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��니카 다수성 벼 드래찬과 보람찬 유래 약배양 계통� �용한 �삭 � 수량 관련 형질 분� � 다수성 계통 육성

Development of High-yielding Rice Lines and Analysis of Panicle and Yield-related Traits Using Doubled Haploid Lines Derived from the Cross between Deuraechan and Boramchan, High-yielding japonica Rice Cultivars in Korea

Korean Journal of Breeding Science 2015;47(4):384-402.
Published online: November 30, 2015

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2 Department of Central Area, NICS, RDA, Suweon 16429, Korea

3 충남대학� �업�명대학

3 Department of Agronomy, Chungnam National University, Daejeon 34134, Korea

*Corresponding author (mayoe@korea.kr, Tel: +82-63-238-5202, Fax: +82-63-238-5205)
• Received: October 12, 2015   • Revised: November 18, 2015   • Accepted: November 18, 2015

© Korean Society of Breeding Science All rights reserved

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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  • A doubled haploid (DH) lines derived from the cross between high-yielding japonica rice cultivars ‘Deuraechan’ and ‘Boramchan’ was developed to increase diversity of panicle and yield-related traits and select high-yielding lines. Panicle and yield-related traits of DH population consisting of 163 lines and 13 selected high-yielding lines were analyzed using correlation, principal component, and path analysis. Among panicle-related traits, number of spikelets on secondary rachis-branches (SRBs) was more highly correlated with and had more positive effect on number of spikelets (NS) than number of spikelets on primary rachis-branches (PRBs) per panicle. In the DH population, NS had the most positive effect on yield, whereas panicle number per hill (PN) was in the selected lines. PN was significantly positive correlated with ratio of ripened grain (RGG) in both case. The enhancement of ripening and increase of total spikelets number by increasing PN had more effect on yield than increase of total spiekelts number by increasing NS in the selected lines. Among 13 lines, four lines, AC60, AC152, AC156, and AC161 showed higher yield than Deuraechan (481 kg/10a) and Boramchan (558 kg/10a). Deuraechan exhibited panicle-weight type with low tiller and heavy panicles and Boramchan was slightly panicle-number type with more PN and higher RGG. Selected high-yielding lines showed medium characteristics of parents with improved yield potential. These elite lines could be utilized to develop high-yielding japonica rice.
벼 육종� 있어서 수량성 향�� 가장 중요한 목표�다. 벼 � 수량성� �물체� 광합성� �해 �화산물� �성�고 전류�는� 관여하는 능력� source와 종실� 축�� 수 있는 잠재력� sink� �해 결정�다. Source와 sink� �기와 � 들� 균형� �해 수량성� 결정�는� sink가 작으면 높� 수량� 기대하기 어렵고, sink� �기가 �다 해� source가 �를 뒷받침해주지 못하면 수량� 향��지 않는다. Source 는 �물체� 광합성 능력, 초형� �한 수광태세, 엽노화� 등 � �해 �향� 받고 sink� �기는 �게 주당수수와 수당립 수� �한 단위면�당 립수와 종실� �기� �해 결정�며 소수가 종실로 �는 임실률과 종실� 등숙률 � 종실중� � 해 수량� 결정�다(Hirota et al. 1990, Li et al. 1998, Park et al. 2011, Venkateswarlu & Visperas 1987).
벼 �삭� 기본구조는 수축�서 분지한 지경� 소수가 착� �어 있는 것으로 지경� 분지 체계, 지경� 종류와 수, 착� 립� 수와 밀�, �삭 내 위치� 따른 착�립� 분� 등� � 해 �삭� 형태가 구성�다(Park et al. 2010). �삭� 형태� 특성� �태형 � 품종간 다양한 변�를 나타낸다. Matsuo (1952)는 ��니카형� 열대 ��니카나 �디카� 비해 �대 �으로 �삭� 짧고 무거우며 착�밀�가 높� 반면 �디카형 � 다른 �태형보다 � 많� 지경� 가지고 있다고 하였고, Manaka & Matsushima (1971)는 대부분� ��니카 벼 품종 � 1차 지경 착�립수는 5개 �는 6개� 반면 �디카 품종� 다양한 변�를 나타낸다 하였다. Taguch-Shiobara et al. (2011)는 136개 �디카와 156개 ��니카 품종� �삭형질� 대한 주성분 분��서 �디카가 ��니카� 비해 수축� 짧고 1차 지경� 성기게 발�하는�, 1차 지경� 길�가 � 길고 1 차 지경�서 착�� 립과 1차 지경�서 분지한 2차 지경 � 착�립수가 많� �성�며, 2차 지경�서 � 많� 지경�나 착�립� �성한다고 하였다. Park et al. (2010)� 178개 우 리나� 벼 품종� 대�으로 �삭관련 형질� 분류하였는� 통 �형� ��니카 형보다 �삭 �기가 �며 �는 2차 지경수가 많고 2차 지경 착�립수가 많기 때문�� 하였고 2차 지경 � 착�립수가 1차 지경 � 착�립수� 비해 수당립수� �향� 많� 미친다고 하였다.
소수와 지경으로 구성� �삭� 특성� 수당립수를 결정하 고, 등숙률과 종실중� �향� 주는 등 수량성과 밀접한 관련 � 가지고 있어 �삭형질� 벼 육종가�게 중요한 목표 형질 중� 하나�다(Park et al. 2011). 우리나��서� ��니카와 �디카 �배를 통한 통�형 벼 개발, �본� 초다수성 벼 육 종사업, 국제미작연구소� NPT (New Plant Type) 초형, 중 국� F1 hybrid 등 육종� 방법으로 �삭� �기를 늘림으로 � 수량성 향�� 위한 다양한 접근� 시��어 왔으며 여러 품종 � 계통� 육성�었다(Chung & Heu 1980, Kushibuchi 1997, Peng et al. 2008, Yuan 1997). �삭� �기와 무게를 �대시키는 것� 중요하지만 sink �기� �대는 �반�으로 등숙률 �소� �향� 미치고 �삭내 소수� 위치와 지경� 구조� 따� 등숙과 성숙 패턴, 광합성 �화산물� 분배와 최 종 중량� 차�가 나게 �며 �는 1차 지경립� 2차 지경립보 다, �위 지경립� 하위 지경립보다 등숙� 잘�기 때문� � 삭지경� 분지 체계� 대한 고려� 중요하다(Kim et al. 2003, Mo et al. 2012, Nagata et al. 2002, Park et al. 2010, Yamagishi et al. 2003).
�삭형질과 관련� 형질 특히 수당립수� 대한 많� 유전 � � QTL� 보고�었다. �디카 Habataki 품종�서 유래한 Gn1a (Grain number 1a)는 1번 염색체 단완� 위치하면서 사 �토카닌 산화효소를 부호화하며, Gn1a� 발현� 줄어들게 �면 �삭 분열조�� 사�토카닌� 축��고 수당립수가 � 가하게 �다(Ashikari et al. 2005). DEP1 (DENSE AND ERECT PANICLE1)� 9번 염색체� 존재하며 대립유전�� �단 부위 � 절단 변�가 발�하여 phosphatidylethanolamine-binding protein-like domain protein� 기능� �실하게 �면 수당립 수가 �가� 고밀� �립 �삭형태를 가져오게 �다(Huang et al. 2009). 중국 특히 ��지역� ��니카형 다수확 품종 중� 많� 수가 dep1� 가지고 있는�, 1970년대 �탈리아� 서 광범위하게 재배�었� 재래종 Balilla가 1958년� 중국� �입�었고 �를 �용해 개발� Shennong 265를 비롯하여 ��니카 �립 �는 반�립 다수확 품종�서 dep1� 있는 것으로 확��었다(Huang et al. 2009). WFP (WHEALTHY FARMER’S PANICLE)와 IPA1 (IDEAL PLANT ARCHITECTURE)� �� 보 고�었는� 나중� 8번 염색체 장완� 위치하는 같� QTL로 �혀 졌고 부호화하는 유전�는 전사��� OsSPL14 (SQUAMOSA PROMOTER BINDING PROTEIN-LIKE 14)로 WFP는 �디 카 ST-12 품종�서 IPA1� ��니카 Shaoniejing 계통�서 유 래하였다(Jiao et al. 2010, Miura et al. 2010). 초기 ��� 장기� OsSPL14WFP� 과발현� �삭� 지경분화를 촉진시키 는� 특히 1차 지경� 수가 �가하여 수당립수가 �가하게 � 고, OsSLP14ipa1� �양�장기� �소성 발현� 나타내고 � �성장기� 과발현� 나타내는� OsSPL14WFP와 마찬가지로 1차 지경 �가� 따른 수당립수가 �가하나 수수가 줄어들게 �고 줄기가 강해지게 �는� �는 아마 �양�장기� �소 성 발현� 기�한 것으로 ���다 하였다(Miura et al. 2011). APO1 (ABERRANT PANICLE ORGANIZATION 1) � �디카 Habataki� 6번 염색체 QTL� PBN6 (for primary branch number on chromosome 6) �서 �정� 유전�로 1 차 지경� �가시킴으로� 수당립수가 �가�고 수량� �가 하나 수수나 등숙률� �소가 없다고 하였으며, APO1� 발현 � 수당립수� �가 �만 아니� 유관�수� �가를 가져와 �화산물 전류효율� 높여 등숙� 잘�게 한다고 하였다 (Terao et al. 2010). �� 반해 Gn1� 경우 수당립수가 �가 하는� 주로 2차 지경� �가� �한 것으로 �는 같� 지경 내 종실� �화산물 전류� 경합� 가져와서 등숙� 악�향 � 가져온다 하였다(Nagata et al. 2002).
�촌진�청 국립�량과학��서는 ��니카 초다수성 벼 품종으로 준조�종� 남�(2003), 중만�종� 한마�(2004), 드래찬(2008), 보람찬(2009), ��찬(2011년) 등 5개 품종� 개발하였다(Won & Oh 2014). �들 품종들� 수량성� 다비 밀� 조건�서 육성 당시 662 (남�), 643 (한마�), 652 (드 래찬), 733 (보람찬), 618 (��찬) kg/10a 으로 ��니카형 표준 � 비� 품종� 비해서 높� 수량성� 나타냈다. 남�과 드래찬� 소얼 수중형으로 �복� 강하고 수당립수를 �대시 킴으로� 수량성� 향�시키는 방향으로 육성� 품종�고, 한 마�� 립형� �게하고 1차 지경수 � 착�립수를 �가시켜 1차 지경당 2차 지경수 � 착�립수� 비율� 줄임으로� 립 중� �대와 등숙률� �시� 향�시킴으로� 다수성� �달 한 품종�다(Ha et al. 2009, Kim et al. 2003). ��찬� 간 장� 키워 다소 키가 �고 수당립수가 한마�� 비해 많으나 내�복성� 바탕으로 립중� �가시킴으로� 수량성� �대 하고� 육성한 품종�다(Ha et al. 2014). 보람찬� 지금까지 육성� ��니카 품종 중�서 육성 당시 수량성� 가장 높� 품종으로 수수가 �대�으로 다른 품종� 비해 많� 초형으 로 등숙률� 높고 내�복성과 내병성� 함께 갖추고 있다(Ha et al. 2012). 하지만 잠재 수량성� 733 kg/10a� 달하는 보 람찬� 2013년 �가 �산성� 600 kg 수준으로 82% 수준� 었으며 지역별로 수량성 변�� 커 연차 � 지역간 변�와 질 소 시비 조건 등� �해 수량성 변�� � 것으로 나타났다 (Sang et al. 2014).
본 연구는 ��니카 초다수성 품종으로 다수성� 관여하는 형질 특성� 다른 드래찬� 모본으로 보람찬� 부본으로 � 용하여 �공�배와 약배양� 통해 �삭 � 수량 관련 형질 변 �를 �대시킨 고정 집단� 육성하고, 보통기 �반재배�서 집단 내�서 �삭 � 수량 관련 형질들� 특성� 분�하여 다 수성 우량 계통� 선발하고 선발� 계통� 특성� 분�하여 ��니카 수량성 향�� 위한 벼 육종사업� 반�하고� 수 행하였다. Table 1
Table 1.
The abbreviations for panicle and yield-related traits.
Table 1.
Abbreviations:

