{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T16:51:36Z","timestamp":1791305496719,"version":"4.3.1"},"reference-count":68,"publisher":"Springer Science and Business Media LLC","issue":"5","license":[{"start":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T00:00:00Z","timestamp":1791244800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T00:00:00Z","timestamp":1791244800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/501100024586","name":"Human Resource Development Centre, Council of Scientific And Industrial Research","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100024586","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100019180","name":"HORIZON EUROPE European Research Council","doi-asserted-by":"publisher","award":["GA 101075426"],"award-info":[{"award-number":["GA 101075426"]}],"id":[{"id":"10.13039\/100019180","id-type":"DOI","asserted-by":"publisher"}]},{"name":"DST-SERB India"},{"name":"DST-FIST India"},{"name":"DBT-IISc"},{"name":"MoEFCC, India"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Biogeochemistry"],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Large mammalian grazers have regulated biogeochemical cycles across 40% of the Earth\u2019s terrestrial surface for millennia. While grazer effects on organic matter entering the soil are becoming clear, their effect on the downstream microbial foraging through extracellular-enzymes (EE), which link processes at the cellular level to ecosystem-level flow of matter remain less clear. We assessed how large mammalian grazers influence microbial foraging in soil using two lines of evidence. First, utilising a 14-year-old long-term grazer-exclusion experiment in the Trans-Himalayan ecosystem, we find that activities of seven enzymes peak at different months during the growing-season in an asynchronous manner. Microbial deployment of enzymes responded to grazer-exclusion by shifting from nitrogen (N)-acquisition in spring to carbon (C)- and phosphorus (P)-acquisition in summer and autumn. Grazer exclusion increased peroxidase activity by 124.7% and \u03b2-glucosidase by 16.7%. About half of the variation in enzyme portfolios was explained by seasonal differences in grazer effects. Grazer-exclusion increased the variability in the enzyme portfolio, requiring the microbes to forage across a larger portion of the functional space. Structural equation models suggested that grazer-exclusion worsens the quality of biomass input to soil due to an 18% increase in acid-unhydrolysable recalcitrant C. As seen in other ecosystems, this decline in substrate quality raises microbial deployment of generic depolymerizers relative to the specialized enzymes which release assimilable end-products. As greater investment in generic depolymerizing enzymes may reduce microbial carbon use efficiency, it can affect the C cycle. Second, laboratory incubations supported that soil enzymes are indeed inducible\/repressible in response to experimental manipulation of biomass quality. Microbial responses appear predictable and generalisable across spatial scales, from laboratory-incubations to whole ecosystems. Hence, fostering microbial functions can enhance the scope and efficacy of rewilding and restoration for natural climate solutions.<\/jats:p>","DOI":"10.1007\/s10533-026-01378-0","type":"journal-article","created":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T15:44:15Z","timestamp":1791301455000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Large mammalian herbivores influence the deployment of microbial extracellular enzymes to control soil carbon cycling"],"prefix":"10.1007","volume":"169","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-9916-5651","authenticated-orcid":false,"given":"Shamik","family":"Roy","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4841-6748","authenticated-orcid":false,"given":"Sumanta","family":"Bagchi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,10,6]]},"reference":[{"key":"1378_CR1","doi-asserted-by":"publisher","first-page":"229","DOI":"10.1007\/978-3-642-14225-3_12","volume-title":"Soil