全 文 ：作物学报 ACTA AGRONOMICA SINICA 2008, 34(9): 1615−1622　 http://www.chinacrops.org/zwxb/
ISSN 0496-3490; CODEN TSHPA9 E-mail: xbzw@chinajournal.net.cn

Foundation items: The Anhui Provincial Key Science and Technology Project of China (06003010B); the Nature Science Foundation of Anhui Pro-
vincial Education Department, China (2005KJ169)
Biography: WANG Yun-Sheng (1977–), male, PhD candidate, major research projects: crop genetic breeding and plant physiology.
* Corresponding author: YANG Jian-Bo. Tel: +86 (0)551-2160212; E-mail: yjianbo@263.net
Received(收稿日期): 2008-01-04; Accepted(接受日期): 2008-03-26.
DOI: 10.3724/SP.J.1006.2008.01615
Physiological Research on the Difference of Bentazon Tolerance in Wild
Type Rice and Sensitive Lethal Mutants
WANG Yun-Sheng1,2, LU Xu-Zhong2, SUN Ming-Na3, SONG Feng-Shun2, LI Li 2, GAO Tong-Chun3,
and YANG Jian-Bo2,*
(1 College of Biology Science, Anhui Agricultural University, Hefei 230036, Anhui; 2 Rice Research Institute, Anhui Academy of Agricultural Sci-
ences, Hefei 230031, Anhui; 3 Plant Protection Research Institute, Anhui Academy of Agricultural Sciences, Hefei 230031, Anhui, China)
Abstract: Bentazon sensitive lethal (bsl) mutants can be applied in hybrid rice (Oryza sativa L.) breeding for improving seed
production and quality. This study was to elucidate the physiological mechanism of bentazon tolerance in rice. Tolerant rice varie-
ties (W6154s and Norin 8) as well as their corresponding mutants, sensitive to bentazon, were employed in this study. Plant net
photosynthetic rate (Pn), chlorophyll fluorescence, and the level of superoxide radical ( 2O
− ) as well as the contents of chlorophyll
(Chl) and malondialdehyde (MDA), were analyzed for both tolerant and sensitive rice plants treated with bentazon. After treat-
ment, the two sensitive mutants showed a significant reduction in Pn at 0.5 h. A continuous decrease of Chl contents was found
over the first 3 d whereas a significant increase of MDA contents was noticed on the 3rd day and thereafter. Analysis of chloro-
phyll fluorescence revealed a bentazon-induced increase in the proportion of the reduced state of QA. In the early stage after ben-
tazon treatment, wild types and their mutants showed no significant difference in the alteration of Pn and chlorophyll fluorescence.
While these two parameters then increased progressively in both wild types and remained low in the mutants. A significant gen-
eration of 2O
− was found over the 5 d period in the mutants. Both wild types and mutants contained the same level of bentazon
after 2 h of treatment. Bentazon content dropped to barely detectable amount in the wild type varieties at 1 d. However, the mu-
tants retained a substantial amount of the herbicide after 5 d. It is proposed that the herbicide might inhibit rice photophythesis and
accumulation of oxidative stress with the treatment of bentazon in both lethal mutants. The damaging effect on PS II system can
be significantly alleviated in the wild type varieties due to a higher rate of catabolism of the herbicide.
