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1 Journal of Wetlands Research Vol. 17, No. 1, February 2015, pp ISSN (Print) / ISSN (Online) DOI 다양한수위조건에서식물생활형이식물정착에미치는영향 남종민 * 조현승 ** 김재근 *,**, * 서울대학교농업생명과학연구원 ** 서울대학교생물교육과 Effect of Plant Life Cycle on Plant Settlement in Diverse Water Level Jong Min Nam * Hyun Seung Cho ** Jae Geun Kim *,**, * Research Institute of Agriculture and Life Sciences, Seoul National University, Seoul , Korea ** Department of Biology Education, Seoul National University, Seoul , Korea (Received: 21 October 2014, Revised: 28 October 2014, Accepted: 19 December 2014) 요약본연구에서는정수환경에정착한 3 생활형 9 분류군의식물을대상으로 1 년동안정기적으로모니터링을실시하였으며, 이를통해식물생활형이수위조건에따라식물정착에어떠한영향을미치는지밝히고자하였다. 18 개의실험구를셋으로나누어 16 주범람기간동안 0, 20, 60cm 로수위를각각유지하였으며, 범람기간을제외한기간은 0cm 로수위를모두동일하게유지하였다. 1 차년에식재한다년생식물인큰고랭이 (Scirpus tabernaemontani), 줄 (Zizania caduciflora), 부들속 2 종 (Typha angustifolia, T. orientalis) 은종자에의한추가적인정착은없었으며, 수문조건이밀도및초고생장에큰영향을미치지않았다. 그러나 1 년생식물인고마리 (Persicaria thunbergii), 여뀌 (Persicaria hydropiper), 사마귀풀 (Aneilema japonicum) 과 2 년생인벼룩나물 (Stellaria uliginosa), 뚝새풀 (Alopecurus aequalis), 개피 (Beckmannia syzigachne) 는상대적으로수문조건에의한영향을크게받았다. 정수환경에서침수수위및침수기간은상대적으로지하경을형성하지못하는 1, 2 년생식물종들의정착및생육에큰영향을미치는것으로보인다. 핵심용어 : 범람, 수위, 식물생활형, 식물정착 Abstract The purpose of this study is to reveal the effect of plant life cycle to plant settlement by 1 year monitoring. The subjects of monitoring are the plants (3 plant life cycle, 9 taxa) well established in the mesocosm. 18 mesocosms were divided into 3 sets and water levels were maintained at 0, 20 and 60 cm during 16 weeks from mid-may, respectively and at 0 cm except these 16 weeks. Height and population size of transplanted perennial plants (Scirpus tabernaemontani, Zizania caduciflora, Typha (Typha angustifolia and Typha orientalis)) at 1st year were not affected by diverse water level, though any more seedlings of these species were not settled at this