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Research Trends and Prospectives in Maize Anther Cultu


Published online: May 31, 2016

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*Corresponding author (bmlee@dongguk.edu, +82-31-961-5130
• Received: April 8, 2016   • Accepted: May 9, 2016

© The Korean Society of Breeding Science

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  • Anther culture is useful and significant tool for producing haploid or doubled haploid (DH) plants in crop breeding system. Androgenesis is the way of inducing haploid and DH plants from anther (immature pollen) or microspore culture. In vitro androgenesis is efficient technique for introducing complete homozygous lines in one generation, thus less time and expense could be necessary than conventional plant breeding. In maize, anther culture is important system for shortening the breeding cycle and enhancing selection efficiency. Anther culture technique is also applicable to various researches such as molecular genetics, genetic engineering, genomics, and plant biotechnology. We review the past and present studies on anther culture and provide useful information for future researches on androgenesis in maize. The combination of androgenesis with other techniques such as molecular breeding and biotechnology is producing a variety of variety of maize species. In addition, we suggest strategy to develop androgenesis technique adapted to Korean research environment.
약배양은 화분이 들어 있는 약을 채취하여 배양하는 것으로 1964년 흰독말풀에서 어린약을 이용하여 약배양이 시작되었고, 1966년 처음으로 약배양 기술이 보고되었다(Guha & Maheshwari 1964, Guha & Maheshwari 1966). 한편 화분 배양은 꽃밥에 들어 있는 꽃가루를 배양하는 방법으로 약배양과 같이 반수성의 캘러스로부터 반수체 식물을 육성하여 육종에 응용되고 있는 배양 방법이다. 하지만 꽃밥에서 꽃가루를 적출하 거나 꽃가루로부터 캘러스를 유도하여 부정배를 형성시키는 것이 어려우므로 실제로는 약 전체를 배양할 수 있는 약배양을 이용하여 반수체 식물을 만든다. 이때 약(미숙화분) 및 소포자 (microspore)를 이용하여 반수체 식물을 만드는 것을 androgenesis 즉 동정생식이라고 하며, 약배양을 통해 만든 배를 androgenic embryo라고 한다. 동정생식이란 배낭 내부로 들어간 정핵이 난핵과 수정하지 않고 단독으로 분열하여 배를 형성한 것으로 약배양을 인공적인 동정생식이라고 볼 수 있다. 반면 자성생식 (gynogenesis)은 난세포에서 정상적인 수정이 없어도 세포분열 이 일어나 새로운 반수체 개체를 만드는 생식방법이다. 동정생식 과 자성생식은 작물육종가들에게 매우 흥미로운 기술이며, 공통 적으로 육종 프로그램의 시간을 절약하여 한 세대 만에 유전체 동형접합성을 달성시킬 수 있는 유용한 육종법이다(Germanà 2011, Portemer et al. 2015). 자성생식 연구는 1964년에 시작하 여 Tulecke (1964)에 의해 자성배우체로부터 캘러스 형성을 유도하여 처음으로 보고하였으며, 자성생식은 동정생식보다 생 식발달 및 개체 형성이 훨씬 더 느리다. 약배양 기술은 약내의 소포자들이 동정생식을 통하여 소포자 유래 배상구조 (embryo-like structure, ELS)를 유도하는 시스템이며, 배상구 조에서 바로 반수체 식물이 생성되거나 재분화가 가능한 캘러스 를 유도한 후 식물체를 육성할 수 있다. 기본적으로 약배양은 무균배양한 약으로부터 세포분열이 시작하고 캘러스(callus) 또 는 부정배가 형성하여 식물 재분화체를 얻을 수 있다. 일반적으로 기존의 계통육종법은 자가수정을 반복시킴으로써 적어도 6~7세 대가 소요되면서 유전적으로 형질이 고정된 순계인 새로운 품종 이 개발된다(Chang & Coe 2009). 반면 약배양은 반수체 (haploid)나 반수성배(haploidy embryo)를 생산하여 육성과정 을 2~3세대 내에 완전한 순계를 개발하여 육종연한을 단축시키 고 비용도 절감할 수 있으며 유용한 열성 유전자를 가진 식물체를 선발할 수 있다(Geiger & Gordillo 2009). 또한 반수체의 세포를 자연적 및 인위적인 염색체 배가를 통해서 동형접합성 식물체 생산이 가능하다. 특히 근친교배를 여러 번 반복하는 자화수분 작물에서 반수체의 이용은 기존의 재래방법을 대체할 수 있고 작물육종 분야에 큰 잠재력 및 발전 전망을 가지고 있다.
