고온에서의벼생리장해및약발달 서은정, 이성곤, 김동헌, 장안철, 서석철, 정미정 국립농업과학원농업생명자원부 *To whom correspondence should be addressed. E-mail: seji00@korea.kr 목차 Ⅰ. 초록 Ⅱ. 서언 Ⅲ. 식물마다다르게느끼는고온의기준 Ⅳ. 식물이고온을느끼는 mechanism Ⅴ. 벼약에서의고온스트레스 Ⅴ.1. 고온에의한꽃의불임 mechanism Ⅴ.2. 고온에서임성에영향을주는수분 (pollination) Ⅴ.3. 고온에서불임을유발하는습도 (humidity) 의역할 Ⅵ. 맺음말 Ⅶ. 사사 Ⅷ. 인용문헌 고온에서의벼생리장해및약발달 _ 서은정외 1
Ⅰ. 초록 온도상승에따른고온스트레스는전세계많은지역에서농업적으로문제가되고있다. 식물의생육과발달에고온은형태적, 생화학적으로변화를일으키고이로인한경제적인손실도불러오고있다. 식물마다각각의고온기준점을지니고있으며기준점을넘게될때단백질의변성, 집적, 세포막지질의유동성증가, 세포막의변형이이루어진다. 많은식물에서산화스트레스와고온에의한반응이밀접하게관련있다는보고가있다. 식물세포가고온에노출되었을때발현이증가되는전사체들중 5% 의전사체가두배혹은그이상으로고온에반응한다고알려져있는데이중일부만이 molecular chaperones 으로만들어진다. 식물이고온에반응할때고온센서역할을한다고알려진고분자들은다양한온도증가를인지해서다르게반응할뿐만아니라고온에특이적으로반응하여발현이증가되는수백개의고온반응유전자의독특한 signal pathway 를조절하는기능을한다. 벼의약은모든발달단계를통틀어서개화 (anthesis 와 fertilization) 와소포자형성기 (microsporogenesis) 단계가가장온도에민감하다고알려져있다. 벼에서약 (anther) 은고온에서화분팽창이저해되면서개약이불량해지고주두에수정되는화분의양이감소되면서수정률이떨어지게되어결과적으로종자생산량이줄어드는데이때고온과더불어습도나수분환경에따라서벼의임성율이영향을받는다. Ⅱ. 서언 온도는지구상의모든생물의생존에영향을미칠수있는가장중요한요소중의하나로서모든생물들은적온에서부터서서히상승하는온도변화를인지하여대사과정을조절하고이로인해세포가입을수있는고온에의한피해를줄이고자하는방향으로점차적으로진화해왔다. 21 세기말까지지구의온도는평균 2-4 (IPCC 2007) 까지상승할것으로예측되었는데이는인류생존과함께발생한문제와자연생태계적인요소에의해발생한것이다. 농업적인측면에서는온실가스 (CHG) 즉이산화탄소 (CO 2 ), 메탄 (CH 4 ), 일산화이질소 (N 2 O) 등이지구온난화에주요한원인으로밝혀졌다 (Maraseni et al, 2009; Smith and Olesen 2010). IRRI (the International Rice Research Institute, Manila, Philippines) 에서보고한바에따르면 1979 년부터 2003 년사이에매년평균최고기온과평균최저기온을비교하여조사해본결과각각 0.35 와 1.13 였다고한다. 이정도의온도상승은대부분의식물에게는중요한생장기간동안고온에노출되고생장에영향을받는온도로간주되지만불행히도아직온도변화에따른영향을정확히예측하기는불가능한현실이다 (Peng et al., 2004; Watanabe and Kume 2009). 여러작물중전세계적으로주요한식량작물인벼 (Oryza sativa) 는수년간모델식물로많은연구가이루어지긴했으나고온의영향에대한연구는아직시작단계이다 (Nagai and Makino, 2009). 본리뷰에서는식물이온도상승에대해서일반적으로어떻게반응하는지를알아보고식량생산과직결되어있는벼의약이고온에반응하는 mechanism 에대해서기술하고자한 고온에서의벼생리장해및약발달 _ 서은정외 2
