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세대단축을 위한 장일조건에서 트리티케일의 생육특성

차진경1, 박명렬2, 신동진1, 권영호1, 이소명1, 고종민3, 김경민3, 이종희1,*

Growth Characteristics of Triticale under Long-Day Photoperiod for Rapid Generation Advancement

Jin-Kyung Cha1, Myoung-Ryoul Park2, Dongjin Shin1, Youngho Kwon1, So-Myeong Lee1, Jong-Min Ko3, Kyeong-Min Kim3, Jong-Hee Lee1,*
Korean Journal of Breeding Science2 2021;53(3):200-205.
Published online: September 1, 2021

1농촌진흥청 국립식량과학원 남부작물부

2농촌진흥청 국립식량과학원 중부작물부

3농촌진흥청 국립식량과학원

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*Corresponding Author (E-mail: ccriljh@korea.kr, Tel: +82-55-350-1168, Fax: +82-55-352-3059)
• Received: April 19, 2021   • Revised: June 6, 2021   • Accepted: June 22, 2021

Copyright © 2021 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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  • Consumer demand for Triticale cultivars as a winter forage crop in Korea has been increasing because of its ability for high and stable yield. However, more than 10 years are required to develop new varieties with conventional breeding programs. A speed breeding system using long-day photoperiodic treatment has recently been suggested and applied in wheat and barley, but not in the triticale breeding program in Korea. To evaluate the availability of the established speed breeding system for triticale breeding programs in Korea, we used nine domestic triticale cultivars to investigate their growth characteristics under a 22 h photoperiod. The average days to heading (DTH) of the nine cultivars was 38 days, and Gwangyoung and Minpung showed the most delayed DTH at 42 days. Therefore, all nine triticale cultivars were able to shorten the growth duration under the tested photoperiod condition. One productive tiller and more than 10 seeds were obtained from each cultivar. The germination percentage was over 82% when the spikes were harvested 20 days after heading, dried, and chilled for a week to break dormancy. These results suggest that in Korea, the rapid generation advancement system with simple long-day photoperiodic treatment can be applied to triticale breeding programs to reduce the breeding time.
  • Keywords: triticale; breeding; speed breeding; single seed descent
트리티케일은 밀의 품질 및 수량성과 호밀의 내한성 등을 인위적으로 도입한 속간 교잡 작물이다(Lorenz & Pomeranz 1974, Zhu 2018). 트리티케일은 식용⋅사료용⋅바이오 연료용 등 다양한 용도로 사용되며(Biel et al. 2020, Glamočlija et al. 2018, McGoverin et al. 2011, Zhu 2018), 전세계적으로 연간 약 4백만 ha 정도가 재배되고 1,400만 톤이 생산된다(FAOSTAT 2018).
