The unseen iceberg: Plant roots in arctic tundra

New Phytologist

Volumn 205, Issue 1, 2015, Pages 34-58

  Iversen, Colleen M ^a,b   Sloan, Victoria L ^a,b   Sullivan, Patrick F ^c   Euskirchen, Eugenie S ^d   Mcguire, A David ^d   Norby, Richard J ^a,b   Walker, Anthony P ^a,b   Warren, Jeffrey M ^a,b   Wullschleger, Stan D ^a,b  

^a OAK RIDGE NATIONAL LABORATORY   (United States)

^b OAK RIDGE NATIONAL LABORATORY   (United States)

^c UNIVERSITY OF ALASKA ANCHORAGE   (United States)

^d UNIVERSITY OF ALASKA FAIRBANKS   (United States)

Author keywords

Arctic; Fine roots; Model; Plant soil; Root biomass; Root production; Root turnover; Tundra

Indexed keywords

ARCTIC; ATMOSPHERE; BIOLOGICAL MODEL; ICE COVER; PHYSIOLOGY; PLANT ROOT; TUNDRA;

ARCTIC REGIONS; ATMOSPHERE; ICE COVER; MODELS, BIOLOGICAL; PLANT ROOTS; TUNDRA;

EID: 84911996607     PISSN: 0028646X     EISSN: 14698137     Source Type: Journal    
DOI: 10.1111/nph.13003     Document Type: Review

References (202)

