Acta Ecologica Sinica 34 (2014) 351355

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Acta Ecologica Sinica

j o u r n a l h o m e p a g e : w w w. e l s e v i e r. c o m / l o c a t e / c h n a e s

Allelopathic control of freshwater phytoplankton by the submerged

macrophyte Najas minor All
Tingting Zhang *, Lu Liu, Xiaohui Yang, Shengjuan Zhang, Wentong Xia, Cheng Li
Provincial Laboratory for Conservation and Utilization of Important Biological Resources in Anhui College of Life Sciences, Anhui Normal University, Wuhu,
241000, China

A R T I C L E

I N F O

Article history:
Received 27 May 2013
Revised 29 July 2014
Accepted 29 September 2014
Available online
Keywords:
Najas minor
Allelopathy
Phytoplankton
Reciprocal response
Growth-inhibition

A B S T R A C T

Water blooms in eutrophic waters have been serious environmental problems in recent years. To explore
effective measures to control this issue has been an interest of research. Our current study was designed to investigate the effects of submerged macrophyte Najas minor All. exudates on the growth of
four freshwater phytoplankton species, toxic Microcystis aeruginosa, toxic Anabaena os-aquae, Chlorella
pyrenoidosa and Scenedesmus obliquus as well as natural phytoplankton assemblages of pond water. We
also conducted a reciprocal response between N. minor and toxic M. aeruginosa using coexistence experiments. Our results showed that: (1) N. minor exudates signicantly inhibited the growth of toxic
M. aeruginosa, toxic A. os-aquae and S. obliquus, with M. aeruginosa being the most sensitive, followed
by toxic A. os-aquae, and S. obliquus the least. N. minor exudates did not show inhibitory effect on
C. pyrenoidosa; (2) N. minor and toxic M. aeruginosa have reciprocal inhibitory effect, and the allelopathic interactions between the two different organisms are density dependent and affect their mutual
growth; (3) N. minor exudates also can induce a decrease in chlorophyll a content and an inhibition in
total dehydrogenase activity of the phytoplankton assemblages. Our present studies indicated the submerged macrophyte N. minor might be a potential useful tool to control phytoplankton blooms.
2014 Published by Elsevier B.V. on behalf of Ecological Society of China.

1. Introduction
In recent years, phytoplankton blooms occurred frequently due
to eutrophication of water, and have caused serious environmental problems all over the world. To effectively utilize water resources
and manage aquatic environment, an important mission for investigators is to nd effective measures to control phytoplankton growth.
In addition to control and reduce those activities that can cause
water pollution, looking for natural resources to treat phytoplankton blooms is critical in improving water quality. Submerged
macrophytes are of great importance for their biological nature in
maintaining water quality of shallow lakes. They are believed to
play a key role in sustaining the clear water state by competing
with phytoplankton for nutrients and light, and directly inhibiting
phytoplankton growth by allelopathy [1]. Certain macrophytes, including Myriophyllum spicatum [2,3], Ceratophyllum demersum, Elodea
spp. [4], and more recently, chara vulgaris [5], Thalia dealbata [6],

* Corresponding author. College of Life Science, Anhui Normal University, The Key
Lab. of Biotic Environment and Ecological Safety in Anhui Province, Wuhu, Anhui
241000, China. Tel.: 86-553-3869297; fax: 86-553-3869571.
E-mail address: cyhztt@mail.ahnu.edu.cn (T. Zhang).
http://dx.doi.org/10.1016/j.chnaes.2014.09.003
1872-2032/ 2014 Published by Elsevier B.V. on behalf of Ecological Society of China.