우리나라에서 트리티케일은 단백질 함량과 조사료로서의 영양학적 가치가 높아 주로 동계 사료 작물로 재배되고 있으며(Han et al. 2017), 최근 이상기후에서도 안정적인 생산량을 보여 그 수요는 지속적으로 증가하고 있는 추세이다(Han et al. 2012).
국내에서 육성된 트리티케일 품종은 세영, 조영, 신성 및 광영 등이 있고, 이들 품종의 평균 육성 기간은 15년 정도로(Han et al. 2016, 2017, 2019), 같은 자식성 작물인 벼⋅밀 등에 비해 긴데, 이는 벼나 밀의 경우 전통 육종법과 더불어 세대단축이나 반수체 육종법을 활용하여 육종 연한을 7~9년 정도로 줄이기도 한 반면(Baek et al. 2019, Kang et al. 2012), 트리티케일은 전통 육종 방법만을 이용해 1년에 1세대를 전개하여 계통을 육성하였기 때문이다(Han et al. 2016, 2017, 2019). 현재 국내에서는 트리티케일 품종 육성을 위해 교배 집단 및 후대 계통을 포장에 전개하여 선발하고 있으며, 일반적으로 10월 중하순에 파종하여 이듬해 6월 상중순에 수확하므로 연 1회 세대를 진전할 수 있다.
트리티케일의 육종 연한을 단축하는 방법으로 약배양 또는 소포자 배양 등을 이용한 반수체 육종법이 주로 이용되고 있으나, 다른 작물들과 마찬가지로 백화현상(albinism)이 발생하고, 조직배양 과정에서 돌연변이 발생 등으로 인한 염색체 이수성(aneuploidy)이 빈번하게 나타나는 등 육종 효율이 낮은 문제점이 있다(Oleszczuk et al. 2004, Oleszczuk et al. 2011, Wędzony et al. 2015). Tuvesson et al. (2003)의 연구 결과에 따르면 트리티케일의 약배양을 통해 하나의 이삭에서 정상적인 반수체가 0~31.3개 형성되었으며, 그중 다시 식물체로 재분화하는 비율은 평균 4.9%에 불과하였다. 따라서 다양한 교배 집단을 작성하여 대량으로 계통을 육성하는 국내 육종 현장에서는 실제로 활용하기 어려운 상황이다.
최근 식량 작물의 육종 연한을 단축하기 위해 일장 조절을 통해 출수를 촉진하는 스피드 브리딩 방법이 제안되었다(Watson et al. 2018, Ghosh et al. 2018). Watson et al. (2018) 등은 동계 작물인 춘파용 밀(spring wheat, Triticum aestivum), 듀럼밀(T. durum), 보리(Hordeum vulgare) 등을 기존의 12시간 일장 조건인 온실에서 재배할 경우 연 2-3세대를 전개할 수 있지만, 22시간 장일처리하고 조기수확하여 휴면타파하는 스피드 브리딩 방법을 활용할 경우 연간 최대 6세대까지 진전할 수 있다고 보고하였다. 또한 국내에서도 스피드 브리딩 방법을 활용하여 국내 밀 품종의 세대단축 가능성을 검토하였는데, 조경, 백강 등 춘파형 밀 품종의 경우 파종일로부터 출수기까지 약 44일이 소요되어 장일처리를 이용한 세대단축이 가능하였다(Cha et al. 2020). 스피드 브리딩 방법을 이용하여 1개체 1계통법(SSD, single seed descent)으로 세대를 전개하면 대량의 교배 후대 집단을 유전적으로 빠르게 고정할 수 있을 뿐만 아니라, 유전형 분석(genotyping)⋅유전체 선발(genomic selection) 등의 분자육종 기술과 접목할 경우 품종 개발의 신속성과 효율성을 극대화할 수 있다(Watson et al. 2018).
따라서 본 연구에서는 트리티케일의 육종 연한을 단축하여 신품종을 조기에 보급할 수 있도록 하고자 국내 품종의 장일 처리를 통한 세대단축 가능성을 검토하였다.
시험 재료 및 재배 조건
시험 재료로 광영, 민풍, 세영, 신성, 신영, 신조성, 옹골진, 조성, 조영 등 트리티케일 국내 육성 품종 9종을 공시하였다. 균일한 발아를 위해 각 품종의 종자를 4℃에서 3일 간 침종한 후 상온에서 12시간 정도 발아시켜 72구 육묘상자(34 mL/구)에 구당 1립씩 파종하였다. 토양은 수도용 상토(Punong, Korea)와 원예용 상토(Seoul Bio, Korea)를 2:1 비율로 혼합하여 사용하였다.
일장 조절
식물생장실에서 파종 직후부터 장일 처리 하였으며, 일장은 Watson et al. (2018)의 스피드 브리딩 방법과 동일하게 22시간 명/2시간 암 조건으로 설정하였다. 인공조명은 식물생장용 LED 조명(Full Spectrum LED PLANT 50WB_SPOT_Full)을 사용하였고, 조명은 식물체가 놓인 선반으로부터 2,672 lx 이상의 조도와 54 umol/m2⋅s 이상의 광합성광량자속밀도로 설치되었다. 식물생장실 내의 온도는 명/암 조건에 따라 22℃/17℃로 변온 처리 하였다.
생육 조사 및 발아율 검정
