{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,5,11]],"date-time":"2026-05-11T11:47:59Z","timestamp":1778500079857,"version":"3.51.4"},"reference-count":60,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2023,11,27]],"date-time":"2023-11-27T00:00:00Z","timestamp":1701043200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Ecol. Evol."],"abstract":"<jats:p>The loss of plant productivity with declining diversity is well established, exceeding other global change drivers including drought. These patterns are most clearly established for aboveground productivity, it remains poorly understood whether productivity increases associated with diversity are replicated belowground. To address this gap, we established a plant diversity-manipulation experiment in 2018. It is a full factorial manipulation of plant species richness and community composition, and precipitation. Three and five years post-establishment, two bulk soil cores (20cm depth) were collected and composited from each plot and were processed for roots to determine belowground biomass as root standing crop. We observed a strong positive relationship between richness and aboveground production and belowground biomass, generating positive combined above and belowground with diversity. Root standing crop increased 1.4-fold from years three to five. Grass communities produced more root biomass (monoculture mean 463.9 \u00b1 410.3g m<jats:sup>\u22122<\/jats:sup>), and the magnitude of the relationship between richness and root standing crop was greatest within those communities. Legume communities produced the fewest roots (monoculture mean 212.2\u00a0\u00b1\u00a0155.1g m<jats:sup>\u22122<\/jats:sup>), and belowground standing crop was not affected by diversity. Root standing crops in year three were 1.8 times higher under low precipitation conditions, while in year five we observed comparable root standing crops between precipitation treatments. Plant family was a strong mediator of increased belowground biomass observed with diversity, with single family grass and aster families generating 1.7 times greater root standing crops in six compared to single species communities, relationships between diversity and aboveground production were consistently observed in both single-family and multiple family communities. Diverse communities with species from multiple families generated only 1.3 times the root standing crop compared to monoculture average root biomass. We surprisingly observe diverse single family communities can generate increases in root standing crops that exceed those generated by diverse multiple family communities, highlighting the importance of plant richness within plant family for a given community. These patterns have potential implications for understanding the interactions of multiple global change drivers as changes in both precipitation and plant community composition do alter whether plant production aboveground is translated belowground biomass.<\/jats:p>","DOI":"10.3389\/fevo.2023.1259809","type":"journal-article","created":{"date-parts":[[2023,11,27]],"date-time":"2023-11-27T06:56:41Z","timestamp":1701068201000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":10,"title":["Plant diversity and grasses increase root biomass in a rainfall and grassland diversity manipulation"],"prefix":"10.3389","volume":"11","author":[{"given":"Laura Y.","family":"Podzikowski","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Megan M.","family":"Heffernan","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"James D.","family":"Bever","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1965","published-online":{"date-parts":[[2023,11,27]]},"reference":[{"key":"B46","doi-asserted-by":"crossref","first-page":"1","DOI":"10.18637\/jss.v067.i01","article-title":"Fitting linear mixed-effects models using lme4","volume":"67","author":"Bates","year":"2015","journal-title":"J. Stat. Software"},{"key":"B41","doi-asserted-by":"publisher","first-page":"2934","DOI":"10.1016\/j.soilbio.2006.05.004","article-title":"Soil organic matter dynamics and land use change at a grassland\/forest ecotone","volume":"38","author":"Billings","year":"2006","journal-title":"Soil Biol. Biochem."