1. Aerts R. 2006. The freezer defrosting: global warming and litter decomposition rates in cold biomes. Journal of Ecology 94: 713-724.
2. Aerts R, Bakker C, Decaluwe H. 1992. Root turnover as determinant of the cycling of C, N, and P in a dry heathland ecosystem. Biogeochemistry 15: 175-190.
3. Allessio ML, Tieszen LL. 1975. Patterns of carbon allocation in an arctic tundra grass, Dupontia fischeri (Gramineae), at Barrow, Alaska. American Journal of Botany 62: 797-807.
4. Atkin OK. 1996. Reassessing the nitrogen relations of Arctic plants: a mini-review. Plant, Cell & Environment 19: 695-704.
5. Atkin OK, Cummins WR. 1994. The effect of root temperature on the induction of nitrate reductase activites and nitrogen uptake rates in arctic plant species. Plant and Soil 159: 187-197.
6. Atkin OK, Villar R, Cummins WR. 1993. The ability of several high arctic plant species to utilize nitrate-nitrogen under field conditions. Oecologia 96: 239-245.
7. Bardgett RD, van der Wal R, Jonsdottir IS, Quirk H, Dutton S. 2007. Temporal variability in plant and soil nitrogen pools in a high-Arctic ecosystem. Soil Biology & Biochemistry 39: 2129-2137.
8. Barry RG, Courtin GM, Labine C. 1981. Tundra climates. In: Bliss LC, Heal OW, Moore JJ, eds. Tundra ecosystems: a comparative analysis. New York, NY, USA: Cambridge University Press, 81-114.
9. 3- uptake kinetics. New Phytologist 133: 423-430.
10. Bell KL, Bliss LC. 1978. Root growth in a polar semidesert environment. Canadian Journal of Botany 56: 2470-2490.
11. Bilbrough CJ, Welker JM, Bowman WD. 2000. Early spring nitrogen uptake by snow-covered plants: a comparison of arctic and alpine plant function under the snowpack. Arctic, Antarctic and Alpine Research 32: 404-411.
12. Billings WD, Mooney HA. 1968. Ecology of arctic and alpine plants. Biological Reviews of the Cambridge Philosophical Society 43: 481-529.
13. Billings WD, Peterson KM, Shaver GR. 1978. Growth, turnover, and respiration rates of roots and tillers in tundra graminoids. In: Tieszen LL, ed. Vegetation and production ecology of an Alaskan arctic tundra. New York, NY, USA: Springer-Verlag, 415-434.
14. Billings WD, Peterson KM, Shaver GR, Trent AW. 1977. Root growth, respiration, and carbon-dioxide evolution in an arctic tundra soil. Arctic and Alpine Research 9: 129-137.
15. Billings WD, Shaver GR, Trent AW. 1973. Temperature effects on growth and respiration of roots and rhizomes in tundra graminoids. In: Bliss LC, Wielgolaski FE, eds. Primary production and production processes. Stockholm, Sweden: IBP Tundra Biome Steering Committee, 57-63.
16. Billings WD, Shaver G, Trent AW. 1976. Measurement of root growth in simulated and natural temperature gradients over permafrost. Arctic and Alpine Research 8: 247-250.
17. Björk RG, Majdi H, Klemedtsson L, Lewis-Jonsson L, Molau U. 2007. Long-term warming effects on root morphology, root mass distribution, and microbial activity in two dry tundra plant communities in northern Sweden. New Phytologist 176: 862-873.
18. Bliss LC, Svoboda J. 1984. Plant-communities and plant-production in the Western Queen Elizabeth Islands. Holarctic Ecology 7: 325-344.
19. Bliss LC, Svoboda J, Bliss DI. 1984. Polar deserts, their plant cover and plant-production in the Canadian High Arctic. Holarctic Ecology 7: 305-324.
20. Borden PW, Ping CL, McCarthy PJ, Naidu S. 2010. Clay mineralogy in Arctic Tundra Gelisols, northern Alaska. Soil Science Society of America Journal 74: 580-592.
21. Brown J, Ferrians OJ Jr, Heginbottom JA, Melnikov ES, eds. 1997. Circum-Arctic map of permafrost and ground-ice conditions. Washington, DC, USA: US Geological Survey in Cooperation with the Circum-Pacific Council for Energy and Mineral Resources. Circum-Pacific Map Series CP-45, scale 1 : 10, 000, 000.
22. Cahoon SMP, Sullivan PF, Shaver GR, Welker JM, Post E. 2012. Interactions among shrub cover and the soil microclimate may determine future Arctic carbon budgets. Ecology Letters 15: 1415-1422.
23. Caldwell MM. 1987. Competition between root systems in natural communities. In: Gregory P, Lake J, Rose D, eds. Root development and function. Cambridge, UK: Cambridge University Press, 167-186.
24. Campioli M, Michelsen A, Demey A, Vermeulen A, Samson R, Lemeur R. 2009. Net primary production and carbon stocks for subarctic mesic-dry tundras with contrasting microtopography, altitude, and dominant species. Ecosystems 12: 760-776.
25. Chanton JP, Martens CS, Kelley CA, Crill PM, Showers WJ. 1992. Methane transport mechanisms and isotopic fractionation in emergent macrophytes of an Alaskan tundra lake. Journal of Geophysical Research 97: 16681-16688.
26. Chapin FS. 1974. Morphological and physiological mechanisms of temperature compensation in phosphate absorption along a latitudinal gradient. Ecology 55: 1180-1198.
27. Chapin FS. 1978. Phosphate uptake and nutrient utilization by Barrow tundra vegetation. In: Tieszen LL, ed. Vegetation and production ecology of an Alaskan arctic tundra. New York, NY, USA: Springer-Verlag New York Inc, 483-507.
28. Chapin FS. 1981. Field measurements of growth and phosphate absorption in Carex aquatilis along a latitudinal gradient. Arctic and Alpine Research 13: 83-94.
29. Chapin FS. 1983. Direct and indirect effects of temperature on arctic plants. Polar Biology 2: 47-52.
30. Chapin FS, Barsdate RJ, Barel D. 1978. Phosphorus cycling in Alaskan coastal tundra: hypothesis for the regulation of nutrient cycling. Oikos 31: 189-199.
