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Chpter 3 Tolernce to Drought in Leguminous Plnts Medited y Rhizoium nd Brdyrhizoium Alln Klynger d Silv Loto, Joquim Alenísio Gomes d Silveir, Roerto Cezr Loo d Cost nd Cândido Ferreir de Oliveir Neto Additionl informtion is ville t the end of the chpter http://dx.doi.org/10.5772/54094 1. Introduction The wter vilility is considered the climtic fctor with lrge effect on griculturl productivity, eing responsile to determine species distriution in different climte zones round the gloe [1]. Effects of drought depend of plnt development stge, intensity, nd durtion of the wter restriction. In other hnd, plnt dptive strtegies will determine the tolernce level, nd consequently your survivl on these conditions of indequte wter supply [2]. Wter deficit is n iotic fctor tht ffects the griculturl production with high frequency nd intensity, influencing spects relted to plnt development, such s decrese in photosynthesis rte, reduction in lef re [3], nd stomt closing [4]. Crops normlly present performnce ffected y wter deficiency, which cn cuse lower growth nd development (Figure 1), with progressive reduction in lef dry mtter [5] nd consequent repercussion on production prmeters, such s numer of grins nd pods per plnt. Root system presents complex strtegy iming to mintin wter supply in conditions of wter deficit, y incresing the root elongtion rte nd completely inhiiting the shoot [6]. On the other hnd, plnts growing in low wter potentils normlly present root thinner [7], nd this morphologicl modifiction is n dpttion to increse wter sorption efficiency. Therefore, comintion of chnges in morphologicl, physiologicl nd iochemicl levels re necessry to plnt survivl in environments ffected y drought. 2013 d Silv Loto et l.; licensee InTech. This is n open ccess rticle distriuted under the terms of the Cretive Commons Attriution License (http://cretivecommons.org/licenses/y/3.0), which permits unrestricted use, distriution, nd reproduction in ny medium, provided the originl work is properly cited.
50 Responses of Orgnisms to Wter Stress control drought Figure 1. Visul spect of shoot in Phseolus vulgris plnts exposed to drought y four dys. The iologicl fixtion of nitrogen is the cpcity of n orgnism to divide the molecule of nitrogen (N 2 ) nd to comine hydrogen toms (H + ), forming mmonium (NH 4+ ) [8], eing crried out y distinct group of microorgnisms, singly or under symiosis. The Brdyrhizoium nd Rhizoium genders re descried s soil cteri tht hve ility to infect root hir of leguminous plnts, nd it to induce nodule formtion (Figure 2), with susequent fixtion of nitrogen [9]. Figure 2. Visul spect of root system of Vign unguicult plnts inoculted with Brdyrhizoium. The red rrows indicte nodules formed fter infection process.
Tolernce to Drought in Leguminous Plnts Medited y Rhizoium nd Brdyrhizoium http://dx.doi.org/10.5772/54094 51 The persistence of rhizoil strins, nd their symiotic performnce in current nd susequent sesons re ffected y numerous iotic nd iotic fctors [10], with drought stress nd nitrogen deprivtion, eing mong the most significnt in mny prts of the world [11]. Other importnt fctor is the root exudtion ility, which it will determine plnt microe ssocitions so tht the survivl nd tolernce of rhizoi during wter restriction. Molydenum is n essentil element for soil microorgnisms, since it serves s cofctor for different enzymes involved in the metolism of nitrogen, cron nd sulfur. Before the synthesis of molydoenzymes, uptke of molydte, which is the more stle form of molydenum), its ctivtion to n pproprite form, nd its incorportion into the orgnic frction of the molydenum-cofctors, re required [12]. The presence of molydenum is necessry during formtion of severl proteins, including the nitrogense, the molydoenzyme tht reduces tmospheric dinitrogen (N 2 ) into mmoni (NH 4+ ) [13]. This cterium is lso cple of denitrifiction, vi nitric oxide (NO) nd nitrous oxide (N 2 O) to N 2, when the cells re cultured under oxygen-limiting conditions [14]. The first rection of denitrifiction, is crried out y the periplsmic Mo-contining nitrte reductse [15]. In ddition, the rection under norml conditions is descried s N 2 + 8 e + 8 H + + 16 MgATP 2 NH 3 + H 2 + 16 MgADP + 16Pi. The enzyme mechnism requires reduction of the Fe protein y electron donors such s ferredoxin nd flvodoxin, trnsfer of single electrons from the Fe protein to the MoFe protein (which is dependent on MgATP hydrolysis) nd, finlly, internl electron trnsfer in the MoFe protein y the P cluster to the FeMo cofctor sustrte-inding site. Ech electrontrnsfer step requires n oligtory cycle of ssocition of the Fe nd MoFe proteins to form complex (Figure 3), fter which the two components dissocite [16]. Figure 3. Schemtic representtion of the nitrogense Fe protein cycle. The Fe protein dimer is shown in light lue with the cue representing the [4Fe 4S] cluster coloured lck to indicte the reduced form nd red to represent the oxidized form. The α nd β suunits of the MoFe protein re depicted s ornge nd pink, respectively, the yellow squres represent the P cluster nd the lck dimond represents the FeMo cofctor. Chnges in the oxidtion stte of the MoFe protein re not shown [16].
