全 文 :Drought Resistance of Slope Pioneer Plant
Magnolia multiflora
Pianpian XU, Jianzhu WANG*
College of Biological and Pharmaceutical Sciences, China Three Gorges University, Yichang 443002, China
Supported by National Natural Science Foundation of China (No. 51179094).
*Corresponding author. E-mail: 262800715@qq.com
Received: July 29, 2016 Accepted: August 31, 2016A
Agricultural Science & Technology, 2016, 17(9): 2037-2041, 2045
Copyright訫 2016, Information Institute of HAAS. All rights reserved Agronomy and Horticultrue
D evelopment and constructionof hydropower projects bringenormous social and eco-
nomic benefits. But in the process, due
to the excavation of cuttings and con-
struction of embankment, a large
number of forest vegetation, grass and
farmland are inevitably destroyed, re-
sulting in appearance of large areas of
bare slope on both sides of roads and
around reservoirs[1]. This directly or in-
directly brings amount a series of neg-
ative ecological and environmental
problems, such as water pollution, soil
erosion, biodiversity loss and large
area of slope landslides. Slope vege-
tation restoration is a necessary mean
for protect water environment safety,
prevent soil erosion and maintain re-
gional biodiversity [2]. Vegetation recov-
ery is the basis for ecosystem
restoration. Recovery and reconstruc-
tion of degraded slope vegetation is a
complicated systematic project, and it
requires overall planning, scientific
design and construction, so as to en-
sure the smooth progress of the re
covery process [3 ] . There have been
currently a large number of studies and
reports on screening of suitable organ-
isms and plants for slope, debris and
other engineering disturbed zones [4 -5].
However, researches on hydropower
slope vegetation recovery in dry and
hot valleys have not been effectively
carried out [ 4 ] . There are many re-
searches about the influence of soil
nutrients on vegetation recovery, in-
stead of influence of soil moisture on
vegetation restoration in disturbed
zones [6]. There are many researches
on influence of soil moisture changes
on plant biological characteristics, but
there are rare researches on suitable
Abstract [Objective] This study aimed to reveal responses of Magnolia multiflora to
soil drought stress. [Method] Pot experiment was employed to simulate drought
stress, and photosynthetic and physiological indices of M. multiflora were deter-
mined. [Result] The net photosynthetic rate (Pn) of M. multiflora did not change
significantly under mild drought stress, and reduced significantly under moderate and
severe drought stress. Drought stress reduced stomatal conductance (Gs), transpira-
tion rate (Tr) and intercellular CO2 concentration (Ci), and compared with those in
the control group, the Pn, Gs, Tr and Ci under severe drought stress declined by
61.04% , 86.27% , 87.77% and 42.63% , respectively. The malondialdehyde (MDA)
content in M. multiflora leaves did not increase significantly under mild drought
stress, and increased significantly under moderate and severe drought stress. The
MDA content in M. multiflora leaves under severe drought stress was 1.63 times as
high as that in the control group. The proline (Pro) and soluble sugar contents of
M. multiflora increased significantly with the aggravation of drought stress, and those
under severe drought stress were 8.06 times and 3.16 times respectively higher
than those in the control group. [Conclusion] M. multiflora has a strong drought tol-
erance, and is suitable for growing in relatively arid environment. It can be used as
candidate for vegetation restoration in hydropower engineering slope.
Key words Hydropower engineering; Drought stress; Magnolia multiflora; Photosyn-
thetic characteristics; Physiological characteristics
边坡先锋植物多花木兰的抗旱
性研究
许翩翩, 王建柱* (三峡大学生物与制药学
院,湖北宜昌 443002)
摘 要 [目的]揭示多花木兰对土壤干旱胁迫
的响应。 [方法]采用盆栽控水法模拟干旱条件,
测定多花木兰光合、生理指标。 [结果]多花木兰
的净光合速率在轻度干旱胁迫下变化不明显,
而在中度和重度干旱胁迫下显著降低;干旱胁
迫使气孔导度、蒸腾速率、胞间 CO2 浓度整体
呈现下降趋势, 在重度干旱胁迫下净光合速
率、气孔导度、蒸腾速率、胞间 CO2浓度分别较
对 照 组 下 降 了 61.04% 、86.27% 、87.77% 和
42.63%;多花木兰叶片丙二醛含量在轻度干旱
胁迫下增加不显著,在中度和重度干旱胁迫下
显著增加,重度干旱胁迫下丙二醛含量是对照
组的 1.63 倍;多花木兰脯氨酸和可溶性糖含量
随干旱胁迫的加剧均呈显著增加趋势,在重度
干旱胁迫下其脯氨酸和可溶性糖含量分别是
对照组的 8.06 倍和 3.16 倍。 [结论]多花木兰耐
旱性较强,适宜于偏干旱环境生长 ,可作为水
电边坡植被修复物种。
关键词 水电工程;干旱胁迫;多花木兰;光合
特性;生理特性
基 金 项 目 国 家 自 然 科 学 基 金 (No.
