Erythrina speciosa (Leguminosae-Papilionoideae) under soil water saturation: morphophysiological and growth responses

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1 Annls of Botny 1: , 29 doi:1.193/o/mcp159, ville online t Erythrin specios (Leguminose-Ppilionoidee) under soil wter sturtion: morphophysiologicl nd growth responses Cmilo L. Medin 1, Mri Cristin Snches 2, *, Mri Luiz S. Tucci 3, Crlos A. F. Sous, Gerldo Rogério F. Cuzzuol 5 nd Crlos A. Joly 1 1 Deprtmento de Biologi Vegetl, IB, Universidde Estdul de Cmpins, CP 619, Cmpins, SP, Brzil, 2 Universidde Federl de Uerlândi, Instituto de Biologi, CP 593, Uerlândi, MG, Brzil, 3 CHORT, Instituto Agronômico (IAC), CP 28, Cmpins, SP, Brzil, EMBRAPA Meio-Norte, Av. Duque de Cxis, CP 565, Teresin, PI, Brzil nd 5 Universidde Federl do Espírito Snto, Av. Fernndo Ferrri 51, Cmpos de Goieirs, ES, Brzil Received: 18 Ferury 29 Returned for revision: 18 Mrch 29 Accepted: 1 June 29 Pulished electroniclly: 5 July 29 Bckground nd Aims Erythrin specios is Neotropicl tree tht grows minly in moist hitts. To chrcterize the physiologicl, morphologicl nd growth responses to soil wter sturtion, young plnts of E. specios were sujected experimentlly to soil flooding. Methods Flooding ws imposed from 2 to cm ove the soil surfce in wter-filled tnks for 6 d. Non-flooded (control) plnts were well wtered, ut never flooded. The net CO 2 exchnge (A CO2 ), stomtl conductnce (g s ) nd intercellulr CO 2 concentrtion (C i ) were ssessed for 6 d. Solule sugr nd free mino cid concentrtions nd the proportion of free mino cids were determined t, 7, 1, 21, 28 nd 5 d of tretments. After 28, 5 nd 6 d, dry msses of leves, stems nd roots were determined. Stem nd root cross-sections were viewed using light microscopy. Key Results The A CO2 nd g s were severely reduced y flooding tretment, ut only for the first 1 d. The solule sugrs nd free mino cids incresed until the tenth dy ut decresed susequently. The content of sprgine in the roots showed drstic decrese while those of lnine nd g-minoutyric incresed shrply throughout the first 1 d fter flooding. From the 2th dy on, the flooded plnts reched A CO2 nd g s vlues similr to those oserved for non-flooded plnts. These events were coupled with the development of lenticels, dventitious roots nd erenchym tissue of honeycom type. Flooding reduced the growth rte nd ltered cron lloction. The iomss llocted to the stem ws higher nd the root mss rtio ws lower for flooded plnts when compred with non-flooded plnts. Conclusions Erythrin specios showed 1 % survivl until the 6th dy of flooding nd ws le to recover its metolism. The recovery during soil flooding seems to e ssocited with morphologicl ltertions, such s development of hypertrophic lenticels, dventitious roots nd erenchym tissue, nd with the mintennce of neutrl mino cids in roots under long-term exposure to root-zone O 2 deprivtion. Key words: Erythrin specios, erenchym, mino cid content, iomss lloction, photosynthesis, flooding dpttions, stomtl conductnce, O 2 deficiency, g-minoutyric cid (GABA). INTRODUCTION The occurrence of flooding events is common in severl Brzilin ecosystems. Their effect on physiologicl nd morphologicl responses in plnts is consequence of prtil decline of O 2 concentrtion in the soil. Under this condition, the decrese in the rtes of eroic root respirtion is detectle nd hence ATP production is impired nd metolism is disrupted (Lmers et l., 1998). Among the min consequences of oxygen depletion on plnts re decrese in stomtl conductnce nd photosynthesis, chnges in metolite levels, plnt hormonl imlnces, nd deficient wter nd nutrient uptke with deleterious effects on plnt growth nd survivl (Lopez nd Kursr, 23). However, some plnts hve cquired some chrcteristics to survive nd grow under neroic conditions. The mechnisms of tolernce to flooding conditions re multiple nd complex, * For correspondence. E-mil snchesmc@yhoo.com.r including responses t ll levels of orgniztion from iochemicl to ntomicl nd morphologicl ltertions. Some works hve pointed out decrese in shoot growth in severl species sumitted to flooding conditions, with flooding suppressing lef formtion nd expnsion of leves nd internodes, nd inducing premture lef senescence nd scission (Tng nd Kozlowski, 1982; Tsukhr nd Kozlowski, 1986; Kozlowski, 1997), while others (Loo nd Joly, 1998; Joly, 199; Lopez nd Kursr, 1999; Andrde et l., 1999) hve shown tht flooding did not lter growth responses, in either tolernt or non-tolernt species. Lenticel nd stem hypertrophy, erenchym formtion nd dventitious roots re mong the min morphologicl ltertions tht llow woody plnts to continue growing nd surviving in flooding environments (Prolin, 21). Wter, nutrients nd hormones supplied y dventitious roots to eril orgns provide the conditions for continued growth, nd in some species the concomitnt lenticel hypertrophy will further contriute to porosity increment t the insertion of dventitious roots, enhncing the # The Author 29. Pulished y Oxford University Press on ehlf of the Annls of Botny Compny. All rights reserved. For Permissions, plese emil: journls.permissions@oxfordjournls.org Downloded from y guest on 1 Ferury 218

