ARBUSCULAR MYCORRHIZAL FUNGUS ENHANCES MAIZE (ZEA MAYS L.) GROWTH AND ORGANIC CARBON SEQUESTRATION IN WHEAT (TRITICUM AESTIVUM L.) STRAW-AMENDED SOILS

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1 Hu et l.: Arbusculr mychorrhizl fungus enhnces mize (Ze Mys L.) growth nd orgnic crbon sequestrtion in whet (Triticum Aestivum L.) strw-mended soils ARBUSCULAR MYCORRHIZAL FUNGUS ENHANCES MAIZE (ZEA MAYS L.) GROWTH AND ORGANIC CARBON SEQUESTRATION IN WHEAT (TRITICUM AESTIVUM L.) STRAW-AMENDED SOILS HU, J.L. 1,2,3 CUI, X.C. 1,2 LIN, X.G. 1,2 * DAI, J. 1,2 CHEN, R.R. 1,2 WANG, J.H. 1,2 WONG, M.H. 2,3 1 Stte Key Lbortory of Soil nd Sustinble Agriculture, Institute of Soil Science,Chinese Acdemy of Sciences Est Beijing Rod 71, Nnjing , P. R. Chin (phone: ; fx: ) 2 Joint Open Lbortory of Soil nd the Environment, Hong Kong Bptist University & Institute of Soil Science, Chinese Acdemy of Sciences Est Beijing Rod 71, Nnjing , P. R. Chin 3 Croucher Institute for Environmentl Sciences, Deprtment of Biology, Hong Kong Bptist University Kowloon Tong, Hong Kong SAR, P. R. Chin *Corresponding uthor e-mil: jlhu@isss.c.cn; xglin@isss.c.cn (Received 18 th Nov 2011; ccepted 22 nd July 2014) Abstrct. A pot experiment ws conducted to investigte the effects of whet (Triticum estivum L.) strws nd rbusculr mycorrhizl (AM) fungi on mize (Ze mys L.) growth nd orgnic crbon (C) sequestrtion in sterilized sndy lom soil. The experiment included three tretments: control, mendment with whet strws (S), nd mendment with whet strws plus inocultion with AM fungus Glomus cledonium (S+M). Compred with the control, the mendment of whet strws significntly (P<0.05) incresed soil lkline phosphtse (ALP) ctivity in mize seedling period (30 d) nd plnt phosphorus (P)-uptke nd shoot biomss t hrvest time (110 d), nd lso tended to increse soil orgnic C content s well s produced low but demiurgic crop yield t hrvest time. In whet strw-mended soils, G. cledonium inocultion significntly (P<0.05) decresed soil ph nd significntly (P<0.05) incresed plnt P-uptke s well s soil ALP ctivity in seedling period, nd tended to increse shoot biomss nd grin yield in seedling nd hrvest periods, respectively. Furthermore, G. cledonium inocultion tended to increse soil orgnic C content t hrvest time, which ws significntly (P<0.05) higher thn tht of the control, suggesting potentil ppliction of AM fungi in fcilitting crop production nd C sequestrtion in strw-returned fields. Keywords: lkline phosphtse, Glomus cledonium, P-uptke, rhizosphere cidifiction, strw return ISSN (Print) ISSN (Online) DOI: /eer/1203_ , ALÖKI Kft., Budpest, Hungry

2 Hu et l.: Arbusculr mychorrhizl fungus enhnces mize (Ze Mys L.) growth nd orgnic crbon sequestrtion in whet (Triticum Aestivum L.) strw-mended soils Introduction Soil orgnic crbon (C), n importnt component of terrestril ecosystems, is crucil regultor of C fluxes between the biosphere nd the tomosphere (Zhu nd Mille, 2003). Soil C storge depends on the blnce between C input nd output. Optimizing griculturl mngements cn increse soil C by incresing C input nd/or decresing C output (Lu et l., 2009). Orgnic mendments re used primrily to increse nutrient vilbility to plnts, nd re essentil for improving soil orgnic C content s well (Zhong et l., 2010). For instnce, strw, the dry stlks of cerel plnts, is n griculturl by-product fter the grin nd chff hve been removed. Since recycling of nutrients is the key to nutrient mngement in sustinble griculturl system, the mngement of strw in the field s crop residue for returning the orgnic mtter to the soils is significnt in griculturl production (Tuyen nd Tn, 2001). Although strw hs vlue from both the nutrients removed nd the orgnic mtter ddition it will