Changes in resource allocation among vegetative organs during the clonal growth of Polygonatum humile (Liliaceae) grown in a temperate forest gap

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1 Min Journl o Eology n Environmnt (17) 41:3 DOI /s y Journl o Eology n Environmnt RESEARCH Opn Ass Chngs in rsour llotion mong vgttiv orgns uring th lonl growth o Polygontum humil (Lili) grown in tmprt orst gp Byong-M Min Astrt Bkgroun: Th im o this stuy ws to xmin th rsour llotion mong th orgns n th morphology o Polygontum humil. Th t wr ollt rom Jun 14 to My 15 in nturl P. humil stn o tmprt orst gp. Rsults: Th ry wight o nw rhizom (R ) shrply rs to th tim o nw shoot sprouting in nxt yr n ws onstnt or ~ 3 yrs. Howvr, ry wight o root on R inrs to th n o growth sson n, thrtr, rs slowly long th tim lps. Th orrltion oiints twn th rhizom n th sizs wr.5 or th lst yr s rhizom (R 1 ) n.6 or R n wr signiint t.1% lvl or R. Th inrs o on plnt l to inrs o th totl r, totl wight, n stm lngth. Morovr, th orgn sizs irns twn two plnts o th on irn wr lmost signiint t 1% lvl. In 11- plnt, th lngth/with rtio n spii l r inrs to th son, rltiv r to th ourth n rltiv ry wight to th ith, n thrtr, rs to th lst. Th irns in th vlus o ths our prmtrs twn two lvs on stm wr lmost signiint t 5% lvl. Conlusion: P. humil xmin show th los rltionships twn th lst yr s rhizom or this yr s onn th shoot systm ( n stm) sizs. Th numr shoot grtly inlun spii r. P. humil s might show ivrs morphology with rnk on stm in orst gp nvironmnt. Kywors: Dry wight, Forst gp,, Polygontum humil, Rhizom, Spii r, Stm Bkgroun In tmprt iuous orsts, unrstory hrous plnts r mor snsitiv thn thos in th opn r to sunlight n us ivrs strtgis or pttion to thir nvironmnt. Sh-tolrnt plnts tht n omplt li yl unr th nopy slowly grow. Howvr, lightmning plnts n gps to grow to ull plnt siz (Lgnr t l. 13). Stuis or th plnt popultion in gp unr th nopy hv n prorm or mny tims (Krn t l. 13). Plnts in prnnil hr popultions in gps ompt with h othr or sunlight n or minrls n wtr in th soil. Enrgy is n or unrstory hrous plnt popultion to sustin thir Corrsponn: mmin@hnmil.nt Dprtmnt o Sin Eution, Dnkook Univrsity, Yongin 1689, Kor sxul or sxul rproution. Howvr, photosynthsis is limit y low irrin, n plnt popultions shoul pply propr strtgis to nsur rproution. Asxul rproution y vgttiv growth is gnrlly us y unrstory lonl hrous plnts in tmprt orsts (Cook 1983, Go t l. 1). In ths plnts, shoot grmintion onsums root systm nrgy in th rly growth sson, whrs photosynthtts prou y lvs r trnsport to th unrgroun prt in th lt growth sson. In ths prosss, root systm iomss o th prvious yr irtly inluns th shoot siz n, in turn, th lttr trmins th nwly grmint ormr. Howvr, it is unlr whthr r or wight hs grtr inlun on iomss o th nw root systm. To urtly stimt iomss o th Th Author(s). 17 Opn Ass This rtil is istriut unr th trms o th Crtiv Commons Attriution 4. Intrntionl Lins ( whih prmits unrstrit us, istriution, n rproution in ny mium, provi you giv pproprit rit to th originl uthor(s) n th sour, provi link to th Crtiv Commons lins, n init i hngs wr m. Th Crtiv Commons Puli Domin Dition wivr ( pplis to th t m vill in this rtil, unlss othrwis stt.

