EFFECTS OF SHADING ON SOME MORPHOLOGICAL AND PHYSIOLOGICAL CHARACTERISTICS OF BEGONIA SEMPERFLORENS

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1 Pk. J. Bot., 5(6): 73-79, 8. EFFECTS OF SHADING ON SOME MORPHOLOGICAL AND PHYSIOLOGICAL CHARACTERISTICS OF BEGONIA SEMPERFLORENS YUANBING ZHANG, AIRONG LIU *, XUEPING ZHANG AND SHOUCHENG HUANG Anhui Sin n Thnology Univrsity, Fngyng 33, P.R. Chin *Corrsponing uthor s mil: liur@hstu.u.n Astrt Pott Bgoni smprflorns wr xpos to iffrnt lvls of shing (5%, 5%, 75%, 87%, n 93%) to invstigt th rspons of th morphologil n physiologil hrtristis of this plnt. Rsults rvl tht, s th lvl of shing inrs, th lf r, wtr ontnt, suproxi nion (O - ) proution rt, mlonilhy (MDA) ontnt n plsm mmrn prmility xhiit inrsing trns whn ompr with nturl light. By ontrst, th rltiv ontnts of nthoynin, solul sugr, strh n th suproxi ismuts (SOD) tivity isply rsing trns. Howvr, numr of flowrs pr plnt, hlorophyll ontnt, nitrt ruts (NR), n proxis (POD) tivitis initilly inrs ut susquntly rs. Comprhnsiv nlysis init tht iffrnt light intnsitis xrt iffrnt grs of influn on th morphologil hrtristis, ornmntl qulity n physiologil hrtristis of B. smprflorns. This plnt n pt from full illumintion to 87% shing, n th numr of flowrs inrs unr 5 75% shing, t whih th st ornmntl qulity ws lso osrv. Ky wors: Shing, Bgoni smprflorns, Antioxint pity, Morphologil hrtristis, Physiologil hrtristis. Introution Light is n importnt nvironmntl ftor tht ffts plnt growth, promots plnt photosynthsis in th form of nrgy, n rgults morphognsis in th form of signls (Smith, ). Th growth nvironmnts of plnts vry us of iffrnt light intnsitis (Shi t l.,3), whih xrt vrying influns on th shp, strutur, photosynthti hrtristis, growth, n vlopmnt of plnts (Xi, 3; Liu t l., 5). For xmpl, shing inrss plnt hight ut rss stm imtr, lf r inx, n lf thiknss (Shi t l., 6). In ition, shing n rgult th ron n nitrogn mtolism of plnts (Evns & Poortr, ; Zhi t l., ; Song t l., ; Zho & Ci, ; Chn t l., 6). At prsnt, groun ovr plnts with fst growth, strong rsistn, rih olor, n high ornmntl vlu hv om inispnsl prts of morn grn lnsping n grning sign. Suh plnts r typilly plnt on r groun, sloping lns, or forst gps unr iffrnt lighting onitions. Hn, thir growth is fft y vrying lvls of shing. Thrfor, stuying th ptility or sh-tolrn ility of groun ovr plnts to light intnsity is of grt thortil signifin n n srv s prtil guin for th sintifi onfigurtion of grn plnts. Bgoni smprfluorns is prnnil hrous grou ovr plnt n is th most ommon spis of th gnus Bgoni. It is hrtriz y roun plnt typ with flowrs tht r opn for four ssons, n its viwing prio is long. Both th flowrs n osrvl lvs of B.smprfluorns n sily plnt with othr flowr plnts. It is minly foun in wrm nvironmnts, n it isplys grt rsistn to ht, moistur, n rought. Howvr, it is not ol-rsistnt n is snsitiv to sunlight. B. smprfluorns hv om inrsingly utiliz in southrn Chin. B. smprfluorns grow grn lvs n r oppr lvs, n th lttr is onsir ttr in right summr nvironmnts. Anthoynin synthsis is inu primrily y th umultion