Advertisement

Plant and Soil

, Volume 371, Issue 1–2, pp 435–446 | Cite as

Biomass of extramatrical ectomycorrhizal mycelium and fine roots in a young Norway spruce stand — a study using ingrowth bags with different substrates

  • Jonny Neumann
  • Egbert Matzner
Regular Article

Abstract

Background and aims

The partitioning of below ground carbon inputs into roots and extramatrical ectomycorrhizal mycelium (ECM) is crucial for the C cycle in forest soils. Here we studied simultaneously the newly grown biomass of ECM and fine roots in a young Norway spruce stand.

Methods

Ingrowth mesh bags of 16 cm diameter and 12 cm height were placed in the upper soil and left for 12 to 16 months. The 2 mm mesh size allowed the ingrowth of fungal hyphae and roots whereas a 45 μm mesh size allowed only the ingrowth of hyphae. The mesh bags were filled with either EA horizon soil, pure quartz sand or crushed granite. Controls without any ingrowth were established for each substrate by solid tubes (2010) and by 1 μm mesh bags (2011). The fungal biomass in the substrates was estimated by the PLFA 18:2ω6,9 and ECM biomass was calculated as difference between fungal biomass in mesh bags and controls.

Results

The maximum ECM biomass was 438 kg ha−1 in October 2010 in 2 mm mesh bags with EA substrate, and the minimum was close to zero in 2011 in 45 μm mesh bags with quartz sand. The high P content of the crushed granite did not influence the ECM biomass. Fine root biomass reached a maximum of 2,343 kg ha−1 in October 2010 in mesh bags with quartz sand after 16 months exposure. In quartz sand and crushed granite, ECM biomass correlated positively with fine root biomass and the number of root tips, and negatively with specific root length.

Conclusion

The ratio of ECM biomass/fine root biomass in October ranged from 0.1 to 0.3 in quartz sand and crushed granite, but from 0.7 to 1.8 in the EA substrate. The results for the EA substrate suggest a large C flux to ECM under field conditions.

Keywords

Biomass Ectomycorrhizal mycelium Fine roots Ingrowth bags Substrate quality Norway spruce 

Notes

Acknowledgments

We thank the Department of Micrometeorology of the University Bayreuth of Prof. Thomas Foken for meteorological data, the Department of Analytic Chemistry of the Bayreuth Center of Ecology and Environmental Research (BAYCEER) for elemental analysis, Uwe Hell for support at the field site and Petra Eckert for laboratory work. Annalena Fischer provided the data on standing fine root biomass in the forest floor. We are thankful for the generally positive and constructive comments made by two anonymous reviewers.