다. Ⅲ. 식물마다다르게느끼는고온의기준 고온스트레스란식물의생장과발달에비가역적인영향을일으키는데충분한정도의일정시간과기준이상의온도상승에노출되었을때식물이받는스트레스를의미한다 (Wahid et al, 2007). 일반적으로는적정온도에서 10-15 상승했을때 heat shock 혹은 heat stress 라고말할수있다. 그러나고온스트레스란것이아주복합적이여서, 고온의정도, 기간, 온도상승률등에따라전혀다르게나타날수있다. 일반적으로말하는고온저항성이란식물이고온에서도정상적으로생장하고경제적으로가치가있는생산이가능한능력을보이는것을의미한다 (Wahid et al, 2007). 매우높은고온에서는수분내에세포가손상을입고심지어는세포사멸에이르러서궁극적으로는식물이죽게된다고알려져있는데 (Schöffl et al., 1999) 적절한고온에서손상혹은사멸현상은오랜기간동안고온에노출되어있을때만발생한다. 고온에의한직접적인손상로는단백질의변성, 집적, 세포막지질의유동성증가등을포함한다. 비간접적인혹은천천히발생하는손상으로는엽록체와미토콘드리아의효소가불활성화되고, 단백질합성이저해되며합성된단백질이분해되고세포막이찌그러지기시작한다 (Howarth, 2005). 식물이고온에반응하는온도는종간, 종내에서도차이가있는데식물에따른고온의기준점을찾기위해서높은기준점혹은낮은기준점에서의생장발달을연구해왔다. 여기서말하는기준온도란생장에서관찰가능한감소가시작되는매일의평균온도를의미한다. 다시말해서낮은기준점은식물의생장이멈추는온도를의미하고마찬가지로높은기준점도역시생장이멈추는기준이되는것이다. 기준이되는온도는식물마다차이가있는데일반적으로동절기작물의경우는 0 가낮은기준점이된다고할수있다 (Miller et al., 2001). 동절기와온대기후에서자라는식물은상대적으로열대작물에비해낮은기준점을가지고있다고할수있다. 높은기준점온도역시종간, 종내에서도차이가있는데지속적으로유지가능한고온기준을정하는것은식물의생장이환경의영향에따라달라지기때문에어렵다고볼수있다. 예를들어토마토의경우생육적온인 35 를넘어서자라게되면종자발아, 실생묘의영양생장. 개화, 착과, 과실성숙등의발달이모두영향을받게된다. 식물에따른고온기준점이 (Table 1) 에제시되어있는데대부분의작물들의기준점이대체적으로열매나종자를생산하는발달시점과일치하는것을볼수있다. 일반적으로고온에대한민감도는특히열대, 아열대기후에서중요한데이지역에서는고온스트레스가밭작물생산에주요한요인이될수있기때문이다 (Wahid et al, 2007). 종자가발달하는과정동안에짧은기간의고온에노출되었을때노화가촉진되고종자착상이나무게가감소되어결과적으로는생산량이줄어든다는보고가있다 (Siddique et al., 1999). 식물이고온환경에처하게되면고온을극복하기위해서자기가가진자원을전환하게되는데이때광합성산물이생식기관으로이동하는것을제한하게된다. 많은식물에서밝혀진또다른주요한고온의영향 고온에서의벼생리장해및약발달 _ 서은정외 3
은고온이개약직전이나개약기간동안에노출되었을때불임을유발한다는것이다. Bean 의일종인 Pulse regume 의경우특히개화기때고온에민감한데 30-35 의고온에단며칠만노출되어도꽃이떨어지고꼬투리가발육이정지되어서심각한수량저하를보이게된다. (Siddique et al., 1999). 일반적으로높거나낮은기준온도는식물이자라는환경에따라차이를보이고생육적온이아닌곳에서재배될경우나새로운품종의경우적절한기준온도를찾아재배하는것이바람직하다. Ⅳ. 식물이고온을느끼는 mechanism 고온스트레스는식물의발달, 생장, 생산그리고수확량등거의모든면에서불리한영향을주는요소이다 (Ahuja et al. 2010). 식물에고온은다양한종류의단백질, 막, RNA및세포골격을이루는구조의안전성에각각다르게영향을미치고그결과세포에서효소반응의능률을변화시켜대사활동의불균형상태를만든다 (McClung and Davis, 2010; Ruelland and Zachowski, 2010; Suzuki et al. 2011). 대부분의세포반응은동시에일어나기때문에대사산물의안정상태를파괴하는것이결과적으로는부산물에의한독성물질의축적즉 reactive oxygen species (ROS; 활성산소 ) 를발생시키게된다. (McClung and Davis, 2010; Miller et al., 2009). 많은식물에서산화스트레스와고온에의한반응이밀접하게관련있다는보고가있는데적온에서고온으로온도변화가생길때식물내의 transcriptome, proteome, metabolome, lipidome에급격한변화가발생한다. 