(Japanese Journal of Conservation Ecology) 20 : 167-179 (2015) 1,2 3 3 3 3 1 2 3 A method for the early detection of vegetation degradation caused by Sika deer Takeshi Osawa 1,2, Gen Igehara 3, Chiaki Ito 3, Shizuka Michimata 3 and Daiki Sugiyama 3 1 National Institute for Agro-Environmental Sciences, 2 Hakone Park Volunteer, 3 Former Hakone Active Ranger Cervus nippon Temminck 5 5 Bray-Cutis index Abstract: Feeding by invasive Sika deer (Cervus nippon Temminck) causes severe damage to natural vegetation, especially on the forest floor. Although several studies have examined the effects of Sika deer, few have focussed on the early stages of deer invasion. Here, we examine the effects of Sika deer feeding on forest floors in Hakone, Kanagawa, Japan, which are in the early stages of deer invasion. We established five deer exclosure fences and monitored differences in vegetation inside and outside the fences from 2010 to 2014. We found clear differences between vegetation inside and outside the exclosures at some sites, but not at others. Thus, our approach was partially effective at detecting the early effects of feeding by Sika deer. Commencing management efforts in the early stages of invasion is a cost-effective strategy for minimizing damage to natural vegetation. Our approach will aid in detecting vegetation damage and establishing a management plan in the early stages of Sika deer invasion. Keywords: Bray-Curtis dissimilarity index, containment, decision-making, invasion, management plan, monitoring Cervus nippon Temminck 2004 http://www.env.go.jp/ press/100922.html 2015 5 1 2010 1 305-8604 3-1-3 National Institute for Agro-Environmental Sciences, 3-1-3, Kannondai, Tsukuba, Ibaraki 305-8604, Japan e-mail: arosawa@gmail.com 2015 5 14 2015 8 24 167
2005 2003 2008 2013 2006 2013 2011 Iijima et al. 2013 Ohashi et al. 2014 2006 2005 2008, 2010 Nagaike et al. 2014 2007 2005 2008, 2010 2007 2003 Nagaike et al. 2014 Akashi et al. 2011 2013 Iijima and Nagaike 2015 2012 2007 2013 Seki et al. 2014 Takagi and Miyashita 2012 2013 2006 2011 Iijima et al. 2013 Ohashi et al. 2014 2009 2013 Iijima and Nagaike 2015 1989 2008 2014 2010 調査地 2,000 2008 2006 2008 2008 2009 2009 http://www. pref.kanagawa.jp/cnt/f986/p10114.html 2015 7 7 柵の設置と調査区の設定 2010 6 5 1 38.1 mm 180 cm 168