enzymology","author":"SD Allison","year":"2010","unstructured":"Allison SD, Weintraub MN, Gartner TB, Waldrop MP (2010) Evolutionary-economic principles as regulators of soil enzyme production and ecosystem function. In: Shukla G, Varma A (eds) Soil enzymology. Springer, Berlin, Heidelberg, pp 229\u2013243"},{"key":"1378_CR2","doi-asserted-by":"publisher","first-page":"314","DOI":"10.1139\/m75-045","volume":"21","author":"JPE Anderson","year":"1975","unstructured":"Anderson JPE, Domsch KH (1975) Measurement of bacterial and fungal contributions to respiration of selected agricultural and forest soils. Can J Microbiol 21:314\u2013322. https:\/\/doi.org\/10.1139\/m75-045","journal-title":"Can J Microbiol"},{"key":"1378_CR3","doi-asserted-by":"publisher","first-page":"215","DOI":"10.1016\/0038-0717(78)90099-8","volume":"10","author":"JPE Anderson","year":"1978","unstructured":"Anderson JPE, Domsch KH (1978) A physiological method for the quantitative measurement of microbial biomass in soils. Soil Biol Biochem 10:215\u2013221. https:\/\/doi.org\/10.1016\/0038-0717(78)90099-8","journal-title":"Soil Biol Biochem"},{"key":"1378_CR4","doi-asserted-by":"publisher","first-page":"1075","DOI":"10.1007\/s00442-010-1690-5","volume":"164","author":"S Bagchi","year":"2010","unstructured":"Bagchi S, Ritchie ME (2010a) Herbivore effects on above- and belowground plant production and soil nitrogen availability in the Trans-Himalayan shrub-steppes. Oecologia 164:1075\u20131082. https:\/\/doi.org\/10.1007\/s00442-010-1690-5","journal-title":"Oecologia"},{"key":"1378_CR5","doi-asserted-by":"publisher","first-page":"959","DOI":"10.1111\/j.1461-0248.2010.01486.x","volume":"13","author":"S Bagchi","year":"2010","unstructured":"Bagchi S, Ritchie ME (2010b) Introduced grazers can restrict potential soil carbon sequestration through impacts on plant community composition. Ecol Lett 13:959\u2013968. https:\/\/doi.org\/10.1111\/j.1461-0248.2010.01486.x","journal-title":"Ecol Lett"},{"key":"1378_CR6","doi-asserted-by":"publisher","first-page":"119","DOI":"10.1111\/j.1600-0706.2010.18932.x","volume":"120","author":"S Bagchi","year":"2011","unstructured":"Bagchi S, Ritchie ME (2011) Herbivory and plant tolerance: experimental tests of alternative hypotheses involving non-substitutable resources. Oikos 120:119\u2013127. https:\/\/doi.org\/10.1111\/j.1600-0706.2010.18932.x","journal-title":"Oikos"},{"key":"1378_CR7","doi-asserted-by":"publisher","first-page":"199","DOI":"10.1016\/j.agee.2017.01.033","volume":"239","author":"S Bagchi","year":"2017","unstructured":"Bagchi S, Roy S, Maitra A, Sran RS (2017) Herbivores suppress soil microbes to influence carbon sequestration in the grazing ecosystem of the Trans-Himalaya. Agric Ecosyst Environ 239:199\u2013206. https:\/\/doi.org\/10.1016\/j.agee.2017.01.033","journal-title":"Agric Ecosyst Environ"},{"key":"1378_CR8","doi-asserted-by":"publisher","DOI":"10.1002\/eap.3091","author":"P Baidya","year":"2025","unstructured":"Baidya P, Roy S, Karapurkar J, Bagchi S (2025) Replacing native grazers with livestock influences arthropods to have implications for ecosystem functions and disease. Ecol Appl. https:\/\/doi.org\/10.1002\/eap.3091","journal-title":"Ecol Appl"},{"key":"1378_CR9","doi-asserted-by":"publisher","first-page":"1023","DOI":"10.1016\/S0038-0717(97)00030-8","volume":"29","author":"T Beck","year":"1997","unstructured":"Beck T, Joergensen RG, Kandeler E et al (1997) An inter-laboratory comparison of ten different ways of measuring soil microbial biomass C. Soil Biol Biochem 29:1023\u20131032. https:\/\/doi.org\/10.1016\/S0038-0717(97)00030-8","journal-title":"Soil Biol Biochem"},{"key":"1378_CR10","doi-asserted-by":"publisher","first-page":"641","DOI":"10.1023\/A:1018430201230","volume":"11","author":"GE Belovsky","year":"1997","unstructured":"Belovsky GE (1997) Optimal foraging and community structure: the allometry of herbivore food selection and competition. Evol Ecol 11:641\u2013672. https:\/\/doi.org\/10.1023\/A:1018430201230","journal-title":"Evol