51179094)。
作者简介 许翩翩(1987-),女,湖北松滋人 ,
在读硕士,研究方向为恢复生态学。 *通讯作
者,副教授,博士,主要从事生态学研究,E-mail:
262800715@qq.com。
收稿日期 2016-07-29
修回日期 2016-08-31
DOI:10.16175/j.cnki.1009-4229.2016.09.012
Agricultural Science & Technology 2016
Different lowercase letters stand for significant differences at the
0.05 level.
Fig.1 Comparison of Pn of M. multiflora leaves among different lev-
els of drought stress
Different lowercase letters stand for significant differences at the
0.05 level.
Fig.2 Comparison of Gs of M. multiflora leaves among different lev-
els of drought stress
Different lowercase letters stand for significant differences at the
0.05 level.
Fig.3 Comparison of Tr of M. multiflora leaves among different lev-
els of drought stress
Different lowercase letters stand for significant differences at the
0.05 level.
Fig.4 Comparison of Ci of M. multiflora leaves among different lev-
els of drought stress
growth environment for plants from the
point of view of biochemistry, let alone
influence of drought stress on plants[5].
Magnolia multiflora is an impor-
tant tropical perennial legume with
well-developed root system and strong
resistance. Root nodules of M. multi-
flora can fix nitrogen in the air, in-
creasing soil nitrogen content and im-
proving soil quality. M. multiflora can
grow in relative barren soil. Therefore,
M. multiflora is often applied as pio-
neer plant to soil fixation, slope pro-
tection and ecological restoration.
Light and moisture are important eco-
logical factors. Plant photosynthetic
and physiological characteristics can
reflect is growing status and coordina-
tion with growing environment[7-9], and
can provide reference for judging pio-
neer community stability and succes-
sion[10-11]. In this study, starting from the
actual situation of hydropower engi-
neering disturbed zone, different
drought stress conditions were simu-
lated by pot experiment to study the
photosynthetic and physio-biochemi-
cal characteristics of M. multiflora,
thereby providing theoretical basis for
vegetation restoration and construc-
tion in hydropower engineering slope.
Materials and Methods
Materials
The experiment was conducted in
the ecology test center of China Three
Gorges University in Yichang, Hubei
(111° 1864 E, 30° 4344 N). The
seeds of M. multiflora were purchased
from the seedling base in Chengdu,
Sichuan. Full, uniform-size and un-
damaged seeds were selected. They
were planted in pots that were placed
in a thermostatic and transparent
plastic greenhouse. The tested soil
was typical yellow-brown slope soil
which was crushed and then placed
into 40 pots. In each pot, total 10
seeds were sowed. Irrigation was car-
ried out regularly. After a certain time,
2 seedlings with robust and consistent
growth (seeding height of about 20
cm) were retained in each pot.
Experimental design
The 40 pots were randomly divid-
ed into four groups. Total four treat-
ment groups were designed, including
the control group (soil moisture was
75%-80% of field capacity) and three
drought stress treatment groups (T1,
mild drought stress, soil moisture was
60%-75% of field capacity; T2, moder-
ate drought stress, soil moisture was
50%-55% of field capacity; T3, severe
drought stress, soil moisture was
35%-40% of field capacity). The soil
moisture content was controlled by
weighing method. The pots were
weighed, and their moisture contents
were adjusted according to the experi-
mental design once a day. After 15 d,
various photosynthetic and physio-bio-
chemical indices of young seedlings of
M. multiflora were determined.
Indicator measurement and meth-
ods
The net photosynthetic rate (Pn),
stomatal conductance (Gs), transpira-
tion rate (Tr) and intercellular CO2con-
centration (Ci) of M. multiflora leaves
were determined respectively using Li-
6400 portable photosynthesis analyz-
er. In addition, the water use efficiency
of M. multiflora was calculated (Pn/Tr).