31. Chapin FS, Bloom A. 1976. Phosphate absorption: adaptation of tundra graminoids to a low temperature, low phosphorus environment. Oikos 27: 111-121.
32. Chapin FS, BretHarte MS, Hobbie SE, Zhong HL. 1996. Plant functional types as predictors of transient responses of arctic vegetation to global change. Journal of Vegetation Science 7: 347-358.
33. Chapin FS, Fetcher N, Kielland K, Everett KR, Linkins AE. 1988. Productivity and nutrient cycling of Alaskan tundra: enhancement by flowing soil-water. Ecology 69: 693-702.
34. Chapin FS, Johnson DA, McKendrick JD. 1980b. Seasonal movement of nutrients in plants of differing growth form in an Alaskan tundra ecosystem: implications for herbivory. Journal of Ecology 68: 189-209.
35. Chapin FS, Moilanen L, Kielland K. 1993. Preferential use of organic nitrogen for growth by a nonmycorrhizal arctic sedge. Nature 361: 150-153.
36. Chapin FS, Shaver GR. 1996. Physiological and growth responses of arctic plants to a field experiment simulating climatic change. Ecology 77: 822-840.
37. Chapin FS, Slack M. 1979. Effect of defoliation upon root growth, phosphate absorption and respiration in nutrient-limited tundra graminoids. Oecologia 42: 67-79.
38. Chapin FS, Tieszen LL, Lewis MC, Miller PC, McCown BH. 1980a. Control of tundra plant allocation patterns and growth. In: Brown J, Miller PC, Tieszen LL, Bunnell FL, eds. An arctic ecosystem: the coastal tundra at Barrow, Alaska. Stroudsburg, PA, USA: Dowden, Hutchinson & Ross, Inc., 140-185.
39. Chapin FS, Tryon PR. 1982. Phosphate absorption and root respiration of different plant growth forms from northern Alaska. Holarctic Ecology 5: 164-171.
40. Chapin FS, van Cleve K, Tieszen LL. 1975. Seasonal nutrient dynamics of tundra vegetation at Barrow, Alaska. Arctic and Alpine Research 7: 209-226.
41. Cheng WX, Parton WJ, Gonzalez-Meler MA, Phillips R, Asao S, McNickle GG, Brzostek E, Jastrow JD. 2014. Synthesis and modeling perspectives of rhizosphere priming. New Phytologist 201: 31-44.
42. Clemmensen KE, Michelsen A, Jonasson S, Shaver GR. 2006. Increased ectomycorrhizal fungal abundance after long-term fertilization and warming of two arctic tundra ecosystems. New Phytologist 171: 391-404.
43. Clemmensen KE, Sorensen PL, Michelsen A, Jonasson S, Strom L. 2008. Site-dependent N uptake from N-form mixtures by arctic plants, soil microbes and ectomycorrhizal fungi. Oecologia 155: 771-783.
44. Cornwell WK, Cornelissen JHC, Amatangelo K, Dorrepaal E, Eviner VT, Godoy O, Hobbie SE, Hoorens B, Kurokawa H, Perez-Harguindeguy N et al. 2008. Plant species traits are the predominant control on litter decomposition rates within biomes worldwide. Ecology Letters 11: 1065-1071.
45. 2 emission from a northern peatland: root respiration, exudation, and decomposition. Ecology 86: 1825-1834.
46. Dennis JG. 1977. Distribution patterns of belowground standing crop in arctic tundra at Barrow, Alaska. Arctic and Alpine Research 9: 113-127.
47. Dennis JG, Johnson PL. 1970. Shoot and rhizome-root standing crops of tundra vegetation at Barrow, Alaska. Arctic and Alpine Research 2: 253-266.
48. Dennis JG, Tieszen LL, Vetter MA. 1978. Seasonal dynamics of above- and belowground production of vascular plants at Barrow, Alaska. In: Tieszen LL, ed. Vegetation and production ecology of an Alaskan arctic tundra. New York, NY, USA: Springer-Verlag New York Inc, 113-140.
49. Deslippe JR, Simard SW. 2011. Below-ground carbon transfer among Betula nana may increase with warming in Arctic tundra. New Phytologist 192: 689-698.
50. Edwards KA, Jefferies RL. 2010. Nitrogen uptake by Carex aquatilis during the winter-spring transition in a low Arctic wet meadow. Journal of Ecology 98: 737-744.
51. Eissenstat DM, Yanai RD. 1997. The ecology of root lifespan. In: Begon M, Fitter AH, eds. Advances in ecological research, Vol. 27. New York, NY, USA: Academic Press, 1-60.
52. Ellis BA, Kummerow J. 1982. Temperature effect on growth rates of Eriophorum vaginatum roots. Oecologia 54: 136-137.
53. Epstein HE, Walker MD, Chapin FS, Starfield AM. 2000. A transient nutrient-based model of Arctic plant community response to climatic warming. Ecological Applications 10: 824-841.
54. Eskelinen A, Stark S, Mannisto M. 2009. Links between plant community composition, soil organic matter quality and microbial communities in contrasting tundra habitats. Oecologia 161: 113-123.
55. Euskirchen ES, McGuire AD, Chapin FS, Yi S, Thompson CC. 2009. Changes in vegetation in northern Alaska under scenarios of climate change, 2003-2100: implications for climate feedbacks. Ecological Applications 19: 1022-1043.
56. Fan Z, McGuire AD, Turetsky MR, Harden JW, Waddington JM, Kane ES. 2013. The response of soil organic carbon of a rich fen peatland in interior Alaska to projected climate change. Global Change Biology 19: 604-620.
57. Freschet GT, Aerts R, Cornelissen JHC. 2012a. A plant economics spectrum of litter decomposability. Functional Ecology 26: 56-65.
58. Freschet GT, Aerts R, Cornelissen JHC. 2012b. Multiple mechanisms for trait effects on litter decomposition: moving beyond home-field advantage with a new hypothesis. Journal of Ecology 100: 619-630.
59. Freschet GT, Cornelissen JHC, van Logtestijn RSP, Aerts R. 2010a. Evidence of the 'plant economics spectrum' in a subarctic flora. Journal of Ecology 98: 362-373.