The Ecological Society of Japan 植生変化の早期検出 図 1 箱根町の全体図と柵の設置場所 いたずら等を避けるため おおまかな設置場所のみ示した 金のネット 目あい 15 cm を張り巡らせた 10 m 四方 の正方形で シカの成獣であっても内部への侵入は不可 能であると考えられる 図 2a 柵内および柵外の隣接 地に柵と同じ大きさ 10 m 10 m の方形区を設定し それぞれの内部に 2 m 2 m のサブプロットを 5 か所ず つ設置した 図 2b サブプロットは調査のたびに区内 図 2 植生保護柵の全景および 植生調査用プロットの様子 柵の様子がわかるように全景は冬季に 植生の様子がわか るようにプロットは調査時期に撮影したものを示している でランダムに設置した なお 2014 年 11 月から 2015 年 3 月に実施された自 動撮影カメラを利用した調査により これら 5 か所には 2010 年から 2014 年まで毎年 9 月に植生調査を実施し 実際にシカが存在していることが確認されている 環境 た 調査方法は サブプロット内における高さ 50 省関東地方環境事務所 2015 特に長尾峠では 1 日平均 150 cm 程度の 植生調査においてしばしば草本層と呼 0.23 頭と 他 4 地域 1 日平均 0.1 頭未満 に比べて圧 ばれる範囲における植物を対象に 種ごとで地表面の植 倒的に多数のシカが撮影されている 環境省関東地方環 被率 パーセント被度 を記録するものである 調査の 境事務所 2015 単位は 1 とし それ未満の植物については参考として 種名を記録した 調査の実施 解析デザイン 国立公園である箱根地域では多数の認定パークボラン ティアを擁しており ボランティアらは活動の一環とし 調査の主体となったパークボランティアは 地域の植 て 地 域 の 植 物 調 査 を 行 っ て い る 大 澤 猪 原 2008 物について日常的に観察を行っているものの 必ずしも Osawa 2013 このため パークボランティアの多くは 専門的な知識を持つわけではない そのため 植物の同 箱根地域における植物相に関して一定以上の知識を持っ 定能力等がメンバー内で大きく異なると考えられる 植 ている そこで パークボランティアが主体となって植 被率についても 調査者によって判断基準が異なること 生調査を実施することとした パークボランティアをは が考えられる これはボランティア等 市民科学者が主 じめ 地域の一般人による調査は 調査の継続性という 体になって行われる植物調査においてしばしば起こる問 点で大きな利点がある 大澤ほか 2013 題である よって 植生調査結果の解析には これら調 169
5 5 5 0 5 <10 5 10 <15 10 5 Bray-Cutis BC 2 2011 1 C ij 2 i a b 20 10 j 15 a 20-15=5 b 10-15=-5 5 5 C ij 10 S i S j S i+s j=20+10+15+15=60 BC 2 0 1 BC BC 5 5 解析 1 i BC o BC i-o BC i o 5 5C2=10 i-o 5 5 25 i o i-o 1989 2008 2014 o i i-o i o i - o i - i-o o - i-o t 解析 2 5 1 i o i-o BC BC i BC o BC i-o BC BC 0-1 - BC 5 1 2 170
R3.0.2 R development core team 2013 解析 1 BC 3 a-b A-B α-β a-b A-B α-β 5 i o i-o 2014 i o 3 i o i-o i o 3 2011 o i-o 3 3 i i o i-o 3 解析 2 i i-o i-o BC 1 1 i 1 o 1 i-o 1 o i 1 i 1 1 2011 o i-o 2005 1 2010 i o BC o BC 2014 i o i o i-o 171
3 BC 50 75 25 90 10 a-b A-B α-β 5 172
1 Bray-Cutis i p<0.001 n.s. o n.s. - n.s. i-o p<0.001 n.s. i p<0.05 n.s. o p<0.05 n.s. i-o p<0.001 n.s. i n.s. - n.s. o n.s. - n.s. i-o p<0.05 - i n.s. n.s. n.s. o n.s. n.s. n.s. i-o n.s. n.s. n.s. i n.s. - - o n.s. n.s. n.s. i-o n.s. n.s. n.s. n.s. - i-o i o i, Aster ageratoides Turcz. var. sawadanus Kitam. 2014 1 1989 2014 BC BC 1 2014 2015 2015 2 1 2 2005 5 2013 Kishimoto et al. 2010 2013 173
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1 b2.3 http://foj.c.u-tokyo.ac.jp/gl/ 1 0 GreenList Abelia spathulata Siebold et Zucc. 0 1 0 1 1 Acer carpinifolium Siebold et Zucc. 0 0 0 1 0 Acer crataegifolium Siebold et Zucc. 0 0 0 1 0 Acer diabolicum Blume ex K.Koch 0 0 0 1 0 Acer micranthum Siebold et Zucc. 1 0 0 1 0 Acer palmatum Thunb. 1 1 1 1 0 Acer pictum Thunb. subsp. dissectum (Wesm.) H.Ohashi 1 0 0 0 0 Acer pictum Thunb. subsp. dissectum (Wesm.) H.Ohashi f. dissectum (Wesm.) H.Ohashi 1 0 0 1 0 Acer rufinerve Siebold et Zucc. 0 0 0 1 0 Aconitum japonicum Thunb. subsp. japonicum 0 0 0 0 1 Actinidia arguta (Siebold et Zucc.) Planch. ex Miq. 0 0 0 0 1 Ainsliaea apiculata Sch.Bip. 0 1 0 0 0 Ajugoides humilis (Miq.) Makino 0 1 0 0 0 Akebia quinata (Houtt.) Decne. 