Effect of phosphorus nutrition and grain position within maize cob on grain phosphorus accumulation

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1 Instituto Nionl de Investigión y Tenologí Agrri y Alimentri (INIA) Spnish Journl of Agriulturl Reserh (2): ISSN: X eissn: SHORT COMMUNICATION OPEN ACCESS Effet of phosphorus nutrition nd grin position within mize o on grin phosphorus umultion Muhmmd Ndeem 1, 2, 3, *, Alin Mollier 1, 2, Alin Vives 1, 2, Loï Prud Homme 1, 2, Sylvie Niollet 1, 2 nd Sylvin Pellerin 1, 2 1 INRA, UMR 1391 ISPA, F Villenve d Ornon, Frne. 2 Bordeux Sienes Agro, UMR 1391 ISPA, F Grdignn, Frne. 3 Deprtment of Environmentl Sienes, COMSATS Institute of Informtion Tehnology, Vehri-61100, Pkistn Astrt Nutritionl sttus of grins my vry due to externl nutrient supply nd their position within prent mize o. Phosphorus (P) is the lest moile nutrient in the soil nd therefore newly growing seedlings re lrgely dependent on the stored grin P ontents whih re umulted during the rop mturity period. Ojetive of this study ws to ess the effets of different P pplitions nd grin positions on P nd dry mtter ontents in grins. Phosphorus pplition nd grin position hs signifint (p <0.05) effets on P ontents in grins wheres dry weight nd P ontent re highly orrelted. Grin weight nd P ontents deresed linerly from se to pil position possily due to flow of nutrients from se towrds pil position within o. Signifintly higher grin dry weight (0.35 ± 0.01 g) nd P ontents (962 ± 57 µg P) re reorded in high P pplition (92.50 kg h 1 ) rte on se position wheres minimum grin dry weight (0.14 ± 0.01 g) nd P ontents (219 ± 11 µg P) were reorded on pil grin position in low P pplition (5.60 kg h 1 ) rte. The results suggest tht for etter seedling P nutrition espeilly in soils of low inherent P, mize grins should e seleted from se or middle position where mximum dry weight nd P ontents re onentrted to support the seedlings to reh t growth t whih roots re ple of externl P uptke. Additionl key words: grin P; grin position; P ontents; P pplition. Phosphorus is the seond most importnt mronutrient for the plnt growth nd omprises pproximtely 0.2% of plnt s dry weight (Shhtmn et l., 1998). It plys ritil role in plnt metolism, ellulr energy trnsfer, respirtion nd photosynthesis (Glss et l., 1980; Oznne, 1980; Deleens et l., 1984; He & Burris, 1992; Usud & Shimogwr, 1993; Bewley, 1997; Bthellier et l., 2007). In seeds, P is stored primrily in the form of phytte (Lott et l., 1995; Prk et l., 2006; Ndeem et l., 2011). Phytte is onsidered s hving nti-nutrient hrteristis when onsumed y non ruminnt nimls (Roy et l., 1989). Phytte ontent of erels is highly orrelted with totl P (Lokhrt & Hurt, 1986), nd P onentrtions in seeds of given speies my vry with ultivr, inherent soil P sttus, nd limti onditions (Miller et l., 1980; Roy et l., 1990; Horvti & Blint, 1996). Seed vigor is n importnt determinnt of finl rop hrvest nd depends upon the sttus of the stored nutrients in seeds. Seed vigor dels with the ility of seed to grow rpidly nd uniformly under unfvorle onditions (Hr & Toriym, 1998). Therefore, the rte of initil seedling growth is often used s n index to quntify seed vigor (Seshu et l., 1988), whih implies tht the elertion of the rte of germintion is importnt to enhne the seedling estlishment. It is generlly reognized tht inrese of seed dry weight nd other nutrients improves the seed vigor (Seshu et l., 1988; Thomson & Bolger, 1993; Ros et l., 1997; Modi & Asnzi, 2008). Seed germintion, seed vigor nd seedling estlishment re therefore three *Corresponding uthor: muh.ndeem@gmil.om Reeived: Aepted: Arevitions used: HP (high phosphorus ontent); IP (intermedite phosphorus ontent); LP (low phosphorus ontent).

