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국내 재래종 벼 유전자원의 중배축 신장 특성 평가

박소연1, 장성규1, 이주현2, 권순욱1,*

Evaluation of Mesocotyl Elongation Ability in Korean Rice Landraces (Oryza sativa L.)

So-Yeon Park1, Seong-Gyu Jang1, Joohyun Lee2, Soon-Wook Kwon1,*
Korean Journal of Breeding Science 2019;51(4):351-356.
Published online: December 1, 2019

1부산대학교 식물생명과학과

2건국대학교 식량자원과학과

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* Corresponding Author (E-mail: swkwon@pusan.ac.kr, Tel: +82-55-350-5506, Fax: +82-55-350-5509)
• Received: August 3, 2019   • Revised: September 23, 2019   • Accepted: October 10, 2019

Copyright © 2019 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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  • Rice landrace germplasms are highly valuable because of their wide genetic variation. Their traits have not been selected by modern breeders but by nature, thus, these traits and the gene associated with them have tremendous potential to improve modern rice cultivars. The first step to utilize this potential is to conduct a thorough evaluation of the target traits to select superior germplasm for a breeding program. Here, with 386 Korean landrace germplasms, we evaluated mesocotyl elongation traits which possibly promote direct seedling cultivation in rice. Mesocotyl length ranged from 0 mm to 28 mm. Among the tested landraces, we selected and reported the top 20 performing landraces whose mesocotyl length were larger than 16 mm. The previously reported agronomic traits for the selected 20 landraces were also listed for breeders to promote the utilization of these germplasms in breeding programs.
  • Keywords: Rice landrace; Mesocotyl elongation; Directly seedling; Germplasm
재래종은 육종기술의 발전 이전에 자연 또는 재배자의 무의식적인 선택으로 고정된 유전자원이다(Camacho Villa et al. 2005). 재래종은 우수한 유전적 다양성을 지니고 있는데, 벼에서는 분자마커를 이용한 다양성 검정을 통해 국내 재래종 집단이 육성종 집단보다 더 높은 다양성을 가지고 있음을 확인한 바 있다(Kwon et al. 2000). 또한 국내 재래종 유전자원 76점에 대해 유전다양성을 분석한 결과 마커 별 평균 PIC 값이 0.58로 보고되었는데(Lee et al. 2015), 이는 729개 벼 유전자원에서의 PIC 값이 0.29이고, 아시아 국가의 유전자원을 평가한 PIC값이 0.50으로 보고(Lee et al. 2018)한 것보다 높은 다양성을 보인 것이다. 국내 재래종 벼 유전자원 394점에 대해 집단구조 분석을 수행한 결과 3개의 집단으로 구분되었으며, 집단 1과 3은 대부분의 자포니카형을, 집단 2는 인디카형을 포함하고 있었다(Li et al. 2014).
