성되어 최대 10,000여개 포자로 구성된 농포가 전체 식물에 형성된다(Singh et al. 2015). 줄기녹병의 감염은 종실 크기 감소와 도복으로 인한 수확량 감소로 최대 80%의 손실을 유발한다(Leonard & Szabo. 2005, Figueroa et al. 2016).
줄기녹병 저항성 기작
줄기녹병 저항성 유전자는 잎녹병 및 줄녹병 저항성 유전자와 마찬가지로 특정 균계에만 저항성 반응을 보이는 종특이적(race-specific) 저항성 유전자와 다수의 균계 또는 병원균에 반응하는 종비특이적(race non-specific) 저항성 유전자로 구분할 수 있다. 종특이적 저항성 유전자의 대부분이 유묘기 저항성(seedling resistance; SR) 유전자 또는 주동 저항성(major resistance) 유전자로도 구분되며 저항성 기작은 병원균의 비병원성 유전자(Avr gene)에 저항성 유전자가 1:1로 반응하는 gene-for-gene 모형으로 설명된다(Zambino et al. 2000, Lowe et al. 2011, Periyannan et al. 2017). 대부분의 종특이적 저항성 유전자는 NBS-LRR (nucleotide binding site-leucine rich repeat) 형태의 단백질을 형성하는 것으로 알려져 있으며, 동정이 이루어진 줄기녹병 저항성 유전자인 Sr13, Sr21, Sr22, Sr33, Sr35, Sr45, Sr50를 비롯한 많은 유전자가 이에 해당된다(Chen et al. 2018, Periyannan et al. 2013, Saintenac et al. 2013, Mago et al. 2015, Steuernagel et al. 2016, Zhang et al. 2017). 종특이적 저항성 유전자는 특정 균계에 대하여 강한 저항성 반응을 보이지만 균계의 빠른 진화로 인하여 수년 안에 저항성이 무너지는 경우가 많다(Duplessis et al. 2011, Ellis et al. 2014).
종비특이적 저항성 유전자는 대부분 성체 저항성(adult plant resistance; APR) 유전자 또는 미동 저항성(minor resistance) 유전자로 구분되며 종특이적 저항성 유전자에 비하여 훨씬 광범위한 균계에 저항성을 보이고 저항성이 지속되는 기간도 보다 길다(Schnurbusch et al. 2004, Singh et al. 2005). 대표적인 종비특이적 줄기녹병 저항성 유전자는 Sr2로, 밀 육종에 오랫동안 활용되어 왔지만 단일 유전자로는 충분한 저항성이 보장되지 않기 때문에 다른 저항성 유전자와의 집적을 통하여 주로 활용된다(Samsapour et al. 2010, Lagudah 2011, Singh et al. 2015). 또다른 종비특이적 줄기녹병 저항성 유전자인 Sr55/Lr67/Yr46/Pm46, Sr57/Lr34/Yr18/Pm38, Sr58/Lr46/Yr29/Pm39의 경우 다면발현으로 잎녹병, 줄녹병, 흰가루병에도 동시에 저항성을 나타낸다(Mateos-Hernandez et al. 2006, Krattinger et al. 2009, Moore et al. 2015). 종비특이적 저항성의 구체적인 기작은 아직 알려진 것이 별로 없는데, 동정이 이루어진 Sr55/Lr67/Yr46/Pm46는 hexose transporter 단백질을 만들고 세포벽의 육탄당/자당(hexose/sucrose) 비율을 병원균 생존에 불리하게 조절함으로써 다수의 병원균 및 다양한 균계에 대한 저항성을 가지는 것으로 추측되며(Moore et al. 2015), Sr57/Lr34/Yr18/Pm38은 ATP-binding cassette transporter 단백질을 형성하며 엽 노화(senescence)와 유사한 기작으로 광범위 저항성을 부여하는 것으로 추측된다(Krattinger et al. 2009).
줄기녹병 저항성 유전자
줄기녹병 저항성 유전자는 Sr로 표기하며 현재까지 보고된 60여개의 저항성 유전자 중 11개(Sr13, Sr21, Sr22, Sr33, Sr35, Sr45, Sr46, Sr50, Sr55, Sr57, Sr60)가 동정되었다(Table 1). 줄기녹병 저항성 증진을 위해 1920년대에 60종 이상의 줄기녹병 균계에 대하여 종비특이적으로 부분 저항성을 나타내는 APR 유전자 Sr2가 발견되어 저항성 품종 육성에 광범위하게 사용되었고, 1970년대에는 호밀에서 유래한 줄기녹병 저항성 유전자인 Sr26과 Sr31이 많이 사용되었는데, 특히 Sr31은 전세계적으로 사용되어 많은 봄밀과 겨울밀 품종에 줄기녹병 저항성의 기초를 제공하였다(Pretorius et al. 2000). 하지만 Sr31을 침해하는 악성 병원성인 Ug99(TTKSK) 균계가 1998년 우간다에서 발견되어 아프리카 전역과 중동으로 확장되었고, 그 후 TTKST나 TTTSK와 같은 Ug99 계열의 변종들이 생겨나면서 육종에 널리 활용되는 Sr24, Sr36, Sr38의 저항성도 무너뜨려 전세계 밀 생산을 크게 위협하였다(Singh et al. 2015).
