Freshwater Biology (1997) 38, 447–471
A model approach to planktonic stoichiometry and
consumer-resource stability
DAG O. HESSEN* AND BIRGER BJERKENG²
*University of Oslo, Department of Biology, PO Box 1027 Blindern, N-0316 Oslo, Norway
²Norwegian Institute for Water Research, PO Box 173 Kjelsås, N-0411 Oslo, Norway
SUMMARY
1. The model explores stoichiometric feedback effects between an alga and a grazer
(Daphnia) in a simplified chemostat-type system in stagnant conditions or with fixed
dilution rate.
2. When running the model with fixed stoichiometry and P-sufficient food, the grazer
with highest requirements for phosphorus (P) will exert the most efficient control of
algal biomass owing to more P being allocated to zooplankton biomass and less P
recycled.
3. When including potential P-limitation of the grazer, the grazer with high P
requirements (high P : C ratio) will be the least efficient grazer in a system with
fluctuating and temporarily low P : C ratio in algae (Q
a
).
4. Qualitatively deficient food will yield decreased growth efficiency in zooplankton. As
Q
a
decreases, the grazer isocline for zero net growth is shifted upwards, and the
required algal biomass for positive growth increases. There may then be a critical level
of Q
a
below which the grazer with high P : C suffers negative population growth
regardless of algal biomass. In cases with low minimum Q
a
and a P-demanding grazer,
this may cause the system to enter an irreversible stage with high biomass of P-deficient
phytoplankton which do not support zooplankton growth.
5. Cumulative primary production for scenarios with continuous P input is, in general,
higher the more Q
a
is allowed to drop below saturation values, and highest when this is
combined with a grazer with a high P : C ratio. The lower growth rate of P-deficient
phytoplankton was compensated for by reduced success of the P-limited grazer, yielding
low grazing pressure and resulting in larger phytoplankton biomass.
Introduction
Models of autotroph–herbivore interactions, from
Lotka–Volterra models and onwards, traditionally
focus on direct interactions and biomasses and flows
in terms of carbon (C). In the grazing process there is
a strong recirculation component embedded, however,
which may be an important regulating factor for
primary production and thus imply important feed-
backs for the grazer itself. Recent works underscore
that foodweb dynamics depend on more than the
direct interactions between trophic levels, populations
or individuals, and that there is a strong stoichiometric
component in foodweb interactions (Reiners, 1986;
DeAngelis et al., 1989; Sterner, Elser & Hessen, 1992;
© 1997 Blackwell Science Ltd 447
Sterner & Hessen, 1994). A grazer (or predator) will
probably extract a maximum amount of whatever
biochemical compounds and elements that are in
short supply relative to its own demands. Surplus
components will be released in some proportion to
their relative abundance in food (Olsen et al., 1986;
Hessen & Andersen, 1992). The release of the nutrient
elements phosphorus (P) and nitrogen (N) calls for
special attention because they are the most important
limiting elements for primary production in aquatic
and terrestrial ecosystems (Hecky & Kilham, 1988;
Vitousek & Howarth, 1991). The quantitative role of
nutrient release from grazers is well recognized in