재래종 벼 유전자원에 대한 특성 평가는 농촌진흥청에서 출수기, 초형 등 기본 농업형질과 미질 특성, 영양성분 및 기능성 성분 함량 등 활용도 증진을 위한 특성 평가가 수행된 바 있으며(Kang et al. 2008), 흰잎마름병 저항성을 평가하여 저항성 유전자원을 보고한 바 있다(Li et al. 2011). 최근에는 재래종 유전자원의 향 특성 등 최근의 소비자 요구에 부합되는 다양한 형질에 대한 평가가 이루어 진 바 있으며(He & Park 2015), 다수의 우수 자원이 선발되었다. 이러한 결과는 국내 재래종 유전자원들의 잠재적 가치를 보여준다고 할 수 있다. 따라서 재래종 집단을 이용한다면 목적 형질을 갖는 우수한 자원을 선발할 수 있을 것이며, 최근에는 생물 다양성 협약으로 인한 유전자원 활용의 제한이 생기면서 재래종 활용의 중요성은 더욱 높아질 것이다. 재래종 유전자원 중 떡 가공적성이 우수한 재래종 유전자원 ‘돈나’를 활용하여 ‘효원2호’와 ‘효원4호’ 벼를 육성한 바 있으며(Lee et al. 2016, Jang et al. 2017), ‘자광도’를 활용한 ‘자광찰’ (Kim et al. 2005), ‘중모1033’(Jeong et al. 2019)이 육성되었다. 또한 ‘가위찰’을 활용한 ‘흑가위찰’(Kim et al. 2006)이 보고된 바 있으나, 여전히 그 활용이 미미한 실정이다.
벼는 국내에서 농업 및 식량안보에서 큰 비중을 차지하고 있다. 하지만 도시 유입 인구의 증가에 따라 농촌 인구는 지속적으로 감소하고 고령화 되고 있으며 벼 재배에 필요한 노동력의 부족과 사회 여건의 변화에 따라 국내 벼 재배 면적은 지속적으로 감소하고 있다. 또한 인건비와 유류비의 상승으로 쌀의 생산비용은 올라갔지만 쌀 소비량 감소로 인해 쌀 가격은 하락하게 되어 쌀 생산 농가에 부담이 늘어나고 있다. 이러한 상황에서 쌀 생산 단가를 줄이기 위한 방법 중 하나로 직파재배가 대두되고 있다. 직파재배는 이앙재배와 달리 육묘, 이앙과정이 생략되므로 노동력 소모가 적다. 따라서 직파재배를 이용하면 투여되는 노력과 소모비용을 줄일 수 있는데, 조사된 바에 따르면 건답직파의 경우 20%, 담수 직파에서는 32%의 생산비 절감 효과를 보인다고 했다(Lee et al. 1996). 직파재배에서는 이앙재배와 달리 벼 종자가 일정기간 생육을 거치지 않고 곧바로 본답에 노출되기 때문에 직파재배에 사용되는 벼 품종은 이앙재배와는 다른 특성이 요구되는데, 대표적으로 중배축 신장성이 있다. 직파재배에서 중배축의 신장은 입모율에 큰 영향을 미치는 요인으로 작용하는데(Lee et al. 2017), 일반적으로 인디카 품종이 자포니카 품종에 비해 중배축의 신장이 더 뚜렷하며, 그 변이 또한 인디카 품종군에서 더 크게 나타나는 것으로 보고된 바 있다(Takahashi 1978). 또한 밭벼 품종들이 논벼 품종들보다, 서남아시아 품종들이 동아시아 품종들보다 신장성이 강한 것으로 보고되었다(Wu et al. 2005, Takahashi 1995, Chang and Vergara 1975).
중배축 신장에 관여하는 QTLs는 2개의 자포니카 품종 교배후대의 고세대 RIL집단으로부터 5개의 QTLs가 보고되었으며(Huang et al. 2010), Kasalath와 Nipponbare 교배 후대 집단에서 qMEL-1, qMEL-3 그리고 qMEL-6 세 개의 QTLs가 보고되었다(Lee et al. 2017). 그리고 최근 유전체 분석 기술의 발달로 전장유전체 연관분석(GWAS)이 이루어짐에 따라 중배축 관련 다수의 유전자좌 영역이 보고되고 있다. 170개 미니코어 집단과 100개 내한발성과 관련된 자원을 이용하여 중배축 신장성에 대해 연관 분석하여 1, 3, 4, 5, 6과 9번 염색체에서 유의한 영역이 보고되었고(Wu et al. 2015), 469개 인디카 유전자원을 이용한 중배축 신장성과 관련된 유전자로 Os01g392100, Os01g0904700, Os04g0630000, Os07g0615000가 보고되었으며(Lu et al. 2016), 3,000 Rice Genome Project가 완료됨에 따라 이를 이용한 GWAS 결과들이 보고되고 있다(Zhao et al. 2018). 하지만 국내 자원의 경우, 중배축 신장성에 대한 자원 평가 및 활용 가능한 자원 선발이 부족한 실정이다.
본 연구에서는 재래종 벼 유전자원에 대한 중배축 신장성 검정을 통해 유용한 자원을 선발하여 육종 및 유전자 연구를 위한 자원 선발을 수행하고, 생물다양성 협약에 대응한 우리나라의 유전자원 개발 및 활용에 기초 자료를 얻고자 수행되었다.
식물 재료
농촌진흥청 농업유전자원센터로부터 분양 받은 재래종 벼 유전자원 394점을 부산대학교에서 증식하여 사용하였고, 이 중 증식되지 않은 8점을 제외한 386점에 대한 중배축 신장성 평가를 수행하였다. 대조 품종인 Nipponbare, Kasalath는 충남대학교 농업생명과학대학에서 분양 받아 사용하였다.
중배축 신장 평가