현재까지 보고된 줄기녹병 저항성 유전자들 중 Sr2, Sr13, Sr14, Sr15, Sr22, Sr25, Sr26, Sr27, Sr28, Sr29, Sr32, Sr33, Sr35, Sr37, Sr39, Sr40, Sr42, Sr43, Sr44, Sr45, Sr46, Sr47, Sr48, Sr50, Sr51, Sr52, Sr53, Sr55, Sr56, Sr57, Sr58 유전자는 Ug99 계열의 균계에 효과적이라고 보고되었다(Singh et al. 2015, Aktar-Uz-Zaman et al. 2017). 하지만 최근에는 Ug99 계열과 다른 줄기녹병 균계인 RRTTF, TKTTF 등이 새롭게 발생하여 동아프리카, 중동, 남아시아 지역에서 피해를 발생시키고 있다(Khan et al. 2013, Singh et al. 2015). 특히 TKTTF 균계의 경우 Ug99 계열에 저항성을 보유하여 에티오피아와 케냐 등 아프리카 지역에서 널리 재배되는 ‘Digalu’, ‘Robin’과 같은 주요 품종의 저항성을 침해하면서 큰 피해를 초래하였다(Bhattacharya 2017, Olivera Firpo et al. 2017). 위에 서술한 Ug99 계열에 효과적인 저항성을 보이는 유전자들 중 Sr13, Sr14, Sr15, Sr25, Sr27, Sr28, Sr35, Sr42, Sr43, Sr44, Sr46, Sr50의 경우 이미 새로운 균계에 의하여 저항성이 침해되는 사례가 보고되었다(Singh et al. 2015). 이러한 사례는 줄기녹병의 균계 변화와 지역적 분포를 지속적으로 모니터링하고 새로운 저항성 유전자원을 탐색하여 재배품종으로 신속하게 도입하는 것이 매우 중요함을 시사한다.
줄기녹병은 잎녹병이나 줄녹병에 비하여 발병 피해가 크지 않은 것으로 인식되는 경향이 있었으나 20세기 후반에 신균계인 Ug99가 발생하여 전세계 밀 생산을 크게 위협하면서 줄기녹병 저항성 품종 육성의 중요성이 다시 부각되고 있다. 특히 줄기녹병으로 인한 전세계 식량안보 위협이 계기가 되어 밀 육종가이자 노벨평화상 수상자인 Norman Borlaug 박사의 이름을 따 2005년에 창립된 BGRI (Borlaug Global Rust Initiative)는 30여개 나라와 국제공조를 통하여 매년 수십만 점의 밀 유전자원에 대한 스크리닝을 진행하고 줄기녹병 저항성 자원을 탐색하고 있다(Singh et al. 2015).
미국의 경우 Ug99로 인한 대규모 피해사례는 없었음에도 불구하고 피해를 사전에 예방하기 위하여 BGRI와의 협력으로 자국 주요 밀 품종의 줄기녹병 저항성을 평가하여 Ug99에 대한 저항성을 보유한 ‘Tom’, ‘Thatcher’ 등의 품종을 선발하였으며, 주요 저항성 유전자를 신속히 미국 품종에 이전시키기 위한 분자표지도 개발하였다(Rouse et al. 2012, Singh et al. 2015). 또한 멕시코에 위치한 국제옥수수밀연구소(CIMMYT)는 줄기녹병 저항성 강화를 위하여 2006년부터 멕시코-케냐 순환육종(shuttle breeding) 프로그램을 시작하여 저세대 육종집단부터 줄기녹병 저항성을 평가함으로써 줄기녹병에 안정적인 저항성을 보유한 우수 품종을 개발, 보급하고 있다(Singh et al. 2006, 2015).
반면 우리나라는 해외에서 도입한 줄기녹병 저항성 유전자원의 국내환경 적응성에 대한 평가결과가 일부 보고된 바 있으나 아직은 연구 초기단계이다(Shin et al. 2014). 하지만 Ug99 균계의 무서운 전파력과 빠른 진화를 통한 균계 분화 사례에서 보듯 유전적 저항성을 보유하지 못한 상황에서 갑자기 병이 발생할 경우 농가가 큰 타격을 입을 수 있으므로 사전에 줄기녹병 저항성을 확보해야 한다. 이를 위하여 한반도와 주변 지역의 줄기녹병 발생 현황 및 균계 분포를 모니터링하고 주요 균계에 대하여 국내 육성 품종과 재래종을 중심으로 저항성 보유 여부를 평가해야 한다. 또한 기존에 저항성 양상이 분석되고 동정이 이루어져 정밀 분자표지 운용이 가능한 유전자를 대상으로 해당 유전자원을 확보하고 분자표지 활용 여교배(marker assisted backcrossing)를 통하여 국내 품종에 신속히 이전 및 집적 시켜야 한다. 장기적으로는 균계분화로 인하여 기존 저항성 유전자가 침해되는 것에도 자체 역량으로 대응할 수 있도록 다양한 유전자원으로부터 새로운 저항성 유전자를 확보하고 이를 활용한 품종 및 중간모본을 개발하여 국내 밀 육종 집단의 줄기녹병 저항성 스펙트럼을 다양화해야 한다.