작물학적 특성은 농촌진흥청 연구조사 분석기준(RDA 2012)에 따라 출수일수, 간장, 수장, 수수, 망장 및 1수립수를 조사하였으며, 품종당 3반복으로 측정하였다. 출수일수는 파종기로부터 이삭의 40% 이상이 출현한 날까지의 소요 일수를 계산하였다. 간장은 지표면으로부터 이삭목까지의 길이를, 수장은 이삭목부터 이삭 끝까지의 길이를 조사하였으며, 망장은 이삭 끝부터 이삭 수염의 길이를 측정하였다. 수수는 한 식물체의 이삭 수를 조사하였으며, 1수립수는 하나의 이삭에서 얻은 총 종자 수를 조사하였다.
발아율 조사를 위해 각 품종의 이삭을 출수기로부터 14일, 17일, 20일 후에 각각 수확하여 이삭 채로 35℃에서 4일 간 건조하였다. 건조 후 탈곡한 종자를 페트리 디쉬에 침종하여 4℃에서 3일 간 저온 처리 하였다. 이후 종자를 상온에서 8일 간 발아시킨 후 발아율을 조사하였다.
통계 분석
통계 분석은 SAS 9.2 (SAS Institute In., Cray, NC, USA) 프로그램을 사용하여 α=0.05 유의성 수준에서 분산분석(One-way ANOVA) 후 던컨다중검정(Dancan’s multiple test)하였다.
장일 처리에 따른 국내 품종의 출수 반응 및 생육 특성
트리티케일의 스피드 브리딩 시스템 적용을 위해 22시간 장일 처리한 결과, 공시 품종들의 평균 출수일수는 38일이었으며, 가장 출수가 빠른 품종인 신조성은 파종 후 33일만에 출수하였다. 출수일수가 가장 길었던 품종은 광영과 민풍이며, 파종일로부터 출수기까지 42일이 소요되었다(Table 1, Fig. 1). Watson et al. (2018)은 밀, 보리 등의 춘파 맥류를 여름철 자연광을 이용해 유리온실에서 포트에 재배할 경우 출수까지 70일 이상 소요되는 반면, 인공조명을 이용해 22시간 장일처리할 경우에는 출수까지 30-40일이 소요된다고 하였다. 본 연구 결과는 Watson et al. (2018)의 장일처리 실험과 유사한 결과를 보였으며, 따라서 국내 육성 9품종을 이용하여 교배한 후대 집단은 장일처리에 의한 세대단축이 가능할 것으로 판단되었다. 한편 트리티케일의 경우 국내 육성 품종의 파성은 검정되어 있지 않으나, Ju et al. (2010)은 신영의 파성을 III으로 추정하였는데, 본 시험에서 신영의 출수일수는 40일이었으며, 이는 Cha et al. (2020)의 밀을 이용한 세대단축 시험에서 파성 III 이상인 품종은 출수일수가 75일 이상으로 지연되었다는 보고와는 다소 다른 결과였다. Hwang et al. (1986)은 맥류 파성 정도 분류 시험에서 춘화처리를 하지 않고 24시간 장일처리만 했을 때 밀의 경우 파성 IV 이상인 품종들은 출수하지 않은 반면, 트리티케일은 파성이 IV인 품종의 지엽전개일수가 파성 II인 품종과 10일 정도의 차이만 나타났다고 보고하였는데, 이에 따라 트리티케일의 장일 조건에서 파성에 따른 출수 반응은 밀과 상이한 것으로 생각된다.
장일처리 조건에서의 작물학적 특성을 조사한 결과 간장은 조성이 약 59 cm로 가장 길었고, 민풍, 세영, 신영이 46~47 cm로 가장 짧은 것으로 나타났다(Table 1). 수장은 조영과 신영이 각각 6.2 cm, 6.1 cm로 가장 길었으며, 광영, 세영, 신조성이 4.4~4.7 cm로 가장 짧았다. 망장은 조영이 6.2 cm로 가장 길었고, 세영이 2.0 cm로 가장 짧게 나타났다. 이처럼 장일 조건에서 72구 육묘상자에 재배할 경우 간장, 수장, 망장 등의 작물학적 특성은 품종 간 통계적으로 유의한 차이가 있는 것으로 나타났다. 수수는 모든 품종에서 1개가 얻어져 통계적 차이는 없었으며, 이는 Cha et al. (2020)과 Watson et al. (2018)의 밀을 이용한 세대 촉진 시험 결과와 같았다. 반면 1수립수는 평균 11개로, 조영이 16개로 가장 많고 신영이 7개로 가장 적었으나 분산분석결과에서는 품종 간 유의성이 인정되지 않았고, SSD를 활용하여 세대를 진전하는 데 충분한 종자가 확보되었다.
수확 시기에 따른 종자 특성 및 발아율 변화
트리티케일의 세대단축 시 수확 적기를 구명하고 등숙기 생육 일수를 단축하기 위해 출수 14일, 17일 및 20일 후에 이삭을 수확하였으며, 건조 후 종자 특성과 발아율을 조사하였다. 출수 14일 또는 17일 후 수확한 이삭은 건조 후에도 비교적 녹색을 띤 반면(Fig. 2A, C), 출수기로부터 20일 후에 수확하여 건조한 이삭은 일반적으로 포장에서 성숙한 이삭과 유사한 모습을 보였다(Fig. 2E). 종자는 출수 14일 후에 수확하여 건조할 경우 형태만 갖추고 등숙이 불량하여 충실도가 낮은 상태였다(Fig. 2B). 출수 17일 또는 20일 후 수확하였을 때 건조 전 종자는 형태만 갖추고 등숙하지 않은 상태였으나(Fig. 2G), 건조 후에는 정상적으로 등숙한 종자와 유사한 형태를 나타내었다(Fig. 2D, F).