52 Responses of Orgnisms to Wter Stress Severl leguminous such s Vign unguicult nd Cicer rietinum re considered tolernt to wter deficit, nd importnt mechnisms were developed y this species to tolerte indequte wter supply. For exmple, iochemicl modifictions in cron metolism, such s increse in sucrose [17], s well s significnt interference in nitrogen metolism, like reduction of solule proteins [5] nd increse in totl mino cids [18] contriute to osmotic djustment of these plnts (Figure 4). Figure 4. Glutmine synthetse ctivity (), totl solule mino cids () nd totl solule proteins (c) in Vign unguicult plnts cv. Vit 7 sujected to 4 dys of wter restriction nd 2 dys of rehydrtion. Mens followed y the sme letter re not significntly different y the Tukey test t 5% of proility. The rs represent the men stndrd error nd the rrow the rrow indictes the rehydrtion point [5].
Tolernce to Drought in Leguminous Plnts Medited y Rhizoium nd Brdyrhizoium http://dx.doi.org/10.5772/54094 53 2. Ojective Aims of this chpter is to define (i) wter deficiency nd iologicl fixtion of nitrogen, to explin (ii) s this symiotic process cn promote eneficil repercussions to plnt nd microorgnism, nd to present (iii) the ttenution of negtive impcts on nodule nd plnt, esides nitrogen compounds nd morphologicl prmeters of plnts exposed to wter restriction. 3. Wter mintennce in lef nd nodule produced y inocultion Drought is environmentl component tht ffect crop yields worldwide. In nture, this stress is multifceted prolems tht re usully ssocited with other dverse circumstnces, which limit plnt performnce such s wter shortge nd nutrient deficits. In order to ssess the osmotic stress, Sssi et l. [19] monitored two Phseolus vulgris cultivrs inoculted with Rhizoium, eing cvs. Flmingo (tolernt) nd Cv. Coco Blnc (sensitive). Lef osmotic potentil (Ψo) decresed in stressed plnts in oth cultivrs. A minimum vlue of 2.3 MP ws reched in Cv. Flmingo plnts under mnnitol-induced osmotic stress (Figure 5 A). Ψo decresed in stressed nodules, reching 1.3 MP in Cv. Coco Blnc nd 1.7 MP in Cv. Flmingo (Figure 5 B). Therefore, Cv. Flmingo showed etter osmotic djustment response to osmotic stress oth in leves nd nodules [19]. Figure 5. Vrition of osmotic potentil (Ψo) in response to osmotic stress in leves (A) nd nodules (B) medited y 50 mm mnnitol. Vlues represents men ± SE (n=6) [19]. In control leves of oth cultivrs, RWC remined close to 80% (Figure 6 A). After 15 dys of osmotic tretment, RWC ws 65%in mnnitol-treted plnts of Cv. Flmingo, nd only
