Hisae Nagashima* and Kouki Hikosaka

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1 Annls of Botny 108: , 2011 doi: /o/mcr109, ville online t Plnts in crowded stnd regulte their height growth so s to mintin similr heights to neighours even when they hve potentil dvntges in height growth Hise Ngshim* nd Kouki Hikosk Grdute School of Life Sciences, Tohoku University, Ao, Sendi , Jpn * For correspondence. E-mil hise@ceres.ocn.ne.jp Received: 5 Jnury 2011 Returned for revision: 15 Ferury 2011 Accepted: 23 Mrch 2011 Pulished electroniclly: 11 My 2011 Bckground nd Aims Although eing tll is dvntgeous in light competition, plnt height growth is often similr mong dominnt plnts in crowded stnds (height convergence). Previous theoreticl studies hve suggested tht plnts should not overtop neighours ecuse greter lloction to supporting tissues is necessry in tller plnts, which in turn lowers lef mss frction nd thus cron gin. However, this model ssumes tht competitor hs the sme potentil of height growth s their neighours, which does not necessrily ccount for the fct tht height convergence occurs even mong individuls with vrious iomss. Methods Stnds of individully potted plnts of Chenopodium lum were estlished, where trget plnts were lifted to overtop neighours or lowered to e overtopped. Lifted plnts were expected to keep overtopping ecuse they intercept more light without incresed lloction to stems, or to regulte their height to similr levels of neighours, sving iomss lloction to the supporting orgn. Lowered plnts were expected to e suppressed due to the low light vilility or to increse height growth so s to hve similr height to the neighours. Key Results Lifted plnts reduced height growth in spite of the fct tht they received higher irrdince thn others. Lowered plnts, on the other hnd, incresed the rte of stem elongtion despite the reduced irrdince. Consequently, lifted nd lowered plnts converged to the sme height. In contrst to the expecttion, lifted plnts did not increse lloction to lef mss despite the decresed stem length. Rther, they llocted more iomss to roots, which might contriute to improvement of mechnicl stility or wter sttus. It is suggested tht decresed lef mss frction is not the sole cost of overtopping neighours. Wind lowing, which my enhnce trnspirtion nd drg force, might constrin growth of overtopping plnts. Conclusions The results show tht plnts in crowded stnds regulte their height growth to mintin similr height to neighours even when they hve potentil dvntges in height growth. This might contriute to voidnce of stresses cused y wind lowing. Key words: Height growth, stem elongtion, plsticity, light competition, neighour effect, stem dimeter growth, iomss prtitioning, Chenopodium lum. INTRODUCTION Light competition is criticl for survivl, growth nd reproduction of individuls in dense stnds. Plnts tht could not receive sufficient light stop their growth nd often die (Weiner et l., 1990; Ngshim et l., 1995; Mtsumoto et l., 2008). As light is unidirectionl resource, tller plnts intercept more irrdince nd overshde shorter competitors (Schmitt et l., 1995; Dudley nd Schmit, 1996; Huer nd Wiggermnn, 1997; Anten nd Hirose, 1998; Huer et l., 1998). It is well known tht mny herceous plnts enhnce elongtion of stems when the stnd density or the lef re index (LAI) is high (e.g. Bllré et l., 1990; Weiner et l., 1990; Weiner nd Thoms, 1992; Weiner nd Fishermn, 1994; Ngshim, 1999; Nishimur et l., 2010). This response hs een regrded s shde voidnce (Smith, 1982). Height growth is, however, not lwys competitive ut often pprently co-cpertive. In mny plnt communities, height is similr mong plnts tht reched the top of the cnopy (e.g. Koym nd Kir, 1956; Kuroiw, 1960; Ford, 1975; Hutchings nd Brkhm, 1976; Weiner nd Thoms, 1992; Weiner nd Fishmn, 1994; Ngshim nd Tershim, 1995; Ngshim et l., 1995; Vermeulen et l., 2008), which is clled height convergence (Ngshim nd Tershim, 1995). Since tll plnts receive high light levels, one my consider tht eing tll is dvntgeous in light competition. However, plnts do not necessrily keep this dvntge. Height convergence is often oserved mong upper plnts with different iomss, even though plnts with lrger iomss my hve higher potentil for growth (Weiner nd Fishmn, 1994; Ngshim nd Tershim, 1995). Why do such plnts not keep overtopping others? Although eing tll is dvntgeous in light competition, it entils costs for plnts. As lef height increses, plnts need to invest iomss more thn proportiontely in the stem to support their own weight, which in turn reduces the frction of lef mss in the plnt (Givnish, 1982; Nikls, 1992). Givnish (1982) proposed gme-theoretic model of plnt height growth. He ssumed tht overtopping the other plnts # The Author Pulished y Oxford University Press on ehlf of the Annls of Botny Compny. All rights reserved. For Permissions, plese emil: journls.permissions@oup.com