P. graminis f. sp. tritici (Pgt)에 의하여 발생하는 줄기녹병은 따뜻하고 습한 환경에서 주로 발생하며 기존에는 잎녹병이나 줄녹병에 비해서 피해가 적은 것으로 오랫동안 인식되어 왔다. 그러나 20세기 후반 아프리카 지역에서 전염력이 강한 Ug99 균계가 출현하여 전세계적으로 줄기녹병으로 인한 피해가 급격히 증가하면서 새롭게 주목을 받고 있다. 국제적으로는 줄기녹병 저항성에 대한 많은 연구가 진행되어 현재까지 60여개의 저항성 유전자가 보고되었고 그 중 11개(Sr13, Sr21, Sr22, Sr33, Sr35, Sr45, Sr46, Sr50, Sr55, Sr57, Sr60)가 동정되었다. 하지만 지속적인 균계 분화로 인하여 기존에 육종에 널리 활용되던 많은 줄기녹병 저항성 유전자가 침해되면서 피해는 더 확대되고 있다. 따라서 미국을 비롯한 주요 밀 생산국과 국제옥수수밀연구소(CIMMYT)에서는 국제 공조를 통하여 대규모 유전자원의 줄기녹병 저항성을 스크리닝하고 유망 자원을 활용하여 저항성 품종 개발을 진행하고 있다. 아직 줄기녹병 저항성 연구가 초기단계인 우리나라 역시 유전육종 연구 강화를 통하여 대규모 발병으로 인한 피해를 미리 예방해야 한다. 이를 위해서는 줄기녹병 균계 분포와 변화에 대한 모니터링과 함께 주요 균계에 대한 국내 밀 품종 및 육성 계통의 저항성 평가가 우선적으로 진행되어야 한다. 이를 통하여 유전적 취약성을 파악하고 기존에 밝혀진 다양한 줄기녹병 저항성 유전자를 분자표지를 활용하여 신속히 국내 품종으로 이전하고 집적 시켜야 한다. 또한 새로운 저항성 유전자를 지속적으로 발굴하고 이를 육종에 활용하여 균계 분화로 인한 저항성 침해에도 미리 대비해야 할 것이다.
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 graminis. Image courtesy: USDA Agricultural Research Service (http://www.ars.usda.gov/Main/docs.htm?docid=10787). As indicated in the copyright policy (https://www.ars.usda.gov/oc/images/copyright/), images in the USDA-ARS Image Gallery are available free of charge and are copyright-free, public domain, images unless otherwise indicated.
KJBS-52-225-f1.gif
Fig. 2
Symptoms of wheat stem rust disease caused by Puccinia graminis f. sp. Tritici. Photo courtesy: Yue Jin, USDA Agricultural Research Service, Image Number K11192-1 (http://www.ars.usda.gov/is/graphics/photos/jun04/k11192-1.htm). As indicated in the copyright policy (https://www.ars.usda.gov/oc/images/copyright/), images in the USDA-ARS Image Gallery are available free of charge and are copyright-free, public domain, images unless otherwise indicated.
KJBS-52-225-f2.gif
Table 1
Summary of major stem rust (Sr) resistance genes.
Table 1
Locus Accz Source Chry Typex Referencew
Sr2 KP244323 Triticum aestivum 3BS APR Spielmeyer et al. 2003, Hayden et al. 2004, Mago et al. 2011, 2014
Sr9a - Triticum aestivum 2BL - Tsilo et al. 2007
Sr9h - Triticum aestivum 2BL SR Rouse et al. 2014
Sr11 - Triticum aestivum 6BL SR Nirmala et al. 2016
Sr12 - Triticum aestivum 3BL SR Hiebert et al. 2016
Sr13 KY825229 Triticum dicoccoides 6AL SR Admassu et al. 2011, Simons et al. 2011, Zhang et al. 2017
