Aquatic Botany 88 (2008) 121–126

www.elsevier.com/locate/aquabot

The effect of different submerged macrophyte species and biomass

on sediment resuspension in a shallow freshwater lake
En-Hua Li a,b, Wei Li a,*, Gui-Hua Liu a, Long-Yi Yuan a,b
a
Laboratory of Aquatic Plant Biology, Wuhan Botanical Garden, Chinese Academy of Sciences, Wuhan 430074, China
b
Graduate School, Chinese Academy of Sciences, Beijing 100039, China
Received 9 August 2006; received in revised form 11 June 2007; accepted 3 September 2007
Available online 7 September 2007

Abstract
Our study aim was to elucidate the effects of different species of submerged macrophytes and biomass levels on sediment resuspension. For this
purpose experiments were conducted in four different enclosures (Potamogeton maackianus enclosure-PE, Vallisneria spinulosa enclosure-VE,
manipulated enclosure-ME and aquaculture enclosure-AE). A sediment trap method was employed and the experiments were conducted from
summer to winter in a shallow freshwater lake located in central China. A total of 813, 1277, 613 and 693 g DW m2 of sediment was resuspended
in VE, AE, ME and PE, respectively. Our results showed that P. maackianus was more effective than V. spinulosa in restraining sediment
resuspension. Macrophytes reached their maximum effectiveness of reducing resuspension at a certain species-specific biomass threshold above
which biomass effects on resuspension were negligible. The threshold biomass was estimated as 300 g m2 for P. maackianus. Accordingly, within
a lake management and aquaculture aspect, we conclude that as long as biomass does not fall below this threshold its consumption will not
influence sediment resuspension. In the mid-lower reaches of the Yangtze River macrophyte coverage protects the lake sediment against adverse
effects of monsoon wind; if the vegetation is eroded aquaculture sediment resuspension increases significantly.
# 2007 Elsevier B.V. All rights reserved.

Keywords: Macrophyte; Biomass; Sediment resuspension; Sediment trap; Freshwater lake; Aquaculture

1. Introduction
Shallow lakes are strongly influenced by sediment
resuspension which has long been recognized as an important
internal physical process affecting many ecological aspects of
lakes (Bloesch, 1994; Madsen et al., 2001; Pluntke and
Kozerski, 2003). Wind is considered the most important factor
promoting sediment resuspension through wave action. The
entire lake may be affected, implying resuspension of the
surface layer of sediment (Bengtsson and Hellstrom, 1992).
This results in an immediate increase in the concentration of
total suspended solids, total phosphorus (TP) and other
parameters within the overlying water column (Newman and
Reddy, 1992). The internal phosphorus (P) loading induced by
resuspension may reach rates 20–30 times higher than those
released from undisturbed sediment (Søndergaard et al., 1992),
and the potentially available P induced by sediment resuspen-
* Corresponding author. Tel.: +86 27 87510140; fax: +86 27 87510251.
E-mail addresses: liwei@rose.whiob.ac.cn, lapb@public.wh.hb.cn (W. Li).
sion would exceed 20 times that of external loading during the
open water season (Krogerus and Ekholm, 2003). Fish feeding
behavior in the bottom can also contribute to sediment
resuspension (Zambrano et al., 2001). In fact, all processes
resulting in or strengthening the water movement at the
sediment–water interface will cause resuspension.
Submerged macrophytes can reduce current and wave
energies and shelter the sediment from erosion and resuspen-
sion (James et al., 2004). Submerged macrophytes can also
promote sedimentation and serve as effective sediment traps by
intercepting suspended sediment. Therefore, macrophytes play
an important role in stabilizing the sediment and reducing
turbidity in the shallow lake ecosystems (Barko and James,
1998; Madsen et al., 2001; Kufel and Kufel, 2002; Pluntke and
Kozerski, 2003).
Although a macrophyte-dominated lake may remain in a
clear state via stabilizing mechanisms (Scheffer, 1998), effects
of submerged macrophytes on sediment resuspension have so
far only been poorly studied (Horppila and Nurminen, 2003).
As the architecture and distribution of plant tissue over the
water column vary substantially among species and growth