54 Responses of Orgnisms to Wter Stress 45% in Cv. Coco Blnc. These results indicte tht osmotic stress cused n importnt reduction in shoot wter supply. The sme trend ws oserved in nodules (Figure 6 B). Indeed, dt showed decresed nodule RWC in oth stressed cultivrs. This decrese ws higher in Cv. Coco Blnc treted nodules [19]. Figure 6. Effect of mnnitol-induced osmotic stress on reltive wter content (RWC) in Flmingo nd Coco lnc en cultivr leves (A) nd nodules (B). Vlues represents men ± SE (n=6) [19]. Mnnitol-induced wter deficit produced sustntil dehydrtion tht led to decresing Ψo (Figure 6). The decrese in Ψo is considered potentil mechnism of cellulr drought resistnce s it enles turgor mintennce nd growth continution [20]. Cv. Flmingo exhiited lower Ψo under osmotic tretment. It ws le to uptke more wter nd then grow more when exposed to decresed Ψo, thus it turned out to e etter drought tolernt cultivr thn Cv. Coco Blnc [21]. This my e ttriuted to mintennce of the lef nd nodule wter sttus under stressed conditions (Figure 5). Severl mechnisms could e involved in contriuting to wter retention. 4. Brdyrhizoium meliortes negtive effects in plnts exposed to drought The reltionship etween the wter sttus in the plnt nd N2 fixtion, minly under wter stress, nd the chnges in nodule morphology hve een studied in some temperte legumes [22]. However, tropicl legumes growing in rid regions, hve not received dequte ttention. Even where informtion is ville, the degree of wter stress in the plnts ws not clerly defined, which mkes it difficult to mke comprisons. The structurl sis for the differ
Tolernce to Drought in Leguminous Plnts Medited y Rhizoium nd Brdyrhizoium http://dx.doi.org/10.5772/54094 55 ence in sensivity of N2 fixtion in tropicl legumes, under wter stress, is not clerly understood [23]. Bsed in these prolems reported, Figueiredo et l. [24] investigted Vign unguicult plnts exposed to 3 inocultion forms (BR-2001, EI-6, nd control) comined with 6 different degrees of wter stress (-1.5, -2.0, -4.0, -6.0, -8.0, nd -10.0 kp). Wter deficit response in cowpe ppers to e directly relted to reduction in nodule mss (Tle 1), which my (fter severe stress, S6) hve ffected nodule structurl constituents. However, in moderte stress (S3) the impct on nodule wter content ws higher thn on the chnges in nodule mss [24]. (1) For S1 to S6 see Tle 1. *, **Significnt t the 0.05 nd 0.01 proility level. In ech column (lower letters) nd in ech line (cpitl letters), the mens followed y the sme letter do not differ sttisticlly (p< 0.05) from ech other, ccording to Tukey s test Tle 1. Nodule dry mtter (NDM) nd nodule wter content (NWC) in cowpe with (BR-2001 nd EI-6) nd without (C) Brdyrhizoium spp. inocultion t different degrees of wter stress [24]. 5. Interference positive on nitrogen compounds of plnts inoculted nd exposed to wter deficit Cowpe (Vign unguicult [L.] Wlp.) is leguminous with high protein content, lrge cpcity of fixtion of the tmospheric nitrogen (N 2 ) nd low requirements to soil fertility [25], eing frequently cultivted y frmers in Northern nd Northestern regions of the Brzil. This species constitutes the min susistence culture, eing the grin used s protein source in feeding [26]. Cowpe presents importnt gronomicl chrcteristics, such s rusticity nd precocity, esides eing considered plnt dpted to conditions of limited or insufficient wter vilility [27].