대조품종을 포함하여 총 386개 자원의 중배축 신장성 평가는 Lee et al. (2012)의 방법에 따라 72.00×72.00×100.00 (mm)의 Incu tissue (SPL Life science Co, Kyeonggi, Korea)에 0.3%의 Agar (Duchefa Biochemi, Haarlem, NLD) gel 배지에서 수행하였다. 공시 된 재래종 유전자원들과 대조품종은 육안으로 건전 종자를 선종하여 1%의 차아염소산나트륨에 10분간 침지 소독하고, 증류수로 세척하였다. 준비된 종자는 자원 당 20립씩 0.3% Agar gel에 치상하였고, 14일 간 30℃의 암조건에서 생육 후 중배축 길이를 측정하였다(Fig. 1). 중배축 길이는 치상 된 20립 중 임의 5개체를 대상으로 종자의 배반과 초엽(coleoptile) 사이의 길이를 측정하여 이들의 평균값을 이용하였고, 재래종 386점과 대조품종에 대해 생물학적 2반복을 수행하였다.
Fig. 1
Methods of evaluating mesocotyl elongation. (A) 20 rice seeds inside of the agarose gel for each magenta box. (B) Dark treatment by covering aluminum foil for 14 days.
KJBS-51-351-f1.jpg
중배축 신장성 다양성
중배축 길이 신장의 형질은 토양 속에서 생장하는 중배축의 특성상 환경의 영향을 많이 받는다. Wu et al. (2015)은 중배축 길이에 대한 변이의 폭과 반복 간 차이가 수경재배 조건에서 토양 재배 조건에 비해 변이 폭이 크고 반복 간 낮은 상관을 보고 하였다. 또한 토양 재배 시 복토 깊이에 따라 품종 간 중배축 신장성의 변이도 크다고 보고하여 중배축 신장성 평가는 환경적 요인 혼재 시, 유전자 효과의 검정이 어려운 것으로 보고하였다. 본 실험에서는 균일한 환경을 제공하고자 토양 조건과 유사하면서 압력이 0.3Mpa 이하로서 뿌리 및 지상부 신장을 억제하지 않는 것으로 보고된(Clark et al. 1999) 0.3% agar gel에서의 실험을 수행하였다. 반복간 상관계수는 R2=0.771로 다소의 변이는 존재하였으나 반복간 재현성이 있음을 확인하였다. 즉, 본 실험에 활용된 중배축 길이평가방법으로 각각의 유전자원의 중배축 신장에 관여하는 유전적인 능력을 평가할 수 있음을 확인하였다. 직파재배 적응성 형질로 알려진 중배축 신장성에 대한 정확한 유전자의 탐색 및 이를 활용한 육종 소재의 개발에는 인공 토양에 기반을 두는 정밀한 환경조절 실험과 실제 포장조건에서의 평가가 동반되어야 할 것이며, 환경변이에 대한 정밀한 평가를 위하여 충분한 반복 실험이 진행되어야 할 것이다.
대조 품종 Nipponbare는 평균 1 mm, Kasalath는 13 mm로 중배축 신장을 보였다(Fig. 2). 재래종 386점의 평균 표현형은 0 mm~28 mm 범위로 나타났다. 길이별 빈도 수 측정결과 중배축이 자라지 않은 품종이 42종(10.8%)으로 조사되었으며, 0.0 mm~ 5.0 mm 구간에서 184 accessions (47.4%), 5.0 mm~10.0 mm 내 97 accessions (25%), 10.0 mm~20.0 mm 58 accessions (15%), 20.0 mm~30.0 mm 내 7 accessions (1.8%)가 포함되었다. 표현형 길이 평가 시 평균, 중앙값, 최빈값의 순서로 작아지는 정적 비대칭을 띄었고, 왜도 값 1.41로 표현형들 간 비 정규분포를 보였다(Fig. 3). 평가된 재래종 유전자원 386점 중 요남조(要南租, IT007740), 인부지나도(隣不知나稻, IT008199), 쇠벤치기(IT006776), 대골도(大骨稻, IT005742) 등을 포함하는 42개 품종은 중배축의 신장이 관찰되지 않았으며, 대부분의 유전자원들이 중배축 신장성이 약한 경향을 가졌으나 일부 유전자원들은 강한 중배축 신장성을 나타내었다. ‘앉은뱅이(IT007486)’, ‘육성재래 陸城在來, IT010555) ‘백곡나(IT006302)’, ‘덕적조도(IT005994)’, ‘긴까락샤레(IT173446)’, ‘한양조(漢陽租, IT009060)를 포함하는 6개의 유전자원은 공통적으로 20 mm 이상의 강한 중배축 신장성을 나타내었다. Li et al. (2014)는 국내 재래종 유전자원의 집단구조 분석결과 3개의 집단으로 구분하였는데, 자포니카 품종은 집단 1 (178점)과 집단 3 (108점)으로 구분되었고, 인디카 품종은 집단 2 (38점)에 속한다고 보고하였다. 각각 집단의 평균 중배축 길이는 각각 6.16 mm, 8.11 mm, 3.61 mm로 조사되어 인디카 그룹(집단 2)에서 중배축 신장성이 높게 나타났다. 이는 Takahashi (1978)의 결과와 일치하였다.
Fig. 2
Measuring of Mesocotyl length of the control varieties. (A) a low vigor control variety ‘Nipponbare’, (B) a high-vigor control variety ‘Kasalath’