0304-3770/$ – see front matter # 2007 Elsevier B.V. All rights reserved.
doi:10.1016/j.aquabot.2007.09.001
122 E.-H. Li et al. / Aquatic Botany 88 (2008) 121–126
forms, the effects of submerged macrophytes on the hydro-
dynamics and, consequently, on sediment resuspension also
differ between plant stands (Vermaat et al., 2000). Though
James et al. (2004) found that a much higher shear stress was
required to resuspend sediments when high biomass sheltered
the sediment surface from wave action, only limited informa-
tion is available on the impacts of biomass levels of submerged
species on sediment resuspension.
Before the 1990s aquatic macrophytes were once the main
primary producers in most lakes of the mid-lower reaches of the
Yangtze River. However, because of anthropogenic effects, an
increasing number of lakes have become turbid with a resultant
decline in or loss of macrophytes. In fact, lake management in
the district is facing a constant dilemma. The lakes will rapidly
turn into swamps or even land if strong control of the rapid
development of macrophytes is not performed, but will turn
turbid if macrophytes are over-exploited and replaced by
phytoplankton (Li et al., 2002). How to balance macrophyte
conservation and exploitation is a central question in lake
management. We hope that our study on the effects of
macrophytes on sediment resuspension will allow us to offer
some suggestions.
We undertook a 7-month study to reveal the effects of
different species of submerged macrophyte and levels of
biomass on sediment resuspension. Our aims were threefold:
first, what are the effects of the monsoon on sediment
resuspension in lakes of the mid-lower reaches of the Yangtze
River. The monsoon obviously affects the water level as do the
seasonally determined changes in wind direction. Second, what
are the effects of different species and biomasses of submerged
macrophytes on sediment resuspension? Can an appropriate
biomass of aquatic macrophytes be inferred for sustainable
utilization of lake? Third, what are the effects of freshwater
aquaculture on lakes? Fish is an important protein source for the
Chinese population, and since the 1990s China has had the
world’s highest freshwater aquaculture output. However, little
information is available about the environmental consequences
of this large-scale aquaculture on the environmental state of the
lakes, although its destructive effects on aquatic macrophytes
are well known.
2. Materials and methods
2.1. Study area
The study was conducted in Lake Xiliang with an area
of 72 km2, situated in the middle reaches of the Yangtze
River (298510 –308020 N, 1148000 –1148100 E). The mean and
Table 1
The name, dominant species, location and distance to the lake shore of the four enclosures
Enclosure name Dominant species
Vallisneria spinulosa enclosure (VE) V. spinulosa
Aquaculture enclosure (AE) P. maackianus
P. maackianus enclosure (PE) P. maackianus
Manipulated enclosure (ME) P. maackianus
maximum depths of the lake were 1.94 and 4.1 m, respectively.
The dominant wind direction in the area was north to northwest
in winter and south to southwest in summer. In summertime,
total nitrogen (TN) and TP concentrations in the water column
were 522–2405 and 15–45 mg L1, respectively. The concen-
tration of suspended solids (SS) was about 10 mg L1 and
Secchi depth usually ca. 1.5 m in the open water area. Due to
the lake’s relatively clear water and its large stand of submerged
macrophytes, large-scale production of crab and grass carp has
taken place since 1991. At the end of 2003, aquaculture
enclosures were established in a large area of the lake, and
vegetation has almost entirely disappeared (Peng et al., 2003).
2.2. Data collection
Three enclosures (30 m  30 m each) were constructed
from bamboo poles and fish nets. Two hosted mono-dominant
macrophyte stands: Potamogeton maackianus (PE) and
Vallisneria spinulosa (VE), while one enclosure was manipu-
lated (ME). ME was dominated by P. maackianus, which was,
however, removed with iron rakes. The mesh size of the fish net
was 3 cm2 in order to prevent crab and grass carp from entering
the enclosure. The top of the net was placed above the water
surface and the net foot was deeply embedded into the
sediment. A P. maackianus stand located in the central of a
large aquaculture enclosure was chosen as the forth enclosure
(AE). Here, plants were almost depleted by grass carps and
crabs, and the stand was not enclosed in order to maintain the
very low plant biomass level (see Table 1 for details).
The study was initiated on 15 June 2003. Five sediment traps
were placed randomly in each enclosure. The traps were
constructed of PVC pipe with an inside diameter of 7 cm. The
height of each trap was 37 cm, the height:width ratio thus being
>5 (Evans, 1994). The bottom of the trap was closed and
weighed down with a brick. The upper end was tied with a
nylon rope to a pole placed in the sediment, and the trap was
accordingly vertically suspended in the water. The trap bottom
was 10 cm above the sediment surface and did therefore not
affect the sediment resuspension process (Bloesch, 1994).
Samples were collected every month. In each enclosure,
three replicate water samples were taken collected for analysis
of suspended solids (SS) (filtered through Whatman GF/C,
dried at 105 8C to constant weight). The mean vegetation
biomass was determined by collecting the macrophytes 10
times randomly using a 20 cm  20 cm clamp in each
enclosure. After collection, the macrophytes were rinsed and
oven-dried at 80 8C to constant weight. Three replicate 5 cm
top sediment cores were collected, air-dried and ground for
Inferior species Location Distance to
lake shore (m)
Potamogeton maackianus 298350 1600 N, 1148040 2300 E 500
V. spinulosa, Myriophyllum spicatum 298350 0200 N, 1148040 0800 E 1000
298340 5200 N, 1148030 5300 E 1500
298340 5200 N, 1148030 5200 E 1500
 E.-H. Li et al. / Aquatic Botany 88 (2008) 121–126 123