즉식물체가자기체내의전사체나단백질, 대사물질과지질의구성을바꿈으로서변화에대응한다는것이다 (Figure 1). 그러한변화는새로운안정적인대사균형상태를만들기위한것으로이러한변화를통해고온에서도식물체가기능을하고생존하여심지어는생산까지가능할수있게한다. 주요한 molecular chaperones family, 예를들어 heat shock protein (HSP) HSP100s, HSP90s, HSP60s, HSP40s, smallhsps (shsp) 이식물대사과정에서고온에대한영향을전달하는데중요한역할을한다고알려져있지만 (Queitsch et al., 2000; Su and Li, 2008; Yamada et al., 2007; Dafny-Yelin et al., 2008) 식물세포가고온에노출되었을때발현이증가되는대다수의전사체들이 molecular chaperones을인지하고있지는않다. 약 5% 의전사체가두배혹은그이상고온에반응한다고보고되었으며 (Larkindale and Vierling, 2008; Qin et al. 2008) 이중일부만이 molecular chaperones으로만들어진다고한다. 나머지전사체들은칼슘시그널전달, 단백질인산화, 광호르몬전달, 당과지질의시그널전달및대사과정, RNA 대사, 번역, 일차및이차대사, 생물적혹은비생물적스트레스반응조절관련전사체들이라고한다. 애기장대가온도가상승되는환경에놓였을때모든세포안에있는거의모든고분자 (macromolecules) 들, 막, 핵산합성관련중합체들이동시에열을감지하고있다고한다 (Mittler et al., 2012). 고분자들의움직임이활발해진다는것은가역적인물리적인변화를수반하게되는데즉막들의유동성, DNA와 RNA의부분적인해체, 단백질 subunit의해체및단백질의소수성 (hydrophobic) 부분이 고온에서의벼생리장해및약발달 _ 서은정외 4
노출되는현상이다 (Fig. 1). 지금까지알려진바로는모든고분자들은일시적인기능을상실함으로써고온센서로서의역할을하게된다 (Saidi and Goloubinoff, 2011). 그래서식물이어떻게고온에반응하는가하는문제는식물세포내에서많은고온반응형고분자의활동을확인하는것으로부터시작된다고할수있다. 고온센서로서의고분자들은다양한온도증가를인지해서다르게반응할뿐만아니라고온에특이적으로반응하여발현이증가되는수백개의고온반응유전자의독특한 signal pathway 도조절하는기능을가진다 (Mittler et al., 2012). 식물에서고온스트레스를인지하는것은다양한경로를통해이루어지는데고온센서분자들은직접적으로는온도인지에의해변하는네개의삼차원구조때문에고온에의해영향을받고또한간접적으로는세포내의다른구성요소들이고온에영향을받을때생기는파급효과로인해영향을받기도한다. 예를들어 membrane protein 은 membrane fluidity 에변화가생김으로써고온에영향을받는다. 식물대사에서고온에관련된변화, 예를들자면대사과정의흐름의변화, 활성산소의축적, 세포로부터 ATP 의방출및에너지레벨의저하등도고온센서분자를활성화시킨다. 또다르게는세포내에서단백질이나 RNA 의 unfolding 으로고온이인지되기도한다 (Saidi et al., 2011). Ⅴ. 벼약에서의고온스트레스 벼는세계적으로주요한식량작물중하나로서특히아시아권에서는주요탄수화물공급원이며최근에는아프리카나라틴아메리카에서도재배가증가하고있다. 작물의생육에있어온도는일장과더불어작물발달에주요한원동력으로작용한다 (Kropff et al., 1995). 최근에전세계기후변화로빠르게상승하는온도가금세기말까지열대및아열대에서의작물수확량에심각하게영향을미칠것으로예측되고있으며 (Battisti and Naylor, 2009) 이에따른영향으로 Peng 등 (2004) 이보고한바에따르면 1 상승에따라약 10% 까지벼수확량이감소할것으로추측한바있다. 벼의적정한생육온도는 27-32 로 (Yin et al., 1996) 고온은벼의대부분의성장단계-출수에서성숙, 수확에영향을미치는데출수기때 40 에노출되면서출수가지연되고감소하며 (Yosida, 1978) 개화시에평균 35 의온도에서약간의노출로도불임이유발된다고한다 (Satake and Yosida, 1978). 