Hydrilla verticillata [(Linn. f.) Royle] [7], Typha angustifolia L. [8],

Eichharnia crassipes [9], and Stratiotes aloides [10] have been reported to strongly inhibit the growth of freshwater planktonic
cyanobacterial. The inhibitory effects of macrophytes are believed
to be mediated by their anti-phytoplankton and anti-cyanobacterial
components [1].
Najas are submerged macrophytes and grow in fresh water or
salt water around the world, and some species of Najas have also
been shown to have inhibitory effects on phytoplankton. Gross et al.
[11,12] reported that Najas marina ssp. intermedia has allelopathic activity on lamentous or chroococcal cyanobacteria. He et al.
[13] showed an inhibitory activity of Najas minor on S. obliquus. But,
even with aforementioned studies, the knowledge to Najas is still
very limited and what effect of these organisms on other planktons
especially on bluegreen algae remains unclear. To further provide
insights into the N. minor in its potential application in biological
control of phytoplankton blooms, in this study, we tested the effect
of N. minor All. on four common phytoplankton species: the
cyanobacteria toxic M. aeruginosa, the toxic A. os-aquae, the
chlorophytes C. pyrenoidosa, S. obliquus (for comparison) and natural
phytoplankton assemblages; and also carried out coexistence experiments between M. aeruginosa and N. minor. Our ndings
suggested that N. minor, a still little known allelochemical-producing
macrophyte, is a useful tool in controlling planktonic blooms, especially cyanobacteria blooms.

352

T. Zhang et al./Acta Ecologica Sinica 34 (2014) 351355

2. Materials and methods

2.1. Materials
The four phytoplankton species toxic M. aeruginosa FACHB942, toxic A. os-aquae FACHB-245, C. pyrenoidosa FACHB-9 and
S. obliquus Ktz FACHB-416 were purchased from Institute of Hydrobiology, the Chinese Academy of Sciences. Prior to the initiation
of the experiments, the four phytoplankton species were inoculated separately into 250 mL sterilized asks lled with 200 mL
medium (M. aeruginosa and A. os-aquae: BG-11 medium;
C. pyrenoidosa and S. obliquus: HB-4 medium), and cultured for 7 days
in an incubator with a light intensity of 100 mol m2 s1 at 25 1 C
and photoperiod 14:10 h L:D. All the asks were shaken four times
each day.
Submerged macrophyte N. minor All. was collected from a clearwater pond in a suburb of Wuhu City, Anhui Province, China. The
young sprouts (shoots and roots) of N. minor were cleaned in running
water to remove debris and washed in sterile water for three times,
and immersed in 10% sodiumhypochloride solution for 10 s and
then washed in sterile water three times again [14].
The natural phytoplankton assemblages were collected from
Chaohu Lake in Anhui Province, the fth largest freshwater lake in
China. The water sample was ltered with plastic yarn (100 mesh)
so as to remove the groups of algae and small aquatic animals,
and the identication of species, which were dominated by the
genera Microcystis and Anabaena, were conducted under a light
microscope.
2.2. Exudate preparation and N. minor culture
The sterilized living N. minor (100 g) was cultured for 7 days in
a beaker containing 2 L Hoagland medium (0.1) as described previously [15]. The beakers were covered by sterilized lter paper and
then placed in an incubator using light intensity of 120 mol m2 s1
at 25 1 C and photoperiod 14L:10D [5]. Then the culture solution (containing exudates of N. minor) was collected and concentrated
to 200 mL by rotary evaporation, and ltered with Millipore glass
lter (0.22 m pore size). The cultured N. minor was also collected
for latter coexistence experiments.
2.3. Monospecic experiments
The concentrated N. minor exudates 0, 0.1, 0.2, 0.3, 0.4, 0.5 mL
were respectively added to the four monospecic phytoplankton
in the exponential growth phase in 100 mL sterilized Erlenmeyer
asks containing fresh medium as mentioned earlier. The concentrations of N. minor exudates by volume (v/v) were 0, 0.2%, 0.4%, 0.6%,
0.8% and 1.0% (correspondent to 1.0, 2.0, 3.0, 4.0, 5.0 g/L N. minor
living weight, respectively) in the asks. The initial densities of the
four mono-specic phytoplanktons were approximately 105106 cells/
mL, which were cultivated under the aforementioned conditions for
7 days. All the asks were shaken four times each day. To observe
the effect of N. minor exudates on the growth of phytoplankton, 1 mL
culture solution was collected from each of the asks daily, and the
phytoplankton cells were counted under Olympus microscope under
magnications of 400 times [6,7].
2.4. Phytoplankton assemblage experiments
Sterilized asks were lled with 200 mL of new BG-11 medium
and 600 mL of phytoplankton assemblages, then the concentrated
N. minor exudates were added to make the concentration 0, 0.8%
1.0% and 1.2% mL/L (v/v) respectively. All experiments were conducted in triplicate, and the cultural conditions were the same as
those mentioned earlier. During the experiments, a certain amount

