Aboveground overyielding in grassland mixtures is associated with reduced biomass partitioning to belowground organs.

We investigated effects of plant species richness in experimental grassland plots on annual above- and belowground biomass production estimated from repeated harvests and ingrowth cores, respectively. Aboveground and total biomass production increased with increasing plant species richness while belowground production remained constant. Root to shoot biomass production ratios (R/S) in mixtures were lower than expected from monoculture performance of the species present in the mixtures, showing that interactions among species led to reduced biomass partitioning to belowground organs. This change in partitioning to belowground organs was not confined to mixtures with legumes, but also measured in mixtures without legumes, and correlated with aboveground overyielding in mixtures. It is suggested that species-rich communities invest less in belowground biomass than do monocultures to extract soil resources, thus leading to increased investment into aboveground organs and overyielding.

[1]  Angela Hodge,et al.  The plastic plant: root responses to heterogeneous supplies of nutrients , 2004 .

[2]  E. Schulze,et al.  Adaptive survival mechanisms and growth limitations of small-stature herb species across a plant diversity gradient. , 2008, Plant biology.

[3]  W. Stanley Harpole,et al.  Mechanisms responsible for the positive diversity–productivity relationship in Minnesota grasslands , 2004 .

[4]  N. Buchmann,et al.  Positive interactions between nitrogen-fixing legumes and four different neighbouring species in a biodiversity experiment , 2007, Oecologia.

[5]  F. Bazzaz,et al.  Interactive effects of diversity, nutrients and elevated CO2 on experimental plant communities , 2002 .

[6]  D. Tilman Distinguishing between the effects of species diversity and species composition , 1997 .

[7]  C. Körner,et al.  Above-ground resource use increases with plant species richness in experimental grassland ecosystems , 2000 .

[8]  J. Wilson A review of evidence on the control of shoot: root ratio , 1988 .

[9]  C. Körner,et al.  Plant diversity effects on soil heterotrophic activity in experimental grassland ecosystems , 2000, Plant and Soil.

[10]  R. B. Jackson,et al.  A global analysis of root distributions for terrestrial biomes , 1996, Oecologia.

[11]  M. Scherer‐Lorenzen,et al.  Inorganic soil nitrogen under grassland plant communities of different species composition and diversity. , 2005 .

[12]  A. Troumbis,et al.  The role of legumes as a component of biodiversity in a cross‐European study of grassland biomass nitrogen , 2002 .

[13]  E. Schulze,et al.  Soil and Plant Nitrogen Pools as Related to Plant Diversity in an Experimental Grassland , 2007 .

[14]  P. Balvanera,et al.  Quantifying the evidence for biodiversity effects on ecosystem functioning and services. , 2006, Ecology letters.

[15]  Catherine Potvin,et al.  BIODIVERSITY AND ECOSYSTEM FUNCTIONING: IMPORTANCE OF SPECIES EVENNESS IN AN OLD FIELD , 2000 .

[16]  F. Woodward,et al.  ECOSYSTEM EFFECTS OF BIODIVERSITY MANIPULATIONS IN EUROPEAN GRASSLANDS , 2005 .

[17]  Y. Crozat,et al.  Assessment of Root System Dynamics of Species Grown in Mixtures under Field Conditions using Herbicide Injection and 15N Natural Abundance Methods: A Case Study with Pea, Barley and Mustard , 2005, Plant and Soil.

[18]  B. Olson,et al.  Response of the invasive Centaurea maculosa and two native grasses to N-pulses , 2003, Plant and Soil.

[19]  Ian T. Carroll,et al.  Impacts of plant diversity on biomass production increase through time because of species complementarity , 2007, Proceedings of the National Academy of Sciences.

[20]  D. Hooper THE ROLE OF COMPLEMENTARITY AND COMPETITION IN ECOSYSTEM RESPONSES TO VARIATION IN PLANT DIVERSITY , 1998 .

[21]  F. Chapin,et al.  EFFECTS OF BIODIVERSITY ON ECOSYSTEM FUNCTIONING: A CONSENSUS OF CURRENT KNOWLEDGE , 2005 .

[22]  E. Schulze,et al.  The role of biodiversity for element cycling and trophic interactions: an experimental approach in a grassland community , 2004 .

[23]  N. Buchmann,et al.  Diversity‐dependent productivity in semi‐natural grasslands following climate perturbations , 2005 .

[24]  P. Reich,et al.  [Letters to nature] , 1975, Nature.

[25]  Z. Rengel,et al.  Mechanistic simulation models for a better understanding of nutrient uptake from soil. , 1999 .

[26]  C. Körner,et al.  A LONG‐TERM FIELD STUDY ON BIODIVERSITY × ELEVATED CO2 INTERACTIONS IN GRASSLAND , 2001 .

[27]  M. Biondini,et al.  Root morphological plasticity and nitrogen uptake of 59 plant species from the Great Plains grasslands, U.S.A. , 2001 .