동정생식은 일부 작물 종에서 배양 기술 확립이 잘 구축되어 손쉽게 이용되고 있으며 약배양 및 소포자 배양을 통하여 반수성 세포에서 식물체를 유도하는 연구가 활발히 진행되고 있다(브로 콜리: Farnham 1998, 아스파라거스: Feng et al. 1991, 유채: Gu et al. 2004, 배추: Jo et al. 2012, Seo et al. 2014 담배: Touraev et al. 1996, 포도: Dhekney et al. 2009, 밀: Islam 2010a, b). 반면 옥수수는 동정생식을 이용한 반수체 및 배가반수 체(doubled haploid, DH) 유도 및 형성이 다른 작물들에 비하여 효율성이 낮은 편이며, 약배양 및 소포자 배양은 옥수수 품종 및 유전형에 따라 영향을 많이 받기 때문에 배양 기술이 확립되어 있지 않다(Genovesi & Collins 1982). 하지만 옥수수 육종에서 배가반수체는 육종 효율을 높이는데 중요한 배양 기술이며, 옥수 수 약배양을 이용한 반수체 배양기술은 DH 집단 생산을 위하여 효과적인 기술이다(Mohammadi et al. 2007). 약배양은 공여식 물의 유전자형(genotype), 소포자 발달 단계, 환경요인 등에 영향을 미치며, 특히 약배양에서 약의 유도와 재분화체 생산을 위한 배지 조성 및 배양 조건, 물리적 및 화학적 조건은 매우 중요한 요인이다(Kahrizi et al. 2000, Spitkó et al. 2006). 따라서 약배양은 육종적 이용 가치가 높고 조직배양의 한 기술로서 다양한 품종에 적용 및 응용할 수 있으며, 돌연변이 육종과 형질 전환 기술에도 응용할 수 있는 육종기술로서 매우 유용하다 (Karsai & Bedö 1998).
옥수수 약배양에 미치는 요인
옥수수 약배양은 중국에서 401 research group(1975)에 의해 시작하여 Kuo et al. (1978)이 다양한 옥수수 품종으로부터 약배양 효율성을 검정하여 발표하였으며, 옥수수 소포자(microspore) 배양은 Nitsch (1977)에 의해서 처음으로 보고되었다. 이후 약배 양에 적합한 옥수수 품종을 선발하고 약배양의 효율성을 높이기 위하여 다양한 연구가 시작되었다(Dieu & Beckert 1986, Genovesi & Collins 1982). 일반적으로 옥수수를 포함한 대부분 의 작물에서 약배양 기술 시스템은 대체로 유사한 실험과정과 방법으로 연구가 이루어지고 있으며, Fig. 1에서 약배양의 각 단계별 및 작업 과정을 확인할 수 있다. 옥수수 약배양 기술의 최대 장점은 배가반수체 식물에서 열성유전자의 동형접합성 계통 개발이 전통 육종기술보다 빠르며, 식물 육종가와 유전학자 들에게 매우 가치 있는 기술이란 점이다(Jumpatong et al. 1996). 현재 약배양 기술은 상당히 발전하였으나, 옥수수 약배양 연구는 다른 작물에 비해 효율성이 비교적 낮기 때문에 반수체와 DH 기술을 이용하기 위하여 다양한 조건 및 요인들이 고려된다.
Fig. 1.
Schematic diagram of haploid and doubled haploids selection following anther culture technique in maize.