2 Short ommunition. Effets of P nutrition on mize grin P umultion 487 importnt hrters whih depend on the seedling ility to utilize stored seed reserves more effiiently (Bedi et l., 2009). Size nd shpe vry widely within th of grins/seeds. Spe etween grins on o dereses from the se to top, therefore ffeting the shpe of grins nd their nutritionl sttus s well (Pommel et l., 1995). In mize onstrints experiened y the grin position during o formtion suh s nutritionl defiienies my derese the size of the grin on o. The resulted grins re less in nutritive vlues (Pommel et l., 1995) to support the seedlings during erly ontogeny. Consequently it is possile tht grin position on o my ffet the P sttus of seeds tht it produes (Pommel et l., 1995). For optimum rop yield, plnts require dequte P from very erly growth stges (Brry & Miller, 1989; Brry et l., 1989; Grnt et l., 2001). The suess or filure of the erly growth stge of developing seedlings is diretly relted to suessful remoiliztion of stored grin P reserves tht were umulted throughout the ripening period (Gurdiol & Sutliffe, 1971; Le Deunff, 1975; Lwrene et l., 1990; Leonov et l., 2010; Ndeem et l., 2011, 2012). As the hydrolysis of grin P during germintion is mjor soure of inorgni P to support the growing seedlings during erly growth stges (Ndeem et l., 2011; 2012) therefore the present study ws onduted to investigte the role of P nutrition to the prent plnt on the dry weight nd P reserves umultion in mize grins for etter seletion of seeds. The irrigted mize rop (Ze mys L. v. DK 5783, Dekl, Monsnto Agrioltur SpA, Lodi, Itly) ws ultivted t the experimentl site of Pierroton, Bordeux, in southwest Frne (44 44' 30" N; 0 46' 59" W; lt. 55 m) during The mize rop ws fertilized with triple superphosphte (45% P 2 O 5 ). Phosphorus ws pplied in three P rtes s low P (LP: 5.6 kg h 1 ), intermedite P (IP: kg h 1 ) nd high P (HP: kg h 1 ) to get three levels in mize seeds. Soil nlysis showed tht ville Olsen-P in ploughed soil lyer ( m) ws 5.50 mg P kg 1, 9.90 mg P kg 1 nd mg P kg 1 of dry soil for LP, IP nd HP, respetively. At mturity, the mize os were hrvested from the field nd seprted in LP, IP nd HP fertiliztion tretments. Five mize os were seleted from eh tretment rndomly to selet homogenous grins from these os. The P tretments were not replited nd one plot orresponds to one P tretment. For eh P tretment, 5 plnts were rndomly seleted (with t lest 5 m etween eh plnt). One o per plnt ws hrvested for P nlysis. So the 5 smples orrespond to the 5 hrvested os per P tretment. For eh o, severl grins were olleted ording their position, ut they were onsidered s susmples. During the study, the mize grins were seprted into three tegories nmely pil, middle nd se grins s shown in Fig. 1. Eh position on o orresponds to 1/3 of the totl o length. Out of these three tegories, homogenous mize grins were seprted nd totl 27 grins were seleted nd pooled for the three grin positions respetively nd for the five os per P tretment. The seleted grins were dried in oven t 130 C for 24 h nd therefter these grins were pled in desitor. Grin dry weight ws mesured fter lyophiliztion for 24 h. Phosphorus ontents were determined in mize grins fter minerliztion (see Ndeem et l., 2011, for detils) y n dpttion of mlhite green olorimetri tehnique (Vn Veldhoven & Mnnerts, 1987). The resulted dt were nlyzed y ANOVA using the R environment for sttistil omputing nd grphis, version (R Development Core Tem, 2009). Mens were ompred using Tukey s test t the 0.05 risk level. Fig. 1 shows tht plots (ssumed to minly differ in phosphorus nutrition) long with grin position within mize o hs signifint (p < 0.05) effets on dry weight umultion in grins t mturity. At eh position on mize o, grin weight inreses linerly from low P pplition