},{"key":"B48","doi-asserted-by":"publisher","first-page":"56","DOI":"10.32614\/RJ-2017-046","article-title":"Visualization of regression models using visreg","volume":"9","author":"Breheny","year":"2017","journal-title":"R J."},{"key":"B45","doi-asserted-by":"publisher","first-page":"261","DOI":"10.1177\/0049124104268644","article-title":"Multimodel inference: understanding AIC and BIC in model selection","volume":"33","author":"Burnham","year":"2004","journal-title":"Sociol. Methods Res."},{"key":"B60","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/s43705-023-00237-5","article-title":"Rapid differentiation of soil and root microbiomes in response to plant composition and biodiversity in the field","volume":"3","author":"Burrill","year":"2023","journal-title":"ISME Commun."},{"key":"B27","doi-asserted-by":"publisher","first-page":"1039","DOI":"10.1007\/s10021-013-9666-z","article-title":"Contrasting effects of precipitation manipulations on production in two sites within the central grassland region, USA","volume":"16","author":"Byrne","year":"2013","journal-title":"Ecosystems"},{"key":"B43","doi-asserted-by":"publisher","first-page":"1","DOI":"10.18637\/jss.v034.i12","article-title":"glmulti: An R Package for Easy Automated Model Selection with (Generalized) Linear Models","volume":"34","author":"Calcagno","year":"2010","journal-title":"Journal of Statistical Software"},{"key":"B10","doi-asserted-by":"publisher","first-page":"683","DOI":"10.1038\/s41586-021-03871-y","article-title":"Fine-root traits in the global spectrum of plant form and function","volume":"597","author":"Carmona","year":"2021","journal-title":"Nature"},{"key":"B42","doi-asserted-by":"publisher","first-page":"2265","DOI":"10.1890\/14-1770.1","article-title":"Grass invasion effects on forest soil carbon depend on landscape-level land use patterns","volume":"96","author":"Craig","year":"2015","journal-title":"Ecology"},{"key":"B28","doi-asserted-by":"publisher","first-page":"317","DOI":"10.1007\/s11258-016-0690-x","article-title":"Drought timing differentially affects above- and belowground productivity in a mesic grassland","volume":"218","author":"Denton","year":"2017","journal-title":"Plant Ecol."},{"key":"B14","doi-asserted-by":"publisher","first-page":"132","DOI":"10.1111\/nph.16001","article-title":"Changes in root-exudate-induced respiration reveal a novel mechanism through which drought affects ecosystem carbon cycling","volume":"224","author":"de Vries","year":"2019","journal-title":"New Phytologist."},{"key":"B22","doi-asserted-by":"publisher","first-page":"6015","DOI":"10.1111\/gcb.15270","article-title":"The response of soil respiration to precipitation change is asymmetric and differs between grasslands and forests","volume":"26","author":"Du","year":"2020","journal-title":"Global Change Biol."},{"key":"B58","doi-asserted-by":"publisher","first-page":"44641","DOI":"10.1038\/srep44641","article-title":"Root biomass and exudates link plant diversity with soil bacterial and fungal biomass","volume":"7","author":"Eisenhauer","year":"2017","journal-title":"Sci. Rep. Uk"},{"key":"B25","doi-asserted-by":"publisher","first-page":"20","DOI":"10.1007\/s10021-012-9593-4","article-title":"Carbon and nitrogen decoupling under an 11-year drought in the shortgrass steppe","volume":"16","author":"Evans","year":"2013","journal-title":"Ecosystems"},{"key":"B33","doi-asserted-by":"publisher","first-page":"10572","DOI":"10.1073\/pnas.1712381114","article-title":"Hydrologic regulation of plant rooting depth","volume":"114","author":"Fan","year":"2017","journal-title":"P Natl. Acad. Sci. U.S.A."},{"key":"B29","doi-asserted-by":"publisher","first-page":"40","DOI":"10.1016\/j.agrformet.2013.01.002","article-title":"Response of plant biomass and soil respiration to experimental warming and precipitation manipulation in a Northern Great Plains grassland","volume":"173","author":"Flanagan","year":"2013","journal-title":"Agr. For. Meteorol."},{"key":"B19","doi-asserted-by":"publisher","first-page":"48","DOI":"10.1111\/j.1365-2745.2008.01453.x","article-title":"Linkages between plant functional composition, fine root processes and potential soil N mineralization rates","volume":"97","author":"Fornara","year":"2009","journal-title":"J. Ecol."