KJBS-48-2-93_F1.gif
옥수수 유전자형
옥수수 약배양에 알맞은 품종 및 유전자형의 부족으로 약배양 연구가 제한되어 있으며, 이를 보완하기 위하여 약배양과 관련된 계통들을 교배하여 약배양에 적합한 유전자형을 선별하는 것이 가장 중요하다(Genovesi & Collins 1982, Petolino & Jones 1986). 이러한 유전자형은 androgenesis(동정생식)에 대하여 민감한 염색체 단편들이 존재하며, 특히 염색체 1번, 3번, 5번, 7번, 8번 등이 옥수수 소포자로부터 배(embryo) 발생이 유도되 는 것과 관련 있는 것으로 보고되었다(Beaumont et al. 1995, Marchic et al. 1998). 보통 대부분의 옥수수 유전자형은 약배양 유도가 어렵기 때문에 배양 효율성이 낮거나 반응이 전혀 나타나 지 않는 경우가 발생할 수 있다(Genovesi & Collins 1982, Spitkó et al. 2006).
염색체 배가 및 DH 생산
한편 동정생식을 통한 반수체 캘러스와 식물체는 자연적으로 염색체 복제 및 배가가 나타나는 경우는 매우 낮다(Chase 1969). 일반적으로 약배양에서 DH 생산 및 염색체를 배가시키기 위하 여 인위적으로 colchicine(체세포분열 억제)을 주로 사용하고 있다. 하지만 colchicine은 암유발성이 매우 큰 물질이므로 취급 및 사용 후에는 폐기가 중요하다. 이러한 단점을 보완하고 colchicine을 대체하기 위하여 제초제를 사용하여 염색체 배가 연구가 진행되고 있으며, Wan et al. (1991)은 옥수수 약배양에 서 유도된 캘러스로부터 DH 식물을 생산하기 위하여 제초제 (amiprophosmethyl/AMP, pronamide, oryzalin, trifluralin)를 반수체 캘러스에 처리하였으며, AMP와 pronamide를 처리한 실험구에서 배가배수체를 유도하는데 유용하다고 보고하였다. 실제로 제초제가 체세포분열의 효과적인 억제제로 보고되고 있으며(Häntzschel & Weber 2010), 제초제는 colchicine에 비 하여 가격 비용이 저렴하며 독성은 거의 없고 취급이 용이하다.
옥수수 수꽃의 전처리
약배양에서 스트레스 전처리는 약배양 과정 중 소포자 발달단 계를 촉진하는 유도인자로 매우 중요한 요인이며(Pechan & Smykal 2001), 스트레스 전처리를 통해 평범한 꽃가루는 배 발생이 가능한 캘러스를 형성하고 이 과정은 동정생식에 있어서 중요하다(Uváčková et al. 2012). 약배양은 웅성화 배아 (androgenic embryo)로부터 완벽한 동형접합성 반수체 및 배가 반수체 작물의 생산이 가능하다. 약배양의 효율성을 높이기 위하 여 배양 전 수꽃 및 약에 질소가스 노출, 생장호르몬 분무, 수분 부족 등의 다양한 스트레스를 가하여 약으로부터 배 발생을 유도한다. 하지만 대부분의 작물에서 약 및 소포자 배양을 위해 온도 전처리를 통하여 배양의 효율성을 증진시킨다. 배추속 식물 과 담배의 약과 소포자 배양에서 화서 및 화뢰에 32~35°C의 고온처리는 배 발생에 효과적이며(Seo et al. 2014, Touraev et al. 1996), 옥수수와 같은 화본과 및 단자엽 식물은 저온 처리가 더 효과적이다. 특히 옥수수는 약배양 전 수꽃에 주로 7~10°C의 저온 처리(Genovesi & Collins 1982, Hosseini et al. 2015, Ismaili & Mohammadi 2016)를 통하여 약배양에 이용하고 있으며, 과거 몇몇 연구에서는 저온과 고온을 혼합 처리(Coumans et al. 1989, Genovesi 1990)하여 약배양의 효율 성을 높였으나 현재는 저온 처리를 주로 사용한다(Table 1). 한편 약배양 초기과정에 colchicine을 처리하여 약배양의 효율성을 높이는 것을 확인하였으며(Barnabás et al. 1999, Mohammadi et al. 2007, Obert & Barnabás 2004, Wan et al. 1989), 또 다른 연구에서는 저온 처리 및 스트레스 전처리 과정을 생략하고 다양한 배지를 조성하여 바로 약배양 과정을 거쳐 배 발생이 유도되었다 (Barnabás et al. 1987, Dieu & Beckert 1986, Hassan et al. 2001, Nitsch et al. 1982). 하지만 약배양 전에 수꽃 및 화서에 고온과 저온의 전처리 과정이 소포자의 분화 및 발생에 있어서 영향을 미치는 것에 대하여 현재까지 정확하게 밝혀진 연구가 많지 않으나, 밀 약배양 연구에서 생리학적 요인들 (약배양과 재분화 배지)과 고온 등에 의해서 알비노 빈도를 증가 시킬 수 있다고 보고되었다(Indrianto et al. 2001, Islam 2010a). 또한 Torp & Andersen (2009)은 알비노 식물 생산의 빈도율을 감소시키기 위해서 장기적인 저온 처리는 피하라고 제안하였으 며, 단기간(3~4일)의 저온 처리를 통하여 소포자 생존율을 증가 시켜 알비노 빈도율을 줄일 수 있었다.