to higher P pplition nd se position of grins. Mximum grin dry weight (0.35 ± 0.01 g) t mturity ws reorded t se position followed y the middle grins position (0.31 ± 0.01 g) when high P ws pplied to rop. Grin weight dereses with the low P fertilizer pplitions nd minimum grin weight ws oserved t pil position (0.14 ± 0.01 g) when rop ws fertilized with low P inputs. No inrese in grin weight t middle position ws oserved (0.20 ± 0.02 g) in the intermedite-p plot ompred the high P plot with (Fig. 1). Signifint differenes were oserved mong plots nd grin positions on finl grin P ontents, s shown in Fig. 1. Higher P pplition used n inresed P ontents in mize grins followed y IP nd LP pplitions. Similrly grin P ontents deresed linerly from se to pil positions. Mximum grin P ontents (962 ± 57 µg) were

3 488 M. Ndeem et l. / Spn J Agri Res (2014) 12(2): ) ) ) Grin weight (g) Apil position Phosphorus ontents (μg) Apil position 0.4 Middle position 1,000 Middle position Grin weight (g) Phosphorus ontents (μg) Bse position Bse position Grin weight (g) Phosphorus ontents (μg) 1, Phosphorus pplitions 0 Phosphorus pplitions Figure 1. Seletion of mize grins on o () for phosphorus. Effet of different phosphorus pplitions nd grin position on the finl grin weight (in grms) () nd on the finl grin phosphorus ontents (µg) (). Dt re mens nd vertil rs indite ±SE for n = 5 replitions. Different letters indite signifint differene etween P level (Student s t-test, p < 0.05) oserved t se position followed y middle position (883 ± 33 µg) when rop reeived HP pplitions s shown in Fig. 1. Minimum grin P ontents were notied t pil grin position (219 ± 11 µg) followed y se (242 ± 22 µg) nd middle positions (295 ± 8 µg) when LP ws pplied during the growing sesons s indited in Fig. 1 (pil, se nd middle, respetively). Phosphorus eing omponent of living ells is essentil to sustins life (Rengel & Zhng, 2011; Six, 2011). Phosphorus umultes rpidly in grins during ripening long with other sustnes suh s lipids nd strh. In seeds, phytte is the min stored form of P (Roy et l., 1989; Rvindrn et l., 1994; Lott et l., 1995; Prk et l., 2006; Ndeem et l., 2011) long with inorgni nd ellulr P (Roy et l., 2001). Externl P supply hs signifint effets on P nd dry mtter umultion in grins phosphorus ontents re signifintly higher in grins treted with HP vilility s ompred to IP or LP. Phosphorus ontent in seeds depends oth on externl P vilility nd grin iomss umultion. A reltionship ws

4 Short ommunition. Effets of P nutrition on mize grin P umultion 489 IP: y = 2507x 130 R 2 = 0.88 HP: y = 2817x + 6 R 2 = 0.97 LP: y = 946x + 3 R 2 = 0.76 et l., 2012) espeilly on soils with low inherent P levels. Higher P ontents per seed were oserved in sl grins in HP tretment. The results suggest tht tking into onsidertion the position of grins in the o nd their P ontents ould e n importnt tool for seletion of grins produing vigorous seedling growth. The study ws rried out only in one growing seson, one plnting dte nd one lotion, onsequently the results presented need to e verified in wider rnge of field onditions. Aknowledgements Figure 2. Reltionship etween dry weight nd phosphorus umultion per grin when irrigted mize rop ws treted with three P nutritionl levels. Dt re mens nd vertil rs indite ±SE for n = 5 replitions. oserved etween grin dry mtter nd P ontent within eh P vilility level s shown in Fig. 2. As grin weight deresed from se towrds pil position, the umultion of P ws negtively ffeted s less P ws umulted in IP nd LP (Fig. 2). So, phosphorus ontent in grins is driven y the dry iomss umultion nd is dependent on the plnt P nutrition. Grin position within the mize o lso ffets the P ontents. Higher grin P ontents were oserved in grins loted t se position s ompred to middle or pil