condition. In contrast, water level condition strong influenced annual and biennial plant, relatively. As a result, timing and duration of flooding have great effect on successful settlement of annual and biennial plant without rhizome. Key words : Flooding, Plant Life Cycle, Plant Settlement, Water Level 1. 서론 1) 우리나라에서습지는해충과병원균들의발생원으로지목되어식물천이과정이나인위적인간섭에의해훼손되거나소멸되었으나최근에는습지의높은종-다양성, 높은생산성, 수질정화, 홍수방지, 하안침식방지, 지하수량조절등다양한역할들 (Mitsch and Gosselink, 2000) 에대한중요성이대두되어다양한지역에서습지를복원하거나조 To whom correspondence should be addressed. Department of Biology Education, Seoul National University, Seoul, , Korea jaegkim@snu.ac.kr 성하고있다 (Kim, 2003; Ministry of environment, 2004). 그러나효과적인습지의보존과복원및조성을위해서는습지식물의정착과정과적정서식환경이무엇인지정확히알고있어야함에도불구하고, 적정서식환경에대한연구는일부종에한해서만진행되었을뿐정착과정에대한연구는거의이루어지지않았다 (Kwon et al., 2007; Lee et al., 2007; Jeon et al., 2013; Yoon et al., 2011). 그결과습지의복원을위한설계단계에서의목표와실제자연환경에서정착하고우점하는식물종사이에차이가큰것으로나타났다 (Hong et al., 2005; Kim, 1999). 습지식물의정착과생장은수문주기, 영양염류함량, 토양의미지형, 염도, 기질산화환원전위등다양한습지환

2 20 남종민 조현승 김재근 경의영향을받는것으로알려져있으며 (Brock et al. 2005; Hong and Kim 2013; van der Valk 1981; Vivian-smith 1997; Wetzel and van der Valk 1998), 그중에서도수문조건 (flooding regime) 은습지식물의생육에매우중요한영향을미친다 (Mitsch and Gosselink, 2000; Nam et al., 2014). 자연환경에서는수문주기, 범람의빈도등다양한수문조건들이식물군락의생육에영향을미치며, 특정시점의수위보다누적된수문변화에영향이상대적으로큰것으로알려져있다 (Casanova and Brock, 2000; Cronk and Fennessy, 2001). 이러한특성들을고려하면특정시기에정기적으로시행하는야외조사방법보다식물성장과수문조건을조절할수있는습지실험구 (mesocosm) 를이용한연구가효과적이다. 또한습지식물이해당수위환경에서정착하여군집을형성하기위해서는수문내성뿐만아니라식물간경쟁능력도중요하다 (Carter and Grace, 1990). 선행연구들에의하면식물간경쟁능력이수문내성보다도성장에중요한경우가보고되었으며, 각식물종의주요한특징들 ( 생체량, 초고, 생식체량, 생장형, 영양염류흡수능, 뿌리지역산화능 ) 이종간경쟁능력과밀접한관련이있다고하였다 (Cronk and Fennessy, 2001; Gaudet and Keddy, 1988). 식물특징들중에서도식물생활환 (Plant life cycles) 은봄에습지식물의성장기반이무엇인지구별할수있는특성이기때문에매우중요하다. 1년생식물은성장기반이종자이기때문에매년개체군이갱신되어환경조건이급격히변화할경우쇠퇴나번성이매우빠르게진행되는반면다년생식물의경우뿌리나지하경을통해수년간생존할수있는것으로알려져있으며 (van der Vark, 1981), 지하경의중요성은다년생식물이지하경형성후생장량이증가한다는것을통해밝혀진바있다 (Boutin and Keddy, 1993). 