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Soil Biology & Biochemistry 36 (2004) 11911193 www.elsevier.



Fixed nitrogen in sustainable farming systems: a symposium examining factors inuencing the extent of biological nitrogen xation and its role in southern Australian agricultural systems. Setting the scene

The principal factor governing the potential productivity of southern Australian agricultural systems is the amount and timing with which water is made available to growing crops. Nitrogen supply is arguably the next most limiting factor with some of the best gains in wheat yield (40 kg ha21 y21) over the last 60 y occurring in areas where soil fertility has been enhanced through inclusion of legumes in rotations (Hamblin and Kyneur, 1993) and more recently by the application of fertiliser nitrogen (Angus, 2001). Increased amounts of plant available nitrogen in soils during the growing season subsequent to legume production attest to the positive effect that legumes can have on the availability of nitrogen to subsequent non-leguminous crops (Peoples et al., 1998; Peoples and Baldock, 2001). Such increases in availability arise from either or both of the following processes: (1) an enhanced release of inorganic nitrogen through the mineralisation of the relatively nitrogen rich legume residues in response to summer or early season rainfalls (Peoples et al., 1998) or (2) a nitrogen sparing effect where inorganic nitrogen not required by the legume crop accumulates within the soil root zone and remains available to subsequent crops (Chalk, 1998; Ahmad et al., 2001). All legumes (and their rhizobia) that have been signicantly exploited in Australian farming systems are exotic to the Australian ora. They have been introduced accidentally, or through plant introduction programs, a process that continues to this day. Early recordings of exotic legumes occurred in the late 1800s and by the 1930s early selections of sub-clover (Trifolium subterraneum), annual medic (Medicago spp.) and eld pea (Pisum sativum) were being recommended to farmers (Cocks et al., 1980; Gladstones and Collins, 1983; Hawthorne et al., 2003). A spectacular adoption of pastures followed in the period from 1940 and 1970 (Donald, 1965; Blyth and Menz, 1987). More legume options were subsequently provided to farmers with the introduction and development of several new genera of pulses including narrow leaf lupin (Lupinus angustifolius), faba bean (Vicia faba), lentil (Lens culinaris) and chickpea (Cicer arietinum). In the last decade there has

also been rapid increase in the use of the pasture legumes serradella (Ornithopus spp.) and biserrula (Biserrula pelecinus), especially on the lighter soils of Western Australia. Collectively, the pulses are now sown on nearly 2 M ha annually (OConnell, 2001), but still occupy only a fraction of the pasture area. Hill and Donald (1998) estimate that sub-clover and annual medics occur on 29 M ha and 24 M ha respectively, although some overlap in the distribution of the two genera undoubtedly occurs. Since none of the aforementioned legumes are able to nodulate with Australias indigenous microora, compatible strains of rhizobia have had to be provided to ensure good nodulation when the legume is sown in a soil for the rst time, or re-sown into a hostile soil (e.g. medics on acidic soils). High biological nitrogen xing capacity with the plant host is a priority in these rhizobial selection programs. More than 30 strains of rhizobia are currently produced as commercial inoculants in Australia by Bio-Care Technology Pty Ltd (www.bio-care.com.au). There has been a strong focus on the quality of inoculants. Accordingly, all batches are tested to ensure they meet standards of efcacy, rhizobial number and purity set by the Australian Legume Inoculants Research Unit (NSW Department of Agriculture). Despite having had the opportunity to carefully manage the introduction of many of legume and rhizobial genotypes released in Australia, the effectiveness of the symbioses that x atmospheric nitrogen into plant dry matter is often suboptimal. It can vary signicantly as a function of legume species, rhizobia ecology, soil properties and environmental characteristics. Estimates of the amount of biologically xed nitrogen for Australian pasture legumes based on the application of 15N natural abundance measurements to shoot dry matter range from 2 153 kg N ha21 for subterranean clover, 2 220 kg N ha21 for annual medics, 72 160 for vetch pastures, and 50 195 kg N ha21 for several other clover species (Peoples and Baldock, 2001). When these amounts of biologically xed nitrogen were expressed per unit of pasture shoot dry matter, values from 9 to 36 kg xed N t21 dry matter, with a mean of approximately 25 kg xed

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J.A. Baldock, R.A. Ballard / Soil Biology & Biochemistry 36 (2004) 11911193

N t21 dry matter, were obtained (Peoples and Baldock, 2001) indicating the highly variable efciency of biological nitrogen xation. Additionally, studies also suggest that 40 50% of the total nitrogen xed by pastures may reside below ground (Zebarth et al., 1991; McNeill et al., 1997; Jrgensen and Ledgard, 1997). As a result, total inputs and the variation in inputs of biologically xed nitrogen could be up to two times larger than values obtained by measuring shoots alone. Amounts of biologically xed nitrogen found in pulse legumes are even more variable than those noted for pastures legumes. For example, ranges for the amounts of nitrogen xed in the shoots of soybean, faba bean and lupin were found to be 0 450, 12 330 and 19 327 kg N ha21, respectively (Unkovich and Pate, 2000). These data demonstrate the potential benecial role that biological nitrogen xation by the legume/rhizobia symbiosis can have on the input of nitrogen to Australian agricultural systems. The data also demonstrate the large variation in the effectiveness of this symbiosis, suggesting the presence of gaps in our understanding of the factors controlling the efciency of biological N2 xation and/or our implementation of management strategies that allow this symbiosis to operate at its potential capacity. Contributions of biologically xed nitrogen can also be derived from a range of non-symbiotic heterotrophic organisms. The presence of nif genes combined with their genetic diversity allows these organisms to exist in a broad range of habitats. Although positive effects of adding heterotrophic organisms on the growth and N status of a variety of plants have been recorded, there are still no examples of where these organisms have been exploited or managed in the agricultural systems of southern Australia. Important areas of research that will improve our ability to maximise contributions of biologically xed N to agricultural systems include: (1) Improved methodologies for assessing in situ the nitrogen xation capacity of various rhizobia/host legume combinations. (2) Selection of legume and rhizobial genotypes that are better adapted to the various environmental niches experienced in Australia. (3) Overcoming the competition presented by ineffective strains of rhizobia. (4) Understanding the mechanisms that lead to the development diverse populations of rhizobia in soil. (5) Development of inoculation technologies that improve the survival of rhizobia and deliver higher numbers of viable cells to the rhizosphere. (6) Quantication of the inuence of improved legume/rhizobia combinations on the net nitrogen balance of rotations. (7) Understanding agronomic practices (e.g. herbicide application) that depress or enhance the symbiosis. (8) Understanding the social psychology needed to encourage legume adoption.

(9) Dening tangible opportunities to exploit nonsymbiotic N2 xers. We add a note of caution. While there are many benecial effects of growing legumes in farming systems, the asynchrony between the release of xed nitrogen and its consumption means that the potential for nitrate leaching exists in many soils. It has the potential to cause serious offsite impacts. The scope of the problem is likely to be large and probably not dissimilar, but less visible, than that of salinity. This being the case, strategies to utilise the enhanced soil nitrogen fertility generated by legumes probably deserves more focus. This issue of Soil Biology and Biochemistry presents a collection of some of the papers presented at the 13th Australian Nitrogen Fixation Conference in Adelaide, South Australia in September 2002 that address the identied areas of research pertaining to biological nitrogen xation.

Ahmad, T., Hafeez, F.Y., Mahmood, T., Malik, K.A., 2001. Residual effect of nitrogen xed by mungbean (Vigna radiata) and blackgram (Vigna mungo) on subsequent rice and wheat crops. Australian Journal of Experimental Agriculture 41, 245248. Angus, J.F., 2001. Nitrogen supply and demand in Australian agriculture. Australian Journal of Experimental Agriculture 41, 277288. Blyth, M.J., Menz, K.M., 1987. The role of economics in pasture research evaluation. In: Wheeler, J.L., Pearson, C.J., Robards, G.E. (Eds.), Temperate pastures: their production, use and management, Australian Wool Corporation/CSIRO, Victoria, pp. 586 589. Chalk, P.M., 1998. Dynamics of biologically xed N in legume-cereal rotations: a review. Australian Journal of Agricultural Research 49, 303 316. Cocks, P.S., Mathison, M.J., Crawford, E.J., 1980. From wild plants to pasture cultivars: annual medic and Subterranean clover in Southern Australia. In: Summereld, R.J., Bunting, A.H. (Eds.), Advances in Legume Science, Royal Botanic Gardens, London. Donald, C.M., 1965. The progress of Australian agriculture and the role of pastures in environmental change. Australian Journal of Science 27, 187 198. Gladstones, J.S., Collins, W.J., 1983. Subterranean clover as a naturalised plant in Australia. The Journal of the Australian Institute of Agricultural Science 49, 191202. Hamblin, A., Kyneur, G., 1993. Trends in wheat yields and soil fertility in Australia, Australian Government Publishing Service, Canberra. Hawthorne, W., Day, T., Pritchard, I., Ali, M., Sykes, J., Armstrong, E., Brouwer, J., Bretag, T., Davidson, J., 2003. Pea history calendar of events. In: Regan, K., Harries, M., Pritchard, I. (Eds.), Proceedings of the Field Pea Focus 2003, Pulse Australia, West Australia, pp. 52 62. Hill, M.J., Donald, G.E., 1998. Australian Temperate Pastures Database. Compact Disc, CSIRO Animal Production, Perth. Jrgensen, R.V., Ledgard, S.T., 1997. Contribution from stolons and roots to estimates of the total amount of N2 xed by white clover (Trifolium repens L). Annals of Botany 80, 641 648. McNeill, A.M., Zhu, C., Fillery, I.R.P., 1997. Use of in situ 15N-labelling to estimate the total below-ground nitrogen of pasture legumes in intact soil/plant systems. Australian Journal of Agricultural Research 48, 295 304.

J.A. Baldock, R.A. Ballard / Soil Biology & Biochemistry 36 (2004) 11911193 OConnell, L., 2001. In: OConnell, L., (Ed.), Australian Grain Yearbook, Australian Grain, Toowoomba, p. 4. Peoples, M.B., Baldock, J.A., 2001. Nitrogen dynamics of pastures: nitrogen xation inputs, the impact of legumes on soil nitrogen fertility, and the contributions of xed nitrogen to Australian farming systems. Australian Journal of Experimental Agriculture 41, 327346. Peoples, M.B., Gault, R.R., Scammell, G.J., Dear, B.S., Virgona, J., Sandral, G.A., Paul, J., Wolfe, E.C., Angus, J.F., 1998. The effect of pasture management on the contributions of xed N to the N-economy of ley-farming systems. Australian Journal of Agricultural Research 49, 459474. Unkovich, M., Pate, J.S., 2000. An appraisal of recent eld measurements of symbiotic N2 xation by annual legumes. Field Crops Research 65, 211228.


Zebarth, B.J., Alder, V., Sheard, R.W., 1991. In situ labelling of legume residues with a foliar application of a 15N-enriched urea solution. Communications in Soil Science and Plant Analysis 90, 103 108.

J.A. Baldock* CSIRO Land and Water, PMB 2, Glen Osmond, Australia SA 5064 E-mail address: jeff.baldock@csiro.au R.A. Ballard South Australian Research and Development Institute, P.O. Box 397, Adelaide, Australia SA 5001

* Corresponding author. Tel.: 618-8303-8537; fax: 618-8303-8550.