2 672 Medin et l. Erythrin specios under wter sturtion rhizosphere oxidtion (Armstrong et l., 199; Vrtpetin nd Jckson, 1997; Colmer, 23). From physiologicl perspective, reduced CO 2 ssimiltion hs een shown for gret numer of species sumitted to neroic conditions (Mielke et l., 23; Fernndez, 26). Such reductions re often ssocited with stomtl limittions, which restricted CO 2 vilility in the mesophyll nd t the sites of croxyltion (Lopez nd Kursr, 23). Non-stomtl limittions re lso oserved which hve een ttriuted to strch ccumultion in the leves nd/or drop in RUBISCO ctivities (Pezeshki, 21). In ddition, pronounced chnges in the pool of severl metolites, prticulrly free mino cids, hve een detected (Sous nd Sodek, 23; Koppitz et l., 2; Thoms et l., 25), s well s in the crohydrte vilility to the root system under hypoxic conditions (Hung nd Johnson, 1995). Although the literture on plnt responses to flooding is undnt for temperte species, there re few reports out the flooding effect on net CO 2 ssimiltion nd growth in tropicl wild species (Lopez nd Kursr, 1999, 23; Prolin, 21; Mielke et l., 23). For exmple, Genip mericn (Ruicee), widely distriuted neotropicl species found in flood-prone hitts in Brzil, ut most commonly occurring on terr firme elsewhere, is le to survive nd to grow under soil wterlogging despite decreses in net cron uptke (Mielke et l., 23). Moreover, nlogies etween flooding tolernce mechnisms in temperte nd tropicl plnts my e misleding since plnts in temperte res typiclly experience flooding during the winter nd erly spring, period of miniml physiologicl ctivity, while tropicl plnts hve to fce flooding events during summer when ir temperture is higher nd physiologicl ctivities more intense (Joly, 199; Lopez nd Kursr, 1999). Erythrin specios, neotropicl tree species distriuted throughout southern nd south-estern Brzil (Krukoff, 1939; Lorenzi, 1992), is typicl in fluvil forest nd moist plins s well s in flooded-prone hitts, lwys in open nd secondry formtions. In the lst two decdes, studies of the flooding effects on some species of the genus Erythrin hve een relted to metolic fetures t erly seedling estlishment nd with physiologicl spects of growth, cron ssimiltion nd the photosynthetic mchinery itself (Smll et l., 1989; Muthuchelin et l., 1995). For E. specios, Kol et l. (22) cme to the conclusion tht the mintennce of n dequte fermenttive metolic energy is importnt for seedling dpttion to flooded hitts. The ojective of the present study ws to evlute concomitntly lef gs exchnge, solule sugr content, free mino cid content nd the proportion of free mino cids in root tissue, nd morphologicl responses nd growth in young individuls of E. specios under soil flooding. Tking into ccount the distriution pttern of the species, it ws hypothesized tht E. specios is le to show morphologicl nd physiologicl ltertions to cope with soil flooding. Experimentl conditions MATERIALS AND METHODS The experiment ws crried out from Mrch to My 1999, in glsshouse t Universidde Estdul de Cmpins, São Pulo Stte, Brzil, t 2285 S; 785 W nd 67 m.s.l. According to Köppen, the climte is Cw, with wrm nd riny seson from Octoer to Mrch (temperture rnge from 18 to 3 8C), nd drier seson from April to Septemer (temperture rnge from 12 to 27 8C). The men nnul rinfll in the region is 157 mm precipittion (Ortolni et l., 1995). Seeds of Erythrin specios Andrews from ripe fruits were collected from dult individuls growing in flood re ner the University. After clening, the seeds were scrified nd germinted in Petri dishes t 25 8C under continuous white fluorescent light. Germinted seeds were trnsplnted to plstic pots (1 L) filled with snd nd supplied with 2 ml of Hoglnd solution per week. After 5 d the seedlings with one pir of cotyledonry leves nd three fully expnded leves were selected nd ssigned to one of two tretments: flooding nd non-flooding (control). Flooding ws imposed in wter-filled tnks for 6 d, providing wter level from 2 to cm ove the soil surfce. Non-flooded plnts were well wtered ut never flooded. Lef gs exchnge mesurements CO 2 nd H 2 O gs exchnge ws mesured from 9 to 11 h. Two fully expnded nd helthy leves from nodes 2 3 from the pex of the shoots from eight different individuls in ech tretment (flood nd non-flooding) were used. The net CO 2 uptke per unit lef re (A CO2 ), stomtl conductnce to wter vpour (g s ), trnspirtion rte (E H2 ), lef temperture (T lef ) nd intercellulr CO 2 concentrtion (C i ) were determined with portle open-system infrred gs nlyser (IRGA) (LCA-; Anlyticl Development Co. Ltd, Hoddesdon, UK) connected to 6.25 cm 2 PLC (B) lef chmer. The ir entering y the lef chmer ws drwn from 3 m ove-ground nd pssed through 15 cm 3 uffer vessel to void mient CO 2 concentrtion (C ) fluctution. Wter use efficiency (WUE) ws clculted s the rtio A CO2 /E H2O (Noel, 25). The IRGA llowed the mesurement of T lef nd photosynthetic photon flux density (PPFD, l ¼ 7 nm). During the mesurements the plnts were trnsferred to n open re where temperture nd reltive humidity rnged from 2 to 25 8C, nd from 75 to 78 %, respectively. The PPFD ws out 11 mmol m 22 s 21, high enough to sturte photosynthesis, s previously determined in n dditionl experiment (dt not shown). Amino cids nd solule sugrs in roots Immeditely fter hrvest, the root system ws ground with liquid N 2 nd lyophilized. For the metolite nlyses, only the originl, pre-existing roots were smpled. Amino cids nd solule sugrs were extrcted with MCW (methnol : chloroform : wter, 12 : 5 : 3, v/v/v) (dpted from Bieleski nd Turner, 1966), for 2 h, using 1 ml g 21 fresh weight of roots. The queous phse ws recovered following phse seprtion on stnding fter ddition of chloroform (1 vol.) nd wter (1.5 vol.) to vols of superntnt. The queous phse ws then reduced to known volume y evportion t 38 8C nd kept frozen until nlysis. Solule sugrs were determined ccording to Grhm nd Smydzuc (1965). Amino cids were determined using the method of Moore Downloded from y guest on 1 Ferury 218

