II - Jurnal Inter 1

sustainability
Article
Effects of Harvesting Intensity on the Growth of
Hydrilla verticillata and Water Quality
Shunmei Zhu 1 , Xiaodong Wu 1,2, *, Mengdie Zhou 1 , Xuguang Ge 1 , Xingqiang Yang 1 , Nuoxi Wang 1 ,
Xiaowen Lin 1 and Zhenguo Li 3
1 College of Urban and Environmental Sciences, Hubei Normal University, Huangshi 435002, China
2 Huangshi Main Laboratory of Biodiversity and Environmental Conservation in Lake Watersheds, Hubei
Normal University, Huangshi 435002, China
3 School of Earth Sciences and Spatial Information Engineering, Hunan University of Science and Technology,
Xiangtan 411201, China
* Correspondence: wuxd@hbnu.edu.cn
Abstract: The effects of harvesting intensity on the growth of Hydrilla verticillata (L. fil.) Royle as
well as water quality were studied in controlled experiments to provide a reference for managing
submerged vegetation and purifying the water. The results showed that harvesting had a significant
effect on the recovery of shoot growth and H. verticillata height. The harvested group recovered
completely or mostly after two harvests, but the recovery time was significantly longer than the
control group. The final biomasses of the harvested groups (15%, 30%, 45%, 60%, and 75% harvested)
decreased to 66.61%, 49.13%, 43.95%, 43.77%, and 29.94% of the control group, respectively. The
greater the harvesting intensity, the fewer the winter buds. Harvesting reduced the number of
H. verticillata branches. Repeated harvesting at medium and low intensities during the rapid growth
of H. verticillata effectively improved the water quality and inhibited the propagation and growth of
Citation: Zhu, S.; Wu, X.; Zhou, M.; phytoplankton. These results show that harvesting controlled the growth of H. verticillata, and that
Ge, X.; Yang, X.; Wang, N.; Lin, X.; Li, medium and low harvesting intensities were best when considering water quality.
Z. Effects of Harvesting Intensity on
the Growth of Hydrilla verticillata and Keywords: H. verticillata; harvesting; growth; water quality
Water Quality. Sustainability 2022, 14,
15390. https://doi.org/10.3390/
su142215390
Academic Editors: Wen Xiong, Peter 1. Introduction

A. Bowler, Zhongxin Wu, Xiaoyu Li Submerged macrophytes are the main primary producers in shallow lake ecosystems
and Dongkui Gao where they help maintain the stability of the ecosystems [1]. Their growth and reproduction
Received: 21 September 2022
absorb large amounts of nutrients from the environment and provide space for microor-
Accepted: 16 November 2022
ganisms. In addition, submerged macrophytes improve the surrounding environment
Published: 18 November 2022
by secreting oxygen and organic acids [2]. Water quality has improved and the aquatic
ecosystem has been restored by planting submerged plants in West Lake [3], Donghu [4],
Publisher’s Note: MDPI stays neutral
and Qinhu [5,6]. Submerged macrophytes have also been used as an ecological remediation
with regard to jurisdictional claims in
measure to control eutrophic lakes, and reasonable harvesting of submerged macrophytes
published maps and institutional affil-
is an important part of management [7]. Reasonable harvesting of plants during growth can
iations.
effectively reduce the concentrations of nutrients in lakes, thereby improving lake water
quality [8,9]. At the same time, harvesting can effectively control secondary pollution and
swamping in lakes caused by the excessive growth of submerged macrophytes [10].
Copyright: © 2022 by the authors.
Managing the harvesting of submerged macrophytes to control lake water quality has
Licensee MDPI, Basel, Switzerland. attracted the attention of researchers worldwide [11–13]. Harvesting has been used as an
This article is an open access article effective measure to manage submerged macrophytes in developed countries, such as the
distributed under the terms and United States and the United Kingdom [14]. The harvesting and management technology
conditions of the Creative Commons for submerged macrophytes involves harvesting the pant at different harvesting intensities
Attribution (CC BY) license (https:// and transporting them to the shore through mechanized or manual labor [15]. Mechan-
creativecommons.org/licenses/by/ ical harvesting is a fast and efficient way to remove harmful macrophytes and improve
4.0/). water quality in shallow lakes [8]. However, the artificial harvesting of lake-submerged
Sustainability 2022, 14, 15390. https://doi.org/10.3390/su142215390 https://www.mdpi.com/journal/sustainability

Sustainability 2022, 14, 15390 2 of 21
macrophytes only removes the top parts of the plants, and the harvesting effect lasts only
for a short time but has a high labor cost. Additionally, the harvesting season must be
carefully determined to avoid the growing season of the plants. Harvesting management of
submerged macrophytes in different akes types should be flexible so that the appropriate
harvesting method can be selected according to the need [16]. Different types of submerged
macrophytes have different growth characteristics, different effects on bodies of water, and
different optimal harvesting times and intensities. Previous research on three aquatic plant
species showed that aquatic plants differ in their ability to absorb and control the effects of
nitrogen and phosphorus, and they also have different optimal harvesting times. Increas-
ing the number of harvests can make a significant difference in water quality and plant
growth [17]. Harvesting will affect the competition among different species; thus, affecting
the growth and reproduction of the aquatic plants. After the high-intensity harvesting
of Potamogeton crispus L., the dominant benthic species changed from a clean species to
a medium-polluting species. Over-intensive harvesting leads to higher nutrient levels in
the water and increases the distribution and number of medium-polluting species [18].
The growth of P. crispus is inhibited under medium-intensity harvesting; thus, promot-
ing the growth of Elodea nuttallii (Planch.) H. St. John, and improving its interspecicific
competition position [19]. Harvesting at medium and low intensities from August to late
September did not significantly affect the asexual reproduction of Myriophyllum spicatum L.,
indicating that it would not affect the long-distance expansion of their population, but
high-intensity harvesting significantly inhibited the production of M. spicatum asexual
propagules which would limit population expansion [20]. The greater the harvesting inten-
sity of Myriophyllum aquaticum (Vell.) Verdc., the shorter the remaining main stem, the more
restricted the growth, and the fewer new buds, resulting in a decrease in the number of
branches and buds. Furthermore, plant heights were shorter and plants had fewer branches
with higher harvesting intensities [7]. In addition, sustainable harvesting of submerged
macrophytes helps manage phytoplankton biomass and reduces the accumulation of waste
at the bottom of the lake [21].
Hydrilla verticillata (L. fil.) Royle is often used in ecological restoration projects, and
recent research has focused on its growth characteristics, purification ability, and resistance
to stress [22,23]. The effect of biosorption of heavy metals has also been studied [24], but
less has been reported on harvesting. This study investigated the changes in H. verticillata
growth indices as well as the environmental water quality characteristics before and after
harvesting. This empirical study will serve as a reference for the development of submerged
macrophytes management strategies.
2. Research Methods
2.1. Experimental Design
The H. verticillata seedlings used in the experiment were collected from the shallow
lakes in the middle reaches of the Yangtze River Basin in China. The experiments were
conducted outdoors on the water environment ecological restoration platform of Hubei
Normal University from June to November 2018. The experiments were carried out in
uncovered round translucent plastic vats (70 cm in height and 40 cm in diameter). Each
vat was planted with 40 H. verticillata sprigs (10 ± 1 cm). Qingshan Lake sediment (total
phosphorus: 1201.77 mg/kg, total nitrogen: 620.74 mg/kg, organic matter: 71.32 mg/kg)
was used as the substrate, and the vat was slowly filled with lake water (mean water
temperature: 25.24 ◦ C, dissolved oxygen: 16.81 mg/L; specific conductance: 226.71 µS/cm;
salinity: 0.34 psu; oxidation-reduction potential: 121.51_mV; hydrogen potential: 8.15)
filtered of algae. The experimental harvesting intensities were 15%, 30%, 45%, 60%, and
75% of the plant height. A non-harvested group was maintained as the control group.
Three parallel vats were set up for each treatment group, and a total of 18 plastic vats were
used throughout the experiment (Figure 1).
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2022, 14, FOR PEER REVIEW 3 3ofof26
21
(a) (b)
Figure
Figure1.1.Diagram
Diagram ofof
thethe
H. H.
verticillata experimental
verticillata growth
experimental process.
growth (a,b) show
process. H. verticillata
(a,b) show at differ-
H. verticillata at
ent harvesting intensities.
different harvesting intensities.
The top
The branchesofofH.H.verticillata
topbranches verticillata were
were planted
planted to to growth.
growth. WhenWhenthe the coverage
coverage in
in the
the bucket reached 50%, the H. verticillata growth index was comprehensively
bucket reached 50%, the H. verticillata growth index was comprehensively assessed, and the assessed,
and
H. the H. verticillata
verticillata was harvested
was harvested by heightby height according
according to the harvesting
to the harvesting intensity.intensity. Any
Any residual
residual
plant plant
debris wasdebris was after
removed removed after harvesting.
harvesting. Once the plant Once the plant
coverage in coverage in the vat
the vat returned to
returned
50%, to 50%, itThe
it recovered. recovered. The growth
growth indices of H. indices of H.
verticillata verticillata
were were comprehensively
comprehensively assessed be-
assessed
fore before
the next the next harvesting;
harvesting; the number the number
of H. of H.winter
verticillata verticillata
budswinter buds was
was recorded at recorded
matura-
at maturation and the biomass of H. verticillata was measured upon
tion and the biomass of H. verticillata was measured upon entering the dying stage. H. entering the dying
ver-
stage. H. verticillata was harvested twice (6 August and 15 September 2018)
ticillata was harvested twice (6 August and 15 September 2018) and the relevant experi- and the relevant
experimental
mental datarecorded
data were were recorded for analysis.
for analysis.
2.2. Determination
2.2. Determination ofof H.
H. verticillata
verticillata Growth
Growth Indices
Indices
Nine H. verticillata plants were selected from each vat, and their growth indices
Nine H. verticillata plants were selected from each vat, and their growth indices were
were determined separately. The initial plant height was measured before planting, and
determined separately. The initial plant height was measured before planting, and again
again every 5 d after harvesting. The number of branches was counted every 5 d. Finally,
every 5 d after harvesting. The number of branches was counted every 5 d. Finally, nine H.
nine H. verticillata were randomly selected to count the number of winter buds. The number
verticillata were randomly selected to count the number of winter buds. The number of dead
of dead plants was recorded every 10 days. The fresh and dry weights of H. verticillata were
plants was recorded every 10 days. The fresh and dry weights of H. verticillata were meas-
measured on 6 August, 15 September, and 16 December 2018.
ured on 6 August, 15 September, and 16 December 2018.
2.3. Determination of Water Quality Indicators
2.3. Determination of Water Quality Indicators
The indicators included total nitrogen (TN), total phosphorus (TP), orthophosphate
(PO4The3 -P),indicators included index
the permanganate total nitrogen
(CODMn(TN), total phosphorus
), chlorophyll a (Chl-a),(TP), orthophosphate
colored dissolved or-
(PO 4 -P), the permanganate index (CODMn), chlorophyll a (Chl-a), colored dissolved organic
ganic
3
matter (CDOM), and suspended solids (SS). Water quality was assessed before
matter (CDOM),
harvesting, and and suspendedwere
the indicators solidsmeasured
(SS). Water quality
again was30,
on day assessed
60, andbefore harvesting,
90 after harvest.
and the indicators were measured again on day 30, 60, and 90 after harvest. The specific
The specific methods for determining the water quality indicators can be found in “Water meth-
ods for determining the water quality indicators can be found in “Water
and Wastewater Monitoring and Analysis Methods” (4th edition) (Table 1) [25]. and Wastewater
Monitoring and Analysis Methods” (4th edition) (Table 1) [25].
Table 1. Measurement methods for the water quality indicators.
Table 1. Measurement methods for the water quality indicators.
Measurement Indicators Measurement Methods
Measurement Indicators Measurement Methods
Alkaline potassium persulfate oxidation-UV spectro photo
TN TN Alkaline potassium persulfate oxidation-UV spectro photo metric method
metric method
TP Potassium persulfate digestion method
TP
PO43-P 3
Potassium persulfate digestion method
Ammonium molybdate spectrophotometry
CODMn 4 -P
PO Ammonium molybdate
Acidic method
spectrophotometry
CODMn
Chl-a 90% acetone Acidic method
extraction method
CDOMChl-a Spectral90% acetone
coefficient extraction
absorption method
method
SS CDOM Spectral coefficient
Gravimetric methodabsorption method
SS Gravimetric method
TN: total nitrogen; TP: total phosphorus; PO43-P: orthophosphate; CODMn: permanganate index;
TN: total nitrogen; TP: total phosphorus; PO 3 -P: orthophosphate; COD
Chl-a: chlorophyll a; CDOM: chromophoric 4 Mn : permanganate
dissolved organic matter; index;
SS: suspended Chl-a: chloro-
matter.
phyll a; CDOM: chromophoric dissolved organic matter; SS: suspended matter.
2.4. Data Processing