PL Panicle length
TNN Thickness of neck node
PRB Primary rachis-branch
SRB Secondary rachis-branch
NR Number of rachis-branches per panicle
NS Number of spikelets per panicle
NPr Number of PRBs per panicle
NSPr Number of spikelets on PRBs per panicle
MNSPr Mean number of spikelets on a PRB
NSr Number of SRBs per panicle
NSSr Number of spikelets on SRBs per panicle
MNSSr Mean number of spikelets on a SRB
Sr/Pr Ratio of SRBs to PRBs
SSr/SPr Ratio of spikelets on SRBs to spikelets on PRBs
CL Culm length
FL Flag leaf length
PN Panicle number per hill
TGW 1,000 grain weight
RRG Ratio of ripened grain
BRR Brown/rough rice ratio
RRY Rough rice yield
BRY Brown rice yield
MRY Milled rice yield
DAS Day after seedling
시험재료 � 재배방법
�촌진�청 국립�량과학��서 ��니카형 초다수 벼 품 종으로 개발� 드래찬(Ha et al. 2009)과 보람찬(Ha et al. 2012)� 모부본으로 하여 �공�배� F1�물체를 �용하여 약배양� 수행하였다(Table 2). 약배양 유래 220개 계통� 2011년 국립�량과학� �산 벼 시험�장� 재�거리 30 × 15 cm로 계통당 1주 1본으로 30주씩 �앙하였고 � 중 출수 기가 모부본과 비슷하고 �업형질� 양호한 163개 계통� 선 발하여 �삭 � 수량 관련 형질� 대해서 조사하였다. 2012년 선발� 163개 계통과 모부본� 3반복으로 재�거리 30 × 15 cm로 1주 1본씩 30주씩 �앙하여 �삭 � 수량 관련 형질� 대해서 조사하였다. 2013년 모부본과 선발� 13개 다수성 계 통� 대해서 재�거리 30 × 15 cm로 주당 3본씩 구당 180주 를 3반복으로 �앙하여 �산력 검정시험� 수행하였다. 2014 년� 모부본과 최종 선발� 4개 우량계통� 대해서 2013년과 ��하게 �산력 검정시험� 수행하여 수량성 � �업형질� 대해서 확�하였다. 시비량� N-P2O5-K2O를 90-45-57 kg/ha 으로 질소는 기비 : 분얼비 : 수비를 50 : 20 : 30 비율로 분시 하였고, �산� 전량 기비로, 칼륨� 기비 : 수비를 70 : 30 비율로 분시하였다. 기타 재배관리는 �촌진�청 표준 재배법 � 준하여 실시하였다.
Table 2.
Anther culture ability and doubled haploid lines derived from the cross of Deuraechan/Boramchan.
Table 2.
Anther cultured (no.) Cali induced (no. and %) Plant regeneration (no. and %)z Green plants (no. and %)y Albino (no. and %)x Doubled haploid lines (no.)w

10,600 3,528(33.3) 450(12.8) 245(54.4) 215(45.6) 220

zPercent plant regeneration is the number of plants regenerated over number of cali plated

yPercent green is the number of green plants regenerated over total plants regenerated

xPercent albino is the number of albino plants regenerated over total plants regenerated

wDouble haploid lines are harvested plants having fertile grain

�업형질 조사
2011, 2012년� 수행� 시험� 대해서 공시 재료� 출수기 를 조사하고 계통 당 �균� �는 5주� 대해서 간장, 수장, 지엽장 � 수수를 실측하였다. �삭관련 형질� 황숙기 �후 � � 계통� 3주� 주간�서 �삭� 채취하여 �삭길�, � 삭목�께, 한 �삭�서 분지한 전체 지경� 수(�지경수), 한 �삭� 착�한 전체 종실� 수(�립수; 수당립수), 1차 지경 수, 1차 지경� 착�한 전체 종실� 수(1차 지경 착�립수), 1차 지경� 착�한 종실� �균립수(1차 지경 �균립수), 2차 지경수, 2차 지경� 착�한 전체 종실� 수(2차 지경 착�립 수), 2차 지경� 착�한 종실� �균립수(2차 지경 �균립수), �삭 내 1차 지경수� 대한 2차 지경수 비율(1차 지경당 2차 지경수), �삭 내 1차 지경 착�립� 대한 2차 지경 착�립� 비율(1차 지경 착�립수당 2차 지경 착�립수)� 조사하였다. 천립중과 등숙률� 3주를 예취하여 간� 천립중과 등숙률� 조사하였고, 수량� 20주를 예취하여 정조 간�수량� �용 하였다. 2013, 2014년� 수행� �산력 검정시험�서는 출수 기를 조사하고 성숙기� 시험구별로 10개체� 대해서 간장, 수장, 수수를 측정하였다. 출수 후 50�� 3주를 예취하여 등 숙률 � 수당립수를 측정하고, 수량� 100주를 예취하여 정 조수량� 측정한 다� � 중 1.5 kg� 대해서 수량조사현미기 (LST, Gwangyang, Korea)로 제�하여 정현비율� 측정하 고, 100주 정조수량� 정현비율� 곱하여 현미수량� 구하고 10a당 수량으로 환산하였다. 현미수량� �반� 현백률� 0.92를 곱하여 백미수량� 구하였고 제현� 현미를 가지고 천립중� 측정하였다. 2013년� 공시� 재료� 대해서 수확 기� 3주를 수확하여 모든 �삭� 대해서 2011, 2012년� 조 사한 �삭관련 형질� 조사하여 형질 값으로 �용하였다.
통계 분�
통계분�� SAS 프로그램(Version 9.2, SAS Institute Inc., Cary, NC, USA)� �용하였다. �업형질� 대한 �균� 기술 통계법으로 구하였고, �균간 비�는 PROC ANOVA로 분산 분� 후 유�성� 있� 경우 5% 유�수준�서 Duncan’s Multiple Range test (DMRT)로 분�하였다. PROC CORR로 �관분�� 실시하여 형질들� 관련성� 알아보고 �들 형질 � 공시재료� 구조� 관계를 분�하고� PROC PRINCOMP 로 주성분분�� 수행하였다. �삭 � 수량 관련 형질 값� � 용하여 PROC CLUSTER� �균연결법으로 군집분�� 수행 하여 공시재료를 분류하였다. 수량성� 대한 형질들� 기여� 를 알아보기 위해서 PROC TCALIS로 경로계수분�� 수행 하였다.
약배양 유래 집단�서� �삭 � 수량 관련 형질 분�
2011, 2012년� 드래찬과 보람찬 � 163개 약배양 계통으 로 구성� 집단� 대해서 �삭길�(PL; x1), �삭목�께 (TNN; x2), �지경수(NR; x3), �립수(NS; x4), 1차 지경수 (NPr; x5), 1차 지경 착�립수(NSPr; x6), 1차 지경 �균립수 (MNSPr; x7), 2차 지경수(NSr; x8), 2차 지경 착�립수 (NSSr; x9), 2차 지경 �균립수(MNSSr; x10), 1차 지경당 2 차 지경수(Sr/Pr; x11), 1차 지경 착�립수당 2차 지경 착�립 수(SSr/SPr; x12), 간장(CL; x13), 지엽장(FL; x14), 수수 (PN; x15), 천립중(TGW; x16), 등숙률(RRG; x17) � 수량 (Yield; x18)� 측정하였다. 약배양 유래 집단� 대해서 조사 � 18개 형질들간� �관관계를 분�하였다(Table 3). 집단 내�서 �삭길�(x1)는 �삭목�께(x2)와 정� �관(r=0.547; 2011, 0.752; 2012년)� 나타냈고 �지경수(x3; 0.472, 0.706)와 �립수(x4; 0.515, 0.659)와 정� �관� 나타내었 으며 1차 지경 관련 형질(x5, x6, x7)� 2차 지경 관련 형질 (x8, x9, x10)보다 높� �관� 나타냈다. 지엽장(x14; 0.556, 0.549)과 현미천립중(x16; 0.381, 0.314)과는 정� �관� 나 타냈고 수수(x15; -0.392, -0.807)와 등숙률(x17; -0.210, -0.329)과는 부� �관� 나타냈다. �삭목�께(x2)는 �지경 (x3; 0.858, 0.833)과 �립수(x4; 0.884, 0.846)와 높� 수준 � 정� �관� 나타냈으며 2차 지경 관련 형질(x8, x9, x10) � 1차 지경 관련 형질(x5, x6, x7)보다 높� �관� 나타냈 다. 지엽장(x14; 0.513, 0.432)과는 정� �관� 수수(x15; -0.325, -0.762)와는 부� �관� 나타냈다. 지경과 착�립수 와� 관련성�서 2차 지경 관련 형질(x8, x9, x10)� 1차 지 경 관련 형질(x5, x6, x7)보다 �지경수(x3), �립수(x4)와 높 � 정� �관� 나타냈으며 1차지경당 2차지경 형질� 대한 비율� 1차 지경당 2차 지경수(x11)와 1차 지경 착�립수당 2차 지경 착�립수(x12)� �지경수 � �립수와 �관계수 0.770�서 0.862사�� 높� 정� �관� 나타냈다. 간장 (x13)� 2011년�는 천립중(x16; 0.386)과 정� �관� 나타 냈으나 다른 형질들과는 �관� �� 편�었으며, 2012년� 천립중(x16; 0.322)과 등숙률(x17; 0.317)과는 정� �관� 나타냈고 2차 지경 �균립수(x10; -0.424), 1차 지경당 2차지 경수(x11; -0.415), 1차 지경 착�립수당 2차 지경 착�립수 (x12; -0.376)와는 부� �관관계를 나타냈다. 지엽장(x14)� �삭길�(x1; 0.556, 0.549)와 가장 �관� 높았으며 지경과 착�립 관련 형질들과 모� 정� �관� 나타냈으며 그 중 1 차 지경 관련 형질(x5, x6, x7)� 2차 지경 관련 형질(x8, x9, x10)보다 �관� 높았다. 수수(x15)는 통계�으로 유�한 수 준�서2012년 등숙률(x17; 0.378)� 제외한 다른 형질들과 부� �관� 나타냈으며 1차 지경 관련 형질(x5, x6)� 2차 지경 관련 형질(x8, x9)보다 높� 부� �관� 나타냈다. 천 립중(x16)� 등숙률(x17; 0.405, 0.458), 간장(x13; 0.386, 0.322), �삭길�(x1; 0.381, 0.314)와 정� �관� 나타냈으 며 1차 지경 관련 형질(x5, x6)과는 낮� 수준� 정� �관� 나타냈으나 2차 지경 관련 형질(x8, x9)과는 관련성� 없었 다. 등숙률(x17)� 지경과 착�립 관련 형질들과 대부분 부� �관관계를 나타냈는� 특히 2차 지경 관련 형질(x8, x9, x10)� 1차 지경 관련 형질(x5, x6, x7)보다 높� 수준� 부 � �관� 나타냈고 1차 지경당 2차 지경 형질� 대한 비율� 1차 지경당 2차 지경수(x11)와 1차 지경 착�립수당 2차 지 경 착�립수(x12)� 부� �관관계를 나타냈다. 등숙률과 수 량(x18; -0.169, -0.323)과는 부� �관관계를 나타냈는� 그 수준� 낮았으며, 천립중(x17; 0.405, 0.458)과는 정� �관 � 수수(x15)와는 2012년� 정� �관� 나타냈다. 수량 (x18)� �삭목�께(x2; 0.376, 0.261)와 정� �관� 나타냈 으며 지경과 착�립 관련 형질들과 정� �관� 나타냈으며 2차 지경 관련 형질(x8, x9, x1)� 1차 지경 관련 형질(x5, x6, x7)보다 �관� 높았고 1차 지경당 2차 지경 형질� 대한 비율� 1차 지경당 2차 지경수(x11)와 1차 지경 착�립수당 2차 지경 착�립수(x12)� 정� �관관계를 나타냈다.
Table 3.
Correlation coefficient among panicle and yield related traits in 2011 and 2012 (n=163).
Table 3.
Traitsz PLy TNN NR NS NPr NSPr MNSPr NSr NSSr MNSSr Sr/Pr SSr/SPr CL FL PN TGW RRG Yield
(x1) (x2) (x3) (x4) (x5) (x6) (x7) (x8) (x9) (x10) (x11) (x12) (x13) (x14) (x15) (x16) (x17) (x18)