3 Medin et l. Erythrin specios under wter sturtion 673 nd Stein (198) nd seprted y reverse-phse HPLC of their o-phthlldehyde (OPA) derivtives, s descried y Puitti nd Sodek (1999). The totl dissolved mino cids re given s the sum of ll detected nd quntified mino cids. The contents re given in solute (mmol g 21 d. wt.) nd in reltive mol percentge of totl units. All these evlutions were mde using three plnts per tretment, t (non-flooded plnts), 7, 1, 21, 28 nd 5 d of the experiment. Growth nd morphologicl mesurements The stem height nd the dimeter t 2 cm ove the soil were mesured weekly for ten flooded nd ten non-flooded plnts. After 28, 5 nd 6 d, six plnts from ech tretment were hrvested, the roots were thoroughly wshed nd the plnts were seprted into leves, stems nd roots. These smples were dried t 7 8C for 8 h for dry mss determintion. From the primry dt the following prmeters were derived: root mss rtio (RMR: root mss per whole plnt mss, units g g 21 ), stem mss rtio (SMR: stem nd petiole mss per unit whole plnt mss, units g g 21 ) nd lef mss rtio (LMR: lef mss per whole plnt mss, units g g 21 ). The reltive growth rte (RGR: dry mss increment per unit totl plnt mss per unit time) ws clculted ccording to Hunt (1982) s: RGR ¼ [ln (finl dry mss) ln (initil dry mss)]/time. For histologicl nlysis, smll pieces of root nd stem were cut with lmin rzor from flooded nd non-flooded plnts nd fixed in 5 FAA (formldehyde : 5 % ethnol : cetic cid, 18 : 1 : 1) nd stored in 7 % lcohol. Hnd-cut trnsverse sections were tken, stined y str lue-sic fuchsin, nd photogrphed using compound microscope optic. Testing for oxygen diffusion from the shoot to the root system ws performed fter 1 d on flooded nd non-flooded plnts y the sumergence of the root system into gel gr (1 %), 15 mg L 21 sodium dithionite nd 1 mg L 21 methylene lue for colorimetric determintion ccording to Joly (1991). Sttistics The effects of flooding on gs exchnge, solule sugrs, free mino cids, proportion of mino cids, nd growth were nlysed in fully rndomized design. The dt were suject to n nlysis of vrince (ANOVA), nd when F ws significnt the tretment mens were compred y Tukey s test t 5 % proility (Snedecor nd Cochrn, 1967). The dt of free mino cids nd solule sugrs were expressed s the mens nd stndrd error. RESULTS Lef gs exchnge responses The lef gs exchnge vriles were drsticlly ffected y flooding tretments. After d of flooding, A CO2 ssimiltion rtes were reduced y 9 % from control vlues. The A CO2 vlues from control plnts were out 5.88 mmol m 22 s 21 while the minimum A CO2 vlue from flooded plnts ws.79 mmol m 22 s 21. The stomtl conductnce nd WUE closely prlleled chnges in A CO2 ssimiltion rtes. Leves from control plnts exhiited mximum g s vlue of.12 mmol m 22 s 21 while for flooded plnts it ws.1 mol m 22 s 21. After the fifth dy grdul recupertion ws detected, nd t the end of the experiment flooded nd nonflooded plnts showed similr vlues of A CO2 nd g s. However, the WUE presented tendency to decline t the end of the experiment, indicting possile non-stomtl limittion of the photosynthesis (Fig. 1). A tendency for increses in C i vlues ws oserved t the eginning of the experiment (Fig. 2). Amino cids nd solule sugrs in the root system The totl free mino cid concentrtion nd the solule sugrs in the roots of E. specios re shown in Figs 3 nd. Although the vlues of free mino cids nd solule sugrs g s (mol m 2 s 1 ) A CO2 (µmol m 2 s 1 ) Wter use efficiency (µmol CO2 mmol 1 H2O ) A B C Dys of tretment Control Flooded F IG. 1. Photosynthetic rtes (A CO2 ), stomtl conductnce (g s ) nd wter use efficiency (WUE) in Erythrin specios under control nd flooding conditions. Dt re expressed s the men + s.e.; n ¼ leves per tretment, ech lef from different individul. Different letters indicte significnt difference etween mens (P,.5 %; Tukey test). Downloded from y guest on 1 Ferury 218

4 67 Medin et l. Erythrin specios under wter sturtion C i (µmol mol 1 ) Dys of tretment Control Flooded F IG. 2. Internl CO 2 concentrtion (C i )inerytrin specios under control nd flooding conditions. Dt re expressed s the men + s.e.; n ¼ leves per tretment, ech lef from different individul. Different letters indicte significnt difference etween mens (P,.5 %; Tukey test). Free mino cids (µmol g 1 f. wt) Dys of tretment F IG. 3. Chnges in totl free mino cids in root tissue of E. specios plnts sujected to flooding of the root system. Dt re the mens + s.e. of three replictes. Solule sugrs (µmol g 1 f. wt) Dys of tretment F IG.. Chnges in solule sugrs in root tissue of E. specios plnts sujected to flooding of the root system. Dt re the mens + s.e. of three replictes. were not significntly different, oth incresed slightly until the tenth dy of imposition of the flooding tretment nd fter tht presented tendency to decrese until dy 5. The composition of free mino cids is presented in Tle 1. At dy (non-flooded plnts), sprgine (Asn) ws the mino cid found in the highest proportion in roots of E. specios, followed y g-minoutyric cid (GABA), glutmine (Glu), sprtic cid (Asp) nd lnine (Al). Although GABA is non-protein mino cid, it lredy showed elevted vlues on dy. Under flooded conditions, Asn levels fell shrply nd remined low until dy 21 of the experiment. Concomitntly increses were oserved in Al content tht lsted until dy 