2 Min Journl o Eology n Environmnt (17) 41:3 Pg o 1 root systm, prtinnt initor is n (Klinkhmr t l. 199). Almost ll lvs on wooy twig or on stm o n hrous plnt vlop t th sm tim in tmprt orst (Min 1995, Ishiok t l. 13). Ths lvs r xpos to n lmost th sm mironvironmnt, so thir morphologis or sizs shoul similr. Howvr, th lvs on stm r morphologilly n rmrkly irnt rom h othr (Min 1995). Morovr, it is unlr whthr or not this issimilrity is signiint n whthr it is us minly y gnti tors or y th mironvironmnts. Polygontum humil (Lili) is gnrlly istriut in grsslns or t orst g in Kor (Choung 1991, Jng ), or in grsslns n sn uns in Jpn (Hsgw n Kuo 5). This spis sxully or sxully rprous (L 3). In ition, this spis lowrs in My to Jun (L 3). Asxul rproution is omplish y rhizom (Jng ), whih grows in soil horizontlly. Th rhizom orms on min no pr yr n thus its g n sily ount. Roots r orm on th rhizom n thir numr is rs y thir th with tim. Th shoot pprs on th tip o th rhizom n its rport hight is 6 ± 1.1 m (Jng t l. 1998), 3 5 m (Choung 1991), m (Jng ), or 15 3 m (L 3, Hsgw n Kuo 5). Th lvs o P. humil r ovl n hv n ntir mrgin (Choung 1991). Th rport numr o lvs is 5 8 (Choung 1991) or 13 (Jng t l. 1998). Th lngth n with r 1 ± 1.6 n 4.4 ± 6.7 m (Jng t l. 1998) or 4 7 n m (L 3), rsptivly. Th numr o rhizoms nwly vlop rom shoot is 1 (Choung 1991), (Jng t l. 1998), or 4 (Jng ), n thir lngth is m (Choung 1991). Ths t suggst tht P. humil hs th ro morphologil rng or is vry snsitiv to th mironvironmnt. Th proprtis tht P. humil is ompos o on rhizom on shoot n its r n sily stimt (ovl n ntir) r onvnint or stuying th rltionship twn th root n shoot systms n th min tors o vrition, without stroying th whol plnt. Morovr, nrgy shiting mong roots, rhizom, n shoot ( n stm) in th non-lowring P. humil is simpl n lr, n nsity is vry high in nrrow sps. In Kor, mny stuis o Polygontum hv n prorm in trms o its us in phrmology or griultur (So t l. 11), ut w ologil stuis r vill. Morovr, stuis or P. humil rsrnthrronlytworportsochoung (1991) n Jng t l. (1998) or Kor. Th im o this stuy ws to xmin th hngs in iomss o vgttiv prts n th rspons to th nvironmnt in nturl P. humil popultion, th rltionship twn th numr o lvs in plnt, n thir proprtis long th rnk on stm. Mthos ThlotionothstuyrwstYonghung-ri, Yongwl-up, Yongwl-gun, Gngwon provin ( N, E) (Fig. 1). Th stuy sit ws 385 m.s.l., (slop, 5 ; irtion, 175 ). Th nnul mn ir tmprtur n pripittion in n vrg yr wr 1.8 C n 14.1 mm, rsptivly, n thos in 15 wr 1.5 C n 676 mm on Yongwl Mtorologil Sttion. P. humil grw in gp rt to mov grv to nothr pl (rius, 1 m) n ws momntrily sh y trs in th south (Fig. ). P. humil s pth ws out 5 m 6 m irl. Expt or P. humil, th hr lyr ws 5% in ovrg n ompos o Astr sr, Crx lnolt, Fstu ovin, n Zoysi jponi. Th surrouning orst h our lyrs o vgttion. Th tr lyr ws ompos o Pinus nsilor, Qurus mongoli, n Q. ntt n h 9% ovrg. Thr wr P. nsilor, Q. srrt, Prunus srgntii, n Euonymus siolin n 4% ovrg in th sutr lyr. Thr wr Exohor srrtioli, Pourthi villos, nrhoonron muronultum in th shru lyr n Crx lnolt, Linizi nnri, n Astr sr in th hr on. Th ovrg ws 1% in