return to the soil, the vlue of the orgnic mtter tht strw returns to the soil is much more difficult to clculte, nd it is the ctivity of microbe nd enzyme tht regultes C ccumultion vi minerliztion nd immobiliztion of plnt derived residues in the soil. Phosphorus (P) is one of the mjor essentil mcronutrients limiting plnt growth owing to its low biovilbility in soils (Feng et l., 2004), nd improving plnt cquisition of P from soil is n obvious lterntive to the mngement of those low P soils (Zhu et l., 2003). It is commonly known tht rbusculr mycorrhize (AMs) provide direct link between soil nd roots: AM fungi help plnts to cpture wter nd nutrients (notbly P) from the soil, nd in return, the plnt provides the fungus with reltively constnt nd direct ccess to crbohydrte (Smith nd Red, 2008), which re trnslocted from their source to root tissue nd on to fungl prtners. It is lso generlly ccepted tht AM fungi receive ll their crbohydrte from host plnt nd tht the ssocition could crete sink demnd for crbohydrte, which could result in 4 20% drin of C from the host plnt nd could indirectly influence C storge in soils (Grhm, 2000). Although strw return cn increse C input to soil from crop residues, uncertinties still remin bout the impcts of AMs on crop production nd soil orgnic C sequestrtion in strw-returned soils. Then, the objectives of this study were to investigte the effects of AMs on mize (Ze mys L.) growth nd orgnic C sequestrtion in whet (Triticum estivum L.) strws-mended soil, nd find out the min fctors tht influence these prmeters. Mterils nd methods Soil nd mycorrhizl inoculum preprtion A subsurfce soil smple ws collected from n rble griculturl lnd t Fengqiu County (35º00 N, 114º24 E), Henn Province, Chin. The ir-dried soil smple ws ground with wooden pestle, homogenized by sieving through 5 mm sieve, utoclved (121 C for 1 h on three successive dys), nd stored for the pot experiment. ISSN (Print) ISSN (Online) DOI: /eer/1203_ , ALÖKI Kft., Budpest, Hungry

3 Hu et l.: Arbusculr mychorrhizl fungus enhnces mize (Ze Mys L.) growth nd orgnic crbon sequestrtion in whet (Triticum Aestivum L.) strw-mended soils The soil is derived from lluvil sediments of the Yellow River nd clssified s quic inceptisol with the following properties: soil ph (soil: wter rtio of 1: 2.5) 8.6, 3.66 g kg -1 orgnic C, 0.35 g kg -1 totl N, 0.55 g kg -1 totl P, nd 22 g kg -1 totl K. Glomus cledonium (Nicol. & Gerd.) Trppe & Gerdemnn ws isolted from fluvo-quic soil in Hennn Province, Chin (Lio et l., 2003), nd deposited t the Institute of Soil Science, Chinese Acdemy of Sciences, Nnjing, Chin. The AM inocul were propgted on white clover (Trifolium repens L.) grown in n utoclved (121 C for 1 h on 3 successive dys) substrte for two successive propgtion cycles (4 months ech). The inocul were mixture of rhizospheric soil contining spores, hyphe, nd mycorrhizl root frgments, nd were ir-dried nd sieved (2 mm). At the sme time, the non-mycorrhizl inoculum ws lso prepred with the sme sterilized substrtum on which white clover ws cultivted under the sme conditions. Pot experiment Three tretments were estblished in the pot experiment: control, mendment with whet strw (S), nd mendment with whet strw plus inocultion with G. cledonium (S+M). Ech squre polyvinyl chloride (PVC) pot ( cm) contined 7.5 kg of soil, which ws mixed with 375 g of mycorrhizl/nonmycorrhizl inoculum, nd ech strw mendment tretment ws lso mixed with 37.5 g of whet strw (< 1 cm). Six mize seeds were sown into ech box, nd three seedling per box were left fter emergence. Plnts were grown in sunlit glsshouse with 30/22 C dy/night temperture, 40-60% reltive humidity nd 60-70% wter-holding cpcity. Pots were rndomly rrnged with four replictes per