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… On Predicting Root Decomposition Kim H. Ludovici USDA Forest Service, SRS-4160 Abstract Table 1. Summary of literature review including root size and species, and decomposition rate constants (k-values) when published. Ref. # Date 1955 1973 1982 1982 1983 1983 1983 1984 1984 1985 1986 1987 1988 1988 1991 1992 1993 1993 1993 1993 1994 1994 1994 1995 1996 1997 1997 1997 1997 1997 1998 1998 1998 1998 1999 2000 2000 2000 2000 2000 2001 2001 2001 2001 2001 2002 2002 2002 2002 2003 2004 2004 2005 2006 2006 2006 2006 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 Determining what we know about root decomposition means going back through the literature, to see what has really been done, what assumptions have been made, what species and environmental conditions are included, and how the work was implemented. A literature search for peer-reviewed publications reguarding “tree and root and decomposition” identified only 58 articles published in the past 40 years (Table 1). Of those 58, only 24 publications included within-species manipulations and/or direct tests of site effects on root decomposition rate (Table 2). Post-harvest decomposition (Fig. 1) is virtually unstudied, even though large roots persist for many years (Fig. 2), and contribute a sizable pool of C to the developing forest (Fig. 3) Methodologies have varied widely, but most often utilize live, excised roots that are washed and dried prior to decomposition (Fig. 4). Studies incorporating season and root hydration (Fig. 5) suggest additional sources of variability in root decomposition rates. 23 24 < 2 mm < 1.5 mm < 3 mm < 2 mm < 2 mm < 19 cm > 10 mm < 2 mm < 2 mm < 5 mm > 10 mm cellulose < 2 mm < 5 mm < 2 mm < 1 mm < 2 mm < 2 mm < 3 cm < 5 mm < 2 mm < 3 mm < 3 mm < 2 cm < 2 mm < 2 mm <10 mm review 1 year 2 years 1 year 1 year 2 years 5.5 years 3 years 5 years 2 years 1 year 2 years 2 years 1 year 1 year 1 year 3.5 years 1 year 1 year 1 year chrono 5 years 0.75 yr 1.5 years 3 years 2+ years 0.75 yr 1 time 1 year < 2 mm 2.5 years < 2 mm 1 year < 2 mm 2 years stumps 60 years < 2 mm 1 year < 10 mm 2 years < 2 mm 0.15 yr < .5 mm 0.25 yr < 3 mm 0.5 year < 10 mm 1 year < 10 mm 1.5 years > 2 mm 1 time 0.60 - 0.83 Species Pinus Pinus Pinus, HW Picea Abies, Pinus, Pseud. Quercus mixed deciduous Pinus Pinus, Quercus, Acer Pinus Pinus Acer Pinus Prestoea, Dacryodes Acer Pinus Dacryodes HW HW Fagus, Fraxinus Picea 0.4 - 0.9 0.001 - 0.01 0.261 - 0.483 0.032 - 3.734 0.10 - 0.715 0.17 - 0.22 0.156 - 0.467 0.81 - 6.17 0.023 - 0.055 0.669 - 0.709 0.001 - 0.002 0.001 - 0.857 Spartina Pinus boreal forest Festuca Pinus, Alnus Acacia Gutierrezia Liquidambar 4 species combined Metrosideros Picea, Pseud., Pinus Pinus, Drypetes Juglans, Quercus Quercus, Pinus Fagus Quercus Quercus, Fagus HW Pinus published data Picea, Pseud, Pinus Acer Pinus Pinus Quercus, Fagus Dichrostachys Acer, Pinus Populus, Acer Eucalyptus Quercus Pinus Pinus, HW Author Orlov Head Yavitt & Fahey McClaugherty et. al. Alexander & Fairley Fogel Thompson & Boddy McClaugherty et. al. Persson Aber et. al. Gholz et. al. Nambiar Fahey et. al. Hunt et. al. Faber et. al. Ruark & Proe Bloomfield et. al. Hendrick & Pregistzer Ruark Silver & Vogt Fahey & Hughes Fahey & Arthur Scheu & Schauermann Lohmus & Ivask Arunachalam et. al. Conn & Day King et. al. Kurka & Starr Robinson et. al. Steele et. al. Lehmann & Zech Mun & Whitford Price & Hendrick Singh Ostertag & Hobbie Chen et. al. Gholz Jose et. al. Usman et. al. Zeller et. al. Dilustro et. al. Dress & Boerner Gaudinski et. al. John et. al. Silver & Miya Chen et. al. Dornbush et. al. John et. al. Ludovici & Richter Dress & Boerner Manlay et. al. Rotkin-Ellman et. al. King et. al. Binkley et. al. Fritz et. al. Ludovici & Kress Miller et. al. Results Discussion Summarization across species, ecotomes and methodologies suggest: Scores Soilsnot 0-20differ cm Root decomposition rate All does by soil depth Soil temperature may, or may not be positively related to decomposition rate Root decomposition is affected by soil texture Site fertility is not a predictor of decomposition rate Soil biota are always important for root decomposition Ambient temperature and CO2 level do not strongly impact root decomposition rates Decomposition is negatively related to root diameter Nutrient concentrations in roots may or may not be related to decomposition rate Decomposition is negatively related to root lignin and carbohydrate PC1 concentrations Root decomposition rates vary widely between species Research studies have been conducted on every continent, and on many of the most economically important species. The 2 phase pattern of decomposition, often reported for fine roots, supports the idea that root structure and complexity control nutrient release rates from