2 208 Ngshim & Hikosk Height growth regultion in crowded stnd is dvntgeous in light cpture nd thus leds to higher production rte per unit lef mss, ut the lef mss frction in shoot decreses with incresing plnt height. His model predicted existence of the evolutionrily stle height (ESH) in stnd t given density: no other height (neither tller nor shorter) is dvntgeous in iomss production in the stnd in which plnts hve the ESH. His model lso predicted tht the ESH increses with incresing cnopy cover. This ws well supported y forest fors (Givnish, 1982). Enhncement of height growth in response to crowding seems to e consistent with the prediction. Height convergence my e result of evolutionry stle growth of competing plnts. Although the model of Givnish (1982) cn provide importnt insights into plnt growth under competition, it is still questionle whether his model fully ccounts for the height convergence oserved in plnt communities. First, his model ssumed tht competing plnts hve equl iomss to ech other, which is not the sme s field situtions where plnts with different iomss hve similr height (Weiner nd Fishmn, 1994; Ngshim nd Tershim, 1995). Secondly, it is unknown whether the constrints ssumed in the model re pproprite to explin the plstic height growth of plnts found in open hitt. The model ssumed tht plnts hve constnt mechnicl stility nd ccordingly the iomss lloction to leves depends only on plnt height. There my e, however, nother disdvntge in eing tll. When plnts overtop the neighouring plnts, they my e exposed to stronger wind thn the neighours, which might entil negtive effects on plnt growth due to excessive trnspirtion nd mechnicl stress (Drke et l., 1970; Grce, 1974; Putz et l., 1983). If this is the cse, eing tll my e more disdvntgeous thn tht considered in Givnish (1982). In the present study, n experiment ws designed to exmine whether plnts void overtopping nd whether the ehviour is ccording to the Givnish model. A stnd of potted plnts ws mde nd the height of the pot (ove or lower thn neighouring pots) ws mnipulted. The lifting tretment will enefit plnts in light cpture without ny reduction in lef mss frction. This is in contrst to the sitution of overtopping plnts in the Givnish model, where overtopping plnts increse light interception ut reduce iomss lloction to leves. The lowering tretment will decrese light vilility of the plnt despite the sme lef mss frction s the neighours. We postulted two hypotheses for response to the tretments. (1) Lifted plnts will keep themselves overtopping their neighours ecuse there is no disdvntge in overtopping. Lowered plnts my e suppressed due to the low light vilility. (2) Treted plnts my regulte their height growth so tht their height ecomes similr to the neighours. Lifted plnts will decrese the height growth nd consequently e cught up y the neighours. Lowered plnts my increse height growth so s to hve similr height to the neighours. Biomss lloction chnges cused y the tretments were lso investigted. First it is expected, s hs een ssumed y Givnish (1982), tht iomss lloction to leves depends on stem length irrespective of tretments. If reltionship etween lloction to leves nd stem length is ffected y the tretment, constrints other thn tht ssumed in Givnish (1982) re suggested in the competing plnts. The experimentl results were lso compred with the theoreticl prediction using the model of Givnish (1982). MATERIALS AND METHODS The experiment ws conducted with Chenopodium lum, rod-leved summer nnul, in n experimentl grden of Tohoku University, Sendi, Jpn (38825 N, E). Monthly men ir tempertures during the experiment were 18.0, 22.5 nd C in June, July nd August, respectively. Seeds of C. lum were otined from plnts in nturl popultion in Three hundred nd twelve plstic pots, ech 12.5 cm in dimeter nd 20 cm high, filled with mixture of 7 : 3 vermiculite nd Akdm (grnulr lomy soil cm in dimeter), were tightly rrnged on ench 0.9m wide, 5.4 m long nd 0.5 m high nd plced outdoors. On 5 June 1996, out five seeds of C. lum were sown in ech pot. Pots were wtered t h nd h every dy y n utomtic wtering system. After emergence, ech pot ws fertilized with 100 ml of nutrient solution of 0.01 : 0.02 : 0.01 NPK every week (10 mg N, 20 mg P nd 10 mg K week 21 per pot) from 26 June until 24 July, nd from 31 July with 100 ml of nutrient solution of 0.02 : 0.04 :0.02 NPK (20 mg N, 40 mg P nd 20 mg K week 21 per pot). Seedlings were thinned to leve one per pot y 16 July: the plnt density ws 77 plnts m 22. On 2 August, the stnd ws surrounded y 80 % shde cloth to the height of the top of the plnts to reduce edge effects. The height of the shde cloth ws chnged ccording to the growth of the plnts. On 3 August, the length nd sl dimeter of the stems of ll plnts were mesured. The stem length ws mesured from the se to the terminl shoot pex to the nerest 1 mm, nd the sl dimeter ws mesured to the nerest 0.1 mm in the middle of the first internode. Cre ws tken to llot plnts of similr size for 21 trget nd 126 neighouring plnts. Pots were rrnged to hve trget plnt surrounded y six neighouring plnts s shown in Fig. 1B. The height differences etween the trget nd neighouring plnts were creted on 4 August, 60 d fter sowing, when the men plnt height ws 37 cm nd LAI ws 2.3. Seven trgetplnt pots were lifted y 7 cm (lifted plnts), nother six trget-plnt pots were lowered y 7 cm (lowered plnts), nd the other six trget-plnt pots were kept t the sme level s the neighouring pot (control; Fig. 1A). The height nd dimeter of trget plnts were mesured in the sme wy s ove. The height of six neighouring plnts ws lso mesured for ech trget plnt. Photosyntheticlly ctive rdition (PAR) ws mesured with quntum sensors (LI-190SA, LI-COR Inc., Nersk, Lincoln, USA) t the top of the plnt nd outside the stnd. For ech plnt, PAR ws verged from three mesurements. One nd two weeks fter the tretment, the height nd dimeter of trget plnts nd the height of the neighouring plnts were mesured. In the experimentl period two neighours were replced with mrginl plnts to mke the neighourhood condition of trget plnts even. After mesurements, trget plnts were hrvested nd seprted into orgns (lef, stem nd root), nd dried t 80 8C for 3 d nd weighed. LAI incresed to 2.9 nd 4.5 in 1 nd 2 weeks fter tretments, respectively.