x1 0.547** 0.472** 0.515** 0.544** 0.589** 0.354** 0.415** 0.396** 0.229** 0.241** 0.209** 0.209** 0.556** -0.392** 0.381** -0.210** 0.198*
x2 0.752** 0.858** 0.884** 0.655** 0.674** 0.294** 0.816** 0.803** 0.549** 0.629** 0.602** 0.005ns 0.513** -0.325** 0.159* -0.313** 0.376**
x3 0.706** 0.833** 0.987** 0.571** 0.571** 0.188* 0.988** 0.965** 0.587** 0.859** 0.823** -0.052ns 0.412** -0.302** 0.044ns -0.364** 0.468**
x4 0.659** 0.846** 0.990** 0.609** 0.630** 0.286** 0.967** 0.958** 0.640** 0.820** 0.793** -0.039ns 0.442** -0.297** 0.059ns -0.378** 0.456**
x5 0.794** 0.765** 0.765** 0.817** 0.966** 0.218** 0.437** 0.367** 0.002ns 0.075ns 0.031ns 0.194* 0.476** -0.343** 0.395** -0.156* 0.118ns
x6 0.803** 0.745** 0.746** 0.812** 0.979** 0.462** 0.443** 0.380** 0.050ns 0.093ns 0.033ns 0.224** 0.515** -0.347** 0.388** -0.170* 0.126ns
x7 0.561** 0.424** 0.423** 0.516** 0.579** 0.732** 0.165* 0.169* 0.185* 0.086ns 0.009ns 0.171* 0.325** -0.128ns 0.118ns -0.098ns 0.07ns
x8 0.594** 0.799** 0.991** 0.969** 0.674** 0.656** 0.367** 0.988** 0.643** 0.926** 0.896** -0.094ns 0.361** -0.266** -0.026ns -0.370** 0.491**
x9 0.580** 0.792** 0.982** 0.966** 0.649** 0.633** 0.360** 0.994** 0.744** 0.942** 0.933** -0.130ns 0.335** -0.226** -0.073ns -0.388** 0.497**
x10 0.401** 0.610** 0.739** 0.759** 0.418** 0.417** 0.281** 0.762** 0.821** 0.713** 0.785** -0.267** 0.180* 0.010ns -0.283** -0.347** 0.382**
x11 0.298** 0.590** 0.825** 0.770** 0.288** 0.256** 0.058ns 0.888** 0.907** 0.806** 0.986** -0.182* 0.204** -0.164ns -0.194* -0.354** 0.495**
x12 0.346** 0.620** 0.862** 0.807** 0.338** 0.324** 0.161* 0.921** 0.927** 0.757** 0.986** -0.222** 0.166* -0.119ns -0.234** -0.369** 0.492**
x13 0.061ns -0.160* -0.221** -0.194ns 0.060ns 0.079ns 0.114ns -0.266** -0.293** -0.424** -0.415** -0.376** -0.075ns -0.047ns 0.386** 0.045ns -0.100ns
x14 0.549** 0.432** 0.413** 0.421** 0.420** 0.426** 0.289** 0.388** 0.369** 0.189* 0.237** 0.279** -0.006ns -0.418** 0.285** -0.163* 0.241**
x15 -0.807** -0.762** -0.733** -0.750** -0.726** -0.713** -0.417** -0.693** -0.678** -0.498** -0.478** -0.515** 0.148ns -0.548** -0.361** 0.091ns 0.142ns
x16 0.314** 0.043ns -0.088ns -0.055ns 0.200* 0.194* 0.107ns -0.142ns -0.158* -0.252** -0.310** -0.291** 0.322** 0.375** -0.180* 0.405** -0.080ns
x17 -0.329** -0.398** -0.538** -0.527** -0.321** -0.308** -0.162* -0.552** -0.562** -0.511** -0.550** -0.545** 0.317** -0.164* 0.378** 0.458** -0.169*
x18 0.196* 0.261** 0.410** 0.413** 0.270** 0.271** 0.187* 0.415** 0.427** 0.427** 0.402** 0.400** -0.14ns 0.254** -0.202** 0.051ns -0.323**

zUpper and lower diagonal indicate the value in 2011 and 2012, respectively

yAbbreviations are listed in table 1

주성분 분�� 통해 형질들� 차�� 줄�고 구조� 관계 를 분�하였다(Table 4, Fig. 1). 축소� 형질들� 주성분 1� 전체 형질 특성� 46.0(2011)와 55.3% (2012년)� 설명할 수 있으며 주성분 2는 18.9(2011)와 16.6%(2012년)� 설명 할 수 있어 �들 2개 주성분으로 전체 형질� 64.8 (2011)와 71.8%(2012년)� 설명할 수 있었다. 주성분 1� �립수 (r=0.987; 2011, 0.987; 2012년), �지경수(0.975, 0.979), 2 차 지경수(0.964, 0.963), 2차 지경 착�립수(0.960, 0.962), �삭목�께(0.895, 0.870), 1차 지경 착�립수당 2차 지경 착 �립수(0.816, 0.820), 1차 지경당 2차 지경수(0.841, 0.787), 2차 지경 �균립수(0.676, 0.779)와 높� 정� �관� 나타냈 고 1차 지경 착�립수(0.582, 0.781), 1차 지경수(0.554, 0.786), �삭길�(0.567, 0.743), 지엽장(0.511, 0.484), 간� 수량(0.507, 0.446)와 중간 수준� 정�관� 나타냈다. 등숙 률과는 �관계수 -0.425(2011년)와 -0.580(2012)로 중간 수 준� 부� �관� 나타냈고 수수와는 2011년� �관계수 -0.333으로 낮� 부� �관� 나타냈는� 2012년�는 -0.801 로 높� 부� �관� 나타냈다. 주성분 2는 현미천립중 (0.726, 0.718)과 높� 정� �관� 나타냈으며 �삭길� (0.511, 0.516), 간장(0.472, 0.570), 지엽장(0.436, 0.381)과 는 중간 수준� 정� �관� 나타냈다. �지경수와 �립수와 는 �관 관계가 없었고 1차 지경 관련 형질� 1차 지경수 (0.680, 0.484), 1차 지경 착�립수(0.709, 0.523), 1차 지경 �균립수(0.352, 0.476)와는 정� �관� 나타낸 반면 2차 지 경 관련 형질� 2차 지경수(-0.158, 0.171), 2차 지경 착�립 수(-0.235, -0.209)와는 거� �관관계가 없었으며 2차 지경 �균립수(-0.465, -0.366)와는 낮� 수준� 부� �관� 나타 냈다. 1차 지경당 2차 지경 형질� 대한 비율� 1차 지경당 2차 지경수(-0.456, -0.545)와 1차 지경 착�립수당 2차 지경 착�립수(-0.525, -0.474)와는 중간 수준� 부� �관� 나타 냈다. 주성분 분��를 통해 형질들� 구조� 관계를 보면 주 성분 1� 기준으로 지경 � 착�립수 관련 형질� �지경수, �립수, 1�2차 지경수 � 착�립수, �균립수 � 비율과 � 삭목�께, �삭길�, 지엽장 � 간�수량� 수수와 등숙률과 다른 방향성� 나타내고 있으며, 주성분 2를 기준으로는 천립 중, 1차 지경 관련 형질(지경수, 착�립수, �균립수), �삭길 �, 지엽장, 간장, 등숙률� 1차 지경당 2차 지경 형질� 대한 비율(1차 지경당 2차 지경수, 1차 지경 착�립수당 2차 지경 착�립수)과 2차 지경 �균립수 � 수수와 다른 방향성� 나 타냈다.
Table 4.
Eigenvectors for the principal components of traits and correlation coefficient between the traits and principal components.
Table 4.
Trait 2011 2012

Eigenvector Correlation coefficient Eigenvector Correlation coefficient

Prin1z Prin2 Prin1 Prin2 Prin1z Prin2 Prin1 Prin2

PLy 0.197 0.277 0.567** 0.511** 0.236 0.299 0.743** 0.516**
TNN 0.311 0.102 0.895** 0.189* 0.276 0.092 0.870** 0.159*
NR 0.339 -0.015 0.975** -0.028ns 0.310 -0.036 0.979** -0.062ns
NS 0.343 0.012 0.987** 0.023ns 0.313 0.010 0.987** 0.017ns
NPr 0.193 0.369 0.554** 0.680** 0.249 0.280 0.786** 0.484**
NSPr 0.202 0.385 0.582** 0.709** 0.248 0.303 0.781** 0.523**
MNSPr 0.102 0.191 0.292** 0.352** 0.158 0.276 0.498** 0.476**
NSr 0.335 -0.086 0.964** -0.158* 0.305 -0.099 0.963** -0.171*
NSSr 0.334 -0.128 0.960** -0.235** 0.305 -0.121 0.962** -0.209**
MNSSr 0.235 -0.252 0.676** -0.465** 0.247 -0.212 0.779** -0.366**
Sr/Pr 0.292 -0.248 0.841** -0.456** 0.249 -0.316 0.787** -0.545**
SSr/SPr 0.284 -0.285 0.816** -0.525** 0.260 -0.275 0.820** -0.474**
CL -0.023 0.256 -0.067ns 0.472** -0.079 0.330 -0.249** 0.570**
FL 0.178 0.236 0.511** 0.436** 0.153 0.220 0.484** 0.381**
PN -0.116 -0.235 -0.333** -0.434** -0.254 -0.167 -0.801** -0.288**
TGW 0.016 0.394 0.045ns 0.726** -0.017 0.416 -0.053ns 0.718**
RRG -0.148 0.110 -0.425** 0.203** -0.184 0.215 -0.580** 0.370**
Yield 0.176 -0.133 0.507** -0.245** 0.141 -0.072 0.446** -0.125ns
Eigen value 8.27 3.40 9.95 2.98
Contribution (%) 46.0 18.9 55.3 16.6
Cumulative contribution (%) 46.0 64.8 55.3 71.8

zPrin1 and Prin2 indicate the first and second component, respectively

yAbbreviations are listed in table 1

Fig. 1.
Relationship among traits in the principal component analysis (PCA) diagram composed of component 1 and 2 in 2011 (A) and 2012 (B). Abbreviations are listed in table 1.
KJBS-47-384_F1.gif
약배양 유래 집단�서� 형질 변� � 다수성 계통 선발
2012년� 조사� 형질들� 변�를 살펴보면 집단 내 계통 들� 형질 값� 대부분 정규분�를 나타냈다(Fig. 2). 모부본 � 드래찬과 보람찬� 형질들 중�서 간장� 비슷하였고 드래 찬� �삭길�, �삭목�께, �지경수, �립수, 1차 지경수, 1 차 지경 착�립수, 1차 지경 �균립수, 2차 지경수, 2차 지경 착�립수, 2차 지경 �균립수, 지엽장, 천립중� 형질 값� 보 람찬보다 컸고 나머지 형질� 1차 지경당 2차 지경수, 1차 지 경 착�립수당 2차 지경 착�립수, 수수, 등숙률, 수량� 형질 값� 보람찬� 드래찬보다 컸다(Table 5). 약배양 유래 집단 (n=163) 내�서 2차 지경 착�립수(변�계수 23.0%), 2차 지 경수(20.0), 1차 지경 착�립수당 2차 지경 착�립수(20.0) 등 � 변�가 컸으며, 1차 지경 �균립수(3.4), 간장(3.5), 2차 지 경 �균립수(3.6) 등� 변�가 작았다. 2012년 조사� 형질 값 � �용하여 군집분�� 실시하여 약배양 집단내 계통들� 분 류하였다(Fig. 3). � 3개� 그룹으로 분류가 �었는� 그룹 A, B, C� �� 108, 42, 13개 계통� �함�었다. 그룹 C는 수량� 가장 높았고 모부본과 간장� 비슷하였고 등숙률� 낮 았으며 2차 지경 관련 형질� 2차 지경수, 2차 지경 착�립수, 2차 지경 �균립수는 드래찬과 비슷하였고 나머지 형질� 드 래찬과 보람찬� 중간 값� 나타냈다. 수량성� 높� 그룹 C � 13개 계통� 다수성 계통으로 선발하였다(Table 5).
Fig. 2.
Distribution of panicle and yield-related traits in 2012. PL (A), TNN (B), NR (C), NS (D), NPr (E), NSPr (F), MNSPr (G), NSr (H), NSSr (I), MNSSr (J), Sr/Pr (K), SSr/ SPr (L), CL (M), FL (N), PN (O), TGW (P), RRG (Q), Yield (R). Abbreviations are listed in table 1.
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Fig. 3.
Cluster analysis of DH lines based on panicle and yield-related traits in 2012.
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Table 5.
Average and range of traits value of the parents and groups divided by cluster analysis in 2012.
Table 5.
Group n PLz (cm) TNN (mm) NR NS NPr NSPr MNSPr NSr NSSr MNSSr Sr/Pr SSr/SPr CL (cm) FL (cm) PN TGW (g) RRG (%) Yield (g)