th ormr n 1% in th lttr, rsptivly. Littr lyr ws 1 m p n A lyr o soil ws 1 m p. Howvr, th soil o th r tht P. humil popultion ws istriut h n istur y work rlot to grv ut h sttl y th tim o th stuy. Plnt shoots wr prtly tth to h othr n thir rhizoms ovrlpp or wr lot losly (Fig., ). Fil survy ws rri out six tims rom Jun, 14 to My 6, 15. All plnts wr ug out togthr with soil rom two 5 5 m qurts, n soil ws slowly wsh out y running wtr to prvnt roots rom rking. Th g o rhizom ws up to 4 yrs. Howvr, th numrs o 4-yr n 3-yr rhizom smpls wr vry smll. Thus, ths smpls wr not nlyz. Only unmg nw rhizoms (R )wrount.rhizoms longr thn 6 m wr us to ollow hngs in root systm iomss uring th growth sson n to ssss th rltionship twn th root n shoot systms. Aoring to Choung (1991), plnts with rhizom longr thn 6 m i not hng thir siz lss in th nxt yr. Howvr, to nlyz th totl numr n lngth o rhizoms pr shoot, ll rhizoms wr ount. In th l, plnts wr ivi into th root n shoot systms. Roots wr sprt oring to th g o rhizom n rhizoms wr numr o R (this yr s rhizom),r 1 (th lst yr s rhizom), n R ( yrs go rhizom). L rnks on stm wr numr rom th lowst or th irst on stm (L 1 ) to th highst (th lst on stm) (Fig. 3). Th numr o lvs on stm ws rom 3 to 13. Howvr, 1- plnt (1-L) n 13- plnt (13-L) wr not suiintly smpl n wr xlu

3 Min Journl o Eology n Environmnt (17) 41:3 Pg 3 o 1 Fig. 1 Stuy r Fig. Nturl Polygontum humil popultion in stuy r on April 6, 15. Totl r, los-up photogrphing, nturl orm o rhizoms in th soil

4 Min Journl o Eology n Environmnt (17) 41:3 Pg 4 o 1 Tl 1 Th numr n rt o rhizoms nwly grmint rom shoot Numr o rhizom 1 3 Totl Rt (numr) 16.5% (118) 8.1% (1) 56.% (395) 1% (715) Fig. 3 Shmti igrm o P. humil morphology. R, L, n F wr rhizom,, n ruit, rsptivly. R,R 1,R n R 3 wr rhizoms grown in this yr (14), lst yr (13), th yr or lst (1), n 3 yrs go (11), rsptivly rom nlysis. Lngth, with n/or r in rhizoms, lvs n stms wr msur, ollow y ovn-rying t 85 C or 48 h n wighing on Hnsung Anlytil n Prision Bln, mol MARK M (1 mg unit) n mol YMJ-C igitl r tr (.1 m unit). Vlus o siz long th rnk wr lult or h rltiv to thos o L 1. Signiin o th irns ws vrii y Stunt t-tst, n orrltion oiints (CCs) twn th two tors wr lult y Prson s qution(y =x +). Rsults Rltionships twn th root n th shoot s systms Th numr o nw rhizoms (R ; > 1 m in lngth) pr shoot ws 1,, or 3. Shoots orming thr R s wr most rprsnt (Tl 1). Th lrgst R ws 5.5 m in lngth, with ry wight o 16 mg. Th mximum root ry wight on n R 1 (th lst yr s rhizom) or R ws 165 mg n thus ws rmrkly smllr thn tht o rhizom. Th mximum umultiv lngth o R pr shoot ws 38.5 m (Tl ). R iomss ws 5.1 ± 11. mg plnt 1 y Jun, 14 n thn ± 6.7 mg plnt 1 y August 8, 14; it ws rltivly onstnt or th shoot sprouting in th spring o th nxt yr. Howvr, tr shoot sprouting, R 1 (R in 14) iomss shrply rs to 84.6 ± 88.5 mg plnt 1 on April 6, 15. This vlu ws similr to tht o R (6.7 ± 6. mg plnt 1 )(R 1 in 14) (6.7 ± 6. mg plnt 1 ; Fig. 4). Atr April 6, 15, R 1 iomss ws onstnt or slightly s. Th pttrn o hngs in rhizom thiknss (ry wight/rhizom lngth) ws similr to tht o rhizom iomss. Howvr, th root