参考文献:Annicchiarico P, Proietti S (2010) White clover selected for enhanced competitive ability widens the compatibility with grasses and favours the optimization of legume content and forage yield in mown clover-grass mixture. Grass Forage Sci 65:318–324. doi:10.1111/j.1365-2494.2010.00749.x CrossRef Annicchiarico P, Tomasoni C (2010) Optimizing legume content and forage yield of mown white clover-Italian ryegrass mixtures through nitrogen fertilization and grass row spacing. Grass Forage Sci 65:220–226. doi:10.1111/j.1365-2494.2010.00740.x CrossRef Badar R, Qureshi SA (2012) Comparative effect of Trichoderma hamatum and host-specific Rhizobium species on growth of Vigna mungo. J Appl Pharm Sci 2(4):128–132. doi:10.7324/JAPS.2012.2409 CrossRef Baker R, Elad Y, Chet I (1984) The controlled experiment in the scientific method with special emphasis on biological control. Phytopathology 74:1019–1021. doi:10.1094/Phyto-74-1019 CrossRef Borrero C, Trillas MI, Delgado A, Avilés M (2012) Effect of ammonium/nitrate ratio in nutrients solution on control of Fusarium wilt of tomato by Trichoderma asperellum T34. Plant Pathol 61:132–139. doi:10.1111/j.1365-3059.2011.02490.x CrossRef Carlsson G, Palmborg C, Jummpponen A, Scherer-Lorenzen M, Högberg P, Huss-Danell K (2009) N2 fixation in three perennial Trifolium species in experimental grasslands of varied plant species richness and composition. Plant Ecol 205:87–104. doi:10.1007/s11258-009-9600-9 CrossRef Chang YC, Chang YC, Baker R, Kleifeld O, Chet I (1986) Increased growth of plants in the presence of the biological-control agent Trichoderma harzianum. Plant Dis 70(2):145–148. doi:10.1094/PD-70-145 CrossRef Contreras-Cornejo HA, Macías-Rodríguez L, Cortés-Penagos C, López-Bucio J (2009) Trichoderma virens, a plant beneficial fungus, enhances biomass production and promotes lateral root growth through an auxin-dependent mechanism in Arabidopsis. Plant Physiol 149:1579–1592. doi:10.1104/pp. 108.130369 PubMed PubMedCentral CrossRef Crotty FV, Adi SM, Blackshaw RP, Murray PJ (2012) Using stable isotopes to differentiate trophic feeding channels within soil food webs. J Eukaryot Microbiol 59(6):520–526. doi:10.1111/j.1550-7408.2011.00608.x PubMed CrossRef de Santiago A, Quintero JM, Avilés M, Delgado A (2009) Effect of Trichoderma asperellum strain T34 on iron nutrition in white lupin. Soil Biol Biochem 41:2453–2459. doi:10.1016/j.soilbio.2009.07.033 CrossRef de Santiago A, Quintero JM, Avilés M, Delgado A (2011) Effect of Trichoderma asperellum strain T34 on iron, copper, manganese, and zinc uptake by wheat grown on a calcareous medium. Plant Soil 342:97–104. doi:10.1007/s11104-010-0670-1 CrossRef Frame J, Newbould P (1986) Agronomy of white clover. Adv Agron 40:1–88. doi:10.1016/S0065-2113(08)60280-1 CrossRef Frankow-Lindberg BE, Frame J (1996) Developments and challenges in sustainable grassland production. Grassland Sci Eur 1:337–345 Frey B, Schüepp H (1992) Transfer of symbiotically fixed nitrogen from berseem (Trifolium alexandrinum L.) to maize via vesicular-arbuscularmycorrhizal hyphae. New Phytol 122:447–454. doi:10.1111/j.1469-8137.1992.tb00072.x CrossRef Gierus M, Kleen J, Loges R, Taube F (2012) Forage legume species determine the nutritional quality of binary mixture with perennial ryegrass in the first