Deuraechan 3 26.3ay 1.74a 42a 166a 14a 86a 6.1a 28a 80a 2.9a 1.9ab 0.9bc 69.1a 32.4a 9c 25.0a 87.3ab 535bc
Boramchan 3 21.0d 1.53bc 35bc 129b 11d 60d 5.7d 24ab 69ab 2.8ab 2.3a 1.1a 67.6a 30.3b 11a 24.3a 91.3a 551b
a 108 24.7ab 1.63ab 40ab 156a 13b 79b 6.0ab 27a 78a 2.9a 2.1ab 1.0abc 68.7a 32.3a 9bc 24.2a 84.1b 520c
(20.1~27.1) (1.29~1.88) (26~51) (107~197) (11~15) (65~93) (5.7~6.3) (15~38) (41~118) (2.6~3.1) (1.3~3.1) (0.6~1.6) (62.3~74.3) (28.3~35.7) (8~13) (21.7~27.2) (74.1~93.0) (477~554)
b 42 22.6cd 1.49c 33c 127b 12c 69c 5.8c 21b 58b 2.7b 1.8b 0.8c 69.5a 31.0ab 11ab 24.0a 87.3ab 467d
(19.2~26.5) (1.32~1.77) (24~44) (97~172) (10~14) (57~85) (5.6~6.1) (13~31) (35~90) (2.5~3) (1.2~2.6) (0.5~1.3) (63.5~75.0) (26.9~33.5) (9~12) (21.0~27.1) (69.4~94.9) (410~501)
c 13 23.7bc 1.59bc 40ab 156a 13bc 75bc 5.9bc 28a 81a 2.9a 2.2a 1.1ab 68.2a 31.8ab 10abc 24.2a 83.9c 581a
(20.7~26.5) (1.25~1.80) (27~46) (108~179) (11~14) (62~85) (5.7~6.1) (16~33) (42~102) (2.7~3.1) (1.4~2.7) (0.6~1.4) (63.9~72.1) (28.8~35.0) (8~13) (21.9~25.7) (75.2~92.4) (555~629)
Total 163 24.1 1.59 38.2 149 13 76 5.9 26 73 2.8 2.0 1.0 68.8 31.9 10 24.1 84.9 512
C.V.(%)x 163 8.3 8.2 15.4 15.1 8.7 10.3 3.4 20.0 23.0 3.6 15.0 20.0 3.5 5.0 11.3 4.5 5.3 7.2

zAbbreviations are listed in table 1

yMeans with same letter are not significantly different at p < 0.05 (ANOVA followed by DMRT)

xC.V. = coefficient of variation

선발 계통 내� 형질 특성
선발� 13개 계통과 모부본� 드래찬과 보람찬� 대해서 �산력 검정시험� 수행하여 �삭 � 수량 관련 형질� 조사 하였다(Table 6). 공시 재료� 형질 변�를 보면 2차 지경 착 �립수(변�계수 40.3%), 2차 지경수(36.4), 1차 지경 착�립 수당 2차 지경 착�립수(30.0) 등� 변�가 컸으며, 간장 (3.5), 현미 천립중(4.1), 1차 지경 �균립수(5.2) 등� 변�가 작았다. 모부본과 선발� 13개 다수성 계통� 대해서 18개 � 삭 � 수량 관련 형질들간� �관관계를 분�하였다(Table 7). �삭길�(x1)� 1차 지경 착�립수(x6; r=0.907), 1차 지 경수(x5; 0.902)와 매우 높� 정� �관� 천립중(x16; 0.802), 2차 지경 �균립수(x7; 0.728), �삭목�께(x2; 0.612), �립수(x4; 0.558)와 높� 정� �관� 나타냈으며, 수수(x15; -0.918)와는 매우 높� 부� �관� 나타냈고 수량 (x18; -0.647)과 등숙률(x17; -0.531)과는 부� �관� 나타 냈다. �삭목�께(x2)는 1차 지경수(x5; 0.584), 천립중(x16; 0.571), �지경수(x3; 0.568), �립수(x4; 0.564), 1차 지경 착 �립수(x6; 0.519) 순으로 정� �관� 나타냈고 수수(x15; -0.684)와는 높� 부� �관� 나타냈다. �지경수(x3)와 � 립수(x4)는 1차 지경 관련 형질(x5, x6, x7)보다 2차 지경 관 련 형질(x8, x9, x10)과 높� 정� �관관계를 나타냈으며, 수 수(x15)와는 �관계수 -0.570, -0.619로 높� 부� �관� 나 타냈고 수량과는 �관관계가 없었다. 1차 지경수(x5) � 착� 립수(x6)는 천립중(x16; 0660, 0.663), 지엽장(x14; 0.549, 0.558)과는 정� �관� 나타냈고 수수(x15; -0.882, -0.847), 수량(x18; -0.650, -0.591), 등숙률(x17; -0.548, -0.539)과는 부� �관� 나타냈다. 2차 지경수(x8) � 착�립수(x9)는 � 지경수(x3; 0.975, 0.952)와 �립수(x4; 0.932, 0.914)와 매우 높� 정� �관� 나타냈으나 수수(x15), 등숙률(x17), 수량 (x18)과는 통계�으로 유�한 �관관계가 없었다. 1차 지경 �균립수(x8)� �삭길�(x1; 0.728)와 천립중(x16; 0.541)과 정� �관� 나타냈고 수수(x15; -0.536)와는 부� �관� 수 량(x18)과는 무�관� 나타낸 반면 2차 지경 �균립수(x10) 는 �삭길�, 천립중, 수수와 무�관� 나타냈고 1차 지경당 2차 지경수(x11; 0.871), 1차 지경 착�립수당 2차 지경 착� 립수(x12; 0.857)와는 높� 정� �관� 수량(0.540)과는 정 � �관� 나타냈다. 간장(x13)� 다른 형질들과 �관관계가 없었고 지엽장(x14)� �지경수(x3; 0.523), �립수(x4; 0.554), 1차 지경수(x5; 0.549), 1차 지경 착�립수(x6; 0.558)와 정� �관� 나타냈다. 수수(x15)는 �삭길�(x1; -0.918), 1차 지경 관련 형질(x5; -0.882, x6; -0.847, x7; -0.536), 천립중(x16; -0.743), �삭목�께(x2; -0.684), �지 경수(x5; -0.570), �립수(x6; -0.619)와 부� �관관계를 나 타냈으나 수량(x18; 0.679) � 등숙률(x17; 0.551)과는 정� �관� 나타냈다. 수량(x18)� 수수(x15)와 �관계수 0.679 로 가장 높� 정� �관� 나타냈고 1차 지경당 2차 지경수 (x11; 0.598), 1차 지경 착�립수당 2차 지경 착�립수(x12; .0563), 2차 지경 �균립수(x13; 0.540)순으로 정� �관� 나타냈으며, 1차 지경수(x5; -0.650), �삭길�(x1; -0.647), 1 차 지경 착�립수(x6; -0.591), 천립중(x16; -0.584)순으로 부 � �관� 나타냈다.
Table 6.
Panicle and yield-related traits of parents and selected DH lines at yield trial in 2013.
Table 6.
Var./lines PLz (cm) TNN NR NS NPr NSPr MNSPr NSr NSSr MNSSr Sr/Pr SSr/SPr CL (cm) FL PN TGW (g) RRG (%) MRY (kg/10a)

Deuraechan (DR) 26aby 1.76ab 32cde 122bcd 11abcd 67bcd 5.9bcde 20de 55efgh 2.7ef 1.7ef 0.8ghi 73ab 36.0ab 10efg 25.3bc 91.2a 512b
Boramchan (BR) 21e 1.50def 31cde 111de 9h 51h 5.5h 22cde 60def 2.8cde 2.3abc 1.2bc 71bcd 32.1de 13abc 24.0ef 91.4a 567ab
HR28832-AC60-1-1 23cd 1.62bcde 31cde 117bcde 10fgh 56gh 5.8defg 21cde 61def 2.8c 2.2bcd 1.1cd 72bc 34.2abcde 12bcd 23.7fg 91.9a 596a
(AC60)
HR28832-AC105-1-1 24cd 1.69abc 35abc 132abc 11bcde 65cde 5.8cdefg 23bcd 67bcde 2.8c 2.0de 1.0def 68e 33.5bcde 12abcd 24.3e 84.1bcd 559ab
(AC105)
HR28832-AC107-1-1 26ab 1.68abc 39a 149a 12a 73a 6.0ab 27ab 75ab 2.8cd 2.2bcd 1.0de 73ab 37.2a 9h 25.7ab 83.8bcd 516b
(AC107)
HR28832-AC109-1-1 23de 1.34g 26f 101e 10gh 57g 5.9abc 16f 44h 2.7ef 1.6fg 0.7hi 72bc 31.0e 13ab 24.5de 89.4ab 561ab
(AC109)
HR28832-AC112-1-1 23de 1.47efg 31cde 117bcde 11def 60gef 5.7fg 21de 57efg 2.7def 1.9def 0.9efg 71bcd 32.0de 12cde 23.2g 92.2a 549ab
(AC112)
HR28832-AC123-1-1 27a 1.62bcde 34abc 133ab 12abc 71abc 6.0a 23cd 62cdef 2.8cdef 1.9def 0.9fgh 73abc 32.6cde 10gh 25.7ab 85.3bcd 541ab
(AC123)
HR28832-AC130-1-1 25ab 1.66abc 29ef 115cde 12abcd 67bcd 5.8cdef 18ef 47gh 2.7ef 1.5g 0.7i 73ab 34.1abcde 10fgh 25.5abc 87.6abc 516b
(AC130)
HR28832-AC134-1-1 26ab 1.58cdef 37ab 143a 12ab 72ab 6.0a 25abc 71abcd 2.8bc 2.0cd 1.0def 75a 35.8abc 10gh 25.0cd 82.7cd 500b
(AC134)
HR28832-AC149-1-1 23cd 1.62bcde 35abc 132abc 10efg 58gf 5.7g 25abc 74abc 2.9ab 2.4ab 1.2ab 68de 31.3e 11cdef 23.3g 85.0bcd 564ab
(AC149)
HR28832-AC152-1-1 23cd 1.65abcd 38a 148a 11cde 66cde 6.0ab 27ab 82a 3.0a 2.4ab 1.2ab 72bcd 35.9ab 11cdef 24.0ef 89.5ab 595a
(AC152)
HR28832-AC156-1-1 22de 1.46fg 29def 110de 10fgh 56g 5.7efg 20def 54fgh 2.7cdef 2.0de 0.9defg 71bcd 35.5abc 14a 23.7fg 90.9a 590a
(AC156)
HR28832-AC161-1-1 25bc 1.73abc 38a 146a 11de 63def 5.9bcd 28a 83a 3.0a 2.6a 1.3a 69de 35.1abcd 11defg 25.5abc 84.9bcd 605a
(AC161)
HR28832-AC178-1-1 25abc 1.80a 33bcde 122bcd 11cde 63def 5.8cdefg 22cd 58efg 2.7f 2.0de 0.9defg 70cde 33.8bcde 10gh 26.1a 80.7d 507b
(AC178)
Mean 24 1.61 33 125 11 62 5.8 62 22 2.8 2.0 1.0 71 34 11 24.6 87.5 552
C.V. 7.5 8.7 27.3 28.0 18.2 21.0 5.2 36.4 40.3 7.1 30.0 30.0 3.5 6.9 13.5 4.1 5.3 8.2

zAbbreviations are listed in table 1

yMeans with same letter are not significantly different at p < 0.05 (ANOVA followed by DMRT)