이러한선행연구들을고려하면정수환경에서식물생활형이수문환경에따라식물의정착및생육에미치는영향을파악하기위해서는수년간수문환경을동일하게유지시킨후성공적으로정착한종들을대상으로한연구가필요하다. 본연구는 3년간수문환경을일정하게유지한소규모습지실험구에적응하여정착한식물을대상으로연구를진행하였다. 총 3생활형 9분류군의식물을대상으로 1년동안정기적으로모니터링을실시하였으며, 이를통해식물생활형이수위조건에따라식물정착에어떠한영향을미치는지밝히고자하였다. 본연구의결과는습지의복원및조성후식생의변화상을예측하는데큰도움을줄수있을것이다. 2. 연구방법 2.1 실험구의설계 습지실험구를이용하여수위를제외한다른변수조건들을통제하였다. 실험구용기는직경 1m, 높이 0.9m인원통 형태의플라스틱고무통을사용하였다. 매질은논의표토를사용하였으며, 각용기에약 30cm 높이로채웠다. 실험기간동안증발산으로인해손실된수량은수돗물로주기적으로보충하였으며, 강수로인하여설정된수위보다넘치는부분은용기측면의작은구멍을통하여배출되도록하여수위를일정하게유지하였다 년 4월큰고랭이 (Scirpus tabernaemontani), 줄 (Zizania caduciflora), 부들속 2종 (Typha angustifolia, Typha orientalis) 의지하경을실험구에식재하였다. 모든실험구에큰고랭이 10개체, 줄 6개체, 부들속식물 8개체씩지하경을식재하였으며, 종별개체수의비율은야외조사결과를참고하였다. 식재한다년생식물을제외한다른식물종들은매질의토양종자은행과주변지역에서의종자유입을통해정착하였다. 2.2 실험의설계 2006년부터 2009년까지유지된수문조건은 Table 1과같다. 5월중순부터 16주인범람기간동안 WL 1은표층수가거의없도록유지하되수위를지표면에가깝게유지하였으며, WL 2와 3은범람기간동안각각 20cm와 60cm 의수위를유지하였다. 범람기간을제외한나머지기간은모든수위조건을 WL 1 조건과동일하게유지하였다. 처리구의배치는완전임의배치법 (randomized block design) 을사용하였으며각처리구별로 6반복총 18개의실험구를설치하였다. Table 1. Flooding regimes of 3 sets of mesocosms from 2006 to 2009 Label Flooding regime Water level Flooding duration WL 1 No surface water cm - WL 2 Low standing water 20 cm May October WL 3 High standing water 60 cm May October 2.3 식생조사방법식재한다년생식물종 [ 큰고랭이 (Scirpus tabernaemontani), 줄 (Zizania caduciflora), 부들속 2종 (Typha angustifolia, Typha orientalis)] 들과 2008년까지개체수가가장많이발견된 6종 [1년생: 고마리 (Persicaria thunbergii), 여뀌 (Persicaria hydropiper), 사마귀풀 (Aneilema japonicum), 2년생 : 벼룩나물 (Stellaria uliginosa), 뚝새풀 (Alopecurus aequalis), 개피 (Beckmannia syzigachne)] 을대상으로 2009 년 5월부터 2주간격으로종별개체수및초고를조사하였다. 실험조건별평균개체수는 6개의실험구전체를대상으로계산하였으며, 평균초고는해당식물종이발견된실험구만을대상으로계산하였다. 식물종의동정은이창복도감을참조하였다 (Lee, 2003). 한국습지학회제 17 권제 1 호, 2015