KJBS-51-351-f2.jpg
Fig. 3
Distribution of mesocotyl length of 386 Korean rice landrace.
KJBS-51-351-f3.jpg
재래종 유전자원의 활용성
실험에 사용한 한국 재래종 자원 중 높은 중배축 신장성을 가진 상위 약 5%에 해당하는 20개 자원을 Table 1에 정리하였다. 재래종 유전자원의 중배축 신장성과 간장과 수장(농업유전자원센터)과의 상관분석 결과, 중배축과 간장 간 상관계수는 0.217로 1%수준에서 유의한 정의 상관을 보였으나, 중배축과 수장 간에는 유의성이 인정되지 않았다. 다만 선발된 상위 5% 유전자원에서는 중배축 신장과 간장간 상관성이 인정되지 않았고, 품종육성 측면에서 중배축 신장성이 양호하면서 초장이 작은 자원이 유리할 것으로 사료된다. 현재 국내에서 이용되고 있는 직파재배 적응성 품종으로는 농안벼, 주안벼, 안산벼, 동안벼 등이 보급되어 있으며, 직파재배 적응성 품종으로 저온발아성이 높으며 높은 초기 신장률로 입모율이 안정적인 품종이 장려되고 있다. 또한 미질 및 수량성이 좋은 품종을 선호하기 때문에 직파재배 외에도 다른 농업 형질이 중요하게 고려되고 있다. 상위 약 5%에 해당하는 20개 품종 중 우수한 농업형질을 나타내는 육성재래, 쥐입파리벼, 찰벼, 밭나락 품종은 중배축 신장에 의한 직파재배 적합성 뿐만 아니라 양질의 품질 및 수량성을 나타내어 추후 교배 모본으로 이용하기에 적합할 것으로 보인다. 그러나 대부분 재래종 유전자원이 도열병 등 각종 병해충에 이병성이고 도복성에 취약하므로 품종 개발 단계에서 저항성 확보에 대한 고려가 필요할 것이다.
Table 1
Some agricultural traits of high mesocotyl elongation ability rice landrace.
Table 1
IT Number Variety Mesocotyl Length (mm) Grain length (mm)Z Grain width (mm)Z Awny Heading Datey Culm Length (cm)y Panicle Length(cm)y Panicles/ planty Lodgey Leaf Blasty Amylose Contentsy
IT007486 Anjeunbaengi 28.0 6.8 3.3 None 8.25 70 24 10 Rx - 0.19
IT010555 Yukseongjaerae 26.5 7.0 3.4 None 8. 2 88 - 8 R M 0.25
IT006302 Baekgokna 24.5 7.3 3.5 Short/Few 8. 6 110 23 5 M S 0.06
IT005994 Deokjeokjodo 24.5 7.0 3.6 Existence 7.31 101 21 12 M M 0.17
IT173446 Ginggalagsyale 24.5 7.3 2.8 None 8. 3 114 22 - - 0.21
IT009060 Hanyangjo 23.0 7.8 3.4 None 8.14 102 23 13 R S 0.31
IT009187 Hwangtodo 20.5 6.4 3.5 Existence 8.16 95 20 6 M M 0.19
IT010627 Joseokjo 20.0 7.4 3.6 Short/Few 7.29 92 21 8 R S 0.16
IT010374 Sando 20.0 7.8 3.5 Existence 8. 5 95 21 7 R S 0.18
IT005044 Guwangdo 19.0 6.2 3.4 Short/Lot 8.27 107 30 8 M - 0.19
IT010612 Jodo 19.0 7.2 3.6 None 8. 2 100 20 6 R S 0.17
IT006556 Sando 19.0 7.2 3.6 Existence 8. 5 83 22 7 M S 0.18
IT006596 Samkyeongjo 19.0 7.1 3.4 Short/Lot 8. 5 85 17 12 S - 0.24
IT008700 Jwiipparibyeo 18.5 6.9 3.4 Short/Few 8.12 99 20 15 S - 0.17
IT005504 Noinjo 17.5 6.9 3.4 Existence 8.19 109 22 8 M S 0.17
IT006372 Baekjo 17.0 6.9 3.6 Existence 8.16 99 24 10 M S 0.18
IT008749 Chalbyeo 16.5 7.7 3.3 Short/Lot 8. 6 92 24 10 M S 0.06
IT007807 Yue 16.0 6.7 3.5 Short/Lot 8.19 116 24 16 S - 0.17
IT006078 Monajo 16.0 6.9 3.6 Short/Lot 8.16 93 20 8 S S 0.05
IT010340 Batnarak 16.0 7.2 3.6 None 8. 7 97 20 9 M S 0.22