60. Freschet GT, Cornelissen JHC, van Logtestijn RSP, Aerts R. 2010b. Substantial nutrient resorption from leaves, stems and roots in a subarctic flora: what is the link with other resource economics traits? New Phytologist 186: 879-889.
61. Freschet GT, Cornwell WK, Wardle DA, Elumeeva TG, Liu WD, Jackson BG, Onipchenko VG, Soudzilovskaia NA, Tao JP, Cornelissen JHC. 2013. Linking litter decomposition of above- and below-ground organs to plant-soil feedbacks worldwide. Journal of Ecology 101: 943-952.
62. Gebauer RLE, Tenhunen JD, Reynolds JF. 1996. Soil aeration in relation to soil physical properties, nitrogen availability, and root characteristics within an arctic watershed. Plant and Soil 178: 37-48.
63. Gill RA, Jackson RB. 2000. Global patterns of root turnover for terrestrial ecosystems. New Phytologist 147: 13-31.
64. Grogan P, Jonasson S. 2003. Controls on annual nitrogen cycling in the understory of a subarctic birch forest. Ecology 84: 202-218.
65. Guo DL, Xia MX, Wei X, Chang WJ, Liu Y, Wang ZQ. 2008. Anatomical traits associated with absorption and mycorrhizal colonization are linked to root branch order in twenty-three Chinese temperate tree species. New Phytologist 180: 673-683.
66. Harden JW, Koven CD, Ping CL, Hugelius G, McGuire AD, Camill P, Jorgenson T, Kuhry P, Michaelson GJ, O'Donnell JA et al. 2012. Field information links permafrost carbon to physical vulnerabilities of thawing. Geophysical Research Letters 39: 6.
67. Harmon ME, Silver WL, Fasth B, Chen H, Burke IC, Parton WJ, Hart SC, Currie WS. 2009. Long-term patterns of mass loss during the decomposition of leaf and fine root litter: an intersite comparison. Global Change Biology 15: 1320-1338.
68. Harms TK, Jones JB. 2012. Thaw depth determines reaction and transport of inorganic nitrogen in valley bottom permafrost soils. Global Change Biology 18: 2958-2968.
69. Hartley IP, Garnett MH, Sommerkorn M, Hopkins DW, Fletcher BJ, Sloan VL, Phoenix GK, Wookey PA. 2012. A potential loss of carbon associated with greater plant growth in the European Arctic. Nature Climate Change 2: 875-879.
70. Henry GHR, Svoboda J, Freedman B. 1990. Standing crop and net production of sedge meadows of an ungrazed polar desert oasis. Canadian Journal of Botany 68: 2660-2667.
71. Henry HAL, Jefferies RL. 2003. Plant amino acid uptake, soluble N turnover and microbial N capture in soils of a grazed Arctic salt marsh. Journal of Ecology 91: 627-636.
72. Hernandez RR, Allen MF. 2013. Diurnal patterns of productivity of arbuscular mycorrhizal fungi revealed with the Soil Ecosystem Observatory. New Phytologist 200: 547-557.
73. 14C. Global Change Biology 19: 649-661.
74. Hill GB, Henry GHR. 2011. Responses of High Arctic wet sedge tundra to climate warming since 1980. Global Change Biology 17: 276-287.
75. Hinzman LD, Kane DL, Gieck RE, Everett KR. 1991. Hydrologic and thermal properties of the active layer in the Alaskan arctic. Cold Regions Science and Technology 19: 95-110.
76. Hobbie SE. 1996. Temperature and plant species control over litter decomposition in Alaskan tundra. Ecological Monographs 66: 503-522.
77. Hobbie SE, Chapin FS. 1996. Winter regulation of tundra litter carbon and nitrogen dynamics. Biogeochemistry 35: 327-338.
78. 2 flux to experimental warming. Ecology 79: 1526-1544.
79. 15N in symbiotic fungi and plants estimates nitrogen and carbon flux rates in Arctic tundra. Ecology 87: 816-822.
80. Hobbie SE, Schimel JP, Trumbore SE, Randerson JR. 2000. Controls over carbon storage and turnover in high-latitude soils. Global Change Biology 6: 196-210.
81. Hugelius G, Bockheim JG, Camill P, Elberling B, Grosse G, Harden JW, Johnson K, Jorgenson T, Koven CD, Kuhry P et al. 2013. A new data set for estimating organic carbon storage to 3 m depth in soils of the northern circumpolar permafrost region. Earth System Science Data 5: 393-402.
82. 2 concentration in forested ecosystems. New Phytologist 186: 346-357.
83. Iversen CM, Murphy MT, Allen MF, Childs J, Eissenstat DM, Lilleskov EA, Sarjala TM, Sloan VL, Sullivan PF. 2012. Advancing the use of minirhizotrons in wetlands. Plant and Soil 352: 23-39.
84. Iversen CM, Sloan VL, Sullivan PF, Euskirchen ES, McGuire AD, Norby RJ, Walker AP, Warren JM, Wullschleger SD. 2014. Plant roots in arctic tundra. Next generation ecosystem experiments arctic data Collection. Oak Ridge, Tennessee, USA: Carbon Dioxide Information Analysis Center, Oak Ridge National Laboratory. Dataset doi: 10.5440/1114222.
85. Jackson RB, Canadell J, Ehleringer JR, Mooney HA, Sala OE, Schulze ED. 1996. A global analysis of root distributions for terrestrial biomes. Oecologia 108: 389-411.
86. Jackson RB, Schenk HJ, Jobbágy EG, Canadell J, Colello GD, Dickinson RE, Field CB, Friedlingstein P, Heimann M, Hibbard K et al. 2000. Belowground consequences of vegetation change and their treatment in models. Ecological Applications 10: 470-483.
87. Joabsson A, Christensen TR. 2001. Methane emissions from wetlands and their relationship with vascular plants: an Arctic example. Global Change Biology 7: 919-932.
88. Jobbágy EG, Jackson RB. 2000. The vertical distribution of soil organic carbon and its relation to climate and vegetation. Ecological Applications 10: 423-436.
89. Jobbágy EG, Jackson RB. 2001. The distribution of soil nutrients with depth: global patterns and the imprint of plants. Biogeochemistry 53: 51-77.
90. Jonasson S, Callaghan TV. 1992. Root mechanical properties related to disturbed and stressed habitats in the Arctic. New Phytologist 122: 179-186.