출수 14일, 17일, 20일 후에 수확할 경우 평균 발아율은 각각 75.8%, 77.5%, 82.1%로 20일 후 수확했을 때 가장 높았다(Table 2). 신성은 출수 20일 후 수확할 경우 100%로 가장 높은 발아율을 보인 반면 광영의 경우 57.1%로 가장 낮은 발아율을 보였는데, 이는 오히려 출수 14일 후 수확했을 때(63.6%)보다 낮은 수치였다. Watson et al. (2018)과 Cha et al. (2020)의 시험에서 밀, 보리 등을 출수 20일 후 수확하였을 경우 대부분의 품종이 80% 이상의 발아율을 보인 것에 비해 본 시험에서 일부 트리티케일 품종의 발아율은 다소 낮았다. 특히 품종 간 발아율 차이가 크게 나타났는데, 신성은 수확 시기 별 평균 발아율 96.5%를 보인 반면 광영은 54.5%로 발아율이 가장 낮았다. 따라서 전체적인 발아율 향상을 위해서는 수확 시기를 1~2주 정도 늦추는 등의 추가적인 실험이 필요하다.
또한, 종자는 수확 후 건조 방법에 따라 발아율의 차이를 보였는데, 수확한 이삭 채로 건조 후 탈곡한 종자는 정상적으로 발아를 하였으나, 수확 직후 이삭에서 분리하여 종자 상태로 건조할 경우 수확시기에 상관없이 건조 후에도 대부분 녹색을 유지하였고, 이 경우에는 저온 처리를 하여도 발아하지 않았다(Fig. 2H). 이에 대한 명확한 원인을 구명되지 않았으며, 추가적인 반복 실험과 종자 생리학적인 연구가 필요한 것으로 판단된다.
장일처리를 이용한 트리티케일의 육종 연한 단축 가능성을 검토하기 위해 조영, 광영, 조성 등 국내 품종 9종을 공시하여 각 품종 별 출수 및 생육 특성을 분석하였다. 22시간 장일조건에서 9품종의 평균 출수일수는 38일이었고, 출수가 가장 느린 품종인 광영과 민풍도 42일 내외로 출수하여 세대단축이 가능하였다. 본 시험에서 조사한 출수일수⋅간장⋅수장⋅망장 등의 작물학적 특성은 각 품종 간 유의한 차이가 있는 것으로 나타났다. 반면 수수의 경우 72구 육묘상자에서 품종에 상관없이 단 한 개의 유효분얼을 가졌는데, 1수립수는 평균 11개로 1개체 1계통법으로 세대를 진전하기에는 안정적인 양의 종자가 확보되었다. 출수기로부터 20일 후 수확하여 건조 및 저온처리 등을 통해 휴면 타파할 경우 평균 발아율은 82.1%였으며, 본 연구 결과는 트리티케일의 육종 연한을 단축하는 데 활용할 수 있을 것으로 생각된다.
본 논문은 농촌진흥청 연구사업(세부과제명: 스피드 육종방법을 활용한 식량작물 세대단축 시스템 구축, 연구과제번호: PJ014502)의 지원에 의해 이루어진 것임.
Fig. 1
Phenotype of the triticale cultivars under 22-hour photoperiod. The photo was taken 41 days after sowing. 1. Gwangyoung 2. Shinjoseong 3. Minpung 4. Saeyoung 5. Shinseong 6. Shinyoung 7. Onggoljin 8. Joseong and 9. Choyoung.
kjbs-53-200-f1.tif
Fig. 2
Spikes and seeds harvested 14 days (A, B), 17 days (C, D, G, H), and 21 days (E, F) after heading. The photos were taken before (G) and after drying at 35℃ for 3 days with (H) or without (A, B, C, D, E, F) threshing.
kjbs-53-200-f2.tif
Table 1
Days to heading and other agronomic traits of the nine triticale cultivars under 22-hour photoperiod condition.
Table 1
Cultivar Days to
heading		 Culm length
(cm)		 Spike length
(cm)		 Awn length
(cm)		 No. of
spikes/plant		 No. of
grains/spike
Choyoung 37.0bcd 52.6c 6.2a 6.2a 1.0ns 16.0ns
Gwangyoung 42.0a 54.5bc 4.6d 3.0bc 1.0ns 10.7ns
Joseong 38.0abc 59.2a 5.9ab 2.3cde 1.0ns 11.7ns
Minpung 42.0a 47.1d 4.9cd 3.0bc 1.0ns 8.0ns
Onggoljin 34.0cd 56.8ab 5.6abc 2.9bcd 1.0ns 9.7ns
Saeyoung 36.0bcd 47.4d 4.4d 2.0e 1.0ns 9.7ns
Shinjoseong 33.0d 52.0c 4.7d 3.6b 1.0ns 14.0ns
Shinseong 37.0bcd 51.1c 5.1bcd 2.2de 1.0ns 13.0ns
Shinyoung 40.0ab 45.6d 6.1a 2.5cde 1.0ns 7.0ns