of low-tmprtur rohyrts in utumn. Illumintion is nothr inispnsl ftor in th inution of nthoynin synthsis (Zhng t l., 3; Zhng t l., 6). Howvr, fw rports hv invstigt th ffts of iffrnt light intnsitis on th morphologil n physiologil hrtristis of B. smprfluorns. Thrfor, in our stuy, pott B. smprfluorns plnts wr trt with iffrnt lvls of shing, n thir morphologil hrtristis, orgni ontnt of smll moluls, n ntioxint pity wr ssss to trmin th light intnsity suitl for thir growth. Morovr, th physiologil mhnisms of light rsistn wr stui. Th finings of this stuy provi insights into th physiologil mhnism of lightfstnss n rfrns for plnting ornmntl grssln rsours with xllnt ornmntl qulity, goo growth, n strong rsistn to strss in lnsping prtis. Mtrils n Mthos Cultivtion of B. smprfluorns: B. smprfluorns slings wr provi y Shnghi Yunyi Sling Co., Lt. On Otor, 6, th B. smprfluorns slings wr grown unr th sm onition n trnsplnt into plsti pots (3 m high n 8 m in imtr), whih wr fill with th sm mount of wsh fin sn. Th xprimnt onsist of 9 pots ontining two plnts h. Th pots wr pl in sh, wtr with tp wtr for ys, n thn trt with/ onntrtion of Hogln nutrint solution. Aftr ys, th pott plnts wr trnsfrr to grnhous n trt with Hogln nutrint solution. Th mngmnt msurs wr th sm, n th plnts wr grown until April, 7. Thn, th plnts wr mov unr nturl light n pross on April, 7.

2 7 YUANBING ZHANG ET AL., Diffrnt shing trtmnt: Shing nts of iffrnt spifitions wr us to implmnt iffrnt lvls of shing. Th shing nts wr st t hight of.5 m from th groun. Th xprimnt ws st up with six trtmnts-on for 5 pots. Th ontrol ws nturl light (full light, % light trnsmission, i.., CK). Th iffrnt lvls of shing onsist of 5% shing (75% trnsmittn), 5% shing (5% trnsmittn), 75% shing (5% trnsmittn), 87% shing (3% light trnsmittn), n 93% shing (7% trnsmittn). Th light intnsity ws msur with YF-7 igitl illuminomtr t fiv points for h trtmnt: 6, 9,, 5, n 8. Light trnsmittn=light intnsity of shing trtmnt/nturl light %, shing = (-light trnsmittn) %. All othr mngmnt msurs wr onstnt uring th prossing, n msurmnt ws onut ftr 5 ys. Msurmnt mtho: Photos of th trt plnts n thir lvs wr ptur. Th numr of flowrs pr plnt ws ount for 6 plnts pr trtmnt. Th rlvnt initors wr msur using th lvs of th sm lf position, n th msurmnt ws rpt thri for h trtmnt. Th physiologil initors, suh s frsh wight, Dry wight, hlorophyll n nthoynin ontnt, solul sugr ontnt, strh ontnt, nitrt ruts (NR) tivity, SOD tivity, POD tivity n MDA ontnt wr trmin oring to prvious mtho (Liu & Zho, 5; Liu & Zho 5; Liu t l., 6). Dt nlysis: Th rsults wr xprss s th stnr vition of th mn of thr rplits, n multipl omprisons wr prform using DPS t prossing softwr. Rsults Shing ffts on plnt growth, numr of flowrs, lf r, lf wtr ontnt, hlorophyll ontnt, n rltiv ontnt of nthoynin: As shown in Fig., th ontrol plnts h th lowst plnt hight n th most ompt plnt typ in ll trtmnts. Th ontrol plnts isply th rkst r lvs, th smllst r, n upwr symmtry with th miris. As th lvl of shing ws inrs, th plnt hight inrs, th plnt typ grully loosn, th lf r grully inrs, th lf surf tn to flttn, n th lf olor grully hng from r to grn. As shown in Tl, ompr with th ontrol group, th numr of flowrs pr plnt initilly inrs n susquntly rs unr 5% 75% shing. Th numr of flowrs pr plnt ws highr thn tht of th ontrol, showing rution from.79% to 3.