91. Jones DL. 1998. Organic acids in the rhizosphere - a critical review. Plant and Soil 205: 25-44.
92. Keuper FK. 2012. Direct and indirect effects of climatic changes on vegetation productivity and species composition of permafrost peatlands. PhD Thesis, VU University, Amsterdam, the Netherlands.
93. Keuper FK, van Bodegom PM, Dorrepaal E, Weedon JT, van Hal J, van Logtestijn RSP, Aerts R. 2012. A frozen feast: thawing permafrost increases plant-available nitrogen in subarctic peatlands. Global Change Biology 18: 1998-2007.
94. Kielland K. 1994. Amino-acid absorption by arctic plants: implications for plant nutrition and nitrogen cycling. Ecology 75: 2373-2383.
95. Kielland K, Chapin FS. 1994. Phosphate uptake in arctic plants in relation to phosphate supply: the role of spatial and temporal variability. Oikos 70: 443-448.
96. Kjelvik S, Kärenlampi L. 1975. Plant biomass and primary production of Fennoscandian subarctic and subalpine forests and of alpine willow and heath ecosystems. In: Wielgolaski FE, ed. Fennoscandian tundra ecosystems. New York, NY, USA: Springer-Verlag, 111-120.
97. Koch GW, Bloom AJ, Chapin FS. 1991. Ammonium and nitrate as nitrogen sources in two Eriophorum species. Oecologia 88: 570-573.
98. Koelbener A, Strom L, Edwards PJ, Venterink HO. 2010. Plant species from mesotrophic wetlands cause relatively high methane emissions from peat soil. Plant and Soil 326: 147-158.
99. Koven CD, Riley WJ, Subin ZM, Tang JY, Torn MS, Collins WD, Bonan GB, Lawrence DM, Swenson SC. 2013. The effect of vertically-resolved soil biogeochemistry and alternate soil C and N models on C dynamics of CLM4. Biogeosciences Discussions 10: 7201-7256.
100. Koven CD, Ringeval B, Friedlingstein P, Ciais P, Cadule P, Khvorostyanov D, Krinner G, Tarnocai C. 2011. Permafrost carbon-climate feedbacks accelerate global warming. Proceedings of the National Academy of Sciences, USA 108: 14769-14774.
101. Krinner G, Viovy N, de Noblet-Ducoudre N, Ogee J, Polcher J, Friedlingstein P, Ciais P, Sitch S, Prentice IC. 2005. A dynamic global vegetation model for studies of the coupled atmosphere-biosphere system. Global Biogeochemical Cycles 19: 1015.
102. Kummerow J, Ellis BA, Kummerow S, Chapin FS. 1983. Spring growth of shoots and roots in shrubs of an Alaskan muskeg. American Journal of Botany 70: 1509-1515.
103. Kummerow J, Krause D. 1982. The effects of variable nitrogen and phosphorus concentrations on Eriophorum vaginatum tillers grown in nutrient solutions. Holarctic Ecology 5: 187-193.
104. Kummerow J, McMaster GS, Krause DA. 1980. Temperature effect on growth and nutrient contents in Eriophorum vaginatum under controlled environmental conditions. Arctic and Alpine Research 12: 335-342.
105. Kummerow J, Russell M. 1980. Seasonal root growth in the arctic tussock tundra. Oecologia 47: 196-199.
106. Kutzbach L, Wagner D, Pfeiffer EM. 2004. Effect of microrelief and vegetation on methane emission from wet polygonal tundra, Lena Delta, northern Siberia. Biogeochemistry 69: 341-362.
107. Laanbroek HJ. 2010. Methane emission from natural wetlands: interplay between emergent macrophytes and soil microbial processes. A mini-review. Annals of Botany 105: 141-153.
108. Le Dizes S, Kwiatkowski BL, Rastetter EB, Hope A, Hobbie JE, Stow D, Daeschner S. 2003. Modeling biogeochemical responses of tundra ecosystems to temporal and spatial variations in climate in the Kuparuk River Basin (Alaska). Journal of Geophysical Research-Atmospheres 108: D2.
109. Limbach WE, Oechel WC, Lowell W. 1982. Photosynthetic and respiratory responses to temperature and light of three Alaskan tundra growth forms. Holarctic Ecology 5: 150-157.
110. Loya WM, Johnson LC, Kling GW, King JY, Reeburgh WS, Nadelhoffer KJ. 2002. Pulse-labeling studies of carbon cycling in arctic tundra ecosystems: contribution of photosynthates to soil organic matter. Global Biogeochemical Cycles 16: 1101.
111. Mack MC, Schuur EAG, Bret-Harte MS, Shaver GR, Chapin FS. 2004. Ecosystem carbon storage in arctic tundra reduced by long-term nutrient fertilization. Nature 431: 440-443.
112. Maessen O, Freedman B, Nams MLN, Svoboda J. 1983. Resource allocation in High Arctic vascular plants of differing growth form. Canadian Journal of Botany 61: 1680-1691.
113. 15N in tussock tundra ecosystems. Holarctic Ecology 10: 230-234.
114. McConnell NA, Turetsky MR, McGuire AD, Kane ES, Waldrop MP, Harden JW. 2013. Controls on ecosystem and root respiration across a permafrost and wetland gradient in interior Alaska. Environmental Research Letters 8: 045029.
115. McCormack ML, Thomas SA, Smithwick EAH, Eissenstat DM. 2012. Predicting fine root lifespan from plant functional traits in temperate trees. New Phytologist 195: 823-831.
116. McGraw JB, Chapin FS. 1989. Competititive ability and adaptation to fertile and infertile soils in two Eriophorum species. Ecology 70: 736-749.
117. McGuire AD, Anderson LG, Christensen TR, Dallimore S, Guo LD, Hayes DJ, Heimann M, Lorenson TD, Macdonald RW, Roulet N. 2009. Sensitivity of the carbon cycle in the Arctic to climate change. Ecological Monographs 79: 523-555.
118. McGuire AD, Christensen TR, Hayes D, Heroult A, Euskirchen E, Kimball JS, Koven C, Lafleur P, Miller PA, Oechel W et al. 2012. An assessment of the carbon balance of Arctic tundra: comparisons among observations, process models, and atmospheric inversions. Biogeosciences 9: 3185-3204.