s iomss ws smll rom Jun to August o th irst yr; it strongly inrs or shoot sprouting th nxt yr n ontinuously rs thrtr. Th root/rhizom ry wight rtio inrs until th rly growth sson o nxt yr n ws lmost onstnt thrtr. In 15, root n shoot systms ry wights wr 93.8 ± 33.9 n ± 11.1 mg plnt 1 on Mrh 16, 15 (or shoot sprouting) n 91. ± 8.8 n 84.6 ± 88.5 mg plnt 1 on April 6, 15 (tr shoot sprouting), rsptivly (Tl 3). Th totl ry wight o plnt (root systm + shoot systm) ws smllr thn or shoot sprouting (Fig. 5). In th mil o th growth sson (Jun, 14), th most CC (orrltion oiint) vlus twn th R (R n R 1 ) n th shoot systm siz wr ovr.4 n signiint t.1% lvl (Tl 4). Thus, siz ws propr initor or stimting o th totl plnt siz. Th lrgst CC vlu ws.753 twn ry wight o R n tht o lvs (Fig. 6). Rltionships twn th numr n th proprtis o lvs P. humil h shoot (or stm) on R 1 n thr wr rom th plnts with 3 lvs (3-L plnt) to th plnts with 11 lvs (11-L plnt). For th 11 plnt proprtis th Tl Mximum vlus o h proprty in Polygontum humil s root systm grown pr yr ( rhizom mns stion grown or yr) Proprty Vlu Lngth o rhizom 5.5 m Totl lngth o rhizoms grown rom shoot 38.5 m Dry wight o rhizom 16 mg Totl ry wight o rhizoms grown rom shoot 154 mg Dry wight o roots on rhizom 163 mg Totl ry wight o root systm grown rom shoot 1669 mg

5 / Min Journl o Eology n Environmnt (17) 41:3 Pg 5 o 1 ry wight Rhizom R R1 Rhizom(R 1-R ) Rhizom(R -R 1) Jun Jul Aug Mr Apr My R 1 R Rhizom ry wight pr unit lngth (mg/m) R R1 Jun Jul Aug Mr Apr My R 1 R Rhizom(R 1-R ) Rhizom(R -R 1) 15 Rhizom(R 1-R ) Rhizom(R -R 1) 1 Rhizom(R 1-R ) Rhizom(R -R 1) ry wight Root 1 5 R R1 R 1 R ry wight Totl R R1 R R 1 Jun Jul Aug Mr Apr My Jun Jul Aug Mr Apr My systm root root 4 3 R1 Rhizom(R 1-R ) Rhizom(R -R 1) R 1 o Rt 1 R R Jun Jul Aug Mr Apr My Fig. 4 Chngs o th lngth, th ry wight n thiknss o rhizom, root ry wight, totl root systm (root + rhizom) ry wight n th root/root systm ry wight rtio rom Jun, 14 to My 8, 15. Brs init th stnr vition totl r n ry wight pr stm, mn r n ry wight pr, spii r (SLA), lngth n ry wight o R n R 1, lngth n ry wight o stm ssss on Jun, 14, ll vlus inrs with th numr o lvs xpt or SLA (Fig. 7). Morovr, n inrmnt o only on rsult in onspiuous inrs in orgn proprty vlus. O th totl 9 stps rom 3-L plnt to 11-L plnt, th mximum numr o stps with th irn in vlus o plnt proprtis twn two typ plnts signiint t 1% lvl ws 9 or totl r pr stm. Signiint irns t 1% lvl wr rvl t th 8 stps or stm lngth n wight n t 7 stps or th totl wight pr stm n mn r. Thus, th vlus o ths iv proprtis onspiuously inrs with th inrs in numr. Signiint irns t 1% lvl wr osrv t 4 stps or R 1 wight n lngth n t 3 stps or R wight, so ths thr proprtis wr loosly rlt to numr. O th 11 proprtis, only SLA rs with th inrs in numr. SLA signiintly ir t 1% lvl t 5 stps, n no signiint irns in SLA wr oun in plnts tht h < 7 lvs. Tl 3 Dry wight o h plnt orgn in Polygontum humil popultion (unit; mg) Dt Numr Root systm L Stm Shoot systm Mr. 16, ± 6.5 Apr. 6, ± ± ± ± 1.4 Bor th P. humil shoot pprn on th groun Atr th P. humil shoot pprn on th groun

