Ecological Modelling 189 (2005) 447–453
Symmetry of belowground competition in a spatially explicit
model of nutrient competition
Xavier Raynaud
∗
, Paul W. Leadley
Laboratoire
´
Ecologie, Syst´ ematique et
´
Evolution, UMR 8079 Universit´ e Paris-Sud XI, Bˆ at. 362 ORSAY, F-91405 Cedex, France
Received 3 September 2004; received in revised form 18 February 2005; accepted 29 March 2005
Available online 23 May 2005
Abstract
We studied the symmetry of belowground competition at the level of species using a model of plant competition at the scale
of individual roots. This model predicts the nutrient uptake of plants with varying root densities and uptake kinetics, competing
for the same resource, at equilibrium between resource supply and plant uptake. A symmetry parameter is derived from this
model. The analysis of the model behaviour suggests that the symmetry of belowground competition depends on the diffusive
supply of the nutrient (buffer capacity × effective diffusion coefficient) for which competition occurs. As the diffusive supply is
determined to a greater extent by soil water content and soil type rather than by nutrient type, this means that the symmetry of
belowground competition should vary with soil water content and soil type. In particular, our model suggests that competition
for nutrients that are primarily supplied by diffusion through the soil is (1) symmetrical at high diffusive supply when root
uptake kinetics are identical or at low diffusive supply when roots are randomly distributed in the soil whatever their uptake
kinetics are, (2) size-asymmetrical at high diffusive supply when root uptake kinetics are different, (3) partially symmetric or
size-asymmetric at low diffusive supply when root are not randomly distributed in the soil.
© 2005 Elsevier B.V. All rights reserved.
Keywords: Belowground competition; Symmetry; Nutrient uptake model
1. Introduction
The symmetry or asymmetry of plant competition
is thought to play a key role in determining competi-
tive outcomes in natural ecosystems (Schwinning and
Weiner, 1998), in part, because competitive displace-
∗
Corresponding author. Tel.: +33 169155701.
E-mail addresses: xavier.raynaud@ese.u-psud.fr (X. Raynaud),
paul.leadley@ese.u-psud.fr (P.W. Leadley).
ment may be much more rapid under size-asymmetric
competition than under size-symmetric competition.
Symmetry and asymmetry of competition have
generally been defined by studying the relationships
between plant size (usually measured as plant height or
plant biomass) and plant resource acquisition (usually
measured as plant growth rates or nutrient uptake)
(Schwinning and Fox, 1995). When resources are taken
up in proportion to plant size, competition is defined
as being size-symmetric. When larger plants acquire
0304-3800/$ – see front matter © 2005 Elsevier B.V. All rights reserved.
doi:10.1016/j.ecolmodel.2005.03.008