1 1 1 0 0 Akebia trifoliata (Thunb.) Koidz. 1 1 1 0 1 Alangium platanifolium (Siebold et Zucc.) Harms var. trilobum (Miq.) Ohwi 0 1 0 0 0 Ampelopsis glandulosa (Wall.) Momiy. var. heterophylla (Thunb.) Momiy. 0 0 1 0 0 Amphicarpaea bracteata (L.) Fernald subsp. edgeworthii (Benth.) H.Ohashi 0 1 1 0 0 Angelica hakonensis Maxim. 0 0 0 0 1 Angelica pubescens Maxim. 0 0 1 0 0 Arisaema angustatum Franch. et Sav. 1 1 0 0 1 Aristolochia kaempferi Willd. 1 1 0 0 0 Artemisia indica Willd. var. maximowiczii (Nakai) H.Hara 0 0 0 0 1 Aster ageratoides Turcz. var. ageratoides 1 0 0 1 0 Aster ageratoides Turcz. var. sawadanus Kitam. 0 0 0 0 1 Aster scaber Thunb. 1 0 1 0 0 Astilbe thunbergii (Siebold et Zucc.) Miq. var. thunbergii 0 0 1 0 0 Aucuba japonica Thunb. 1 0 1 0 0 Berberis thunbergii DC. 1 0 0 0 0 Berchemia racemosa Siebold et Zucc. 0 0 1 1 0 Bistorta tenuicaulis (Bisset et S.Moore) Nakai var. tenuicaulis 0 0 0 1 0 Boenninghausenia albiflora (Hook.) Rchb. ex Meisn. var. japonica (Nakai) Suzuki 0 1 0 0 0 Brachypodium sylvaticum (Huds.) P.Beauv. var. miserum (Thunb.) Koidz. 1 0 0 0 1 Calamagrostis hakonensis Franch. et Sav. 1 0 1 0 0 Callicarpa japonica Thunb. 1 0 0 0 0 Cardiocrinum cordatum (Thunb.) Makino var. cordatum 1 0 1 0 0 Carex alterniflora Franch. 0 0 0 1 0 Carex aphanolepis Franch. et Sav. 0 0 1 0 0 Carex candolleana H.Lev. et Van. 1 0 1 0 0 Carex filipes Franch. et Sav. 0 0 1 0 0 Carex hakonemontana Katsuy. 0 0 1 0 0 Carex japonica Thunb. 1 0 0 0 0 Carex multifolia Ohwi 0 1 0 0 0 Carex sachalinensis F.Schmidt var. iwakiana Ohwi 0 0 1 0 0 Carpesium abrotanoides L. 1 0 0 0 0 Carpesium rosulatum Miq. 1 0 0 0 0 Carpinus cordata Blume 0 0 0 1 0 Carpinus japonica Blume 0 0 0 1 0 Carpinus tschonoskii Maxim. 1 0 1 1 0 Celastrus orbiculatus Thunb. 1 1 1 0 1 Chamaele decumbens (Thunb.) Makino 0 1 1 0 1 Chrysosplenium album Maxim. var. stamineum (Franch.) H.Hara 0 0 0 0 1 Circaea erubescens Franch. et Sav. 0 1 0 0 0 Circaea mollis Siebold et Zucc. 0 1 0 0 0 Clematis apiifolia DC. 1 0 1 0 0 Clematis apiifolia DC. var. biternata Makino 1 0 1 0 0 Clematis japonica Thunb. 1 0 0 0 1 Clematis terniflora DC. 1 0 1 0 0 Clematis tosaensis Makino 0 0 0 1 0 Clerodendrum trichotomum Thunb. 0 1 0 0 0 Clethra barbinervis Siebold et Zucc. 0 0 0 1 1 Clinopodium multicaule (Maxim.) Kuntze 0 1 0 1 0 Cocculus trilobus (Thunb.) DC. 1 0 1 0 0 Comanthosphace japonica (Miq.) S.Moore 0 0 1 0 0 Cornus controversa Hemsl. ex Prain 1 0 1 1 0 176