119. McGuire AD, Hayes DJ, Kicklighter DW, Manizza M, Zhuang Q, Chen M, Follows MJ, Gurney KR, McClelland JW, Melillo JM et al. 2010. An analysis of the carbon balance of the Arctic Basin from 1997 to 2006. Tellus Series B-Chemical and Physical Meteorology 62: 455-474.
120. McKane RB, Johnson LC, Shaver GR, Nadelhoffer KJ, Rastetter EB, Fry B, Giblin AE, Kielland K, Kwiatkowski BL, Laundre JA et al. 2002. Resource-based niche provides a basis for plant species diversity and dominance in arctic tundra. Nature 415: 68-71.
121. 15N natural abundance in heath and forest tundra ecosystems is closely correlated with presence and type of mycorrhizal fungi in roots. Oecologia 115: 406-418.
122. 15N abundance of subarctic plants provides field evidence that ericoid, ectomycorrhizal and non- and arbuscular mycorrhizal species access different sources of soil nitrogen. Oecologia 105: 53-63.
123. Miller OK. 1982. Mycorrhizae, mycorrhizal fungi and fungal biomass in subalpine tundra at Eagle Summit, Alaska. Holarctic Ecology 5: 125-134.
124. Miller PA, Smith B. 2012. Modelling tundra vegetation response to recent arctic warming. Ambio 41: 281-291.
125. Miller PC, Mangan R, Kummerow J. 1982. Vertical distribution of organic matter in eight vegetation types near Eagle Summit, Alaska. Holarctic Ecology 5: 117-124.
126. Miller PC, Miller PM, Blakejacobson M, Chapin FS, Everett KR, Hilbert DW, Kummerow J, Linkins AE, Marion GM, Oechel WC et al. 1984. Plant-soil processes in Eriophorum vaginatum tussock tundra in Alaska: a systems modeling approach. Ecological Monographs 54: 361-405.
127. Miller PC, Stoner WA, Ehleringer JR. 1978. Some aspects of water relations of arctic and alpine regions. In: Tieszen LL, ed. Vegetation and production ecology of an Alaskan arctic tundra. New York, NY, USA: Springer-Verlag, 343-369.
128. Mokany K, Raison RJ, Prokushkin AS. 2006. Critical analysis of root : shoot ratios in terrestrial biomes. Global Change Biology 12: 84-96.
129. Muc M. 1977. Ecology and primary production of sedge-moss meadow communities, Truelove Lowland. In: Bliss LC, ed. Truelove Lowland, Devon Island, Canada: a high arctic ecosystem. Edmonton, AB, Canada: University of Alberta Press, 157-184.
130. Myhre G, Shindell D, Bréon F-M, Collins W, Fuglestvedt J, Huang J, Koch D, Lamarque J-F, Lee D, Mendoza B et al. 2013. Anthropogenic and natural radiative forcing. In: Stocker TF, Qin D, Plattner G-K, Tignor M, Allen SK, Boschung J, Nauels A, Xia Y, Bex V, Midgley PM, eds. Climate change. The physical science basis. Contribution of Working Group I to the fifth assessment report of the Intergovernmental Panel on Climate Change. New York, NY, USA: Cambridge University Press, 659-740.
131. 15N natural abundances and N use by tundra plants. Oecologia 107: 386-394.
132. Nadelhoffer KJ, Giblin AE, Shaver GR, Laundre JA. 1991. Effects of temperature and substrate quality on element mineralization in six arctic soils. Ecology 72: 242-253.
133. Nadelhoffer KJ, Johnson L, Laundre J, Giblin AE, Shaver GR. 2002. Fine root production and nutrient content in wet and moist arctic tundras as influenced by chronic fertilization. Plant and Soil 242: 107-113.
134. Näsholm T, Kielland K, Ganeteg U. 2009. Uptake of organic nitrogen by plants. New Phytologist 182: 31-48.
135. Newsham KK, Upson R, Read DJ. 2009. Mycorrhizas and dark septate root endophytes in polar regions. Fungal Ecology 2: 10-20.
136. Nowinski NS, Trumbore SE, Schuur EAG, Mack MC, Shaver GR. 2008. Nutrient addition prompts rapid destabilization of organic matter in an arctic tundra ecosystem. Ecosystems 11: 16-25.
137. Oleson KW, Lawrence DM, Bonan GB, Drewniak B, Huang M, Koven CD, Levis S, Li F, Riley WJ, Subin ZM et al. 2013. Technical description of version 4.5 of the Community Land Model (CLM). NCAR/TN-503+STR NCAR Technical Note.
138. 14C pulse-labelling. Soil Biology & Biochemistry 36: 245-253.
139. Öquist MG, Svensson BH. 2002. Vascular plants as regulators of methane emissions from a subarctic mire ecosystem. Journal of Geophysical Research-Atmospheres 107: 4580.
140. Orwin KH, Kirschbaum MUF, St John MG, Dickie IA. 2011. Organic nutrient uptake by mycorrhizal fungi enhances ecosystem carbon storage: a model-based assessment. Ecology Letters 14: 493-502.
141. Parton W, Silver WL, Burke IC, Grassens L, Harmon ME, Currie WS, King JY, Adair EC, Brandt LA, Hart SC et al. 2007. Global-scale similarities in nitrogen release patterns during long-term decomposition. Science 315: 361-364.
142. Ping CL, Bockheim JG, Kimble JM, Michaelson GJ, Walker DA. 1998. Characteristics of cryogenic soils along a latitudinal transect in Arctic Alaska. Journal of Geophysical Research-Atmospheres 103: 28917-28928.
143. Ping CL, Michaelson GJ, Packee EC, Stiles CA, Swanson DK, Yoshikawa K. 2005. Soil catena sequences and fire ecology in the boreal forest of Alaska. Soil Science Society of America Journal 69: 1761-1772.
144. Pregitzer KS. 2002. Fine roots of trees - a new perspective. New Phytologist 154: 267-270.
145. 2, climate, and N-deposition. Tree Physiology 9: 101-126.
146. Read DJ, Leake JR, Perez-Moreno J. 2004. Mycorrhizal fungi as drivers of ecosystem processes in heathland and boreal forest biomes. Canadian Journal of Botany 82: 1243-1263.
147. Riley WJ, Subin ZM, Lawrence DM, Swenson SC, Torn MS, Meng L, Mahowald NM, Hess P. 2011. Barriers to predicting changes in global terrestrial methane fluxes: analyses using CLM4Me, a methane biogeochemistry model integrated in CESM. Biogeosciences 8: 1925-1953.