Ecol"},{"key":"1378_CR11","doi-asserted-by":"publisher","first-page":"1419","DOI":"10.1016\/j.soilbio.2004.12.012","volume":"37","author":"REJ Boerner","year":"2005","unstructured":"Boerner REJ, Brinkman JA, Smith A (2005) Seasonal variations in enzyme activity and organic carbon in soil of a burned and unburned hardwood forest. Soil Biol Biochem 37:1419\u20131426. https:\/\/doi.org\/10.1016\/j.soilbio.2004.12.012","journal-title":"Soil Biol Biochem"},{"key":"1378_CR12","doi-asserted-by":"publisher","first-page":"229","DOI":"10.1111\/1365-2745.12507","volume":"104","author":"MA Bradford","year":"2016","unstructured":"Bradford MA, Berg B, Maynard DS et al (2016) Understanding the dominant controls on litter decomposition. J Ecol 104:229\u2013238. https:\/\/doi.org\/10.1111\/1365-2745.12507","journal-title":"J Ecol"},{"key":"1378_CR13","doi-asserted-by":"publisher","DOI":"10.1201\/9780203904039","volume-title":"Enzymes in the environment: activity, ecology, and applications","author":"R Burns","year":"2002","unstructured":"Burns R, Dick R (2002) Enzymes in the environment: activity, ecology, and applications. CRC Press"},{"key":"1378_CR14","doi-asserted-by":"publisher","first-page":"10892","DOI":"10.1038\/srep10892","volume":"5","author":"W Chen","year":"2015","unstructured":"Chen W, Huang D, Liu N et al (2015) Improved grazing management may increase soil carbon sequestration in temperate steppe. Sci Rep 5:10892. https:\/\/doi.org\/10.1038\/srep10892","journal-title":"Sci Rep"},{"key":"1378_CR15","doi-asserted-by":"publisher","first-page":"202","DOI":"10.1111\/ele.12722","volume":"20","author":"LC Cline","year":"2017","unstructured":"Cline LC, Zak DR, Upchurch RA et al (2017) Soil microbial communities and elk foraging intensity: implications for soil biogeochemical cycling in the sagebrush steppe. Ecol Lett 20:202\u2013211. https:\/\/doi.org\/10.1111\/ele.12722","journal-title":"Ecol Lett"},{"key":"1378_CR16","doi-asserted-by":"publisher","first-page":"989","DOI":"10.1038\/s41561-019-0484-6","volume":"12","author":"MF Cotrufo","year":"2019","unstructured":"Cotrufo MF, Ranalli MG, Haddix ML et al (2019) Soil carbon storage informed by particulate and mineral-associated organic matter. Nat Geosci 12:989\u2013994. https:\/\/doi.org\/10.1038\/s41561-019-0484-6","journal-title":"Nat Geosci"},{"key":"1378_CR17","doi-asserted-by":"publisher","first-page":"1369","DOI":"10.1016\/j.scitotenv.2018.01.012","volume":"628\u2013629","author":"I Djukic","year":"2018","unstructured":"Djukic I, Kepfer-Rojas S, Schmidt IK et al (2018) Early stage litter decomposition across biomes. Sci Total Environ 628\u2013629:1369\u20131394. https:\/\/doi.org\/10.1016\/j.scitotenv.2018.01.012","journal-title":"Sci Total Environ"},{"key":"1378_CR18","doi-asserted-by":"publisher","first-page":"107111","DOI":"10.1016\/j.agee.2020.107111","volume":"303","author":"S Dong","year":"2020","unstructured":"Dong S, Li Y, Ganjurjav H et al (2020) Grazing promoted soil microbial functional genes for regulating C and N cycling in alpine meadow of the Qinghai-Tibetan plateau. Agric Ecosyst Environ 303:107111. https:\/\/doi.org\/10.1016\/j.agee.2020.107111","journal-title":"Agric Ecosyst Environ"},{"key":"1378_CR19","doi-asserted-by":"publisher","first-page":"1034","DOI":"10.1126\/science.1153213","volume":"320","author":"PG Falkowski","year":"2008","unstructured":"Falkowski PG, Fenchel T, Delong EF (2008) The microbial engines that drive earth\u2019s biogeochemical cycles. Science 320:1034\u20131039. https:\/\/doi.org\/10.1126\/science.1153213","journal-title":"Science"},{"key":"1378_CR20","doi-asserted-by":"publisher","first-page":"97","DOI":"10.1111\/grs.70000","volume":"71","author":"J Feng","year":"2025","unstructured":"Feng J, Sun Z, Tang S et al (2025) Effect of grazing on soil enzyme activities in grassland ecosystems: a meta-analysis. Grassl Sci 71:97\u2013105. https:\/\/doi.org\/10.1111\/grs.70000","journal-title":"Grassl Sci"},{"key":"1378_CR21","first-page":"329","volume-title":"Gastrointestinal microbiology","author":"CW Forsberg","year":"1997","unstructured":"Forsberg