%. Unr th 87% 93% shing rts, th numr of flowrs pr plnt ws only 65.3% n 5.% of th ontrol, whih init signifint iffrn. As th lvl of shing ws inrs, oth th lf r n th lf wtr ontnt isply inrsing trns y 6.99% 9.86% n.39%.7%, rsptivly, unr 5% 93% shing. Th iffrns wr signifint. Unr 5% 87% shing, th hlorophyll ontnt show n inrsing trn y 69.86% 87.5% rltiv to th ontrol. Th hlorophyll ontnt unr 93% shing ws lowr thn tht unr 87% shing ut ws still grtr thn th ontrol y 3.95%. Th rltiv ontnt of nthoynin rs with inrsing lvl of shing. Unr 5% 93% shing, th rltiv ontnt of nthoynin rs y 66.87% 96.% rltiv to th ontrol, n th iffrn ws signifint. Effts of shing on solul sugr ontnt, strh ontnt n NR tivity: As shown in Fig., th solul sugr, n strh ontnts xhiit ownwr trn s th lvl of shing inrs, rsing y 9.6% 7.5% n.7% 77.8%, rsptivly, rltiv to th ontrol. Th iffrn in solul sugr ontnt unr 5% 5% shing n th ontrol ws insignifint, whrs tht unr 75% 93% shing signifintly iffr from th ontrol. Th strh ontnt unr 5% 93% shing signifintly iffr from th ontrol. As th lvl of shing inrs, th NR tivity initilly inrs n susquntly rs. Unr 5% 5% shing, NR tivity inrs from 7.79% to 5.%. Th NR tivity unr 75% 93% shing ws 3.8% 7.6% lowr thn th ontrol. Fig.. Growth of B. smprfluorns plnts n thir lvs unr full light n sh trtmnts.

3 Nitrt ruts tivity [µg/(g.h)] Strh ontnt (mg/g) Solul sugr ontnt (mg/g) SHADING INFLUENCE THE GROWTH OF B. SEMPERFLORENS 75 Tl. Effts of shing on flowr numr, lf r, lf wtr ontnt, hlorophyll ontnt, n rltiv ontnt of nthoynin. Shing prnt Flowr numr Lf r Wtr ontnt Chlorophyll Rltiv ontnts (numr/plnt) (m ) of lf (%) ontnt (mg/g) of nthoynins f % % % % % Fig.. Effts of shing on nitrt ruts tivity, solul sugr, n strh ontnts. 5% 5% 75% 87% 93% Shing prntg 5% 5% 75% 87% 93% Shing prntg 5% 5% 75% 87% 93% Shing prntg Shing ffts on SOD n POD tivitis, O - proution rt, MDA ontnt, n plsm mmrn prmility: Compr with th ontrol, th SOD tivity ws ru y 8.3% 89.38% unr 5% 93% shing, n th iffrn ws signifint (Fig. 3). As th lvl of shing ws inrs, th POD tivity initilly inrs n susquntly rs, n it ws highr thn th ontrol. Th POD tivity ws strongst unr 75% shing, rhing 39.7 U/(g min), whih ws 88.38% highr thn tht of th ontrol, n th iffrn ws signifint. Th POD tivity unr 93% shing rh only 6.53 U/(g min) ut ws 3.8% highr thn tht of th ontrol, n th iffrn ws signifint. Compr with th orrsponing ontrol, O - proution rt, MDA ontnt, n plsm mmrn prmility inrs y 59.83% 56.75%,.