3 다양한수위조건에서식물생활형이식물정착에미치는영향 연구결과및논의 3.1 1년생식물 고마리, 여뀌, 사마귀풀 고마리는 WL 1에서평균 20개체내외로유지되었으며, 다른수위조건에비해상대적으로개체수가많았으나최종초고는수위조건에의한영향을받지않았다 (Fig. 1-B). 비슷한초고성장이가능함에도불구하고 WL 2와 3이 WL 1보다개체수가적은이유는 WL 2, 3 조건은침수에의해광합성을할수있는잎과꽃이맺힐수있는공간이제한되어세대가지날수록종자의숫자가감소했기때문이다. 특히, 고마리는이종과를형성하는습생식물로지상부와지하부에각각의종자를맺는것으로알려져있는데 (Choung et al., 2012; Kawano et al., 1990), 자가수분을하는지하부종자의경우습지의지표아래에서종자를맺기때문에상대적으로침수환경에서종자를맺는데큰문제가없으나, 곤충등에의해타가수분을주로하는지상부꽃의경우침수환경에서성숙할수없기때문이다. 반면, 기존연구들에따르면고마리유묘는 4월에발아하며약 10cm이상의수위에서는생존하기힘든것으로알려져있으나 (Choo et al., 2015; Kim et al., 2012), 본실험에서는고마리가 20cm와 60cm의수위를극복하고초고가 90cm 이상으로성장하였다. 이는약 5cm이하의본잎이몇장없는초기유묘를대상으로한기존연구들 (Choo et al., 2015; Kim et al., 2012) 과달리본실험에서는침수되기전 1주차에이미고마리의평균초고가약 30cm로, 침수시기의유묘상태에차이가있었기때문이다. 고마리는수변지역에서주로생육하며, 예측할수없는수위환경에서안정적인종자생산을위하여범람이잦은여름시기를견디고가을에꽃을피워종자를맺는데 (Kawano et al., 1990; Satake et al., 2001), 이러한번식전략이효과적인이유는고마리가생육기간동안침수환경에서장시간버티고성장할수있는능력이있기때문이다. 게다가 WL 2는식물체전체가침수된상황이아니었으며, WL 3에서는빠른초고성장을통해 4주차에식물체일부가침수상태를벗어났다. 이는 WL 2와 3이고마리의초고성장이중단될만큼장기간동안식물체전체가침수된것이아니라는것 을의미한다. 여뀌는고마리와달리 WL 2, 3에서 WL 1보다개체수가많고초고도컸다 (Fig. 1-A). 여뀌가고마리와달리침수에의한피해를받지않는 WL 1이다른조건에비해개체수가적은이유는 1년생과 2년생식물종들대부분이 WL 1 에서개체수및초고생장이좋아종간경쟁이가장컸기때문이다. 또한여뀌는습생식물이지만 (Choung et al., 2012), 고마리와달리침수수위가높을수록최종초고가커서수면위로잎이발달할수있는공간이충분히확보되었고, 꽃은줄기끝부분에형성되기때문에침수에의해꽃이맺힐수있는공간에대한제약이거의없었다. 이처럼침수환경에적응하여 60cm의수위조건에서도생육이가능하다는점은여뀌가수위변화가큰환경에서경쟁력이있다는것을의미하며, 이는수위변화가심한환경에서생육한다는기존연구들과일치하는결과이다 (Lee and Park, 2005; Seo et al., 2012). 사마귀풀은위두종과달리침수정도에따라다른반응을보인다 (Fig. 1-C). WL 2에서는초고성장이촉진되어 4주차부터식물체일부분이수면위로노출되고, 최종적으로는평균초고가약 80cm로다른수위조건에비해가장높은초고를보였다. 그러나 WL 3에서는 6주차까지초고가성장하지만침수상태를벗어나지못했으며, 그결과 6 주부터 12주까지초고가성장하지못하고 14주에는대부분고사하였다. 이처럼침수정도에따라다른반응을보이는이유는사마귀풀종자의휴면이타파되기위해서는 20일이상의침종기간이필요하여 (Im et al., 2007) 다른두종에비해상대적으로발아시기가늦기때문이다. 발아시기가늦다는것은상대적으로유묘가성장할수있는시간이부족하다는것을의미하며, 이는침수가시작되는시기에다른종들에비해유묘의초고가상대적으로작은결과를초래하였다. 이러한결과들은비록사마귀풀의초고범위가 10~30cm으로알려져있지만 (Lee, 2006), 침수환경이유묘의초고성장을촉진하기때문에침수시점에침수환경하에서의잠재적성장능력에따라극복할수있는수위가달라질수있다는것을의미한다. 또한초고성장은정체되어있지만 12주까지개체수가감소하지않는다는것은식물체전체가침수된상태에서약 10주정도버틸수있다는것 Fig. 1. Density and height of annual plants depending on water level. A: Persicaria thunbergii, B: Persicaria hydropiper, C: Aneilema keisak. Vertical bars show standard error.