3 Ngshim & Hikosk Height growth regultion in crowded stnd 209 A Pot-height tretment where P mx nd P min re the photosynthetic rte per unit lef mss of unshded nd shded leves, respectively; o is the proility of horizontl overlp of leves; leves re distriuted on stems rndomly within the rnge of h i (1 + s). When n invder is shorter thn the neighours (h 1, h 2 ), { } [ h 1 (1 + s) h 2 (1 s) ] 2 g(h 1, h 2 )=P min + 4h 1 h 2 s ( 1 o ) (P mx P min ) (2) 2 For n explntion of how the equtions re derived see Givnish (1982), ut note tht eqn 2 is in different form from the eqution given in tht pper. Lef mss frction, f(s), is given s function of stem length (s), where nd re constnt. Control (C) Lowered (Lo) Lifted (Li) B Pot rrngement f (s) = s (3) Effects of the tretment on plnt growth were nlysed y the generlized liner model with norml error distriution nd identity link (JMP sttisticl softwre; SAS Institute Inc., Cry, NC, USA). Tukey Krmer honestly significnt difference test ws used for post-hoc pirwise comprisons. Simultion C Lo Li FIG. 1. Experimentl design: tretment of height difference (A) nd rrngement of pots (B). Trget plnts were lifted, lowered, or remined the sme level s surrounding plnts, s indicted. Optiml lef height of lifted or lowered plnts ws simulted ccording to the model of Givnish (1982). The model ssumed single-lef plnts ut the model cn e extended to multi-lef plnts like C. lum. Here photosynthetic production of n invder plnt tht competes with neighours for light is considered. Plnt photosynthetic production is otined y photosynthetic production per unit lef mss (g) multiplied y lloction of shoot mss to leves [ f(s)], where s is stem length, g is function of the men height of the leves of trget (h 1 ) nd neighouring (h 2 ) plnts. When n invder is tller thn the neighours (h 1. h 2 ) { g(h 1, h 2 )=P mx [h 2(1 + s) h 1 (1 s)] 2 } 4h 1 h 2 s 2 o 2 (1) (P mx P min ) C Lo Li C In simultions, nd mmol CO 2 g 21 lef dry mss s 21 were dopted for P mx nd P min, respectively, which were clculted from the top (1.68 g N m 22 ) nd ottom leves (0.44 g N m 22 ) of dominnt individuls in monospecific stnd of Chenopodium lum estlished in the sme experimentl grden in 1999 (for detils, see Hikosk et l., 2003). Vlues of nd were 0.73 nd , respectively, which were otined from control plnts in the present study (r 2 ¼ 0.97), s is 0.48 from the verge of control plnts, nd o ws regrded s one s the LAI exceeded one. Simultions were conducted in the cse of n invder 7 cm higher or lower thn neighours (simultion A); nd n invder whose pot is lifted or lowered y 7 cm (simultion B). In simultion A, height (h) nd stem length (s) of n invder re identicl vlues, while in simultion B, h nd s re different vlues from ech other. The evolutionrily stle height (h*), which is defined s the height of stnd where no invder with ny other height hs higher photosynthetic productivity (g f ), is given s follows (Givnish, 1982): h = (/) o [1 (P min /P mx )] 2s +(1 s) o [1 (P min /P mx )] In simultion B, n optiml height mximizing their photosynthetic production ws numericlly found out for lifted or lowered invders. TABLE 1. Photosyntheticlly ctive rdition (PAR) t the top of the plnt reltive to outside of stnd just fter the height difference tretment (for control, lifted nd lowered plnts, see Fig. 1) (4) Control Lifted Lowered Reltive PAR t the top (%) Different superscript letters indicte significnt difference (P, 0.05) ssessed y Tukey Krmer test (n ¼ 7 in ech group).