% 39.6%, n.9% 8.3%, rsptivly, unr 5% 93% shing. Th O - proution rt unr 5% 93% shing ws signifintly highr thn tht of th ontrol. MDA ontnt n plsm mmrn prmility unr 5% shing wr not signifintly iffrnt from thos of th orrsponing ontrol, whrs thos unr 5% 93% shing signifintly iffr from th ontrol. Disussion Aftr 5 ys of trtmnt, th B. smprfluorns plnts m ompt n short unr full light, isplying smll lvs n symmtril sinusoil upturns in th miri. Thus, th vrtil projtion r ws ru, n th lf olor hng from grn to r to ru th rint nrgy of full illumintion n voi th slf-prottion ginst th mg us y strong light. This phnomnon is morphologil ptiv hng. As th lvl of shing, th plnt hight inrs, th rown with inrs, th plnt typ loosn, th lvs xpn n flttn, n th lf olor hng from r to grn. Thn, th light intnsity grully wkn. This phnomnon srvs s mns to ptur mor solr rition nrgy to pt to th morphologil rspons of th shing nvironmnt. Thrfor, this hng fft th morphologil hrtristis. Plnt typ, rown with, plnt hight, lf r, lf olor, n lf xtnsion wr snsitiv to iffrnt light intnsitis. Rlt stuis suggst tht unr shing onitions, plnts inrs thir hight n rown with to xpn th r of light rption in th vrtil n horizontl irtions n thus otin mor light. Morovr, suh rspons nhns th plnt s ility to riv iffus n sttr light. Th prsnt stuy is onsistnt with th litrtur(mols, 998; Prlt t l., ).

4 Mmrn prmility (%) O -gnrtion rt MDA ontnt(nmol/g) SOD tivity (U/g) POD tivity (U/g min) 76 YUANBING ZHANG ET AL., f 5 5% 5% 75% 87% 93% 5% 5% 75% 87% 93% Shing prntg Shing prntg % 5% 75% 87% 93% 5% 5% 75% 87% 93% Shing prntg Shing prntg 8 6 5% 5% 75% 87% 93% Shing prntg Fig. 3. Effts of shing on SOD n POD tivitis, O - proution rt, MDA ontnt, n plsm mmrn prmility. Anthoynins r flvonois, n thir synthsis rquirs phnyllnin mmoni-lys (PAL) to triggr. PAL is light-inu nzym, n its tivity is positivly orrlt to light, suggsting tht light promots th synthsis of nthoynins (Ju t l., 995). Prvious stuis suggst tht strong light signifintly inu th xprssion of nthoynin-synthsis-rlt gns n inrs th umultion of nthoynin in lvs. Low light or rknss inhiits th synthsis n umultion of nthoynins (Mol t l., 996). Th iosynthsis of hlorophyll rquirs light, n th xtnt of plnt s hlorophyll ontnt rflts th strngth of its photosynthti pity (Hssnin t l., 3). This xprimnt ws prform to xmin th hngs in th numr of flowrs, lf r, lf wtr ontnt, rltiv nthoynin ontnt, n hlorophyll ontnt of B. smprfluorns unr h trtmnt. Brifly, nthoynin synthsis ws promot unr full light. Howvr, th inhiition of hlorophyll synthsis rsult in th wkst photosynthti ility. Th onsumption of mor

5 SHADING INFLUENCE THE GROWTH OF B. SEMPERFLORENS 77 photosynthti prouts y th nthoynin synthsis l to th rution of th photosynthti prouts rquir for flowr orgn growth. Consquntly, th numr of flowrs pr plnt ws ru, n th plnt growth ws wkn. Th low wtr ontnt in th lvs ws u to th high tmprtur of th surf lyr unr strong light n strong trnspirtion fft. Mort light ttnution (5% 75% shing) inhiit nthoynin synthsis n promot hlorophyll synthsis, thry nhning th photosynthti pity. Givn tht th mount of photosynthti prouts us for nthoynin synthsis ws signifintly ru, th numr of flowrs ws inrs n ws vn grtr thn tht unr full light. In ition, th plnt growth ws nhn, n th wtr ontnt ws inrs u to th ru surf tmprtur n th wkn trnspirtion. Unr 87% 93%, lthough th