zEvaluation average of 20 seeds measured by Vernier calipers

yThese data acquired from the database of National Agrobiodiversity Center (http://genebank.rda.go.kr/simpleSearch.do)

xR: Resistant, M: Moderate, S: Susceptible

재래종 유전자원은 다양한 유전적 다양성 때문에 높은 가치를 지니고있다. 재래종 자원은 자연변이에 의해 선택되어 현대 품종 개선과 관련된 높은 잠재성을 가지고 있다. 이러한 잠재성을 이용하기 위한 첫 번째 단계는 유용 자원을 선발하는 과정이다. 본 연구에서는 벼 직파재배 적응성을 확대할 수 있는 중배축 신장성에 대해 국내 재래종 벼 유전자원 386점에 대해 평가하였다. 직파재배는 쌀 생산 단가와 노동력 감소에 효과적인 재배 방법으로 대두되고 있으나, 잡초경합과 초기 입모율 확보의 어려움이 있다. 중배축 신장성은 초기 입모율 확대에 큰 영향을 미치는 요인으로 직파재배 확대에 중요한 형질이다. 본 연구에서 암조건에서 평가된 재래종 유전자원의 중배축 길이는 0 mm~28 mm의 범위를 보였다. 특히 16 mm 이상의 중배축 신장성이 큰 재래종 유전자원 20점(5%)을 선발하였고, 이들의 출수기, 간장, 주당 분얼 수 등 기본 농업 형질과 종실 특성 및 까락 여부, 생태형 등을 검토하여 유용 육종소재를 제공하였다.
KJBS-51-351_Supple.pdf
이 논문은 정부의 재원으로 한국연구재단의 지원을 받아 수행된 기초연구사업(NRF-2017R1E1A1A01075282, NRF-2015R1C1A1A01054699)의 성과임.

보충자료

본문의 Supplementary Table 1은 한국육종학회지 홈페이지에서 확인할 수 있습니다.