4 210 Ngshim & Hikosk Height growth regultion in crowded stnd RESULTS The PAR t the top of the plnts ws significntly lower in lowered plnts, though the difference ws smll (P ¼ 0.03, ANOVA). No significnt difference ws found etween lifted nd control plnts (Tle 1). In lifted plnts, however, out nine expnded leves were positioned higher thn the cnopy surfce, while in control plnts only three expnded leves were exposed to the top of the cnopy. Thus light interception ws incresed y the lifting tretment. Control plnts exhiited the height growth rte of 23 cm per week during the experiment. Lifted plnts slowed stem elongtion (Tle 2 nd Fig. 2A). Two weeks fter the onset of the tretment, the difference in pprent height from neighouring plnts ecme 1 cm (not significntly different from zero) in lifted plnts (Fig. 2B). Lowered plnts ccelerted stem elongtion (Tle 2 nd Fig. 2A), nd the difference in pprent height from neighours ws reduced from 7 to 3.5 cm during the experiment (Fig. 2B). However, their height ws still lower thn tht of their neighours (Fig. 2B). Totl iomss ws significntly different mong tretments (Tle 2). However, the iomss of lifted plnts ws not greter thn tht of control plnts (Tle 2), even though lifted plnts might hve received greter irrdince. Lowered plnts hd smller iomss thn the other two (Tle 2). There ws no significnt effect of the tretment on lef mss nd lef mss frction (lef/totl mss) despite the differing stem lengths mong the tretments (Tle 2). The tretment significntly ffected the root mss nd root mss frction (Tle 2). Stem mss nd the stem mss frction showed mrginlly significnt difference (Tle 2). These results suggest tht there were chnges in lloction etween roots nd stems cross the tretments: lifted nd lowered plnts tended to invest more iomss in roots nd stems, respectively. Specific lef re (SLA) incresed more in lowered plnts thn in the others, resulting in greter lef re rtio (LAR) in lowered plnts (Tle 2). Lifted plnts slowed stem elongtion ut incresed stem thickness enlrgement, while lowered plnts ccelerted elongtion ut tended to reduce dimeter growth (Tle 2). Tle 3 summrizes the simultion results. Here it is ssumed tht neighours hve n evolutionrily stle height (h*) (eqn 4). When stnd is composed of pots t the sme height level (simultion A; no lifting or lowering), plnt tller or shorter thn h* hs lower photosynthetic production thn the neighours with h*. This is ecuse, for exmple, in higher plnt the positive effect y the increse in g ws smller thn the negtive effect y the decrese in f. In simultion B, on the other hnd, lifted or lowered plnt hs the sme stem length s neighours, ut is 7 cm tller or shorter, respectively, thn the neighours. A lifted plnt hs greter photosynthetic production thn the neighours ecuse they increse g without decrese in f. If the lifted plnts decrese their stem length nd increse f, photosynthetic production further increses. Photosynthetic production of the lifted plnts is mximized when they re 2 cm higher thn the neighours, suggesting tht keeping 2 cm tller thn neighours is dvntgeous for lifted plnt. A lowered plnt in simultion B produces less photosyntheticlly thn the neighours irrespective of stem length. Photosynthetic production of the lower plnt is mximized when the height is 4 cm lower thn the neighours. DISCUSSION The present study showed tht plnts in crowded stnds regulte their height growth so s to mintin similr height to neighours: plnts whose top hd een lowered reltive to their neighours ccelerted the elongtion rte while plnts whose top hd een lifted higher thn the neighours reduced the elongtion rte (Fig. 2). Lifted plnts did not keep themselves overtopping neighours, even though they hd two dvntges t the onset of TABLE 2. Stem dimensions, plnt dry mss, dry mss prtitioning, lef re, growth prmeters of trget plnts 2 weeks fter tretments (see Fig. 1) Control Lifted Lowered P-vlue Stem length (m) c < Height increment (m) c < Dimeter (mm) Dimeter increment (mm) Length/dimeter (m m 21 ) c < Totl mss (g) Root mss (g) Stem mss (g) Lef mss (g) Root/totl mss (g g 21 ) Stem/totl mss (g g 21 ) Lef/totl mss (g g 21 ) Lmin re (cm 2 ) SLA (m 2 kg 21 ) < LAR (m 2 kg 21 ) < Vlues re mens + s.e. Different superscript letters indicte significnt differences (P, 0.05) mong tretments ccording to Tukey Krmer test (n ¼ 7 in ech group). P-vlues re the result of generlized liner model nlyses (d.f. ¼ 2). Significnt vlues (P, 0.05) re highlighted in old. SLA, specific lef re (lmin re/lmin mss); LAR, lef re rtio (lmin re/totl mss).