tretment. Two mize seedlings per pot ws hrvested fter growing for 30 dys, nd the other one ws hrvested fter growing for 110 dys. At ech smpling time, soil smples were lso collected. Mycorrhizl coloniztion, plnt nd soil nlysis Whet plnts were divided into roots, shoots, nd grins if possible. Fresh roots were used for mycorrhizl coloniztion ssessment by the grid-line intersect method (Giovnnetti nd Mosse, 1980) fter rinsing with tp wter, clering with 10% KOH nd stining with cid fuchsin (Phillips nd Hymn, 1970). Shoots nd grins were weighed fter oven drying t 70 C for 48 h, nd subsmples of dried nd ground shoots nd grins were tken for immedite nitric-perchloric cid digestion in HNO 3 (70%): HClO 4 (70%) mixture (6:1 v/v) (Zhu et l., 2003), followed by molybdenum-scorbic cid spectrophotometry (Hnson, 1950), to mesure tissue P concentrtion. To this end, the individul P-uptke by mize plnt (shoot plus grin) ws clculted. Soil smples were ir-dried nd homogenized by sieving through 2 mm mesh sieve. Soil ph ws determined with glss electrode using soil-to-wter rtio of 1:2.5. Soil lkline phosphtse ctivity ws determined by incubtion t 37 C with borte buffer (ph 9) ccording to the method of Tbtbi (1982), nd is given in units of mg p-nitrophenol produced g -1 soil 24 h -1. Then, the enzyme ctivity ws expressed on n oven-dried soil ISSN (Print) ISSN (Online) DOI: /eer/1203_ , ALÖKI Kft., Budpest, Hungry

4 Hu et l.: Arbusculr mychorrhizl fungus enhnces mize (Ze Mys L.) growth nd orgnic crbon sequestrtion in whet (Triticum Aestivum L.) strw-mended soils weight bsis by correcting for wter content in the soil (105 C, 24 h). To this end, soil orgnic C content ws determined by dichromte oxidtion (Mebius, 1960). Sttisticl nlysis The mens nd stndrd devitions of four replictes were computed. An nlysis of vrince ws crried out using the one-wy procedure with SPSS softwre while the comprison of men effects ws bsed on lest significnt difference (LSD) multiple-comprison tests (P < 0.05). Results Mycorrhizl coloniztion, shoot biomss, grin yield, nd P uptke of mize plnt Mycorrhiztion in mize roots ws shown in the G. cledonium-inoculted tretment (Tble 1), nd the coloniztion rtes were 31.7±3.1% nd 50.3±2.9% in seedling (30 d) nd hrvest (110 d) periods, respectively. The mendment of whet strw significntly (P < 0.05) incresed mize shoot biomss nd P-uptke t hrvest time but not in seedling period, nd produced low but demiurgic crop yield, which pproched zero in the control soils. In whet strw-mended soils, G. cledonium inocultion significntly (P < 0.05) incresed mize P-uptke nd tended to increse mize shoot biomss in seedling period s well, but hd no significnt effects on them t hrvest time, except for trend towrds higher grin yield. Tble 1. Mycorrhizl coloniztion, shoot biomss, grin yield, nd P-uptke of mize plnt under different tretments. Smpling Tretment Mycorrhizl Individul Individul Individul time coloniztion shoot biomss grin yield P-uptke (%) (g) (g) (mg) 30 d Control 0.56 (0.11) b 0.39 (0.24) b S 0.64 (0.14) b 0.59 (0.09) b S+M 31.7 (3.1) 0.84 (0.10) 1.05 (0.20) 110 d Control 5.10 (0.40) b 3.09 (0.36) b S 8.28 (1.38) 0.07 (0.12) (1.22) S+M 50.3 (2.9) 8.35 (2.55) 2.28 (2.44) (1.21) Control, without inocultion/mendment; S, mendment with whet strw; S+M, mendment with whet strw plus inocultion with AM fungi. Stndrd devitions re given in prentheses. Vlues within the sme column nd the sme cultivtion time not followed by the sme letter differ significntly (P < 0.05). ISSN (Print) ISSN (Online) DOI: /eer/1203_ , ALÖKI Kft., Budpest, Hungry

