Interactions among nutrients govern the global grassland biomass-precipitation relationship

Philip A Fay; Laureano A Gherardi; Laura Yahdjian; Peter B Adler; Jonathan D Bakker; Siddharth Bharath; Elizabeth T Borer; W Stanley Harpole; Erika Hersch-Green; Travis E Huxman; Andrew S MacDougall; Anita C Risch; Eric W Seabloom; Sumanta Bagchi; Isabel C Barrio; Lori Biederman; Yvonne M Buckley; Miguel N Bugalho; Maria C Caldeira; Jane A Catford; QingQing Chen; Elsa E Cleland; Scott L Collins; Pedro Daleo; Christopher R Dickman; Ian Donohue; Mary E DuPre; Nico Eisenhauer; Anu Eskelinen; Nicole Hagenah; Yann Hautier; Robert W Heckman; Ingibjörg S Jónsdóttir; Johannes M H Knops; Ramesh Laungani; Jason P Martina; Rebecca L McCulley; John W Morgan; Harry Olde Venterink; Pablo L Peri; Sally A Power; Xavier Raynaud; Zhengwei Ren; Christiane Roscher; Melinda D Smith; Marie Spohn; Carly J Stevens; Michelle J Tedder; Risto Virtanen; Glenda M Wardle; George R Wheeler

doi:10.1073/pnas.2410748122

Interactions among nutrients govern the global grassland biomass-precipitation relationship

Philip A Fay^*, Laureano A Gherardi^*, Laura Yahdjian^*, Peter B Adler, Jonathan D Bakker, Siddharth Bharath, Elizabeth T Borer, W Stanley Harpole, Erika Hersch-Green, Travis E Huxman, Andrew S MacDougall, Anita C Risch, Eric W Seabloom, Sumanta Bagchi, Isabel C Barrio, Lori Biederman, Yvonne M Buckley, Miguel N Bugalho, Maria C Caldeira, Jane A CatfordQingQing Chen, Elsa E Cleland, Scott L Collins, Pedro Daleo, Christopher R Dickman, Ian Donohue, Mary E DuPre, Nico Eisenhauer, Anu Eskelinen, Nicole Hagenah, Yann Hautier, Robert W Heckman, Ingibjörg S Jónsdóttir, Johannes M H Knops, Ramesh Laungani, Jason P Martina, Rebecca L McCulley, John W Morgan, Harry Olde Venterink, Pablo L Peri, Sally A Power, Xavier Raynaud, Zhengwei Ren, Christiane Roscher, Melinda D Smith, Marie Spohn, Carly J Stevens, Michelle J Tedder, Risto Virtanen, Glenda M Wardle, George R Wheeler

^*Corresponding author for this work

Research output: Contribution to journal › Article › Academic › peer-review

Abstract

Ecosystems are experiencing changing global patterns of mean annual precipitation (MAP) and enrichment with multiple nutrients that potentially colimit plant biomass production. In grasslands, mean aboveground plant biomass is closely related to MAP, but how this relationship changes after enrichment with multiple nutrients remains unclear. We hypothesized the global biomass-MAP relationship becomes steeper with an increasing number of added nutrients, with increases in steepness corresponding to the form of interaction among added nutrients and with increased mediation by changes in plant community diversity. We measured aboveground plant biomass production and species diversity in 71 grasslands on six continents representing the global span of grassland MAP, diversity, management, and soils. We fertilized all sites with nitrogen, phosphorus, and potassium with micronutrients in all combinations to identify which nutrients limited biomass at each site. As hypothesized, fertilizing with one, two, or three nutrients progressively steepened the global biomass-MAP relationship. The magnitude of the increase in steepness corresponded to whether sites were not limited by nitrogen or phosphorus, were limited by either one, or were colimited by both in additive, or synergistic forms. Unexpectedly, we found only weak evidence for mediation of biomass-MAP relationships by plant community diversity because relationships of species richness, evenness, and beta diversity to MAP and to biomass were weak or opposing. Site-level properties including baseline biomass production, soils, and management explained little variation in biomass-MAP relationships. These findings reveal multiple nutrient colimitation as a defining feature of the global grassland biomass-MAP relationship.