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Evaluation of Mesocotyl Elongation Ability in Korean Rice Landraces (Oryza sativa L.)
Korean. J. Breed. Sci.. 2019;51(4):351-356.   Published online December 1, 2019
DOI: https://doi.org/10.9787/KJBS.2019.51.4.351
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Evaluation of Mesocotyl Elongation Ability in Korean Rice Landraces (Oryza sativa L.)
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DOI: https://doi.org/10.9787/KJBS.2019.51.4.351
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Evaluation of Mesocotyl Elongation Ability in Korean Rice Landraces (Oryza sativa L.)
Image Image Image
Fig. 1 Methods of evaluating mesocotyl elongation. (A) 20 rice seeds inside of the agarose gel for each magenta box. (B) Dark treatment by covering aluminum foil for 14 days.
Fig. 2 Measuring of Mesocotyl length of the control varieties. (A) a low vigor control variety ‘Nipponbare’, (B) a high-vigor control variety ‘Kasalath’
Fig. 3 Distribution of mesocotyl length of 386 Korean rice landrace.

Fig. 1

Fig. 2

Fig. 3

Evaluation of Mesocotyl Elongation Ability in Korean Rice Landraces (Oryza sativa L.)

Some agricultural traits of high mesocotyl elongation ability rice landrace.

IT Number Variety Mesocotyl Length (mm) Grain length (mm)Z Grain width (mm)Z Awny Heading Datey Culm Length (cm)y Panicle Length(cm)y Panicles/ planty Lodgey Leaf Blasty Amylose Contentsy
IT007486 Anjeunbaengi 28.0 6.8 3.3 None 8.25 70 24 10 Rx - 0.19
IT010555 Yukseongjaerae 26.5 7.0 3.4 None 8. 2 88 - 8 R M 0.25
IT006302 Baekgokna 24.5 7.3 3.5 Short/Few 8. 6 110 23 5 M S 0.06
IT005994 Deokjeokjodo 24.5 7.0 3.6 Existence 7.31 101 21 12 M M 0.17
IT173446 Ginggalagsyale 24.5 7.3 2.8 None 8. 3 114 22 - - 0.21
IT009060 Hanyangjo 23.0 7.8 3.4 None 8.14 102 23 13 R S 0.31
IT009187 Hwangtodo 20.5 6.4 3.5 Existence 8.16 95 20 6 M M 0.19
IT010627 Joseokjo 20.0 7.4 3.6 Short/Few 7.29 92 21 8 R S 0.16
IT010374 Sando 20.0 7.8 3.5 Existence 8. 5 95 21 7 R S 0.18
IT005044 Guwangdo 19.0 6.2 3.4 Short/Lot 8.27 107 30 8 M - 0.19
IT010612 Jodo 19.0 7.2 3.6 None 8. 2 100 20 6 R S 0.17
IT006556 Sando 19.0 7.2 3.6 Existence 8. 5 83 22 7 M S 0.18
IT006596 Samkyeongjo 19.0 7.1 3.4 Short/Lot 8. 5 85 17 12 S - 0.24
IT008700 Jwiipparibyeo 18.5 6.9 3.4 Short/Few 8.12 99 20 15 S - 0.17
IT005504 Noinjo 17.5 6.9 3.4 Existence 8.19 109 22 8 M S 0.17
IT006372 Baekjo 17.0 6.9 3.6 Existence 8.16 99 24 10 M S 0.18
IT008749 Chalbyeo 16.5 7.7 3.3 Short/Lot 8. 6 92 24 10 M S 0.06
IT007807 Yue 16.0 6.7 3.5 Short/Lot 8.19 116 24 16 S - 0.17
IT006078 Monajo 16.0 6.9 3.6 Short/Lot 8.16 93 20 8 S S 0.05
IT010340 Batnarak 16.0 7.2 3.6 None 8. 7 97 20 9 M S 0.22
Table 1 Some agricultural traits of high mesocotyl elongation ability rice landrace.

Evaluation average of 20 seeds measured by Vernier calipers

These data acquired from the database of National Agrobiodiversity Center (http://genebank.rda.go.kr/simpleSearch.do)

R: Resistant, M: Moderate, S: Susceptible

Table 1

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