5 Ngshim & Hikosk Height growth regultion in crowded stnd 211 Apprent height difference from neighouring plnts (m) Stem length (m) A B c experiment: incresed light interception nd less cost of tissues to support leves t higher positions. At the end of the experiment, the height difference etween lifted nd the neighour plnts ecme 1 cm (Tle 2). The vlue ws similr to tht expected from simultion B (Tle 3). However, growth ws quite different from tht expected from the Givnish model. First, frction of leves in the plnt mss ws not different mong the tretments despite the different stem lengths (Tle 2). This result suggests tht the cost of competition my not e fully ccounted for y reduction in lef mss frction with n increse of plnt height (stem mss frction). Secondly, lifted plnts did not increse growth, even though they hd received more light thn their neighours (Tle 2). This suggests tht overtopping is not dvntgeous for photosynthetic rte per lef mss. Some other costs or constrints therefore need to e incorported for understnding height growth regultion in crowded stnds. Wht ws the cost or constrint of overtopping, then? Constrint of eing tll hs een considered minly sed on sfety ginst uckling: stem ends due to its own nd lef c Weeks fter tretment Control Lifted Lowered FIG. 2. Growth of stem length of trget plnts (A) nd difference in height of the top of plnts etween trget nd neighouring plnts tht ws otined s [height of trget plnt (+7 cm, for lifted plnts; 7 cm, for lowered plnts)] (men height of six djcent plnts) (B), fter pot-elevtion tretment (see Fig. 1). Different letters indicte significnt differences (P, 0.05) mong tretments ccording to Tukey Krmer test (n ¼ 7 in ech group). Brs represent + s.e. c c Height of invder plnt (cm) TABLE 3. Simultion results Photosynthetic production (mmol CO 2 g 21 d 21 ) Simultion A Unchnged pot height h* h* h* Simultion B 7 cm lifted h* h* h* + 2 (optiml) cm lowered h* h* h* 4 (optiml) Photosynthetic production per ove-ground iomss is clculted s the product of photosynthetic production per lef mss (g) nd lef mss frction ( f ). In every simultion, height of neighouring plnts is n evolutionrily stle one (h* ¼ 153 cm). In simultion A, the height of n invder plnt is ltered from the neighours with n identicl pot height to the neighours. h* + 7 mens tht the height of n invder is 7 cm higher/lower thn h*. In simultion B, the pot height of n invder plnt is lifted up or lowered y 7 cm compred with neighours. h* + 7 in 7 cm lifted in simultion B mens tht the height of n invder is 7 cm higher thn h* due to pot elevtion ut the stem length is the sme s the neighours. h* + 2 mens tht height nd stem length re 2 cm higher nd 5 cm lower thn the neighours nd the invder tht hs this height hs higher photosynthetic production thn those t other heights. Similrly, 7 cm lowered mens tht the pot height of n invder is 7 cm lower thn the neighours. weight (e.g. McMhon, 1973; King, 1986; Nikls, 1993,, 1994, ; Csl et l., 1994; King et l., 2009). However, uckling sfety does not necessrily gurntee sfety ginst reking dmge cused y wind lowing. Becuse wind speeds increse drmticlly ove the oundry lyer of the cnopy (Goudrin, 1977; Speck, 2003), overtopped plnts my e exposed to stronger winds thn the neighours. As wind speed increses, the risk of mechnicl filure increses (Nikls, 2000). During the present experiment, the mximum instntneous wind speed in the dy ws m s 21 in Sendi (Jpn Meteorologicl Agency). In potted plnt of n erect nnul, Xnthium cndense, wind speed of 20 m s 21 ent the stem y (H. Ngshim, Tohoku University, Sendi, Jpn, unpul. res.). Thus lifted plnts in the present study might hve suffered from wind lowing nd chnged iomss lloction nd morphology to void mechnicl filure. In fct, lifted plnts reduced the height : dimeter rtio of the stem nd incresed the root mss frction (Tle 2), oth of which might hve contriuted to mechnicl stility (Nikls, 1992; Henry nd Thoms, 2002; Anten et l., 2005, 2006, 2009). Another constrint my e trnspirtion. It is notle tht lifted plnts did not hve greter iomss thn control plnts (Tle 2), though the intercepted light might hve een greter. This suggests tht light use efficiency in photosynthesis (cron gin per unit sored light; Hikosk et l., 1999) ws lower in lifted plnts. Grce (1974) showed tht n increse in wind speed from 1 to 3.5ms 21 enhnced trnspirtion nd reduced wter content of leves in Festuc