production year. An Feed Sci Technol 172:150–161. doi:10.1016/j.anifeedsci.2011.12.026 CrossRef Harman GE (2000) Myths and dogmas of biocontrol. Changes in perceptions derived from research on Trichoderma harzianum T-22. Plant Dis 84(4):377–393. doi:10.1094/PDIS.2000.84.4.377 CrossRef Harman GE, Howell CR, Viterbo A, Chat I, Lorito M (2004) Trichoderma species - opportunistic, avirulent plant symbionts. Nature Rev Microbiol 2:43–56 Harrod TR, Hogan DV (2008) The soils of North Wyke and Rowden. Soil survey of England and Wales. Rothamsted Research, Okehampton, UK http://www.rothamsted.ac.uk/sites/default/files/SoilsNWRowden.pdf Accessed 22 July 2014 Harvey A, Parsons AJ, Rook AJ, Penning PD, Orr RJ (2000) Dietary preference of sheep for perennial ryegrass and white clover at contrasting sward surface heights. Grass Forage Sci 55:242–252CrossRef Haynes RJ (1988) Competitive aspects of the grass-legume association. Adv Agron 33:227–262CrossRef Høgh-Jensen H, Schjoerring JK (2010) Interactions between nitrogen, phosphorus and potassium determine growth and N2-fixation in white clover and ryegrass leys. Nutr Cycl Agroecosyst 87:327–338. doi:10.1007/s10705-009-9341-0 CrossRef Johansen A, Jensen ES (1996) Transfer of N and P from intact or decomposing roots of pea to barley interconnected by an arbuscular mycorrhizal fungus. Soil Biol Biochem 28:73–81. doi:10.1016/0038-0717(95)00117-4 CrossRef Lancashire PD, Bleiholder H, Van Den Boom T, Langeluddeke P, Stauss R, Weber E, Witzenberger A (1991) An uniform decimal code for growth stage crops and weeds. Ann Appl Biol 119:561–601. doi:10.1111/j.1744-7348.1991.tb04895.x CrossRef Melino VJ, Drew EA, Ballard RA, Reeve WG, Thomson G, White RG, O’Hara GW (2012) Identifying abnormalities in symbiotic development between Trifolium spp. and Rhizobium leguminosarum v. trifolii leading to sub-optimal and ineffective nodule phenotypes. Ann Bot 110(1559–15):55 Murray PJ, Clements RO (1998) Transfer of nitrogen between clover and wheat: effect of root herbivory. Eur J Soil Biol 34:25–30. doi:10.1016/S1164-5563(99)80003-X CrossRef Murray PJ, Hatch DJ (1994) Sitona weevils (Coleoptera, Curculionidae) as agents for rapid transfer of nitrogen from white clover (Trifolium repens L) to perennial ryegrass (Lolium perenne L) Ann. Appl Biol 125:29–33. doi:10.1111/j.1744-7348.1994.tb04943.x CrossRef Murray PJ, Clegg CD, Crotty FV, de la Fuente-Martínez N, Williams JK, Blackshaw RP (2009) Dissipation of bacterially derived C and N through the meso and macrofauna of a grassland soil. Soil Biol Biochem 41:1146–1150. doi:10.1016/j.soilbio.2009.02.021 CrossRef Naseby DC, Pascual JA, Lynch JM (2000) Effect of biocontrol strains of Trichoderma on plant growth, Pythium ultimum populations, soil microbial communities and soil enzyme activities. J Appl Microbiol 88:161–169. doi:10.1046/j.1365-2672.2000.00939.x PubMed CrossRef Nesheim L, Boller BC (1991) Nitrogen-fixation by white clover when competing with grasses at moderately low-temperatures. Plant Soil 133:47–56. doi:10.1007/BF00011898 CrossRef Nyfeler D, Huguenin-Elie O, Suter M, Frossard E, Lüscher A (2011) Grass-legume mixture can yield more nitrogen than legume pure stands due to mutual stimulation of nitrogen uptake from symbiotic and non-symbiotic source. Agric Ecosyst Environ 