CW, Cheng K-J, White BA (1997) Polysaccharide degradation in the rumen and large intestine. In: Mackie R, White B (eds) Gastrointestinal microbiology. Springer Netherlands, Dordrecht, pp 329\u2013381"},{"key":"1378_CR22","doi-asserted-by":"publisher","first-page":"14","DOI":"10.1016\/j.apsoil.2013.12.003","volume":"76","author":"G Francini","year":"2014","unstructured":"Francini G, Liiri M, M\u00e4nnist\u00f6 M et al (2014) Response to reindeer grazing removal depends on soil characteristics in low Arctic meadows. Appl Soil Ecol 76:14\u201325. https:\/\/doi.org\/10.1016\/j.apsoil.2013.12.003","journal-title":"Appl Soil Ecol"},{"key":"1378_CR23","doi-asserted-by":"publisher","first-page":"513","DOI":"10.2307\/1313313","volume":"48","author":"DA Frank","year":"1998","unstructured":"Frank DA, McNaughton SJ, Tracy BF (1998) The ecology of the Earth\u2019s grazing ecosystems. Bioscience 48:513\u2013521. https:\/\/doi.org\/10.2307\/1313313","journal-title":"Bioscience"},{"key":"1378_CR24","doi-asserted-by":"publisher","first-page":"3201","DOI":"10.1890\/09-1831.1","volume":"91","author":"DA Frank","year":"2010","unstructured":"Frank DA, Pontes AW, Maine EM et al (2010) Grassland root communities: species distributions and how they are linked to aboveground abundance. Ecology 91:3201\u20133209. https:\/\/doi.org\/10.1890\/09-1831.1","journal-title":"Ecology"},{"key":"1378_CR202","doi-asserted-by":"publisher","first-page":"1387","DOI":"10.1016\/j.soilbio.2011.03.017","volume":"43","author":"DP German","year":"2011","unstructured":"German DP, Weintraub MN, Grandy AS, et al (2011) Optimization of hydrolytic and oxidative enzyme methods for ecosystem studies. Soil Biol Biochem 43:1387\u20131397. https:\/\/doi.org\/10.1016\/j.soilbio.2011.03.017","journal-title":"Soil Biol Biochem"},{"key":"1378_CR25","doi-asserted-by":"publisher","first-page":"904","DOI":"10.1126\/science.adu0703","volume":"389","author":"C Geremia","year":"2025","unstructured":"Geremia C, Hamilton EW, Merkle JA (2025) Yellowstone\u2019s free-moving large bison herds provide a glimpse of their past ecosystem function. Science 389:904\u2013908. https:\/\/doi.org\/10.1126\/science.adu0703","journal-title":"Science"},{"key":"1378_CR26","doi-asserted-by":"publisher","first-page":"9641","DOI":"10.1073\/pnas.1707186114","volume":"114","author":"TJ Hammer","year":"2017","unstructured":"Hammer TJ, Janzen DH, Hallwachs W et al (2017) Caterpillars lack a resident gut microbiome. Proc Natl Acad Sci U S A 114:9641\u20139646. https:\/\/doi.org\/10.1073\/pnas.1707186114","journal-title":"Proc Natl Acad Sci U S A"},{"key":"1378_CR27","doi-asserted-by":"publisher","first-page":"695","DOI":"10.2307\/3802368","volume":"60","author":"NT Hobbs","year":"1996","unstructured":"Hobbs NT (1996) Modification of ecosystems by ungulates. J Wildl Manage 60:695\u2013713. https:\/\/doi.org\/10.2307\/3802368","journal-title":"J Wildl Manage"},{"key":"1378_CR28","doi-asserted-by":"publisher","first-page":"400","DOI":"10.1111\/J.1574-6941.2008.00524.X","volume":"65","author":"P Iovieno","year":"2008","unstructured":"Iovieno P, B\u00e5\u00e5th E (2008) Effect of drying and rewetting on bacterial growth rates in soil. FEMS Microbiol Ecol 65:400\u2013407. https:\/\/doi.org\/10.1111\/J.1574-6941.2008.00524.X","journal-title":"FEMS Microbiol Ecol"},{"key":"1378_CR29","doi-asserted-by":"publisher","first-page":"144","DOI":"10.1016\/j.apsoil.2016.05.018","volume":"107","author":"B Khalili","year":"2016","unstructured":"Khalili B, Ogunseitan OA, Goulden ML, Allison SD (2016) Interactive effects of precipitation manipulation and nitrogen addition on soil properties in California grassland and shrubland. Appl Soil Ecol 107:144\u2013153. https:\/\/doi.org\/10.1016\/j.apsoil.2016.05.018","journal-title":"Appl Soil Ecol"},{"key":"1378_CR30","doi-asserted-by":"publisher","first-page":"323","DOI":"10.1007\/s10533-018-0448-8","volume":"138","author":"JA Kristensen","year":"2018","unstructured":"Kristensen JA, Metcalfe DB, Rousk J (2018) The biogeochemical consequences of litter transformation by insect