1 GreenList Cornus kousa Buerger ex Hance 0 0 0 1 0 Cryptotaenia canadensis DC. subsp. japonica (Hassk.) Hand.-Mazz. 0 0 1 0 0 Daphne pseudomezereum A.Gray 1 1 0 0 0 Deutzia crenata Siebold et Zucc. 1 0 0 1 1 Dioscorea japonica Thunb. 1 0 1 1 0 Dioscorea tenuipes Franch. et Sav. 1 0 0 0 0 Dioscorea tokoro Makino 1 1 1 1 0 Disporum sessile D.Don ex Schult. et Schult.f. var. sessile 1 1 0 1 1 Dumasia truncata Siebold et Zucc. 1 1 0 0 0 Elaeagnus matsunoana Makino 0 0 0 1 0 Elaeagnus montana Makino 0 0 0 0 1 Elaeagnus umbellata Thunb. 1 0 0 0 0 Elatostema japonicum Wedd. var. japonicum 0 1 0 0 0 Elatostema japonicum Wedd. var. majus (Maxim.) H.Nakai et H.Ohashi 0 1 0 1 0 Eleutherococcus spinosus (L.f.) S.Y.Hu 0 1 0 0 0 Eleutherococcus spinosus (L.f.) S.Y.Hu var. japonicus (Franch. et Sav.) H.Ohba 1 0 0 0 0 Eleutherococcus trichodon (Franch. et Sav.) H.Ohashi 0 0 0 1 0 Euonymus alatus (Thunb.) Siebold 0 0 1 0 0 Euonymus alatus (Thunb.) Siebold f. striatus (Thunb.) Makino 1 1 1 1 1 Euonymus fortunei (Turcz.) Hand.-Mazz. 1 1 1 1 1 Euonymus oxyphyllus Miq. 1 1 0 1 0 Euonymus sieboldianus Blume 1 1 1 0 1 Fagus crenata Blume 1 0 0 1 0 Filipendula multijuga Maxim. 0 0 0 0 1 Fraxinus lanuginosa Koidz. f. serrata (Nakai) Murata 0 0 0 1 1 Galium shikokianum Nakai 0 0 0 1 0 Galium trifidum L. var. brevipedunculatum Regel 0 0 1 1 0 Geranium thunbergii Siebold ex Lindl. et Paxton 0 0 1 0 0 Geum japonicum Thunb. 0 0 1 0 0 Glechoma hederacea L. subsp. grandis (A.Gray) H.Hara 0 0 1 0 0 Gynostemma pentaphyllum (Thunb.) Makino 1 1 1 0 1 Hedera rhombea (Miq.) Bean 1 1 0 0 0 Helwingia japonica (Thunb.) F.Dietr. 1 1 1 1 0 Houttuynia cordata Thunb. 0 1 0 0 0 Hydrangea hirta (Thunb.) Siebold et Zucc. 0 0 0 1 0 Hydrangea involucrata Siebold var. involucrata 0 1 0 0 0 Hydrangea petiolaris Siebold et Zucc. 0 1 0 1 1 Hydrangea scandens (L.f.) Ser. 0 0 0 0 1 Hydrangea serrata (Thunb.) Ser. var. serrata 0 1 0 1 1 Hylodesmum podocarpum (DC.) H.Ohashi et R.R.Mill var. japonicum (Miq.) H.Ohashi 1 1 0 0 0 Ilex crenata Thunb. 1 1 1 1 1 Ilex macropoda Miq. 0 0 0 1 0 Ilex serrata Thunb. 1 0 0 0 0 Isodon umbrosus (Maxim.) H.Hara 0 0 0 1 1 Kerria japonica (L.) DC. 0 1 0 0 0 Kuhlhasseltia nakaiana (F.Maek.) Ormerod 0 1 0 0 0 Laportea bulbifera (Siebold et Zucc.) Wedd. 0 1 0 0 1 Ligustrum obtusifolium Siebold et Zucc. 1 0 1 0 0 Ligustrum tschonoskii Decne. 1 1 1 1 1 Lilium auratum Lindl. var. auratum 1 0 0 0 0 Lindera praecox (Siebold et Zucc.) Blume 1 1 1 1 1 Lindera umbellata Thunb. 1 1 0 1 0 Liriope minor (Maxim.) Makino 1 0 1 0 0 Lonicera gracilipes Miq. var. glabra Miq. 1 0 0 1 0 Lonicera gracilipes Miq. var. gracilipes 1 0 0 0 0 Lonicera japonica Thunb. 