28. However, the GABA content ws high when compred with tht of other mino cids t dy 5, nd Al recovered vlues close to its initil vlues. In contrst to GABA nd Al, the Glu nd Asp levels decresed on the tenth dy of flooding imposition, ut fter this recovery of the content of these mino cids ws oserved. Growth nd morphologicl responses Flooding hd no effect on growth height. Both control nd flooded plnts hd reched heights of out 13 cm y the end of the experiment. In contrst, from the 1th to the th dy, growth in stem dimeter ws 2 % higher in the flooding tretments compred with the control (Fig. 5). The RGR ws not ffected until the 28th dy of tretment, ut significnt decreses with flooding were oserved etween the 5th nd the 6th dy (Fig. 6). The pttern of cron lloction ws lso ffected y flooding tretment. The iomss frction llocted to the stem ws higher on the 28th nd 5th dys, nd the RMR nd LMR were lower on the 5th dy for flooded plnts when compred with the non-flooded controls (Fig. 7). In generl, the lef, root nd totl iomss, s well s the root/ shoot rtio, were ffected y flooding round the 5th dy; fter tht only root nd totl iomss differed etween control nd flooded plnts (Tle 2). Increses in cron lloction to the stem oserved t the th dy of flooding were coupled with the formtion of hypertrophic lenticels (Fig. 8), dventitious roots nd erenchym tissue (Fig. 9). The ltter seems to e of the honeycom type (Justin nd Armstrong, 1987; Sego et l., 25). The development of lenticels ws oserved 2 d fter the tretment imposition nd ws prlleled y the emergence of dventitious roots, which proliferted throughout the experiment. Fifteen dys fter the eginning of the experiment, the erenchym tissue on flooded plnts ws lredy widely developed (Fig. 9). Oxygen diffusion from roots in gel gr ws detected 1 d fter flooding, mening tht the morphologicl modifictions oserved in flooded plnts were contriuting to etter ertion of the root system. DISCUSSION Under flood, E. specios went through intricte processes involving iochemicl nd ntomicl, s well s morphologicl effects. This species ws highly sensitive to root oxygen deprivtion considering tht 9 % decrese in g s vlues ws oserved immeditely fter exposure to tretments. Decreses in g s nd A CO2 ssimiltion rtes re frequent Downloded from y guest on 1 Ferury 218

5 Medin et l. Erythrin specios under wter sturtion 675 TABLE 1. Amino cid composition (mol %) of roots of Erythrin specios plnts sujected to flooding of the root system Time fter induction of flooding (d) Amino cid* Asp Glu Asn Ser Al c c c GABA For the metolite nlyses, only the originl pre-existing roots were smpled. n ¼ 3 replictes. * The following mino cids were detected: Gln, His, Gly, Thr, Arg, Tyr, Met, Phe, Ile nd Leu; however t concentrtions lower thn 3%. Mens followed y sme letter in row do not differ y Tukey test t the 5 % level of proility. Height (cm) Stem dimeter (mm) response to soil flooding conditions in oth temperte (Grvtt nd Kiry, 1998; Co nd Korner, 1999) nd tropicl species (Lopez nd Kursr, 1999, 23; Núñez-Elise et l., 1999; Mielke et l., 23). According to Grvtt nd Kiry (1998), shrp decline of stomtl conductnce t the eginning of exposure to low O 2 vilility is typicl for sensitive species. These results indicte tht E. specios experienced some level of stress, even though it is normlly plnt of flood-prone hitts. The stomtl closure under flooded conditions might e relted to decrese in permeility nd root hydrulic conductnce under neroic conditions, which could hve induced internl wter stress, leding to Dys of tretment Control Flooded FIG. 5. Height nd stem dimeter for E. specios under control nd flooding conditions. Dt re expressed s men + s.e.; n ¼ 1 plnts per tretment. Different letters indicte significnt difference etween mens (P,.5 %; Tukey test). RGR (mg g 1 d 1 ) decresing lef turgor nd stomtl conductnce (Else et l., 21; Pezeshki, 21; Mielke et l., 23). The decrese in root hydrulic conductnce my e relted to fewer wter chnnel proteins in the plsm memrnes of the roots, due to lck of metolic energy to sustin their synthesis (Lmers et l., 1998). The other possiility is relted to some chemicl signls, prticulrly scisic cid (ABA), which my e synthesized in the root tips sumitted to hypoxi nd trnsported to leves, or in wilted leves nd susequently trnsported to young turgid leves (Zhng nd Zhng, 199; Jckson, 22). The mrked reduction in g s vlues ws followed y pronounced decline in net A CO2 uptke nd WUE. Stomtl closure impirs the CO 2 diffusion from the tmosphere into the mesophyll, leding to the reduction of sustrte vilility (C i ) for RUBISCO ctivities nd thus reductions in photosynthetic rtes (Chves, 1991). Nevertheless, tendency for n increse in C i ws detected 5 d fter flooding, indicting lso tht decreses in net CO 2 uptke might e relted to nonstomtl limittions. The incresing C i coincided with phse when decreses in A CO2 ssimiltion were more severely ffected y flooding tretment. At the sme time, the flooding tretment lso dversely ffected the WUE once the flooding Dys of tretment Control Flooded FIG. 6. Reltive growth rte of E. specios under control nd flooding conditions. Dt re expressed s the men + s.e.; n ¼ 6 plnts in ech tretment. Different letters indicte significnt difference etween mens (P,.5 %; Tukey test). Downloded from y guest on 1 Ferury 218