6 Min Journl o Eology n Environmnt (17) 41:3 Pg 6 o 1 wight Dry *** Mrh 16 April 6 root rhizom stm totl Fig. 5 Dry wight o h orgn or (Mrh 16) n tr (April 6) th shoot sprouting. Br inits stnr vition. *** Signiint t.1% lvl in irn twn ry wights o rhizom on Mrh 16 n April 6 Chngs in our proprtis long th rnk on stm n th numr o lvs Th plnts us or th rnk on stm nlysis wr rom plnts with 4 lvs (4-L plnt) to 11 lvs (11-L plnt). O ths plnts, I onsir 11-L plnt s th most rlil initor to show th stnr hrtr (Fig. 8). For 11-L, two typs o hngs in our proprtis long th rnk on stm rom L 1 (th irst on stm) to L 11 (th 11th on stm) wr osrv, on prouing rgulr urv n th othr on n irrgulr urv. In th ormr s, th vlus irst inrs n thn rs, with th mximum t L (th son on stm) or rltiv r, L 4 (th ourth on stm) or rltiv ry wight n L 4 n L 5 (th ith on stm) or SLA. Th urv show th positiv skwnss. O th 55 pirs o th rltiv r, th irns twn th lvs on stm wr signiint t.1% in 39 n t 1 ~ 5% lvl in 4. For rltiv ry wight, th irns wr signiint t th.1% lvl in pirs n t 1 5% lvl in 18 pirs. Howvr, or SLA th irns lmost rh signiin t 5% lvl in 5 pirs. Th lttr irrgulrly lutut n ws th lngth/with rtio. O th 55 pirs, th irns twn th Tl 4 Corrltion oiints twn th rhizom n shoot systm in Polygontum humil popultion. R 1 n R r th lst yr rhizom n this yr on, rsptivly. Numr is smpls (n). All orrltion oiints r signiintly irnt t n 1% lvl Rhizom Shoot systm Numr Lngth o stm Wight o stm Totl wight o shoot Lngth o R Wight o R Lngth o R Wight o R * lvs on stm wr signiint only t th 5% in our pirs. In plnts with lvs 1, two typs o hngs in proprtis long th rnk on stm wr osrv. For th lngth/with rtio, th vlus long th rnk on shoot show norml istriution urv in plnts with 3 8 lvs n lutut in plnts with 9 or 1 lvs. Howvr, th irns twn th lvs on stm wr not signiint or most proprtis. For th othr thr proprtis, th lous o showing th mximum vlu mov to th L 1 irtion with th rs in th totl numr o lvs stm. In othr wors, in plnts with mor lvs, th lvs with th mximum vlus wr lot urthr rom L 1 thn in plnts with wr lvs. Morovr, with th inrs in th numr o lvs pr plnt, th irns twn th lvs on stm inrs n thir signiin t.1% lvl inrs. For xmpl, or th rltiv r o 5-L plnt, th numr o rltionships showing signiint irn t.1% lvl ws 1 o 1 pirs; howvr, it ws 1 o 8 pirs in 7-L plnt n 6 o 38 pirs in 9-L plnt. Howvr, th numrs o pir in whih th irn ws signiint t.1% lvl wr issimilr to h othr mong th thr proprtis or th sm plnts. In th plnts with smll numr o lvs (4 to 7 lvs), proprtis hng long th rnk on stm; howvr, th pttrns o ths hngs wr irrgulr n most o th irn ws not signiint. Disussion In northrn tmprt iuous orst, unrstory prnnil hrs riv littl sunlight us thy r sh y th nopy (Emrog 1998, Whinghm 4). Thus, ths plnts shoul rspon n limt to lowlight nvironmnt. In this nvironmnt, gp provis goo rug or ths plnts (Km t l. 13). In this stuy, P. humil popultion show morphologil ivrsity or sunlight. First, P. humil s rhizom grw up to 5.5 m in lngth. This is in ontrst to th t o Choung (1991), who rport tht th mximl growth o P. humil s rhizom ws 9. m unr nrly ull sunlight. Thus, th growth o P. humil