herbivory in the Subarctic: a microcosm simulation experiment. Biogeochemistry 138:323\u2013336. https:\/\/doi.org\/10.1007\/s10533-018-0448-8","journal-title":"Biogeochemistry"},{"key":"1378_CR31","doi-asserted-by":"publisher","first-page":"117","DOI":"10.1016\/j.tree.2021.09.006","volume":"37","author":"JA Kristensen","year":"2022","unstructured":"Kristensen JA, Svenning J-C, Georgiou K, Malhi Y (2022) Can large herbivores enhance ecosystem carbon persistence? Trends Ecol Evol 37:117\u2013128. https:\/\/doi.org\/10.1016\/j.tree.2021.09.006","journal-title":"Trends Ecol Evol"},{"key":"1378_CR32","doi-asserted-by":"publisher","first-page":"573","DOI":"10.1111\/2041-210X.12512","volume":"7","author":"JS Lefcheck","year":"2016","unstructured":"Lefcheck JS (2016) piecewiseSEM: piecewise structural equation modelling in r for ecology, evolution, and systematics. Methods Ecol Evol 7:573\u2013579. https:\/\/doi.org\/10.1111\/2041-210X.12512","journal-title":"Methods Ecol Evol"},{"key":"1378_CR33","doi-asserted-by":"publisher","first-page":"7871","DOI":"10.1073\/pnas.1915769117","volume":"117","author":"EJ Lundgren","year":"2020","unstructured":"Lundgren EJ, Ramp D, Rowan J et al (2020) Introduced herbivores restore Late Pleistocene ecological functions. Proc Natl Acad Sci U S A 117:7871\u20137878. https:\/\/doi.org\/10.1073\/pnas.1915769117","journal-title":"Proc Natl Acad Sci U S A"},{"key":"1378_CR34","doi-asserted-by":"publisher","first-page":"246","DOI":"10.1038\/s41559-025-02955-6","volume":"10","author":"AS MacDougall","year":"2026","unstructured":"MacDougall AS, Vanzant B, Sulik J et al (2026) The global extent of the grassland biome and implications for the terrestrial carbon sink. Nat Ecol Evol 10:246\u2013257. https:\/\/doi.org\/10.1038\/s41559-025-02955-6","journal-title":"Nat Ecol Evol"},{"key":"1378_CR35","doi-asserted-by":"publisher","first-page":"1247","DOI":"10.3389\/fmicb.2016.01247","volume":"7","author":"AA Malik","year":"2016","unstructured":"Malik AA, Chowdhury S, Schlager V et al (2016) Soil fungal:bacterial ratios are linked to altered carbon cycling. Front Microbiol 7:1247. https:\/\/doi.org\/10.3389\/fmicb.2016.01247","journal-title":"Front Microbiol"},{"key":"1378_CR36","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-017-01418-8","author":"O Margalef","year":"2017","unstructured":"Margalef O, Sardans J, Fern\u00e1ndez-Mart\u00ednez M et al (2017) Global patterns of phosphatase activity in natural soils. Sci Rep. https:\/\/doi.org\/10.1038\/s41598-017-01418-8","journal-title":"Sci Rep"},{"key":"1378_CR37","doi-asserted-by":"publisher","first-page":"e0268461","DOI":"10.1371\/journal.pone.0268461","volume":"17","author":"K Min","year":"2022","unstructured":"Min K, Choi M-A (2022) Resource landscape, microbial activity, and community composition under wintering crane activities in the demilitarized zone, South Korea. PLoS One 17:e0268461. https:\/\/doi.org\/10.1371\/journal.pone.0268461","journal-title":"PLoS One"},{"key":"1378_CR38","doi-asserted-by":"publisher","first-page":"e01777","DOI":"10.1002\/ecs2.1777","volume":"8","author":"LC Moorhead","year":"2017","unstructured":"Moorhead LC, Souza L, Habeck CW et al (2017) Small mammal activity alters plant community composition and microbial activity in an old-field ecosystem. Ecosphere 8:e01777. https:\/\/doi.org\/10.1002\/ecs2.1777","journal-title":"Ecosphere"},{"key":"1378_CR39","doi-asserted-by":"publisher","first-page":"e2211317119","DOI":"10.1073\/pnas.2211317119","volume":"119","author":"DGT Naidu","year":"2022","unstructured":"Naidu DGT, Roy S, Bagchi S (2022) Loss of grazing by large mammalian herbivores can destabilize the soil carbon pool. Proc Natl Acad Sci U S A 119:e2211317119. https:\/\/doi.org\/10.1073\/pnas.2211317119","journal-title":"Proc Natl Acad Sci U S A"},{"key":"1378_CR40","doi-asserted-by":"publisher","first-page":"642","DOI":"10.1111\/1365-2664.14100","volume":"59","author":"KO Reinhart","year":"2022","unstructured":"Reinhart KO, Sanni Worogo HS, Rinella MJ (2022) Ruminating on the science of carbon ranching. J