organic content analysis (Bengtsson and Hellstrom, 1992). The
contents of the traps were also filtered through Whatman GF/C,
dried at 105 8C and weighed as gross sedimentation. The
organic fractions of surface sediment, entrapped material and
suspended seston (SS) were calculated from the residue of a
sample ignited at 550 8C for 4 h (Bloesch, 1994).
2.3. Rate of sediment resuspension calculation
wavelengths within different enclosures were compared with
ANCOVA by using the water depth from each enclosure as
covariates. Analysis of covariance was also performed to
compare the rate of sediment resuspension in the four
enclosures by using water depth, average and maximum
wavelengths for each enclosure as covariates (Horppila and
Nurminen, 2005). All data analyses were performed in SPSS
for Windows.

The rate of sediment resuspension was estimated using the 3. Results

method developed by Gasith (1975) for shallow water bodies
with assumption that the organic content of the bottom
sediment differs from that of SS. The method uses the
equation:
fS  fT
R¼S ; exceeded 5 m s1 (Fig. 1). Although the maximum wavelength
fR  fT
where R is the resuspended bottom sediment, S the entrapped
material, f S the organic fraction of S, f R the organic fraction of
surface sediment, and f T is the organic fraction of SS. The
values of gross sedimentation rate were corrected by subtract-
ing the dry weight of suspended matter contained by the water
volume from the gross dry weight in each trap.
3.1. Wind effects
The mean wind velocity was 2.5 m s1 during the study
period, and the maximum wind velocity only occasionally
from June to October occasionally reached or exceeded the
water depth, increased wave activity affected the sediments at
the time of declining water depths from November to January
(Fig. 2). Both maximum and average wavelength in AE were
similar to those in PE and ME, which were significantly higher
than those in VE ( p < 0.05, ANCOVA).

2.4. Wind velocity and wavelength

Resuspension occurs when deep water waves enter water

shallower than one-half of the wavelength (Evans, 1994). To
study the role of wind velocity and wavelength in the rate of
sediment resuspension, the relationship between wavelength
and resuspension rate was explored. The theoretical wave-
lengths were calculated using the equations presented by
Carper and Bachmann (1984): Fig. 1. Maximal and average wind velocity (m s1) in Lake Xiliang, central
China, during the study period.
gT 2
L¼ ;
2p
where L is the wavelength (m), g the gravitational constant and
T is the wave period (s). The value of T is given by
  0:25 
gT gF
¼ 1:20 tanh 0:077 ;
2pU U2
where U is the wind velocity (m s1) and F is the effective fetch
(m). The effective fetch for each wind direction was calculated
according to Carper and Bachmann (1984):
P
xi cos li
F¼ ;
13:5
where xi is the distance from the sampling enclosure to land for
every deviation angle li (up 428). Wind velocity and direction
data were obtained from the Weather Station of Jiayu County
situated 20 km west of the lake.

2.5. Data analyses

The values of f R were compared with those of f T for each Fig. 2. Water depth, one-half of the maximal and average wavelength (cm) in
enclosure using Paired-sample t-test. Average and maximum different enclosures during the study period in Lake Xiliang, central China.
124 E.-H. Li et al. / Aquatic Botany 88 (2008) 121–126

Fig. 3. Biomass (g DW m2) of macrophytes in different enclosures in Lake

Xiliang, central China, during the study period (mean  S.E., N = 10).
Fig. 4. The average daily rate (mean  S.E., N = 5) of sediment resuspension
3.2. Vegetation changes (g DW m2 day1) in different enclosures of Lake Xiliang, central China.