The data were analyzed using SPSS 23.0 statistical software (SPSS Inc., Chicago, IL,
USA). One-way analysis of variance (ANOVA) was performed to compare the groups. A
p-value < 0.05 was considered significant, and p < 0.01 was considered extremely signifi-
cantly different. The Shapiro-Wilk test and Levene’s test for normality and the chi-square
hypothesis (p > 0.05) for the morphological and water quality data of H. verticillata were
performed before the statistical analysis. If the assumptions of normality or the chi-square
could not be met, the data were transformed and ANOVA was followed by Tukey’s multiple
comparison test.
3. Results and Analysis

3.1. Changes in Growth Indicators
3.1.1. Branch Recovery and Biomass
Harvesting compromised the growth of H. verticillata, with significant effects on
biomass, plant height, and shoot recovery (Figure 2). The growth of H. verticillata branches
was affected to different degrees at all harvesting intensities, and the main branches stopped
growing, which led to the germination of new buds on side branches. On day 20 after
the first harvest, the low-intensity group had recovered and exceeded its pre-harvest
level, with recovery lengths of 19.85 cm and 20.70 cm, respectively, but no canopy had
formed. The 45% intensity harvesting group displayed full branch recovery. However,
branch recovery was significantly slower in the high-intensity harvesting group than in the
medium and low-intensity harvesting groups, with recovery rates of 72% and 66% in the
high-intensity harvesting group, respectively. On day 40 after the first harvest, the branches
of the 15–60% harvesting groups had completely recovered to their pre-harvest levels, but
the plant heights showed a decreasing trend with increasing harvesting intensity at 83, 73,
71, and 63 cm, respectively. The 75% harvesting group had a branch recovery rate of 97.74%.
On day 40 after the second harvest, the final plant height of the control group was 113 cm,
and a thick canopy had formed on the surface of the water. All H. verticillata branches in the
15% harvesting group had recovered to their pre-harvesting state, and plant height reached
88 cm, which was significantly different from the control group (p < 0.05), but no canopy
had formed on the water surface. The recovery rate of the branches in the 30%, 45%, and
60% harvesting groups was 100%, but the plant heights decreased sequentially according
to harvesting intensity and were 78, 71, and 64 cm, respectively. Notably, the recovery rate
of branches in the 75% harvesting group was only 96.55% and plant height was 56 cm.
Significant differences were observed in the biomasses of the harvested groups and
control group (p < 0.05) (Figure 3). By the end of the second harvest, the biomass (dry
weight) of the 15% harvesting group had increased to 7.03 times its initial weight, while the
biomasses of the 30–75% harvesting groups were significantly lower at only 3.54, 2.42, 1.15,
and 0.97 times their initial weights, respectively. The increased biomasses in the harvesting
groups were significantly less than that in the control group. The final H. verticillata biomass
decreased significantly with increasing harvesting intensity, and the final biomass in the
control group was the largest. The final biomasses of the 15–75% harvesting groups were
66.61%, 49.13%, 43.95%, 43.77%, and 29.94%of the control group, respectively.
3.1.2. Number of Branches

The number of H. verticillata branches differed under the different harvesting intensities
(Figure 4). The number of branches in the control group was the highest, reaching 2.78 per
plant, while the numbers of branches in the 15–75% harvesting groups were 2.39, 1.83, 1.67,
0.94, and 0.17 per plant, respectively. The greater the harvesting intensity, the fewer main
stems remained and the fewer new buds sprouted, resulting in a decrease in the number of
branch buds. Furthermore, re-germinated H. verticillata from the roots grew at a slow rate.
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Recovery rate of branches (%)

120 (a) 1st20d 100 120 (b) 100.0
1st20d 1st40d
Plant height (cm)

100
Plant height (cm)

100 1st40d 99.5
90
80 80
99.0
60 80 60
98.5
40 40
70 98.0
20 20
0 60 0 97.5
0 15 30 45 60 75 0 15 30 45 60 75
Harvesting intensity (%) Harvesting intensity (%)
140