따라서식물이고온에노출되는발달단계가식물이입는피해의정도를결정한다고할수있다 (Wahid et al., 2007). 벼에서는모든발달단계를통틀어서개화 ( 개약 : anthesis 와수정 : fertilization) 와소포자형성기 (microsporogenesis) 단계가가장온도에민감하다고알려져있다 (Satake and Yosida 1978; Farrell et al., 2006). 이시기에 41 온도에서 4시간노출될경우비가역적인손상을입게되고식물은완전히불임이되는반면 (IRRI 1976) 개화후 1일혹은개화1일전의경우에는같은온도에노출되어도전혀화분의임성 (fertility) 에는영향을주지않는다고한다 (Yosida et al., 1981). 또고온처리 1시간전에자가수분했거나고온스트레스처리한주두에고온을받지않은화분을처리한경우에도전혀임성에문제가발생하지않았다. 이러한결과로추론해볼때벼이삭이가장고온에민감한시기는개화직전과직후로추정된다. 고온에서의벼생리장해및약발달 _ 서은정외 5
고온에서벼이삭의불임성의차이는벼의유전적변이에따라각각다른온도기준점을가지는데일반적으로 indica 벼들이 japonica 벼에비해더고온에저항성을지니고있다 (Satake and Yosida, 1978; Nakagawa et al., 2003). Ⅴ.1. 고온에의한꽃의불임 mechanism 고온에서벼가불임이되는가장중요한 mechanism 은화분의팽창을저해하면서화분낭 (thecae) 의개약 (dehiscence) 이잘일어나지않게된다는것이다 (Matsui et al., 2000). 일반적으로화분주머니에서화분이팽창하게되고이로인해약의개약이일어나게된다 ((Matsui et al., 1999). Endo 등 (2009) 은고온에노출된화분이정상적인화분과유사하게둥근모양을이루긴하지만화분이주두에붙어발아되는과정에문제를일으킨다고한다. 또다른가설로는꽃에서발현되는호르몬의균형이깨져서고온에서이삭의임성이감소하는보고도있고 (Michael and Beringer 1980), 고온에서는꽃눈에탄수화물이부족해서이거나 (Dinar and Rudich 1985) starch 와 sucrose 생합성효소활성에변화가생겨서일것이라는가설도발표되었다 (Keelimg et al., 1994; Singletary et al., 1994). 외부에서생장호르몬을처리했을때이삭의불임 (sterility) 과화분발달에긍정적인영향이있다고하는데 (Mohammed and Tarpley 2009) 이는외부에서처리한호르몬이내생항산화물질의발현을증가시켜벼에있는세포막들이산화반응에입을수있는손상을방지하는것으로확인되었다. Ⅴ.2. 고온에서임성에영향을주는수분 (pollination) 식물의생식과정에서수분과정 ( 화분생성, 화분의활력, 수분 ) 은매우중요한과정이다. 일반적으로벼에서의웅성생식과정은고온에매우민감하다고알려져있다 (Wassmann et al., 2009). Prasad 등 (2006) 은벼의개화기동안의고온스트레스는화분의생산을감소시키고화분을쓸모없게만든다고한다. 그이유로는고온이화분의팽창을저해하고약의개약이일어나지않아화분방출이거의일어나지않는데이로인해아주작은수의화분만이주두에수분되기때문으로보인다 (Matsui et al., 2000, 2005). 생리적으로고온에서생산이감소된화분의경우소포자의모세포분열에문제를일으키며실제로개약때혹은직후의고온이화분발아와화분관신장을저해한다고한다 (Takeoka et al., 1992). 고온에노출된화분이 10 분내로화분의활력을잃어버린다는보고도있는데 (Song et al., 2001) 결과적으로고온에노출된화분으로인해수분이불량해져수확량의감소로이어진다 (Wassmann and Dobermann 2007). 주두는약에비해고온에덜민감하고고온에노출된주두에정상적인화분을수분했을시임성을회복한다고한다 (Yoshida et al., 1981). Ⅴ.3. 고온에서불임을유발하는습도 (humidity) 의역할 습도는벼의생산에중요한영향을미치는요인으로개화기에고온에노출 고온에서의벼생리장해및약발달 _ 서은정외 6