1 0 1 0 0 Lonicera ramosissima Franch. et Sav. ex Maxim. 0 0 0 1 1 Luzula plumosa E.Mey. subsp. plumosa 0 0 0 0 1 Lysimachia japonica Thunb. 1 0 0 0 0 Magnolia kobus DC. 1 0 0 0 0 Malus toringo (Siebold) Siebold ex de Vriese 0 0 1 0 0 Miscanthus sinensis Andersson 0 0 1 0 0 Morus australis Poir. 1 1 1 0 0 Neillia incisa (Thunb.) S.H.Oh var. incisa 0 1 1 1 0 Neomolinia japonica (Franch. et Sav.) Honda 0 0 1 0 0 Neottia makinoana (Ohwi) Szlach. 0 1 0 0 0 177
1 GreenList Ophiopogon japonicus (Thunb.) Ker Gawl. var. umbrosus Maxim. 1 0 1 0 0 Ophiopogon planiscapus Nakai 1 0 1 0 0 Oplismenus undulatifolius (Ard.) Roem. et Schult. var. japonicus (Steud.) Koidz. 1 1 1 1 0 Oplismenus undulatifolius (Ard.) Roem. et Schult. var. undulatifolius 0 0 1 0 0 Oxalis griffithii Edgew. et Hook.f. var. kantoensis (Terao) T.Shimizu 0 1 0 1 0 Paederia scandens (Lour.) Merr. 1 1 1 0 0 Panax japonicus (T.Nees) C.A.Mey. var. japonicus 0 0 0 0 1 Parasenecio delphiniifolius (Siebold et Zucc.) H.Koyama 0 1 0 1 0 Persicaria debilis (Meisn.) H.Gross ex W.T.Lee 0 1 0 0 1 Persicaria filiformis (Thunb.) Nakai ex W.T.Lee 0 1 1 0 0 Petasites japonicus (Siebold et Zucc.) Maxim. 0 0 1 0 0 Phryma leptostachya L. subsp. asiatica (H.Hara) Kitam. 1 0 0 0 0 Pieris japonica (Thunb.) D.Don ex G.Don 0 1 0 0 1 Platanthera ussuriensis (Regel et Maack) Maxim. 1 1 0 0 0 Pleioblastus chino (Franch. et Sav.) Makino 0 0 1 0 0 Polygonatum falcatum A.Gray 1 0 0 0 0 Potentilla freyniana Bornm. 1 0 0 0 0 Potentilla hebiichigo Yonek. et H.Ohashi 0 0 1 0 0 Prunus incisa Thunb. 1 1 0 1 1 Pseudopyxis heterophylla (Miq.) Maxim. 0 1 0 0 0 Quercus acuta Thunb. 1 1 0 1 0 Quercus crispula Blume 1 0 0 0 0 Rhamnus japonica Maxim. var. decipiens Maxim. 0 0 1 0 0 Rhododendron dilatatum Miq. 0 0 0 0 1 Rosa multiflora Thunb. var. multiflora 1 1 1 0 0 Rubia argyi (H.Lév. et Vaniot) H.Hara ex Lauener 1 0 1 0 0 Rubus illecebrosus Focke 0 0 1 0 1 Rubus palmatus Thunb. var. coptophyllus (A.Gray) Kuntze ex Koidz. 1 1 1 1 0 Sambucus racemosa L. subsp. sieboldiana (Miq.) H.Hara 0 1 0 0 0 Sanicula chinensis Bunge 1 0 1 0 0 Sasa borealis (Hack.) Makino et Shibata 0 1 0 1 1 Sasa shimidzuana Makino 0 0 0 1 0 Schisandra repanda (Siebold et Zucc.) Radlk. 0 0 0 1 0 Schizocodon ilicifolius Maxim. var. australis T.Yamaz. 0 0 0 0 1 Schizophragma hydrangeoides Siebold et Zucc. 1 1 0 1 1 Skimmia japonica Thunb. var. intermedia Komatsu f. repens (Nakai) Ohwi 1 1 0 1 1 Smilax china L. 1 1 1 1 0 Smilax riparia A.DC. 1 0 1 0 0 Smilax riparia A.DC. var. ussuriensis (Regel) H.Hara et T.Koyama f. stenophylla (H.Hara) T.Koyama 1 0 0 0 0 Staphylea bumalda DC. 1 0 1 0 0 Stellaria diversiflora Maxim. 