Sr15 - Triticum aestivum 2BL SR Jayatilake et al. 2013, Babiker et al. 2015, Gao et al. 2019
Sr21 MG582649 Triticum monococcum 2AL SR Chen et al. 2015, Chen et al. 2018
Sr22 MH546067 Triticum monococcum 7AL SR Olson et al. 2010a, Steuernagel et al. 2016
Sr24 X95647 Agropyron elongatum 3DL SR Mago et al. 2005a, Pretorius et al. 2012
Sr25 - Thinopyrum ponticum 7DL SR Ayala-N. et al. 2007, Liu et al. 2010, Yadav et al. 2015, Li et al. 2016
Sr26 GQ899187 Agropyron elongatum 6AL SR Mago et al. 2005a, Dundas et al. 2007, Liu et al. 2010, Li et al. 2016
Sr27 - Secale cereale 3AS SR Acosta 1963
Sr28 - Triticum aestivum 2BL SR Rouse et al. 2012, Babiker et al. 2017
Sr31 DQ849629 Secale cereale 1BL SR Mago et al. 2005b, Das et al. 2006, Pretorius et al. 2012
Sr32 - Triticum speltoides 2DS SR Mago et al. 2013, Li et al. 2016
Sr33 KF031303 Aegilops tauschii 1DS SR Sambasivam et al. 2008, Periyannan et al. 2013
Sr35 KC573058 Triticum monococcum 3AL SR Saintenac et al. 2013
Sr36 - Triticum timopheevi 2BS - Tsilo et al. 2008, Olson et al. 2010b, Chemayek et al. 2017
Sr37 - Triticum timopheevii 4BL SR Zhang et al. 2012
Sr38 - Triticum ventricosa 2AS SR Helguera et al. 2003, Li et al. 2016
Sr39 - Triticum speltoides 2BS SR Gold et al. 1999, Mago et al. 2009, Chemayek et al. 2017
Sr40 - Triticum timopheevii ssp. araraticum 2BS - Wu ea al. 2009
Sr42 - Triticum aestivum 6DS SR Olson et al. 2013
Sr43 - Thinopyrum ponticum 7DL - Niu et al. 2014
Sr44 - Thinopyrum intermedium 7DS - Liu et al. 2013
Sr45 LN883757 Aegilops tauschii 1DS SR Sambasivam et al. 2008, Periyannan et al. 2014, Steuernagel et al. 2016
Sr46 - Aegilops tauschii 2DS - Yu et al. 2015, Arora et al. 2018
Sr47 - Triticum speltoides 2BL - Klindworth et al. 2012
Sr49 - Triticum aestivum 5BL - Bansal et al. 2015
Sr50 KT725818 Secale cereale 1DL SR Mago et al. 2015
Sr51 - Aegilops geniculata 5DL SR Liu et al. 2011a
Sr52 - Dasypyrum villosum 6AL - Qi et al. 2011
Sr53 - Aegilops geniculata 5DL - Liu et al. 2011b
Sr55/Yr46/Lr67 KR604818 Triticum aestivum 4DL APR McIntosh 2012, Moore et al. 2015
Sr56 - Triticum aestivum 5BL APR Bansal et al. 2014
Sr57/Yr18/Lr34 FJ436983 Triticum aestivum 7DS APR Lagudah et al. 2006, 2009, Krattinger et al. 2009
Sr58 - Triticum aestivum 1BL APR Mateos-Hernandez et al. 2006
Sr59 - Secale cereale 2RL - Rahmatov et al. 2016
Sr60 MK629708 Triticum monococcum 5AS - Chen et al. 2020