56 Responses of Orgnisms to Wter Stress Beneficil effects proportioned y the inocultion on growth prmeters s lef, stem nd root re lrgely explored nd well known in leguminous plnts [28-30], ut informtions more specific of this symiotic process on essentil compounds such s mino cids nd proteins re limited. Figueiredo et l. [24] report tht inocultion using Brdyrhizoium cn llevite the negtive consequences in Vign unguicult plnts induced to wter deficiency, ut study conducted y Serrj nd Sinclir [31] reveled tht wter supply presents repercussion on symiotic efficiency. Bsed on this overview, Bros et l. [32] crried out study iming to investigte if nitrogen compounds exercise influence on ccumultion of dry mtter in Vign unguicult plnts exposed to comined ction of inocultion nd wter deficit. The concentrtion of totl solule mino cids in plnts sujected to inocultion ws higher only in tolernt plnts, if compred with sme tretments of plnts non-inoculted (Figure 7 A). Wter deficit promoted significnt increse in this vrile to ll tretments. The tolernt cultivr presented lower chnges, in comprison with sme tretments in sensitive cultivr. Totl solule proteins of inoculted plnts presented higher vlues (Figure 7 B), when compred to sme tretments in non-inoculted plnts. Wter deficit cused significnt decrese in oth cultivrs, presenting higher vrition in sensitive plnts. For proline the inoculted plnts presented higher vlues, compring with sme tretments in non-inoculted plnts (Figure 7 C). The two cultivrs demonstrted higher vlues in wter deficit, when compred with respective controls. These results present greter vrition in tolernt plnts, if compred with sme tretments in sensitive plnts. Tolernt plnts sumitted to inocultion presented significnt increse in mino cids, nd these results re ttriuted to iologicl fixtion of nitrogen. The nitrogense enzyme promotes the nitrogen sorption in form of nitrogen gs (N 2 ) nd conversion to mmonium (NH 4+ ). In ddition, the higher formtion of mino cids proly is linked to increse in ctivity of enzymes glutmine synthetse (GS), eing your ctivity depending of ATP (denosine-5'-triphosphte), nd glutmte synthse (GOGAT). In ddition, the increse in mino cids of plnts exposed to inocultion is due to greter flux nd etter ssimiltion of nitrogen in form of mmonium, concomitntly with higher ctivity of GS nd GOGAT enzymes. Rmos et l. [33] evluting the responses in Glycine mx plnts under wter deficit nd inocultion of Brdyrhizoium jponicum oserved lso n increse in concentrtion of totl solule mino cids. The concentrtion of totl solule mino cids in plnts under wter deficit incresed in ll tretments. This increment occurred proly due to increse in ctivity of protese enzymes, responsile y rekdown of proteins iming to djust osmoticlly the plnt [34]. Similr results on increse in mino cids were otined to Cost et l. [35] investigting Vign unguicult plnts. Delfini et l. [36] evluting the responses of two Archis hypoge cultivrs sumitted to inocultion of Brdyrhizoium sp. showed significnt increse in mino cids.
Tolernce to Drought in Leguminous Plnts Medited y Rhizoium nd Brdyrhizoium http://dx.doi.org/10.5772/54094 57 Totl solule mino cids (µmol g DM -1 ) 200 150 100 50 wter deficit c cd control c cd A 0 Totl solule proteins (mg g DM -1 ) 12 10 8 6 4 2 c dc B 0 12 10 C Proline (µmol g DM -1 ) 8 6 4 d c d ce d ce 2 0 tolernt sensitive tolernt sensitive inoculted non-inoculted Figure 7. Totl solule mino cids (A), totl solule proteins (B), nd proline (C) in two contrsting Vign unguicult plnts under wter deficit nd sujected to inocultion. Mens followed y the sme letter re not significntly different y the Scott-Knott test t 5% of proility. The rs represent the men stndrd error [32]. The increse showed in totl solule proteins induced y inocultion suggests tht cteri ction resulted in increse in nitrogen supply through secondry route, tht is regulted y the nitrogense [37], ecuse in this study ws not verified increse in ctivity of the nitrte reductse fter inocultion. Hristozkov et l. [38] evluting the responses