(c) 100
120 2nd40d
Plant height (cm)
100 2nd40d
99
80
60 98
40
97
20
0 96
0 15 30 45 60 75
Harvesting intensity (%)
Figure Branchrecovery
Figure 2. Branch recoveryand
andthe finalH.H.verticillata
thefinal verticillata plant
plant height
height under
under different
different harvesting
harvesting intensi-
inten-
Sustainability 2022, 14, x FOR PEER REVIEW
sities.The
ties. Thebar
bargraph
graph represents
represents plant
plantheight,
height,the
theline graph
line represents
graph thethe
represents recovery rate rate
recovery 6 of 26
of branches,
of branches,
and the
and the error
error bars
barsrepresent
representthethestandard
standarddeviations.
deviations.(a–c) indicates
(a–c) the the
indicates H. verticillata rate rate
H. verticillata of branch
of branch
recovery and plant height under different harvesting intensities on day 20 and day 40 after the first
recovery and plant height under different harvesting intensities on day 20 and day 40 after the first
harvest, and day 40 after the second harvest, respectively.
harvest, and day 40 after the second harvest, respectively.
20 30
(a) 1st (b) 2nd
Dry weight removal (g)
25
15
20
10 15
10
5
5
0 0
15 30 45 60 75 15 30 45 60 75
Intensities of harvest (%) Intensities of harvest (%)
100
90 (c) Final biomass
80
Final biomass (g)
70
60
50
40
30
20
10
0
0 15 30 45 60 75
Intensities of harvest (%)
Figure 3. H. verticillata biomasses under different harvesting intensities. Error bars represent standard
Figure standard
deviations. (a,b) denotes the H. verticillata
deviations. verticillata dry weight
weight removal
removal under
under different
harvesting intensities
intensities
after the
after the first
first and
and second
second harvests,
harvests, respectively.
respectively. (c)
(c) denotes
denotes the
the H.
H. verticillata
verticillata final
final biomass
biomass under
under
different harvesting intensities after the first and second harvests.
different harvesting intensities after the first and second harvests.
Sustainability 2022, 14, x FOR PEER REVIEW 7 of 26
3.0
Branch number (number/plant)

2.5
2.0
1.5
1.0
0.5
0.0
0 15 30 45 60 75
Figure 4. The number of H. verticillata branches under
under the
the different
harvesting intensities.
intensities.
3.1.3.
3.1.3. Number
Number of of Winter
Winter Buds
Buds
Harvesting affected the numberofofH.
Harvesting affected the number H.verticillata
verticillata winter
winter buds
buds (Figure
(Figure 5). 5).
By By
thethe
endend
8of
of
ofthe
26
the experiment, the control group had the largest number of winter buds at
experiment, the control group had the largest number of winter buds at 1.44 per plant; while1.44 per plant;
while the 15–75%
the 15–75% harvesting
harvesting groups groups had 0.96,
had 1.19, 1.19, 0.78,
0.96, 0.74,
0.78, and
0.74,0.30
andper
0.30plant,
per plant, respectively.
respectively. The
The number of H. verticillata winter buds decreased significantly with increasing
number of H. verticillata winter buds decreased significantly with increasing harvesting harvesting
inten-
intensity, andnumber
sity, and the the number of winter
of winter buds
buds in in the
the 75% 75% harvested
harvested group
group was thewas the lowest. .
lowest.
1.6
Winter bud number (number/plant)
1.4
1.2
1.0
0.8
0.6
0.4
0.2
0.0
0 15 30 45 60 75
Figure 5. The number of H. verticillata winter buds under the different harvesting
harvesting intensities.
intensities.
3.2. Changes in the Suspended Solids in Water

Harvesting H.
Harvesting H. verticillata
verticillatahad
hadaasignificant
significanteffect
effectonon
SSSS (Figure
(Figure 6).6). Significantly
Significantly fewer
fewer SS
SS were
were detected
detected in the
in the control
control group
group thanthan in the
in the harvesting
harvesting groups,
groups, and and the greater
the greater the har-
the harvest-
vesting intensity,
ing intensity, the greater
the greater the increase
the increase in SS. in
TheSS.SSThe SS concentration
concentration changed changed significantly
significantly during
during harvesting. The SS concentrations in the medium and high-intensity
harvesting. The SS concentrations in the medium and high-intensity harvesting groups were harvesting
groupsthan
higher werethat
higher
in thethan that in thegroup,
low-intensity low-intensity
and the SS group, and the in
concentration SSthe
concentration in the
harvesting groups
harvesting
increased groups
with time increased with time
after harvesting. The SSafter harvesting.increased
concentration The SS concentration increased
significantly with the in-
significantly
crease with theofincrease
in the number harvests in
overthetime.
number of harvests
The increases over
in SS time. The increases
concentrations in SS
in the harvest-
ing groups from the second harvest until the end of the experiment were significantly higher
than those after the first harvest; the SS concentration in the high-intensity harvesting group
was always higher than that of the low-intensity harvesting group; the SS concentration in the
low-intensity harvesting group tended to decrease after harvesting.
concentrations in the harvesting groups from the second harvest until the end of the ex-
periment were significantly higher than those after the first harvest; the SS concentration
in the high-intensity harvesting group was always higher than that of the low-intensity
harvesting group; the SS concentration in the low-intensity harvesting group tended to
decrease after harvesting.
25 25
1st4d (a) 1st20d (b)
20 20
SS (mg/L)
SS (mg/L)
15 15
10 10
5 5
0 0
0 15 30 45 60 75 0 15 30 45 60 75
45
2nd5d (c)
40
35
30
SS (mg/L)
25
20
15
10
5
0
0 15 30 45 60 75
(a)
Figure 6.6. Suspended
Figure Suspended solids
solids (SS)
(SS) concentrations
concentrations in
in water
water under
under different
different H.
H. verticillata
verticillata harvesting
harvesting
intensities. Error
intensities. Errorbars
barsrepresent
representstandard
standarddeviations.
deviations.(a–c)
(a–c) indicates
indicates thethe suspended
suspended solids
solids concen-
concentra-
trations
tions in the
in the water
water under
under different
harvesting intensities
intensities ononday
day4 4and
andday
day20
20after
afterthe
thefirst
first harvest,
harvest,
andday
and day55after
afterthe
thesecond
secondharvest,
harvest,respectively.
respectively.
3.3. Changes
3.3. Changes inin the
the Water
Water Nutrients
Nutrients
3.3.1. Total Nitrogen
3.3.1. Total Nitrogen
The
The different harvesting
harvestingintensities
intensitieshad had significant
significant effects
effects on the
on the TN concentration
TN concentration (Fig-
(Figure 7). TN remained low in the control group and
ure 7). TN remained low in the control group and was lower than that was lower than that in the harvesting
the harvesting
groups
groups throughout
throughout the theexperiment.
experiment.The Thelow-harvesting
low-harvesting intensity
intensity groups
groups (15%(15%
andand
30%)30%)had
had the lowest
the lowest TN concentrations,
TN concentrations, and
and the the medium
medium and high-intensity
and high-intensity harvesting
harvesting groupsgroups
(>45%)
(>45%) had significantly
had significantly higher TN higher TN concentrations
concentrations than the low-intensity
than the low-intensity harvest
harvest groups. groups.
After har-
After harvesting,
vesting, the TN the TN concentrations
concentrations in the in the medium
medium and low-intensity
and low-intensity harvesting
harvesting groups
groups de-
decreased slightly for some time. These results show that the higher the
creased slightly for some time. These results show that the higher the harvesting intensity, harvesting intensity,
the
the higher
higher the
the TNTN concentration,
concentration, and and the
the high-intensity
high-intensity harvesting
harvesting groupgroup maintained
maintained the the
highest TN concentration.
highest TN concentration.
On
On day
day 44after
afterthe
thefirst
firstharvest,
harvest,thetheTNTNconcentrations
concentrations in inthe
thelow-intensity
low-intensityharvesting
harvesting
groups
groups were 0.34 and
were 0.34 and0.350.35mg/L,
mg/L, andandthethe
TNTN concentrations
concentrations in theinmedium
the medium and high-
and high-inten-
intensity harvesting groups were 0.42, 0.49, and 0.57 mg/L, respectively.
sity harvesting groups were 0.42, 0.49, and 0.57 mg/L, respectively. On day 40 after harvest- On day 40 after
harvesting, the TN concentrations in all harvesting groups were significantly
ing, the TN concentrations in all harvesting groups were significantly higher than those higher thanon
those on day 4 after harvesting. The concentrations in the low-intensity
day 4 after harvesting. The concentrations in the low-intensity harvesting groups were 0.94 harvesting groups
were 0.94 mg/L,
and 0.96 and 0.96 mg/L,
which werewhich weretheir
2.7 times 2.7 times
initialtheir initial concentrations,
concentrations, while the TN while the TN
concentra-
concentrations in the medium and high-intensity harvesting groups
tions in the medium and high-intensity harvesting groups were 1.53, 1.58, and 1.75 mg/L, were 1.53, 1.58, and
1.75 mg/L, which increased by 3.6, 3.2 and 3.1 times, respectively. On day 20 after the
which increased by 3.6, 3.2 and 3.1 times, respectively. On day 20 after the second harvest,
second harvest, the TN concentrations in the medium and low-intensity harvesting groups
the TN concentrations in the medium and low-intensity harvesting groups were lower than
were lower than the pre-harvest levels, with concentrations of 0.53, 0.62, and 0.89 mg/L,
the pre-harvest levels, with concentrations of 0.53, 0.62, and 0.89 mg/L, respectively, while the
TN concentrations in the 60% and 75% intensity harvesting group were 1.30 and, 2.20 mg/L
respectively, which was the highest of all of the groups.