Table 7.
Correlation coefficient among panicle-related traits and agronomic traits in 2013 (n=15).
Table 7.
Traits PLz TNN NR NS NPr NSPr MNSPr NSr NSSr MNSSr Sr/Pr SSr/SPr CL FL PN TGW RRG MRY
(x1) (x2) (x3) (x4) (x5) (x6) (x7) (x8) (x9) (x10) (x11) (x12) (x13) (x14) (x15) (x16) (x17) (x18)

x1 0.612* 0.464ns 0.558* 0.902** 0.907** 0.728** 0.279ns 0.218ns -0.008ns -0.265ns -0.213ns 0.420ns 0.473ns -0.918** 0.802** -0.531* -0.647**
x2 0.568* 0.564* 0.584* 0.519* 0.209ns 0.483ns 0.443ns 0.251ns 0.197ns 0.210ns -0.233ns 0.486ns -0.684** 0.571* -0.457ns -0.289ns
x3 0.984** 0.581* 0.582* 0.423ns 0.975** 0.952** 0.761** 0.647** 0.692** -0.109ns 0.523* -0.570* 0.232ns -0.518* 0.010ns
x4 0.671** 0.687** 0.560* 0.932** 0.914** 0.739** 0.543* 0.588* -0.023ns 0.554* -0.619* 0.301ns -0.532* -0.040ns
x5 0.987** 0.713** 0.384ns 0.320ns 0.059ns -0.232ns -0.182ns 0.341ns 0.549* -0.882** 0.660** -0.548* -0.650**
x6 0.815** 0.390ns 0.334ns 0.092ns -0.232ns -0.175ns 0.393ns 0.558* -0.847** 0.663** -0.539* -0.591*
x7 0.285ns 0.272ns 0.164ns -0.212ns -0.143ns 0.450ns 0.463ns -0.536* 0.541* -0.375ns -0.247ns
x8 0.992** 0.847** 0.798** 0.835** -0.217ns 0.442ns -0.404ns 0.082ns -0.437ns 0.190ns
x9 0.907** 0.835** 0.861** -0.249ns 0.407ns -0.330ns 0.021ns -0.390ns 0.278ns
x10 0.871** 0.857** -0.301ns 0.244ns -0.047ns -0.211ns -0.164ns 0.540*
x11 0.992** -0.471ns 0.102ns 0.108ns -0.318ns -0.103ns 0.598*
x12 -0.426ns 0.157ns 0.060ns -0.269ns -0.148ns 0.563*
x13 0.354ns -0.328ns 0.249ns 0.187ns -0.468ns
x14 -0.466ns 0.380ns -0.160ns -0.158ns
x15 -0.743** 0.551* 0.679**
x16 -0.579* -0.584*
x17 0.409ns

zAbbreviations are listed in table 1

주성분 분�� 통해 형질들� 차�� 줄�고 구조� 관계 를 분�하였다(Table 8, Fig. 4A). 축소� 형질들� 주성분 1 � 전체 형질 특성� 45.8%를 설명할 수 있었으며 주성분 2 는 32.0%를 설명할 수 있어 �들 2개 주성분으로 전체 형질 � 77.8%를 설명할 수 있었다. 주성분 1� �립수(r=0.937), �지경수(0.898), 1차 지경 착�립수(0.842), 1차 지경수 (0.838), �삭길�(0.789), 2차 지경수(0.788), �삭목�께 (0.707), 1차 지경 �균립수(0.653), 지엽장(0.635) 순으로 높 � 정� �관� 나타냈고 천립중과는 중간 수준� 정�관� 나타냈다. 수수와 등숙률과는 �관계수 -0.825와 -0.645로 높 � 부� �관� 나타냈다. 주성분 2는 1차 지경당 2차 지경수 (0.933), 1차 지경 착�립수당 2차 지경 착�립수(0.906), 수 량(0.795), 2차 지경 �균립수(0.791), 2차 지경 착�립수 (0.659), 2차 지경수(0.598)와는 높� 정� �관� 나타냈으 며 간장(-0.589), 천립중(-0.579), �삭길�(-0.551)과는 부� �관� 나타냈다. 주성분 분��를 통해 형질들� 구조� 관 계를 보면 주성분 1� 기준으로 수수와 등숙률� 지경 � 착 �립수와 다른 방향성� 나타내고 있었고 2차 지경 관련 형질 보다 1차 지경 관련 형질� 그 거리가 � 멀게 분�하였으며 �삭길�, 지엽장, 천립중과 다른 방향성� 나타냈다. 주성분 2를 기준으로는 1차 지경당 2차 지경 형질� 대한 비율(1차 지경당 2차 지경수, 1차 지경 착�립수당 2차 지경 착�립수), 수량 � 2차 지경 관련 형질(지경수, 착�립수, �균립수)� 천립중, �삭길�와 다른 방향성� 나타냈다.
Fig. 4.
Relationship among traits (A) and lines (B) in the principal component analysis (PCA) diagram composed of component 1 and 2 in 2013. Abbreviations are listed in table 1 and table 6.
KJBS-47-384_F4.gif
Table 8.
Eigenvectors for the principal components of traits and correlation coefficient between the traits and principal components.
Table 8.
Trait Eigenvector Correlation coefficient

Prin1z Prin2 Prin3 Prin1 Prin2 Prin3

PL 0.275 -0.230 -0.011 0.789** -0.551* -0.014ns
TNN 0.246 -0.015 -0.375 0.707** -0.037ns -0.451ns
NR 0.313 0.172 0.030 0.898** 0.412ns 0.036ns
NS 0.326 0.127 0.087 0.937** 0.306ns 0.105ns
NPr 0.292 -0.198 0.027 0.838** -0.476ns 0.032ns
NSPr 0.293 -0.195 0.108 0.842** -0.468ns 0.130ns
MNSPr 0.228 -0.150 0.342 0.653** -0.359ns 0.411ns
NSr 0.275 0.249 0.027 0.788** 0.598* 0.032ns
NSSr 0.260 0.274 0.053 0.745** 0.659** 0.064ns
MNSSr 0.176 0.329 0.145 0.506ns 0.791** 0.174ns
Sr/Pr 0.103 0.389 -0.036 0.297ns 0.933** -0.043ns
SSr/SPr 0.123 0.377 -0.014 0.352ns 0.906** -0.016ns
CL 0.040 -0.245 0.569 0.114ns -0.589* 0.684**
FL 0.221 -0.034 0.313 0.635* -0.082ns 0.376ns
PN -0.287 0.178 0.112 -0.825** 0.427ns 0.134ns
TGW 0.203 -0.241 -0.227 0.582* -0.579* -0.273ns
RRG -0.225 0.044 0.422 -0.645** 0.105ns 0.507ns
MRY -0.109 0.331 0.181 -0.312ns 0.795** 0.218ns
Eigen value 8.24 5.77 1.45
Contribution (%) 45.8 32.0 0.08
Cumulative contribution (%) 45.8 77.8 85.8

zPrin1 and Prin2 indicate the first and second component, respectively

수량� 대한 형질� 기여� 분�
약배양 유래 집단(2012)과 선발� 계통(2013년)�서 수량� 관여하는 형질� 수당립수, 수수, 등숙률, 천립중� 수량� 미치 는 기여�를 경로계수 분�� 통해 분�하였다(Fig. 5). 경로계 수� 모형�서 모형� �합�를 결정하는 기준 중 Chi-square 값� 유�하여 기준� 맞지 않으나 �외� 기준� �합�지수 (GFI; Goondess of Fit Index)와 표준�합지수(NFI; Normed Fit Index), 비��합� 지수(CFI; Comparative Fit Index)가 0.9보다 �고 �소간 �균차�(RMSR; Root Mean Square Residual)� 0� 가�게 작� 값으로 전반�으로 모형� �합 하였다. 약배양 집단�서 수당립수가 표준화 회귀계수 0.554로 수량성� �가시키는 방향으로 기여�가 가장 컸으며 �후 수 수(0.383), 천립중(0.294) 순�었고, 등숙률(-0.308)� 수량� �소시키는 방향으로 작용하였다. 2차 지경 착�립수가 1차 지 경 착�립수� 비해 수당립수 �가� 기여하는 정�가 컸으며, 간접� 경로를 통해 2차 지경 착�립수가 수량 향�� 미치는 �향� 0.418로 수수� 비해서 컸다. 선발� 계통� 분��서는 수수가 표준화 계수 0.639로 수량성� 기여�가 가장 컸고 수 당립수는 0.329로 집단�서보다 비해 기여�가 줄었으며 천립 중과 등숙률� 통계�으로 기여하는 바가 없었다. 2차 지경 착 �립수가 1차 지경 착�립수� 비해 수당립수� 기여하는 바가 컸으나 그 정�는 집단�서보다 줄었고 간접�으로 수량성� 기여하는 바� 줄었다.
Fig. 5.
Path analysis of causal relationships among traits to explain the variation of yield in 2012 (A) and 2013 (B). Continuous arrows and dotted arrows indicate the significant and no significant effect, respectively. The arrow line thickness means proportion of the effect. ** and * mean significant at 1 and 5% probability level, respectively.
KJBS-47-384_F5.gif
다수성 우량계통 선발 � 특성 분�
드래찬� 보람찬보다 �삭길�, �삭목�께, �지경수, � 립수, 1차 지경수, 1차 지경 착�립수, 1차 지경 �균립수, 간 장, 지엽장, 천립중� 형질 값� 컸고, 2차 지경수, 2차 지경 착�립수, 2차 지경 �균립수 1차 지경당 2차 지경수, 1차 지 경 착�립수당 2차 지경 착�립수, 수수, 백미 수량� 형질 값 � 보람찬� 드래찬보다 컸다(Table 6). 선발� 13개 계통 중 �서 HR-28832-AC161-1-1 (�하 AC161), AC60, AC152, AC156 계통� 수량성� 605, 596, 595, 590 kg/10a로 드래 찬(512)과 보람찬(567 kg/10a)보다 높았다. AC152과 AC161� 모부본보다 �지경수, �립수, 2차 지경 관련 형질 (지경수, 착�립수, �균립수) � 1차 지경당 2차 지경 형질� 대한 비율(1차 지경당 2차 지경수, 1차 지경 착�립수당 2차 지경 착�립수)� 컸고 �삭길�, �삭목�께, 지엽장, 수수, 천립중� 모부분� 중간 값� 나타냈으며 1차 지경 관련 형질 (지경수, 착�립수, �균립수)� 드래찬과 비슷하였고 간장� 작� 편으로 등숙률� 낮았다. AC60� 모부본과 등숙률� 비슷하고, �삭길�, �삭목�께, 간장, 지엽장, 수수와 �지 경수 � �립수, 1차 지경 관련 형질(지경수, 착�립수, �균 립수), 1차 지경당 2차 지경 형질� 대한 비율(1차 지경당 2 차 지경수, 1차 지경 착�립수당 2차 지경 착�립수)� 모부 본� 중간 값� 나타냈으며 2차 지경 관련 형질(지경수, 착� 립수, �균립수)과 천립중� 보람찬과 비슷하였다. AC156� 모부본과 등숙률� 비슷한 편으로 수수가 많았고 �삭길�, 1 차 지경 관련 형질(지경수, 착�립수, �균립수), 1차 지경당 2차 지경 형질� 대한 비율(1차 지경당 2차 지경수, 1차 지경 착�립수당 2차 지경 착�립수), 지엽장� 모부본� 중간 값 � 나타냈으며, 2차 지경 관련 형질(지경수, 착�립수, �균립 수)는 드래찬과 비슷하였고 천립중� 작았다. 주성분 값� 기 준으로 한 공시재료들� 구조� 관계를 보면 AC152와 AC161� 모부본� 드래찬과 보람찬� 중간� 위치하고 있으 며 AC156� 보람찬� 근접하였고AC60� 보람찬� 가�게 AC152, AC161과 AC156� 중간� 위치하고 있다(Fig. 4B).
수량성� 모부본보다 많았� AC60, AC152, AC156, AC161� 대해서 2014년� 수량성� 대해서 재검정하였다 (Table 9). �들 계통들� 출수기는 모부본과 비슷하였고 간 장� AC152가 68 cm로 드래찬(69)과 보람찬(69)가 비슷하 였고 나머지 계통� 작았으며, �삭길�가 AC60, AC152, AC161� 22 cm로 드래찬(24)과 보람찬(20)� 중간 값� 나 타냈고 AC156� 20 cm보람찬과 비슷하였다. 수수는 보람찬 과 AC156� 12개로 많았고 AC60, AC152, AC161� 11개, 드래찬� 9개였다. �립수는 AC161 (143), 드래찬(142) AC152 (140개) 많았고 AC60� 129개, 보람찬과 AC156� 108개 순�었다. 등숙률� AC156 (94.8), 보람찬(94.5%)로 높았고 드래찬(91.1), AC60 (91.0), AC152 (90.5)는 중간� 었고 AC161 (86.3%)는 낮았다. 천립중� AC161 (25.9), 드 래찬(25.8 g)� 무거웠으며 AC152 (24.7), AC60 (24.5), 보 람찬(24.4), AC156 (24.0 g) 순�었다. 정현비율� AC156� 83.2%로 가장 높았고 보람찬(82.5)� 중간 값� 나타냈고 AC152 (81.6), 드래찬(81.6), AC160 (81.1) AC60 (81.0%) 는 비슷하였다. 수량성� 드래찬(백미 수량; 481 kg/10a)� 낮았고 보람찬(558), AC152 (558), AC161 (559), AC156 (567), AC60 (575 kg/10a) 순으로 높았다.
Table 9.
Yield-related traits of parents and selected DH lines at yield trial in 2014.
Table 9.
Var./lines Heading date (DASz) CL (cm) PL (cm) PN NS RRG (%) TGW (g) RRY (kg/10a) BRR (%) BRY (kg/10a) MRY (kg/10a) Index of MRY (%)