References

  1. Agerer R (2001) Exploration types of ectomycorrhizae. A proposal to classify ectomycorrhizal mycelial systems according to their patterns of differentiation and putative ecological importance. Mycorrhiza 11:107–114CrossRefGoogle Scholar
  2. Allen MF (2007) Mycorrhizal fungi: highways for water and nutrients in arid soils. Vadose Zone J 6:291–297CrossRefGoogle Scholar
  3. Alt D, Peters I (1992) Analyis of macro- and trace-elements in Horticultural substrates by the CaCl2/DTPA Method. 1. phosphorus, potassium and nitrogen. Agribiol Res 45:204–214Google Scholar
  4. Antubus RK, Sinsabaugh RL (1993) The extraction end quantification of ergosterol from ectomycorrhizal fungi and roots. Mycorrhiza 3:137–344CrossRefGoogle Scholar
  5. Arnebrant K (1994) Nitrogen amendments reduce the growth of extramatrical ectomycorrhizal mycelium. Mycorrhiza 5:7–15CrossRefGoogle Scholar
  6. Borken W, Kossmann G, Matzner E (2007) Biomass, morphology and nutrient contents of fine roots in four Norway spruce stands. Plant Soil 292:79–93CrossRefGoogle Scholar
  7. Brunner I, Bakker MR, Björk RG, Hirano Y, Lukac M, Aranda X et al (2012) Fine-root turnover rates of European forests revisited: an analysis of data from sequential coring and ingrowth cores. Plant Soil 362:357–372Google Scholar
  8. Chapin FS, McFarland J, McGuire AD, Euskirchen ES, Ruess RW, Kielland K (2009) The changing global carbon cycle: linking plant-soil carbon dynamics to global consequences. J Ecol 97:840–850CrossRefGoogle Scholar
  9. R Core Team (2012) R. A language and enviroment for statistical computingGoogle Scholar
  10. Ekblad A, Nasholm T (1996) Determination of chitin in fungi and mycorrhizal roots by an improved HPLC analysis of glucosamine. Plant Soil 178:29–35CrossRefGoogle Scholar
  11. Ekblad A, Wallander H, Carlsson R, Huss-Danell K (1995) Fungal biomass in roots and extramatrical mycelium in relation to macronutrients and plant biomass of ectomycorrhizal Pinus sylvestris and Alnus incana. New Phytol 131:443–451CrossRefGoogle Scholar
  12. Ekblad A, Wallander H, Godbold DL, Cruz C, Johnson D, Baldrian P et al. (2013) The production and turnover of extramatrical mycelium of ectomycorrhizal fungi in forest soils: role in carbon cycling. Plant Soil. doi: 10.1007/s11104-013-1630-3
  13. Frostegård Å, Bååth E (1996) The use of phospholipid fatty acid analysis to estimate bacterial and fungal biomass in soil. Biol Fertil Soils 22:59–65CrossRefGoogle Scholar
  14. Gerstberger P, Foken T, Kalbitz K (2004) The Lehstenbach and Steinkreuz catchments in NE Bavaria, Germany. In: Matzner E (ed) Biogeochemistry of forested catchments in a changing environment: A german case study, Ecol Stud 172. Springer, Berlin, pp 15–41Google Scholar
  15. Godbold DL, Fritz H-W, Jentschke G, Meesenburg H, Rademacher P (2003) Root turnover and root necromass accumulation of Norway spruce (Picea abies) are affected by soil acidity. Tree Physiol 23:915–921PubMedCrossRefGoogle Scholar
  16. Godbold D, Hoosbeek M, Lukac M (2006) Mycorrhizal hyphal turnover as a dominant process for carbon input into soil organic matter. Plant Soil 281:15–24CrossRefGoogle Scholar
  17. Güselwell S (2004) N:P ratios in terrestrial plants: variation and functional significance. New Phytol 164:243–266CrossRefGoogle Scholar
  18. Hagerberg D, Wallander H (2002) The impact of forest residue removal and wood ash amendment on the growth of the ectomycorrhizal external mycelium. FEMS Microbiol Ecol 39:139–146PubMedCrossRefGoogle Scholar
  19. Hedh J, Wallander H, Erland S (2008) Ectomycorrhizal mycelial species composition in apatite amended and non-amended mesh bags buried in a phosphorus-poor spruce forest. Mycol Res 112:681–688PubMedCrossRefGoogle Scholar
  20. Helmisaari H, Hallbäcken L (1999) Fine-root biomass and necromass in limed and fertilized Norway spruce (Picea abies [L.] Karst.) stands. For Ecol Manag 119:99–110CrossRefGoogle Scholar