148. 2 feedback. Biogeosciences 8: 2137-2157.
149. Rosswall T, Veum AK, Kärenlampi L. 1975. Plant litter decomposition at Fennoscandian tundra sites. In: Wielgolaski FE, ed. Fennoscandian tundra ecosystems. New York, NY, USA: Springer-Verlag, 268-278.
150. Schaefer K, Collatz GJ, Tans P, Denning AS, Baker I, Berry J, Prihodko L, Suits N, Philpott A. 2008. Combined Simple Biosphere/Carnegie-Ames-Stanford Approach terrestrial carbon cycle model. Journal of Geophysical Research-Biogeosciences 113: G3.
151. Schimel JP. 1995. Plant transport and methane production as controls on methane flux from arctic wet meadow tundra. Biogeochemistry 28: 183-200.
152. + nitrogen in situ: plants compete well for amino acid N. Ecology 77: 2142-2147.
153. Schuur EAG, Bockheim J, Canadell JG, Euskirchen E, Field CB, Goryachkin SV, Hagemann S, Kuhry P, Lafleur PM, Lee H et al. 2008. Vulnerability of permafrost carbon to climate change: implications for the global carbon cycle. BioScience 58: 701-714.
154. Segal AD, Sullivan PF. 2014. Identifying the sources and uncertainties of ecosystem respiration in Arctic tussock tundra. Biogeochemistry. doi: 10.1007/s10533-014-0017-8.
155. Shaver GR, Billings WD. 1975. Root production and root turnover in a wet tundra ecosystem, Barrow, Alaska. Ecology 56: 401-409.
156. Shaver GR, Billings WD. 1977. Effects of daylength and temperature on root elongation in tundra graminoids. Oecologia 28: 57-65.
157. Shaver GR, Chapin FS III. 1991. Production : biomass relationships and element cycling in contrasting arctic vegetation types. Ecological Monographs 61: 1-31.
158. Shaver GR, Chapin FS III. 1995. Long-term responses to factorial, NPK fertilizer treatment by Alaskan wet and moist tundra sedge species. Ecography 18: 259-275.
159. Shaver GR, Chapin FS, Billings WD. 1979. Ecotypic differentiation in Carex aquatilis on ice-wedge polygons in the Alaskan coastal tundra. Journal of Ecology 67: 1025-1046.
160. Shaver GR, Chapin FS III, Gartner BL. 1986. Factors limiting seasonal growth and peak biomass accumulation in Eriophorum vaginatum in Alaskan tussock tundra. Journal of Ecology 74: 257-278.
161. Shaver GR, Cutler JC. 1979. Vertical distribution of live vascular phytomass in cottongrass tussock tundra. Arctic and Alpine Research 11: 335-342.
162. Sloan VL. 2011. Plant roots in arctic ecosystems: stocks and dynamics, and their coupling to above-ground parameters. PhD Thesis, University of Sheffield, Sheffield, UK.
163. Sloan VL, Fletcher BJ, Press MC, Williams M, Phoenix GK. 2013. Leaf and fine root carbon stocks and turnover are coupled across Arctic ecosystems. Global Change Biology 19: 3668-3676.
164. Smith B, Prentice IC, Sykes MT. 2001. Representation of vegetation dynamics in the modelling of terrestrial ecosystems: comparing two contrasting approaches within European climate space. Global Ecology and Biogeography 10: 621-637.
165. Sorensen PL, Clemmensen KE, Michelsen A, Jonasson S, Strom L. 2008. Plant and microbial uptake and allocation of organic and inorganic nitrogen related to plant growth forms and soil conditions at two subarctic tundra sites in Sweden. Arctic, Antarctic, and Alpine Research 40: 171-180.
166. Stewart J, Freedman B. 1994. Biomass allocation in ten Saxifraga species in the high arctic. In: Svoboda J, Freedman B, eds. Ecology of a polar oasis: Alexandra Fiord, Ellesmere Island, Canada. New York, ON, Canada: Captus Press Inc., 123-136.
167. Ström L, Ekberg A, Mastepanov M, Christensen TR. 2003. The effect of vascular plants on carbon turnover and methane emissions from a tundra wetland. Global Change Biology 9: 1185-1192.
168. Ström L, Mastepanov M, Christensen TR. 2005. Species-specific effects of vascular plants on carbon turnover and methane emissions from wetlands. Biogeochemistry 75: 65-82.
169. Ström L, Tagesson T, Mastepanov M, Christensen TR. 2012. Presence of Eriophorum scheuchzeri enhances substrate availability and methane emission in an Arctic wetland. Soil Biology & Biochemistry 45: 61-70.
170. Sturm M. 2010. Arctic plants feel the heat. Scientific American 302: 66-73.
171. Sullivan PF, Arens SJT, Chimner RA, Welker JM. 2008. Temperature and microtopography interact to control carbon cycling in a high arctic fen. Ecosystems 11: 61-76.
172. Sullivan PF, Sommerkorn M, Rueth HM, Nadelhoffer KJ, Shaver GR, Welker JM. 2007. Climate and species affect fine root production with long-term fertilization in acidic tussock tundra near Toolik Lake, Alaska. Oecologia 153: 643-652.
173. Sullivan PF, Welker JM. 2005. Warming chambers stimulate early season growth of an arctic sedge: results of a minirhizotron field study. Oecologia 142: 616-626.
174. Tarnocai C, Canadell JG, Schuur EAG, Kuhry P, Mazhitova G, Zimov S. 2009. Soil organic carbon pools in the northern circumpolar permafrost region. Global Biogeochemical Cycles 23: 11.
175. Tieszen LL, Lewis MC, Miller PC, Mayo J, Chapin FS III, Oechel WC. 1981. An analysis of processes of primary production in tundra growth forms. In: Bliss LC, Heal OW, Moore JJ, eds. Tundra ecosystems: a comparative analysis. New York, NY, USA: Cambridge University Press, 647-684.
176. Tikhomirov BA, Shamurin VF, Aleksandrova VD. 1981. Phytomass and primary production of tundra communities, USSR. In: Bliss LC, Heal OW, Moore JJ, eds. Tundra ecosystems: a comparative analysis. New York, NY, USA: Cambridge University Press, 227-238.