Keywords: Oryza sativa; Bentazon; Sensitive lethal mutant; Tolerance; Physiological research
水稻及其敏感突变体苯达松抗性的生理生化差异研究
王云生 1,2 陆徐忠 2 孙明娜 3 宋丰顺 2 李 莉 2 高同春 3 杨剑波 2,*
(1 安徽农业大学生命科学学院, 安徽合肥 230036; 2 安徽省农业科学院水稻研究所, 安徽合肥 230031; 3 安徽省农业科学院植物保护
研究所, 安徽合肥 230031)
摘 要: 苯达松敏感致死(bentazon sensitive lethal, bsl)基因在杂交水稻(Oryza sativa L.)混播制种和杂交稻种纯度鉴定
等方面具有广阔的应用前景。以水稻品种农林 8 号(N8)、W6154s 和其对应的 bsl 突变体农林 8 号 m (N8m)和 8077s
为材料, 分析了苯达松处理对叶片中叶绿素(Chl)含量、丙二醛(MDA)含量、氧自由基含量、净光合速率(Pn)、叶绿素
荧光参数等生理生化指标的影响及苯达松含量的变化, 旨在揭示水稻苯达松抗性差异的生理机制。结果表明, 苯达松
处理使 bsl 突变体叶片光系统 II 中还原性 QA组分积累, 光合电子传递受阻, 光合能力丧失, 氧自由基伤害积累, Chl
降解、质膜氧化加剧, 植株死亡。叶片中苯达松残留含量分析表明, 较强的苯达松代谢能力是抗性品种免受苯达松伤
害的主要原因。
关键词: 水稻; 苯达松; 敏感致死突变体; 抗性; 生理
Hybrid rice technology is feasible and readily
successful for raising rice yield potential. However,
hybrid rice production is very labor intensive because
the false hybrid seeds (self-pollination of sterile lines)
1616 作 物 学 报 第 34卷

have to be identified and removed, usually by hand.
Chemical, morphological, and genetic markers have
been investigated to identify the purity of rice hybrid
seeds, even at the seedling stage[1-3].
Bentazon [3-isopropyl-1H-2,1,3-benzothiadiazine-
4(3H)-one 2,2-dioxide], known as basagran commer-
cially, is used as a post-emergence herbicide with high
selectivity for killing broadleaf weed. In general,
common rice varieties (wild types) are all tolerant to
bentazon, and can survive under foliar spray of 0.5%
bentazon solution during the whole life cycle[3-4].
Norin 8m (N8m) is a bentazon sensitive lethal (bsl)
mutant, obtained by γ-ray radiation from a japonica
rice, Norin 8 (N8)[5]. The 8077s is also a γ-ray induced
mutant from an indica photoperiod-temperature sensi-
tive genic male sterile (P/TGMS) line, W6154s[4,6].
Previous genetic studies on the cross-pollinated and
self-pollinated progenies of the two mutants have con-
firmed that the bsl phenotype of N8m or 8077s is con-
trolled by a single recessive gene[4-6].
In hybrid breeding system, tagging the bsl gene to
the male sterile line can ensure the hybrid seeds free of
false hybrids (self-pollination of sterile lines) by
spraying bentazon at the seedling stage. On the other
hand, if the bsl gene is transferred into a restorer line,
the restorer line is easily killed by spraying bentazon
immediately after pollination, while the sterile line
maintains normal growth and development. Therefore,
the bsl gene can be used an reasonable selective
chemical marker in hybrid rice production so as to
reduce labor cost[4,6].
Many researches have been performed to find
molecular markers that are linked with the bsl gene,
and to provide an effective and powerful tool for
marker-assisted selection for breeding purposes, in
addition to the efforts to tag bsl gene in map-based
cloning[6-7]. The results of map-based gene cloning
showed that these mutants were caused by the muta-
tion of the gene CYP81A6 encoding cytochrome P450
monooxygenases[8]. However, there is little informa-
tion about physiological basis of bentazon resistance
or sensitivity in wild type and bentazon-lethal mutants.
The objectives of this study were to clarify physio-
logical mechanisms of different responds to bentazon
treatment between the wild type and their mutants. The
results provided sufficient evidence for further bsl
gene isolation and characterization as well as its app-
lication in rice breeding.
1 Materials and methods
1.1 Plant materials
Seeds of N8m and its wild type N8 were kindly
provided by Dr. Taketa from Japan. Seeds of mutant
8077s and its wild type W6154s were obtained from
Dr. Ji-Wen Zhang, Hubei Academy of Agricultural
Sciences, China. Surface-sterilized rice seeds were
placed on moistened filter paper in a germination dish
and incubated at (28±2)˚C for one week. Seedlings
were transplanted into plastic trays (80 cm × 40 cm ×
10 cm) filled with sandy loam soil. Plants (40 per tray)
were grown under an irradiance of 800 μmol m−2 s−1
photosynthetically active radiation (PAR) from growth
lamps (400–700 nm) and a 10/14 h (light/dark) photo-
period. Temperature in the greenhouse was maintained
at (30±2)˚C/(20±2)˚C (light/dark). At six-leaf stage, a
thin layer of an aqueous solution containing 400 mg
L−1 bentazon (Sigma Co., USA) and 0.25% (V/V)
Tween 20 was daubed on the surface of the fourth leaf
with a fur brush. Distilled water containing 0.25%
(V/V) Tween 20 was daubed as control.