  21. Hendricks JJ, Mitchell RJ, Kuehn KA, Pecot SD, Sims SE (2006) Measuring external mycelia production of ectomycorrhizal fungi in the field: the soil matrix matters. New Phytol 171:179–186PubMedCrossRefGoogle Scholar
  22. Hobbie EA (2006) Carbon allocation to ectomycorrhizal fungi correlates with belowground allocation in culture studies. Ecology 87:563–569PubMedCrossRefGoogle Scholar
  23. Högberg MN, Högberg P (2002) Extramatrical ectomycorrhizal mycelium contributes one-third of microbial biomass and produces, together with associated roots, half the dissolved organic carbon in a forest soil. New Phytol 154:791–795CrossRefGoogle Scholar
  24. Hughes J, Hodge A, Fitter A, Atkin O (2008) Mycorrhizal respiration: implications for global scaling relationships. Trends Plant Sci 13:583–588PubMedCrossRefGoogle Scholar
  25. IUSS Working Group WRB (2006) 2006 World reference base for soil resourcesGoogle Scholar
  26. Jentschke G, Drexhage M, Fritz H, Fritz E (2001) Does soil acidity reduce subsoil rooting in Norway spruce (Picea abies)? Plant Soil 237:91–108CrossRefGoogle Scholar
  27. Korkama T, Fritze H, Pakkanen A, Pennanen T (2007) Interactions between extraradical ectomycorrhizal mycelia, microbes associated with the mycelia and growth rate of Norway spruce (Picea abies) clones. New Phytol 173:798–807PubMedCrossRefGoogle Scholar
  28. Lukac M, Godbold DL (2001) Short Communication A modification of the ingrowth-core method to determine root production in fast growing tree species. J Plant Nutr Soil Sci 164:613–614CrossRefGoogle Scholar
  29. Majdi H, Truus L, Johansson U, Nylund J-E, Wallander H (2008) Effects of slash retention and wood ash addition on fine root biomass and production and fungal mycelium in a Norway spruce stand in SW Sweden. For Ecol Manag 255:2109–2117CrossRefGoogle Scholar
  30. Makkonen K, Helmisaari H (1999) Assessing fine-root biomass and production in a Scots pine stand–comparison of soil core and root ingrowth core methods. Plant Soil 210:43–50CrossRefGoogle Scholar
  31. Matzner E, Murach D, Fortmann H (1986) Soil acidity and its relationship to root growth in declining forest stands in Germany. Water Air Soil Pollut 31:273–282CrossRefGoogle Scholar
  32. Matzner E, Köstner B, Lischeid G (2004) Biogeochemistry of two forested catchments in a changing environment: A synthesis. In: Matzner E (ed) Biogeochemistry of forested catchments in a changing environment: A german case study, Ecol Stud 172. Springer, Berlin, pp 457–489Google Scholar
  33. Mellert K, Göttlein A (2012) Comparison of new foliar nutrient thresholds derived from van den Burg’s literature compilation with established central European references. Eur J Forest Res 131:1461–1472Google Scholar
  34. Nilsson LO, Wallander H (2003) Production of external mycelium by ectomycorrhizal fungi in a norway spruce forest was reduced in response to nitrogen fertilization. New Phytol 158:409–416CrossRefGoogle Scholar
  35. Nilsson LO, Giesler R, Bååth E, Wallander H (2005) Growth and biomass of mycorrhizal mycelia in coniferous forests along short natural nutrient gradients. New Phytol 165:613–622PubMedCrossRefGoogle Scholar
  36. Nilsson LO, Bååth E, Falkengren-Grerup U, Wallander H (2007) Growth of ectomycorrhizal mycelia and composition of soil microbial communities in oak forest soils along a nitrogen deposition gradient. Oecologia 153:375–384PubMedCrossRefGoogle Scholar
  37. Nilsson LO, Wallander H, Gundersen P (2012) Changes in microbial activities and biomasses over a forest floor gradient in C-to-N ratio. Plant Soil 355:75–86CrossRefGoogle Scholar
  38. Ostonen I, Helmisaari H (2011) Fine root foraging strategies in Norway spruce forests across a European climate gradient. Glob Change Biol 17:3620–3632CrossRefGoogle Scholar
  39. Ostonen I, Lõhmus K, Pajuste K (2005) Fine root biomass, production and its proportion of NPP in a fertile middle-aged Norway spruce forest: comparison of soil core and ingrowth core methods. For Ecol Manag 212:264–277CrossRefGoogle Scholar