of microalgae suspension was collected from each ask every other

day to measure the contents of chlorophyll a using the colorimetric method [16] and TTC reduction, respectively. TTC, a watersoluble compound, can be changed into triphenyl formazan (TPF,
a red-colored compound) by reduction through the catalysis of dehydrogenase [17]. Three milliliter of 0.6% TTC was added into 10 mL
of microalgae suspension with 10 mL of 0.5 M Na2HPO4KH2PO4
(pH 7.4) buffer solution, extracted by ethyl acetate, followed by OD485
measurement. A standard curve was made, and the TPF of treatment groups were calculated based on the curve [6,18].

2.5. Coexistence experiments

Three hundred milliliter of M. aeruginosa in three different initial
densities (low-, intermediate-, and high-density: 103, 105, and
107 cells/mL, respectively) was added into three separate 500 mL
Erlenmeyer asks with sterile silica gel-stoppered. N. minor (5 gliving weight) was added and co-cultured with the M. aeruginosa.
For control, M. aeruginosa with the three densities mentioned earlier
and N. minor (5 g living weight) were cultured individually. One milliliter concentrated stock solution (100) of Hoagland medium was
added to all the aks each day to provide sucient nutrient to the
culture [5]. Triplicates of all the experiments were performed in an
incubator with a light intensity of 120 mol m2 s1 at 25 1 C and
photoperiod 14L:10D for 12 days. M. aeruginosa density was monitored every other day. To evaluate reciprocal effect between
M. aeruginosa and N. minor, at the end of the coexistence experiments, N. minor was separated from M. aeruginosa and cleaned
with distilled water. The biomass of N. minor (living weight) was
measured [5].

2.6. Statistical analysis

Data are expressed as mean SD and analyzed by SPSS 13.0. The
inhibitory ratio (IR) was calculated based on the formula: IR = [1 (Nt/
No)] 100. Here Nt is the density of phytoplankton in treatment and
No is the density of phytoplankton in control. The half inhibitory
concentration (EC50, i.e., the concentration of N. minor exudates that
inhibits normal growth of phytoplankton by 50%, v/v) was obtained using a regression equation of N. minor exudate concentration
and the inhibitory ratio. One-way ANOVA followed by LSD test was
used to test for the differences between individual means. The 0.05
level or less was selected as the point of minimum statistical signicance in every comparison.

3. Results
3.1. Allelopathic effects of N. minor exudates on
monospecic phytoplankton
N. minor exudates signicantly inhibited the growth of
M. aeurginosa, A. os-aquae and S. obliquus in a concentrationdependent manner (Fig. 1). The maximal growth inhibition was
achieved at the exudate concentration of 1.0% (correspondent to
5.0 g/L N. minor living weight) on day 7, with IRs being 94.1%, 88.0%
and 26.9% for M. aeurginosa (Fig. 1A), A. os-aquae (Fig. 1B), and
S. obliquus (Fig. 1C), respectively. The EC50 on the seventh day for
M. aeurginosa and A. os-aquae were 0.29% and 0.34% of N. minor exudates (correspondent to 1.5 g/L and 1.7 g/L N. minor living weight).
In contrast, N. minor exudates at these concentrations had no
signicant effect on C. pyrenoidosa (Fig. 1D). The detailed data on
day 7 was listed in Table 1, in which the IRs for each concentration of N. minor exudate (from 0.2% to 1.0%) were included.

14
M.aeruginosa

10

12

6
4
2
0

6
4
2
0
0 1 2 3 4 5 6 7 8
Time (d)

Algal density (10 cells/mL)

S.obliquus

10

Algal density (10 cells/mL)

8
6
4
2
0

A. flos-aquae

0 1 2 3 4 5 6 7 8
Time (d)

14

10

12

353

14
A

Algal density (10 cells/mL)

12

Algal density (10 cells/mL)

T. Zhang et al./Acta Ecologica Sinica 34 (2014) 351355

14
12

D C.pyrenoidosa

10
8
6
4
2
0
0 1 2 3 4 5 6 7 8

0 1 2 3 4 5 6 7 8
Time (d)

Time (d)

Fig. 1. Allelopathic effects of N. minor exudates at indicated concentrations (v/v) on four freshwater phytoplankton. Results are means SDs (bars) of triplicate experiments. (A) Effect of N. minor on M. aeruginosa; (B) effect of N. minor on A. os-aquas; (C) effect of N. minor on S. obliquus; (D) effect of N. minor on C. pyrenoidos. () Control,
() 0.2%, () 0.4%, () 0.6%, (*) 0.8%, () 1.0%.