respectively, while the TN concentrations in the 60% and 75% intensity harvesting group
were 1.30 and, 2.20 mg/L respectively, which was the highest of all of the groups.
2.4
2.2 1st4d
1st40d
2.0
2nd20d
1.8
1.6
TN (mg/L)
1.4
1.2
1.0
0.8
0.6
0.4
0.2
0.0
0 15 30 45 60 75
Figure 7.
Figure 7.TN
TNconcentrations
concentrationsin water withwith
in water H. verticillata harvested
H. verticillata at different
harvested intensities.
at different Error barsError bars
intensities.
represent standard deviations.
represent standard deviations.
3.3.2. Total Phosphorus
3.3.2. Total Phosphorus
Harvesting H. verticillata had effects on the TP concentrations of the experimental wa-
Harvesting
ter, and the changes H. in
verticillata had effects
TP were different on the
under TP concentrations
the different harvesting of the experimental
intensities (Figure water,
and the harvesting,
8). After changes inthe TPTPwere different in
concentration under the different
the control group was harvesting intensities
significantly lower than (Figure 8).
After
those inharvesting,
the groups. the
The TP concentration
greater the harvesting in intensity,
the controlthe group was
higher the TPsignificantly
concentrationlower
in than
those in the
the water. groups.
Initially, The greater
harvesting caused thea harvesting
slight decreaseintensity, theconcentration,
in the TP higher the TP butconcentration
the TP in
concentration
the increased
water. Initially, with increasing
harvesting causedharvesting intensityin
a slight decrease and
thetheTPeffect became more
concentration, but the TP
pronounced over
concentration time. The with
increased TP concentrations in the medium
increasing harvesting and low-intensity
intensity harvesting
and the effect became more
groups were significantly
pronounced over time. lower
The TP than that in the high-intensity
concentrations harvesting
in the medium and group. In the low-
low-intensity harvesting
intensitywere
groups harvesting group (≤30%),
significantly lower thethan
ratesthat
of decrease
in the in TP reached 83%harvesting
high-intensity and 84%, and the In the
group.
TP rate of decrease was lower in the medium and high-intensity harvesting groups. 11 of 26
low-intensity harvesting group (≤30%), the rates of decrease in TP reached 83% and 84%,
and the TP rate of decrease was lower in the medium and high-intensity harvesting groups.
3.3.3. Orthophosphate
Different harvesting intensities had different effects on PO43-P concentration in the exper-
0.7
imental water (Figure
1st4d 9). The PO43-P concentration increased with increasing harvesting in-
tensity,
0.6 and the PO43-P concentrations in the medium and low-intensity harvesting groups
1st40d
were significantly lower than that in the high-intensity harvesting group, but lower than the
2nd20d
control
0.5 group. After the first harvest, the PO43-P concentration in the harvesting groups exhib-
2nd40d
ited an upward trend and peaked on day 20 after harvesting. On day 20 after the second har-
TP (mg/L)
vest, the change in PO43-P concentrations in the harvested groups exhibited the same trend as
0.4
after the first harvest, and the PO43-P concentration in the harvesting groups peaked at a sim-
ilar maximum value. After harvesting, the PO43-P concentration trended upward with har-
0.3
vesting intensity, i.e., higher harvesting intensities produced more of a change. The concen-
trations of PO43-P in the medium and low-intensity harvesting groups were always lower than
0.2
that in the high-intensity harvesting group. By the end of the experiment, the PO43-P concen-
tration in the 75% intensity harvesting group was 0.078 mg/L, while the concentrations in the
0.1
other harvesting groups and the control group were significantly lower.
0.0
0 15 30 45 60 75
Figure 8.
Figure 8. TP
TPconcentrations
concentrationswith H. verticillata
with harvested
H. verticillata at different
harvested intensities.
at different Error bars
intensities. represent
Error bars represent
standard deviations.
standard deviations.
3.3.3. Orthophosphate
Different harvesting intensities had different effects on PO4 3 -P concentration in the
experimental water (Figure 9). The PO4 3 -P concentration increased with increasing harvest-
ing intensity, and the PO4 3 -P concentrations in the medium and low-intensity harvesting
groups were significantly lower than that in the high-intensity harvesting group, but lower
than the control group. After the first harvest, the PO4 3 -P concentration in the harvesting
groups exhibited an upward trend and peaked on day 20 after harvesting. On day 20 after
the second harvest, the change in PO4 3 -P concentrations in the harvested groups exhibited
the same trend as after the first harvest, and the PO4 3 -P concentration in the harvesting
groups peaked at a similar maximum value. After harvesting, the PO4 3 -P concentration
trended upward with harvesting intensity, i.e., higher harvesting intensities produced more
of a change. The concentrations of PO4 3 -P in the medium and low-intensity harvesting
groups were always lower than that in the high-intensity harvesting group. By the end
3
of the experiment, the PO4 -P concentration in the 75% intensity harvesting group was
0.078 mg/L, while the concentrations in the other harvesting groups and the control group
were significantly lower.
0.10
2nd20d
0.08
PO43-P(mg/L)
0.06
0.04
0.02
0.00
0 15 30 45 60 75
Figure 9.
Figure PO4433-P
9. PO -Pconcentrations
concentrationsin
inthe
thewater
waterof H.verticillata
ofH. verticillata harvested
harvested at
at different
different intensities.
intensities. Error
Error
bars represent standard deviations.
3.4. Changes
3.4. Changes in
in the
the Water
Water Organic
Organic Matter
Matter
3.4.1. Permanganate
3.4.1. PermanganateIndex
Index(COD
(CODMn ))
Mn
The COD
The CODMn index was significantly different under the different harvesting intensities
Mn index was significantly different under the different harvesting intensities
(Figure10).
(Figure 10).The
Thechange
changein in
thethe
CODCOD Mn index
Mn index in theinlow-intensity
the low-intensity harvesting
harvesting group
group was was
greater
greater than that in the medium and high-intensity harvesting groups. The higher the
than that in the medium and high-intensity harvesting groups. The higher the harvesting in-
harvesting intensity, the higher the CODMn index. The CODMn index decreased slightly
tensity, the higher the CODMn index. The COD Mn index decreased slightly after harvesting in
after harvesting in the medium and low-intensity harvesting groups, but the CODMn index
the medium and low-intensity harvesting groups, but the CODMn index generally trended up-
generally trended upward with harvesting time. After two harvests, the CODMn index
ward with harvesting time. After two harvests, the CODMn index of the high-intensity harvest
of the high-intensity harvest group increased significantly and remained above 14 mg/L.
group increased significantly and remained above 14 mg/L. The CODMn index of the medium
The CODMn index of the medium and low-intensity harvesting groups tended to increase,
and low-intensity harvesting groups tended to increase, but changed little overall. The CODMn
but changed little overall. The CODMn index also changed with harvesting intensity. The
index also changed with harvesting intensity. The CODMn index value of the low harvesting
CODMn index value of the low harvesting intensity group was always lower than that of
intensity group was always lower than that of the medium and high-intensity harvesting
the medium and high-intensity harvesting groups, and the higher the harvesting intensity,
groups, and the higher the harvesting intensity, the higher the CODMn index.
the higher the CODMn index.
40
1st20d
35 1st40d
2nd20d
30 2nd40d
CODMn(mg/L) 25
20
15
10
0 15 30 45 60 75
Figure10.
Figure 10.COD
COD Mn indexofofthe
Mnindex thewater
water with
with H.H. verticillata
verticillata harvested
harvested under
under different
different intensities.
intensities. ErrorError
3.4.2.
3.4.2.Colored
ColoredDissolved
DissolvedOrganic
OrganicMatter
Matter
The
TheCDOM
CDOMconcentration
concentrationwas
wassignificantly
significantlydifferent
differentunder
underthe
thedifferent
differentharvesting
harvesting in-
intensities. TheCDOM
tensities. The CDOM concentrations
concentrationsininthe themedium
medium and high-intensity
and harvesting
high-intensity groups
harvesting groups
were significantly higher than that in the low-intensity harvesting group (Figure 11).
were significantly higher than that in the low-intensity harvesting group (Figure 11). The The
CDOM
CDOM concentration
concentration in
in the 75% intensity
the 75% intensity harvesting
harvesting group
group was
was the
thehighest,
highest,reaching
reaching32.50
32.50 µg/L, whereas the CDOM concentrations of the 15%, 30%, 45%, and, 60% intensity
µg/L, whereas the CDOM concentrations of the 15%, 30%, 45%, and, 60% intensity harvest-
harvesting groups were 13.23, 13.93, 15.28, and 15.35 µg/L, respectively, while the control
ing groups were 13.23, 13.93, 15.28, and 15.35 μg/L, respectively, while the control group
group was only 13.00 µg/L. The CDOM concentrations in the harvesting groups increased
was only 13.00 µg/L. The CDOM concentrations in the harvesting groups increased by 2.1%,
by 2.1%, 7.2%, 17.5%, 18.1%, and 150.0%, respectively compared with the control group.
7.2%, 17.5%, 18.1%, and 150.0%, respectively compared with the control group.
35
30
25
CDOM (μg/L)
20
15
10
0
0 15 30 45 60 75
Figure 11. CDOM
Figure 11. CDOMconcentrations
concentrationsininwater
waterof of
H.H. verticillata
verticillata harvested
harvested at different
at different intensities.
intensities. ErrorError
3.5.Chlorophyll
3.5. Chlorophyllaa
Harvesting H.
Harvesting H.verticillata
verticillata resulted
resultedininaasignificantly
significantlyfluctuating
fluctuatingconcentration
concentrationof ofChl-a
Chl-a
in the experimental water (Figure 12). The Chl-a concentration increased initially
in the experimental water (Figure 12). The Chl-a concentration increased initially after the after the
first harvest, but decreased over time. In the low-intensity harvesting group,
first harvest, but decreased over time. In the low-intensity harvesting group, the Chl-a the Chl-a con-
centration remained
concentration remainedlowlowduring the experimental
during the experimental period. The concentrations
period. of Chl-a
The concentrations in the
of Chl-a
medium and high-intensity harvesting groups showed a periodic change
in the medium and high-intensity harvesting groups showed a periodic change of “low- of “low-high-
low”, and were
high-low”, significantly
and were different
significantly from those
different fromof the medium
those and low-intensity
of the medium harvest-
and low-intensity
ing groups.groups.
harvesting After the second
After harvest,harvest,
the second the Chl-atheconcentration in the high-intensity
Chl-a concentration harvest-
in the high-intensity
ing group remained
harvesting at a consistently
group remained high levelhigh
at a consistently andlevel
was significantly higher thanhigher
and was significantly those in the
than
medium and low-intensity harvesting groups. By the end of the experiment,
those in the medium and low-intensity harvesting groups. By the end of the experiment, the Chl-a concen-
trations
the Chl-ainconcentrations
the water of theinharvesting
the watergroups had increased
of the harvesting by 2.56%,
groups 2.03%, 1.15%,
had increased 6.76%,
by 2.56%,
and 17.89%,
2.03%, 1.15%,respectively, compared
6.76%, and 17.89%, with the control
respectively, comparedgroup.
with the control group.
325
300 1st24d
275 1st40d
250 2nd20d
225 2nd40d
Chl-a (mg/L)
200
175
150
125
100
75
50
25
0
0 15 30 45 60 75
Figure12.
Figure 12.Chl-a
Chl-aconcentrations
concentrationsin
inwater
waterof
ofH.
H.verticillata
verticillataharvested
harvestedat
atdifferent
differentintensities.
intensities.Error
Errorbars
bars
representstandard
represent standarddeviations.
deviations.
4.4.Discussion
Discussion
4.1.
4.1.Effects
EffectsofofHarvesting
Harvestingon
onthe
theGrowth
Growthand
andReproduction
ReproductionofofH.
H.verticillata
verticillata
Harvesting intensityand
Harvesting intensity andharvest
harvest frequency
frequency willwill affect
affect the growth
the growth and recovery
and recovery of
of H. ver-
H. verticillata. A total of two harvests were carried out during the experiment.
ticillata. A total of two harvests were carried out during the experiment. No death of H. vertic- No death of
H. verticillata
illata occurredoccurred
in any ofinthe any of the harvesting
harvesting groups ingroups
the firstinharvest,
the firstbut
harvest, but there
there were were
differences
differences in the recovery ability of H. verticillata under the different harvesting
in the recovery ability of H. verticillata under the different harvesting intensities (Figures 13 intensities
(Figures
and 14).13 Inand
our 14). In our experiment,
experiment, H. could
H. verticillata verticillata could
recover orrecover
exceed theor exceed the pre-harvest
pre-harvest levels in a
levels
relatively short period of time under moderate and low-intensity harvesting,harvesting,
in a relatively short period of time under moderate and low-intensity which was
which
similarwas similar
to the study toby
theZuo
study by[26].
et al. Zuo In et addition,
al. [26]. Inthe
addition,
recovery theofrecovery
submerged of submerged
plants after
plants after harvest is closely related to the growing season, climate,
harvest is closely related to the growing season, climate, resources, growth characteristics, resources, growth
characteristics, nutrient reserves and other factors [21,26,27]. The temperature
nutrient reserves and other factors [21,26,27]. The temperature of submerged plants changes of submerged
plants
slowlychanges slowly
and stably. Theand stably. The
influence influence of on
of temperature temperature
submerged onplants
submerged
is lessplants is less
than that on
than that on land plants, but its influence on the seasonal growth of submerged plants is
land plants, but its influence on the seasonal growth of submerged plants is still obvious.
still obvious. The study of Chen et al. showed that the growth of H. verticillata was vigorous
The study of Chen et al. showed that the growth of H. verticillata was vigorous at 25 and 35
at 25 and 35 ◦ C, but was inhibited at low temperatures (5 ◦ C) [28–30]. In our experiment,
°C, but was inhibited at low temperatures (5 °C) [28,29,30]. In our experiment, the recovery
the recovery ability of H. verticillata was significantly decreased, and the recovery time
ability of H. verticillata was significantly decreased, and the recovery time was significantly
was significantly prolonged after the second harvest. The reason may be that the first
prolonged after the second harvest. The reason may be that the first harvest was in early
harvest was in early August and the second harvest was in mid-September. The difference
August and the second harvest was in mid-September. The difference in harvest time led to
in harvest time led to differences in water temperature, air temperature, and other factors,
differences in water temperature, air temperature, and other factors, which affect the growth
which affect the growth recovery of H. verticillata after the second harvest [20]. The choice of
recovery of H. verticillata after the second harvest [20]. The choice of harvesting time had a
harvesting time had
significant effect on thea significant
growth and effect on the
recovery ofgrowth and recovery
H. verticillata. of H. verticillata.
When harvesting was con- When
harvesting
ducted during was conducted
a suitable growthduring
period,a suitable growth
with suitable period,
water and airwith suitable water
temperatures, and air
H. vertic-
illata was able H.
temperatures, verticillata
to quickly was able
recover to quickly recover
to pre-harvest to pre-harvest
levels after harvesting.levels
Whenafter harvesting.
harvesting
When harvesting
was conducted wasthe
during conducted duringperiod
slowly growing the slowly growing period
of H. verticillata, H. verticillata, or in
or in theoflow-temperature
the low-temperature
season, environmental season, environmental
factors such factors suchhave
as water temperatures as water temperatures
not reached have not
the optimal
reached the optimal
combination [31], andcombination
high-intensity[31], and high-intensity
harvesting harvesting
made it difficult made it for
or impossible difficult
the or
growth of H. verticillata to recover.
impossible for the growth of H. verticillata to recover.
100
(a) 1st20d 100 (b) 1st40d
Plant height (cm)