4 22 남종민 조현승 김재근 을의미하며, 해당기간내에수위가낮아져식물체일부가침수상태를벗어날경우생존이가능할것으로추정된다. 위의결과들을종합해보면 1년생식물들은침수시기및침수수위가식물종의정착및생육과밀접한관련이있다. 1년생식물들은종자로부터 1년의생활사가시작되기때문에침수시기가침수전유묘가성장할수있는시간을결정하며, 유묘의성장단계에따라침수환경에서의잠재적성장능력에차이를보이기때문이다. 또한, 침수수위는침수환경에서초고성장에의해식물체일부가침수환경을벗어날수있느냐를결정하기때문에중요하다. 침수는차광효과처럼단기적으로는식물의초고성장을촉진시키지만약 4주간의단기적초고성장에도불구하고식물체전체가침수상태를벗어날수없을경우성장이정체되어침수상태를벗어날수없으며침수기간이더오랫동안지속될경우종자를맺지못하고생활사가종료되기때문이다 (Kim et al., 2012) 년생식물 뚝새풀, 벼룩나물, 개피본실험에서군집을형성한 2년생식물의경우뚝새풀과벼룩나물처럼 1년생식물에비해상대적으로초기성장및생활사가빠르게진행되는종들이많았다. Fig. 2-A와 Fig. 2-B에서보이는바와같이습생식물인뚝새풀은침수시기보다상대적으로개화시기나종자가성숙되는시기가빨라 WL 2, 3에서도종이유지될수있었으며, 벼룩나물은절대육상식물임에도불구하고 WL 2에서일부개체가발견되었다 (Choung et al., 2012; Kim et al., 1996). 그러나개피의경우개화시기가 5~6월이기때문에침수시기인 5 월중순까지종자가성숙하기에는상대적으로시간이부족했다 (Lee, 2006). 또한, 개피는양생식물로주로건조한개방지에서생육하는종이기때문에 20cm이상의수위가지속적으로유지되는환경에서는생육할수없는종으로추정된다 (Choung et al., 2012). 그결과 WL 1에서는초고가 90cm이상성장하며성공적으로종자가성숙하였으나 WL 2, 3에서는 4년동안수위조건이유지된결과총 12 개의실험구에서 1개체도발견되지않았다 (Fig. 2-C). 이처럼극명한결과를보이는이유는수명이짧은식물종들의경우개체군갱신이빠르게진행되어환경조건에따른쇠퇴 가매우빠르게진행되기때문이다 (van der Vark, 1981). 2년생식물의경우가을에발아하여뿌리와잎이있는상태에서월동하기때문에종자발아부터시작하는 1년생종들에비해초기성장이빠른편이다 (Seong et al., 2012; Kim et al., 2010). 그러나 2년생식물들도개화및종자가성숙되는시기가종별로다르며, 범람이시작되는시기와의상대적차이에따라종자결실률에차이를보인다. 본실험은 5월중순부터침수가시작되기때문에종자를빨리맺는종들도종자가성숙할수있는시간이부족했고, 그결과침수피해를받지않은대조군에비해상대적으로개체군크기가작았다. 그러나장마기간이 6월이후인것을고려하면 (KMA) 2년생식물들의종자를빨리성숙시킴으로써범람의피해를최소화하는전략은자연환경에서매우효과적일것으로판단된다 (Satake et al., 2001). 3.3 다년생식물 줄, 큰고랭이, 애기부들줄은모든수위조건에서다년생식물중가장높은밀도를보였으며, 모든식물종중에서초고가가장큰것으로나타났다 (Fig. 3-A). 다년생식물인줄과큰고랭이및부들속식물 ( 애기부들, 부들 ) 들의지하경을 3:5:4의비율로식재한것과각식물종의주요한특징들 ( 생체량, 초고, 생식체량, 생장형, 영양염류흡수능, 뿌리지역산화능 ) 중에서생체량과초고가종간경쟁에서가장중요한것을고려하면 (Cronk and Fennessy 2001, Gaudet and Keddy 1988), 줄은모든수위조건에서우점한것으로판단되며, 이는정수조건에서는수위조건에상관없이줄이애기부들등다른다년생식물에비해상대적으로생육이빠르고생장량이많다는선행연구결과와일치한다 (Hong et al., 2014). 한편, 줄은 WL 1에서다른조건들에비해상대적으로개체밀도및초고가작았다. 이러한결과는여뀌를제외한다른 1, 2년생식물들이 WL 1 조건에서개체수가가장많았으며, 초고도대체적으로컸다는것을고려하면, 다른수위조건에비해 WL 1이종간경쟁이심하여줄이흡수할수있는양분이상대적으로적었기때문으로추정된다. 큰고랭이의경우모든수위조건에서 16주차에개체수가급감하였는데 (Fig. 3-B), 이는큰고랭이가다른다년생식물종들에비해상대적으로생활사가빨리종료되었기때문이 Fig. 2. Density and height of biennial plants depending on water level. A: Alopecurus aequalis, B: Stellaria alsine, C: Beckmannia syzigachne. Vertical bars show standard error. 한국습지학회제 17 권제 1 호, 2015

5 다양한수위조건에서식물생활형이식물정착에미치는영향 23 Fig. 3. Density and height of perennial plants depending on water level. A: Zizania latifolia, B: Scirpus tabernaemontani, C: Typha angustifolia and Typha orientalis. Vertical bars show standard error. 다. 반면, 부들속식물은생육특성이다른애기부들과부들 2종으로구성되어있어, 다른다년생식물에비해상대적으로개체수및초고의편차가컸다 (Fig. 3-C). 이러한결과는애기부들의경우최대초고로성장한후분얼을시작하지만부들은초고성장도중에분얼을시작하여종에따라시기별개체수의편차가크고, 두종의평균초고는통계적으로차이를보이지않으나분포범위에차이가있어종에따라초고의분포가다르기때문이다 (Inoue and Tsuchiya, 2006; Kim et al. 2002). 