6 676 Medin et l. Erythrin specios under wter sturtion LMR (g g 1 ) SMR (g g 1 ) A B Control Flooded TABLE 2. Growth nd iomss chrcteristics of Erythrin specios seedlings grown under soil flooding nd control conditions Time (d) Prmeter Tretment Stem iomss (g) Control Flooded Lef iomss (g) Control Flooded Root iomss (g) Control Flooded Totl iomss (g) Control Flooded Root/shoot rtio Control Flooded n ¼ 6 replictes. Mens followed y sme letter in column do not differ y Tukey test t the 5 % level of proility. RMR (g g 1 ) C A Dys of tretment F IG. 7. (A) Lef mss rtio (LMR; lef dry mss/totl dry mss); (B) stem mss rtio (SMR; stem dry mss/totl dry mss); (C) root mss rtio (RMR; root dry mss/totl dry mss) in E. specios under control nd flooding conditions. Dt re expressed s the men + s.e.; n ¼ 6 plnts in ech tretment. Different letters indicte significnt difference etween mens (P,.5 %; Tukey test). tretment decresed net CO 2 uptke to greter extent thn trnspirtion rtes. Furthermore, in some circumstnces stem photosynthesis my ecome sustntil component of cron gin during the period of cute stress imposed y flooding (Armstrong nd Armstrong, 25; Teskey et l., 28). This spect deserves future investigtion, since some dult individuls of E. specios exhiits green woody tissues just elow the outer rk. In their study with severl trees from sesonlly flooded forest t Venezuel, Fernndez et l. (1999) showed tht the reduction in net CO 2 uptke induced y flooding ws ssocited with generl inhiition of metolism cused y hypoxic soil conditions. As consequence of energy shortge, the generl inhiition of metolism in E. specios ws perceived y increses in free mino cid content in their root tissue. The decline in the rte of protein synthesis leding to lower demnd for mino cids in the hypoxic conditions could explin the ccumultion of free mino cids oserved during erly tretment imposition (Reggini nd Bertni, 23). B FIG. 8. Stem trnsverse sections of E. specios under (A) control conditions nd (B) fter 6 d of flooded conditions. Note the lenticel hypertrophy on plnts under flooded conditions. Scle rs ¼ 1 mm. From the tenth to the 2th dy fter the eginning of the experiment, E. specios ws le to recover stomtl conductnce nd the net CO 2 uptke. The sme pttern ws descried for Txodium distichum (Pezeshki, 1993; Pezeshki et l., 1996) nd Melleuc quinquinervi (Senn-Gomes nd Kozlowski, 198), ut not for G. mericn (Mielke et l., 23), which did not exhiit complete stomtl reopening nd net CO 2 ssimiltion recovery fter 63 d of hypoxi, in spite of the high rte of seedling survivl under these conditions (Andrde et l., 1999; Mielke et l., 23). Downloded from y guest on 1 Ferury 218

7 Medin et l. Erythrin specios under wter sturtion 677 F IG. 9. Root trnsverse sections of Erythrin specios under (A) control conditions nd (B) fter 6 d of flooded conditions. Root sections were mde etween 2 nd 3 cm from the root tip, in roots with length of no more thn 5 6 cm. Note the erenchym development in flooded plnts. Scle rs ¼ 1 mm. Some uthors consider the cpcity for lef conductnce (g s ) resumption fter flooding to e the physiologicl trit tht confers flood tolernce, once it llows the net CO 2 exchnge to continue, with the consequent plnt growth nd survivl under hypoxic conditions (Pezeshki, 1993; Li et l., 2). This fct ws oserved for Priori copifer, tree species tht forms monodominnt stnds in sesonlly flooded hitts in Pnm tropicl forests (Lopez nd Kursr, 1999). In prllel with photosynthesis recovery, on the 21st dy decreses in mino cid concentrtions were A B oserved. It is possile tht these decreses were cused y greter demnd for mino cids during flooding for the iosynthesis of neroiclly induced enzymes (Vrtpetin nd Jckson, 1997; Kreuzwieser et l., 22). Likewise, quntittive chnges, i.e. shift in composition of mino cids, re usully oserved in plnts under O 2 deprivtion (Kreuzwieser et l., 22; Reggini nd Bertni, 23; Sous nd Sodek, 23; Thoms et l., 25). Under flooding, the Asn, Asp nd Glu concentrtions decresed while the Al nd GABA concentrtions incresed on the seventh dy of the experiment. For instnce, on the tenth dy, the Al nd GABA composition represented up to 56 % of the totl mino cid content. These results reflect the interconversions etween mino cids tht help the mintennce of cell ph (Reggini et l., 1988; Crwford et l., 199; Fn et l., 1997; Reggini nd Bertni, 23). These mino cids formed were neutrl or less cidic, nd therefore this could counterct the elevtion of the cid metolite pool produced y fermenttive rections, such s tht of lctte. Our findings lso indicte tht increses in Al were proportionlly higher when compred with increses in GABA on the tenth dy of flooding imposition. Indeed, severl studies hve reported tht pronounced increse in Al in root tissues is typicl plnt response to O 2 deprivtion (Drew, 1997; Sous nd Sodek, 23). Considering tht Al is produced from the rection of pyruvte with mino-n (Good nd Muench, 1993; Reggini nd Bertni, 23), increses in Al in roots of E. specios my e consequence of pyruvte ccumultion, which could reflect poorly coordinte rections etween glycolysis nd lcoholic fermenttion (Kreuzwieser et l., 22). In ddition, it is noteworthy tht even the initil vlues of GABA concentrtion were high. The mount of GABA declined on the 28th dy nd therefter remined t elevted vlues until the 5th dy. GABA ccumultion hs lso een oserved in severl trees nd herceous species sumitted to flooding, nd in some cses hs een considered s n importnt physiologicl response to tolernce to hypoxi (Drew, 1997, Kreuzwieser et l., 22; Koppitz et l., 2). Besides the chnges in photosynthetic ctivities nd in the concentrtion of free mino cids, E. specios