Original language	English
Article number	e2410748122
Journal	Proceedings of the National Academy of Sciences of the United States of America
Volume	122
Issue number	15
DOIs	https://doi.org/10.1073/pnas.2410748122
Publication status	Published - Apr 2025

Bibliographical note

Publisher Copyright:
Copyright © 2025 the Author(s).

Funding

This work was conducted within the Nutrient Network (http://www.nutnet.org) experiment, funded at the site-scale by individual researchers. We thank the Minnesota Supercomputer Institute for hosting project data, the University of Minnesota Institute on the Environment for hosting Nutrient Network meetings, and each site investigator for funding their site-level operations. Network coordination and data management were supported by funds from the NSF Research Coordination Network (NSF-DEB-1042132) to E.T.B. and E.W.S., from the Long Term Ecological Research program (NSF-DEB-1831944) to the Cedar Creek LTER, and from the Institute on the Environment (DG-0001-13). The Co-authors acknowledge funding from the United States Department of Agriculture (USDA) (USDA-NIFA-2010-65615-20632) and the Swiss Federal Institute for Forest, Snow, and Landscape Research. The USDA is an equal opportunity employer and provider (P.A.F.). Consejo Nacional de Investigaciones Cient\u00EDficas y T\u00E9cnicas and Agencia Nacional de Promoci\u00F3n de la Investigaci\u00F3n, el Desarrollo Tecnol\u00F3gico y la Innovaci\u00F3n, Argentina PICT 2014-3026 (L.Y.). University of Iceland Research Fund (2015), Orkuranns\u00F3knasj\u00F3\u00D0ur Landsvirkjunnar N\u00DDR-09-2017, N\u00DDR-14-2018, N\u00DDR-12-2019 (I.C.B.). Co-Centre award number 22/CC/11103, managed by Science Foundation Ireland, Northern Ireland\u2019s Department of Agriculture, Environment and Rural Affairs and UK Research and Innovation, and supported via UK\u2019s International Science Partnerships Fund, and the Irish Government\u2019s Shared Island initiative (Y.M.B.). Funda\u00E7\u00E3o para a Ci\u00EAncia e Tecnologia funding to research units Forest Reseach Center (UIDB/00239/2023) and Research Network in Biodiversity and Evolutionary Biology (UID/BIA/50027/2023) and to Rui Alves for granting access to study site of Companhia das Lez\u00EDrias (M.N.B. and M.C.C.). European Research Council under the European Union\u2019s Horizon 2020 research and innovation programme grant agreement No. [101002987] (J.A.C.). Bush Heritage Australia and the Wangkamadla people for access to the sites (C.R.D. and G.M.W.). iDiv funded by the German Research Foundation DFG\u2013FZT 118, 202548816, and DFG Ei 862/29-1 (N.E.). Academy project no 351089 (A.E.). USDA Forest Service, Rocky Mountain Research Station (R.W.H.). University of Iceland Research Fund (I.S.J.). CEREEP-Ecotron IleDeFrance Centre National de la Recherche Scientifique/Ecole Normale Superior UMS 3194 (X.R.).