3.2. Allelopathy of N. minor exudates on the natural

phytoplankton assemblage
As seen in Fig. 2, the N. minor exudates also showed an inhibitory effect on the natural phytoplankton assemblage with Microcystis
sp. being dominating parts of the lake water in a concentrationdependent manner, as indicated by signicant decrease of the
chlorophyll a and TFP contents with increased N. minor exudate concentrations (compared with the control groups, p < 0.05). The
chlorophyll a contents stand for the phytoplankton biomass, and
the TPF contents can reect the total dehydrogenase activity of the
phytoplankton assemblages, and the decrease of chlorophyll a contents as well as the total dehydrogenase activity might be a reection
of cell structure and function damage due to N. minor exudate
exposure.

3.3. Mutual inhibition between N. minor and M. aeruginosa

To test whether N. minor and M. aeruginosa impact mutually, the
coexistence experiments were conducted. First, the effect of N. minor

on M. aeruginosa was observed. At low density of M. aeruginosa,

N. minor did not show obvious allelopathic inhibition on
M. aeruginosa during the rst 8-day testing period. With the
increase in M. aeruginosa density, the allelopathic inhibition of
N. minor became signicant (p < 0.05 vs. control). The inhibitory effect
of N. minor on the M. aeruginosa growth was most potent at
intermediate-density, which lasted through the whole experimental period. However, at high-density of the M. aeruginosa, the
allelopathic inhibition of N. minor decreased sharply (Fig. 3). The
IRs on day 12 were 42.4%, 89.2% and 29.7% for low-, intermediate-,
and high-densities of M. aeruginosa respectively in coexistence experiments. The difference between intermediate-density and other
two density groups was statistically signicant (p < 0.05).
Next, we studied the inuence of M. aeruginosa on N. minor. The
biomass of N. minor was measured on day 12, the last day of the
experiments with M. aeruginosa at low, intermediate or high densities. At low- and intermediate- densities, M. aeruginosa did not show
signicant impact on N. minor, reected by comparable increases
of biomass as in control. In contrast, at high density (107 cells/
mL), M. aeruginosa was found to have a signicant inhibitory effect
on N. minor (Fig. 4).

Table 1
The IRs (%) of N. minor exudate at different concentrations on different phytoplankton species on day 7.
Phytoplankton species

M. aeruginosa
A. os-aquae
S. obliquus
C. pyrenoidosa

Different concentrations of N. minor exudate (v/v)

0.2%

0.4%

0.6%

0.8%

1.0%

39.7 4.6Aa
27.8 3.8Ab
4.6 0.9Ac
2.8 0.6Ad

70.4 6.8Ba
62.0 5.4Ba
12.0 1.6Bb
0.9 0.4Bc

80.7 5.9Ba
78.7 8.8Ca
11.1 2.3Bb
1.9 0.7Bc

88.9 8.7Ca
84.3 5.9Ca
15.7 2.8Bb
6.4 1.6Cc

94.1 7.1Ca
88.0 7.6Ca
26.9 3.5Cb
4.6 1.1Cc

Different capital letters in same row (AC) and different lowercase letters in same list (ad) represented signicant differences, p < 0.05.

T. Zhang et al./Acta Ecologica Sinica 34 (2014) 351355

Chlorophyll a (left) and TPF(right)

contents (mg/L)

354

2.5
control
0.8mL/L
1.0mL/L
1.2mL/L

2.0
1.5

1.0

* *

*
**

0.5
0.0
0

Time (d)
Fig. 2. Effects of N. minor exudates on chlorophyll a and TPF contents of the natural phytoplankton assemblage from lake water. Results are means SDs (bars) of triplicate
experiments, *P < 0.05 vs. control.