The top 3rd or 4th fully expanded mature
leaf of each plant was sampled. A
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Agricultural Science & Technology2016
Different lowercase letters stand for significant differences at the 0.05 level.
Fig.5 Comparison of water use efficiency of M. multiflora leaves among different levels of
drought stress
Table 1 Comparisons of physio-biochemical indicators of M. multiflora leaves among
different levels of drought stress
Treatment MDA content//mmol/g Pro content//μg/g Soluble sugar content//mg/g
CK 12.36±0.36 c 21.35±3.62 d 2.65±0.12 c
T1 13.57±0.76 bc 76.63±4.21 c 2.89±0.21 c
T2 15.89±0.54 b 133.86±9.62 b 6.62±0.11 b
T3 20.12±1.07 a 172.11±11.25 a 8.37±0.24 a
Different lowercase letters in the same column stand for significant differences at the 0.05
level.
piece of 2 cm × 3 cm was cut off. In a
standard red and blue light leaf cham-
ber, the Pn, Tr, Gs and Ci of each M.
multiflora leaf piece were determined.
Total 10 plants were determined for
each treatment. Before the determina-
tion, the collected M. multiflora leaves
were first placed under saturated light
for 30 min. During the determination,
the conditions of the leaf chamber
were set as follows: temperature 25
℃, saturation intensity 1 000 μmol/(m2·
s) andCO2concentration 400 μmol/mol.
All the determination was carried out
during 9:00 -11:00 am with outdoor
temperature of 25 ℃ . Malondialde-
hyde (MDA) content was determined
by thiobarbituric acid (TBA) method;
Proline (Pro) content was determined
using the ninhydrin method; and solu-
ble sugar content was determined by
anthrone method. For the determina-
tion of physio-biochemical indices,
each index was determined three
times, and total 5 M. multiflora leaves
with similar position and maturity to
those for the determination of photo-
synthetic parameters were determined
for each treatment.
Data processing
Differences in various photosyn-
thetic and physio-biochemical indices
of M. multiflora among different levels
of drought stress were compared with
single factor analysis of variance. Sig-
nificance of differences among differ-
ent treatments was analyzed by LSD
tests. All statistics and analysis was
performed using SPSS 17.0 and Excel
2003.
Results and Analysis
Effects of soil drought stress on
photosynthetic characteristics of M.
multiflora
Comparison of Pn among different
treatments
The Pn of M. multiflora leaves re-
duced gradually with the aggravation
of drought stress (Fig.1). The severer
the drought stress was, the larger the
decrement of Pn was. Under normal
water supply, the Pn of M. multiflora
leaves was 17.42 μmol/(m2·s); under
mild drought stress, the Pn of M. multi-
flora leaves did not change significant-
ly compared with that of the control
group; under moderate and severe
drought stress, the Pn of M. multiflora
leaves reduced by 38.42% and
61.04% respectively compared with
that of the control group (P<0.05). It in-
dicated that mild drought stress had no
significant inhibitory effect on Pn of M.
multiflora leaves, and moderate and
severe drought stress significantly in-
hibited the Pn ofM. multiflora leaves.
Comparison of Gs among different
treatments
The Gs of M. multiflora leaves re-
duced significantly under different lev-
els of drought stress, and it reduced
significantly with the aggravation of
drought stress (Fig.2). Under normal
water supply, the Gs of M. multiflora
leaves was 0.306 mol/(m2·s); the Gs
of M. multiflora leaves declined signif-
icantly with decreased soil moisture
content, and under mild, moderate and
severe drought stress, it was reduced
by 30.39% , 65.69% and 86.27% re-
spectively compared with that of the
control group. It indicated that different
levels of drought stress significantly
inhibited stomatal activity of M. multi-
flora leaves to avoid excessive loss of
nutrients.