다년생식물인줄과큰고랭이및부들속식물은 2006년부터 2009년까지매년꽃이피고종자를맺었음에도불구하고종자를통한유묘의정착은확인되지않았으며이는다년생식물들도침수환경에서는종자정착이힘들다는선행연구와일치하는결과이다 (van der Vark, 1981). 그러나다년생식물들은지하경을통해모든수위조건에서생존하였으며다른생활형식물들에비해수위조건에따른영향이상대적으로크지않았다. 이러한결과는줄과큰고랭이및부들속식물이다양한수위환경에서생육한다고한기존연구와일치하는결과이다 (Kwon et al., 2007; Lee et al., 2007). 다년생식물의지하경은무성번식기관인동시에지하경에저장한양분을바탕으로환경조건이좋지않을시에도생장속도를빠르게하여해당환경을극복할수있게하므로지하경의존재유무는다양한수위환경에적응하는데매우중요한요인으로판단된다 (Grace and Wetzel, 1981; Hong et al., 2012). 4. 결론 다년생식물들은지하경이형성될경우 1, 2년생에비해상대적으로수위변화에대한영향을적게받으므로, 성공적인정착을위해서는지하경의식재가필요하며, 반대로다년생식물의유입을막기위해서는지하경이형성되지않도록종자단계에서수위조절을통해정착을차단해야한다. 1, 2년생식물종들의경우침수수위및기간에의한영향을크게받으며, 그중에서도유묘및개화시기의수위는종의정착및유지에매우큰영향을미치는것으로나타났다. 이러한결과는시기별로수위를조절할수있을 경우 1, 2년생식물종의정착을통제하거나예측할수있다는것을의미한다. 사사 이논문은정부 ( 교육인적자원부 ) 의재원으로한국연구재단 (NRF-2012R1A ) 과환경부 차세대에코이노베이션기술개발사업 ( , 과제명 : 습지생태계조성및자연생태회복기술개발 ) 의지원을받아수행된연구임. References Boutin, C and Keddy, PA (1993). A functional classification of wetland plants, J. of Vegetation Science, 4(5), pp Brock, MA, Nielsen, BL and Crossle, K (2005). Changes in biotic communities developing from freshwater wetland sediments under experimental salinity and water regimes, Freshwater Biology, 50(8), pp Carter, MF and Grace, JB (1990). Relationships between flooding tolerance, life history, and short-term competitive performance in three species of polygonum, American J. of Botany, 77(3), pp Casanova, MT and Brock, MA (2000). How do depth, duration and frequency of flooding influence the establishment of wetland plant communities?, Plant Ecology, 147(2), pp Choo, Y-H, Kim, H-T, Nam, JM, and Kim, JG (2014). Flooding effects on seed production of the amphicarpic plant Persicaria thunbergii, Aquatic Botany, 119(1), pp Choo, YH, Nam, JM, Kim, JH, and Kim JG (2015). Advantages of amphicarpy of Persicaria thunbergi in the early life history, Aquatic Botany, 121, pp doi: /j.aquabot Choung, Y, Lee, WT, Cho, K-H, Joo, KY, Min BM, Hyun, J-O and Lee, KS (2012). Categorizing Vascular Plant

6 24 남종민 조현승 김재근 Species Occurring in Wetland Ecosystems of the Korean Peninsula, Center for Aquatic Ecosystem Restoration. [Korean Literature] Cronk, JK and Fennessy, MS (2001). Wetland Plants: Biology and Ecology, CRC press, Boca Raton, Florida. Gaudet, CL and Keddy, PA (1988). A comparative approach to predicting competitive ability from plant traits, Nature, 334(3), pp Grace, JB and Wetzel, RG (1981). Habitat partitioning and competitive displacement in cattails (Typha): experimental field studies, The American Naturalist, 118(4), pp Hong, M-G and Kim, JG (2013). Inhabitation characteristics of Sphagnum palustre in abandoned paddy terrace wetland: a case report in Ansan, J