hlorophyll ontnt ws high, lss photosynthti prouts wr umult us th light intnsity ws too wk. Consquntly, th numr of flowrs ws mrkly rs, n th ovrshowing signifintly ru th surf tmprtur, furthr wkning th trnspirtion n inrsing th wtr ontnt in th plnt. Strong light promots nthoynin synthsis n inhiits hlorophyll synthsis, whrs wk light hs th opposit fft (Mninlli t l., 975). Th osrv hngs in th rltiv ontnts of nthoynin n hlorophyll in this xprimnt wr onsistnt with th finings of prvious stuis. In ition, this xprimnt show tht svrly fiint light (93% shing) xrt n inhiitory fft on hlorophyll iosynthsis. Anothr stuy suggst tht low tmprturs fvor nthoynin synthsis (Zhng t l., ), whrs high tmprturs supprss th gn xprssion n ru th ontnt of nthoynins (Islm t l., 5). In our stuy, ompr with full light, th tmprtur rs n nthoynin synthsis ws inhiit s th lvl of shing ws rs. Ths rsults wr inonsistnt with prvious stuis, suggsting tht th nthoynin synthsis in B. smprflorns is mor snsitiv to strong light thn low tmprtur. At prsnt, nthoynin umultion is liv to isply ths thr spts: nthoynins n fn ginst photo-strutiv ffts, suh s soring visil light to ttnut inint light nrgy n soring ultrviolt light to ru its mg (Zhng t l., ); thy t s ntioxints to fftivly svng rtiv oxygn spis (ROS); n thy t s solul osmorgultorsin vuols (Zhng t l., ). Th lrg umultion of nthoynins unr full light my lso prsnt ths physiologil ttriuts. Thrfor, th light nrgy sor y th B. smprflorns lvs xs th rng in whih photosynthsis n utiliz. To voi th mg inu y xss light nrgy, lrg mount of nthoynins is synthsiz n umult. This rspons is physiologilly ptiv hng to full light strss. Solul sugrs n strh r ky initors of ron mtolism in plnts. Sugr n promot th iosynthsis of nthoynins (Zhng t l., 3). In this stuy, solul sugr, strh, n nthoynin wr umult unr full light, n shing ru th solul sugr n strh ontnts. Ths rsults wr onsistnt with prvious stuis (Wng & Fng, ). Howvr, th hlorophyll ontnt ws th lowst unr full light onitions, initing tht th photosynthti pity wk. Thus, th mhnisms y whih th solul sugr n strh umult r unlr. Svrl stuis hv invstigt th spil physiologil rsponss of plnts unr strss onitions (Zhng & Xu, 3). Plnt growth is rltivly slow unr vrs onitions, thry ruing th n for photosynthti prouts, suh tht lrg mounts of solul sugr n strh r umult in th lvs (Munns, 993). This rsult my lso th omin fft of th ov mntion two ftors, n it is physiologil rspons to totl light strss. Mortly ru light (5% 75% shing) my l to inrs plnt growth n inrs numr of flowrs, whih onsum mor nutrints, rsulting in rs solul sugr n strh ontnts. As illumintion ws furthr wkn (87% 93% shing), th solul sugr n strh ontnts wr mrkly ru proly u to th svr lk of light. - NR is th first ky nzym tht tlyzs NO 3 rution. It is light-inu nzym tht is losly rlt to th us of nitrt nitrogn y plnts. Illumintion not only rgults NR tivity ut lso influns th nitrogn mtolism y inuing its synthsis (Mglhs & Hur, 99). In this xprimnt, th sh NR tivity initilly inrs n susquntly rs, whih ws inonsistnt with th rs NR tivity in sh