옥수수 약배양 배지 조성
옥수수 약배양에서는 일반적으로 N6 (Chu et al. 1975)와 YP (Ku et al. 1977, Genovesi & Collins 1982)를 기본배지로 이용하 여 식물생장조절제 조성, sucrose, 활성탄(activated charcoal), gelling agent (phyto agar, phytagel) 및 아미노산 등을 첨가하여 사용하며, 특히 sucrose는 옥수수 약배양 유도에 있어서 매우 중요하다. Barnabás et al. (1987)은 15%의 고농도 sucrose를 첨가하여 옥수수 약배양의 효율성을 확인하였으며, Pescitelli et al. (1990)은 다양한 sucrose의 농도(4.5~18%)를 조성하여 소포자 배양에 검증한 결과 비교적 높은 농도인 8~9.5% 사이에서 소포자 유래 배상구조(embryo-like structure, ELS)가 다른 농도 에서보다 높게 반응하였다. 대부분의 옥수수 약배양은 9%의 sucrose를 주로 사용하지만(Büter et al. 1991, Büter et al. 1993, Saisingtong et al. 1996), 옥수수 유전자형에 따라 비교적 낮은 6%의 농도와 식물생장조절제의 조성으로 약배양의 배 발생 효율성이 높게 나타났다(Hosseini et al. 2014). 최근에는 YP배 지에 6%의 sucrose 및 maltose와 0.0125 M의 polyethylene glycol (PEG)를 첨가하여 약배양의 효율성을 평가하였으며, 결과 적으로 약배양 배지에 낮은 수준의 PEG 첨가는 옥수수 약배양을 향상시킬 수 있는 요인으로 확인되었다(Ismaili & Mohammadi 2016). 또한 Kasha et al. (1990)은 화본과 작물의 약배양 및 소포자 배양에서 maltose을 사용하면 약으로부터 배 발생 및 재분화체 유도 효율성이 높았으며, sucrose와 maltose는 품종의 유전형에 따라 배 발생을 억제하거나 촉진시키는 것으로 나타났 다. 한편 약배양 배지에 활성탄을 첨가하면 배 발생률이 증가하여 대부분의 연구자들이 약배양 및 소포자 연구에 활용하고 있다 (Büter et al. 1993, Büter et al. 1991, Obert & Barnabás 2004, Pescitelli et al. 1990, Saisington et al. 1996, Spitkó et al. 2006, Wan et al. 1989). 활성탄은 배지 내에서 가스 및 고체 화합물을 흡수시키기 때문에 세포 발달에 저해될 수 있는 각종 물질들을 억제할 수 있으며, 식물생장조절제 중 auxin과 cytokinin 등이 과잉으로 작동하면 바로 흡착하여 약배양의 효율성을 높일 수 있다(Büter et al. 1993).