ws le to increse, lthough t lower rte, the vilility of solule sugrs in roots t the onset of flooding imposition. We suggested tht in E. specios the increse in solule sugrs could hve resulted from degrdtion of reserves present in roots or s function of cellulr ph cidifiction cused y L-lctte ccumultion (dt not shown). According to Roerts et l. (1992), the L-lctte ccumultion during hypoxi is enough to depress the metolism due to cytoplsmic cidosis nd consequently to reduce the consumption of the sustrtes used in respirtion. The solule sugrs re of specil importnce for plnts growing under O 2 deprivtion conditions due to their direct reltionship with respirtion. The switch from eroic towrd neroic metolism yields very few molecules of ATP. Hence lrger sugr pool tht cn e redily metolized is crucil for fermenttive metolism to e crried out properly (Grvtt nd Kiry, 1988; Chen et l., 22; Sirm et l., 28). This hypothesis ws elucidted in severl experiments where plnts kept under O 2 shortge fed with exogenous sugrs could mintin glycolysis, fermenttion nd root Downloded from y guest on 1 Ferury 218

8 678 Medin et l. Erythrin specios under wter sturtion longevity (Vrtpetin nd Jckson, 1997, nd references therein). Islm et l. (2) oserved tht lthough root sugrs declined in Pice mericn nd Lrix lricin, the ltter mintined higher sugr content throughout the experiment. These differences coincided with the production of dventitious roots during flooding in L. lricin ut not in P. mericn, conferring greter tolernce to hypoxi in the former species (Islm et l., 2). Proly similr event my hve hppened in E. specios since the content of solule sugrs declined fter morphologicl modifictions were induced y flooding. These results indicte tht the reserves were used to form dventitious roots nd hypertrophied lenticels s well s to sustin fermenttive metolism. The dventitious roots, the hypertrophied lenticels nd the erenchym my llevite the hypoxi stress, y fcilitting oxygen diffusion through low resistnt internl pthwy etween shoot nd root extremities, s well s higher flow of wter, nutrients nd hormones to the shoot offsetting the reduced metolic ctivities in the originl root system (Joly, 1996; Vrtpetin nd Jckson, 1997; Colmer, 23; Rengifo et l., 25). The honeycom erenchym s oserved on E. specios under flooded conditions (Fig. 9) ws proly developed y expnsion of intercellulr spces into lcune y cell division, wll seprtions nd cell expnsion, with no lysigeny or cell deth (Justin nd Armstrong, 1987; Sego et l., 25). Note tht some intercellulr spces re lso oserved in root cortex tissue from control plnts. This spect shows tht the precursor of erenchym structure is present even efore the occurrence of the flooding events. This might e n importnt trit in E. specios, indicting its likely tolernce to hypoxic conditions. In some species, erenchymtous dventitious roots ecme woody even during the flooding events (Armstrong nd Armstrong, 25). Bsed on our field oservtions, the dventitious roots in E. specios re ephemerl, i.e. fter the flooding episode they shrivel nd re lost. The development of hypertrophied lenticels, dventitious roots nd erenchym coincided with the stomtl reopening nd the recovery of photosynthetic rtes. It ws oserved tht the lenticels do not promote enough ertion to resume the eroic metolism in plnts sujected to flooding conditions (Joly, 1991), ut for E. specios in the present study, the lenticels role in ertion ws verified using the gel gr technique. The O 2 diffusion from the se of the stem into the root system through erenchym is common trit for mny flood-tolernt plnts (Colmer, 23; Armstrong nd Armstrong, 25). Flooding generlly decreses the reltive growth rte nd ffects the prtitioning of cron (Lopez nd Kursr, 23; Mielke et l., 23), nd indeed the present study shows reductions in the RGR of the flooded plnts. These reductions were due to differentil pttern of photossimilte distriution. The RMR were lower in flooded compred with control plnts, which could hve een due to ltertions in metolic ctivities cused y O 2 deprivtion in the root environment, since lck of oxygen locks mitochondril electron trnsport, the oxidtion of NADH þ H þ nd ATP synthesis (Drew, 1997; Mielke et l., 25). On the other hnd, increses in SMR could e explined y the increment in stem dimeter in flooded plnts compred with the control. The increse in the stem dimeter is contriuted to y the production of the hypertrophied lenticels. The pttern of lloction of photosynthtes to root iomss (RMR) in E. specios ws higher when compred with G. mericn, neotropicl species tht shows wide distriution in flood-prone hitts (Mielke et l., 23). The root/shoot rtio ws lso higher in E. specios when compred with severl other tropicl tree species, from sesonlly flooded nd flood-free forests (Lopez nd Kursr, 1999, 23). As pointed out y these uthors, the higher investment in the root iomss in plnts sujected to flooding could e n importnt morphologicl dpttion. In some ecosystems, drought my follow flooding, nd species tht re le to withstnd some level of wter limittion fter period of flooding will proly successfully occupy environments which re sesonlly flooded (Lopez nd Kursr, 1999). Erythrin specios showed 1 % survivl throughout 6 d under soil flooding. Although decreses of 86 % in the photosynthetic rtes were oserved t the eginning of the experiment, E. specios showed complete recovery of lef gs exchnge rtes. The proportion etween cidic mino cids nd GABA ws mintined under long-term