rhizom ws longr t this stuy sit thn in othr stuy. This strtgy might th mns to voi sh n is in lonl plnts (Go t l. 1). Morovr, th numr o P. humil s rhizoms gnrt on shoot ws 1 3 in most plnts, rthr thn 4 y Jng (). In gnrl, orging o rhizom is osrv in nvironmnt with nutrint rsour htrognity (Dong n Altn 1999). Son, th shoots o P. humil sprout in spring using root systm nrgy. Thus, root systm iomss rs in orn with shoot sprouting. In this stuy, root systm ry wight rs to 44% (375.6 ± mg plnt 1 on April 6, 15 n 85.6 ± 6.5 mg plnt 1 on Mrh 16, 15) tr th

7 Min Journl o Eology n Environmnt (17) 41:3 Pg 7 o 1 ) (m r L 1 r =.395 o wight 5 r = Lngth o R 1 (mm) Dry Lngth o R 1 (mm) L r (m ) r = Dry wight o R 1 5 o wight Dry r = Dry wight o R 1 5 L nght o R (mm ) r =.5639 L nght o R (mm ) r = L r (m ) Dry wight o D ry wight o R (mg ) r = L r (m ) D ry wight o R (mg ) r = Dry wight o Fig. 6 Rltionship twn th rhizom proprty n th shoot on. R n R 1 init th rhizom o this n lst yr, rsptivly shoot sprout. Morovr, rhizom ry wight rs to 37% (rom ± 11.1 to 84.6 ± 88.5 mg plnt 1 ) tr shoot sprouting. Th totl ry wight o plnt (root systm + shoot systm) ws smllr thn or shoot sprouting. This rsult inits onsirl nrgy xpnitur or rspirtion uring shoot sprouting. It my impossil or th sm rhizom to prou shoots twi us o insuiint nrgy. No shoots wr oun on R (rhizom tht vlop yrs go) or R 3 (rhizom tht vlop 3 yrs go) whih tth to R 1 (Hsgw n Kuo 5). Unr optiml onitions, ospring proution os not pn on th strting pitl in th hirnl; ut unr growth-limiting onitions, thik hirnl my mor sussul in Hlinthus x ltiolius (Vrurg t l. 1996). Thir, th siz o h plnt orgn inrs with th numr o lvs pr plnt n th irn orrsponing to on- inrmnt ws signiint t 1 5% lvl. Thus, th numr o lvs ws goo initor o plnt siz. Although rhizom siz inrs with th numr o lvs, most irns wr not signiint; in prtiulr, plnts with smll numr o lvs (3 to 7 lvs) show no signiint irns, likly us smll plnts oul not ully riv sunlight lik th lrgr plnts. Th rson or this inrn ws tht smll plnts h low SLA vlus n, s rsult, grw mor slowly thn lrg plnts. As xpt, plnts with mny lvs wr tllr thn plnts with wr lvs. Thus, P. humil pprs to sns n to rspon to mirolimt. Fith, th CC vlus twn rhizom n shoot sizs wr signiint t 1% lvl. In gnrl, orgni mttr is trnsport rom R 1 to th stm n lvs, whrs

8 Min Journl o Eology n Environmnt (17) 41:3 Pg 8 o 1 ) (m plnt r pr Totl 3 1 i h g Numr o plnt ) r (m Mn 3 1 g Numr o plnt plnt pr wight Totl g Numr o plnt wight Mn Numr o plnt Spii r (m /g) Numr o plnt L ngth o R 1 (m ) Numr o plnt D ry wight o R 1 (mg ) Numr o plnt L ngth o R (m ) Numr o plnt D ry wight o R (mg ) Numr o plnt Lngth o stm (m) Numr o plnt g h stm 6 4 g h o Dry wight Numr o plnt Fig. 7 Proprtis h orgn long th numr o lvs. Br inits stnr vition. Lttrs rom to h init tht irns mong th plnts long th numr o r signiint t 1% lvl