  40. Ostonen I, Püttsepp Ü, Biel C, Alberton O, Bakker MR, Lohmus K et al (2007) Specific root length as an indicator of environmental change. Plant Biosyst 141:426–442CrossRefGoogle Scholar
  41. Parrent JL, Vilgalys R (2007) Biomass and compositional responses of ectomycorrhizal fungal hyphae to elevated CO2 and nitrogen fertilization. New Phytol 176:164–174PubMedCrossRefGoogle Scholar
  42. Persson H (1980) Spatial distribution of fine-root growth, mortality and decomposition in a young Scots pine stand in Central Sweden. Oikos 34:77–87CrossRefGoogle Scholar
  43. Pritchard SG, Strand AE, McCormack ML, Davis MA, Oren R (2008) Mycorrhizal and rhizomorph dynamics in a loblolly pine forest during 5 years of free-air-CO2-enrichment. Glob Change Biol 14:1252–1264CrossRefGoogle Scholar
  44. Schramel P, Wolf A, Seif R, Klose B-J (1980) Eine Apparatur zur Druckveraschung von biologischem Material. Fresenius’ Z Anal Chem 302:62–64CrossRefGoogle Scholar
  45. Schramel P, Wendler I, Knapp G (1996) Total digestion of silicate containing matrices (plants, soil, sludges) using a pressure ashing device with PFA-vessels. Fresenius’ Z Anal Chem 356:512–514Google Scholar
  46. Setala H, Kulmala P, Mikola J, Markkola AM (1999) Influence of ectomycorrhiza on the structure of detrital food webs in pine rhizosphere. Oikos 87:113CrossRefGoogle Scholar
  47. Taylor A, Martin F, Read D (2000) Fungal diversity in ectomycorrhizal communities of Norway spruce (Picea abies [L.] Karst.) and beech (Fagus sylvatica L.) along north-south transects in Europe. In: Schulze ED (ed) Carbon and nitrogen cycling in european forest catchments, Ecol Stud 142. Springer, Heidelberg, pp 343–365Google Scholar
  48. Vogt K, Persson H (1991) Measuring growth and development of roots. In: Lassoie JP, Hinkley TM (eds) Techniques and approaches in forest tree. CRC Press, Boca Raton, pp 477–501Google Scholar
  49. Vogt K, Vogt D, Bloomfield J (1998) Analysis of some direct and indirect methods for estimating root biomass and production of forests at an ecosystem level. Plant Soil 200:71–89CrossRefGoogle Scholar
  50. Wallander H (2000) Uptake of P from apatite by Pinus sylvestris seedlings colonised by different ectomycorrhizal fungi. Plant Soil 218:249–256CrossRefGoogle Scholar
  51. Wallander H, Nylund J-E (1992) Effects of excess nitrogen and phosphorus starvation on the extramatrical mycelium of ectomycorrhizas of Pinus sylvestris L. New Phytol 120:495–503CrossRefGoogle Scholar
  52. Wallander H, Thelin G (2008) The stimulating effect of apatite on ectomycorrhizal growth diminishes after PK fertilization. Soil Biol Biochem 40:2517–2522CrossRefGoogle Scholar
  53. Wallander H, Nilsson LO, Hagersberg D, Bååth E (2001) Estimation of the biomass and seasonal growth of externalmycelium of ectomycorrhizal fungi in the field. New Phytol 151:753–760CrossRefGoogle Scholar
  54. Wallander H, Göransson H, Rosengren U (2004) Production, standing biomass and natural abundance of 15N and 13C in ectomycorrhizal mycelia collected at different soil depths in two forest types. Oecologia 139:89–97PubMedCrossRefGoogle Scholar
  55. Wallander H, Johansson U, Sterkenburg E, Durling MB, Lindahl BD (2010) Production of ectomycorrhizal mycelium peaks during canopy closure in Norway spruce forests. New Phytol 187:1124–1134PubMedCrossRefGoogle Scholar
  56. Wallander H, Ekblad A, Bergh J (2011) Growth and carbon sequestration by ectomycorrhizal fungi in intensively fertilized Norway spruce forests. For Ecol Manag 262:999–1007CrossRefGoogle Scholar
  57. Wallander H, Ekblad A, Godbold DL, Johnson D, Bahr A, Baldrian P et al. (2012) Evaluation of methods to estimate production, biomass and turnover of ectomycorrhizal mycelium in forests soils – a review. Soil Biol Biochem. doi: 10.1016/j.soilbio.2012.08.027

Copyright information

© Springer Science+Business Media Dordrecht 2013

Authors and Affiliations

  1. 1.Soil Ecology, Bayreuth Center of Ecology and Environmental Research (BayCEER)University of BayreuthBayreuthGermany

Personalised recommendations