exposure to O 2 deprivtion, which proly contriuted to voidnce of cell cidifiction. Proly these responses could explin why the plnts continue their growth under soil flooding, lthough t lower rtes. It is importnt to emphsize tht the tolernce mechnisms exhiited y the young spling stge studied here my not e sufficient to sustin mture trees, which certinly require further studies. In conclusion, the coordinted chnges in physiologicl responses together with the development of typicl morphologicl ltertions (lenticels, dventitious roots, erenchym) when plnts were exposed to deprivtion of O 2 t the rhizosphere indicted tht E. specios showed the ility to thrive in wter sturtion conditions, consistent with its occurrence in flood-prone hitts (Prolin, 21). The high levels of GABA nd the presence of some erenchym even in root tissues of control plnts indictes tht seedlings of E. specios re redy to cope with flooding episodes. ACKNOWLEDGEMENTS We thnk Dr. Sndr M. C. Guerreiro from the Deprtment of Plnt Biology/Stte University of Cmpins (Unicmp) for the lortory fcilities in the morphologicl studies, Miss Jnete Myumi Okmoto for technicl ssistnce during ll the experiments nd Dr Vldenir Queiroz Rieiro from the Emrp Meio-Norte for the sttisticl nlyses. We re lso grteful to Professor Dr Tim Colmer nd nonymous reviewers for constructive comments on the mnuscript. The uthors grtefully cknowledge the finncil support of the Fundção deampro à Pesquis do Estdo de São Pulo FAPESP (grnt 3/ ). Crlos A. Joly ws supported y CNPq Productivity Fellowship (grnt 5233/99-). LITERATURE CITED Andrde ACS, Rmos FN, Souz AF, Loureiro MB, Bstos R Flooding effects in seedlings of Cythrexyllum myrinthum Chm. nd Downloded from y guest on 1 Ferury 218

9 Medin et l. Erythrin specios under wter sturtion 679 Genip mericn L.: responses of two neotropicl lowlnd tree species. Revist Brsileir de Botânic 22: Armstrong W, Armstrong J. 25. Stem photosynthesis not pressurized ventiltion is responsile for light-enhnced oxygen supply to sumerged roots of lder (Alnus glutinos). Annls of Botny 96: Armstrong W, Brndle R, Jckson MB Mechnisms of flood tolernce in plnts. Act Botnic Neerlndic 3: Bieleski RL, Turner NA Seprtion nd estimtion of mino cids in crude plnt extrcts y thin-lyer electrophoresis nd chromtogrphy. Anlyticl Biochemistry 17: Co FL, Conner WH Selection of flood-tolernt Populus deltoides clones for reforesttion projects in Chin. Forest Ecology nd Mngement 117: Chves MM Effects of wter deficits on cron ssimiltion. Journl of Experimentl Botny 2: Chen H, Qulls RG, Blnk RR. 22. Effect of soil flooding on photosynthesis, crohydrte prtitioning nd nutrient uptke in the invsive exotic Lepidium ltifolium. Aqutic Botny 82: Colmer TD. 23. Long-distnce trnsport of gses in plnts: perspective on internl ertion nd rdil oxygen loss from roots. Plnt, Cell nd Environment 26: Crwford LA, Bown AW, Breitkreuz KE, Guinel FC The synthesis of g-minoutyric cid in response to tretments reducing cytosolic ph. Plnt Physiology 1: Drew MC Oxygen deficiency nd root metolism: injury nd cclimtion under hypoxi nd noxi. Annul Review of Plnt Physiology nd Plnt Moleculr Biology 8: Else MA, Couplnd D, Dutton L, Jckson MB. 21. Decresed root hydrulic conductivity reduces lef wter potentil, initites stomtl closure nd slows lef expnsion in flooded plnts of cstor oil (Ricinnus communis) despite diminished delivery of ABA from the roots to shoots in xylem sp. Physiologi Plntrum 111: 6 5. Fn TWM, Higshi RM, Frenkiel TA, Lne AN Aneroic nitrte nd mmonium in flood-tolernt rice coleoptiles. Journl of Experimentl Botny 8: Fernndez MD, Pieters A, Donoso C, et l Sesonl chnges in photosynthesis of trees in the flooded forest of the Mpire River. Tree Physiology 19: Fernndez MD. 26. Chnges in photosynthesis nd fluorescence in response to flooding in emerged nd sumerged leves of Pouteri orinocoensis. Photosynthetic : Good AG, Muench DG Long-term neroic metolism in root tissue. Plnt Physiology 11: Grhm D, Smydzuc J Use of nthrone in the quntittive determintion of hexose phosphtes. Anlyticl Biochemistry 11: Grvtt DA, Kiry CJ Ptterns of photosynthesis nd strch lloction in seedlings of four ottomlnd hrdwood tree species sujected to flooding. Tree Physiology 18: Hung B, Johnson JW Root respirtion nd crohydrte sttus of two whet genotypes in response to hypoxi. Annls of Botny 75: Hunt R Plnt growth curves. The functionl pproch to growth nlysis. London: Edwrd Arnold. Islm MA, Mcdonld SE. 2. Ecophysiologicl dpttions of Blck spruce (Pice mrin) nd tmrck (Lrix lricin) seedlings to flooding. Trees 18: Jckson MB. 22. Long-distnce signlling from roots to shoots ssessed: the flooding story. Journl of Experimentl Botny 53: Joly CA Flooding tolernce in tropicl trees. In: Jckson MB, Dvies DD, Lmers H, eds. Plnt life under oxygen deprivtion. Ecology, physiology nd iochemistry. The Hgue, SPB Acdemic Pulishing, Joly CA Flooding tolernce: reinterprettion of Crwford s metolic theory. Proceedings of the Royl Society of Edinurgh 12B: Joly CA The role of oxygen diffusion to the root system on the flooding tolernce of tropicl trees. Revist Brsileir de Biologi 56: Justin SHFW, Armstrong W The ntomicl chrcteristics of roots nd plnt response to soil flooding. New Phytologist 16: Kol RM, Rwler A, Brendle R. 22. Prmeters ffecting the erly seedling development of four Neotropicl trees under oxygen deprivtion stress. Annls of Botny 89: Koppitz H, Dewender M, Ostendorf W, Schmieder K. 2. Amino cids s indictors in comom reed Phrgmites