4. Discussions

12

1, 2, 3 : 10 , 10 , 10 control groups

10

4, 5, 6 : 10 , 10 , 10 coexistence
experiments, respectively

respectively)

7
5
3

M.aeruginosa density

(10 ,10 ,10 cells/mL,

In water ecosystem, at least 37 species of submerged macrophytes have been shown to have allelopathic effects on phytoplankton
[19]. Our present studies showed that the exudates of submerged macrophyte N. minor can inhibit the growth of cyanobacteria M. aeurginosa,
A. os-aqua and chlorophytes S. obliquus. Our results also demonstrated a species-specic allelopathic inhibition of N. minor on the

2
1
3

6
4

6
4
2

5
0
0

6
8
Time (day)

10

12

14

Fig. 3. Allelopathic inhibition of N. minor on M. aeruginosa at varying densities. Results

are means SDs (bars) of triplicate experiments.

Biomass of N. minor (g)

7
6

4
3
2
1
0
1

3
Different groups

Fig. 4. Effect of M. aeruginosa at different densities (cells/mL) on the biomasses of

N. minor. Results are means SDs (bars) of triplicate experiments. *p < 0.05 vs. control;
#p < 0.05 vs. the initial biomass. (1) Initial biomass, (2) control group, (3) lowdensity coexistence group, (4) intermediate-density coexistence group, (5) highdensity coexistence group.

phytoplankton. Among the four examined fresh-water phytoplankton, cyanobacteria (M. aeurginosa and A. os-aqua) were found to be
more sensitive to N. minor exudate than the green algae (S. obliquus).
C. pyrenoidosa showed no response to the treatment with N. minor
exudate. These results indicated the selectivity of N. minor in its inhibitory effect on phytoplankton, which is similar to that of many other
aquatic macrophytes including M. verticillatum [20] and Chara [21,22].
On the other hand, phytoplankton species may exhibit varying sensitivity in response to macrophyte allelochemicals. In the published studies
on phytoplanktonic species, cyanobacteria were often found to be signicantly inhibited by allelochemicals of submerged macrophytes
[10,23].
N. minor exudates can inhibit the growth of phytoplankton monospecic species, whether it also has the potential to inhibit natural
phytoplankton assemblages in lake water is not known. So, we collected the phytoplankton assemblages from eutrophic lake water
and measured their response to N. minor exudates. As expected, the
chlorophyll a and TFP contents decreased signicantly, which indicated that N. minor exudates stress resulted in a decrease in the
phytoplankton biomass and an inhibition in the total dehydrogenase activity.
Studies of reciprocal allelopathy among organisms in (terrestrial) plant competition have been focused on the growth of the
target organisms (usually algae and/or cyanobacteria). Macrophyte growth has been rarely investigated. In our current study, we
presented evidence that the macrophyte growth (e.g. N. minor) can
be negatively inuenced by high density of M. aeruginosa. It has been
reported that M. aeruginosa at high density may produce
allelochemicals such as microcystin. It might be toxin like this that
caused an inhibition on the growth of N. minor, thus limited its allelopathic effect. We noticed that the reciprocal allelopathic intensity
was correlated to the organism density/biomass. The allelopathic
inhibition of macrophyte N. minor on M. aeruginosa at intermediate density was more signicant than that at low- and highdensities. The mechanisms underlying low- and high density
conditions might be different: at the low density, M. aeruginosa may
be not sucient to stimulate the N. minor to produce and release
allelochemicals that can inhibit the M. aeruginosa; at high density,
the cyanobacteria itself may produce and release toxins inhibiting
the growth of N. minor, as mentioned earlier. Our current results
are in agreement with previous studies showing M. aeruginosa can
be controlled by C. vulgaris in a density-dependent manner. Our
results further conrmed that allelopathic interactions between the
two different organisms are density-dependent, even though the nutrient and the light supply are sucient [4,5,14,15]. Our current
studies suggest that the macrophyte N. minor might be a useful tool
for controlling phytoplankton blooms, esp. cyanobacteria blooms
in the early stage.

T. Zhang et al./Acta Ecologica Sinica 34 (2014) 351355

Acknowledgments
This study was supported by National Natural Science Foundation of China (No.31170443) and The Provincial Laboratory for
Conservation and Utilization of Important Biological Resources and
the Provincial Key Laboratory of Biotic Environment and Ecological Safety in Anhui.
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