Abstract Macrozooplankton may affect algal and mi-
crobial plankton directly through grazing or predation
and indirectly through nutrient regeneration. They may
also affect potential prey positively by removing alterna-
tive predators. Here, we examined the effects of a cla-
doceran (Daphnia) and a calanoid copepod (Eodiapto-
mus) on algal and microbial plankton in a Japanese lake
using in situ experiments in which we manipulated the
nutrient supply and biomass of these macrozooplankton.
The response of algal and microbial plankton to macro-
zooplankton was diverse and varied depending on the
level of nutrient supply. Eodiaptomus seemed to feed
mainly on large algae (>20 μm) and microzooplankton,
while direct grazing by Daphnia on algae, bacteria, het-
erotrophic nanoflagellates (HNF), and microzooplankton
(ciliates, heliozoa, and rotifers) was pronounced. Trophic
linkages within these microbial plankton was also sug-
gested; bacteria were grazed by HNF and these in turn
were grazed by microzooplankton. When the nutrient
supply was high, both HNF and microzooplankton were
exposed to higher amounts of algae and lower bacterial
abundance. Moreover, nutrient regeneration by daphnids
and Eodiaptomus copepods seemed to differentially
stimulate the growth of algae and bacteria. The results
suggest that the relationship between macrozooplankton
and microbial plankton cannot be fully understood with-
out taking into consideration not only the feeding char-
acteristics of the macrozooplankton, but also the food
web structure, the subsidized algal resource, and nutrient
regeneration from the macrozooplankton.
Keywords Trophic interactions · Nutrient regeneration ·
Resource subsidy · Classic and microbial food chains
Introduction
In pelagic communities, the grazing chain from algae to
macrozooplankton such as daphnids and copepods has
traditionally been viewed as a major way to sustain high-
er trophic levels. However, there is growing evidence
that bacteria and protozoan plankton (flagellates and cil-
iates) are equally or sometimes more important constitu-
ents of pelagic food webs. Via bacteria, this trophic link
can transfer substantial amounts of dissolved organic
matter to higher trophic levels. This transfer of dissolved
organic matter to bacteria and protozoan plankton is
called the microbial chain, or the microbial loop (Azam
et al. 1983). Because macrozooplankton feed not only on
algae but also on a wide range of organisms, including
bacteria and protozoans (Hessen et al. 1986; Sanders and
Porter 1990; Wickham et al. 1993; Pace and Vaqué
1994), they play a key role in linking the microbial loop
to the grazing chain (Porter et al. 1979; Carrick et al.
1991; Jürgens 1994). This, in turn, implies that macro-
zooplankton can affect the abundance of organisms in
both the microbial and the grazing chains.
It is well documented that macrozooplankton affect
algal abundance and composition directly through graz-
ing and indirectly through nutrient regeneration (Lehman
and Sandgren 1985; Sterner 1986; Vanni 1987; Elser and
Goldman 1991; Brett et al. 1994; Urabe 1995). In addi-
tion, different macrozooplankton affect algal species
composition in different ways, primarily because of spe-
cies- or taxon-specific characteristics of the feeding
mode (Okamoto 1984; DeMott 1986, 1988) and nutrient
recycling (Elser and Urabe 1999).
Compared with the wealth of knowledge on algal-
macrozooplankton interactions, less is known about in-
teractions between small heterotrophic plankton and
macrozooplankton. A number of studies have examined
the grazing effects of macrozooplankton on bacteria
(Güde 1988; Jürgens 1994), flagellated protozoans
(Riemann 1985; Pace and Funke 1991), ciliated protozo-
ans (Pace and Funke 1991; Wickham and Gilbert 1991),
and rotifers (Gilbert 1988; Pace and Vaqué 1994). How-
T. Yoshida (
✉
) · T.B. Gurung · M. Kagami · J. Urabe
Center for Ecological Research, Kyoto University,
509-3 Kamitanakami-Hirano, Otsu, Shiga 520-2113, Japan
Present address:
T. Yoshida, Department of Ecology and Evolutionary Biology,
Corson Hall, Cornell University, Ithaca, NY 14853, USA,
e-mail: ty59@cornell.edu,
Tel.: +1-607-254-4273, Fax: +1-607-255-8088
Oecologia (2001) 129:602–610
DOI 10.1007/s004420100766
Takehito Yoshida · Tek Bahadur Gurung
Maiko Kagami · Jotaro Urabe
Contrasting effects of a cladoceran (Daphnia galeata)
and a calanoid copepod (Eodiaptomus japonicus) on algal
and microbial plankton in a Japanese lake, Lake Biwa
Received: 15 January 2001 / Accepted: 25 June 2001 / Published online: 31 July 2001
© Springer-Verlag 2001