Funders	Funder number
Fundação para a Ciência e a Tecnologia
École normale supérieure
M.C.C.
Consejo Nacional de Investigaciones Científicas y Técnicas
Swiss Federal Institute for Forest
U.S. Forest Service
IleDeFrance Centre National de la Recherche Scientifique
Science Foundation Ireland
Háskóli Íslands
Rocky Mountain Research Station
European Research Council
Northern Ireland’s Department of Agriculture, Environment and Rural Affairs and UK Research and Innovation
University of Minnesota Institute
Bush Heritage Australia
California Department of Fish and Game	Ei 862/29-1, 351089
National Science Foundation	NSF-DEB-1831944, NSF-DEB-1042132
U.S. Department of Agriculture	USDA-NIFA-2010-65615-20632
Research Network in Biodiversity and Evolutionary Biology	UID/BIA/50027/2023
Institute on the Environment	DG-0001-13
Horizon 2020 Framework Programme	101002987
University of Iceland Research Fund	NÝR-14-2018, NÝR-12-2019, 22/CC/11103, NÝR-09-2017
Forest Reseach Center	UIDB/00239/2023
Deutsche Forschungsgemeinschaft	202548816, FZT 118
Agencia Nacional de Promoción de la Investigación, el Desarrollo Tecnológico y la Innovación	PICT 2014-3026

Keywords

Biodiversity
Biomass
Ecosystem
Fertilizers
Grassland
Nitrogen/metabolism
Nutrients/metabolism
Phosphorus/metabolism
Rain
Soil/chemistry

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10.1073/pnas.2410748122

Cite this

Fay, P. A., Gherardi, L. A., Yahdjian, L., Adler, P. B., Bakker, J. D., Bharath, S., Borer, E. T., Harpole, W. S., Hersch-Green, E., Huxman, T. E., MacDougall, A. S., Risch, A. C., Seabloom, E. W., Bagchi, S., Barrio, I. C., Biederman, L., Buckley, Y. M., Bugalho, M. N., Caldeira, M. C., ... Wheeler, G. R. (2025). Interactions among nutrients govern the global grassland biomass-precipitation relationship. Proceedings of the National Academy of Sciences of the United States of America, 122(15), Article e2410748122. https://doi.org/10.1073/pnas.2410748122

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title = "Interactions among nutrients govern the global grassland biomass-precipitation relationship",

abstract = "Ecosystems are experiencing changing global patterns of mean annual precipitation (MAP) and enrichment with multiple nutrients that potentially colimit plant biomass production. In grasslands, mean aboveground plant biomass is closely related to MAP, but how this relationship changes after enrichment with multiple nutrients remains unclear. We hypothesized the global biomass-MAP relationship becomes steeper with an increasing number of added nutrients, with increases in steepness corresponding to the form of interaction among added nutrients and with increased mediation by changes in plant community diversity. We measured aboveground plant biomass production and species diversity in 71 grasslands on six continents representing the global span of grassland MAP, diversity, management, and soils. We fertilized all sites with nitrogen, phosphorus, and potassium with micronutrients in all combinations to identify which nutrients limited biomass at each site. As hypothesized, fertilizing with one, two, or three nutrients progressively steepened the global biomass-MAP relationship. The magnitude of the increase in steepness corresponded to whether sites were not limited by nitrogen or phosphorus, were limited by either one, or were colimited by both in additive, or synergistic forms. Unexpectedly, we found only weak evidence for mediation of biomass-MAP relationships by plant community diversity because relationships of species richness, evenness, and beta diversity to MAP and to biomass were weak or opposing. Site-level properties including baseline biomass production, soils, and management explained little variation in biomass-MAP relationships. These findings reveal multiple nutrient colimitation as a defining feature of the global grassland biomass-MAP relationship.",