soyn, otton, n Mnhurin sh lvs (Huo, 9; Yu t l., ), initing tht th NR tivity of iffrnt plnts rspons iffrntly to th physiologil rsponss to iffrnt light intnsitis. Thrfor, with rgr to th us of nitrt nitrogn in B. smprflorns, th light my mortly wkn (i.., 5% 5% shing), whih ws fvorl to NR tivtion or th inrs in th xprssion mount. - Thus, th NO 3 ruing ility ws strong, n th N supply ws high, thry inrsing th numr of flowrs pr plnt. Whn th totl illumintion n shing wr 75%, th NR tivtion ws inhiit or th xprssion ws ru. Th ility to ru NO - 3 ws ru, n th N supply ws rltivly wkn, thry rsing th numr of flowrs pr plnt. Unr 87% 93% shing, th NR tivity ws shrply wkn, th N supply ws rstilly ru, n th numr of flowrs pr plnt ws lso shrply ru. In viw of th isturn of th ln systm of th proution n rmovl of ROS in plnts unr vrsity strss, lrg mount of ROS umult, using or xrting mmrn lipi proxition n rsulting in mmrn mg. SOD n POD r ntioxint nzyms in plnts. SOD is isproportiont O - to H O, n O, POD n oxiiz H O into wtr. Ths nzyms work togthr to rmov ROS from lls, ru th toxi ffts of ROS on plnts (Lópzuio t l., ), n prott ll struturs ginst mg (Blokhin t l., 3). Thrfor, thy r oftn us s fftiv initors of plnt ntioxint strss. Prvious

6 78 YUANBING ZHANG ET AL., stuis init tht nthoynins n ru light nrgy ptur n qunh oxygn fr rils (Wng t l., 997). In this xprimnt, th SOD tivity ws th strongst n th rltiv ontnt of nthoynins ws highst unr full light. Although th POD tivity ws th lowst, th proution rt n MDA ontnt of O - wr th lowst, n th prmility of th plsm mmrn ws th smllst. Ths rsults wr prsum minly u to th rmovl of ROS (suh s O - n H O ) y SOD n nthoynins. Thrfor, th mmrn lipi proxition ws th lightst, n th mg to th plsm mmrn ws miniml. Unr 5% 75% shing, th SOD tivity n rltiv ontnt of nthoynin rs, n th POD tivity inrs. Howvr, th O - proution rt, MDA ontnt, n plsm mmrn prmility inrs, initing tht th ility to lr O - ws ru. Givn tht only strong POD tivity n rmov H O, O - still umult. Mmrn lipi proxition n plsm mmrn mg wr lso inrs. Unr 87% 93% shing, th SOD tivity n th rltiv ontnt of nthoynin rs shrply, whrs POD mintin high tivity. Th O - proution rt, MDA ontnt, n plsm mmrn prmility mrkly inrs, initing tht ROS (prtiulrly O - ) ws iffiult to rmov y singl POD, whih is strongly pnnt on tivity. Thus, mmrn lipi proxition n plsm mmrn injury wr grully inrs. Conlusion In summry, unr full light, B. smprflorns xhiit r lf olor, lrg numr of flowrs, upwr lf urling (ov th top), ompt plnt typ, mort rution nitrt pity, n strong ntioxint pity. Howvr, th plnt growth ws wk u to th low hlorophyll ontnt. Unr 5% 5% shing, th lf olor hng from r to grn, n th uppr lf rspons rs. In viw of th strong rution nitrt pity, th ntioxint pity ws strong, n th numr of flowrs ws th grtst unr trtmnt. Morovr, th plnt typ n th ornmntl qulity wr th st. Unr 75% 93% shing, th lvs wr grn n flt. Th rution nitrt pity ws signifintly wkn, th mmrn lipi proxition grully inrs, th numr of flowrs ws grully rs, n th plnt typ tn to loos. Aknowlgmnts This work ws support y th griulturl sin n thnology hivmnts trnsformtion fun projt of Anhui provinil sin n thnology prtmnt (37) n th sintifi n thnologil projt of Anhui provinil sin n thnology prtmnt (333). W lso thnk th ky rsrh n vlopmnt projts of Anhui sin n thnology prtmnt (778) Sin n thnology projt of Chuzhou Muniipl Sin n Thnology Buru of Anhui Provin (76) for thir finil support. Rfrns Blokhin, O., E. Virolinn n K.V. Fgrstt. 