},{"key":"B16","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2111321118","article-title":"Plant biodiversity and the regeneration of soil fertility","volume":"118","author":"Furey","year":"2021","journal-title":"P Natl. Acad. Sci. U.S.A."},{"key":"B17","doi-asserted-by":"publisher","first-page":"101","DOI":"10.1016\/S0929-1393(02)00137-3","article-title":"No consistent effects of plant diversity on root biomass, soil biota and soil abiotic conditions in temperate grassland communities","volume":"24","author":"Gastine","year":"2003","journal-title":"Appl. Soil Ecol."},{"key":"B55","doi-asserted-by":"publisher","first-page":"79","DOI":"10.1046\/j.1466-822X.2001.00267.x","article-title":"Using simple environmental variables to estimate below-ground productivity in grasslands","volume":"11","author":"Gill","year":"2002","journal-title":"Global Ecol. Biogeogr."},{"key":"B53","doi-asserted-by":"publisher","first-page":"3870","DOI":"10.1038\/s41467-020-17688-2","article-title":"Blind spots in global soil biodiversity and ecosystem function research","volume":"11","author":"Guerra","year":"2020","journal-title":"Nat. Commun."},{"key":"B5","doi-asserted-by":"publisher","first-page":"1123","DOI":"10.1126\/science.286.5442.1123","article-title":"Plant diversity and productivity experiments in European grasslands","volume":"286","author":"Hector","year":"1999","journal-title":"Science"},{"key":"B34","doi-asserted-by":"publisher","first-page":"232","DOI":"10.1016\/S0169-5347(98)01328-7","article-title":"Hydraulic lift: a potentially important ecosystem process","volume":"13","author":"Horton","year":"1998","journal-title":"Trends Ecol. Evol."},{"key":"B7","doi-asserted-by":"publisher","first-page":"574","DOI":"10.1038\/nature15374","article-title":"Biodiversity increases the resistance of ecosystem productivity to climate extremes","volume":"526","author":"Isbell","year":"2015","journal-title":"Nature"},{"key":"B23","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0069561","article-title":"Biomass and its allocation in relation to temperature, precipitation, and soil nutrients in inner Mongolia Grasslands, China","volume":"8","author":"Kang","year":"2013","journal-title":"PloS One"},{"key":"B36","doi-asserted-by":"publisher","first-page":"307","DOI":"10.1046\/j.1526-100x.2000.80043.x","article-title":"Land-use history in ecosystem restoration: a 40-year study in the prairie-forest ecotone","volume":"8","author":"Kettle","year":"2000","journal-title":"Restor. Ecol."},{"key":"B49","doi-asserted-by":"publisher","first-page":"2202","DOI":"10.1126\/science.1076347","article-title":"Rainfall variability, carbon cycling, and plant species diversity in a mesic grassland","volume":"298","author":"Knapp","year":"2002","journal-title":"Science"},{"key":"B47","first-page":"1","article-title":"lmerTest Package: Tests in Linear Mixed Effects Models","volume-title":"Journal of Statistical Software","author":"Kuznetsova","year":"2017"},{"key":"B40","doi-asserted-by":"publisher","first-page":"6707","DOI":"10.1038\/ncomms7707","article-title":"Plant diversity increases soil microbial activity and soil carbon storage","volume":"6","author":"Lange","year":"2015","journal-title":"Nat. Commun."},{"key":"B50","volume-title":"Future Global Climate: Scenario-Based Projections and Near-Term Information","author":"Lee","year":"2021"},{"key":"B11","doi-asserted-by":"publisher","first-page":"787","DOI":"10.1890\/12-1399.1","article-title":"Root depth distribution and the diversity-productivity relationship in a long-term grassland experiment","volume":"94","author":"Mueller","year":"2013","journal-title":"Ecology"},{"key":"B57","doi-asserted-by":"publisher","first-page":"1017","DOI":"10.1111\/j.0030-1299.2007.15630.x","article-title":"Soil water partitioning contributes to species coexistence in tallgrass prairie","volume":"116","author":"Nippert","year":"2007","journal-title":"Oikos"},{"key":"B51","doi-asserted-by":"publisher","first-page":"65","DOI":"10.1007\/s11258-005-9052-9","article-title":"Intra-annual rainfall variability and grassland productivity: can the past predict the future","volume":"184","author":"Nippert","year":"2006","journal-title":"Plant Ecol."