Plant height (cm)
80 90
80
60
70
40
60
20 50
0 15 30 45 60 75 90 0 15 30 45 60 75
120
Branch number (number/plant)
3.0
(c) 2nd40d (d)
110 2.5
Plant height (cm)
100
2.0
90
1.5
80
1.0
70
60 0.5
50 0.0
0 15 30 45 60 75 0 15 30 45 60 75
Figure 13.
Figure
Sustainability 2022, 14, x FOR PEER REVIEW 13. Relationship
Relationshipofofharvesting
harvestingintensity with
intensity plant
with height
plant andand
height the the
number of branches.
number of of 26 (a,b) (a,b)
17 branches.
(c) indicate
(c) indicate relationship
relationship between
betweenharvesting
harvestingintensity
intensityand
andH.H.verticillata
verticillataplant
plantheight
heightonon
day 20,20,
day dayday 40
40 after the first harvest, and day 40 after the second harvest, respectively. (d) indicates the relation-
after the first harvest, and day 40 after the second harvest, respectively. (d) indicates the relationship
ship between harvesting intensity and H. verticillata number of branches.
between harvesting intensity and H. verticillata number of branches.
18
(a) 1st 24 (b) 2nd
16
14 22
12
20
10
8 18
6
16
4
2 14
15 30 45 60 75 15 30 45 60 75
Winter bud number (number/plant)
2.0
52 (c) (d)
48
Final biomass (g)
44 1.5
40
36 1.0
32
28
24 0.5
20
16 0.0
0 15 30 45 60 75 0 15 30 45 60 75
Figure 14.
Figure 14.Relationships of harvesting
Relationships intensity
of harvesting with biomass
intensity withand numberand
biomass of winter
numberbudsof
in winter
H. ver- buds in
ticillata. (a,b) indicates the relationship between harvesting intensity and H. verticillata dry weight
H. verticillata. (a,b) indicates the relationship between harvesting intensity and H. verticillata dry
removal after the first and second harvests. (c) indicates the relationship between harvesting inten-
weight
sity andremoval afterfinal
H. verticillata the biomass.
first and(d)
second harvests.
indicates (c) indicates
the relationship the relationship
between between
harvesting intensity andharvesting
intensity
H. verticillata H. verticillata
andwinter final biomass. (d) indicates the relationship between harvesting intensity
bud number.
and H. verticillata winter bud number.
No mass mortality of H. verticillata was observed under different harvesting intensities,
but a significant effect on plant height and branch growth (Figure 13). branches accelerated
and new apical branches were formed. When the harvesting intensity is high, H. verticillata
mainly sprouted new branches from the base to replace the original main lotus to form new
top branches [31], and the branches that sprout from the base after harvesting become the
most vigorous parts [26]. Because harvesting causes damage to plant tissue and the har-
No mass mortality of H. verticillata was observed under different harvesting intensities,