177. Torn MS, Chapin FS. 1993. Environmental and biotic controls over methane flux from arctic tundra. Chemosphere 26: 357-368.
178. 15N added to high Arctic tundra to mimic increased inputs of atmospheric nitrogen released from a melting snowpack. Global Change Biology 11: 1640-1654.
179. Valenzuela-Estrada LR, Vera-Caraballo V, Ruth LE, Eissenstat DM. 2008. Root anatomy, morphology, and longevity among root orders in Vaccinium corymbosum (Ericaceae). American Journal of Botany 95: 1506-1514.
180. Vartapetian BB, Jackson MB. 1997. Plant adaptations to anaerobic stress. Annals of Botany 79: 3-20.
181. van der Wal R, van Lieshout SMJ, Loonen M. 2001. Herbivore impact on moss depth, soil temperature and arctic plant growth. Polar Biology 24: 29-32.
182. Walker DA, Raynolds MK, Daniels FJA, Einarsson E, Elvebakk A, Gould WA, Katenin AE, Kholod SS, Markon CJ, Melnikov ES et al. 2005. The Circumpolar Arctic vegetation map. Journal of Vegetation Science 16: 267-282.
183. Walker JKM, Egger KN, Henry GHR. 2008. Long-term experimental warming alters nitrogen-cycling communities but site factors remain the primary drivers of community structure in high arctic tundra soils. ISME Journal 2: 982-995.
184. Walker MD, Wahren CH, Hollister RD, Henry GHR, Ahlquist LE, Alatalo JM, Bret-Harte MS, Calef MP, Callaghan TV, Carroll AB et al. 2006. Plant community responses to experimental warming across the tundra biome. Proceedings of the National Academy of Sciences, USA 103: 1342-1346.
185. Wallén B. 1986. Above- and belowground dry mass of the three main vascular plants on hummocks on a sub-arctic peat bog. Oikos 46: 51-56.
186. Wania R, Ross I, Prentice IC. 2009. Integrating peatlands and permafrost into a dynamic global vegetation model: 2. Evaluation and sensitivity of vegetation and carbon cycle processes. Global Biogeochemical Cycles 23: GB3015.
187. Wania R, Ross I, Prentice IC. 2010. Implementation and evaluation of a new methane model within a dynamic global vegetation model: LPJ-WHyMe v1.3.1. Geoscientific Model Development 3: 565-584.
188. Warren JM, Hanson PJ, Iversen CM, Kumar J, Walker AP, Wullschleger SD. 2014. Root structural and functional dynamics in terrestrial biosphere models - Evaluation and recommendations. New Phytologist, in press. doi: 10.1111/nph.13034.
189. Washburn AL. 1973. Soil deformation resulting from some laboratory freeze-thaw experiments. Proceedings of the 2nd International Permafrost Conference. Washington, DC, USA: National Academy of Sciences, 757-762.
190. Webber PJ. 1977. Belowground tundra research - commentary. Arctic and Alpine Research 9: 105-111.
191. Webber PJ. 1978. Spatial and temporal variation of the vegetation and its productivity. In: Tieszen LL, ed. Vegetation and production ecology of an Alaskan arctic tundra. New York, NY, USA: Springer-Verlag New York Inc, 37-112.
192. Weintraub MN, Schimel JP. 2005. The seasonal dynamics of amino acids and other nutrients in Alaskan arctic tundra soils. Biogeochemistry 73: 359-380.
193. Wielgolaski FE. 1975. Primary productivity of alpine meadow communities. In: Wielgolaski FE, ed. Fennoscandian tundra ecosystems. New York, NY, USA: Springer-Verlag, 121-128.
194. Wielgolaski FE, Bliss LC, Svoboda J, Doyle G. 1981. Primary production of tundra. In: Bliss LC, Heal OW, Moore JJ, eds. Tundra ecosystems: a comparative analysis. New York, NY, USA: Cambridge University Press, 187-225.
195. van Wijk MT, Williams M, Gough L, Hobbie SE, Shaver GR. 2003. Luxury consumption of soil nutrients: a possible competitive strategy in above-ground and below-ground biomass allocation and root morphology for slow-growing arctic vegetation? Journal of Ecology 91: 664-676.
196. Withington JM, Reich PB, Oleksyn J, Eissenstat DM. 2006. Comparisons of structure and life span in roots and leaves among temperate trees. Ecological Monographs 76: 381-397.
197. Wookey PA, Aerts R, Bardgett RD, Baptist F, Brathen KA, Cornelissen JHC, Gough L, Hartley IP, Hopkins DW, Lavorel S et al. 2009. Ecosystem feedbacks and cascade processes: understanding their role in the responses of Arctic and alpine ecosystems to environmental change. Global Change Biology 15: 1153-1172.
198. Wu FZ, Yang WQ, Zhang J, Deng RJ. 2010. Fine root decomposition in two subalpine forests during the freeze-thaw season. Canadian Journal of Forest Research 40: 298-307.
199. Wullschleger SD, Epstein HE, Box EO, Euskirchen ES, Goswami S, Iversen CM, Kattge J, Norby RJ, van Bodegom PM, Xu X. 2014. Plant functional types in Earth System Models: past experiences and future directions for application of dynamic vegetation models in high-latitude ecosystems. Annals of Botany 114: 1-16.
200. Yi S, Li N, Xiang B, Wang X, Ye B, McGuire AD. 2013. Simulating the effects of alpine grassland vegetation cover on soil thermal dynamics for the Qinghai-Tibetan Plateau. Journal of Geophysical Research - Biogeosciences 118: 1186-1199.
201. Yi SH, McGuire AD, Kasischke E, Harden J, Manies K, Mack M, Turetsky M. 2010. A dynamic organic soil biogeochemical model for simulating the effects of wildfire on soil environmental conditions and carbon dynamics of black spruce forests. Journal of Geophysical Research-Biogeosciences 115: G04015.
202. Yuan F-M, Yi S-H, McGuire AD, Johnson KD, Liang J-J, Harden WJ, Kasischke E, Kurz WA. 2012. Assessment of historical boreal forest carbon dynamics in the Yukon River Basin: relative roles of climate warming and fire regime changes. Ecological Applications 22: 2091-2109.