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

yChromosomal location.

xAPR: adult plant resistance, SR: seedling resistance, -: uncertain.

wReference for gene cloning is underlined.

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Research Advances in Wheat Breeding and Genetics for Stem Rust Resistance
Korean. J. Breed. Sci.. 2020;52(3):225-234.   Published online September 1, 2020
DOI: https://doi.org/10.9787/KJBS.2020.52.3.225
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Research Advances in Wheat Breeding and Genetics for Stem Rust Resistance
Korean. J. Breed. Sci.. 2020;52(3):225-234.   Published online September 1, 2020
DOI: https://doi.org/10.9787/KJBS.2020.52.3.225
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Research Advances in Wheat Breeding and Genetics for Stem Rust Resistance
Image Image
Fig. 1 Life cycle of Puccinia graminis. Image courtesy: USDA Agricultural Research Service (http://www.ars.usda.gov/Main/docs.htm?docid=10787). As indicated in the copyright policy (https://www.ars.usda.gov/oc/images/copyright/), images in the USDA-ARS Image Gallery are available free of charge and are copyright-free, public domain, images unless otherwise indicated.
Fig. 2 Symptoms of wheat stem rust disease caused by Puccinia graminis f. sp. Tritici. Photo courtesy: Yue Jin, USDA Agricultural Research Service, Image Number K11192-1 (http://www.ars.usda.gov/is/graphics/photos/jun04/k11192-1.htm). As indicated in the copyright policy (https://www.ars.usda.gov/oc/images/copyright/), images in the USDA-ARS Image Gallery are available free of charge and are copyright-free, public domain, images unless otherwise indicated.

Fig. 1

Fig. 2

Research Advances in Wheat Breeding and Genetics for Stem Rust Resistance

Summary of major stem rust (Sr) resistance genes.