된이삭에게는부정적인영향을줄수있다 (Yan et al., 2010). Matsui 등 (1997) 이보고한바에따르면벼이삭의임성에미치는고온의영향을습도가조절할수있다고한다. 꽃내에서온도의감소는화분의활성을증가시키는데활성화된화분은습도를흡수하고팽창하게된다. 이는화분의개약을촉진시키고개약을통해화분이주두에앉으면서수정된이삭으로발전하는것이다 (Weerakoon et al., 2008). 즉꽃내에서온도가높아지게되면화분의활력이떨어지게되는데이때상대습도가낮을경우화분내습도를방출함으로써화분내온도를낮출수있지만습도가높을경우에는화분내수분방출이어려워결과적으로화분의활력이저해된다고볼수있다. Ⅵ. 맺음말 식물의생장과생식에고온이미치는영향의심각성에대한한사례로 1980 년과 1988 년에미국에 heat wave 가왔을때총농업생산이 550 억과 710 억달러의손실을남겼으며결과적으로경제적, 사회적으로큰재앙이되었다 (Mittler, R. and E. Blumwald 2010; Ahuja et al., 2010). 또한과거 10 년동안 (1980-2008) 고온은전세계의밀과옥수수의생산량을각각 5.5% 와 3.8% 감소시켰다 (Lobell et al., 2011). 최근우리나라의경우에도평년기온이 2 상승하면 10a 당전국의벼수량이평년보다 4.5% 감소하고 3 상승하면 8.2%, 5 상승하면 14.9% 가감소하는것으로예측되면서 (Shim et al., 2009) 온난화현상에대해심각성을알리고있다. 고온의영향에대한 mechanism 에대한연구가아직미비하지만최근에는수확량에결정적인영향을미치는약의고온노출에대한연구가일본과미국등에서활발히이루어지고있으며벼를주식량작물로삼고있는우리나라에서도이에대한연구가앞으로반드시필요할것으로생각된다. Ⅶ. 사사 본연구는농촌진흥청어젠다과제 ( 과제번호 PJ0086162012) 연구비에의해이루어진것임 Ⅷ. 인용문헌 1. Ahuja, I., R. C. de Vos, A. M. Bones and R. D. Hall 2010. Plant molecular stress responses face climate change. Trends Plant Sci 15: 664-674 2. Ashraf, M. and M. Hafeez, 2004. Thermotolerance of pearl millet and maize at early growth stages: growth and nutrient relations. Biol. Plant. 48: 81 86 3. Battisti, D.S. and R.L. Naylor 2009. Historical warnings of future food 고온에서의벼생리장해및약발달 _ 서은정외 7
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Table 1. Threshold high temperature for some crop plants (Wahid et al., 2007) Figure 1. A schematic model for temperature sensing in plants. Pathway (highlighted in yellow and linked by silid arrows) may serve as the primary heat sensing mechanism of plants. Pathway indicated in light turquoise are thought to trigger different signal transduction events and contribute to plant heat acclimation. (Mittler et al., 2012) Keyword: 약 (anther), heat( 고온 ), 임성 (fertility), 벼 (rice) 고온에서의벼생리장해및약발달 _ 서은정외 14
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