6 212 Ngshim & Hikosk Height growth regultion in crowded stnd rudince. It is prole tht wind induced stomtl closure to prevent wter loss (Drke et l., 1970), leding to reduction in photosynthesis rtes. Lifted plnts incresed the root mss frction (Tle 2), which is lso in ccord with generl responses to wter stress (Wtts et l., 1981). Lowered plnts, on the other hnd, ccelerted height growth, nerly reching the cnopy top (Fig. 2). The height difference ws 3.5 cm, which ws close to the prediction y the Givnish model (Tle 3). Enhnced height growth ws relized t the expense of stem dimeter growth, leding to higher height : dimeter rtio of the stems (Tle 2). This chnge in stem morphology is in ccord with the previous studies in which suordinte plnts were more slender thn dominnt nd solitry plnts (Weiner et l., 1990; Weiner nd Thoms, 1992; Weiner nd Fishmn, 1994; Ngshim nd Tershim, 1995). Biomss lloction ws lso chnged (Tle 2); lowered plnts tended to increse stem mss t the expense of root s compred with the other plnts (Tle 2). This lso grees with previous oservtions tht plnts llocte fewer resources to roots when they re shded or in dense stnd compred with plnts tht re exposed or solitry (e.g. Corré, 1983, ; Mlikl et l., 1999; Nishimur et l., 2010). These results imply tht ccelerting height growth is costly: reduced dimeter growth my increse risk of mechnicl filure of plnts nd reduced root mss frction my e disdvntgeous for cquisition of elow-ground resources s well s lower mechnicl stility. However, the elongtion my enefit the plnts y cron gin, since wind speed is lower thn it is on the outside of the cnopy (Goudrin, 1977; Speck, 2003). In the model of Givnish (1982), totl iomss of plnts ws ssumed to e identicl mong individuls for simplifiction. The model showed tht, if stnd consists of individuls with evolutionrily stle height (ESH), plnts t nother height cnnot hve higher iomss production rtes. However, this does not necessrily hold when the stnd consists of individuls with different totl iomss. If n individul hs greter iomss thn neighours, it cn hve higher production rte even though its height is greter or shorter thn the ESH of the neighours (Tle 3). In other words, the Givnish model my not explin why height convergence is often oserved even mong upper plnts with different iomss (Weiner nd Fishmn, 1994; Ngshim nd Tershim, 1995). The present results clerly suggest tht overtopping others is not necessrily dvntgeous even when the individul hs greter potentil for height growth. Plnts keep their height similr to their neighours to void constrints tht rise y overtopping others. Convergence of height growth leds to J-shped or imodl frequency distriution of plnt height (J-shped: the popultion consists of mny similrly tll plnts with smller numer of shorter plnts) (Koym nd Kir, 1956; Kuroiw, 1960; Ford, 1975; Hutchings nd Brkhm, 1976; Weiner nd Fishmn, 1994; Ngshim nd Tershim, 1995; Ngshim et l., 1995). Height convergence lso ccounts for non-liner reltionship etween log-trnsformed height nd dimeter, which hs commonly een oserved in vrious plnt communities; height is positively correlted with dimeter mong suordinte short plnts, while height is reltively similr in dominnt tll plnts irrespective of dimeter (Ogw et l., 1965; Assmnn, 1970; Kir, 1978; Ri, 1979; Kohym et l., 1990; Nikls, 1995; Ai nd Kohym, 1996; Thoms, 1996; Sterck nd Bongers, 1998; Weiner nd Thoms, 1992; Weiner nd Fishmn, 1994; Ngshim nd Tershim, 1995). Suordinte individuls tend to increse their height, while dominnt plnts suppress their height growth to keep their height similr to their neighours even though they hve greter potentil of height growth. Such n pprently co-cpertive growth of height my mke plnt plnt interctions less competitive, which llows coexistence of dominnt plnts. Ngshim et l. (1995) erlier showed in C. lum stnds tht the numer of upper plnts tht rech the cnopy top ws similr irrespective of initil density, which my prtly result from co-cpertive growth of the upper plnts. Severl environmentl cues hve een suggested for the regultion of height growth. In prticulr, the red : fr-red (R/FR) rtio in incident light ffects stem elongtion (for review, see Bllré, 1999, 2009; Smith, 2000). Height convergence ws impeded y tretments to reduce the R/FR effect (Bllré et l. 1994; Aphlo nd Rikl 2006). On the other hnd, mechnicl stimuli such s swying due to wind my lso ffect stem growth (Biro et l., 1980; Telewski, 1990; Jffe nd Fores, 1993; Henry nd Thoms, 2002; Anten et l., 2005, 2006, 2009). Both fctors cn ccount for our results: lifted plnts might hve received higher R/FR rtio in incident light nd stronger winds thn others, leding to suppression of stem growth. This will e nlysed in forthcoming pper. In summry, it ws found tht plnts in crowded stnds regulte their height growth to mintin similr height to their neighours even when there re potentil dvntges to