0 1 0 0 1 Stellaria sessiliflora Y.Yabe 0 0 0 0 1 Stewartia monadelpha Siebold et Zucc. 1 0 0 1 1 Styrax japonica Siebold et Zucc. 1 0 1 1 1 Symplocos coreana (H.Lév.) Ohwi 1 0 1 1 1 Toxicodendron orientale Greene subsp. orientale 1 1 0 1 0 Tripterospermum japonicum (Siebold et Zucc.) Maxim. var. japonicum 1 0 0 1 1 Tylophora aristolochioides Miq. 0 0 0 1 1 Veronica miqueliana Nakai 0 0 0 0 1 Viburnum dilatatum Thunb. 1 1 1 1 1 Viburnum erosum Thunb. 0 0 0 1 0 Viburnum plicatum Thunb. var. tomentosum (Thunb.) Miq. 0 0 1 1 0 Viola bissetii Maxim. 0 1 0 1 0 Viola eizanensis (Makino) Makino 1 0 0 1 0 Viola grypoceras A.Gray 1 0 1 0 0 Viola keiskei Miq. 1 0 1 0 0 Viola obtusa Makino 1 0 1 0 0 Viola shikokiana Makino 0 0 0 1 0 Viola sieboldii Maxim. 1 0 1 0 0 Viola vaginata Maxim. 0 0 1 0 0 Viola verecunda A.Gray 1 0 1 0 0 Wisteria floribunda (Willd.) DC. 1 0 1 0 0 Zanthoxylum piperitum (L.) DC. 1 1 1 0 0 Zanthoxylum schinifolium Siebold et Zucc. 0 0 1 0 0 178
2 Bray-Curtis 2010 i 0.73 ± 0.14 (i)-(i-o)(p<0.05) 0.75 ± 0.19 (o)-(i-o)(p<0.05) 0.64 ± 0.09 n.s. 0.45 ± 0.16 n.s. 0.39 ± 0.13 i-o(p<0.01) (o) 0.79 ± 0.28 0.71 ± 0.16 0.57 ± 0.13 0.69 ± 0.25 0.64 ± 0.11 (i)-(i-o)(p<0.01) i-o 0.91 ± 0.12 0.85 ± 0.11 0.67 ± 0.12 0.62 ± 0.19 0.56 ± 0.17 2011 i 0.7 ± 0.17 (i)-(i-o)(p<0.01) 0.68 ± 0.22 (i)-(i-o)(p<0.001) 0.64 ± 0.18 (o)-(i-o)(p<0.01) 0.54 ± 0.14 n.s. 0.43 ± 0.15 i-o(p<0.05) o 0.56 ± 0.11 0.75 ± 0.19 (o)-(i-o)(p<0.01) 0.47 ± 0.1 0.55 ± 0.13 0.64 ± 0.19 (i)-(i-o)(p<0.05) i-o 0.77 ± 0.21 0.97 ± 0.07 0.72 ± 0.13 0.53 ± 0.16 0.58 ± 0.18 2012 i 0.77 ± 0.23 (o)-(i-o)(p<0.001) 0.72 ± 0.22 (i)-(i-o)(p<0.01) 0.57 ± 0.16 n.s. 0.48 ± 0.15 n.s. 0.37 ± 0.13 i-o(p<0.001) o 0.59 ± 0.14 0.69 ± 0.16 (o)-(i-o)(p<0.001) 0.54 ± 0.19 0.65 ± 0.12 0.73 ± 0.13 (i)-(i-o)(p<0.001) i-o 0.91 ± 0.16 0.92 ± 0.12 0.64 ± 0.16 0.56 ± 0.17 0.65 ± 0.17 2013 i 0.78 ± 0.31 (o)-(i-o)(p<0.01) 0.53 ± 0.26 (i)-(i-o)(p<0.001) 0.43 ± 0.29 n.s. 0.44 ± 0.23 n.s. 0.26 ± 0.1 i-o(p<0.001) o 0.57 ± 0.19 0.64 ± 0.18 (o)-(i-o)(p<0.001) 0.47 ± 0.15 0.65 ± 0.12 0.67 ± 0.16 i-w(p<0.001) i-o 0.85 ± 0.17 0.92 ± 0.11 0.59 ± 0.16 0.59 ± 0.19 0.54 ± 0.15 o-w(p<0.05) 2014 i 0.81 ± 0.09 (i)-(o)(p<0.001) 0.65 ± 0.19 (i)-(i-o)(p<0.001) 0.51 ± 0.24 n.s. 0.49 ± 0.13 n.s. 0.34 ± 0.11 i-o(p<0.01) o 0.54 ± 0.13 (o)-(i-o)(p<0.01) 0.66 ± 0.15 (o)-(i-o)(p<0.001) 0.64 ± 0.24 0.62 ± 0.08 0.63 ± 0.2 (i)-(i-o)(p<0.001) i-o 0.83 ± 0.15 0.92 ± 0.09 0.63 ± 0.17 0.54 ± 0.14 0.64 ± 0.19 (i)-(o) (p) (i)-(i-o) (p) (o)-(i-o) (p) n.s. 179