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author = "Fay, {Philip A} and Gherardi, {Laureano A} and Laura Yahdjian and Adler, {Peter B} and Bakker, {Jonathan D} and Siddharth Bharath and Borer, {Elizabeth T} and Harpole, {W Stanley} and Erika Hersch-Green and Huxman, {Travis E} and MacDougall, {Andrew S} and Risch, {Anita C} and Seabloom, {Eric W} and Sumanta Bagchi and Barrio, {Isabel C} and Lori Biederman and Buckley, {Yvonne M} and Bugalho, {Miguel N} and Caldeira, {Maria C} and Catford, {Jane A} and QingQing Chen and Cleland, {Elsa E} and Collins, {Scott L} and Pedro Daleo and Dickman, {Christopher R} and Ian Donohue and DuPre, {Mary E} and Nico Eisenhauer and Anu Eskelinen and Nicole Hagenah and Yann Hautier and Heckman, {Robert W} and J{\'o}nsd{\'o}ttir, {Ingibj{\"o}rg S} and Knops, {Johannes M H} and Ramesh Laungani and Martina, {Jason P} and McCulley, {Rebecca L} and Morgan, {John W} and {Olde Venterink}, Harry and Peri, {Pablo L} and Power, {Sally A} and Xavier Raynaud and Zhengwei Ren and Christiane Roscher and Smith, {Melinda D} and Marie Spohn and Stevens, {Carly J} and Tedder, {Michelle J} and Risto Virtanen and Wardle, {Glenda M} and Wheeler, {George R}",

note = "Publisher Copyright: Copyright {\textcopyright} 2025 the Author(s).",

year = "2025",

month = apr,

doi = "10.1073/pnas.2410748122",

language = "English",

volume = "122",

journal = "Proceedings of the National Academy of Sciences of the United States of America",

issn = "0027-8424",

publisher = "National Academy of Sciences",

number = "15",

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Fay, PA, Gherardi, LA, Yahdjian, L, Adler, PB, Bakker, JD, Bharath, S, Borer, ET, Harpole, WS, Hersch-Green, E, Huxman, TE, MacDougall, AS, Risch, AC, Seabloom, EW, Bagchi, S, Barrio, IC, Biederman, L, Buckley, YM, Bugalho, MN, Caldeira, MC, Catford, JA, Chen, Q, Cleland, EE, Collins, SL, Daleo, P, Dickman, CR, Donohue, I, DuPre, ME, Eisenhauer, N, Eskelinen, A, Hagenah, N, Hautier, Y, Heckman, RW, Jónsdóttir, IS, Knops, JMH, Laungani, R, Martina, JP, McCulley, RL, Morgan, JW, Olde Venterink, H, Peri, PL, Power, SA, Raynaud, X, Ren, Z, Roscher, C, Smith, MD, Spohn, M, Stevens, CJ, Tedder, MJ, Virtanen, R, Wardle, GM & Wheeler, GR 2025, 'Interactions among nutrients govern the global grassland biomass-precipitation relationship', Proceedings of the National Academy of Sciences of the United States of America, vol. 122, no. 15, e2410748122. https://doi.org/10.1073/pnas.2410748122

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T1 - Interactions among nutrients govern the global grassland biomass-precipitation relationship