Appl Ecol 59:642\u2013648. https:\/\/doi.org\/10.1111\/1365-2664.14100","journal-title":"J Appl Ecol"},{"key":"1378_CR41","doi-asserted-by":"publisher","first-page":"e1400103","DOI":"10.1126\/sciadv.1400103","volume":"1","author":"WJ Ripple","year":"2015","unstructured":"Ripple WJ, Newsome TM, Wolf C et al (2015) Collapse of the world\u2019s largest herbivores. Sci Adv 1:e1400103. https:\/\/doi.org\/10.1126\/sciadv.1400103","journal-title":"Sci Adv"},{"key":"1378_CR42","doi-asserted-by":"publisher","first-page":"165","DOI":"10.2307\/176872","volume":"79","author":"ME Ritchie","year":"1998","unstructured":"Ritchie ME, Tilman D, Knops JMH (1998) Herbivore effects on plant and nitrogen dynamics in oak savanna. Ecology 79:165\u2013177. https:\/\/doi.org\/10.2307\/176872","journal-title":"Ecology"},{"key":"1378_CR43","doi-asserted-by":"publisher","first-page":"4150","DOI":"10.1111\/gcb.13296","volume":"22","author":"K Rousk","year":"2016","unstructured":"Rousk K, Michelsen A, Rousk J (2016) Microbial control of soil organic matter mineralization responses to labile carbon in subarctic climate change treatments. Glob Chang Biol 22:4150\u20134161. https:\/\/doi.org\/10.1111\/gcb.13296","journal-title":"Glob Chang Biol"},{"key":"1378_CR44","doi-asserted-by":"publisher","first-page":"202","DOI":"10.1016\/j.soilbio.2006.07.021","volume":"39","author":"P Rovira","year":"2007","unstructured":"Rovira P, Ram\u00f3n Vallejo V (2007) Labile, recalcitrant, and inert organic matter in Mediterranean forest soils. Soil Biol Biochem 39:202\u2013215. https:\/\/doi.org\/10.1016\/j.soilbio.2006.07.021","journal-title":"Soil Biol Biochem"},{"key":"1378_CR45","doi-asserted-by":"publisher","first-page":"109","DOI":"10.1016\/S0016-7061(01)00143-4","volume":"107","author":"P Rovira","year":"2002","unstructured":"Rovira P, Vallejo VR (2002) Labile and recalcitrant pools of carbon and nitrogen in organic matter decomposing at different depths in soil: an acid hydrolysis approach. Geoderma 107:109\u2013141. https:\/\/doi.org\/10.1016\/S0016-7061(01)00143-4","journal-title":"Geoderma"},{"key":"1378_CR46","doi-asserted-by":"publisher","first-page":"976","DOI":"10.1007\/s10021-021-00696-8","volume":"25","author":"S Roy","year":"2022","unstructured":"Roy S, Bagchi S (2022) Large mammalian herbivores and the paradox of soil carbon in grazing ecosystems: role of microbial decomposers and their enzymes. Ecosystems 25:976\u2013988. https:\/\/doi.org\/10.1007\/s10021-021-00696-8","journal-title":"Ecosystems"},{"key":"1378_CR47","doi-asserted-by":"publisher","first-page":"109225","DOI":"10.1016\/j.soilbio.2023.109225","volume":"187","author":"S Roy","year":"2023","unstructured":"Roy S, Karapurkar J, Baidya P et al (2023a) Community composition, and not species richness, of microbes influences decomposer functional diversity in soil. Soil Biol Biochem 187:109225. https:\/\/doi.org\/10.1016\/j.soilbio.2023.109225","journal-title":"Soil Biol Biochem"},{"key":"1378_CR48","doi-asserted-by":"publisher","first-page":"2141","DOI":"10.1111\/gcb.16600","volume":"29","author":"S Roy","year":"2023","unstructured":"Roy S, Naidu DGT, Bagchi S (2023b) Functional substitutability of native herbivores by livestock for soil carbon stock is mediated by microbial decomposers. Glob Chang Biol 29:2141\u20132155. https:\/\/doi.org\/10.1111\/gcb.16600","journal-title":"Glob Chang Biol"},{"key":"1378_CR203","doi-asserted-by":"publisher","first-page":"1309","DOI":"10.1016\/S0038-0717(02)00074-3","volume":"34","author":"K Saiya-Cork","year":"2002","unstructured":"Saiya-Cork K, Sinsabaugh R, Zak D (2002) The eff ects of long term nitrogen deposition on extracellular enzyme activity in an Acer saccharum forest soil. Soil Biol Biochem 34:1309\u20131315. https:\/\/doi.org\/10.1016\/S0038-0717(02)00074-3","journal-title":"Soil Biol Biochem"},{"key":"1378_CR49","doi-asserted-by":"publisher","first-page":"1052","DOI":"10.1890\/03-0354","volume":"85","author":"M Sankaran","year":"2004","unstructured":"Sankaran M, Augustine DJ (2004) Large herbivores suppress decomposer abundance in a