grins possily due to the flow of nutrients from se position towrds pil. This my lso e relted to the distne etween P soure in root zone nd grin position (P sink) resulting into lrger grins t se position ompred to middle or pil. Previous reserh showed tht size of mize grin n influene the nutritionl sttus (miro nd mronutrients) of grins (Clderini & Ortiz-Monsterio, 2003; Brloto et l., 2005; Hnley et l., 2007; Zhng et l., 2012) nd onsequently the growth of seedlings. Lrger grins with lrger nutritionl reserves re ple of produing vigorous plnts (Howe & Shupp, 1985; Ellison, 1987; Pommel et l., 1995; Ndeem et l., 2012). Seedling emergene ws more rpid in seeds with higher P onentrtions ompred to seeds with low P sttus (De Mro, 1990; Pommel et l., 1995). Crop grins hve lrge enough P reserves to sustin seedling P requirement during few weeks of erly growth (White & Venekls, 2012), however grins with higher indigenous P reserves n support seedling growth for longer period of time (Ndeem This study ws funded y projet grnt y the Higher Edution Commission (HEC), Pkistn nd enefited from finnil support from INRA (Ntionl Institute for Agriulturl Reserh), Frne. Authors thnk the INRA Experimentl Unit (UE570) of Cests- Pierroton nd the INRA long-term P fertiliztion network for providing mize grin from the long-term P fertiliztion experiment. Referenes Brry DAJ, Miller MH, Phosphorus nutritionlrequirement of mize seedlings for mximum yield. Agron J 81: Brry DAJ, Miller MH, Btes TE, Er lef nd seedling-p onentrtion nd dris indexes s inditors of P-nutrition for mize. Commun Soil Si Plnt Anl 20: Brloto C, Forget PM, Golderg DE, Seed mss, seedling size nd neotropil tree seedling estlishment. J Eol 93: Bthellier C, Bdek F, Couzi P, Hrsoët S, Muve C, Ghshghie J, Divergene in δ 13 C of drk respired CO 2 nd ulk orgni mtter ours during the trnsition etween heterotrophy nd utotrophy in Phseolus vulgris plnts. New Phytol 177: Bedi S, Meht S, Shrm S, Vshist KK, Nitrogen nutrition nd effiieny of seed reserve moiliztion during germintion in winter mize v. Bulnd. J New Seeds 10: Bewley JD, Seed germintion nd dormny. Plnt Cell 9: De Mro DG, Effet of seed weight, nd seed phosphorus nd nitrogen onentrtions on the erly growth of whet seedlings. Aust J Exp Agri 30: Deleens E, Gregory N, Bourdu R, Trnsition etween seed reserve use nd photosyntheti supply during development of mize seedlings. Plnt Si Lett 37:

5 490 M. Ndeem et l. / Spn J Agri Res (2014) 12(2): Clderini DF, Ortiz-Monsterio I, Grin position ffets grin mronutrient nd mironutrient onentrtions in whet. Crop Si 43: Ellison AM, Effet of seed dimorphism on the density dependent dynmis of experimentl popultion of Antiplex trinulris (Chenopodiee). Amer J Bot 74: Glss ADM, Beton JD, Bomke A, Role of P in plnt nutrition. Pro Western Cnd Phosphte Symposium. pp: Grnt CA, Flten DN, Tomsiewiz DJ, Shepprd SC, The importne of erly seson phosphorus nutrition. Cn J Plnt Si 81: Gurdiol JL, Sutliffe JF, Moiliztion of phosphorus in the otyledons of young seedlings of the grden pe (Pisum stivum L.). Ann Bot 35: Hnley ME, Cordier PK, My O, Kelly CK, Seed size nd seedling growth: differentil response of Austrlin nd British Fee to nutrient limittion. New Phytol 174: Hr Y, Toriym K, Seed nitrogen elertes the rtes of germintion, emergene, nd estlishment of rie plnts. Soil Si Plnt Nutr 44(3): He LS, Burris JS, Respirtion nd rohydrtemetolism during germintion of sh2 nd Sh2 sweet orn seed. Hortsiene 27: Horvti M, Blint L, Reltionship mong the phyti id nd protein ontent during mize grin mturtion. J Agron Crop Si 176: Howe HF, Shupp EW, Erly onsequene of seed dispersl for neotrophil tree (Viol surinmensis). Eol 66: Lwrene DM, Hlmer P, Bowles DJ, Moilistion of storge reserves during germintion nd erly seedling growth of