Comparison of Tr among different
treatments
The Tr of M. multiflora leaves
showed a significant downward trend
with the aggravation of soil drought
stress. When soil drought stress was
aggravated, in order to effectively re-
duce the leaf water loss, the Tr of M.
multiflora leaves declined in varying
degrees, which was basically the
same with the change trend of Gs. Un-
der the condition of normal soil mois-
ture content, the Tr of M. multiflora
leaves was 3.68 mmol/(m2· s); under
mild drought stress, the Tr of M. multi-
flora leaves was 2.34 mmol/(m2· s);
under moderate drought stress, the Tr
of M. multiflora leaves was 2.21 mmol/
(m2· s); and under severe drought
stress, the Tr of M. multiflora leaves
was 0.45 mmol/(m2· s). With the ag-
gravation of drought stress, differ-
ences were found in downward trend
of Tr of M. multiflora leaves among dif-
ferent treatments. The Tr of M. multiflo-
ra leaves under severe drought stress
was reduced by 87.77% compared
with that of the control group (Fig.3).
Comparison of Ci among different
treatments
The Ci of M. multiflora leaves de-
creased with increased degree of
drought stress, and there were signifi-
cant differences among different treat-
ments. In the control group, the Ci ofM.
multiflora leaves was 230.64 μmol/mol;
under mild drought stress, the Ci of M.
multiflora leaves was 205.65 μmol/mol;
under moderate drought stress, the Ci
of M. multiflora leaves was 183.52
μmol/mol; and under severe drought
stress, the Ci of M. multiflora leaves
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Agricultural Science & Technology 2016
was 132.32 μmol/mol. The Ci of M.
multiflora leaves under severe drought
stress reduced by 42.63% compared
with that of the control group (Fig.4).
Influence of drought stress on wa-
ter use efficiency of M. multiflora
Compared with that of the control
group, the water use efficiency of M.
multiflora leaves under mild drought
stress did not change significantly, and
that under moderate and severe
drought stress declined significantly
(Fig.5). Under normal soil moisture
content, the water use efficiency of M.
multiflora leaves was 4.55 μmol/mmol;
and the water use efficiency of M. mul-
tiflora leaves under moderate and se-
vere drought stress reduced by
35.82% and 62.86% respectively
compared with that of the control
group.
Effects of soil drought stress on
physio-biochemical characteristics
of M. multiflora
Comparison of MDA content among
different treatments
With increased degree of soil
drought stress, the MDA content of M.
multiflora leaves increased continu-
ously. There was no significant differ-
ence in MDA content of M. multiflora
leaves between normal water supply
and mild drought stress (P >0.05),
while there were significant differences
between normal water supply and
moderate, severe drought stress (P<
0.05) (Table 1). Under the condition of
normal water supply, the MDA content
of M. multiflora leaves was 12.36
mmol/g; and the MDA content of M.
multiflora leaves under mild, moderate
and severe drought stress was 1.01
times, 1.28 times and 1.63 times re-
spectively higher than that of the con-
trol group. Under different levels of
drought stress, the membrane system
of M. multiflora was slightly injured.
This might be because that the free
radical scavenging system in M. multi-
flora cells had a high activity, reducing
the injury degree of cell membrane in
membrane lipid peroxidation, thereby
improving the drought tolerance of M.
multiflora.
Comparison of Pro content among
different treatments
With the decrease of soil moisture
content, the Pro content of M. multiflo-
ra leaves showed an upward trend, but
the increasing amplitude differed a-
mong different treatments (Table 1).
The Pro content of M. multiflora leaves
in the control group was 21.35 μg/g;
and that under mild, moderate and se-
vere drought stress was 3.59 times,
6.27 times and 8.06 times respectively
higher than that in the control group.
The Pro accumulation in M. multiflora
increased significantly with the aggra-
vation of drought stress, thereby effec-
tively regulating plant osmotic balance
and avoiding serious water loss from
cells. Thus, M. multiflora adapted to
the arid environment.