ustrlis ffected y extreme flood. Aqutic Botny 79: Kozlowski TT Responses of woody plnts to flooding nd slinity. Tree Physiology Monogrph 1: Kreuzwieser J, Fürniss J, Rennenerg H. 22. Impct of wterlogging on the N-metolism of flood tolernt nd non-tolernt tree species. Plnt, Cell nd Environment 25: Krukoff BA The Americn species of Erythrin. Brittoni 3: Lmers H, Chpin STIII, Pons TJ Plnt physiologicl ecology. Berlin: Springer-Verlg. Li S, Pesezki SR, Goodwin S, Shields FDJ. 2. Physiologicl responses of lck willow (Slix nigr) cuttings to rnge of soil moisture regimes. Photosynthetic 2: Loo PC, Joly CA Tolernce to hypoxi nd noxi in neotropicl tree species. In: Scrno FR, Frnco AC. eds. Ecophysioloicl strtegies of xerophytic nd mphyious plnts in the neotropics. Series Oecologi Brsiliensis, vol. IV. Rio de Jneiro: PPGE-UFRJ, Lopez OR, Kursr TA Flood tolernce of four tropicl tree species. Tree Physiology 19: Lopez OR, Kursr TA. 23. Does flood tolernce explin tree species distriution in tropicl sesonlly flooded hitts? Oecologi 136: Lorenzi H Árvores Brsileirs. Mnul de identificçãp e cultivos de plnts róres ntivs do Brsil. Nov Odess: Editor Plntrum. Mielke MS, Almeid AAF, Gomes FP, Aguilr MAG, Mngeir PAO. 23. Lef gs exchnge, chlorophyll fluorescence nd growth responses of Genip mericn seedlings to soil flooding. Environmentl nd Experimentl Botny 5: Mielke MS, Mtos EL, Couto VB, Almeid AAF, Gomes FP, Mngeir PAO. 25. Some photosynthetic nd growth responses of Annon glr L. seedlings to soil flooding. Act Botnic Brsilic 19: Moore S, Stein WH Photometric ninhydrin method for use in the chromtogrphy of mino cids. Journl of Biology Chemistry 176: Muthuchelin K, Murugn C, Hrigovindn R, Nedunchezhin N, Kulndivelu G Effect of tricontnol in flooded Erythrin vriegt seedlings. 1. Chnges in growth, photosynthetic pigments nd iomss productivity. Photosynthetic 31: Noel PS. 25. Physicochemicl nd environmentl plnt physiology. Sn Diego: Elsevier/Acdemic Press. Núñez-Elise R, Schffer B, Fischer JB, Collis AM, Crne JH Influence of flooding on net CO 2 ssimiltion, growth nd stem ntomy of Annon species. Annls of Botny 8: Ortolni AA, Cmrgo MBP, Pedro MJ Jr Norms climtológics dos pontos meteorológicos do Instituto Agronômico: 1. Centro Exeprimentl de Cmpins. Boletim Técnico, 155. Cmpins: Instituto Agronômico, 13pp. Prolin P. 21. Morphologicl nd physiologicl djustments to wterlogging nd drought in seedlings of Amzonin floodplin trees. Oecologi 128: Pezeshki SR Differences in ptterns of photosynthetic responses to hypoxi in flood tolernt nd flood sensitive tree species. Photosynthetic 28: Pezeshki SR. 21. Wetlnd plnt responses to soil flooding. Environmentl nd Experimentl Botny 6: Pezeshki SR, Prdue JH, Delune RD Lef gs exchnge nd growth of flood-tolernt nd flood sensitive tree species under low soil redox conditions. Tree Physiology 16: Puitti M, Sodek L Wterlogging ffects nitrogen trnsport in the xylem of soyen. Plnt Physiology nd Biochemistry 37: Reggini R, Bertni A. 23. Aneroic mino cid metolism. Russin Journl of Plnt Phsysiology 5: Reggini R, Cntú CA, Brmill I, Bertni A Accumultion nd interconversion of mino cids in rice roots under noxi. Plnt nd Cell Physiology 9: Rengifo E, Tezr W, Herrer A. 25. Wter reltions, chlorophyll fluorescence, nd contents of scchrides in tree species of tropicl forest in response to flood. Photosynthetic 3: Roerts JKM, Hooks MA, Miullis AP, Edwrds S, Wester C Contriution of mlte nd mino cid metolism to cytoplsmic ph regultion in hypoxic mize root tips studied using nucler mgnetic resonnce spectroscopy. Plnt Physiology 98: Sirm RK, Kumuth D, Ezhilmthi K, Deshmukh PS, Srivstv GC. 28. Physiology nd iochemistry of wterlogging tolernce in plnts. Biologi Plntrum 52: Downloded from y guest on 1 Ferury 218

10 68 Medin et l. Erythrin specios under wter sturtion Sego JL Jr, Mrsh LC, Stevens KJ, Soukup A, Votruová O, Enstone DE. 25. A re-exmintion of the root cortex in wetlnd flowering plnts with respect to erenchym. Annls of Botny 96: Senn-Gomes AR, Kozlowski TT Growth responses nd dpttions of Frxinus pennsylvnic seedlingstoflooding. Plnt Physiology 66: Smll JGC, Potgieter GP, Both FC Anoxic seed germintion of Erythrin cffr: ethnol fermenttion nd response to metolic inhiitors. Journl of Experimentl Botny : Snedecor GW, Cochrne WG Sttisticl methods, 6th edn. Ames, IA: Iow Stte University Press. Sous CAF, Sodek L. 23. Alnine metolism nd lnine minotrnsferse ctivity in soyen (Glycine mx) during hypoxi of the root system nd susequent return to normoxi. Environmentl ndexperimentl Botny 5: 1 8. Tng ZC, Kozlowski TT Some physiologicl nd morphologicl responses of Quercus mcrocrp seedlings to flooding. Cndin Journl of Forest Reserch 12: Teskey RO, Sveyn A, Stepper K, McGuire MA. 28. Origin, fte nd significnce of CO 2 in tree stems. New Phytologist 177: Thoms AL, Guerreiro SMC, Sodek L. 25. Aerenchym formtion nd recovery from hypoxi of the flooded root system of nodulted soyen. Annls of Botny 96: Tsukhr T, Kozlowski TT Importnce of dventitious roots to growth of flooded Pltnus occidentlis seedlings. Plnt nd Soil 88: Vrtpetin BB, Jckson MB Plnt dpttions to neroic stress. Annls of Botny 79 (Supplement A): 3 2. Visser EJW, Voesenek ACJ, Vrtpetin BB, Jckson MB. 23. Flooding nd plnt growth. Annls of Botny 91: Zhng J, Zhng X Cn erly wilting of old leves ccount for much of the ABA ccumultion in flooded pe plnts? Journl of Experimentl Botny 5: Downloded from y guest on 1 Ferury 218

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