9 Min Journl o Eology n Environmnt (17) 41:3 Pg 9 o Rt o lngth / with o plnt 7- plnt 9- plnt 11- plnt r Rltiv plnt 7- plnt 9- plnt 11- plnt ry wight Rltiv plnt 7- plnt 9- plnt 11- plnt /g) (m r Spii plnt 7- plnt 9- plnt 11- plnt Rt o lngth / with o plnt 6- plnt 8- plnt 1- plnt r Rltiv plnt 6- plnt 8- plnt 1- plnt ry wight Rltiv plnt 6- plnt 8- plnt 1- plnt /g) (m r Spii plnt 6- plnt 8- plnt 1- plnt Fig. 8 Chngs o proprtis long rnk on stm rom 4-L plnt to n 11-L plnt photosynthtts r trnsport rom lvs to R.Thus, rhizom n shoot r losly rlt. In gnrl, th r is us to stimt o photosynthti proutivity, possily us, in th il, th mount o sunlight riv y h ts ry wight n irt sunlight inus thikning n rus th SLA (Frrir t l. 13). As mntion ov, SLA vlus wr smllr in tll plnts thn in short plnts us th ormr riv mor sunlight thn th lttr. SLA is goo initor or ripint sunlight grint (Dwyr t l. 14) n wight might hng oring to umultiv sunlight xposur. Finlly, in plnts with th sm numr o lvs, th vlus o our proprtis irst inrs n thn rs long th rnk on stm. Howvr, th urvs lutut slightly in plnts with smll numr o lvs. This t inits tht th proprtis o lvs t th sm no r ivrs in smll plnts u to vrition in th mount o riv sunlight. In 11-L plnt, r, SLA, n ry wight inrs rom th L 1 to L or L 3 or L 4 n, thrtr, rs to L 11. Dirns o proprtis twn th lvs on stm wr signiint or most t.1 1% lvl. From th with th mximum vlu, th highr th position, th lss th SLA n th r vlus wr. Thus, hngs in ths proprtis oul n pttion to vritions in sunlight. I liv tht P. humil popultion xmin h goo strtgy or pttion y voiing th sh (Hsgw n Kuo 5). Wooy plnt lvs in

10 Min Journl o Eology n Environmnt (17) 41:3 Pg 1 o 1 tmprt iuous orst show htromorphism long th rnk on nw twig, lthough th tim n or th growth rom th L 1 to th lst is short n h might grow unr th sm sunlight onitions (Min 1995). P. humil lvs on stm lmost grw t th sm tim, so mironvironmnt might similrly t ll lvs. Thus, plnt might show ivrs morphology with rnk on stm or twig unr similr mironvironmnts. Conlusions Rltionships twn th root n th shoot s systms Root systm s iomss rs in orn with shoot sprouting n this rsult rom onsirl nrgy xpnitur or rspirtion uring shoot sprouting. Morovr, th CC vlus twn rhizom n shoot sizs wr signiint t 1% lvl. Thus, rhizom n shoot r losly rlt. Th growth o P. humil rhizom ws longr t this stuy sit thn in othrs. This strtgy o lonl plnts might th mns to voi sh in orst gp. Rltionships twn th numr n th proprtis o lvs Th siz o h plnt orgn inrs with th numr o lvs pr plnt, n th irn orrsponing to on- inrmnt ws signiint t 1 5% lvl. Thus, th numr o lvs ws goo initor o th plnt siz. Although rhizom siz inrs with th numr o lvs, most irns wr not signiint, likly us smll plnts oul not ully riv sunlight lik th lrgr plnts. Th rson or this inrn ws tht smll plnts h low SLA vlus n, s rsult, grw mor slowly thn lrg plnts. Chngs in our proprtis long th rnk on stm n th numr o lvs For 11- plnts, two typs o hngs in our proprtis long th rnk on stm rom L 1 to L 11 wr osrv, on prouing rgulr urv n th othr on n irrgulr urv. In th ormr s, th vlus irst inrs n thn rs long th rnk. Thr wr rltiv r, rltiv ry wight, n SLA in ths proprtis. In th lttr s, th vlu irrgulrly lutut long th rnk n ws th lngth/with rtio. Thus, hngs in th sizs show rgulr trn long th rnk on stm. Howvr, hng o morphology