Copyright © 2005 by the author(s). Published here under license by the Resilience Alliance.
Amemiya, T., T. Enomoto, A. G. Rossberg, N. Takamura and K. Itoh. 2005. Lake restoration in terms of
ecological resilience: a numerical study of biomanipulations under bistable conditions. Ecology and Society
10(2): 3. [online] URL: http://www.ecologyandsociety.org/vol10/iss2/art3/
Research
Lake Restoration in Terms of Ecological Resilience: a Numerical
Study of Biomanipulations under Bistable Conditions
Takashi Amemiya
1
, Takatoshi Enomoto
1
, A. G. Rossberg
1
, Noriko Takamura
2
, and Kiminori Itoh
1
ABSTRACT. An abstract version of the comprehensive aquatic simulation model (CASM) is found to
exhibit bistability under intermediate loading of nutrient input, supporting the alternative-stable-states
theory and field observations for shallow lakes. Our simulations of biomanipulations under the bistable
conditions reveal that a reduction in the abundance of zooplanktivorous fish cannot switch the system from
a turbid to a clear state. Rather, a direct reduction of phytoplankton and detritus was found to be most
effective to make this switch in the present model. These results imply that multiple manipulations may
be effective for practical restorations of lakes. We discuss the present results of biomanipulations in terms
of ecological resilience in multivariable systems or natural systems.
Key Words: alternative stable state; biomanipulation; bistable; comprehensive aquatic simulation model
(CASM); resilience
INTRODUCTION
Several mathematical models of ecosystems have
been proposed and used for understanding complex
aspects of ecosystems (Walker et al. 1981,
DeAngelis et al. 1989, Hulot et al. 2000, Scheffer
2001). Among these models, the comprehensive
aquatic simulation model (CASM) (DeAngelis et
al. 1989) has been found to be useful in applications
for several aquatic ecosystems, and has been used
for ecological risk assessment (Bartel et al. 1999,
Naito et al. 2002). A simplified, more abstract
version of CASM introduced by DeAngelis et al.
(1989), abbreviated as abs.-CASM hereafter, is
more suitable for a theoretical analysis; it was
previously employed for studying stability of
ecosystems (DeAngelis et al. 1989). Abs.-CASM
retains important characteristics of aquatic
ecosystems: material circulation, species interactions
among three trophic levels, and the effects of human
activities on aquatic ecosystems. DeAngelis et al.
(1989) have analyzed abs.-CASM in terms of
resilience, sensu Pimm (1984), i.e., the return time
following perturbations (“engineering resilience,”
Holling 1996). For their analyses, monostable states
were examined as a function of nutrient loading.
Many ecosystems, including shallow lakes, are
known to exhibit bistable states or alternative stable
states (Scheffer et al. 2001). Scheffer (2001) and
Beisner et al. (2003) have shown bistable behavior
in two- or three-variable aquatic models. We show
that abs.-CASM also exhibits bistable behavior
under a realistic range of parameter values, and that
it is useful to study the properties of alternative
stable states.
Ecological resilience, below just referred to as
resilience, is introduced to describe the properties
of alternative stable states in ecosystems (Holling
1973). Resilience concerns large and drastic
changes in ecosystems. It characterizes “the amount
of change the system can undergo and still retain
the same controls on function and structure”
(Holling 1973, 1996), and is used as a novel concept
for ecosystem management (Scheffer et al. 2001,
Bellwood et al. 2004, Carpenter and Brock 2004).
Resilience is usually illustrated by using a sigmoidal
bifurcation curve for a simple one-variable system,
as shown in Fig. 1. The horizontal axis is an
environmental factor, i.e., control parameter, such
1
Yokohama National University,
2
National Institute for Environmental Studies