semiarid grazing ecosystem. Ecology 85:1052\u20131061. https:\/\/doi.org\/10.1890\/03-0354","journal-title":"Ecology"},{"key":"1378_CR50","doi-asserted-by":"publisher","first-page":"74","DOI":"10.1016\/j.apsoil.2018.08.013","volume":"132","author":"M Santalahti","year":"2018","unstructured":"Santalahti M, Sun H, Sieti\u00f6 OM et al (2018) Reindeer grazing alter soil fungal community structure and litter decomposition related enzyme activities in boreal coniferous forests in Finnish Lapland. Appl Soil Ecol 132:74\u201382. https:\/\/doi.org\/10.1016\/j.apsoil.2018.08.013","journal-title":"Appl Soil Ecol"},{"key":"1378_CR51","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1007\/s10533-022-01009-4","volume":"163","author":"KJ Schaap","year":"2023","unstructured":"Schaap KJ, Fuchslueger L, Quesada CA et al (2023) Seasonal fluctuations of extracellular enzyme activities are related to the biogeochemical cycling of C, N and P in a tropical terra-firme forest. Biogeochemistry 163:1\u201315. https:\/\/doi.org\/10.1007\/s10533-022-01009-4","journal-title":"Biogeochemistry"},{"key":"1378_CR52","doi-asserted-by":"publisher","first-page":"108677","DOI":"10.1016\/j.soilbio.2022.108677","volume":"169","author":"J Schimel","year":"2022","unstructured":"Schimel J, Weintraub MN, Moorhead D (2022) Estimating microbial carbon use efficiency in soil: isotope-based and enzyme-based methods measure fundamentally different aspects of microbial resource use. Soil Biol Biochem 169:108677. https:\/\/doi.org\/10.1016\/j.soilbio.2022.108677","journal-title":"Soil Biol Biochem"},{"key":"1378_CR53","doi-asserted-by":"publisher","first-page":"344","DOI":"10.1007\/s10021-013-9715-7","volume":"17","author":"OJ Schmitz","year":"2014","unstructured":"Schmitz OJ, Raymond PA, Estes JA et al (2014) Animating the carbon cycle. Ecosystems 17:344\u2013359. https:\/\/doi.org\/10.1007\/s10021-013-9715-7","journal-title":"Ecosystems"},{"key":"1378_CR54","doi-asserted-by":"publisher","first-page":"eaar3213","DOI":"10.1126\/science.aar3213","volume":"362","author":"OJ Schmitz","year":"2018","unstructured":"Schmitz OJ, Wilmers CC, Leroux SJ et al (2018) Animals and the zoogeochemistry of the carbon cycle. Science 362:eaar3213. https:\/\/doi.org\/10.1126\/science.aar3213","journal-title":"Science"},{"key":"1378_CR55","doi-asserted-by":"publisher","first-page":"263","DOI":"10.1006\/anae.1996.0036","volume":"2","author":"LB Selinger","year":"1996","unstructured":"Selinger LB, Forsberg CW, Cheng K-J (1996) The rumen: a unique source of enzymes for enhancing livestock production. Anaerobe 2:263\u2013284. https:\/\/doi.org\/10.1006\/anae.1996.0036","journal-title":"Anaerobe"},{"key":"1378_CR56","doi-asserted-by":"publisher","first-page":"363","DOI":"10.1890\/08-1034.1","volume":"90","author":"B Shipley","year":"2009","unstructured":"Shipley B (2009) Confirmatory path analysis in a generalized multilevel context. Ecology 90:363\u2013368. https:\/\/doi.org\/10.1890\/08-1034.1","journal-title":"Ecology"},{"key":"1378_CR57","doi-asserted-by":"publisher","first-page":"391","DOI":"10.1016\/j.soilbio.2009.10.014","volume":"42","author":"RL Sinsabaugh","year":"2010","unstructured":"Sinsabaugh RL (2010) Phenol oxidase, peroxidase and organic matter dynamics of soil. Soil Biol Biochem 42:391\u2013404. https:\/\/doi.org\/10.1016\/j.soilbio.2009.10.014","journal-title":"Soil Biol Biochem"},{"key":"1378_CR58","doi-asserted-by":"publisher","first-page":"930","DOI":"10.1111\/ele.12113","volume":"16","author":"RL Sinsabaugh","year":"2013","unstructured":"Sinsabaugh RL, Manzoni S, Moorhead DL, Richter A (2013) Carbon use efficiency of microbial communities: stoichiometry, methodology and modelling. Ecol Lett 16:930\u2013939. https:\/\/doi.org\/10.1111\/ele.12113","journal-title":"Ecol Lett"},{"key":"1378_CR59","doi-asserted-by":"publisher","first-page":"172","DOI":"10.1890\/15-2110.1","volume":"86","author":"RL Sinsabaugh","year":"2016","unstructured":"Sinsabaugh RL, Turner BL, Talbot JM et al (2016) Stoichiometry of microbial carbon use efficiency in soils. Ecol Monogr 