5 Hu et l.: Arbusculr mychorrhizl fungus enhnces mize (Ze Mys L.) growth nd orgnic crbon sequestrtion in whet (Triticum Aestivum L.) strw-mended soils Soil ph, lkline phosphtse ctivity, nd orgnic C content The mendment of whet strw significntly (P < 0.05) incresed soil ALP (Fig. 1b) ctivity in mize seedling period, but hd no significnt effects on soil ph (Fig. 1) nd orgnic C content (Fig. 1c) both in mize seedling nd hrvest periods, except for trend towrds higher orgnic C content t mize hrvest time. In whet strw-mended soils, G. cledonium inocultion significntly (P < 0.05) decresed soil ph nd significntly (P < 0.05) incresed soil ALP ctivity in seedling period, nd thereby tended to increse soil orgnic C content t mize hrvest time, which ws significntly (P < 0.05) higher thn tht of the control. 9.0 Control S S+M Soil ph (H 2 O) b 7.4 b Soil ALP ctivity (mg g -1 24h -1 ) d 110 d Control S S+M b c 30 d 110 d c Soil orgnic C (g kḡ 1 ) Control S S+M b b d 110 d Figure 1. Soil ph (), lkline phosphtse ctivity (b), nd orgnic C content (c) under different tretments. Control, without inocultion/mendment; S, mendment with whet strw; S+M, mendment with whet strw plus inocultion with AM fungi. Verticl T brs indicte stndrd devitions. Brs within the sme cultivtion time not topped by the sme letter indicte significnt difference in vlues (P < 0.05). ISSN (Print) ISSN (Online) DOI: /eer/1203_ , ALÖKI Kft., Budpest, Hungry

6 Hu et l.: Arbusculr mychorrhizl fungus enhnces mize (Ze Mys L.) growth nd orgnic crbon sequestrtion in whet (Triticum Aestivum L.) strw-mended soils Discussion Orgnic mendments ct s source of C nd other nutrients, which my improve soil orgnic C nd fvor plnt growth s well (Albich et l., 2000), owing to the enhnced plnt nutrition. However, mechnisms cusing incresed grin yield upon AM fungl inocultion in strw-mended soils seem to be due to the rhizosphere cidifiction nd enhnced ALP ctivity in the seedling period. On the one hnd, plnt roots lter rhizosphere soil ph by production or consumption of H + or by exudtion of orgnic cids, nd thereby induce chnges in nutrient vilbility (Li et l., 1991), nd such n effect is possibly enhnced by AM fungl inocultion (Bgo et l., 1996). On the other hnd, soil ALP ctivity is lso involved in the mycorrhiztion effect on plnt nutrition, since AM propgules themselves synthesize enzymes, such s ALP within the P-ccumulting vcuoles of AM hyphe (Dong nd Zho, 2004), nd mycorrhizl roots my relese more root exudtes contining enzymes thn non-mycorrhizl roots becuse of the lrger root system nd/or improved nutrition (Wng et l., 2006). It is noteworthy tht crops seedlings growing in strw-mended soils were still P limited in this study, nd the significnt increse of plnt P-uptke in the seedling period upon AM fungl inocultion plyed vitl role in deuteric grin filling. Strw mendment rised grin yield nd plnt biomss, which would lso increse biomss input into soil from crop roots (Lu et l., 2009). However, in ddition to fcilitting plnt uptke of minerl nutrients, AM fungi cn lso impct soil C dynmics directly by contributing to soil C inputs (Jkobsen nd Rosendhl, 1990), nd indirectly by modifying the decomposition of orgnic mterils (Tu et l., 2006). Although they re thought to be unble to decompose orgnic mterils directly due to lck of sprotrophic cpcity (Red nd Perez-Moreno, 2003), AM fungi my lter rhizosphere soil directly or indirectly through chnges in root exudtion ptterns or fungl exudtes (Lindermn, 1992), nd still be involved in decomposition processes. For exmple, AM fungi hve been found to proliferte in decomposing orgnic residues (St John et l., 1983), nd therefore, my enhnce whet strw decomposition by stimulting some enzyme ctivities. Consequently, the combined ppliction of whet strw nd AM fungi reveled the strongest influences on soil orgnic C content, which my be explined by the multitude of fctors