140:155–163. doi:10.1016/j.agee.2010.11.022 CrossRef Oke V, Long SR (1999) Bacteroid formation in the Rhizobium-legume symbiosis. Curr Opts Microbiol 2:641–646. doi:10.1016/S1369-5274(99)00035-1 CrossRef Parsons AJ, Chapman DF (2000) The principles of pasture growth and utilization. In: Hopkins A (ed) Grass, its production & utilization, 3rd edn. British Grassland Society by Blackwell Science, Oxford, pp 31–89 Paynel F, Murray PJ, Cliquet JB (2001) Root exudates: a pathway for short-term N transfer from clover and ryegrass. Plant Soil 229(2):235–243. doi:10.1023/A:1004877214831 CrossRef Peeters A (2009) Importance, evolution, environmental impact and future challenges of grasslands and grassland-based systems in Europe. Grassland Sci 55:113–125. doi:10.1111/j.1744-697X.2009.00154.x CrossRef Penning PD, Newman JA, Parsons AJ, Harvey A, Orr RJ (1997) Diet preferences of adult sheep and goats grazing ryegrass and white clover. Small Rumin Res 24:175–184. doi:10.1016/S0921-4488(96)00930-3 CrossRef Pérez-de-Luque A, Rubiales D, Cubero JI, Press MC, Scholes J, Yoneyama K, Takeuchi Y, Plakhine D, Joel DM (2005) Interaction between Orobanche crenata and its host legumes: unsuccessful haustorial penetration and necrosis of the developing parasite. Ann Bot 95:935–942. doi:10.1093/aob/mci105 PubMed PubMedCentral CrossRef Rassmusen J, Eriksen J, Jensen ES, Esbensen KH, Høgh-Jensen H (2007) In situ carbon and nitrogen dynamics in ryegrass-clover mixture: transfer, deposition and leaching. Soil Biol Biochem 39:804–815. doi:10.1016/j.soilbio.2006.10.004 CrossRef Rassmusen J, Søegaard K, Pirhofer-Walzi K, Eriksen J (2012) N2-fixation and residual N effect of four legume species and four companion grass species. Eur J Agron 36:66–74. doi:10.1016/j.eja.2011.09.003 CrossRef Romão-Dumaresq AS, de Araújo WL, Talbot NJ, Talbot NJ (2012) RNA interference of endochitinases in the sugarcane endophyte Trichoderma virens 223 reduces its fitness as biocontrol agent of pineapple disease. PLoS ONE 7:e47888. doi:10.1371/journal.pone.0047888 PubMed PubMedCentral CrossRef Rook AJ, Harvey A, Parsons AJ, Penning PD, Orr RJ (2002) Effect of long-term changes in relative resource availability on dietary preference of grazing sheep for perennial ryegrass and white clover. Grass Forage Sci 57:54–60. doi:10.1046/j.1365-2494.2002.00301.x CrossRef Ruzin SE (1999) Plant microtechniques and microscopy. Oxford University Press, New York, USA Ryder LS, Harris BD, Soanes DM, Kershaw MJ, Talbot NJ, Thornton CR (2012) Saprotrophic competitiveness and biocontrol fitness of a genetically modified strain of the plant-growth-promoting fungus Trichoderma hamatum GD12. Microbiology 158:84–97. doi:10.1099/mic.0.051854-0 PubMed CrossRef Shaban WI, El-Bramawy MA (2011) Impact of dual inoculation with Rhizobium and Trichoderma on damping off, root rot diseases and plant growth parameters of some legumes field crop under greenhouse conditions. International Res. J Agric Sci Soil Sci 1(3):98–108 Shoresh M, Harman GE, Mastouri F (2010) Induced systemic resistance and plant responses to fungal biocontrol agents. Ann Rev Phytopathol 48:21–43CrossRef Sreenivasaprasad S, Sharada K, Brown A, Mills P (1996) PCR-based detection of Colletotrichum acutatum on strawberry. Plant Pathol 45:650–655CrossRef Studholme D, Harris