Deuraechan (DR) 107ay 69a 24a 9c 142a 91.1ab 25.8a 641b 81.6b 523b 481b 100
Boramchan (BR) 108a 69a 20c 12a 108b 94.5a 24.4b 736a 82.5ab 607a 558a 116
HR28832-AC60-1-1 108a 66bc 22b 11b 129ab 91.0ab 24.5b 772a 81.0b 625a 575a 119
(AC60)
HR28832-AC152-1-1 108a 68ab 22b 11b 140a 90.5ab 24.7b 744a 81.6b 607a 558a 116
(AC152)
HR28832-AC156-1-1 108a 64c 20c 12a 108b 94.8a 24.0c 741a 83.2a 617a 567a 118
(AC156)
HR28832-AC161-1-1 106a 65bc 22b 11b 143a 86.3b 25.9a 748a 81.1b 607a 559a 116
(AC161)

zAbbreviations are listed in table 1

yMeans with same letter are not significantly different at p < 0.05 (ANOVA followed by DMRT)

��니카 초다수 벼 품종으로 다수성� 관여하는 형질 특성 � 다른 드래찬과 보람찬� �용하여 약배양� 통해 �삭 � 수 량 관련 형질 변�를 �대시킨 고정 집단� 육성하고, 보통기 � 반재배�서 �삭 � 수량 관련 형질들� 특성� 분�하여 수량 � 높� 우량계통� 개발하고� 하였다. 드래찬� 소얼 수중형 초형으로 �복� 강하고 수당립수를 �대시킴으로� 수량성� 향�시키는 방향으로 육성� 품종�다. 드래찬� 계보�를 보면 중국 온대 ��니카 품종으로 sd-1� 가지고 있으면서 �복� 매우 강하여 NPT 계통� �복저항성 강화를 위해 �용� Shennong 89-336(Khush & Aquino 1994)과 �를 통해 개발� NPT 계통� IR66738-118-1-2(Shennong 89-336/Soponjono), IR66158-72-2-1-3(Shennong 89-336/Baliontjer), IR65601-35-6-2-3 (Shennong 89-336/K. Rondo Marong)� �용�었다(Fig. 6). �들 ��들� 소얼 수중형으로 수당립수가 많� 특성� 나타낸 다(Yamagishi et al. 2003). 보람찬� 현재까지 개발� 우리나 � ��니카 품종 중�서 잠재 수량성� 가장 높� 품종으로 수 수가 �대�으로 다른 품종� 비해 많� 초형으로 등숙률� 높 고 내�복성� 갖추고 있다(Ha et al. 2012). 2013, 2014년 수행 � �산력 검정시험�서 출수기는 � 품종� 비슷하였고 드래찬 � 간장� �고 수장� 길며 수당립수는 많고 천립중� 무거운 반면 보람찬� 수수가 많고 등숙률과 정현비율� 높았고 수량� 많았고(Table 6, 9) 드래찬� 소얼 수중형, 보람찬� 다소 수수 형� 가까운 초형� 나타냈다.
Fig. 6.
Pedigree diagram of Deuraechan.
KJBS-47-384_F6.gif
약배양 집단� 분��서 지경과 착�립수 관련 형질� 수 량과 높� 정� �관� 나타냈고 2차 지경 형질� 1차 지경 형질보다 수당립수와 수량과 높� 정� �관� 나타냈으며, �삭 관련 형질� 수수와 등숙률과는 높� 부� �관� 나타 냈다(Table 3). 수수와 등숙률� 수량성과 �관� 없거나 낮 � 부� �관� 나타냈으며 주성분 분��서 수량성과 다른 방향성� 나타내(Fig. 1) 집단�서� 다수성� 수당립수와 같 � �삭관련 형질� �해서 � 많� �향� 받� 것으로 �� �었다. 다수성 선발 계통� 대한 분��서는 집단�서와 다 르게 수량과 �지경수, 수당립수, 1차 지경 �균립수는 통계 � 유�성� 없었고, 1차 지경수와 착�립수는 부� �관� 나타냈으며 2차 지경수와 착�립수는 정� �관 정�가 집단 � 비해 �었다(Table 7). 수량과 가장 �관� 높� 형질� 수 수로 약배양 집단 내�서 무�관(2011년) � 낮� 부� �관 (2012)� 나타낸 것과 다른 결과로 주성분 분�� 통한 형질 들� 구조� 관계�서� 수수, 등숙률과 수량� 같� 방향성 � 나타내고 있어(Fig. 4) 수량성� 확보� 계통들�서는 수 수가 수당립수� 비해 수량성과� 관련성� 높� 것으로 나 타났다. 수량 구성 요소들� 수량� 기여하는 정�를 보면 집 단과 선발 계통 모� 2차 지경 착�립수가 1차 지경 착�립수 보다 수당립수� 기여하는 정�가 컸으며, 집단�서는 수당립 수가 선발 계통�서는 수수가 수량� �가시키는 방향으로 기여하는 정�가 가장 컸다(Fig. 5). 집단�서 등숙률� 수량 성� 줄�는 방향으로 작용하였는�, �는 수량성 지표로 � 용한 간�수량� 정조 �태로 정현비율� 대한 고려가 �루 지지 않았으며 �가� 수당립수가 등숙률 저하� �한 수량 �소를 �쇄한 것으로 ��한다. 선발� 계통� 분��서 수 량성� 확보� 계통들� �미 �정 수준� 수당립수를 확보 하고 있어 수수� �가� �한 등숙률 향�과 단위면�당 립 수� �가로 �한 수량성 향�� 수당립수� �가� �한 향 � 효과보다 높� 것으로 ���다.
수량성� 낮� AC107, AC123, AC130, AC134, AC178 � 드래찬과 초형� 비슷하였으며 �삭� 길고 1차 지경수 � 착�립수 많고 수수는 �고 천립중� 무거운 소얼 수중형으 로 드래찬� 비해 등숙률� 떨어지는 특성� 나타냈다(Table 6). NPT 계통� �함하는 소얼 수중형 초형� 문제�으로 수 수가 줄어들고 소수가 �가함� 따� �물체� 바�오매스가 �소하게 �며 �삭 내 소수� 착�밀�가 높아지고 제한� 유관�� �화산물� 제대로 채워주지 못하게 �며 엽노화가 빠르게 진행�어 등숙률� 떨어지는 �� 들었다(Peng et al. 2008, Yamagishi et al. 2003). 드래찬� 등숙률� 양호하는 등 소얼 수중형 초형� 문제�� 다소 해결한 품종�나 보람 찬� 비해 엽노화가 빠르고 수량성� 낮으며, 육성� 소얼 수 중형� 계통들� 수량과 등숙률� 낮는 등 전형�� 소얼 수 중형� 문제�� 발�하였다. 드래찬과 보람찬보다 수량� 높 았� 계통� AC60, AC152, AC156, AC161로 AC152와 AC161� 모부본� 드래찬과 보람찬� 중간형태로 수당립수 가 드래찬과 비슷하게 많지만 수수� 많아 단위면�당 립수 가 드래찬� 비해 늘어 수량� �가하였으나 등숙률� 다소 떨어지는 특성� 나타냈다. AC156� 수당립수와 수수가 보 람찬과 비슷하고 등숙률과 제현율� 높으며 보람찬과 비슷한 초형� 나타냈다. AC60� 수당립수, 등숙률과 수수 등 �업 형질� AC152, AC161과 AC156� 중간 형태� 특성� 나 타내었다. 선발� 4계통 모� 보람찬과 같거나 다소 많� 수 량� 나타내 수량성 �대� 소기� 목�� 달성하였으나 � 하는 만�� 수량 향�� �루지 못하였다.
벼� 수량성� 환경변�� �해 �향� 많� 받으며 여러 형질� 관여하는 양� 형질로서 광합성� �해 �화산물� �성�고 전류�는� 관여하는 능력� source와 종실� 축� � 수 있는 잠재력� sink� �해 결정�다. 수량성 향�� 위 해서는 source와 sink� �기와 균형� 중요하며, sink� 주 요 형질들� 수당립수, 수수, 종실�기 � 등숙률� 서로 �향 � 주며 관련성� 맺고 있기 때문� �들� ���� 조합� 필요하다. 본 연구�서와 같� 종실 �기가 비슷한 우리나� ��니카 품종� 수량성 향�� 위한 초다수성 육종사업� 목표를 달성하기 위해서는 sink 측면�서 등숙률� �소하지 않는 범위�서 수당립수를 �가시키고 �정 수수를 확보하는 방법과 수수를 �가시킴으로� 등숙률� 향�시키는 방향� 서 �정 수당립수를 확보하는 방법� 들 수 있겠고, 광합성� 향�시킬 수 있는 엽색, 수광태세� 좋� 지엽형태나 엽노화 지연 등과 같� source� 능력 �대� 좀 � 관심� 가져야 할 것으로 ���었다. 육성� ��니카 초다수성 품종� � 용하여 약배양� 통해 형질 변�� �� 넓힘으로� 한 단계 향�� 다수성 계통� 육성하고� 하였으나 한계가 있는 것 으로 �단�었고, �러한 한계 극복� 위해서는 극단�� 형 질 변�를 보�는 유전��� 확보, 중간모본 육성 � 품종화 와 같� 새로운 형질 특성 �입� 대한 단계�� 노력� �욱 필요할 것으로 ��한다.
��니카 초다수성 품종으로 다수성� 관여하는 형질 특성 � 다른 드래찬과 보람찬� �용하여 �삭과 수량 관련 형질 � 다양성� �대� 약배양 유래 집단� 육성하고 다수성 계 통� 선발하였다. 163개로 구성� 약배양 유래 집단과 선발 � 다수성 13계통� �삭 � 수량 관련 형질� �관분�, 주 성분분� � 경로계수 분�� 통해 분�하였다. �삭 관련 형 질 중 2차 지경 착�립수가 1차 지경 착�립수� 비해 수당립 수와 높� 정� �관관계를 나타냈으며 수당립수가 �가하는 쪽으로 기여�가 컸다. 형질들� 수량� 기여하는 정�를 보 면 집단�서는 수당립수가 선발 계통�서는 수수가 수량성 �진� 기여�가 가장 컸다. 수수는 집단과 선발 계통�서 모 � 등숙률과 정� �관� 나타냈다. 선발 계통�서 수수 �가 � �한 등숙률� 향�과 단위면�당 립수� �가로 �한 수 량성 향�� 수당립수 �가� �한 향� 효과보다 컸다. 13개 선발 계통 중 AC60, AC152, AC156, AC161 계통� 모부본 � 드래찬(481 kg/10a)과 보람찬(559 kg/10a) 보다 높� 수 량성� 나타냈다. 드래찬� 수수가 �고 수당립수가 많� 소 얼 수중형� 나타냈으며 보람찬� 수수가 많고 등숙률� 높 � 수수형� 가까웠다. 선발� 다수성 계통들� 모부본� 중 간 형태� 형질 특성� 나타내며 개선� 잠재 수량성� 가지 고 있다. �들 우량 계통들� ��니카 다수성 품종 개발� 위해 활용� 수 있� 것�다.
본 논문� �촌진�청 연구사업(과제번호: PJ01020202)� 지�� �해 �루어진 것임
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Development of High-yielding Rice Lines and Analysis of Panicle and Yield-related Traits Using Doubled Haploid Lines Derived from the Cross between Deuraechan and Boramchan, High-yielding japonica Rice Cultivars in Korea
Korean. J. Breed. Sci.. 2015;47(4):384-402.   Published online December 31, 2015
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Development of High-yielding Rice Lines and Analysis of Panicle and Yield-related Traits Using Doubled Haploid Lines Derived from the Cross between Deuraechan and Boramchan, High-yielding japonica Rice Cultivars in Korea
Korean. J. Breed. Sci.. 2015;47(4):384-402.   Published online December 31, 2015
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Development of High-yielding Rice Lines and Analysis of Panicle and Yield-related Traits Using Doubled Haploid Lines Derived from the Cross between Deuraechan and Boramchan, High-yielding japonica Rice Cultivars in Korea
Image Image Image Image Image Image
Fig. 1. Relationship among traits in the principal component analysis (PCA) diagram composed of component 1 and 2 in 2011 (A) and 2012 (B). Abbreviations are listed in table 1.
Fig. 2. Distribution of panicle and yield-related traits in 2012. PL (A), TNN (B), NR (C), NS (D), NPr (E), NSPr (F), MNSPr (G), NSr (H), NSSr (I), MNSSr (J), Sr/Pr (K), SSr/ SPr (L), CL (M), FL (N), PN (O), TGW (P), RRG (Q), Yield (R). Abbreviations are listed in table 1.
Fig. 3. Cluster analysis of DH lines based on panicle and yield-related traits in 2012.
Fig. 4. Relationship among traits (A) and lines (B) in the principal component analysis (PCA) diagram composed of component 1 and 2 in 2013. Abbreviations are listed in table 1 and table 6.
Fig. 5. Path analysis of causal relationships among traits to explain the variation of yield in 2012 (A) and 2013 (B). Continuous arrows and dotted arrows indicate the significant and no significant effect, respectively. The arrow line thickness means proportion of the effect. ** and * mean significant at 1 and 5% probability level, respectively.
Fig. 6. Pedigree diagram of Deuraechan.
Development of High-yielding Rice Lines and Analysis of Panicle and Yield-related Traits Using Doubled Haploid Lines Derived from the Cross between Deuraechan and Boramchan, High-yielding japonica Rice Cultivars in Korea

The abbreviations for panicle and yield-related traits.