The four enclosures showed different macrophyte composi- 4. Discussion

tions and biomass changes (Table 1, Fig. 3), biomass being
highest in PE and lowest in AE ( p < 0.001, ANOVA). In AE
and ME biomasses exhibited quite rapid changes, while the
biomasses of PE and VE were relatively stable or increased
only slowly.
3.3. Resuspension rates
In each enclosure, the value of f T was significantly higher
than that of f R ( p < 0.001) (Table 2), suggesting that the
resuspension rate is reliably calculated using the Gasith method
(1975).
The rate of sediment resuspension (R) rose in the first few
months and then declined gradually in winter in the VE, PE
and ME enclosures, while in AE it increased and peaked at
8 g DW m2 day1 in December. R differed significantly
among the enclosures (d.f. = 3, F = 9.970, p = 0.001). Thus, R
was significantly higher in AE than in the other enclosures
(P < 0.05), R in PE being significantly lower than that in VE
( p = 0.022). No significant difference was found between VE
and ME or between PE and ME ( p > 0.05, ANCOVA)
(Fig. 4).
4.1. Wind effects
Wind is considered the most important factor affecting the
resuspension process in shallow lakes (Bengtsson and
Hellstrom, 1992; Madsen et al., 2001). Although the recorded
wind was not particularly high during the study period (Fig. 1),
it still disturbed the sediments and resuspension occurred
frequently rather than occasionally. However, the resuspension
rates were substantially lower than those recorded in other
lakes, even in the aquaculture enclosure (Kristensen et al.,
1992; Madsen et al., 2001; Horppila and Nurminen, 2003). The
low resuspension rates may owe to three factors. First, the U-
shaped lake morphology shortened the wind fetch, and the
relatively higher water level in summertime (June to October)
prevented wave effects on the sediment. Second, the dense
macrophyte vegetation shielded the sediment from resuspen-
sion (James et al., 2004), and, third, thickly dotted aquaculture
fish pens decreased wind energies.
In wintertime, at declining water levels, sediment resuspen-
sion increased in AE in December. However, in the three
macrophyte enclosures, the percentage of the water column