sugr eet. Physiol Plnt 78: Le Deunff Y, L régultion hormonle de l germintion: le s des éréles. In: L germintion de semenes (Chusst R & Le Deunff Y, eds). Guthier Villrs, Pris. pp: Leonov S, Grimerg A, Mrttil S, Stymne S, Crlsson AS, Moiliztion of lipid reserves during germintion of ot (Aven stiv L.), erel rih in endosperm oil. J Exp Bot 61: Lokhrt HB, Hurt HD, 1986: Nutrition of ots. Am Asso of Cerel Chemists, St. Pul, MN, USA. pp: Lott JNA, Greenwood JS, Btten GD, Mehnisms nd regultion of minerl nutrient storge during seed development. Mrel Dekker, NY. pp: Miller GA, Youngs VL, Oplinger ES, Effet of ville soil-phosphorus nd environment on the phyti id onentrtion in ots. Cerel Chem 57: Modi AT, Asnzi NM, Seed performne of mize in response to phosphorus pplition nd growth temperture is relted to phytte-phosphorus ourrene. Crop Si 48: Ndeem M, Mollier A, Morel C, Vives A, Prud homme L, Pellerin S, 2011: Reltive ontriution of seed phosphorus reserves nd exogenous phosphorus uptke to mize (Ze mys L.) nutrition during erly growth stges. Plnt Soil 346: Ndeem M, Mollier A, Morel C, Vives A, Prud homme L, Pellerin S, Mize (Ze mys L.) endogenous seed phosphorus remoiliztion is not influened y exogenous phosphorus during germintion nd erly growth stges. Plnt Soil 357: Ndeem M, Mollier A, Morel C, Vives A, Prud homme L, Pellerin S, Seed phosphorus remoiliztion is not mjor limiting step for phosphorus nutrition during erly growth of mize. J Plnt Nutr Soil Si 175: Oznne PG, Phosphte nutrition of plnts - A generl tretise. ASA, Mdison, WI, USA. pp: Prk SH, Sung JK, Lee SY, Prk JH, Lee JY, Jng BC, Lee BH, Kim TW, Erly growth, rohydrte, nd phyti id ontents of germinting rie seeds under NCl stress. Koren J Crop Si 51: Pommel B, Goytino B, Bonhomme R, Effets of seed size, seed position on the prent o nd prentl o size on the lef re of mize under field onditions. Eur J Agron 4: R Development Core Tem, R: lnguge nd environment for sttistil omputing. R Foundtion for Sttistil Computing, Vienn, Austri. Aville in [Aessed 4 Fe 2012]. Roy V, Biohemistry nd genetis of phyti id synthesis. In: Inositol metolism in plnts (Morré DJ, Boss WF & Loewus FA, eds). Wiley Liss In, NY. pp: Roy V, Below FE, Dikinson DB, Altertion of mize kernel phyti id levels y reurrent seletion for protein nd oil. J Hered 80: Roy V, Young KA, Dorsh JA, Cook A, Genetis nd reeding of seed phosphorus nd phyti id. J Plnt Physiol 158: Rvindrn V, Rvindrn G, Sivlogn S, Totl nd phytte phosphorus ontents of vrious foods nd feedstuffs of plnt origin. Food Chem 56: Rengel Z, Zhng F, Phosphorus sustins life. Plnt Soil 349: 1-2. Ros C, Bell RW, White PF, Effet of seed phosphorus nd soil phosphorus pplitions on erly growth of rie (Oryz stiv L.) v IR66. Soil Si Plnt Nutr 43: Shhtmn DP, Reid RJ, Ayling SM, Phosphorus uptke y plnts: From soil to ell. Plnt Physiol 116: Seshu DV, Krishnswmy V, Siddique SB, Seed vigour in rie. In: Rie seed helth. Interntionl Rie Reserh Institute, Mnil. pp: Six J, Plnt nutrition for sustinle development nd glol helth. Plnt Soil 339: 1-2. Thomson CJ, Bolger TP, Effets of seed phosphorus onentrtion on the emergene nd growth of suterrnen lover (Trifolium suterrneum L.). Plnt Soil 156:

6 Short ommunition. Effets of P nutrition on mize grin P umultion 491 Usud H, Shimogwr K, Phosphte defiieny in mize. III. Chnges in mounts of surose-phosphte synthse during phosphte deprivtion. Plnt Physiol 102: Vn Veldhoven PP, Mnnerts GP, Inorgni nd orgni phosphte mesurements in the nnomolr rnge. Anl Biohem 161: White PJ, Venekls EJ, Nture nd nurture: the importne of seed phosphorus ontent. Plnt Soil 357: 1-8. Zhng Y, Zhng Y, Liu N, Su D, Xue Q, Stewrt BA, Wng Z, Effet of soure-sink mnipultion on umultion of mironutrients nd protein in whet grins. J Plnt Nutr Soil Si 175: