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REVIEW

Pastures and drought: a review of processes and implications for nitrogen and phosphorus cycling in grassland systems

Gina M. Lucci https://orcid.org/0000-0002-8402-088X
+ Author Affiliations
- Author Affiliations

AgResearch Ruakura Research Centre, Private Bag 3123 Hamilton 3240, New Zealand. Email: Gina.Lucci@agresearch.co.nz

Soil Research 57(2) 101-112 https://doi.org/10.1071/SR18079
Submitted: 21 March 2018  Accepted: 11 January 2019   Published: 11 February 2019

Abstract

The incidence and extent of drought is predicted to increase and therefore understanding the effects on the plant–soil system is important. The objective of this review is to report on the fundamental processes involved in the effects of drought on pasture, soil, and soil microorganisms in grassland systems and evaluate the consequences of drought to determine whether management decisions could mitigate the impact of drought. There are associations within the plant–soil system affecting the flows and cycling of nutrients. Drought conditions often create a flush of nitrogen, carbon, and phosphorus upon rewetting that is at risk of loss to the environment. Prediction of the flush magnitude is difficult because it is influenced by drought characteristics such as duration, soil temperature, degree of drying, and rate at which the rewetting occurs post-drought. Response to drought is also affected by the microbial community population and structure of the soil-related flora and fauna. Increasing pasture diversity and soil organic matter may help to mitigate the effects of drought in grassland systems. More research is needed that incorporates all the components of the plant–soil system to examine the net effects of drought on grassland systems. Better measures are also needed to estimate the consequences for future climate change on nutrient stocks and flows.

Additional keywords: carbon, climate change, drying and rewetting, mineralisation.


References

Appel T (1998) Non-biomass soil organic N – the substrate for N mineralization flushes following soil drying-rewetting and for organic N rendered CaCl2-extractable upon soil drying. Soil Biology & Biochemistry 30, 1445–1456.
Non-biomass soil organic N – the substrate for N mineralization flushes following soil drying-rewetting and for organic N rendered CaCl2-extractable upon soil drying.Crossref | GoogleScholarGoogle Scholar |

Augé RM (2001) Water relations, drought and vesicular-arbuscular mycorrhizal symbiosis. Mycorrhiza 11, 3–42.
Water relations, drought and vesicular-arbuscular mycorrhizal symbiosis.Crossref | GoogleScholarGoogle Scholar |

Austen EA, Sale PWG, Clark SG, Graetz B (2002) A survey of farmers’ attitudes, management strategies and use of weather and seasonal climate forecasts for coping with climate variability in the perennial pasture zone of south-east Australia. Australian Journal of Experimental Agriculture 42, 173–183.
A survey of farmers’ attitudes, management strategies and use of weather and seasonal climate forecasts for coping with climate variability in the perennial pasture zone of south-east Australia.Crossref | GoogleScholarGoogle Scholar |

Bakonyi G, Nagy P, Kovacs-Lang E, Kovacs E, Barabas S, Repasi V, Seres A (2007) Soil nematode community structure as affected by temperature and moisture in a temperate semiarid shrubland. Applied Soil Ecology 37, 31–40.
Soil nematode community structure as affected by temperature and moisture in a temperate semiarid shrubland.Crossref | GoogleScholarGoogle Scholar |

Baskaran S, Bolan N, Rahman A, Tillman R, MacGregor A (1994) Effect of drying of soils on the adsorption and leaching of phosphate and 2,4-dichlorophenoxyacetic acid. Soil Research 32, 491–502.
Effect of drying of soils on the adsorption and leaching of phosphate and 2,4-dichlorophenoxyacetic acid.Crossref | GoogleScholarGoogle Scholar |

Bhatti A, McClean CJ, Cresser MS (2013) Does plant uptake or low soil mineral-N production limit mineral-N losses to surface waters and groundwater from soils under grass in summer? Environmental Pollution 178, 128–134.
Does plant uptake or low soil mineral-N production limit mineral-N losses to surface waters and groundwater from soils under grass in summer?Crossref | GoogleScholarGoogle Scholar | 23562960PubMed |

Birch HF (1964) Mineralisation of plant nitrogen following alternate wet and dry conditions. Plant and Soil 20, 43–49.
Mineralisation of plant nitrogen following alternate wet and dry conditions.Crossref | GoogleScholarGoogle Scholar |

Blackwell MSA, Brookes PC, Fuente-Martinez Ndl, Murray PJ, Snars KE, Williams JK, Haygarth PM (2009) Effects of soil drying and rate of re-wetting on concentrations and forms of phosphorus in leachate. Biology and Fertility of Soils 45, 635–643.
Effects of soil drying and rate of re-wetting on concentrations and forms of phosphorus in leachate.Crossref | GoogleScholarGoogle Scholar |

Blackwell MSA, Carswell AM, Bol R (2013) Variations in concentrations of N and P forms in leachates from dried soils rewetted at different rates. Biology and Fertility of Soils 49, 79–87.
Variations in concentrations of N and P forms in leachates from dried soils rewetted at different rates.Crossref | GoogleScholarGoogle Scholar |

Borken W, Matzner E (2009) Reappraisal of drying and wetting effects on C and N mineralization and fluxes in soils. Global Change Biology 15, 808–824.
Reappraisal of drying and wetting effects on C and N mineralization and fluxes in soils.Crossref | GoogleScholarGoogle Scholar |

Boval M, Dixon RM (2012) The importance of grasslands for animal production and other functions: a review on management and methodological progress in the tropics. Animal 6, 748–762.
The importance of grasslands for animal production and other functions: a review on management and methodological progress in the tropics.Crossref | GoogleScholarGoogle Scholar | 22558923PubMed |

Brock JL (1988) Evaluation of New Zealand bred white clover cultivars under rotational grazing and set stocking with sheep. In ‘Proceedings of the New Zealand Grassland Association’. Vol. 49, pp. 203–206. Available at https://www.grassland.org.nz/publications/nzgrassland_publication_1081.pdf

Brock JL, Kim MC (1994) Influence of the stolon/soil surface interface and plant morphology on the survival of white clover during severe drought. In ‘Proceedings of the New Zealand Grassland Association’. Vol. 56, pp. 187–191. Available at http://www.grassland.org.nz/publications/nzgrassland_publication_767.pdf

Brock JL, Albrecht KA, Tilbrook JC, Hay MJM (2000) Morphology of white clover during development from seed to clonal populations in grazed pastures. The Journal of Agricultural Science 135, 103–111.
Morphology of white clover during development from seed to clonal populations in grazed pastures.Crossref | GoogleScholarGoogle Scholar |

Bünemann EK, Keller B, Hoop D, Jud K, Boivin P, Frossard E (2013) Increased availability of phosphorus after drying and rewetting of a grassland soil: processes and plant use. Plant and Soil 370, 511–526.
Increased availability of phosphorus after drying and rewetting of a grassland soil: processes and plant use.Crossref | GoogleScholarGoogle Scholar |

Butterly CR, Marschner P, McNeill AM, Baldock JA (2010) Rewetting CO2 pulses in Australian agricultural soils and the influence of soil properties. Biology and Fertility of Soils 46, 739–753.
Rewetting CO2 pulses in Australian agricultural soils and the influence of soil properties.Crossref | GoogleScholarGoogle Scholar |

Butterly C, McNeill A, Baldock J, Marschner P (2011a) Rapid changes in carbon and phosphorus after rewetting of dry soil. Biology and Fertility of Soils 47, 41–50.
Rapid changes in carbon and phosphorus after rewetting of dry soil.Crossref | GoogleScholarGoogle Scholar |

Butterly CR, McNeill AM, Baldock JA, Marschner P (2011b) Changes in water content of two agricultural soils does not alter labile P and C pools. Plant and Soil 348, 185
Changes in water content of two agricultural soils does not alter labile P and C pools.Crossref | GoogleScholarGoogle Scholar |

Cabrera ML, Kissel DE (1988) Potentially mineralizable nitrogen in disturbed and undisturbed soil samples. Soil Science Society of America Journal 52, 1010–1015.
Potentially mineralizable nitrogen in disturbed and undisturbed soil samples.Crossref | GoogleScholarGoogle Scholar |

Chaves MM, Maroco JP, Pereira JS (2003) Understanding plant responses to drought – from genes to the whole plant. Functional Plant Biology 30, 239–264.
Understanding plant responses to drought – from genes to the whole plant.Crossref | GoogleScholarGoogle Scholar |

Chepkwony CK, Haynes RJ, Swift RS, Harrison R (2001) Mineralization of soil organic P induced by drying and rewetting as a source of plant-available P in limed and unlimed samples of an acid soil. Plant and Soil 234, 83–90.
Mineralization of soil organic P induced by drying and rewetting as a source of plant-available P in limed and unlimed samples of an acid soil.Crossref | GoogleScholarGoogle Scholar |

Clegg CD, Lovell RDL, Hobbs PJ (2003) The impact of grassland management regime on the community structure of selected bacterial groups in soils. FEMS Microbiology Ecology 43, 263–270.
The impact of grassland management regime on the community structure of selected bacterial groups in soils.Crossref | GoogleScholarGoogle Scholar | 19719687PubMed |

Collis-George N, Greene R (1979) The effect of aggregate size on the infiltration behaviour of a slaking soil and its relevance to ponded irrigation. Australian Journal of Soil Research 17, 65–73.
The effect of aggregate size on the infiltration behaviour of a slaking soil and its relevance to ponded irrigation.Crossref | GoogleScholarGoogle Scholar |

Corleto A, Laude HM (1974) Evaluating growth potential after drought stress. Crop Science 14, 224–227.
Evaluating growth potential after drought stress.Crossref | GoogleScholarGoogle Scholar |

Cumberland GLB, Dyson CB, Honore EN (1971) A comparison of phosphorus responses in pasture and sheep, and effects of increased grazing pressure in a dry season. Proceedings of the New Zealand Society of Animal Production 31, 66–73.

De Boeck HJ, Dreesen FE, Janssens IA, Nijs I (2011) Whole-system responses of experimental plant communities to climate extremes imposed in different seasons. New Phytologist 189, 806–817.
Whole-system responses of experimental plant communities to climate extremes imposed in different seasons.Crossref | GoogleScholarGoogle Scholar | 21054412PubMed |

de Vries FT, Liiri ME, Bjornlund L, Setala HM, Christensen S, Bardgett RD (2012) Legacy effects of drought on plant growth and the soil food web. Oecologia 170, 821–833.
Legacy effects of drought on plant growth and the soil food web.Crossref | GoogleScholarGoogle Scholar | 22555357PubMed |

Dijkstra FA, He M, Johansen MP, Harrison JJ, Keitel C (2015) Plant and microbial uptake of nitrogen and phosphorus affected by drought using 15N and 32P tracers. Soil Biology & Biochemistry 82, 135–142.
Plant and microbial uptake of nitrogen and phosphorus affected by drought using 15N and 32P tracers.Crossref | GoogleScholarGoogle Scholar |

Dinka TM, Morgan CLS, McInnes KJ, Kishné AS, Daren Harmel R (2013) Shrink–swell behavior of soil across a Vertisol catena. Journal of Hydrology 476, 352–359.
Shrink–swell behavior of soil across a Vertisol catena.Crossref | GoogleScholarGoogle Scholar |

Doerr SH, Thomas AD (2000) The role of soil moisture in controlling water repellency: new evidence from forest soils in Portugal. Journal of Hydrology 231–232, 134–147.
The role of soil moisture in controlling water repellency: new evidence from forest soils in Portugal.Crossref | GoogleScholarGoogle Scholar |

Evans SE, Wallenstein MD (2012) Soil microbial community response to drying and rewetting stress: does historical precipitation regime matter? Biogeochemistry 109, 101–116.
Soil microbial community response to drying and rewetting stress: does historical precipitation regime matter?Crossref | GoogleScholarGoogle Scholar |

Fierer N, Schimel JP, Holden PA (2003) Influence of drying-rewetting frequency on soil bacterial community structure. Microbial Ecology 45, 63–71.
Influence of drying-rewetting frequency on soil bacterial community structure.Crossref | GoogleScholarGoogle Scholar | 12469245PubMed |

Forber KJ, Ockenden MC, Wearing C, Hollaway MJ, Falloon PD, Kahana R, Villamizar ML, Zhou JG, Withers PJA, Beven KJ, Collins AL, Evans R, Hiscock KM, Macleod CJA, Haygarth PM (2017) Determining the effect of drying time on phosphorus solubilization from three agricultural soils under climate change scenarios. Journal of Environmental Quality 46, 1131–1136.
Determining the effect of drying time on phosphorus solubilization from three agricultural soils under climate change scenarios.Crossref | GoogleScholarGoogle Scholar | 28991973PubMed |

Ford DJ, Cookson WR, Adams MA, Grierson PF (2007) Role of soil drying in nitrogen mineralization and microbial community function in semi-arid grasslands of north-west Australia. Soil Biology & Biochemistry 39, 1557–1569.
Role of soil drying in nitrogen mineralization and microbial community function in semi-arid grasslands of north-west Australia.Crossref | GoogleScholarGoogle Scholar |

Franzluebbers AJ (1999) Potential C and N mineralization and microbial biomass from intact and increasingly disturbed soils of varying texture. Soil Biology & Biochemistry 31, 1083–1090.
Potential C and N mineralization and microbial biomass from intact and increasingly disturbed soils of varying texture.Crossref | GoogleScholarGoogle Scholar |

Gales K (1979) Effects of water supply on partitioning of dry matter between roots and shoots in Lolium perenne. Journal of Applied Ecology 16, 863–877.
Effects of water supply on partitioning of dry matter between roots and shoots in Lolium perenne.Crossref | GoogleScholarGoogle Scholar |

Goebel MO, Bachmann J, Woche SK, Fischer WR (2005) Soil wettability, aggregate stability, and the decomposition of soil organic matter. Geoderma 128, 80–93.
Soil wettability, aggregate stability, and the decomposition of soil organic matter.Crossref | GoogleScholarGoogle Scholar |

Gordon H, Haygarth PM, Bardgett RD (2008) Drying and rewetting effects on soil microbial community composition and nutrient leaching. Soil Biology & Biochemistry 40, 302–311.
Drying and rewetting effects on soil microbial community composition and nutrient leaching.Crossref | GoogleScholarGoogle Scholar |

Griffiths RI, Whiteley AS, O’Donnell AG, Bailey MJ (2003) Physiological and community responses of established grassland bacterial populations to water stress. Applied and Environmental Microbiology 69, 6961–6968.
Physiological and community responses of established grassland bacterial populations to water stress.Crossref | GoogleScholarGoogle Scholar | 14660337PubMed |

Harrison-Kirk T, Beare MH, Meenken ED, Condron LM (2013) Soil organic matter and texture affect responses to dry/wet cycles: effects on carbon dioxide and nitrous oxide emissions. Soil Biology & Biochemistry 57, 43–55.
Soil organic matter and texture affect responses to dry/wet cycles: effects on carbon dioxide and nitrous oxide emissions.Crossref | GoogleScholarGoogle Scholar |

Hartmann AA, Barnard RL, Marhan S, Niklaus PA (2013) Effects of drought and N-fertilization on N cycling in two grassland soils. Oecologia 171, 705–717.
Effects of drought and N-fertilization on N cycling in two grassland soils.Crossref | GoogleScholarGoogle Scholar | 23297047PubMed |

Hofer D, Suter M, Buchmann N, Lüscher A (2017) Nitrogen status of functionally different forage species explains resistance to severe drought and post-drought overcompensation. Agriculture, Ecosystems & Environment 236, 312–322.
Nitrogen status of functionally different forage species explains resistance to severe drought and post-drought overcompensation.Crossref | GoogleScholarGoogle Scholar |

Hudson BD (1994) Soil organic matter and available water capacity. Journal of Soil and Water Conservation 49, 189–194.

Huygens D, Schouppe J, Roobroeck D, Alvarez M, Balocchi O, Valenzuela E, Pinochet D, Boeckx P (2011) Drying-rewetting effects on N cycling in grassland soils of varying microbial community composition and management intensity in south central Chile. Applied Soil Ecology 48, 270–279.
Drying-rewetting effects on N cycling in grassland soils of varying microbial community composition and management intensity in south central Chile.Crossref | GoogleScholarGoogle Scholar |

Iovieno P, Bååth E (2008) Effect of drying and rewetting on bacterial growth rates in soil. FEMS Microbiology Ecology 65, 400–407.
Effect of drying and rewetting on bacterial growth rates in soil.Crossref | GoogleScholarGoogle Scholar | 18547324PubMed |

IPCC (2007) Climate change 2007: the physical science basis. Contribution of Working Group I to the Fourth Assessment Report of the Intergovernmental Panel on Climate Change. (Eds S Solomon, D Qin, M Manning, Z Chen, M Marquis, KB Averyt, M Tignor, HL Miller) 996 pp. (Cambridge University Press: Cambridge, United Kingdom and New York, NY, USA)

Jin VL, Haney RL, Fay PA, Polley HW (2013) Soil type and moisture regime control microbial C and N mineralization in grassland soils more than atmospheric CO2-induced changes in litter quality. Soil Biology & Biochemistry 58, 172–180.
Soil type and moisture regime control microbial C and N mineralization in grassland soils more than atmospheric CO2-induced changes in litter quality.Crossref | GoogleScholarGoogle Scholar |

Jupp AP, Newman EI (1987) Phosphorus uptake from soil by Lolium perenne during and after severe drought. Journal of Applied Ecology 24, 979–990.
Phosphorus uptake from soil by Lolium perenne during and after severe drought.Crossref | GoogleScholarGoogle Scholar |

Kilpeläinen J, Barbero-López A, Vestberg M, Heiskanen J, Lehto T (2017) Does severe soil drought have after-effects on arbuscular and ectomycorrhizal root colonisation and plant nutrition? Plant and Soil 418, 377–386.
Does severe soil drought have after-effects on arbuscular and ectomycorrhizal root colonisation and plant nutrition?Crossref | GoogleScholarGoogle Scholar |

Kim I, Deurer M, Sivakumaran S, Huh KY, Green S, Clothier B (2011) The impact of soil carbon management and environmental conditions on N mineralization. Biology and Fertility of Soils 47, 709–714.
The impact of soil carbon management and environmental conditions on N mineralization.Crossref | GoogleScholarGoogle Scholar |

Knowles IM, Fraser TJ, Daly MJ (2003) White clover: loss in drought and subsequent recovery. In ‘Legumes for dryland pastures. Proceedings of a New Zealand Grassland Association (Inc.) Symposium’. 18–19 November, 2003, Lincoln University, Lincoln. pp. 37–41. Available at http://www.grassland.org.nz/publications/nzgrassland_publication_1675.pdf

Koch AL (1998) The biophysics of the gram-negative periplasmic space. Critical Reviews in Microbiology 24, 23–59.
The biophysics of the gram-negative periplasmic space.Crossref | GoogleScholarGoogle Scholar | 9561823PubMed |

Kreyling J, Dengler J, Walter J, Velev N, Ugurlu E, Sopotlieva D, Ransijn J, Picon-Cochard C, Nijs I, Hernandez P, Güler B, Gillhaussen P, De Boeck HJ, Bloor JMG, Berwaers S, Beierkuhnlein C, Arfin Khan MAS, Apostolova I, Altan Y, Zeiter M, Wellstein C, Sternberg M, Stampfli A, Campetella G, Bartha S, Bahn M, Jentsch A (2017) Species richness effects on grassland recovery from drought depend on community productivity in a multisite experiment. Ecology Letters 20, 1405–1413.
Species richness effects on grassland recovery from drought depend on community productivity in a multisite experiment.Crossref | GoogleScholarGoogle Scholar | 28941071PubMed |

Kubota T (1972) Aggregate-formation of allophanic soils: effect of drying on the dispersion of the soils. Soil Science and Plant Nutrition 18, 79–87.
Aggregate-formation of allophanic soils: effect of drying on the dispersion of the soils.Crossref | GoogleScholarGoogle Scholar |

Lau JA, Lennon JT (2012) Rapid responses of soil microorganisms improve plant fitness in novel environments. Proceedings of the National Academy of Sciences of the United States of America 109, 14058–14062.
Rapid responses of soil microorganisms improve plant fitness in novel environments.Crossref | GoogleScholarGoogle Scholar | 22891306PubMed |

Lehner B, Döll P, Alcamo J, Henrichs T, Kaspar F (2006) Estimating the impact of global change on flood and drought risks in Europe: a continental, integrated analysis. Climatic Change 75, 273–299.
Estimating the impact of global change on flood and drought risks in Europe: a continental, integrated analysis.Crossref | GoogleScholarGoogle Scholar |

Lemaire G, Denoix A (1987) Summer dry matter accumulation in Festuca arundinacea and Dactylis glomerata populations in Western France. II. Interaction between moisture level and nitrogen nutrition. Agronomie 7, 381–389.
Summer dry matter accumulation in Festuca arundinacea and Dactylis glomerata populations in Western France. II. Interaction between moisture level and nitrogen nutrition.Crossref | GoogleScholarGoogle Scholar |

Lucci G, Shepherd M, Vogeler I (2013) Effects of fertiliser nitrogen management on nitrate leaching risk from grazed dairy pasture in New Zealand. Proceedings of the New Zealand Grassland Association 75, 191–196.

Mariotte P, Vandenberghe C, Kardol P, Hagedorn F, Buttler A (2013) Subordinate plant species enhance community resistance against drought in semi-natural grasslands. Journal of Ecology 101, 763–773.
Subordinate plant species enhance community resistance against drought in semi-natural grasslands.Crossref | GoogleScholarGoogle Scholar |

Mariotte P, Robroek BJM, Jassey VEJ, Buttler A (2015) Subordinate plants mitigate drought effects on soil ecosystem processes by stimulating fungi. Functional Ecology 29, 1578–1586.
Subordinate plants mitigate drought effects on soil ecosystem processes by stimulating fungi.Crossref | GoogleScholarGoogle Scholar |

Mariotte P, Le Bayon R-C, Eisenhauer N, Guenat C, Buttler A (2016) Subordinate plant species moderate drought effects on earthworm communities in grasslands. Soil Biology & Biochemistry 96, 119–127.
Subordinate plant species moderate drought effects on earthworm communities in grasslands.Crossref | GoogleScholarGoogle Scholar |

Mazzarino MJ, Bertiller MB, Sain C, Satti P, Coronato F (1998) Soil nitrogen dynamics in northeastern Patagonia steppe under different precipitation regimes. Plant and Soil 202, 125–131.
Soil nitrogen dynamics in northeastern Patagonia steppe under different precipitation regimes.Crossref | GoogleScholarGoogle Scholar |

McDaniel JP, Barbarick KA, Stromberger ME, Cranshaw W (2013) Survivability of Aporrectodea caliginosa in response to drought stress in a Colorado soil. Soil Science Society of America Journal 77, 1667–1672.
Survivability of Aporrectodea caliginosa in response to drought stress in a Colorado soil.Crossref | GoogleScholarGoogle Scholar |

Mikha MM, Rice CW, Milliken GA (2005) Carbon and nitrogen mineralization as affected by drying and wetting cycles. Soil Biology & Biochemistry 37, 339–347.
Carbon and nitrogen mineralization as affected by drying and wetting cycles.Crossref | GoogleScholarGoogle Scholar |

Miller AE, Schimel JP, Meixner T, Sickman JO, Melack JM (2005) Episodic rewetting enhances carbon and nitrogen release from chaparral soils. Soil Biology & Biochemistry 37, 2195–2204.
Episodic rewetting enhances carbon and nitrogen release from chaparral soils.Crossref | GoogleScholarGoogle Scholar |

Mishra AK, Singh VP (2010) A review of drought concepts. Journal of Hydrology 391, 202–216.
A review of drought concepts.Crossref | GoogleScholarGoogle Scholar |

Murphy DV, Sparling GP, Fillery IRP, McNeill AM, Braunberger P (1998) Mineralisation of soil organic nitrogen and microbial respiration after simulated summer rainfall events in an agricultural soil. Australian Journal of Soil Research 36, 231–246.
Mineralisation of soil organic nitrogen and microbial respiration after simulated summer rainfall events in an agricultural soil.Crossref | GoogleScholarGoogle Scholar |

Navarro-García F, Casermeiro MÁ, Schimel JP (2012) When structure means conservation: effect of aggregate structure in controlling microbial responses to rewetting events. Soil Biology & Biochemistry 44, 1–8.
When structure means conservation: effect of aggregate structure in controlling microbial responses to rewetting events.Crossref | GoogleScholarGoogle Scholar |

Nichols SN, Hofmann RW, Williams WM (2015) Physiological drought resistance and accumulation of leaf phenolics in white clover interspecific hybrids. Environmental and Experimental Botany 119, 40–47.
Physiological drought resistance and accumulation of leaf phenolics in white clover interspecific hybrids.Crossref | GoogleScholarGoogle Scholar |

Orwin KH, Wardle DA (2005) Plant species composition effects on belowground properties and the resistance and resilience of the soil microflora to a drying disturbance. Plant and Soil 278, 205–221.
Plant species composition effects on belowground properties and the resistance and resilience of the soil microflora to a drying disturbance.Crossref | GoogleScholarGoogle Scholar |

Pang J, Yang J, Ward P, Siddique K, Lambers H, Tibbett M, Ryan M (2011) Contrasting responses to drought stress in herbaceous perennial legumes. Plant and Soil 348, 299–314.
Contrasting responses to drought stress in herbaceous perennial legumes.Crossref | GoogleScholarGoogle Scholar |

Puppi G, Bras A (1990) Nutrient and water relations of mycorrhizal white clover. Agriculture, Ecosystems & Environment 29, 317–322.
Nutrient and water relations of mycorrhizal white clover.Crossref | GoogleScholarGoogle Scholar |

Raveh A, Avnimelech Y (1978) The effect of drying on the colloidal properties and stability of humic compounds. Plant and Soil 50, 545–552.
The effect of drying on the colloidal properties and stability of humic compounds.Crossref | GoogleScholarGoogle Scholar |

Ritsema C, Dekker L (1996) Water repellency and its role in forming preferred flow paths in soils. Soil Research 34, 475–487.
Water repellency and its role in forming preferred flow paths in soils.Crossref | GoogleScholarGoogle Scholar |

Sardans J, Peñuelas J, Estiarte M (2008) Changes in soil enzymes related to C and N cycle and in soil C and N content under prolonged warming and drought in a Mediterranean shrubland. Applied Soil Ecology 39, 223–235.
Changes in soil enzymes related to C and N cycle and in soil C and N content under prolonged warming and drought in a Mediterranean shrubland.Crossref | GoogleScholarGoogle Scholar |

Schimel JP, Scott WJ, Killham K (1989) Changes in cytoplasmic carbon and nitrogen pools in a soil bacterium and a fungus in response to salt stress. Applied and Environmental Microbiology 55, 1635–1637.

Schimel J, Balser TC, Wallenstein M (2007) Microbial stress-response physiology and its implications for ecosystem function. Ecology 88, 1386–1394.
Microbial stress-response physiology and its implications for ecosystem function.Crossref | GoogleScholarGoogle Scholar | 17601131PubMed |

Schwalm CR, Anderegg WRL, Michalak AM, Fisher JB, Biondi F, Koch G, Litvak M, Ogle K, Shaw JD, Wolf A, Huntzinger DN, Schaefer K, Cook R, Wei Y, Fang Y, Hayes D, Huang M, Jain A, Tian H (2017) Global patterns of drought recovery. Nature 548, 202–205.
Global patterns of drought recovery.Crossref | GoogleScholarGoogle Scholar | 28796213PubMed |

Sheik CS, Beasley WH, Elshahed MS, Zhou XH, Luo YQ, Krumholz LR (2011) Effect of warming and drought on grassland microbial communities. The ISME Journal 5, 1692–1700.
Effect of warming and drought on grassland microbial communities.Crossref | GoogleScholarGoogle Scholar | 21451582PubMed |

Shepherd M, Lucci G, Vogeler I, Balvert S (2018) The effect of drought and nitrogen fertiliser addition on nitrate leaching risk from a pasture soil; an assessment from a field experiment and modelling. Journal of the Science of Food and Agriculture
The effect of drought and nitrogen fertiliser addition on nitrate leaching risk from a pasture soil; an assessment from a field experiment and modelling.Crossref | GoogleScholarGoogle Scholar | 30191570PubMed | in press.

Sierra J (1997) Temperature and soil moisture dependence of N mineralization in intact soil cores. Soil Biology & Biochemistry 29, 1557–1563.
Temperature and soil moisture dependence of N mineralization in intact soil cores.Crossref | GoogleScholarGoogle Scholar |

Singh DK, Sale PWG (2000) Growth and potential conductivity of white clover roots in dry soil with increasing phosphorus supply and defoliation frequency. Agronomy Journal 92, 868–874.
Growth and potential conductivity of white clover roots in dry soil with increasing phosphorus supply and defoliation frequency.Crossref | GoogleScholarGoogle Scholar |

Singh DK, Sale PWG (2002) Subsoil phosphorus concentration and tolerance of heavily grazed legume-based pastures to dry soil conditions. Wool Technology and Sheep Breeding 50, 499–502.

Singh DK, Sale PWG, McKenzie BM (1997) Water relations of white clover (Trifolium repens L.) in a drying soil, as a function of phosphorus supply and defoliation frequency. Australian Journal of Agricultural Research 48, 675–682.
Water relations of white clover (Trifolium repens L.) in a drying soil, as a function of phosphorus supply and defoliation frequency.Crossref | GoogleScholarGoogle Scholar |

Skinner RH, Comas LH (2010) Root distribution of temperate forage species subjected to water and nitrogen stress. Crop Science 50, 2178–2185.
Root distribution of temperate forage species subjected to water and nitrogen stress.Crossref | GoogleScholarGoogle Scholar |

Skinner RH, Gustine DL, Sanderson MA (2004) Growth, water relations, and nutritive value of pasture species mixtures under moisture stress. Crop Science 44, 1361–1369.
Growth, water relations, and nutritive value of pasture species mixtures under moisture stress.Crossref | GoogleScholarGoogle Scholar |

Smolander A, Barnette L, Kitunen V, Lumme I (2005) N and C transformations in long-term N-fertilized forest soils in response to seasonal drought. Applied Soil Ecology 29, 225–235.
N and C transformations in long-term N-fertilized forest soils in response to seasonal drought.Crossref | GoogleScholarGoogle Scholar |

Sparling G, Ross D (1988) Microbial contributions to the increased nitrogen mineralization after air-drying of soils. Plant and Soil 105, 163–167.
Microbial contributions to the increased nitrogen mineralization after air-drying of soils.Crossref | GoogleScholarGoogle Scholar |

Sparling G, Whale K, Ramsay A (1985) Quantifying the contribution from the soil microbial biomass to the extractable P levels of fresh and air-dried soils. Soil Research 23, 613–621.
Quantifying the contribution from the soil microbial biomass to the extractable P levels of fresh and air-dried soils.Crossref | GoogleScholarGoogle Scholar |

Steenwerth KL, Jackson LE, Calderon FJ, Scow KM, Rolston DE (2005) Response of microbial community composition and activity in agricultural and grassland soils after a simulated rainfall. Soil Biology & Biochemistry 37, 2249–2262.
Response of microbial community composition and activity in agricultural and grassland soils after a simulated rainfall.Crossref | GoogleScholarGoogle Scholar |