Comparison of soluble sugar con-
tent among different treatments
The soluble sugar content of M.
multiflora leaves increased with the
aggravation of drought stress. Under
mild drought stress, the soluble sugar
content of M. multiflora leaves did not
change significantly compared with
that of the control group; and signifi-
cant difference were found in soluble
sugar content of M. multiflora leaves
between moderate, severe drought
stress and normal water supply. The
soluble sugar content of M. multiflora
leaves in the control group was 2.65
mg/g, and that under mild, moderate
and severe drought stress was 1.09
times, 2.50 times and 3.16 times re-
spectively higher than that in the con-
trol group. The soluble sugar content
of M. multiflora leaves increased with
the aggravation of drought stress. M.
multiflora had a strong ability to regu-
late the osmotic effect, and its water
retention capacity was stronger. Thus,
M. multiflora had a stronger adaptabili-
ty to drought stress (Table 1).
Conclusions and Discus-
sion
The arid environment in hy-
dropower engineering slope was sim-
ulated by pot experiment, and the
physiological and ecological adapt-
ability of M. multiflora to different de-
grees of drought stress was studied.
The results showed that M. multiflora
was greatly influenced by changes of
soil moisture content. It had certain
drought tolerance, and was suitable for
growing in mildly or severely arid soil
environment.M. multiflora can be used
as pioneer plant for slope vegetation
restoration.
Effects of soil drought stress on
photosynthetic characteristics of M.
multiflora
Plant growth and stress resis-
tance all directly affected by the plant
photosynthetic capacity, and they can
be reflected by various plant photosyn-
thetic indicators [12]. The results of this
study showed that under different de-
grees of drought stress, the net photo-
synthetic rate, stomatal conductance
and transpiration rate of M. multiflora
showed different variation trends. Un-
der mild drought stress, the variations
of various photosynthetic indicators
were basically the same with those in
the control group; and under moderate
and severe drought stress, various
photosynthetic indicators all reduced
greatly compared with those in the
control group. The appearance of eco-
logical indicators also showed that wa-
ter saturation was the optimal growth
condition for M. multiflora, and the ap-
pearance of various physiological indi-
cators of M. multiflora was the best
under saturated soil moisture content.
Stomatal limitation and non-stomatal
limitation led to reduced photosynthet-
ic rate in plants under drought stress[13].
This study showed that in the very ini-
tial period, stomas restricted the pho-
tosynthesis of M. multiflora, resulting in
reduced net photosynthetic rate. With
the aggravation and extension of
drought stress, the stomatal conduc-
tance and net photosynthetic rate of
plants will reduce significantly. At that
time, non-stomatal limitation is the
main cause of reduced photosynthetic
rate. Transpiration rate is an important
physiological indicator of plants. It can
be used to measure the water balance
of plants, and is the inherent physio-
logical characteristic of plants. Chen et
al.[14] found that the transpiration rate of
peanut and early rice all reduced un-
der mild and moderate drought stress,
which was consistent with the result of
this study. Plants achieve material ex-
change through leaf stomas. The
openness of stomas is stomatal con-
ductance. Plant photosynthesis, tran-
spiration and biomass are also affect-
ed by the stomatal conductance. Zuo
et al. [15] found that stomatal conduc-
tance of cassava was influenced by
soil moisture. When soil moisture con-
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Agricultural Science & Technology2016
tent is below a certain value, relative
soil moisture content is the dominant
factor influencing plant stomatal con-
ductance. Under drought stress,
stomas of M. multiflora leaves will
close to reduce the transpiration water
loss, so as to maintain water balance
in plants. Numerous studies have
shown that reduced photosynthetic
rate of plants is mainly caused by
stomatal closure due to drought stress.
Under drought stress, stomatal limita-
tion and non-stomatal limitation were
the main causes of reduced photosyn-
thetic rate in plant leaves [16]. Farquhar
et al.[17] considered that if stomatal lim-
itation was dominant, reduced net pho-
tosynthetic rate of plants was accom-
panied by a decline in intercellular CO2
concentration; and if non-stomatal
limitation was dominant, reduced net
photosynthetic rate of plants was ac-
companied by a rise in intercellular
CO2 concentration. This study showed
that under mild and moderate drought
stress, the reduced photosynthetic
rate in M. multiflora leaves was mainly
caused by stomatal limitation; and un-
der severe drought stress, the reduced
photosynthetic rate in M. multiflora
leaves was mainly caused by non-
stomatal limitation. Therefore, the de-
gree of drought stress determined that
whether the photosynthesis of M. mul-
tiflora leaves was influenced by stom-
atal limitation or non-stomatal limita-
tion.