옥수수 약배양과 분자육종 및 유전체 연구
분자육종은 분자생물학적 기술을 적용 및 응용 시키는 것으로 분자마커를 활용하여 유용한 작물들을 선발하는데 이용할 수 있으 며, 주로 QTL mapping, marker-assisted selection (MAS), 유전공 학 및 식물형질전환 등의 다양한 기술이 있다. 특히 약배양과 DH 기술은 옥수수 분자육종에 응용 및 활용되어 새로운 계통을 육성할 뿐만 아니라 시간과 노동 등의 절감으로 전통육종 방법보다 효과적 으로 계통 선발이 가능하다(Prasanna et al. 2012). 또한 분자마커를 이용하여 옥수수 병저항성 및 재해내성 등 다양한 유전자들이 포함 된 형질들을 선발하여 품종 육성이 가능하며, 양친의 완벽한 동형접 합성 동형유전자형은 품종보호에 필요한 DUS (distinctness, uniformity, stability; 구별성, 균일성, 안정성) 조건에 부합하여 새로운 품종 육성 연구에 활용할 수 있다(Geiger & Gordillo 2009, Prasanna et al. 2012). 또한 육성된 계통들을 유지하기 위하여 투자되는 시간과 비용 및 노력 등이 경감하여 분자육종과 작물 유전학 연구를 특정한 계절에 상관없이 비교적 쉽게 수행할 수 있는 가능성이 충분하다(Röber et al. 2005). 따라서 옥수수 육종에 있어서 DH 기술과 분자마커의 조합은 분자 육종 연구의 효율성을 증진하고 분자 유전적 결과물에 대하여 발전 가능한 전략이다 (Prasanna et al. 2012). 또한 유전자 지도를 작성하여 표현형의 차이가 나타난 DH 집단의 양친(모계 및 부계에서 발현되는 유전자 원)을 이용할 수 있으며, 분자마커와 형질 연관 관계 분석을 통해 증명된 분자마커들을 MAS에 응용할 수 있다(Chang & Coe 2009). 특히 유용한 유전자 및 대립형질 등을 활용하여 상대적으로 단순하 게 유전되는 형질을 갖는 DH 계통의 MAS와 seed-chipping 기술을 접목하면 직접적으로 DH 계통의 표현형을 비교 및 확인하는 것보 다 비용이 절감되고 빠르며 훨씬 효과적이다(Prasanna et al. 2012). 최근에는 약배양으로부터 배 발생 유도에 중요한 요인인 저온 전처 리를 프로테오믹스(단백질 유전정보학) 연구에 접목해서 소포자 배발생 유도의 분자 메커니즘을 증명하여 새로운 정보를 제공하고 있다(Uváčková et al. 2012). 이 연구에서 중요한 사실은 약배양 유도 중에 과산화 수소(hydrogen peroxide)를 분해하는 효소인 아스 코베이트 퍼록시다아제(ascorbate peroxidase)가 upregulate하였 고, 이것은 단백질 유전 정보 데이터에서 입증되어 약배양에서 동정생식이 유도되는 동안에 퍼록시다아제 활성이 증가되는 것을 확인하였다. 이러한 단백질 유전 정보 데이터는 옥수수 동정생식 유도 중에 산화성 스트레스(oxidative stress)를 조절하는 효소들에 대한 역할을 밝혀내어 새로운 정보를 제공할 수 있다.