Abbreviations:

PL Panicle length
TNN Thickness of neck node
PRB Primary rachis-branch
SRB Secondary rachis-branch
NR Number of rachis-branches per panicle
NS Number of spikelets per panicle
NPr Number of PRBs per panicle
NSPr Number of spikelets on PRBs per panicle
MNSPr Mean number of spikelets on a PRB
NSr Number of SRBs per panicle
NSSr Number of spikelets on SRBs per panicle
MNSSr Mean number of spikelets on a SRB
Sr/Pr Ratio of SRBs to PRBs
SSr/SPr Ratio of spikelets on SRBs to spikelets on PRBs
CL Culm length
FL Flag leaf length
PN Panicle number per hill
TGW 1,000 grain weight
RRG Ratio of ripened grain
BRR Brown/rough rice ratio
RRY Rough rice yield
BRY Brown rice yield
MRY Milled rice yield
DAS Day after seedling

Anther culture ability and doubled haploid lines derived from the cross of Deuraechan/Boramchan.

Anther cultured (no.) Cali induced (no. and %) Plant regeneration (no. and %)z Green plants (no. and %)y Albino (no. and %)x Doubled haploid lines (no.)w

10,600 3,528(33.3) 450(12.8) 245(54.4) 215(45.6) 220

zPercent plant regeneration is the number of plants regenerated over number of cali plated

yPercent green is the number of green plants regenerated over total plants regenerated

xPercent albino is the number of albino plants regenerated over total plants regenerated

wDouble haploid lines are harvested plants having fertile grain

Correlation coefficient among panicle and yield related traits in 2011 and 2012 (n=163).

Traitsz PLy TNN NR NS NPr NSPr MNSPr NSr NSSr MNSSr Sr/Pr SSr/SPr CL FL PN TGW RRG Yield
(x1) (x2) (x3) (x4) (x5) (x6) (x7) (x8) (x9) (x10) (x11) (x12) (x13) (x14) (x15) (x16) (x17) (x18)

x1 0.547** 0.472** 0.515** 0.544** 0.589** 0.354** 0.415** 0.396** 0.229** 0.241** 0.209** 0.209** 0.556** -0.392** 0.381** -0.210** 0.198*
x2 0.752** 0.858** 0.884** 0.655** 0.674** 0.294** 0.816** 0.803** 0.549** 0.629** 0.602** 0.005ns 0.513** -0.325** 0.159* -0.313** 0.376**
x3 0.706** 0.833** 0.987** 0.571** 0.571** 0.188* 0.988** 0.965** 0.587** 0.859** 0.823** -0.052ns 0.412** -0.302** 0.044ns -0.364** 0.468**
x4 0.659** 0.846** 0.990** 0.609** 0.630** 0.286** 0.967** 0.958** 0.640** 0.820** 0.793** -0.039ns 0.442** -0.297** 0.059ns -0.378** 0.456**
x5 0.794** 0.765** 0.765** 0.817** 0.966** 0.218** 0.437** 0.367** 0.002ns 0.075ns 0.031ns 0.194* 0.476** -0.343** 0.395** -0.156* 0.118ns
x6 0.803** 0.745** 0.746** 0.812** 0.979** 0.462** 0.443** 0.380** 0.050ns 0.093ns 0.033ns 0.224** 0.515** -0.347** 0.388** -0.170* 0.126ns
x7 0.561** 0.424** 0.423** 0.516** 0.579** 0.732** 0.165* 0.169* 0.185* 0.086ns 0.009ns 0.171* 0.325** -0.128ns 0.118ns -0.098ns 0.07ns
x8 0.594** 0.799** 0.991** 0.969** 0.674** 0.656** 0.367** 0.988** 0.643** 0.926** 0.896** -0.094ns 0.361** -0.266** -0.026ns -0.370** 0.491**
x9 0.580** 0.792** 0.982** 0.966** 0.649** 0.633** 0.360** 0.994** 0.744** 0.942** 0.933** -0.130ns 0.335** -0.226** -0.073ns -0.388** 0.497**
x10 0.401** 0.610** 0.739** 0.759** 0.418** 0.417** 0.281** 0.762** 0.821** 0.713** 0.785** -0.267** 0.180* 0.010ns -0.283** -0.347** 0.382**
x11 0.298** 0.590** 0.825** 0.770** 0.288** 0.256** 0.058ns 0.888** 0.907** 0.806** 0.986** -0.182* 0.204** -0.164ns -0.194* -0.354** 0.495**
x12 0.346** 0.620** 0.862** 0.807** 0.338** 0.324** 0.161* 0.921** 0.927** 0.757** 0.986** -0.222** 0.166* -0.119ns -0.234** -0.369** 0.492**
x13 0.061ns -0.160* -0.221** -0.194ns 0.060ns 0.079ns 0.114ns -0.266** -0.293** -0.424** -0.415** -0.376** -0.075ns -0.047ns 0.386** 0.045ns -0.100ns
x14 0.549** 0.432** 0.413** 0.421** 0.420** 0.426** 0.289** 0.388** 0.369** 0.189* 0.237** 0.279** -0.006ns -0.418** 0.285** -0.163* 0.241**
x15 -0.807** -0.762** -0.733** -0.750** -0.726** -0.713** -0.417** -0.693** -0.678** -0.498** -0.478** -0.515** 0.148ns -0.548** -0.361** 0.091ns 0.142ns
x16 0.314** 0.043ns -0.088ns -0.055ns 0.200* 0.194* 0.107ns -0.142ns -0.158* -0.252** -0.310** -0.291** 0.322** 0.375** -0.180* 0.405** -0.080ns
x17 -0.329** -0.398** -0.538** -0.527** -0.321** -0.308** -0.162* -0.552** -0.562** -0.511** -0.550** -0.545** 0.317** -0.164* 0.378** 0.458** -0.169*
x18 0.196* 0.261** 0.410** 0.413** 0.270** 0.271** 0.187* 0.415** 0.427** 0.427** 0.402** 0.400** -0.14ns 0.254** -0.202** 0.051ns -0.323**

zUpper and lower diagonal indicate the value in 2011 and 2012, respectively

yAbbreviations are listed in table 1

Eigenvectors for the principal components of traits and correlation coefficient between the traits and principal components.

Trait 2011 2012

Eigenvector Correlation coefficient Eigenvector Correlation coefficient

Prin1z Prin2 Prin1 Prin2 Prin1z Prin2 Prin1 Prin2

PLy 0.197 0.277 0.567** 0.511** 0.236 0.299 0.743** 0.516**
TNN 0.311 0.102 0.895** 0.189* 0.276 0.092 0.870** 0.159*
NR 0.339 -0.015 0.975** -0.028ns 0.310 -0.036 0.979** -0.062ns
NS 0.343 0.012 0.987** 0.023ns 0.313 0.010 0.987** 0.017ns
NPr 0.193 0.369 0.554** 0.680** 0.249 0.280 0.786** 0.484**
NSPr 0.202 0.385 0.582** 0.709** 0.248 0.303 0.781** 0.523**
MNSPr 0.102 0.191 0.292** 0.352** 0.158 0.276 0.498** 0.476**
NSr 0.335 -0.086 0.964** -0.158* 0.305 -0.099 0.963** -0.171*
NSSr 0.334 -0.128 0.960** -0.235** 0.305 -0.121 0.962** -0.209**
MNSSr 0.235 -0.252 0.676** -0.465** 0.247 -0.212 0.779** -0.366**
Sr/Pr 0.292 -0.248 0.841** -0.456** 0.249 -0.316 0.787** -0.545**
SSr/SPr 0.284 -0.285 0.816** -0.525** 0.260 -0.275 0.820** -0.474**
CL -0.023 0.256 -0.067ns 0.472** -0.079 0.330 -0.249** 0.570**
FL 0.178 0.236 0.511** 0.436** 0.153 0.220 0.484** 0.381**
PN -0.116 -0.235 -0.333** -0.434** -0.254 -0.167 -0.801** -0.288**
TGW 0.016 0.394 0.045ns 0.726** -0.017 0.416 -0.053ns 0.718**
RRG -0.148 0.110 -0.425** 0.203** -0.184 0.215 -0.580** 0.370**
Yield 0.176 -0.133 0.507** -0.245** 0.141 -0.072 0.446** -0.125ns
Eigen value 8.27 3.40 9.95 2.98
Contribution (%) 46.0 18.9 55.3 16.6
Cumulative contribution (%) 46.0 64.8 55.3 71.8

zPrin1 and Prin2 indicate the first and second component, respectively

yAbbreviations are listed in table 1

Average and range of traits value of the parents and groups divided by cluster analysis in 2012.

Group n PLz (cm) TNN (mm) NR NS NPr NSPr MNSPr NSr NSSr MNSSr Sr/Pr SSr/SPr CL (cm) FL (cm) PN TGW (g) RRG (%) Yield (g)

Deuraechan 3 26.3ay 1.74a 42a 166a 14a 86a 6.1a 28a 80a 2.9a 1.9ab 0.9bc 69.1a 32.4a 9c 25.0a 87.3ab 535bc
Boramchan 3 21.0d 1.53bc 35bc 129b 11d 60d 5.7d 24ab 69ab 2.8ab 2.3a 1.1a 67.6a 30.3b 11a 24.3a 91.3a 551b
a 108 24.7ab 1.63ab 40ab 156a 13b 79b 6.0ab 27a 78a 2.9a 2.1ab 1.0abc 68.7a 32.3a 9bc 24.2a 84.1b 520c
(20.1~27.1) (1.29~1.88) (26~51) (107~197) (11~15) (65~93) (5.7~6.3) (15~38) (41~118) (2.6~3.1) (1.3~3.1) (0.6~1.6) (62.3~74.3) (28.3~35.7) (8~13) (21.7~27.2) (74.1~93.0) (477~554)
b 42 22.6cd 1.49c 33c 127b 12c 69c 5.8c 21b 58b 2.7b 1.8b 0.8c 69.5a 31.0ab 11ab 24.0a 87.3ab 467d
(19.2~26.5) (1.32~1.77) (24~44) (97~172) (10~14) (57~85) (5.6~6.1) (13~31) (35~90) (2.5~3) (1.2~2.6) (0.5~1.3) (63.5~75.0) (26.9~33.5) (9~12) (21.0~27.1) (69.4~94.9) (410~501)
c 13 23.7bc 1.59bc 40ab 156a 13bc 75bc 5.9bc 28a 81a 2.9a 2.2a 1.1ab 68.2a 31.8ab 10abc 24.2a 83.9c 581a
(20.7~26.5) (1.25~1.80) (27~46) (108~179) (11~14) (62~85) (5.7~6.1) (16~33) (42~102) (2.7~3.1) (1.4~2.7) (0.6~1.4) (63.9~72.1) (28.8~35.0) (8~13) (21.9~25.7) (75.2~92.4) (555~629)
Total 163 24.1 1.59 38.2 149 13 76 5.9 26 73 2.8 2.0 1.0 68.8 31.9 10 24.1 84.9 512
C.V.(%)x 163 8.3 8.2 15.4 15.1 8.7 10.3 3.4 20.0 23.0 3.6 15.0 20.0 3.5 5.0 11.3 4.5 5.3 7.2

zAbbreviations are listed in table 1

yMeans with same letter are not significantly different at p < 0.05 (ANOVA followed by DMRT)

xC.V. = coefficient of variation

Panicle and yield-related traits of parents and selected DH lines at yield trial in 2013.