Locus Accz Source Chry Typex Referencew
Sr2 KP244323 Triticum aestivum 3BS APR Spielmeyer et al. 2003, Hayden et al. 2004, Mago et al. 2011, 2014
Sr9a - Triticum aestivum 2BL - Tsilo et al. 2007
Sr9h - Triticum aestivum 2BL SR Rouse et al. 2014
Sr11 - Triticum aestivum 6BL SR Nirmala et al. 2016
Sr12 - Triticum aestivum 3BL SR Hiebert et al. 2016
Sr13 KY825229 Triticum dicoccoides 6AL SR Admassu et al. 2011, Simons et al. 2011, Zhang et al. 2017
Sr15 - Triticum aestivum 2BL SR Jayatilake et al. 2013, Babiker et al. 2015, Gao et al. 2019
Sr21 MG582649 Triticum monococcum 2AL SR Chen et al. 2015, Chen et al. 2018
Sr22 MH546067 Triticum monococcum 7AL SR Olson et al. 2010a, Steuernagel et al. 2016
Sr24 X95647 Agropyron elongatum 3DL SR Mago et al. 2005a, Pretorius et al. 2012
Sr25 - Thinopyrum ponticum 7DL SR Ayala-N. et al. 2007, Liu et al. 2010, Yadav et al. 2015, Li et al. 2016
Sr26 GQ899187 Agropyron elongatum 6AL SR Mago et al. 2005a, Dundas et al. 2007, Liu et al. 2010, Li et al. 2016
Sr27 - Secale cereale 3AS SR Acosta 1963
Sr28 - Triticum aestivum 2BL SR Rouse et al. 2012, Babiker et al. 2017
Sr31 DQ849629 Secale cereale 1BL SR Mago et al. 2005b, Das et al. 2006, Pretorius et al. 2012
Sr32 - Triticum speltoides 2DS SR Mago et al. 2013, Li et al. 2016
Sr33 KF031303 Aegilops tauschii 1DS SR Sambasivam et al. 2008, Periyannan et al. 2013
Sr35 KC573058 Triticum monococcum 3AL SR Saintenac et al. 2013
Sr36 - Triticum timopheevi 2BS - Tsilo et al. 2008, Olson et al. 2010b, Chemayek et al. 2017
Sr37 - Triticum timopheevii 4BL SR Zhang et al. 2012
Sr38 - Triticum ventricosa 2AS SR Helguera et al. 2003, Li et al. 2016
Sr39 - Triticum speltoides 2BS SR Gold et al. 1999, Mago et al. 2009, Chemayek et al. 2017
Sr40 - Triticum timopheevii ssp. araraticum 2BS - Wu ea al. 2009
Sr42 - Triticum aestivum 6DS SR Olson et al. 2013
Sr43 - Thinopyrum ponticum 7DL - Niu et al. 2014
Sr44 - Thinopyrum intermedium 7DS - Liu et al. 2013
Sr45 LN883757 Aegilops tauschii 1DS SR Sambasivam et al. 2008, Periyannan et al. 2014, Steuernagel et al. 2016
Sr46 - Aegilops tauschii 2DS - Yu et al. 2015, Arora et al. 2018
Sr47 - Triticum speltoides 2BL - Klindworth et al. 2012
Sr49 - Triticum aestivum 5BL - Bansal et al. 2015
Sr50 KT725818 Secale cereale 1DL SR Mago et al. 2015
Sr51 - Aegilops geniculata 5DL SR Liu et al. 2011a
Sr52 - Dasypyrum villosum 6AL - Qi et al. 2011
Sr53 - Aegilops geniculata 5DL - Liu et al. 2011b
Sr55/Yr46/Lr67 KR604818 Triticum aestivum 4DL APR McIntosh 2012, Moore et al. 2015
Sr56 - Triticum aestivum 5BL APR Bansal et al. 2014
Sr57/Yr18/Lr34 FJ436983 Triticum aestivum 7DS APR Lagudah et al. 2006, 2009, Krattinger et al. 2009
Sr58 - Triticum aestivum 1BL APR Mateos-Hernandez et al. 2006
Sr59 - Secale cereale 2RL - Rahmatov et al. 2016
Sr60 MK629708 Triticum monococcum 5AS - Chen et al. 2020
Table 1 Summary of major stem rust (Sr) resistance genes.

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

Chromosomal location.

APR: adult plant resistance, SR: seedling resistance, -: uncertain.

Reference for gene cloning is underlined.

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