옥수수 약배양과 형질전환 연구
약배양은 대량의 배(embryo)가 유도되고 동일한 유전자를 지닌 반수체 및 단세포이므로 형질전환 연구에 적용하면 유용 유전자 선발 및 새로운 품종 개발을 위하여 효율성 높은 기내 배양 기술이다. 약배양 형질전환 연구는 단 한 세대를 통하여 반수체로부터 배가된 형질전환체를 복구 및 재연이 가능하고, 이러한 형질전환체들은 이식 유전자뿐만 아니라 전체 genome 역시 모두 동형접합성이다. Gaillard et al. (1992)은 옥수수 소포자로부터 형질전환에 적합한 소포자의 시기를 확인하기 위해 서 microinjection을 이용한 형질전환을 시도하였으며, 전제 배 발생 구조에서 초반 4~10일 정도된 소포자가 이보다 늦은 단계의 소포자보다 활력이 더 높아 형질전환에 적합하였다. Jardinaud et al. (1995) 또한 옥수수 소포자를 bombardment 기술에 이용하 여 안정적인 형질전환 기술을 확보하였으며, 형질전환된 반수체 로부터 옥수수 재분화체 및 gus (β-glucuronidase) 유전자를 확인 하였다. 한편 Wan et al. (1995)은 약배양으로부터 유래된 캘래 스를 microprojectile bombardment 기술에 적용하였으며, 약배 양에 적합한 유전자형을 선발하여 옥수수 형질전환 효율성을 높이기 위해서 형질전환 연구에 적합한 캘러스를 유도하여 선발 하였다. 일반적으로 약배양은 유전자형에 대한 의존도가 높고 중요하기 때문에 일부 연구에서는 처음부터 androgenic 반응이 매우 높은 유전자형을 선택해서 제초제 저항성 유전자를 지닌 non-androgenic 형질전환체와 교배하여 약배양 형질전환 연구 를 수행하고 있으며, 이 때 non-androgenic 형질전환체는 약배양 을 위하여 공여식물로 사용하였다(Aulinger et al. 2003). 약과 소포자뿐만 아니라 소포자 유래 액체배양으로부터 원형질체를 분리하여 형질전환 연구에 적용한 결과 유전자가 안정적으로 발현되었고 형질전환 식물체도 확인하였다 (Shukhapinda et al. 1993). 국내에서 옥수수 약배양 및 소포자를 이용한 형질전환 연구는 매우 드물고 일부 연구자들에 의해서만 진행되고 있으며, 국내 연구 시설에 적합한 약배양 형질전환 시스템 또한 전무한 실정이다. 옥수수 약배양으로부터 형질전환 효율성을 높이기 위해서 다양한 옥수수 품종을 선발 및 선별하는 것도 중요하지만, 연구의 효율을 높이기 위해서는 일반적으로 잘 알려진 androgenic 반응성이 높은 유전자형을 선택하는 것이 더 중요하 다. 이러한 효율성 좋은 품종과의 교배를 통해 옥수수 육종 시스 템을 발달시킨다면 약배양을 이용한 형질전환 연구는 국내 분자 육종 기술 활성화에 기여하고 발전 가능성이 높다고 생각한다.
옥수수 약배양 기술을 이용하여 DH 계통을 생산하기 위한 전략은 우선 반수체를 생산하여 반수체를 확인한 후, 염색체를 배가시켜 DH 계통을 재현 및 복구하는 것이 약배양의 기본 연구 과정이다. 하지만 현재 우리나라의 옥수수 약배양 기술은 과채류 및 원예작물, 벼, 보리, 밀 등 다른 화본과 작물들에 비하여 연구 업적 및 결과 등이 많지 않으며, 실제로 옥수수 약배양에 적합한 품종 선별 및 선택이 쉽지 않다. 반면 약배양을 통하여 유도된 반수체와 염색체가 배가된 옥수수는 포장에서 수확한 옥수수보다 일반적으로 약하기 때문에 고온 및 저온, 병해충, 제초제 등의 다양한 환경에 대하여 취약한 것이 사실이 다. 하지만 약배양을 이용한 DH 계통 생산은 유용유전자선발이 가능하고 육종 효율성이 높기 때문에, 최종적으로 육성 품종을 신속하게 개발하여 분자육종과 결합하면 연중 육종 시스템을 가동시킬 수 있는 유용한 기내 배양 기술이다. 이러한 옥수수 약배양은 분자육종 및 생명공학에 활용하여 다양하고 우수한 옥수수 신품종을 육성할 수 있는 가능성 높은 기술이며, 국내에 적합한 옥수수 약배양 기술 개발을 위하여 많은 노력이 필요하다.
본 연구는 농림축산식품부 동남아시아 적응 수출용 옥수수 품종 및 수출 촉진화 기술 개발(213001-04-4-SB920)에 의해 이루어진 것임. 본 논문은 농촌진흥청 공동연구사업(과제번호: PJ00993903)의 지원에 의해 이루어진 것임.
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Research Trends and Prospectives in Maize Anther Cultu
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Research Trends and Prospectives in Maize Anther Cultu
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Fig. 1. Schematic diagram of haploid and doubled haploids selection following anther culture technique in maize.
Research Trends and Prospectives in Maize Anther Cultu