but a significant effect on plant height and branch growth (Figure 13). branches accelerated
and new apical branches were formed. When the harvesting intensity is high, H. verticillata
mainly sprouted new branches from the base to replace the original main lotus to form
new top branches [31], and the branches that sprout from the base after harvesting become
the most vigorous parts [26]. Because harvesting causes damage to plant tissue and the
harvested plant will not have the ability to transport nutrients from the stem, which
promotes new branches of H. verticillata will mainly germinate from the base [32]. However,
as H. verticillata is a canopy-submerged plant with rapid growth and most of its biomass is
mainly allocated to photosynthetic tissues [33], a large number of branches and leaves are
easy to form a large canopy on the water surface and have strong competition for light and
space [34–36]. Therefore, harvesting effectively reduces the concentration of H. verticillata
biomass on the surface of the water and plays a better purification and landscape effect.
At the same time, the canopy can reduce the shade of light and promote the growth of
submerged plants in the lower layer [37,38]. The final plant height was highest in the
control group, reaching 113 cm, where a canopy formed on the water surface. However, no
canopy was formed on the water surface in the harvesting group. No branches sprouted at
the bases of the harvested plants in the control group and in the medium and low-intensity
harvesting groups. The number of basal branches after two harvests was higher. in the
high-intensity harvesting group than in the medium and low-intensity harvesting groups
due to fewer remaining residual branches. These results indicated that harvesting intensity
had an inhibitory effect on canopy formation and branch growth.
Medium and low-intensity harvesting did not have a significant effect on the regen-
erating ability of H. verticillata, and these lower harvesting levels were actually beneficial
to the growth of H. verticillata (Figure 14). However, with the increase of harvesting
intensity, the final biomass of H. verticillata decreased as harvesting intensity increased.
In the three harvesting treatments, except for the first time in the high-intensity group,
the maximum fresh weight of the other two occurred in the moderate and low-intensity
harvesting groups, and the fresh and dry weights decreased with increasing harvesting
intensity, which was similar to the study by Xu et al. [39]. At the end of the experiment, the
H. verticillata biomass of all harvested groups exhibited overall decreases in final biomass.
This was similar to the results of Yan et al. [18]. who reported that high-intensity harvesting
inhibits plant biomass accumulation. However, in the study of Zuo et al., it was found that
the relative growth rate of the dry mass of Myriophyllum spicatum L. showed a decreasing
trend after continuous harvesting under different harvesting intensities, which may be
related to continuous harvesting and the gradual decline of temperature [20].
Asexual reproduction is the main mode of reproduction, submerged plants can ac-
quire nutrients through asexual reproduction, and population expansion in submerged
macrophytes. As well, it plays an important role in population continuity [40]. But it is not
the only mode. Vegetative reproduction is also a common reproductive mode in submerged
macrophytes, but it is affected by factors such as water transparency and sediment [41].
The main ways of asexual propagation of H. verticillata are broken branches and winter
buds [42–44]. In this study, H. verticillata mainly relied on the production of winter buds
for asexual reproduction. The correlation coefficient between harvesting intensity and the
number of winter buds was as high as 0.92. It is because harvesting reduced the number
of stem nodes of H. verticillata, and H. verticillata invested most of its energy into plant
recovery after harvesting, which reduced the formation of winter buds. This affected the
number of new winter buds, while it did not directly result in the complete disappearance
of all winter buds. So a certain intensity of harvesting would restrict the expansion of the
H. verticillata population to a certain extent.
4.2. Effects of Harvesting H. verticillata on Water Quality