B, Le Cocq K, Winsbury R, Perera V, Ryder L, Ward JL, Beale MH, Thornton CR, Grant M (2013) Investigating the beneficial traits of Trichoderma hamatum GD12 for sustainable agriculture – insights from genomics. Front Plant Sci 4:1–13. doi:10.3389/fpls.2013.00258 CrossRef Talbot NJ, Salch YP, Ma M, Hamer JE (1993) Karyotypic variation within clonal lineages of the rice blast fungus, Magnaporthe grisea. Appl Environ Microbiol 59:585–593PubMed PubMedCentral Vance CP, Johnson LEB, Hardarson G (1980) Histological comparisons of plant and Rhizobium induced ineffective nodules in alfalfa. Physiol Plant Pathol 17:167–173. doi:10.1016/0048-4059(80)90049-1 CrossRef Vargas WA, Mandawe JC, Kenerley CM (2009) Plant-derived sucrose is a key element in the symbiotic association between Trichoderma virens and maize plants. Plant Physiol 151:792–808. doi:10.1104/pp. 109.141291 PubMed PubMedCentral CrossRef Vasse J, De Billy F, Camut S, Truchet G (1990) Correlation between ultrastructural differentiation of bacteroids and nitrogen fixation in alfalfa nodules. J Bacteriol 8:4295–4306 Viterbo A, Horwitz BA (2010) Mycoparasitism. In: Borkovich KA, Ebbole DJ (eds) Cellular and biology of filamentous fungi. American Society for Microbiology, Washington, pp 676–693CrossRef Walker JA, King JR (2010) Does relative time of emergence affect stand composition and yield in a grass-legume mixture? Kura clover (Trifolium ambiguum)-meadow bromegrass (Bromus biebersteinii) and Kura clover-orchardgrass (Dactylis glomerata) mixtures. Grass Forage Sci 65:237–347. doi:10.1111/j.1365-2494.2010.00742.x CrossRef White TJ, Bruns T, Lee S, Taylor J (1990) Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. PCR Protocols: Guide Methods Appl 18:315–322 Woledge J, Davidson K, Dennis WD (1992) Growth and photosynthesis of tall and short cultivars of white clover with tall and short grasses. Grass Forage Sci 47:230–238. doi:10.1111/j.1365-2494.1992.tb02267.x CrossRef Yedidia I, Srivastva AK, Kapulnik Y, Chet I (2001) Effect of Trichoderma harzianum on microelement concentrations and increased growth of cucumber plants. Plant Soil 235:235–242. doi:10.1023/A:1011990013955 CrossRef
作者单位:C. Alcántara (1) C. R. Thornton (3) A. Pérez-de-Luque (2) K. Le Cocq (4) V. Pedraza (1) P. J. Murray (4)
1. Área de Producción Agraria, IFAPA Centro Alameda del Obispo, Junta de Andalucía, Avda. Menéndez Pidal s/n, 14080, Córdoba, Spain 3. Biosciences, University of Exeter, Geoffrey Pope Building, Stocker Road, Exeter, EX4 4QD, UK 2. Área de Mejora y Biotecnología, IFAPA Centro Alameda del Obispo, Junta de Andalucía, Avda. Menéndez Pidal s/n, 14080, Córdoba, Spain 4. Sustainable Soils and Grassland Systems Department, Rothamsted Research, North Wyke, Okehampton, EX20 2SB, UK
Aim A principal goal of grassland management is to minimize the use of artificial fertilizers by maximising the productivity of nitrogen-fixing leguminous plants such as clovers. The objective of this study was to investigate whether a plant-growth-promoting strain of the free-living rhizosphere fungus Trichoderma hamatum (GD12) could be used as a natural and sustainable means of enhancing the competitiveness of white clover (Trifolium repens) while allowing increased productivity of both clover and ryegrass (Lolium perenne) in mixed species systems.