58 Responses of Orgnisms to Wter Stress in Pisum stivum plnts under inocultion nd molydenum ppliction lso otined increse in protein levels. The decrese in protein levels promoted y the wter deficit is ssocited to decrese of the protein synthesis comined with increse of proteolytic enzymes, responsile y rekdown of solule proteins in plnts [39]. Cost [40] otined similr results studying Vign unguicult sujected to wter deficit, corroorting these results. The increse of proline levels provoked y the inocultion is proly linked to etter mino cids utiliztion such s glutmic cid nd rginine, eing the glutmic cid the precursor of the proline, while rginine cn suffer rection medited y enzyme clled of pyrrolline-5- croxylte reductse (P5CR) nd consequently to lierte proline [33]. Kohl et l. [41] lso oserved higher mounts of proline in Glycine mx plnts inoculted with Brdyrhizoium jponicum, contriuting with results of this study. The increse of proline in plnts under wter deficit is response to loss of cell turgescence [42]. Nogueir et l. [43] descrie tht the proline ccumultion hs een relted with drought tolernce in higher plnts, ctuting s osmoregultor gent with the ojective to keep wter in plnt tissue [44]. Similr ehvior ws descried y González et l. [45] working with Pisum stivum plnts under wter restriction. 6. Brdyrhizoium producing etter performnce on morphologicl prmeters Beneficil effects proportioned y the inocultion on growth prmeters re lrgely explored in crops s Phseolus vulgris nd Glycine mx, ut informtions on dry mtter ccumultion of Vign unguicult under wter deficit is limited. Bros et l. [32] conduced n experiment with 2 cultivrs (tolernt nd sensitive) comined with 2 wter regimes (wter deficit nd control), nd 2 inocultion forms (inoculted nd non-inoculted), totlizing 8 tretments. In shoot dry mtter the inocultion provoked increse, considering sme tretments in plnts non-inoculted (Figure 8 A). However, this increse only ws significnt in tolernt cultivr under inocultion nd irrigtion. Wter deficit occsioned significnt decrese in shoot dry mtter, with exception in tolernt cultivr non-inoculted. The tolernt plnts presented etter results, if compred with sensitive plnts independently of tretment (Figure 8 A). The inocultion provoked increse in plnt dry mtter, with exception in sensitive cultivr under irrigtion (Figure 8 B), compring to sme tretments in plnts non-inoculted. Wter deficiency induced decrese in plnt dry mtter for ll tretments, eing these significnt results when compred with control plnts. Independently of tretments ws showed tht sensitive cultivrs presented lower vlues, if compred to tolernt cultivrs.
Tolernce to Drought in Leguminous Plnts Medited y Rhizoium nd Brdyrhizoium http://dx.doi.org/10.5772/54094 59 30 25 wter deficit control A Shoot dry mtter (g) 20 15 10 5 cd c dc c 0 30 25 B Lef (numer) Plnt dry mtter (g) 20 15 10 5 0 50 40 30 20 10 e d dc c dc c ed cd C c 0 tolernt sensitive tolernt sensitive inoculted non-inoculted Figure 8. Shoot dry mtter (A), plnt dry mtter (B), numer of leves (C) in two contrsting Vign unguicult plnts under wter deficit nd sujected to inocultion. Mens followed y the sme letter re not significntly different y the Scott-Knott test t 5% of proility. The rs represent the men stndrd error [32]. For lef numer occurred increse fter inocultion, with exception in sensitive plnts under wter deficiency (Figure 8 C). Wter deficit proportioned decrese in vlues of lef numer, eing significnt in comprison with control plnts. In tolernt cultivr were otined higher vlues of lef numer, if compred with sensitive cultivr.