of Wetlands Research, 15(1), pp [Korean Literature] Hong, M-G, Nam, JM and Kim, JG (2012). Occupational strategy of runner reed (Phragmites japonica Steud.): change of growth patterns with developmental aging, Aquatic Botany, 97(1), pp Hong, MG, Heo, YJ and Kim, JG (2014). The construction and management of artificial wetland using emergent macrophytes for high biomass production, J of Wetlands Research, 16(1), pp [Korean Literature] Hong, S-K, Kang, HJ, Kim, ES, Kim, JG, Kim, CH, Lee, EJ, Lee, JC, Lee, JS, Lim, BS, Choung, YS and Choung, HL (2005). Restoration Ecology and Engineering-Conservation and Management of Habitats and Ecotope, Life Science Publishing Co, Seoul. [Korean Literature] Im, I-B, Kim, S and Kang, JG (2007). Control and occurrence of Aneilema keisak Hassk. in paddy field, Korean J. Weed Science, 27(1), pp [Korean Literature] Inoue, TM and Tsuchiya, T (2006). Growth strategy of an emergent macrophyte, Typha orientalis Presl. in comparison with Typha latifolia L. and Typha angustifolia L., Limnology, 7(3), pp Jeon, SH, Kim, H, Nam, JM and Kim, JG (2013). Habitat characteristics of sweet flag (Acorus calamus) and their relationship with biomass, Landscape and Ecological Engineering, 9(1), pp Kawano, S, Hara, T, Hiratsuka, A, Matsuo, K and Hirota, I (1990). Reproductive biology of an amphicarpic annual Polygonum thunbergii (Polygonaceae): spatio-temporal changes in growth, structure and reproductive components of a population over an environmental gradient, Plant Species Biology, 5(1), pp Kim, C, Shin, H and Choi, H-K (2003). A phonetic analysis of Typha in Korea and far east Russia, Aquatic Botany, 75(1), pp Kim, DH, Choi, H and Kim, JG (2012). Occupational strategy of Persicaria thunbergii in riparian area: rapid recovery after harsh flooding disturbance, J. of Plant Biology, 55(3), pp Kim, H-Q (1999). A study on the criteria of planting substrate for hydrophytes in creating Man-made Wetland, Master s Thesis, Seoul Universiy. [Korean Literature] Kim, JG (2003). Wetland conservation and restoration, Nature Conservation, 123, pp [Korean Literature] Kim, JS, Hwang, IT and Cho, KY (1996). Effect of storage conditions on the dormancy release and the induction of secondary dormancy in weed seeds, Korean J. Weed Science, 16(3), pp [Korean Literature] Kim, S, Ahn S-H, Im, I-B, Cheong, Y-K and Kim, S-J (2010). Effect of seeding methods to the growing Alopecurus aequalis var. amurensis in wheat field of rice-wheat cropping system, Korean J. Weed Science, 30(3), pp [Korean Literature] Korea Meteorological Administration (KMA) (2009). Kwon, GJ, Lee, BA, Nam, JM and Kim, JG (2007). The