[28]  A. Pfisterer,et al.  Species vs community perspectives in biodiversity experiments , 2003 .

[29]  P. Reich,et al.  Diversity and Productivity in a Long-Term Grassland Experiment , 2001, Science.

[30]  F. Berendse,et al.  Diversity-productivity relationships: initial effects, long-term patterns, and underlying mechanisms. , 2005, Proceedings of the National Academy of Sciences of the United States of America.

[31]  S. Pennings,et al.  PHENOTYPIC PLASTICITY AND INTERACTIONS AMONG PLANTS , 2003 .

[32]  P. Leadley,et al.  No consistent effects of plant diversity on root biomass, soil biota and soil abiotic conditions in temperate grassland communities , 2003 .

[33]  P. Högberg,et al.  How plant diversity and legumes affect nitrogen dynamics in experimental grassland communities , 2002, Oecologia.

[34]  Nina Buchmann,et al.  Niche complementarity for nitrogen: an explanation for the biodiversity and ecosystem functioning relationship? , 2006, Ecology.

[35]  N. Stanton The Underground in Grasslands , 1988 .

[36]  D. Wardle,et al.  Interspecific interactions and biomass allocation among grassland plant species , 2003 .

[37]  Michel Loreau,et al.  From selection to complementarity: shifts in the causes of biodiversity–productivity relationships in a long-term biodiversity experiment , 2007, Proceedings of the Royal Society B: Biological Sciences.

[38]  S. von Felten,et al.  Belowground nitrogen partitioning in experimental grassland plant communities of varying species richness. , 2009, Ecology.

[39]  Michel Loreau,et al.  Partitioning selection and complementarity in biodiversity experiments , 2001, Nature.

[40]  V. Ninkovic Volatile communication between barley plants affects biomass allocation. , 2003, Journal of experimental botany.

[41]  D. Tilman,et al.  Productivity and sustainability influenced by biodiversity in grassland ecosystems , 1996, Nature.

[42]  M. Loreau,et al.  Overyielding in grassland communities: testing the sampling effect hypothesis with replicated biodiversity experiments , 2002 .

[43]  P. Dimitrakopoulos,et al.  Biodiversity effects increase linearly with biotope space , 2004 .

[44]  D. Hooper,et al.  Overyielding among plant functional groups in a long‐term experiment , 2003 .

[45]  J. Coleman,et al.  BIOMASS ALLOCATION IN PLANTS: ONTOGENY OR OPTIMALITY? A TEST ALONG THREE RESOURCE GRADIENTS , 1999 .

[46]  R. B. Jackson,et al.  Global patterns of root turnover for terrestrial ecosystems , 2000 .

[47]  E. Schulze,et al.  Effects of trait plasticity on aboveground biomass production depend on species identity in experimental grasslands , 2008 .

[48]  Joel s. Brown,et al.  Tragedy of the commons as a result of root competition , 2001 .

[49]  D. Wardle,et al.  PLANT REMOVALS IN PERENNIAL GRASSLAND: VEGETATION DYNAMICS, DECOMPOSERS, SOIL BIODIVERSITY, AND ECOSYSTEM PROPERTIES , 1999 .

[50]  H. Mooney,et al.  Resource Limitation in Plants-An Economic Analogy , 1985 .

[51]  Christina E. Wells,et al.  Building roots in a changing environment: implications for root longevity , 2000 .

[52]  E. Schulze,et al.  THE ROLE OF PLANT DIVERSITY AND COMPOSITION FOR NITRATE LEACHING IN GRASSLANDS , 2003 .

[53]  P. Reich,et al.  The Influence of Functional Diversity and Composition on Ecosystem Processes , 1997 .

[54]  Michael A. Huston,et al.  Hidden treatments in ecological experiments: re-evaluating the ecosystem function of biodiversity , 1997, Oecologia.

[55]  F. Bazzaz,et al.  Density may alter diversity-productivity relationships in experimental plant communities , 2005 .

[56]  M. Loreau,et al.  Biodiversity and ecosystem functioning : synthesis and perspectives , 2002 .

[57]  J. Singh,et al.  The structure and function of ten Western North American grasslands: III. Net primary production, turnover and efficiencies of energy capture and water use , 1978 .

[58]  J. Fridley Diversity effects on production in different light and fertility environments: an experiment with communities of annual plants , 2003 .

[59]  J. Fargione,et al.  The role of plant species in biomass production and response to elevated CO 2 and N , 2003 .

[60]  A. Hector The effect of diversity on productivity : detecting the role of species complementarity , 1998 .

[61]  Nina Buchmann,et al.  Overyielding in experimental grassland communities - irrespective of species pool or spatial scale , 2005 .

[62]  Pereira,et al.  Plant diversity and productivity experiments in european grasslands , 1999, Science.