The effects of removing submerged macrophytes on SS occur mainly through the
interception and adsorption of roots, leaves, and resistance to flow resistance, which
reduces the flow velocity and promotes the SS to settle [45]. The harvesting disturbed the
water body, causing the SS to adsorb on the H. verticillata leaves and that settled on the
bottom to re-enter the water body. The greater the harvesting intensity, the greater the
disturbance to the water body. The SS of each harvesting group decreased slightly with the
passage of harvesting time, but by the end of the experiment, the increases in SS reflected
the increases in harvesting intensity by the end of the experiment (Figure 15).
22
18 1st4d (a) 1st20d (b)
20
15 18
16
SS (mg/L)
SS (mg/L)
12
14
9 12
6 10
8
3
6
0 4
15 30 45 60 75 15 30 45 60 75
45
2nd5d (c)
40
35
SS (mg/L)
30
25
20
15
10
15 30 45 60 75
Figure
Figure15.15.Relationship
Relationshipbetween
between harvesting intensity
harvesting intensityof H.
ofverticillata and SS.
H. verticillata and(a–c) indicate
SS. (a–c) the rela-
indicate the
tionship between
relationship harvesting
between intensity
harvesting of H.of
intensity verticillata and suspended
H. verticillata and suspendedsolidssolids
on dayon4day
and4day
and20day
after
20
the first harvest, and day 5 after the second harvest, respectively.
after the first harvest, and day 5 after the second harvest, respectively.
Excesses
Excessesofofnutrients
nutrientssuchsuchasasnitrogen
nitrogenand andphosphorus
phosphorus ininthe
thewater,
water, lead
leadtoto
deterio-
deteri-
rated
oratedwater
waterquality, increased
quality, increased turbidity, andand
turbidity, reduced
reducedlightlight
penetration,
penetration, which can kill
which cansub-
kill
merged macrophytes and further deteriorate the water quality. Nitrogen
submerged macrophytes and further deteriorate the water quality. Nitrogen and phospho- and phosphorus
are
rusessential nutrients
are essential for thefor
nutrients growth of aquatic
the growth plants. plants.
of aquatic Submerged macrophytes
Submerged can remove
macrophytes can
large
removeamounts
large of nitrogen
amounts ofand phosphorus
nitrogen from the water
and phosphorus fromthrough
the waterassimilation, adsorption,
through assimilation,
and synergistic
adsorption, andrelationships with attachedwith
synergistic relationships microorganisms [46]. It has been
attached microorganisms [46].shown that
It has been
shown
most that most macrophytes
submerged submerged macrophytes
will greatly will greatlytocontribute
contribute the removal to the removal of
of nitrogen nitrogen
and phos-
and phosphorus
phorus from eutrophic
from eutrophic lakes [47].lakesThe [47].
studyTheby study
Fu et al.by [48].
Fu etshowed
al. [48]. that,
showedamongthat,various
among
various species of aquatic plants, H. verticillata had the best pollutant-purifying
species of aquatic plants, H. verticillata had the best pollutant-purifying effect towards pol- effect to-
wards in
lutants pollutants
eutrophicinbodies
eutrophic bodies
of water. of water.
Most submergedMostmacrophytes
submerged macrophytes
contribute greatly contribute
to the
greatly tonitrogen
removing the removing nitrogen and
and phosphorus from phosphorus
eutrophic lakesfrom[47].
eutrophic lakes [47].showed
This experiment This experi-
that
ment
the showed that
harvesting of H.the harvesting
verticillata hadofaH. verticillataimpact
significant had a onsignificant impact
the nitrogen, on the nitrogen,
phosphorus, and
phosphorus,
other nutrientsand other
in the nutrients
water in the water
body, which body, which
also significantly also significantly
affected water quality affected
(Figurewater
16),
quality (Figure 16), similar to the study by Zheng and Zhao [49,50].
similar to the study by Zheng and Zhao [49,50]. Harvesting caused significant differences in Harvesting caused
significant
the differences
concentrations of TN,inTP,
theand
concentrations
PO43-P in water.of TN, TP, and
During PO4 3 -P
the entire in water. During
experiment, the greaterthe
entire experiment, the greater the H. verticillata harvesting intensity, the
the H. verticillata harvesting intensity, the higher the nutrient concentrations, but that is justhigher the nutrient
concentrations,
the overall pattern. butRestoring
that is just the overallmacrophytes
submerged pattern. Restoring
combined submerged
with effectivemacrophytes
harvest
combined with
management caneffective harvest management
greatly improve water qualitycan [51]greatly
In thisimprove
experiment water quality
it was found [51] In the
that this
experiment it was found that the nutrients in the harvesting groups increased
nutrients in the harvesting groups increased rapidly and the turbidity of the water increased rapidly and
the turbidity
after harvesting. of the
Then water increased
the nutrient after harvesting.
contents Then theand
decreased steadily nutrient
returned contents
to lowerdecreased
levels,
and water transparency increased. These significant changes were closely related to the re-
lease of nitrogen and phosphorus and the recovery of the growth of H. verticillata. After the
harvesting of H. verticillata, the decay of the plant residues, stems, and leaves in the water
body releases nitrogen, phosphorus, and other substances [50]. Thus, this release of nutri-
ents had a strong effect on the water quality over a short time [52].
steadily and returned to lower levels, and water transparency increased. These significant
changes were closely related to the release of nitrogen and phosphorus and the recovery of
the growth of H. verticillata. After the harvesting of H. verticillata, the decay of the plant
residues, stems, and leaves in the water body releases nitrogen, phosphorus, and other
substances [50]. Thus, this release of nutrients had a strong effect on the water quality over
a short time [52].
0.60 2.0
1st4d (a) 1st40d (b)
0.55 1.8
0.50 1.6
TN (mg/L)
TN (mg/L)
0.45 1.4
0.40 1.2
0.35 1.0
0.30 0.8
15 30 45 60 75 15 30 45 60 75
2.4 0.5
2nd20d (c) 1st4d (d)
2.1
0.4
1.8
TN (mg/L)
TP (mg/L) 0.3
1.5
1.2
0.2
0.9
0.6 0.1
0.3 0.0
15 30 45 60 75 15 30 45 60 75
0.25 2nd40d (e) 0.08 2nd20d (f)
0.20
PO43-P(mg/L)
0.06
TP (mg/L)
0.15
0.04
0.10
0.02
0.05
0.00 0.00
15 30 45 60 75 15 30 45 60 75
Figure Relationshipbetween
16. Relationship
Figure 16. betweenharvesting
harvesting intensity
intensity of H. H. verticillata
of verticillata and and TN,and
TN, TP, TP, PO POin4 3the
and43-P -P water.
in the
water. (a–c) indicates
(a–c) indicates relationship
relationship betweenbetween harvesting
harvesting intensityintensity of H. verticillata
of H. verticillata and TN in and
theTN in the
water onwater
day 4
andday
on day4 40
and after
daythe
40 first
afterharvest,
the firstand day 20and
harvest, after
daythe20second
after theharvest,
secondrespectively. (d,e) indicate(d,e)
harvest, respectively. the
relationship
indicate between harvesting
the relationship betweenintensity of H.
harvesting verticillata
intensity of and TP in the and
H. verticillata waterTPoninday
the4water
after the first
on day
4harvest,
after theand day
first 40 after
harvest, andtheday
second harvest.
40 after (f) indicates
the second harvest.the (f)relationship
indicates thebetween harvesting
relationship betweenin-
tensity of H. verticillata and PO 43-P in water on day 20after the second harvest.
harvesting intensity of H. verticillata and PO4 3 -P in water on day 20 after the second harvest.
Aquatic plants
Aquatic plants affect
affectlight
lighttransmission
transmissionthrough
through coverage,
coverage, which regulates
which regulatesthethe
temper-
tem-
atures at the
peratures at bottom of the
the bottom ofwater bodybody
the water creating a temperature
creating difference
a temperature between
difference the surface
between the
and bottom
surface water. Lower
and bottom water.temperatures will slowwill
Lower temperatures the slow
release
theofrelease
nutrients [53,54]. Harvest-
of nutrients [53,54].
ing reduces reduces
Harvesting the water thesurface
watercoverage by H. verticillata,
surface coverage which increases
by H. verticillata, which the temperature
increases the tem- at
the bottom
perature of the
at the waterofbody
bottom and accelerates
the water body andthe release ofthe
accelerates nutrients
release into the sediment.
of nutrients into the In
our experiment,
sediment. In our harvesting
experiment,disturbs the H.
harvesting verticillata
disturbs the H.and the waterand
verticillata body,
the and
waterthebody,
particles
and
attached
the to the
particles surface to
attached of the
the plants
surfaceand
of in
thethe sediment
plants and re-enter the water,
in the sediment which also
re-enter signif-
the water,
which
icantlyalso significantly
increases increases of
the concentrations thenitrogen,
concentrations
phosphorus,of nitrogen,
and other phosphorus,
substances, andandcauses
other
the water quality to degrade. This experiment also showed that low-intensity harvesting re-
sulted in a smaller disturbance and facilitated the maintenance of good water quality. In ad-
dition, the amount of nitrogen and phosphorus removed by multiple low-intensity harvests
was more than that taken away by one harvest. This finding shows that a lower harvesting
substances, and causes the water quality to degrade. This experiment also showed that
low-intensity harvesting resulted in a smaller disturbance and facilitated the maintenance
of good water quality. In addition, the amount of nitrogen and phosphorus removed by
multiple low-intensity harvests was more than that taken away by one harvest. This finding
shows that a lower harvesting intensity and a reasonable number of harvests result21inofthe
Sustainability 2022, 14, x FOR PEER REVIEW 26
largest water quality improvement, similar to Verhofstad’s study [55]. Therefore, in the
management of submerged plants, it is necessary to carry out reasonable and effective
management of the intensity and frequency of harvesting, which will help improve water
necessary to carry out reasonable and effective management of the intensity and frequency
quality [49,51].
of harvesting,
In addition which will
to the help improve
adsorption water quality
of nutrients, [49,51]. also controlled algal growth
H. verticillata
(FigureIn addition
17). The to Chl-a
the adsorption of nutrients,
concentration in theH. verticillata
water also
serves ascontrolled algal growth
a comprehensive (Fig-
index of
ure
the 17). The Chl-a
number concentration
of algae in a lake andin the
is water
the main serves as a comprehensive
evaluation index of the number
index for eutrophication. This
of algae in a lake
experiment and isthat
showed the the
main evaluation
greater the H. index for eutrophication.
verticillata This experiment
harvesting intensity, the moreshowed
algal
that the greater the H. verticillata harvesting intensity, the more algal blooms
blooms in the water, and the higher the Chl-a concentration. The Chl-a concentration only in the water, and
the higher the Chl-a concentration. The Chl-a concentration only increased
increased temporarily under low-intensity harvesting, and the increase was not obvious. temporarily under
low-intensity
Subsequently,harvesting,
the Chl-aand the increasequickly
concentration was notstabilized
obvious. Subsequently,
and decreased thebelow
Chl-a the
concen-
pre-
tration
harvestquickly
levels.stabilized
The Chl-aand decreased below
concentration underthe pre-harvest
medium levels. The Chl-a
and high-intensity concentration
harvesting was
under medium
significantly and high-intensity
higher harvesting
than in the control was significantly
and low-intensity higher than
harvesting in theand
groups, control and
the high
low-intensity
concentrations harvesting
persistedgroups, and the
for a long time high
andconcentrations
did not returnpersisted for a long time
to the pre-harvest and The
levels. did
firstreturn
not harvest waspre-harvest
to the in Augustlevels.
when The
the temperature
first harvest waswasinhigher,
Augustwhichwhenmeant that the algae
the temperature was
in the medium
higher, which meantand high-intensity
that the algae in harvest groupsand
the medium flourished. Chl-aharvest
high-intensity was well controlled
groups flour-
by theChl-a
ished. secondwas harvest, and water
well controlled byquality
the second recovered
harvest, well.
andHowever, when
water quality H. verticillata
recovered well.
entered a when
However, slow growth period
H. verticillata and the
entered temperature
a slow growth periodwas lower
and the after the second
temperature washarvest,
lower
the improvements
after in water
the second harvest, thequality following
improvements inthe harvest
water were
quality weaker than
following after thewere
the harvest first
harvest.than
weaker This after
further
thedemonstrated
first harvest.thatThisH.further
verticillata can improve
demonstrated water
that quality andcan
H. verticillata inhibit
im-
algae when reasonably managed [39,54,56].
prove water quality and inhibit algae when reasonably managed [39,54,56].
300 1st24d (a) 140 1st40d (b)

120
250
Chl-a (mg/L)
Chl-a (mg/L)
100
200
80
150 60
100 40
20
50
0
15 30 45 60 75 15 30 45 60 75
120
2nd20d (c) 140 2nd40d (d)
100 120
80 100
Chl-a (mg/L)
Chl-a (mg/L)
60 80
60
40
40
20
20
0 0
15 30 45 60 75 15 30 45 60 75
Figure 17.
Figure Relationship between
17. Relationship between harvesting
intensity of H. verticillata
of H. verticillata and
and water
water Chl-a.
Chl-a. (a–d)
(a–d) indicate
indicate
the relationship
the relationship between
between harvesting
intensity of H. verticillata
of H. verticillata and
and Chl-a
Chl-a on
on day
day 24
24 and
and day
day 4040 after
after
the
the first
first harvest,
harvest, and day 20 and day 40 after the second harvest, respectively.
respectively.
Harvesting of submerged macrophytes also affects the release of algae-inhibiting chemi-

cals, thereby weakening the allelopathic effect of the plant. The greater the harvesting inten-
sity, the faster the phytoplankton proliferated and the higher their concentration. This was
reflected by the changes in CDOM, which was released in large amounts during the growth
and death of phytoplankton. Additionally, the degradation of the remnants of the harvested
Harvesting of submerged macrophytes also affects the release of algae-inhibiting