3. Antioxints, oxitiv mg n oxygn privtion strss: A rviw. Ann Bot., 9(): Co, G.H. n R.L. Prior Oxygn ril soring pity of nthoynins. J. Agri. Foo Chm., 5(): Chn, Q., S.W. Yu, X.M. Jing, Y. Zho, X.Y. Mng n X.C. Wn. 6. Efft of sh trtmnt in summr on th xprssion of gns rlt to thnin iosynthsis in t plnts (Cmlli sinnsis). Bull. Bot. Rs., 36(): 6-3. Evns, J.R. n H. Poortr.. Photosynthti limtion of plnts to growth irrin: Th rltiv importn of spifi lf r n nitrogn prtitioning in mximizing ron gin. Plnt Cll Env., (8): Hssnin, R.A., S.A. El-Khws, S.K. Irhim, H.M. El- Bssiouny, H.A. Mostf n A.A.A. El-Monm. 3. Improving th thrmo tolrn of wht plnt y folir pplition of rginin or putrsin. Pk. J. Bot., 5(): -8. Huo, C.F., H.L. Sun, Z.Q. Wng, Z.Q. Fn n X.M. Zho. 9. Effts of light n nitrogn on growth, ron n nitrogn mtolism of Frxinus mnshuri slings. Si. Sil. Sin., 5(7): 38-. Islm, M.S., M. Jlluin, J.O. Grnr, M. Yoshimoto n O. Ymkw. 5. Artifiil shing n tmprtur influn on nthoynin ompositions in swtpotto lvs. Hortsi., 39(): Ju, Z.G., Y.B. Yun, C.L. Liou n S.H. Xin Rltionships mong phnyllnin mmoni-iys tivity, simpl phnol onntrtions n nthoynin umultion in ppl. Si. Hort., 6(3-): 5-6. Liu A.R. n K.F. Zho. 5. Effts of slt strss on th growth n th nitrt ruts tivity in Thllungill hlophil. J. Plnt Physiol. Mol. Bio., 3(5): Liu, A.R. n K.F. Zho. 5. Osmoti umultion n its rol in osmoti justmnt in Thllungill hlophilunr slt strss. J. Plnt Physiol. Mol. Bio., 3(): Liu, A.R., Y.B. Zhng n D.K. Chn. 6. Effts of slt strss on th growth n th ntioxint nzym tivity of Thllungill hlophil. Bull. Bot. Rs., 6(): 6-. Liu, Z.B., R.M. Chng, W.F. Xio, Q.S. Guo n N. Wng. 5. Effts of shing on growth n photosynthti hrtristis of istylium hinns slings. Si. Sil. Sin., 5(): Lópzuio, J., M.F. Nitojoo, V. Rmírzroríguz n L. Hrrrstrll.. Orgni i mtolism in plnts: From ptiv physiology to trnsgni vritis for ultivtion in xtrm soils. Plnt Si., 6(): -3. Mglhs, J.R. n D.M. Hur. 99. Rspons of mmonium ssimiltion nzyms to nitrogn form trtmnts in iffrnt plnt spis. J. Plnt Nutri., (): Mninlli, A.L., C.P. Yng, P. Linquist, O.R. Anrson n I. Rino Photoontrol of nthoynin synthsis: Iii. Th tion of strptomyin on th synthsis of hlorophyll n nthoynin. Plnt Physiol., 55(): Mol, J., G. Jnkins, E. ShãoFr, D. Wiss n V. Wlot Signl prption, trnsution, n gn xprssion involv in nthoynin iosynthsis. Plnt Si., 5(5-6): Mols, J Chngs in morphologil n ntomil strutur of g (Brssi olr L.) outr lvs n in ultrstrutur of thir hloroplsts us y n in vitro xss of nikl. Photosynthti, 3(): Munns, R Physiologil prosss limiting plnt growth in slin soils: Som ogms n hypothss. Plnt Cll Env., 6(): 5-.