},{"key":"B35","doi-asserted-by":"publisher","first-page":"79","DOI":"10.1111\/ppl.12020","article-title":"Commensalism in an agroecosystem: hydraulic redistribution by deep-rooted legumes improves survival of a droughted shallow-rooted legume companion","volume":"149","author":"Pang","year":"2013","journal-title":"Physiol. Plant."},{"key":"B15","doi-asserted-by":"publisher","first-page":"187","DOI":"10.1111\/j.1461-0248.2010.01570.x","article-title":"Enhanced root exudation induces microbial feedbacks to N cycling in a pine forest under long-term CO2 fumigation","volume":"14","author":"Phillips","year":"2011","journal-title":"Ecol. Lett."},{"key":"B18","doi-asserted-by":"publisher","first-page":"669","DOI":"10.1111\/gcb.14777","article-title":"Increased microbial growth, biomass, and turnover drive soil organic carbon accumulation at higher plant diversity","volume":"26","author":"Prommer","year":"2020","journal-title":"Global Change Biol."},{"key":"B59","doi-asserted-by":"publisher","first-page":"1528","DOI":"10.1111\/oik.01502","article-title":"Long-term study of root biomass in a biodiversity experiment reveals shifts in diversity effects over time","volume":"123","author":"Ravenek","year":"2014","journal-title":"Oikos"},{"key":"B44","volume-title":"R: A language and environment for statistical computing. 4.1.1","year":"2022"},{"key":"B3","doi-asserted-by":"publisher","first-page":"589","DOI":"10.1126\/science.1217909","article-title":"Impacts of biodiversity loss escalate through time as redundancy fades","volume":"336","author":"Reich","year":"2012","journal-title":"Science"},{"key":"B13","doi-asserted-by":"publisher","first-page":"2281","DOI":"10.1890\/02-0298","article-title":"Grassroots ecology: Plant-microbe-soil interactions as drivers of plant community structure and dynamics","volume":"84","author":"Reynolds","year":"2003","journal-title":"Ecology"},{"key":"B1","doi-asserted-by":"publisher","first-page":"472","DOI":"10.1038\/461472a","article-title":"A safe operating space for humanity","volume":"461","author":"Rockstrom","year":"2009","journal-title":"Nature"},{"key":"B39","doi-asserted-by":"publisher","first-page":"419","DOI":"10.1111\/j.1461-0248.2005.00736.x","article-title":"Overyielding in experimental grassland communities - irrespective of species pool or spatial scale","volume":"8","author":"Roscher","year":"2005","journal-title":"Ecol. Lett."},{"key":"B30","doi-asserted-by":"publisher","first-page":"255","DOI":"10.1023\/A:1004801718567","article-title":"Root dynamics in a semi-natural grassland in relation to atmospheric carbon dioxide enrichment, soil water and shoot biomass","volume":"223","author":"Sindh\u00f8j","year":"2000","journal-title":"Plant Soil"},{"key":"B8","doi-asserted-by":"publisher","first-page":"5744","DOI":"10.1073\/pnas.1503376112","article-title":"Predicting plant vulnerability to drought in biodiverse regions using functional traits","volume":"112","author":"Skelton","year":"2015","journal-title":"P Natl. Acad. Sci. U.S.A."},{"key":"B31","doi-asserted-by":"publisher","first-page":"1265","DOI":"10.1007\/s10021-021-00714-9","article-title":"Effects of compounded precipitation pattern intensification and drought occur belowground in a mesic grassland","volume":"25","author":"Slette","year":"2022","journal-title":"Ecosystems"},{"key":"B2","doi-asserted-by":"publisher","first-page":"471","DOI":"10.1146\/annurev-ecolsys-120213-091917","article-title":"Biodiversity and ecosystem functioning","volume":"45","author":"Tilman","year":"2014","journal-title":"Annu. Rev. Ecol. Evol. S."},{"key":"B6","doi-asserted-by":"publisher","first-page":"1300","DOI":"10.1126\/science.277.5330.1300","article-title":"The influence of functional diversity and composition on ecosystem processes","volume":"277","author":"Tilman","year":"1997","journal-title":"Science"},{"key":"B38","unstructured":"Web Soil Survey2019"},{"key":"B9","doi-asserted-by":"publisher","first-page":"81","DOI":"10.1111\/j.1365-2745.2009.01603.x","article-title":"Diversity enhances community recovery, but not resistance, after drought","volume":"98","author":"van Ruijven","year":"2010","journal-title":"J. Ecol."