chemicals, thereby weakening the allelopathic effect of the plant. The greater the harvesting
intensity, the faster the phytoplankton proliferated and the higher their concentration. This
was reflected by the changes in CDOM, which was released in large amounts during the
growth and death of phytoplankton. Additionally, the degradation of the remnants of the
harvested H. verticillata released organic matter into the water, which further contributed to
decreasing transparency and changes in water quality [57–59]. In our experiment, when
H. verticillata was harvested in mid-summer, the concentrations of phytoplankton and
CDOM
at the endin of
each harvesting
summer due group increased rapidly,
to temperature and otherbutfactors,
for a short
andtime. The growth of of phy-
the concentrations
H. verticillata slowed at the end of summer due to temperature and other
toplankton and CDOM increased significantly under high-intensity harvesting, factors, and the
maintain-
concentrations of phytoplankton and CDOM increased significantly under high-intensity
ing the higher concentrations for longer (Figure 18).
harvesting, maintaining the higher concentrations for longer (Figure 18).
35
30
CDOM (μg/L)
25
20
15
10
0 15 30 45 60 75
Figure 18. Relationship between harvesting intensity of H. verticillata and phytoplankton and CDOM.
Figure 18. Relationship between harvesting intensity of H. verticillata and phytoplankton and CDOM.
CODMn is an important water quality indicator. Harvesting facilitated algal blooms in
CODMn
the water, is anproduced
which important water
large qualityofindicator.
amounts Harvesting
organic matter through facilitated algal blooms
photosynthesis, and in the
water, which produced
was reflected large amounts
in an immediate of the
increase in organic
CODmatter through
Mn index, whichphotosynthesis,
was also affectedand by was re-
Chl-a. Harvesting
flected increased
in an immediate the turbidity
increase of the
in the COD Mnwater,
index,and the re-released
which organic by
was also affected matter
Chl-a. Har-
promoted
vesting the consumption
increased of potassium
the turbidity permanganate,
of the water, which also resulted
and the re-released organic in the COD
matter promoted
Mn the
index being positively related to the harvesting intensity and the trends
consumption of potassium permanganate, which also resulted in the CODMn index being pos-in algal growth,
i.e., the greater the harvesting intensity, the higher the COD index (Figure 19). The
itively related to the harvesting intensity and the trends in Mn algal growth, i.e., the greater the
CODMn index of the harvesting group was higher in September than in August, which may
harvesting
be related tointensity, the higher
factors such as algalthe CODMn index
reproduction, (Figure 19).conditions,
temperature The CODMn andindex of the harvest-
the growth
ing grouprate
recovery wasofhigher in September than in August, which may be related to factors such as
H. verticillata.
algal reproduction, temperature conditions, and the growth recovery rate of H. verticillata.
Sustainability
Sustainability2022,
2022, 14,
14, x FOR PEER REVIEW
15390 18 of23
21 of 26
10 16
1st20d (a) 1st40d (b)
9 14
CODMn (mg/L)
CODMn (mg/L)
8 12
7 10
6 8
5 6
15 30 45 60 75 15 30 45 60 75
15 2nd20d (c) 22 2nd40d (d)
14 20
13
CODMn (mg/L)
CODMn (mg/L)
18
12
16
11
14
10
9 12
8 10
7 8
6 6
15 30 45 60 75 15 30 45 60 75
Figure19.
Figure 19.Relationship
Relationshipbetween
between harvesting
intensity of verticillata
of H. H. verticillata
and and the COD
the COD Mn index.
Mn index. (a–d)(a–d)
indicate the relationship between harvesting intensity of H. verticillata and the COD
indicate the relationship between harvesting intensity of H. verticillata and the CODMn index on
Mn index on day
20 and day 40 after the first harvest, and day 20 and day 40 after the second harvest, respectively.
day 20 and day 40 after the first harvest, and day 20 and day 40 after the second harvest, respectively.
5. Conclusions
5. Conclusions
The
Theeffects
effectsofofdifferent
differentharvesting
harvestingintensities on on
intensities thethe
growth of H.
growth ofverticillata and and
H. verticillata the the
surrounding
surroundingwater
waterquality were
quality werestudied in in
studied harvesting experiments.
harvesting TheThe
experiments. main conclusions
main conclusions
were
wereasasfollows:
follows:
(1)
(1) Harvesting
Harvestingsignificantly
significantlyaffected
affectedthe growth
the growth of H. verticillata,
of H. limiting
verticillata, the formation
limiting the formation
of
of the canopy.H.
the canopy. H.verticillata
verticillatacould
couldrecover
recover quickly
quickly after harvesting
after at medium
harvesting at medium and and
low intensities but the recovery rate of H. verticillata was significantly
low intensities but the recovery rate of H. verticillata was significantly slower after slower after
two
two high-intensity
high-intensityharvests.
harvests.
(2)
(2) Harvesting reducedthe
Harvesting reduced theaccumulation
accumulation H. H.
of of verticillata biomass.
verticillata biomass.Under medium
Under mediumand and
high-intensity harvesting, H. verticillata plant height and the number of branches
high-intensity harvesting, H. verticillata plant height and the number of branches de-
decreased significantly, resulting in lower final biomass. This finding indicates that
creased significantly, resulting in lower final biomass. This finding indicates that me-
medium and high-intensity harvesting could effectively restrict the accumulation of
dium and high-intensity harvesting could effectively restrict the accumulation of H.
H. verticillata biomass.
verticillata biomass.
(3) Harvesting had a significant effect on water quality. Low-intensity H. verticillata
(3) harvesting
Harvestingimproved
had a significant
the watereffect onwhile
quality, watermedium
quality.andLow-intensity
high-intensityH. harvesting
verticillata har-
vesting
of improved
H. verticillata the waterdeteriorated
significantly quality, whilethe medium and high-intensity harvesting of
water quality.
H. verticillata significantly
(4) Phytoplankton deteriorated
increased significantly inthe
thewater quality. harvesting group, and
high-intensity
(4) the
Phytoplankton increased significantly
CDOM concentrations varied with the in the high-intensity
increase harvesting
in phytoplankton. group,and
Medium and the
CDOM concentrations
low-intensity varied with
harvesting effectively the increase
suppressed the in phytoplankton.
growth Medium
and reproduction and low-
of phyto-
plankton
intensity and the CDOM
harvesting concentration
effectively during the
suppressed the peak
growthH. verticillata growth period.
and reproduction of phyto-
plankton and the CDOM concentration during the peak H. verticillata growth period.
Author Contributions: Conceptualization and methodology, X.W. and X.G.; experiment, M.Z. and
X.Y.;
Authorsoftware and figure images,
Contributions: N.W. and X.L.;
Conceptualization anddata curation, Z.L. X.W.
methodology, and M.Z.;
and writing, S.Z.; reviewM.Z.
X.G.; experiment, and and
editing, X.W. and X.G. All authors have read and agreed to the published version of the manuscript.
X.Y.; software and figure images, N.W. and X.L.; data curation, Z.L., M.Z.; writing, S.Z.; review and
editing, X.W. and X.G. All authors have read and agreed to the published version of the manuscript.
Funding: This research was funded by the Science and Technology Research Project of Education
Department of Hubei Province (No. Q20182502); Graduate Innovative Research Project Construc-
tion of Hubei Normal University (No. 20220454); College Students’ Innovative Entrepreneurial
Training Plan Program (No. D202105061637055045, No. D202205122303594741).
Funding: This research was funded by the Science and Technology Research Project of Education
Department of Hubei Province (No. Q20182502); Graduate Innovative Research Project Construction
of Hubei Normal University (No. 20220454); College Students’ Innovative Entrepreneurial Training
Plan Program (No. D202105061637055045, No. D202205122303594741).
Data Availability Statement: The datasets generated and analyzed during this study are available
from the corresponding author upon request.
Conflicts of Interest: The authors declare no conflict of interest.
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sustainability

Academic Editors: Wen Xiong, Peter 1. Introduction

Sustainability 2022, 14, 15390. https://doi.org/10.3390/su142215390 https://www.mdpi.com/journal/sustainability

2.4. Data Processing

3. Results and Analysis

3.1.2. Number of Branches

Recovery rate of branches (%)

Recovery rate of branches (%)

Plant height (cm)

Plant height (cm)

Recovery rate of branches (%)

Dry weight removal (g)

Sustainability 2022, 14, 15390 6 of 21

Branch number (number/plant)

3.2. Changes in the Suspended Solids in Water

Sustainability 2022, 14, x FOR PEER REVIEW 10 of 26

Sustainability 2022, 14, 15390 10 of 21

Plant height (cm)

Dry weight removal (g)

No mass mortality of H. verticillata was observed under different harvesting intensities,

4.2. Effects of Harvesting H. verticillata on Water Quality

0.25 2nd40d (e) 0.08 2nd20d (f)

300 1st24d (a) 140 1st40d (b)

Harvesting of submerged macrophytes also affects the release of algae-inhibiting chemi-

Harvesting of submerged macrophytes also affects the release of algae-inhibiting

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