height growth. Lifted plnts incresed neither iomss lloction to leves nor iomss production, indicting tht overtopping neighours is not necessrily eneficil in competing plnts. There my e other constrints for overtopping. A likely constrint is wind lowing, which my reduce mechnicl stility nd/or cuse deteriortion in the wter sttus of the plnts. ACKNOWLEDGMENTS We thnk two nonymous referees, nd N. Osd, Y. Osone, H. Tned nd M. Tteno for vlule comments nd suggestions, nd K. Stoh for the experimentl set-up. This work ws supported y Reserch Fellowships of the Jpn Society for the Promotion of Science for Young Scientists (nos 3025 nd 40172) to H.N. nd y KAKENHI (nos nd ) to K.H. LITERATURE CITED Ai S, Kohym T Tree species strtifiction in reltion to llometry nd demogrphy in wrm-temperte rin forest. Journl of Ecology 84: Anten NPR, Hirose T Biomss lloction nd light prtitioning mong dominnt nd suordinte individuls in Xnthium cndense stnds. Annls of Botny 82: Anten NPR, Csdo-Grci R, Ngshim H Effects of mechnicl stress nd plnt density on mechnicl chrcteristics, growth, nd

7 Ngshim & Hikosk Height growth regultion in crowded stnd 213 lifetime reproduction of tocco plnts. The Americn Nturlist 166: Anten NPR, Csdo-Grci R, Pierik R, Pons TL Ethylene sensitivity ffects chnges in growth ptterns, ut not stem properties, in response to mechnicl stress in tocco. Physiologi Plntrum 128: Anten NPR, von Wetterg EJ, Pwlowski M, Huer H Interctive effects of spectrl shding nd mechnicl stress on the expression nd costs of shde voidnce. The Americn Nturlist 173: Aphlo PJ, Rikl R Spcing of silver irch seedlings grown in continers of equl size ffects their morphology nd their vriility. Tree Physiology 26: Assmnn E Principles of forest yield study. Oxford: Pergmon Press. Bllré CL Keeping up with the neighours: phytochrome sensing nd other signlling mechnisms. Trends in Plnt Science 4: Bllré CL Illuminted ehviour: phytochrome s key regultor of light forging nd plnt nti-herivore defence. Plnt, Cell & Environment 32: Bllré CL, Scopel AL, Snchez RA Fr-red rdition reflected from djcent leves: n erly signl of competition in plnt cnopies. Science 247: Bllré CL, Scopel AL, Jordn ET, Vierstr RD Signling mong neighoring plnts nd the development of size inequlities in plntpopultions. Proceedings of the Ntionl Acdemy of Sciences of the USA 91: Biro RL, Hunt ER, Erner Y, Jffe MJ Thigmomorphogenesis: chnges in cell-division nd elongtion in the internodes of mechniclly pertured or ethrel-treted en-plnts. Annls of Botny 45: Csl JJ, Bllré CL, Tourn M, Snchez RA Antomy, growth nd survivl of long-hypocotyl mutnt of Cucumis stivus deficient in phytochrome-b. Annls of Botny 73: Corré WJ Growth nd morphogenesis of sun nd shde plnts. 1. The influence of light-intensity. Act Botnic Neerlndic 32: Corré WJ Growth nd morphogenesis of sun nd shde plnts. 2. The influence of light qulity. Act Botnic Neerlndic 32: Drke BG, Rschke K, Slisury FB Tempertures nd trnspirtion resistnces of Xnthium leves s ffected y ir temperture, humidity, nd wind speed. Plnt Physiology 46: Dudley SA, Schmitt J Testing the dptive plsticity hypothesis: density-dependent selection on mnipulted stem length in Imptiens cpensis. The Americn Nturlist 147: Ford ED Competition nd stnd structure in some even-ged plnt monocultures. Journl of Ecology 63: Givnish TJ On the dptive significnce of lef height in forest hers. The Americn Nturlist 120: Goudrin J Crop micrometeorology: simultion study. Simultion Monogrphs. Wgeningen: Pudoc. Grce J The effect of wind on grsses. 1. Cuticulr nd stomtl trnspirtion. Journl of Experimentl Botny 25: Henry HAL, Thoms SC Interctive effects of lterl shde nd wind on stem llometry, iomss lloction, nd mechnicl stility in Autilon theophrsti (Mlvcee). Americn Journl of Botny 89: Hikosk K, Sudoh S, Hirose T Light cquisition nd use y individuls competing in dense stnd of n nnul her, Xnthium cndense. Oecologi 118: Hikosk K, Ymno T, Ngshim H, Hirose T Light-cquisition nd use of individuls s influenced y elevted CO 2 in even-ged monospecific stnds of Chenopodium lum. Functionl Ecology 17: Huer H, Wiggermn L Shde voidnce in the clonl her Trifolium frgiferum: field study with experimentlly mnipulted vegettion height. Plnt Ecology 130: Huer H, Fijn A, During HJ A comprtive study of spcer plsticity in erect nd stoloniferous hers. Oikos 81: Hutchings MJ, Brkhm JP An investigtion of shoot integrtions in Mercurilis perennis L., rhizomtous hrceous perennil. Journl of Ecology 64: Jffe MJ, Fores S Thigmomorphogenesis: the effect of mechnicl perturtion on plnts. Plnt Growth Regultion 12: King DA Tree form, height growth, nd susceptiility to wind dmge in