Table 2
The organic fraction (%, mean  S.D.) of entrapped material ( fS), suspended seston ( fT), and surface sediment ( fR) in different enclosures during the study period in
Lake Xiliang, central China
July August September October November December January
VE
fS 27.14  1.23 31.50  1.09 29.54  0.38 30.14  0.42 31.30  1.17 28.49   1.19 32.80  0.14
fT 77.49  2.58 83.31  4.97 86.86  2.49 81.26  1.53 79.64  1.91 83.55  5.33 70.20  4.27
fR 11.02  0.69 12.06  1.55 11.65  0.78 12.13  0.96 13.01  1.37 16.60  1.81 17.42  0.54
AE
fS 27.51  3.16 34.77  1.07 25.25  0.22 27.76  0.68 26.43  1.08 33.62  0.71 31.05  4.04
fT 70.60  1.14 71.91  1.96 68.88  0.64 68.93  4.33 66.53  3.53 66.99  2.08 69.33  3.55
fR 11.00  1.45 10.34  0.49 10.75  0.59 12.08  0.51 11.83  0.90 14.54  0.54 15.07  0.88
PE
fS 31.58  1.08 30.13  0.32 31.71  0.94 30.55  0.60 35.54  0.51 30.11  0.74 33.96  1.16
fT 83.27  2.99 87.37  1.11 85.59  3.81 83.90  2.38 82.66  2.40 82.21  2.05 78.07  2.05
fR 13.19  1.73 12.46  0.55 13.67  1.51 12.47  0.43 13.42  0.50 15.73  1.25 14.36  0.25
ME
fS 20.91  1.14 24.99  0.85 36.13  1.15 34.05  0.84 30.94  1.04 35.94  0.61 29.14  1.35
fT 80.84  1.61 82.44  2.08 86.94  2.76 81.51  2.27 80.66  3.98 81.85  2.07 71.73  6.55
fR 10.62  0.80 9.58  0.81 10.96  1.35 9.80  0.58 10.04  0.67 13.40  1.13 10.97  0.28
 E.-H. Li et al. / Aquatic Botany 88 (2008) 121–126
Fig. 5. The biomass (g m2 DW) and resuspension rate (g DW m2 day1) of
the four enclosures during the study period.
infested by submerged macrophytes rose at declining water
depth, enhancing the ability of macrophytes to reduce sediment
resuspension (Madsen et al., 2001), and thus seems to be the
factor responsible for the still decreasing resuspension rates.
4.2. Macrophytes effects
That R was significantly lower in PE than in VE, combined
with the stronger wind effects in PE (Figs. 2 and 4), suggests a
species-specific difference in reducing sediment resuspen-
sion. Thus, P. maackianus seems to have stronger capability
of restraining sediment resuspension than V. spinulosa. We
suggest that the main reason for the observed difference is that
P. maackianus can accumulate more biomass than V.
spinulosa. More biomass seems to entail higher effectiveness
of restraining sediment resuspension. A relatively clear
pattern emerged when all biomass data were pooled and
related to R, while ignoring species and other differences
(Fig. 5). However, even at similar biomass concentrations, P.
maackianus proved to be more effective at restraining
sediment resuspension than V. spinulosa. The biomass of P.
maackianus in ME was almost similar to that of V. spinulosa
in VE during July to September, wind effects being stronger in
ME, however. The insignificant differences in sedimentation
rates show that when biomass levels are comparable, V.
spinulosa was less efficient at reducing sediment resuspen-
sion. This might be the result of a species-specific spatial
distribution of plant biomass.
Although the biomass in PE was significantly larger than in
ME, their sediment resuspension rates were similar, suggesting
the occurrence of a critical point of biomass above which
accumulation of more biomass only has little effect on
restraining sediment resuspension. In our study, this threshold
seemed to be 300 g m2. Even in AE, whose biomass was
below this value, R increased significantly (Fig. 6). However, in
Lake Christina, USA, a biomass of 200 g m2 (Myriophyllum
sibiricum and Chara canescens) was effective at reducing
resuspension (Madsen et al., 2001; James et al., 2004). The
difference in threshold biomass may primarily be determined
by the architecture of the dominant macrophyte species and the
physical properties of the sediments. The variability of wind
field in relation to the morphometric features of the lake may
also be responsible.
During the study period (212 days), a total of 813, 1277, 613
and 693 g DW m2 of sediment was resuspended in VE, AE,
125
Fig. 6. Comparison of resuspension rate (R) (g DW m2 day1) and biomass
(g DW m2) (mean  S.E.) in AE in Lake Xiliang, central China.
PE and ME, respectively. Considering the relatively low
sedimentation rate and the gradually decreasing temperature
during the study period, the sediment traps were exposed for a
relatively long period (Bloesch and Burns, 1980). However, the
comparatively high water temperatures occurring in the
warmest season may lead to mineralization of the organic
content of deposited material in the traps and, with it,
underestimation of the grass sedimentation rate. On the other
hand, the sediment trap method yields higher sedimentation
rates compared with the mass balance method because of the
decreased water turbulence within the traps (Kozerski, 1994).
Our results show the approximate values of sedimentation and
resuspension in Lake Xiliang, but, considering the disturbance
of fish feeding activities to the sediment, wind-induced
resuspension may be overestimated for the aquaculture
enclosure.
4.3. Aquaculture effect
Enclosure culture of herbivorous fishes has proved to be very
effective at controlling the fast development of aquatic
macrophytes, which may create in in-fill processes in lakes
(Wu, 2001; Gu et al., 2006). However, in shallow lake
ecosystems it may trigger a shift from one stable state to
another, which is unfortunate and a challenge to lake managers
whose most common response is to limit the fish yield. Our
study reveals that even at reduced intensity, aquaculture may
have significant effects on sediment resuspension (Fig. 6).
However, if the aquatic macrophyte biomass is maintained at
about 300 g m2, the changes in resuspension rates appear
acceptable.
Acknowledgements
We thank Dr. Erik Jeppesen and Dr. Stephen C. Maberly
for their valuable advice during manuscript preparation.
Three anonymous referees are greatly acknowledged for
their valuable comments. Many thanks to A.M. Poulsen
for her careful linguistic edit. This study was funded by the
State Key Basic Research and Development Plan of
China (2002CB412300), the Innovation Key project of
CAS (KSCX2-1-10) and the Hundred Talents Program
of CAS.
126 E.-H. Li et al. / Aquatic Botany 88 (2008) 121–126
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