relationship of vegetation to environmental factors in Wangsuk stream and Gwarim reservoir in Korea: II. Soil environments, Ecological Research, 22(1), pp Lee, BA, Kwon, GJ and Kim, JG (2007). The optimal environmental ranges for wetland plants: II. Scirpus tabernaemontani and Typha latifolia, J. of Ecology and Field Biology, 30(2), pp Lee, CB (2006). Coloured Flora of Korea, Hyangmunsa. [Korean Literature] Lee, Y-S and Park, J-G (2005), Assessment of water quality impact of submerged lakeside macrophyte, J. of Environmental Impact Assessment, 23(4), pp [Korean Literature] Ministry of Environment (2004). Preliminary Environemental Review Services Manual. [Korean Literature] Mitsch, WJ and Gosselink, JG (2000). Wetlands, John Wiley & Sons, New York. Nam, JM, Jeon, S-H, Choi, H and Kim JG (2014). Effects of micro-topography on vegetation pattern in Dunchondong wetland, J. of Wetlands Research, 16(4), pp [Korean Literature] Satake, A, Sasaki, A and Iwasa, Y (2001). Variable timing of reproduction in unpredictable environments: adaption of flood plain plants, Theoretical Population Biology, 60(1), pp Seo, Y-J, Lee, J-B, Seo, D-C, Kang, S-W, Lee, S-G, Choi, I-W, Lim, B-J, Kim, S-D, Heo, J-S and Cho, J-S (2012). Growth characteristics and nutrient contents under dominant submerged plants in flood control 한국습지학회제 17 권제 1 호, 2015

7 다양한수위조건에서식물생활형이식물정착에미치는영향 25 reservoir around Dongbok lake, Korean J. Soil Science and Fertilizer, 45(5), pp [Korean Literature] Seong, K-Y, Park, T-S, Cho, H-S, Seo, M-C and Jeon, W-T (2012). Weed occurrence according to the density of water foxtail in no-tillage seeding rice paddy fields, Korean J. Weed Science, 32(3), pp [Korean Literature] van der Vark, AG (1981). Succession in wetlands: a gleasonian approach, Ecology, 62(3), pp Vivian-Smith, G (1997). Microtopographic heterogeneity and floristic diversity in experimental wetland communities, J. of Ecology, 85(1), pp Wetzel, P and van der Valk, A (1998). Effects of nutrient and soil moisture on competition between shape Carex stricta, shape Phalaris arundinacea, and shape Typha latifolia, Plant Ecology, 138(2), pp Yoon, J, Kim H, Nam, JM and Kim, JG (2011). Optimal environmental range for Juncus effusus, an important plant species in an endangered insect species (Nannopya pygmaea) habitat in Korea, J. of Ecology and Field Biology, 34(2), pp

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09È«¼®¿µ5~152s Korean Journal of Remote Sensing, Vol.23, No.2, 2007, pp.45~52 Measurement of Backscattering Coefficients of Rice Canopy Using a Ground Polarimetric Scatterometer System Suk-Young Hong*, Jin-Young Hong**,