},{"key":"B12","doi-asserted-by":"publisher","first-page":"20130119","DOI":"10.1098\/rstb.2013.0119","article-title":"Biological nitrogen fixation: rates, patterns and ecological controls in terrestrial ecosystems","volume":"368","author":"Vitousek","year":"2013","journal-title":"Philos. T R Soc. B."},{"key":"B4","doi-asserted-by":"publisher","first-page":"7752","DOI":"10.1038\/s41467-022-35189-2","article-title":"Biodiversity-stability relationships strengthen over time in a long-term grassland experiment","volume":"13","author":"Wagg","year":"2022","journal-title":"Nat. Commun."},{"key":"B37","doi-asserted-by":"publisher","first-page":"2721","DOI":"10.1111\/gcb.16091","article-title":"Microbial mediators of plant community response to long-term N and P fertilization: Evidence of a role of plant responsiveness to mycorrhizal fungi","volume":"28","author":"Wang","year":"2022","journal-title":"Global Change Biol."},{"key":"B32","doi-asserted-by":"publisher","first-page":"561","DOI":"10.1890\/15-1437.1","article-title":"Does ecosystem sensitivity to precipitation at the site-level conform to regional-scale predictions","volume":"97","author":"Wilcox","year":"2016","journal-title":"Ecology"},{"key":"B24","doi-asserted-by":"publisher","first-page":"927","DOI":"10.1111\/j.1365-2486.2010.02302.x","article-title":"Responses of terrestrial ecosystems to temperature and precipitation change: a meta-analysis of experimental manipulation","volume":"17","author":"Wu","year":"2011","journal-title":"Global Change Biol."},{"key":"B26","doi-asserted-by":"publisher","first-page":"1648","DOI":"10.1111\/j.1365-2486.2012.02651.x","article-title":"Interannual variability in responses of belowground net primary productivity (NPP) and NPP partitioning to long-term warming and clipping in a tallgrass prairie","volume":"18","author":"Xu","year":"2012","journal-title":"Global Change Biol."},{"key":"B54","doi-asserted-by":"publisher","first-page":"2753","DOI":"10.1111\/gcb.12248","article-title":"Net primary productivity and rain-use efficiency as affected by warming, altered precipitation, and clipping in a mixed-grass prairie","volume":"19","author":"Xu","year":"2013","journal-title":"Global Change Biol."},{"key":"B20","doi-asserted-by":"publisher","first-page":"718","DOI":"10.1038\/s41467-019-08636-w","article-title":"Soil carbon sequestration accelerated by restoration of grassland biodiversity","volume":"10","author":"Yang","year":"2019","journal-title":"Nat. Commun."},{"key":"B21","doi-asserted-by":"publisher","first-page":"143","DOI":"10.1016\/S0168-1656(01)00342-X","article-title":"Rhizobia from wild legumes: diversity, taxonomy, ecology, nitrogen fixation and biotechnology","volume":"91","author":"Zahran","year":"2001","journal-title":"J. Biotechnol."},{"key":"B52","doi-asserted-by":"publisher","first-page":"3083","DOI":"10.5194\/bg-11-3083-2014","article-title":"Impacts of extreme precipitation and seasonal changes in precipitation on plants","volume":"11","author":"Zeppel","year":"2014","journal-title":"Biogeosciences"},{"key":"B56","doi-asserted-by":"publisher","DOI":"10.1002\/ecy.2828","article-title":"Plants alter their vertical root distribution rather than biomass allocation in response to changing precipitation","volume":"100","author":"Zhang","year":"2019","journal-title":"Ecology"}],"container-title":["Frontiers in Ecology and Evolution"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fevo.2023.1259809\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2023,11,27]],"date-time":"2023-11-27T06:56:49Z","timestamp":1701068209000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fevo.2023.1259809\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2023,11,27]]},"references-count":60,"alternative-id":["10.3389\/fevo.2023.1259809"],"URL":"https:\/\/doi.org\/10.3389\/fevo.2023.1259809","relation":{},"ISSN":["2296-701X"],"issn-type":[{"value":"2296-701X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2023,11,27]]},"article-number":"1259809"}}