Styles D, Coxon C (2006) Laboratory drying of organic-matter rich soils: phosphorus solubility effects, influence of soil characteristics, and consequences for environmental interpretation. Geoderma 136, 120–135.
Laboratory drying of organic-matter rich soils: phosphorus solubility effects, influence of soil characteristics, and consequences for environmental interpretation.Crossref | GoogleScholarGoogle Scholar |

Szukics U, Abell GCJ, Hödl V, Mitter B, Sessitsch A, Hackl E, Zechmeister-Boltenstern S (2010) Nitrifiers and denitrifiers respond rapidly to changed moisture and increasing temperature in a pristine forest soil. FEMS Microbiology Ecology 72, 395–406.
Nitrifiers and denitrifiers respond rapidly to changed moisture and increasing temperature in a pristine forest soil.Crossref | GoogleScholarGoogle Scholar | 20298502PubMed |

Thanh Nguyen B, Marschner P (2005) Effect of drying and rewetting on phosphorus transformations in red brown soils with different soil organic matter content. Soil Biology & Biochemistry 37, 1573–1576.
Effect of drying and rewetting on phosphorus transformations in red brown soils with different soil organic matter content.Crossref | GoogleScholarGoogle Scholar |

Thomas H (1980) Terminology and definitions in studies of grassland plants. Grass and Forage Science 35, 13–23.
Terminology and definitions in studies of grassland plants.Crossref | GoogleScholarGoogle Scholar |

Turner BL, Haygarth PM (2001) Phosphorus solubilization in rewetted soils. Nature 411, 258
Phosphorus solubilization in rewetted soils.Crossref | GoogleScholarGoogle Scholar | 11357117PubMed |

Turner BL, Driessen JP, Haygarth PM, McKelvie ID (2003) Potential contribution of lysed bacterial cells to phosphorus solubilisation in two rewetted Australian pasture soils. Soil Biology & Biochemistry 35, 187–189.
Potential contribution of lysed bacterial cells to phosphorus solubilisation in two rewetted Australian pasture soils.Crossref | GoogleScholarGoogle Scholar |

Wakelin SA, Gregg AL, Simpson RJ, Li GD, Riley IT, McKay AC (2009) Pasture management clearly affects soil microbial community structure and N-cycling bacteria. Pedobiologia 52, 237–251.
Pasture management clearly affects soil microbial community structure and N-cycling bacteria.Crossref | GoogleScholarGoogle Scholar |

Wallis M, Horne D (1992) Soil water repellency. Advances in Soil Science 20, 91–146.
Soil water repellency.Crossref | GoogleScholarGoogle Scholar |

Wedderburn ME, Crush JR, Pengelly WJ, Walcroft JL (2010) Root growth patterns of perennial ryegrasses under well-watered and drought conditions. New Zealand Journal of Agricultural Research 53, 377–388.
Root growth patterns of perennial ryegrasses under well-watered and drought conditions.Crossref | GoogleScholarGoogle Scholar |

Whitehead DC (1995) ‘Grassland nitrogen.’ (CAB International: Wallingford)

Witteveen CFB, Visser J (1995) Polyol pools in Aspergillus niger. FEMS Microbiology Letters 134, 57–62.
Polyol pools in Aspergillus niger.Crossref | GoogleScholarGoogle Scholar |

Wood J (1999) Osmosensing by bacteria: signals and membrane-based sensors. Microbiology and Molecular Biology Reviews 1, 230–262.

Wu J, Brookes PC (2005) The proportional mineralisation of microbial biomass and organic matter caused by air-drying and rewetting of a grassland soil. Soil Biology & Biochemistry 37, 507–515.
The proportional mineralisation of microbial biomass and organic matter caused by air-drying and rewetting of a grassland soil.Crossref | GoogleScholarGoogle Scholar |

Xiang S-R, Doyle A, Holden PA, Schimel JP (2008) Drying and rewetting effects on C and N mineralization and microbial activity in surface and subsurface California grassland soils. Soil Biology & Biochemistry 40, 2281–2289.
Drying and rewetting effects on C and N mineralization and microbial activity in surface and subsurface California grassland soils.Crossref | GoogleScholarGoogle Scholar |

Zhao B, Chen J, Zhang J, Qin S (2010) Soil microbial biomass and activity response to repeated drying-rewetting cycles along a soil fertility gradient modified by long-term fertilization management practices. Geoderma 160, 218–224.
Soil microbial biomass and activity response to repeated drying-rewetting cycles along a soil fertility gradient modified by long-term fertilization management practices.Crossref | GoogleScholarGoogle Scholar |