The different letters are significantly different from each other (p<0.05) according to the DMRT. One-way ANOVA was performed for each characteristic.

Table 2
Different germination percentages with various harvesting timing.
Table 2
Cultivar Harvested day after heading
14 17 20 Average
Seed
total		 Germination rate (%) Seed
total		 Germination rate (%) Seed
total		 Germination rate (%) Seed
total		 Germination rate (%)
Choyoung 30 90.0 34 88.2 32 96.9 96 91.7abc
Gwangyoung 11 63.6 14 42.9 7 57.1 32 54.5e
Joseong 14 64.3 13 92.3 15 66.7 42 74.4cd
Minpung 28 67.9 11 54.5 8 75.0 47 65.8de
Onggoljin 17 76.5 13 76.9 20 80.0 50 77.8bcd
Saeyoung 27 59.3 22 81.8 8 87.5 57 76.2cd
Shinjoseong 13 92.3 16 93.8 16 93.8 45 93.2ab
Shinseong 29 93.1 28 96.4 20 100.0 77 96.5a
Shinyoung 12 75.0 17 70.6 11 81.8 40 75.8cd
Average 181 75.8 168 77.5 137 82.1 486 78.4

The different letters are significantly different from each other (p<0.05) according to the DMRT. One-way ANOVA was performed for each characteristic.

