Ecological Modelling 220 (2009) 2272–2280
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Ecological Modelling
journal homepage: www.elsevier.com/locate/ecolmodel
Foliage profiles of individual trees determine competition, self-thinning, biomass
and NPP of a Cryptomeria japonica forest stand: A simulation study based on a
stand-scale process-based forest model
Motomu Toda
a,∗
, Masayuki Yokozawa
b,1
, Akihiro Sumida
c,2
, Tsutomu Watanabe
c,2
, Toshihiko Hara
a,c,2
a
Pan-Okhotsk Research Center, The Institute of Low Temperature Science, Hokkaido University, Sapporo 060-0819, Japan
b
National Institute for Agro-Environmental Sciences, Ibaraki 305-8604, Japan
c
Cryosphere Science Research Section, The Institute of Low Temperature Science, Hokkaido University, Sapporo 060-0819, Japan
article info
Article history:
Received 19 September 2008
Received in revised form 28 April 2009
Accepted 11 May 2009
Available online 17 June 2009
Keywords:
MINoSGI
Size structure
Competitive asymmetry
Carbon gain of an individual tree
Biomass allocation pattern
Maximum carboxylation velocity
abstract
A simulation study was carried out to investigate simultaneously the effects of eco-physiological param-
eters on competitive asymmetry, self-thinning, stand biomass and NPP in a temperate forest using an
atmosphere–vegetation dynamics interactive model (MINoSGI). In this study, we selected three eco-
physiological relevant parameters as foliage profiles (i.e. vertical distribution of leaf area density) of
individual trees (distribution pattern is described by the parameter ), biomass allocation pattern in indi-
vidual tree growth () and the maximum carboxylation velocity (V
max
). The position of the maximal leaf
area density shifts upward in the canopy with increasing . For scenarios with < 4 (foliage concentrated
in the lowest canopy layer) or > 12 (foliage concentrated in the uppermost canopy layer), a low degree of
competitive asymmetry was produced. These scenarios resulted in the survival of subordinate trees due
to a brighter lower canopy environment when < 4 or the generation of spatially separated foliage profiles
between dominant and subordinate trees when >12. In contrast, competition between trees was most
asymmetric when 4 ≤ ≤ 12 (vertically widespread foliage profile in the canopy), especially when = 8.
In such cases, vertically widespread foliage of dominant trees lowered the opportunity of light acquisition
for subordinate trees and reduced their carbon gain. The resulting reduction in carbon gain of subordinate
trees yielded a higher degree of competitive asymmetry and ultimately higher mortality of subordinate
trees. It was also shown that 4 ≤ ≤ 12 generated higher self-thinning speed, smaller accumulated NPP,
litter-fall and potential stand biomass as compared with the scenarios with < 4 or > 12. In contrast, our
simulation revealed small effects of or V
max
on the above-mentioned variables as compared with those
of . In particular, it is notable that greater V
max
would not produce greater potential stand biomass and
accumulated NPP although it has been thought that physiological parameters relevant to photosynthesis
such as V
max
influence dynamic changes in forest stand biomass and NPP (e.g. the greater the V
max
, the
greater the NPP). Overall, it is suggested that foliage profiles rather than biomass allocation or maximum
carboxylation velocity greatly govern forest dynamics, stand biomass, NPP and litter-fall.
© 2009 Elsevier B.V. All rights reserved.
1. Introduction
Over the past decade, many atmosphere–vegetation dynamics
model studies have been conducted, and one of the significant aims
of the model development is to elucidate the feedback responses
of terrestrial ecosystems to climate systems (e.g. Cox et al., 2000;
Moorcroft, 2003; Sitch et al., 2003; Watanabe et al., 2004; Sato et
al., 2007). The intercomparison of representative models has shown
∗
Corresponding author. Tel.: +81 11 706 3356.
E-mail address: todam@ees.hokudai.ac.jp (M. Toda).
1
Tel.: +81 29 838 8435.
2
Tel.: +81 11 706 5455.
large uncertainties regarding the carbon storage potential of forests
in the future (Cramer et al., 2001; Friedlingstein et al., 2006). The
uncertainties possibly remain due to our inability to accurately cap-
ture ecological dynamics in forest models (Purves and Pacala, 2008),
therefore, suggesting the importance of accurately assessing forest
dynamics that influence the carbon cycle. In the present study, we
assess how the differences in eco-physiological parameters affect
the dynamic changes of plant size structure (frequency distribu-
tion of individual plant size (plant height, stem diameter, mass) in
a plant population) due to competition between individual plants
and the carbon cycles such as NPP (net primary production) in a
forest stand. We dealt with the eco-physiological parameters such
as foliage profiles of individual trees, biomass allocation pattern
in individual plant growth and maximum carboxylation velocity.
0304-3800/$ – see front matter © 2009 Elsevier B.V. All rights reserved.
doi:10.1016/j.ecolmodel.2009.05.011