roots, however, most studies include only roots < 5mm in diameter, and last only a year or two (approximately the duration of a Master’s project). Consideration that most decomposition work is conducted on a small percentage of the root system (just 5 – 15% of the total tree biomass), and has a duration far shorter than the lifespan of a fine root, is necessary. Root decomposition and nutrient release are also traditionally estimated from dried root tissues, and while it is unlikely that roots dehydrate prior to decomposition in-situ, the limited studies disagree on the cause and effect. PC2 Conclusions Table 2. Summary of root decomposition rate response to within-species manipulations and/or direct tests of site effects. Ref Soil H2O Soil depth Soil temp --- Soil text Soil Soil amb fert biota temp CO2 level Root size --+ --- Root nutrients 1 2 O O 3 4 O O 5 + --6 O O 7 + --O+ --- O 8 O ----9 O O --10 + O --+ 11 +O + 13 14 + + --+ 15 O O O 16 + + --+ 17 O O --O 18 19 + 20 --21 O 22 + + --+ 23 + O 24 --O Responses could be + (positive), --- (negative) or O (no significant response). Root lignin +O O --- Root carbo Tree species --- O O+ + We are seriously over estimating the amount of root decomposition that occurs over a forest rotation. Additional studies are required to test the mechanisms of nutrient loss, and the long-term decomposition rates of larger roots. + + --- ----+ --- O + + --- --- + 100 90 A J dry J fresh 80 O dry O fresh 70 Ja dry Ja fresh 60 A dry 50 Large root decomposition Carbon surrounding a stump A fresh 40 1200 160 B 55 tap wt = 119.03e Root supplied C -0.059x lateral wt = 58.715e 80 -0.1073x 2 R = 0.7269 40 1000 total soil C 45 2 R = 0.7962 120 35 800 25 15 0 600 5 0 20 40 60 80 Years since harvest -5 0 10 20 30 40 50 Years since harvest 60 400 70 Ludovici et al. 2002 CJFR Fig. 2. Biomass of mature loblolly pine root systems recovered along a time chronosequence, Durham, NC 200 Fig. 3. Carbon pools in decomposing roots and the soil volume surrounding them, measured along a time chronosequence, Durham, NC 2000 Ju nA 97 ug -9 O 7 ct D 97 ec -9 Fe 7 b9 A 8 pr -9 Ju 8 nA 98 ug -9 O 8 ct D 98 ec -9 Fe 8 b9 A 9 pr -9 9 Introduction and Methods 16 17 18 19 20 21 22 kg/tree Fig. 1 Photograph of recovered root material from 55 to 70-year-old loblolly pine stumps that had been decomposing for (A) 5 years, (B) 20 years, (C) 10 years, and (D) 55 years, on a Kanhapludult in the Piedmont region of North Carolina. Reproduced from Ludovici et al. 2002 Root diam Duration k-values < 3 mm review < 1 cm chrono 0.0415 < 5 cm 1 year < 5 mm 2 years < 2 mm review large < 3 mm 0.193 - 0.696 < 2 mm chrono < 3 mm 1 year < 5 mm 1 year > 5 mm 2.5 years < 10 cm 2 years 0.042 - 0.093 kg / tree Quantification of root decomposition remains controversial because researchers can not control the process. The literature provides examples in which researchers have limited the size, age and species of roots used in studies, or controlled the onset of decomposition. Others have controlled ambient and/or soil conditions during root growth and decomposition. Still, there is not one quintessential root decomposition methodology that can be utilized across species or ecotomes. Using data from available publications, decay curves will be generated with consideration given to tree species, stand age, root size, temperature and methodology. Fig. 4. Photograph of a controlled environment chamber used in root incubations. Fig. 5. Graph of % biomass loss (A) and carbon concentrations (B) in fine roots of Pinus taeda decomposing in-situ. References 1. Yavitt, J.B. and T.J. Fahey. 1982. Loss of mass and nutrient changes of decaying woody roots in lodgepole pine forests, southeastern Wyoming. Can. J. For. Res. 12:745-752 2. McClaugherty, C.A., J.D. Aber and J.M. Melillo. 1984. Decomposition dynamics of fine roots in forested ecosystems. Oikos 42:378-386 3. Fahey, T.J., J.W. Hughes, Mou Pu and M.A. Arthur. 1988. Root Decomposition and Nutrient Flux Following Whole-Tree Harvest of Northern Hardwood Forest. For. Sci. 34(3):744-768 4. Bloomfield, J., K.A. Vogt and D.J. Vogt. 1993. Decay rate and substrate quality of fine roots and foliage of two tropical tree species in the Luquillo Experimental Forest, Puerto Rico. Plant and Soil 150:233-245 5. Ruark, G.A. 1993. Modeling Soil Temperature Effects on In Situ Decomposition Rates for Fine Roots of Loblolly Pine. For. Sci. 39(1):118-129 6. Silver, W.L. and K.A. Vogt. 1993. Fine root dynamics following single and multiple disturbances in a subtropical wet forest ecosystem. J. Ecology. 81:729-738 7. Scheu, S. and J. Schauermann. 1994.Decomposition of roots and twigs: Effects of wood type (beech and ash), diameter, site of exposure and macrofauna exclusion. Plant and Soil 163:13-24 8. Lohmus,K. and M. 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