Acer scchrum. Ecology 67: King DA, Dvies SJ, Tn S, Noor NSM Trees pproch grvittionl limits to height in tll lowlnd forests of Mlysi. Functionl Ecology 23: Kir T Community rchitecture nd orgnic mtter dynmics in tropicl lowlnd rin forests of Southest Asi with specil reference to Psoh Forest, West Mlysi. In: Tomlinson PB, Zimmermnn MH. eds. Tropicl trees s living systems. Cmridge: Cmridge University Press, Koym H, Kir T Intrspecific competition mong higher plnts. VIII. Frequency distriution of individul plnt weight s ffected y the interction etween plnts. Journl of the Institute of Polytechnics, Osk City University Ser. D7: Kohym T, Hr T, Tdki Y Ptterns of trunk dimeter, tree height nd crown depth in crowded Aies stnds. Annls of Botny 65: Kuroiw S Intrspecific competition in rtificil sunflower communities. Botnicl Mgzine Tokyo 73: Mlikl SK, McDonnell K, Dudley SA, Schmitt J Effects of red to fr-red rtio nd plnt density on iomss lloction nd gs exchnge in Imptiens cpensis. Interntionl Journl of Plnt Sciences 160: McMhon T Size nd shpe in iology. Science 179: Mtsumoto Y, Oikw S, Ysumur Y, Hirose T, Hikosk K Reproductive yield of individuls competing for light in dense stnd of n nnul, Xnthium cndense. Oecologi 157: Ngshim H The processes of height-rnk determintion mong individuls nd neighourhood effects in Chenopodium lum L. stnds. Annls of Botny 83: Ngshim H, Tershim I Reltionships etween height, dimeter nd weight distriutions of Chenopodium lum plnts in stnds: effects of dimension nd llometry. Annls of Botny 75: Ngshim H, Tershim I, Ktoh S Effects of plnt density on frequency distriutions of plnt height in Chenopodium lum stnds: nlysis sed on continuous monitoring of height-growth of individul plnts. Annls of Botny 75: Nikls KJ Plnt iomechnics: n engineering pproch to plnt form nd function. Chicgo, IL: The University of Chicgo Press. Nikls KJ The scling of plnt height: comprison mong mjor plnt cldes nd ntomicl grdes. Annls of Botny 72: Nikls KJ Influence of tissue density-specific mechnicl-properties on the scling of plnt height. Annls of Botny 72: Nikls KJ The llometry of sfety-fctors for plnt height. Americn Journl of Botny 81: Nikls KJ Interspecific llometries of criticl uckling height nd ctul plnt height. Americn Journl of Botny 81: Nikls KJ Plnt height nd the properties of some herceous stems. Annls of Botny 75: Nikls KJ Computing fctors of sfety ginst wind-induced tree stem dmge. Journl of Experimentl Botny 51: Nishimur E, Suzki E, Irie M, Ngshim H, Hirose T Architecture nd growth of n nnul plnt Chenopodium lum in different light climtes. Ecologicl Reserch 25: Ogw H, Yod K, Ogino K, Kir T Comprtive ecologicl studies on three min types of forest vegettion in Thilnd. II. Plnt iomss. Nture nd Life in Southest Asi 4: Putz FE, Coley PD, Lu K, Montlvo A, Aiello A Uprooting nd snpping of trees: structurl determinnts nd ecologicl consequences. Cndin Journl of Forest Reserch 13: Ri SN Dimeter/height nd dimeter/girth reltionship of some rin forest tree species of Krntk, Indi. Mlysin Forester 42: Schmitt J, McCormc AC, Smith H A test of the dptive plsticity hypothesis using trnsgenic nd mutnt plnts disled in phytochromemedited elongtion responses to neighors. The Americn Nturlist 146: Smith H Light qulity, photoperception, nd plnt strtegy. Annul Review of Plnt Physiology 33: Smith H Phytochromes nd light signl perception y plnts n emerging synthesis. Nture 407: Speck O Field mesurements of wind speed nd reconfigurtion in Arundo donx (Pocee) with estimtes of drg forces. Americn Journl of Botny 90: Sterck FJ, Bongers F Ontogenetic chnges in size, llometry, nd mechnicl design of tropicl rin forest trees. Americn Journl of Botny 85:

8 214 Ngshim & Hikosk Height growth regultion in crowded stnd Telewski FW Growth, wood density, nd ethylene production in response to mechnicl perturtion in Pinus ted. Cndin Journl of Forest Reserch 20: Thoms SC Asymptotic height s predictor of growth nd llometric chrcteristics in Mlysin rin forest trees. Americn Journl of Botny 83: Vermeulen PJ, Anten NPR, Schieving F, Werger MJA, During HJ Height convergence in response to neighour growth: genotypic differences in the stoloniferous plnt Potentill reptns. New Phytologist 177: Wtts S, Rodriguez JL, Evns SE, Dvies WJ Root nd shoot growth of plnts treted with scisic cid. Annls of Botny 47: Weiner J, Fishmn L Competition nd llometry in Kochi scopri. Annls of Botny 73: Weiner J, Thoms SC Competition nd llometry in three species of nnul plnts. Ecology 73: Weiner J, Berntson GM, Thoms SC Competition nd growth form in woodlnd nnul. Journl of Ecology 78:

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