7 SHADING INFLUENCE THE GROWTH OF B. SEMPERFLORENS 79 Prlt, G., J.L. Pérz-Lloréns, I. Hrnánz n J.J. Vrgr.. Effts of light vilility on growth, rhittur n nutrint ontnt of th sgrss zostr noltii hornm. J. Exp. Mrin Bio. Eo., 69(): 9-6. Shi, J.G., H.Y. Cui, B. Zho, S.T. Dong, P. Liu n J.W. Zhng. 3. Efft of light on yil n hrtristis of grinfilling of summr miz from flowring to mturity. Si. Agr. Sin., 6(): 7-3. Shi, X.D., Z.Q. Wn, Y.F. Liu n W.J. Wng. 6. Effts of shing on growth n photosynthti pilitis of too lvs of igr-wrppr us. At Bot. Borli- Ointli Sin., 6(8): Smith, H.. Phytohroms n light signl prption y plnts--n mrging synthsis. Ntur, 7(68): Song, Y.X., W.Y. Yng, Z.X. Li, T.M. Yong n L. Liu.. Efft of miz-soyn rly ropping sh on nitrogn mtolism of soyn slings. Chin. J. Oil Crop Si., 3(3): Wng, J.F. n Y.L. Fng.. Th fft of light intnsity on iomss llotion, lf morphology n rltiv growth rt of two invsiv plnts. At Phytool. Sin., 8(6): Xi, X.J., X.H. Yng n X.Y. Chn. 3. Effts of shing on lf shp n photosynthti hrtristis of th trnsgni Lspz formos with xprssing BADH gn. Si. Sil. Sin., 9(3): 33-. Yu, S., Y.H. Wng, Z.G. Zhou, F.J. Lü, J.R. Liu, Y.N. M, J. Chn n A. Auurzik.. Efft of sh on nitrogn mtolism n its mhnism in otton plnt t flowring n oll-forming stg. At Agr. Sin., 37(): Zhng, K.M., H.J. Yu, S. Ki, Y.H. Zhou, J.Q. Yu n X.J. Xi.. Photoprottiv rols of nthoynins in Bgoni smprflorns. Plnt Si., 79(3): -8. Zhng, K.M., M.L. Guo, D. H, R.H. Wu n Y.H. Li. 6. Th inhiition fft n xssiv ron flux rsulting from loking nthoynin iosynthsis unr rknss in Bgoni smprflorns. J. Plnt Growth Rg., 35(): -3. Zhng, K.M., Z. Li, Y. Li, Y.H. Li, D.Z. Kong n R.H. Wu. 3. Crohyrt umultion my th proximt triggr of nthoynin iosynthsis unr utumn onitions in Bgoni smprflorns. Plnt Bio., 5(6): 99-. Zhng, L. n J.R. Xu. 3. Stuis on physiologil n iohmil rsponss of Roini Psuoi lons unr wtr strss. Si. Sil. Sin., 39(): Zho, T.T. n X. Ci.. Physiologil n iohmil hrtrs of t-plnts with iffrnt shing grs. Hunn Agr. Si., 5: 38-. Zhi, Z.G., Y.L. Mng n S. Pi.. Efft of shing uring sling prio on th strutur of otton stm n lf n photosynthti prformn of funtionl lf. Si. Agr. Sin., 3(5): (Riv for pulition 9 Dmr 7)

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