tht my ct in different wys, such s direct C sources nd crop root exudtes. Although the results indicted the potentil ppliction of AM fungi in fcilitting C sequestrtion in strw-returned fields, the pplying strtegy still needs n in-depth study. In summry, whet strw incresed mize shoot biomss nd P-uptke s well s soil ALP ctivity, nd tended to increse soil orgnic C content s well s produced low but demiurgic crop yield t hrvest time. AM fungl inocultion gretly incresed mize grin yield nd soil orgnic C content in strw-mended soils, nd seemed to be due to enhnced rhizosphere cidifiction nd incresed nutrients (notbly P) uptke in the seedling period. The results indicted potentil ppliction of AM fungi in fcilitting crop growth nd C sequestrtion in strw-returned fields. ISSN (Print) ISSN (Online) DOI: /eer/1203_ , ALÖKI Kft., Budpest, Hungry

7 Hu et l.: Arbusculr mychorrhizl fungus enhnces mize (Ze Mys L.) growth nd orgnic crbon sequestrtion in whet (Triticum Aestivum L.) strw-mended soils Acknowledgements. This work ws supported by the Ntionl Nturl Science Foundtion (No ), the Ntionl Bsic Reserch Progrm (2011CB100505), nd the Specil Fund of Industril (Agriculture) Reserch for Public Welfre ( ) of Chin. REFERENCES [1] Albich R., Cnet R., Pomres F., Ingelmo F. (2000): Microbil biomss content nd enzymtic ctivities fter the ppliction of orgnic mendments to horticulturl soil. Bioresource Technology 75: [2] Bgo B., Vierheilig H., Piché Y., Azcon-Aguilr C. (1996): Nitrte depletion nd ph chnges induced by the extrrdicl mycelium of the rbusculr mycorrhizl fungus Glomus intrrdices grown in monoxenic culture. New Phytologist 133: [3] de Dtt S.K. (1981): Principles nd Prctices of Rice Production. John Wiley nd Sons, New York. [4] Dong C., Zho B. (2004): Arbusculr mycorrhizl formtion of crucifer lef mustrd induced by flvonoids pigenin nd didzein. Chinese Science Bulletin 49: [5] Feng K., Lu H.M., Sheng H.J., Wng X.L., Mo J. (2004): Effect of orgnic lignds on biologicl vilbility of inorgnic phosphorus in soils. Pedosphere 14: [6] Giovnnetti M., Mosse B. (1980): An evlution of techniques for mesuring vesiculr rbusculr mycorrhizl infection in roots. New Phytologist 84: [7] Grhm J.H. (2000): Assessing the cost of rbusculr mycorrhizl symbiosis in groecosystems. In: Podil G.K., Douds D.D. (eds.) Current Advnces in Mycorrhizl Reserch, The Americn Phytopthologicl Society, St. Pul, MN. [8] Hnson W.C. (1950): The photometric determintion of phosphorus in fertilisers using the phosphovndo molybdte complex. Journl of the Science of Food nd Agriculture 1: [9] Hu J., Lin X., Wng J., Cui X., Di J., Chu H., Zhng J. (2010): Arbusculr mycorrhizl fungus enhnces P-cquisition of whet (Triticum estivum L.) in sndy lom soil with long-term inorgnic fertiliztion regime. Applied Microbiology nd Biotechnology 88: [10] Jkobsen I., Rosendhl L. (1990): Crbon flow into soil nd externl hyphe from roots of mycorrhizl cucumber plnts. New Phytologist 115: [11] Li X.L., George E., Mrschner H. (1991): Phosphorus depletion nd ph decrese t the root-soil nd hyphe-soil interfces of VA mycorrhizl white clover fertilized with mmonium. New Phytologist 119: [12] Lio J.P., Lin X.G., Co Z.H., Shi Y.Q., Wong M.H. (2003): Interctions between rbusculr mycorrhize nd hevy metls under snd culture experiment. Chemosphere 50: [13] Lindermn R.G. (1992): Vesiculr rbusculr mycorrhize nd soil microbiot interctions. In: Bethlenflvy G.J., Lindermn R.G. (eds.) Mycorrhize in Sustinble Agriculture, Americn Society of Agronomy, Mdison, WI. [14] Lu F., Wng X., Hn B., Ouyng Z., Dun X., Zheng H., Mio H. (2009): Soil crbon ISSN (Print) ISSN (Online) DOI: /eer/1203_ , ALÖKI Kft., Budpest, Hungry

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