60 Responses of Orgnisms to Wter Stress Shoot dry mtter ws mximized fter the inocultion procedure, eing this fct linked to proly increse in nodule numer in root (dt not shown), s well s it proportioned higher sorption nd vilility of nitrogen to plnt [46-47]. Similr results linked to shoot dry mtter were found y Figueiredo et l. [24] in reserch with Vign unguicult plnts exposed to inocultion of Brdyrhizoium. The wter deficit reduced the production of shoot dry mtter, with these effects ssocited to negtive interference of wter deficiency on iochemicl processes s nitrte ssimiltion nd iologicl fixtion of nitrogen [35], modifying indirectly the prtitioning of photo-ssimiltes in root nd shoot, nd consequent decrese in ccumultion of shoot iomss [48]. Similr results were found y Mendes et l. [49] working with two Vign unguicult cultivrs sumitted to drought during two stges. The inocultion proportioned increse of totl dry mtter, nd this result must e linked to etter development nd efficiency of root system, in which presents higher nitrogen sorption using the nodultion process. I ddition, normlly the higher nitrogen fixtion will produce increse in mino cids nd lso proteins [36], nd it exercises influence on photossimiltes vilility. Similr responses were descried y Sssi et l. [50] investigting two Phseolus vulgris cultivrs sujected to inocultion with cteri of Rhizoium gender. Plnts under wter deficiency frequently hve the production of dry mtter reduced, eing this decrese relted to fct tht wter deficit ffects severl metolic processes such s sorption of wter nd nutrients, which re fundmentl to keep dequte growth nd development rtes. Nscimento [51] lso reported tht wter deprivtion ffects the osmotic mechnism, nd y consequence reduces the CO 2 supply, tht is essentil in photosynthetic process. Similr results were found y Leite nd Virgens Filho [52] studying Vign unguiculd plnts exposed to wter deficit. The increse in lef numer promoted y inocultion is occsioned y the higher numer of nodules in root, nd consequently due to the etter iologicl fixtion of nitrogen [53]. Arújo et l. [54] studying Vign unguicult nd Leucen leucocephl plnts lso reported n increse of this vrile, confirming the results otined in this work. The lower lef numer fter wter deficiency is cused y the process of lef scission, nd this fct occurs due to sustrte not to present wter nd nutrient sufficient to supply the plnt exigencies [4]. Correi nd Nogueir [55] otined similr results with Archis hypoge plnts under wter deficit. 7. Finl considertions This chpter ws structured with recent informtions on cpcity of Brdyrhizoium nd Rhizoium to medite tolernce in leguminous plnts sumitted to wter deficit, which it cn e used y students, techers, reserchers, scientists nd frmers. It reveled concepts, effects, nd results on wter deficiency nd your consequences on plnts, s well s explored sever
Tolernce to Drought in Leguminous Plnts Medited y Rhizoium nd Brdyrhizoium http://dx.doi.org/10.5772/54094 61 l possiilities linked to symiosis etween plnt-microorgnisms. Additionlly, it presented essentil compounds such s molydenum nd rections during process of iologicl fixtion of nitrogen. Also ws demonstrted the wter mintennce in lef nd nodule produced fter inocultion. Bsed in novel results, were relted interference positives on nitrogen compounds such s totl solule mino cids, proline, nd totl solule proteins. Other results prove the eneficil repercussion produced y inocultion with Brdyrhizoium on morphologicl prmeters. Acknowledgements This chpter hd finncil support from Conselho Ncionl de Pesquis (CNPq/Brzil) nd Coordenção de Aperfeiçomento de Pessol de Nível Superior (CAPES/Brzil) for Loto AKS. Author detils Alln Klynger d Silv Loto 1, Joquim Alenísio Gomes d Silveir 2, Roerto Cezr Loo d Cost 1 nd Cândido Ferreir de Oliveir Neto 1 1 Núcleo de Pesquis Vegetl Básic e Aplicd, Universidde Federl Rurl d Amzôni, Prgomins, Brzil 2 Lortório de Metolismo e Estresse de Plnts, Universidde Federl do Cerá, Fortlez, Brzil References [1] Rockström J, Flkenmrk M. Semirid crop production from hydrologicl perspective: Gp etween potentil nd ctul yield. Crit. Rev. Plnt Sci 2000;19 319-326. [2] Krmer PJ, Boyer JS. Wter reltions of plnt nd soils. Acdemic Press, New York. 1995. [3] Fontn DC, Berlto MA, Bergmschi H. Micrometeorologicl ltertions in soyens grown under different wter regimes. Pesquis Agropecuári Brsileir 1992;27 661-669. [4] Sntos RF, Crlesso R. Wter deficit nd morphologic nd physiologic ehvior of plnts. Revist Brsileir de Engenhri Agrícol e Amientl 1998;2 287-294.
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