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Growth Characteristics of Triticale under Long-Day Photoperiod for Rapid Generation Advancement
Korean. J. Breed. Sci.. 2021;53(3):200-205.   Published online September 1, 2021
DOI: https://doi.org/10.9787/KJBS.2021.53.3.200
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Growth Characteristics of Triticale under Long-Day Photoperiod for Rapid Generation Advancement
Korean. J. Breed. Sci.. 2021;53(3):200-205.   Published online September 1, 2021
DOI: https://doi.org/10.9787/KJBS.2021.53.3.200
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Growth Characteristics of Triticale under Long-Day Photoperiod for Rapid Generation Advancement
Image Image
Fig. 1 Phenotype of the triticale cultivars under 22-hour photoperiod. The photo was taken 41 days after sowing. 1. Gwangyoung 2. Shinjoseong 3. Minpung 4. Saeyoung 5. Shinseong 6. Shinyoung 7. Onggoljin 8. Joseong and 9. Choyoung.
Fig. 2 Spikes and seeds harvested 14 days (A, B), 17 days (C, D, G, H), and 21 days (E, F) after heading. The photos were taken before (G) and after drying at 35℃ for 3 days with (H) or without (A, B, C, D, E, F) threshing.

Fig. 1

Fig. 2

Growth Characteristics of Triticale under Long-Day Photoperiod for Rapid Generation Advancement

Days to heading and other agronomic traits of the nine triticale cultivars under 22-hour photoperiod condition.

Cultivar Days to
heading		 Culm length
(cm)		 Spike length
(cm)		 Awn length
(cm)		 No. of
spikes/plant		 No. of
grains/spike
Choyoung 37.0bcd 52.6c 6.2a 6.2a 1.0ns 16.0ns
Gwangyoung 42.0a 54.5bc 4.6d 3.0bc 1.0ns 10.7ns
Joseong 38.0abc 59.2a 5.9ab 2.3cde 1.0ns 11.7ns
Minpung 42.0a 47.1d 4.9cd 3.0bc 1.0ns 8.0ns
Onggoljin 34.0cd 56.8ab 5.6abc 2.9bcd 1.0ns 9.7ns
Saeyoung 36.0bcd 47.4d 4.4d 2.0e 1.0ns 9.7ns
Shinjoseong 33.0d 52.0c 4.7d 3.6b 1.0ns 14.0ns
Shinseong 37.0bcd 51.1c 5.1bcd 2.2de 1.0ns 13.0ns
Shinyoung 40.0ab 45.6d 6.1a 2.5cde 1.0ns 7.0ns

Different germination percentages with various harvesting timing.

Cultivar Harvested day after heading
14 17 20 Average
Seed
total		 Germination rate (%) Seed
total		 Germination rate (%) Seed
total		 Germination rate (%) Seed
total		 Germination rate (%)
Choyoung 30 90.0 34 88.2 32 96.9 96 91.7abc
Gwangyoung 11 63.6 14 42.9 7 57.1 32 54.5e
Joseong 14 64.3 13 92.3 15 66.7 42 74.4cd
Minpung 28 67.9 11 54.5 8 75.0 47 65.8de
Onggoljin 17 76.5 13 76.9 20 80.0 50 77.8bcd
Saeyoung 27 59.3 22 81.8 8 87.5 57 76.2cd
Shinjoseong 13 92.3 16 93.8 16 93.8 45 93.2ab
Shinseong 29 93.1 28 96.4 20 100.0 77 96.5a
Shinyoung 12 75.0 17 70.6 11 81.8 40 75.8cd
Average 181 75.8 168 77.5 137 82.1 486 78.4
Table 1 Days to heading and other agronomic traits of the nine triticale cultivars under 22-hour photoperiod condition.

The different letters are significantly different from each other (p<0.05) according to the DMRT. One-way ANOVA was performed for each characteristic.

Table 2 Different germination percentages with various harvesting timing.

The different letters are significantly different from each other (p<0.05) according to the DMRT. One-way ANOVA was performed for each characteristic.

Table 1

Table 2

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