Effects of soil drought stress on
physio-biochemical characteristics
of M. multiflora
Under stress, membrane lipid
peroxidation will occur in plant cells,
and the final product is malondialde-
hyde, of which the content often re-
flects the degree of injury in plant
membrane system. The smaller the
increase of plant MDA content was,
the stronger the drought resistance
was, whereas the weaker the drought
resistance was [18]. This study found
that there was no significant difference
in MDA content between normal water
supply and mild drought stress; and
compared with that in the control
group, the MDA content under moder-
ate and severe drought stress
changed significantly. The degree of
membrane lipid peroxidation was in-
creased, but the added values were in
a controllable range, indicating that M.
multiflora had some drought tolerance.
Plant protoplasm and the environment
need to maintain osmotic balance to
maintain normal growth and develop-
ment of plants. In the maintaining pro-
cess of osmotic balance, plants re-
quire a variety of osmotic adjustment
substances, and proline (Pro) is one of
the intracellular osmotic adjustment
substances. Under stress, Pro accu-
mulation in plants will increase in order
to maintain osmotic balance. In an arid
environment, in order to prevent ex-
cessive dehydration, plants will in-
crease Pro accumulation. Thus, the
stronger the plant drought tolerance is,
the higher the plant Pro accumulation
is [19]. This study showed that the Pro
accumulation in M. multiflora leaves
increased significantly under mild,
moderate and severe drought stress to
effectively prevent the loss of body
water and to ensure the normal physi-
ological activities of plants. Soluble
sugar, as a osmotic adjustment sub-
stance, can reflect the osmotic adjust-
ment ability of the plants to a certain
extent. When plants encounter arid en-
vironment, they will adapt to the envi-
ronment through a variety of reactions.
In an arid environment, plants will in-
crease the cell sap concentration
mainly through the accumulation of
various substances, thereby reducing
cellular osmotic potential to increase
the water retention capacity of the
cells. Thus, plants adapt to the arid en-
vironment[20]. The results of this study
showed that the soluble sugar content
of M. multiflora leaves under varying
degrees of drought stress all increased
significantly, thereby effectively reduc-
ing the cell osmotic pressure to ensure
the water content of the cells. It sug-
gested that M. multiflora had certain
adaptability to the arid environment.
At the same time, due to the differ-
ences in habitat, water stress influ-
enced plants from various macro and
micro aspects, such as external mor-
phology, internal physiological and
biochemical characteristics, hormone
regulation and cell ultrastructure.
Therefore, whether M. multiflora is
able to adapt to longer time or deeper
degree of drought stress still needs
more in-depth studies.
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Responsible editor: Yingzhi GUANG Responsible proofreader: Xiaoyan WU
of borer, small brown planthopper and
aphid in seedling stage, and to army-
worm, aphid and millet blast around
heading. Nematod edisease and millet
downy mildew are mainly controlled by
seed pelleting. After thinning, 1 500
times dilution of 4.5% beta-cyperme-
thrin EC is applied for the control of
borer once every 7 d; in case of millet
blast, 500 -800 time dilution of 40%
edifenphos EC or 1 000 times dilution
of 6% kasugamycin WP is sprayed; for
brown streak, 72% agrucultrual strep-
tomycin is sprayed on leaves once ev-
ery 7 d; and 2000 times dilution of 10%
imidaclopridis used for the control of
aphid, and 1 500-2 000 times dilution
of 5% beta-cypermethrinis used for
controlling armyworm.
(6) Harvest at a proper harvest:
For silage, harvest is carried out in
milk-ripe stage generally with a har-
vester, and the foxtail millet for fresh
forage could be cut from heading
stage to flowering stage and could be
fed to animals directly; and for the
preparation of hay, the suitable cutting
period is from full-bloom stage to filling
stage. The forage millet cultivated in
vacant field in autumn could be cut
before early frost. Harvest should be
carried out on sunny days, and drying
or silage operation should be finished
in a short period, so as to avoid rainy
weather, which would affect quality.
Hay should be stacked for storage
timely.
And (7) Sun-drying: For the
preparation of hay, cutting should be
performed on a sunny day, the fresh
grass is sun-dried on the spot, and
when the water content is reduced to
50%, the grass is gathered into smaller
stacks followed by air drying.
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