86:172\u2013189. https:\/\/doi.org\/10.1890\/15-2110.1","journal-title":"Ecol Monogr"},{"key":"1378_CR60","doi-asserted-by":"publisher","first-page":"1274","DOI":"10.1038\/s41564-018-0225-4","volume":"3","author":"LM Solden","year":"2018","unstructured":"Solden LM, Naas AE, Roux S et al (2018) Interspecies cross-feeding orchestrates carbon degradation in the rumen ecosystem. Nat Microbiol 3:1274\u20131284. https:\/\/doi.org\/10.1038\/s41564-018-0225-4","journal-title":"Nat Microbiol"},{"key":"1378_CR61","doi-asserted-by":"publisher","first-page":"18","DOI":"10.1007\/s10021-011-9491-1","volume":"15","author":"S Stark","year":"2012","unstructured":"Stark S, Eskelinen A, M\u00e4nnist\u00f6 MK (2012) Regulation of microbial community composition and activity by soil nutrient availability, soil pH, and herbivory in the tundra. Ecosystems 15:18\u201333. https:\/\/doi.org\/10.1007\/s10021-011-9491-1","journal-title":"Ecosystems"},{"key":"1378_CR62","doi-asserted-by":"publisher","first-page":"313","DOI":"10.1007\/s10530-009-9624-z","volume":"12","author":"ML Stritar","year":"2010","unstructured":"Stritar ML, Schweitzer JA, Hart SC, Bailey JK (2010) Introduced ungulate herbivore alters soil processes after fire. Biol Invasions 12:313\u2013324. https:\/\/doi.org\/10.1007\/s10530-009-9624-z","journal-title":"Biol Invasions"},{"key":"1378_CR63","doi-asserted-by":"publisher","DOI":"10.1007\/s11104-019-04380-8","author":"J Sun","year":"2019","unstructured":"Sun J, Zhan T, Liu M et al (2019) Verification of the biomass transfer hypothesis under moderate grazing across the Tibetan plateau: a meta-analysis. Plant Soil. https:\/\/doi.org\/10.1007\/s11104-019-04380-8","journal-title":"Plant Soil"},{"key":"1378_CR64","doi-asserted-by":"publisher","first-page":"141","DOI":"10.1016\/0168-1656(93)90035-L","volume":"30","author":"P Susmel","year":"1993","unstructured":"Susmel P, Stefanon B (1993) Aspects of lignin degradation by rumen microorganisms. J Biotechnol 30:141\u2013148. https:\/\/doi.org\/10.1016\/0168-1656(93)90035-L","journal-title":"J Biotechnol"},{"key":"1378_CR65","doi-asserted-by":"publisher","first-page":"1631","DOI":"10.1111\/j.1365-2486.2008.01819.x","volume":"15","author":"MD Wallenstein","year":"2009","unstructured":"Wallenstein MD, Mcmahon SK, Schimel JP (2009) Seasonal variation in enzyme activities and temperature sensitivities in Arctic tundra soils. Glob Chang Biol 15:1631\u20131639. https:\/\/doi.org\/10.1111\/j.1365-2486.2008.01819.x","journal-title":"Glob Chang Biol"},{"key":"1378_CR66","doi-asserted-by":"publisher","DOI":"10.1002\/fee.70022","author":"L Yahdjian","year":"2026","unstructured":"Yahdjian L, Campana S, Tognetti PM et al (2026) Insights on global rangeland ecosystem services shaped by grazing and fertilization. Front Ecol Environ. https:\/\/doi.org\/10.1002\/fee.70022","journal-title":"Front Ecol Environ"}],"container-title":["Biogeochemistry"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/article\/10.1007\/s10533-026-01378-0","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s10533-026-01378-0.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s10533-026-01378-0.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T16:00:16Z","timestamp":1791302416000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1007\/s10533-026-01378-0"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,10,6]]},"references-count":68,"journal-issue":{"issue":"5","published-online":{"date-parts":[[2026,10]]}},"alternative-id":["1378"],"URL":"https:\/\/doi.org\/10.1007\/s10533-026-01378-0","relation":{},"ISSN":["1573-515X"],"issn-type":[{"value":"1573-515X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,10,6]]},"assertion":[{"value":"30 May 2026","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"24 September 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"6 October 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors have not disclosed any competing interests.","order":1,"name":"Ethics","label":"Competing interests","group":{"name":"EthicsHeading","label":"Declarations"}}],"article-number":"62"}}