Var./lines PLz (cm) TNN NR NS NPr NSPr MNSPr NSr NSSr MNSSr Sr/Pr SSr/SPr CL (cm) FL PN TGW (g) RRG (%) MRY (kg/10a)

Deuraechan (DR) 26aby 1.76ab 32cde 122bcd 11abcd 67bcd 5.9bcde 20de 55efgh 2.7ef 1.7ef 0.8ghi 73ab 36.0ab 10efg 25.3bc 91.2a 512b
Boramchan (BR) 21e 1.50def 31cde 111de 9h 51h 5.5h 22cde 60def 2.8cde 2.3abc 1.2bc 71bcd 32.1de 13abc 24.0ef 91.4a 567ab
HR28832-AC60-1-1 23cd 1.62bcde 31cde 117bcde 10fgh 56gh 5.8defg 21cde 61def 2.8c 2.2bcd 1.1cd 72bc 34.2abcde 12bcd 23.7fg 91.9a 596a
(AC60)
HR28832-AC105-1-1 24cd 1.69abc 35abc 132abc 11bcde 65cde 5.8cdefg 23bcd 67bcde 2.8c 2.0de 1.0def 68e 33.5bcde 12abcd 24.3e 84.1bcd 559ab
(AC105)
HR28832-AC107-1-1 26ab 1.68abc 39a 149a 12a 73a 6.0ab 27ab 75ab 2.8cd 2.2bcd 1.0de 73ab 37.2a 9h 25.7ab 83.8bcd 516b
(AC107)
HR28832-AC109-1-1 23de 1.34g 26f 101e 10gh 57g 5.9abc 16f 44h 2.7ef 1.6fg 0.7hi 72bc 31.0e 13ab 24.5de 89.4ab 561ab
(AC109)
HR28832-AC112-1-1 23de 1.47efg 31cde 117bcde 11def 60gef 5.7fg 21de 57efg 2.7def 1.9def 0.9efg 71bcd 32.0de 12cde 23.2g 92.2a 549ab
(AC112)
HR28832-AC123-1-1 27a 1.62bcde 34abc 133ab 12abc 71abc 6.0a 23cd 62cdef 2.8cdef 1.9def 0.9fgh 73abc 32.6cde 10gh 25.7ab 85.3bcd 541ab
(AC123)
HR28832-AC130-1-1 25ab 1.66abc 29ef 115cde 12abcd 67bcd 5.8cdef 18ef 47gh 2.7ef 1.5g 0.7i 73ab 34.1abcde 10fgh 25.5abc 87.6abc 516b
(AC130)
HR28832-AC134-1-1 26ab 1.58cdef 37ab 143a 12ab 72ab 6.0a 25abc 71abcd 2.8bc 2.0cd 1.0def 75a 35.8abc 10gh 25.0cd 82.7cd 500b
(AC134)
HR28832-AC149-1-1 23cd 1.62bcde 35abc 132abc 10efg 58gf 5.7g 25abc 74abc 2.9ab 2.4ab 1.2ab 68de 31.3e 11cdef 23.3g 85.0bcd 564ab
(AC149)
HR28832-AC152-1-1 23cd 1.65abcd 38a 148a 11cde 66cde 6.0ab 27ab 82a 3.0a 2.4ab 1.2ab 72bcd 35.9ab 11cdef 24.0ef 89.5ab 595a
(AC152)
HR28832-AC156-1-1 22de 1.46fg 29def 110de 10fgh 56g 5.7efg 20def 54fgh 2.7cdef 2.0de 0.9defg 71bcd 35.5abc 14a 23.7fg 90.9a 590a
(AC156)
HR28832-AC161-1-1 25bc 1.73abc 38a 146a 11de 63def 5.9bcd 28a 83a 3.0a 2.6a 1.3a 69de 35.1abcd 11defg 25.5abc 84.9bcd 605a
(AC161)
HR28832-AC178-1-1 25abc 1.80a 33bcde 122bcd 11cde 63def 5.8cdefg 22cd 58efg 2.7f 2.0de 0.9defg 70cde 33.8bcde 10gh 26.1a 80.7d 507b
(AC178)
Mean 24 1.61 33 125 11 62 5.8 62 22 2.8 2.0 1.0 71 34 11 24.6 87.5 552
C.V. 7.5 8.7 27.3 28.0 18.2 21.0 5.2 36.4 40.3 7.1 30.0 30.0 3.5 6.9 13.5 4.1 5.3 8.2

zAbbreviations are listed in table 1

yMeans with same letter are not significantly different at p < 0.05 (ANOVA followed by DMRT)

Correlation coefficient among panicle-related traits and agronomic traits in 2013 (n=15).

Traits PLz TNN NR NS NPr NSPr MNSPr NSr NSSr MNSSr Sr/Pr SSr/SPr CL FL PN TGW RRG MRY
(x1) (x2) (x3) (x4) (x5) (x6) (x7) (x8) (x9) (x10) (x11) (x12) (x13) (x14) (x15) (x16) (x17) (x18)

x1 0.612* 0.464ns 0.558* 0.902** 0.907** 0.728** 0.279ns 0.218ns -0.008ns -0.265ns -0.213ns 0.420ns 0.473ns -0.918** 0.802** -0.531* -0.647**
x2 0.568* 0.564* 0.584* 0.519* 0.209ns 0.483ns 0.443ns 0.251ns 0.197ns 0.210ns -0.233ns 0.486ns -0.684** 0.571* -0.457ns -0.289ns
x3 0.984** 0.581* 0.582* 0.423ns 0.975** 0.952** 0.761** 0.647** 0.692** -0.109ns 0.523* -0.570* 0.232ns -0.518* 0.010ns
x4 0.671** 0.687** 0.560* 0.932** 0.914** 0.739** 0.543* 0.588* -0.023ns 0.554* -0.619* 0.301ns -0.532* -0.040ns
x5 0.987** 0.713** 0.384ns 0.320ns 0.059ns -0.232ns -0.182ns 0.341ns 0.549* -0.882** 0.660** -0.548* -0.650**
x6 0.815** 0.390ns 0.334ns 0.092ns -0.232ns -0.175ns 0.393ns 0.558* -0.847** 0.663** -0.539* -0.591*
x7 0.285ns 0.272ns 0.164ns -0.212ns -0.143ns 0.450ns 0.463ns -0.536* 0.541* -0.375ns -0.247ns
x8 0.992** 0.847** 0.798** 0.835** -0.217ns 0.442ns -0.404ns 0.082ns -0.437ns 0.190ns
x9 0.907** 0.835** 0.861** -0.249ns 0.407ns -0.330ns 0.021ns -0.390ns 0.278ns
x10 0.871** 0.857** -0.301ns 0.244ns -0.047ns -0.211ns -0.164ns 0.540*
x11 0.992** -0.471ns 0.102ns 0.108ns -0.318ns -0.103ns 0.598*
x12 -0.426ns 0.157ns 0.060ns -0.269ns -0.148ns 0.563*
x13 0.354ns -0.328ns 0.249ns 0.187ns -0.468ns
x14 -0.466ns 0.380ns -0.160ns -0.158ns
x15 -0.743** 0.551* 0.679**
x16 -0.579* -0.584*
x17 0.409ns

zAbbreviations are listed in table 1

Eigenvectors for the principal components of traits and correlation coefficient between the traits and principal components.

Trait Eigenvector Correlation coefficient

Prin1z Prin2 Prin3 Prin1 Prin2 Prin3

PL 0.275 -0.230 -0.011 0.789** -0.551* -0.014ns
TNN 0.246 -0.015 -0.375 0.707** -0.037ns -0.451ns
NR 0.313 0.172 0.030 0.898** 0.412ns 0.036ns
NS 0.326 0.127 0.087 0.937** 0.306ns 0.105ns
NPr 0.292 -0.198 0.027 0.838** -0.476ns 0.032ns
NSPr 0.293 -0.195 0.108 0.842** -0.468ns 0.130ns
MNSPr 0.228 -0.150 0.342 0.653** -0.359ns 0.411ns
NSr 0.275 0.249 0.027 0.788** 0.598* 0.032ns
NSSr 0.260 0.274 0.053 0.745** 0.659** 0.064ns
MNSSr 0.176 0.329 0.145 0.506ns 0.791** 0.174ns
Sr/Pr 0.103 0.389 -0.036 0.297ns 0.933** -0.043ns
SSr/SPr 0.123 0.377 -0.014 0.352ns 0.906** -0.016ns
CL 0.040 -0.245 0.569 0.114ns -0.589* 0.684**
FL 0.221 -0.034 0.313 0.635* -0.082ns 0.376ns
PN -0.287 0.178 0.112 -0.825** 0.427ns 0.134ns
TGW 0.203 -0.241 -0.227 0.582* -0.579* -0.273ns
RRG -0.225 0.044 0.422 -0.645** 0.105ns 0.507ns
MRY -0.109 0.331 0.181 -0.312ns 0.795** 0.218ns
Eigen value 8.24 5.77 1.45
Contribution (%) 45.8 32.0 0.08
Cumulative contribution (%) 45.8 77.8 85.8

zPrin1 and Prin2 indicate the first and second component, respectively

Yield-related traits of parents and selected DH lines at yield trial in 2014.

Var./lines Heading date (DASz) CL (cm) PL (cm) PN NS RRG (%) TGW (g) RRY (kg/10a) BRR (%) BRY (kg/10a) MRY (kg/10a) Index of MRY (%)

Deuraechan (DR) 107ay 69a 24a 9c 142a 91.1ab 25.8a 641b 81.6b 523b 481b 100
Boramchan (BR) 108a 69a 20c 12a 108b 94.5a 24.4b 736a 82.5ab 607a 558a 116
HR28832-AC60-1-1 108a 66bc 22b 11b 129ab 91.0ab 24.5b 772a 81.0b 625a 575a 119
(AC60)
HR28832-AC152-1-1 108a 68ab 22b 11b 140a 90.5ab 24.7b 744a 81.6b 607a 558a 116
(AC152)
HR28832-AC156-1-1 108a 64c 20c 12a 108b 94.8a 24.0c 741a 83.2a 617a 567a 118
(AC156)
HR28832-AC161-1-1 106a 65bc 22b 11b 143a 86.3b 25.9a 748a 81.1b 607a 559a 116
(AC161)

zAbbreviations are listed in table 1

yMeans with same letter are not significantly different at p < 0.05 (ANOVA followed by DMRT)

Table 1. The abbreviations for panicle and yield-related traits.
Table 2. Anther culture ability and doubled haploid lines derived from the cross of Deuraechan/Boramchan.

Percent plant regeneration is the number of plants regenerated over number of cali plated

Percent green is the number of green plants regenerated over total plants regenerated

Percent albino is the number of albino plants regenerated over total plants regenerated

Double haploid lines are harvested plants having fertile grain

Table 3. Correlation coefficient among panicle and yield related traits in 2011 and 2012 (n=163).

Upper and lower diagonal indicate the value in 2011 and 2012, respectively

Abbreviations are listed in table 1

Table 4. Eigenvectors for the principal components of traits and correlation coefficient between the traits and principal components.

Prin1 and Prin2 indicate the first and second component, respectively

Abbreviations are listed in table 1

Table 5. Average and range of traits value of the parents and groups divided by cluster analysis in 2012.

Abbreviations are listed in table 1

Means with same letter are not significantly different at p < 0.05 (ANOVA followed by DMRT)

C.V. = coefficient of variation

Table 6. Panicle and yield-related traits of parents and selected DH lines at yield trial in 2013.

Abbreviations are listed in table 1

Means with same letter are not significantly different at p < 0.05 (ANOVA followed by DMRT)

Table 7. Correlation coefficient among panicle-related traits and agronomic traits in 2013 (n=15).

Abbreviations are listed in table 1

Table 8. Eigenvectors for the principal components of traits and correlation coefficient between the traits and principal components.

Prin1 and Prin2 indicate the first and second component, respectively

Table 9. Yield-related traits of parents and selected DH lines at yield trial in 2014.

Abbreviations are listed in table 1

Means with same letter are not significantly different at p < 0.05 (ANOVA followed by DMRT)