long th rnk on stm ws irrgulr. Arvitions (3 ~ 1)-L: (3 ~ 1)- plnt;.s.l.: Aov s lvl; CC: Corrltion oiint; L (1~1) :L s orinl numr on stm; R : Nw (this yr) rhizom;; R 1 :Thlst yr s rhizom;r : yrs go rhizom; R 3 : 3 yrs go rhizom; SLA: Spii r Aknowlgmnts Not pplil. Funing Not pplil. Avilility o t n mtrils Th tsts nlyz uring th urrnt stuy r irtly vill rom th ils ( nturl popultion o P. humil), s mntion in Mthos stion. Ethis pprovl n onsnt to prtiipt Not pplil. Consnt or pulition Not pplil. Compting intrsts Th uthor lrs tht h hs no ompting intrsts. Pulishr s Not Springr Ntur rmins nutrl with rgr to jurisitionl lims in pulish mps n institutionl ilitions. Riv: Frury 17 Apt: August 17 Rrns Choung, Y.-S. (1991). Growth hrtristis n mogrphy o Polygontum involurtum n Polygontum humil rmt popultion. Korn J Eol, 14, Cook, R. E. (1983). Clonl plnt popultion. Amrin Sintist, 71, Dong, M., & Altn, B. (1999). Clonl plstiity in rspons to rhizom svring n htrognous rsour supply in th rhizomtous grss Psmmohlo villos in n innr Mongolin un, Chin. Plnt Eology, 141, Dwyr, J. M., Hos, R. J., & Myil, M. M. (14). Spii r rsponss to nvironmntl grints through sp n tim. Eology, 95, Emorg, J. (1998). Unrstory light onitions n rgnrtion with rspt to th struturl ynmis o nr-nturl tmprtur iuous orst in Dnmrk. Forst Eology n Mngmnt, 16, Frrir, G. N., Pro Jr., J. A., Shivini, I., & Mlo, C. (13). Plsti rsponss in tr rhittur n spii r o Xylopi romti Annon: Apttions to nvironmnts with irnt light intnsitis. Brz J Bot, 36, Go, Y., Xing, F., Jin, Y., Ni, D., & Wng, Y. (1). Forging rsponss o lonl plnts to multi-pth nvironmntl htrognity: Sptil prrn n tmporl rvrsiility. Plnt n Soil, 359, Hsgw, T., & Kuo, G. (5). Comprisons o growth shul, rproutiv proprty n llotion pttrn mong thr rhizotomous Polygontum spis with rrn to thir hitt typs. Plnt Spis Biology,, 3 3. Ishiok, R., Mullr, O., Hiur, T., & Kuo, G. (13). Rsponss o ing phnology n photosynthsis to soil wrming in orst-loor plnts. At Oologi, 51, Jng, C.-C. (). A txonomi rviw o Korn Polygontum (Rus). Korn Journl o Plnt Txonomy, 3, Jng, K. H., Prk, J. M., Kng, J. H., & L, S. T. (1998). Growth n lowring hrtristis o Polygontum spp. J Miinl Crop Si, 6, Krn, C. C., Montgomry, R. A., Rih, P. B., & Strong, T. F. (13). Cnopy gp siz inluns nih portioning o th groun-lyr plnt ommunity in northrn tmprt orst. Journl o Plnt Eology, 6, Klinkhmr, P. G. L., Jong, T. M., & Mlis, E. (199). How to tst or proportionlity in th rproutiv ort o plnts. Amr Ntl, 135, L, T. B. (3). Colour lor o Kor (p. 676). Soul: Hyngmoon. Lgnr, N., Flk, S., & Lushnr, C. (13). Low light limtion in iv tmprt ro-lv tr spis o irnt sussionl sttus: Th signiin o sh nopy. Annls o Forst Sin, 7, Min, B. M. (1995). Growth n htromorphism o lvs long th rnk (plstohron) o wooy plnts. J Plnt Biol, 38, So, Y.-S., Prk, W.-H., & Ch, Y.-Y. (11). Et o Polygontum oortum on lowring lipi n ntioxition. J Orintl Rh M, 1, Vrurg, R. W., Kwnt, R., & Wrgr, M. J. A. (1996). Th t o plnt siz on vgttiv rproution in psuo-nnul. Vgttio, 15, Whighm, D. F. (4). Eology o wooln hrs in tmprt iuous orsts. Ann Rv Eol Syst, 35,