AU - Fay, Philip A

AU - Gherardi, Laureano A

AU - Yahdjian, Laura

AU - Adler, Peter B

AU - Bakker, Jonathan D

AU - Bharath, Siddharth

AU - Borer, Elizabeth T

AU - Harpole, W Stanley

AU - Hersch-Green, Erika

AU - Huxman, Travis E

AU - MacDougall, Andrew S

AU - Risch, Anita C

AU - Seabloom, Eric W

AU - Bagchi, Sumanta

AU - Barrio, Isabel C

AU - Biederman, Lori

AU - Buckley, Yvonne M

AU - Bugalho, Miguel N

AU - Caldeira, Maria C

AU - Catford, Jane A

AU - Chen, QingQing

AU - Cleland, Elsa E

AU - Collins, Scott L

AU - Daleo, Pedro

AU - Dickman, Christopher R

AU - Donohue, Ian

AU - DuPre, Mary E

AU - Eisenhauer, Nico

AU - Eskelinen, Anu

AU - Hagenah, Nicole

AU - Hautier, Yann

AU - Heckman, Robert W

AU - Jónsdóttir, Ingibjörg S

AU - Knops, Johannes M H

AU - Laungani, Ramesh

AU - Martina, Jason P

AU - McCulley, Rebecca L

AU - Morgan, John W

AU - Olde Venterink, Harry

AU - Peri, Pablo L

AU - Power, Sally A

AU - Raynaud, Xavier

AU - Ren, Zhengwei

AU - Roscher, Christiane

AU - Smith, Melinda D

AU - Spohn, Marie

AU - Stevens, Carly J

AU - Tedder, Michelle J

AU - Virtanen, Risto

AU - Wardle, Glenda M

AU - Wheeler, George R

PY - 2025/4

Y1 - 2025/4

N2 - Ecosystems are experiencing changing global patterns of mean annual precipitation (MAP) and enrichment with multiple nutrients that potentially colimit plant biomass production. In grasslands, mean aboveground plant biomass is closely related to MAP, but how this relationship changes after enrichment with multiple nutrients remains unclear. We hypothesized the global biomass-MAP relationship becomes steeper with an increasing number of added nutrients, with increases in steepness corresponding to the form of interaction among added nutrients and with increased mediation by changes in plant community diversity. We measured aboveground plant biomass production and species diversity in 71 grasslands on six continents representing the global span of grassland MAP, diversity, management, and soils. We fertilized all sites with nitrogen, phosphorus, and potassium with micronutrients in all combinations to identify which nutrients limited biomass at each site. As hypothesized, fertilizing with one, two, or three nutrients progressively steepened the global biomass-MAP relationship. The magnitude of the increase in steepness corresponded to whether sites were not limited by nitrogen or phosphorus, were limited by either one, or were colimited by both in additive, or synergistic forms. Unexpectedly, we found only weak evidence for mediation of biomass-MAP relationships by plant community diversity because relationships of species richness, evenness, and beta diversity to MAP and to biomass were weak or opposing. Site-level properties including baseline biomass production, soils, and management explained little variation in biomass-MAP relationships. These findings reveal multiple nutrient colimitation as a defining feature of the global grassland biomass-MAP relationship.

AB - Ecosystems are experiencing changing global patterns of mean annual precipitation (MAP) and enrichment with multiple nutrients that potentially colimit plant biomass production. In grasslands, mean aboveground plant biomass is closely related to MAP, but how this relationship changes after enrichment with multiple nutrients remains unclear. We hypothesized the global biomass-MAP relationship becomes steeper with an increasing number of added nutrients, with increases in steepness corresponding to the form of interaction among added nutrients and with increased mediation by changes in plant community diversity. We measured aboveground plant biomass production and species diversity in 71 grasslands on six continents representing the global span of grassland MAP, diversity, management, and soils. We fertilized all sites with nitrogen, phosphorus, and potassium with micronutrients in all combinations to identify which nutrients limited biomass at each site. As hypothesized, fertilizing with one, two, or three nutrients progressively steepened the global biomass-MAP relationship. The magnitude of the increase in steepness corresponded to whether sites were not limited by nitrogen or phosphorus, were limited by either one, or were colimited by both in additive, or synergistic forms. Unexpectedly, we found only weak evidence for mediation of biomass-MAP relationships by plant community diversity because relationships of species richness, evenness, and beta diversity to MAP and to biomass were weak or opposing. Site-level properties including baseline biomass production, soils, and management explained little variation in biomass-MAP relationships. These findings reveal multiple nutrient colimitation as a defining feature of the global grassland biomass-MAP relationship.

KW - Biodiversity

KW - Biomass

KW - Ecosystem

KW - Fertilizers

KW - Grassland

KW - Nitrogen/metabolism

KW - Nutrients/metabolism

KW - Phosphorus/metabolism

KW - Rain

KW - Soil/chemistry

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U2 - 10.1073/pnas.2410748122

DO - 10.1073/pnas.2410748122

M3 - Article

C2 - 40215280

SN - 0027-8424

VL - 122

JO - Proceedings of the National Academy of Sciences of the United States of America

JF - Proceedings of the National Academy of Sciences of the United States of America

IS - 15

M1 - e2410748122

ER -

Interactions among nutrients govern the global grassland biomass-precipitation relationship

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