1.2 Determination of chlorophyll and malondi-
aldehyde contents
Changes in chlorophyll (Chl) and malondialde-
hyde (MDA) contents were used as the indices of leaf
injury by bentazon. The two indices were evaluated at
0, 1, 2, 3, 4, and 5 d after treatment. For Chl extraction,
approximately 0.5 g fresh leaves were homogenized in
10 mL of 80% acetone and followed by centrifugation
at 5 000 × g for 5 min. The supernatant was measured
for absorbance at 663 and 645 nm[9]. To extract MDA,
0.5 g fresh leaves were homogenized in 10 mL of 5%
(W/V) trichloroacetic acid (TCA) and the mixture was
centrifuged at 4 000 ×g for 10 min at 4˚C. The MDA
content was determined by measuring the absorbance
at 532 and 600 nm[10].
1.3 Determination of superoxide radical
0.5 g frozen leaf segments were homogenized
with 5 mL of 65 mmol L−1 potassium phosphate buffer
(pH 7.8) and centrifuged at 5 000 × g for 10 min. 2O
−
was measured as described by monitoring the nitrite
formation from hydroxylamine in the presence of 2O
− [11].
A standard curve with NO2− was used to calculate the
production rate of 2O
− from the chemical reaction of
2O
− and hydroxylamine.
1.4 Measurements of photosynthesis and re-
lated parameters
Leaf net photosynthetic rate (Pn) was measured
using a portable photosynthesis system (Model CI-310,
CID Inc., USA) operating in the semi-open mode. Ex-
ternal air was scrubbed of CO2 and mixed with a sup-
ply of pure CO2 to result in a reference CO2 concen-
tration of 350 μL L−1.
Chlorophyll fluorescence was measured using a
portable fluorometer (PAM 2100, Walz, Effeltrich,
Germany). Steady-state fluorescence during diurnal
illumination was measured using a leaf clip (Model
2030-B, Walz). Dark adaptation of leaves also took
place with the leaf clips (Walz) for 20 min before
第 9期 WANG Yun-Sheng et al.: Difference of Bentazon Tolerance in Wild Rice and Lethal Mutants 1617


measurements. The maximum amount of fluorescence
(Fm) was recorded with a saturating flash of light,
while the minimal amount of fluorescence (Fo) was
recorded in the dark. The ratio of variable fluorescence
(Fv) to Fm (Fv/Fm) was calculated by Fv/Fm = (Fm −
Fo)/Fm. Fluorescence yield (ФPSII) was calculated by
ФPSII = (Fm′− Fv)/Fm′, where Fm′ was detected from the
leaf sample after application of a saturating light pulse
of high quantum flux density (5 000 μmol m−2 s−1), and
F was the fluorescence intensity in the steady state of
photosynthesis. Photochemical quenching coefficient
(qp) was calculated by qp = (Fm′ − Fv)/ (Fm′ − Fo′).[12]
1.5 Extraction and quantification of bentazon
High performance liquid chromatography (HPLC)
was performed to determine Bentazon content in treated
leaves[13]. Fresh leaves were sampled once every 2 h after
application of bentazon solution. The leaf tissue (5 g)
was homogenized in 80 mL of methanol. After filtering
the mixture through the filter paper, the volume of the
extraction solution was reduced to 5 mL in a speed vac-
cum concentrator. The concentrated sample was dis-
solved in 80 mL of ethanamide and extracted twice with
30 mL of ethane. A standard calibration was prepared
using the pure bentazon (Sigma Co. USA).
1.6 Statistical analyses
Statistical differences among genotypes and dif-
ferent time periods of bentazon treatment were ana-
lyzed following the Student’s t-test with SPSS soft-
ware (Chicago, IL, USA). Differences were considered
to be significant at P<0.05 probability level.
2 Results
2.1 Injury indices of bentazon
N8 and its mutant N8m had lower chlorophyll
contents [(1.81 ± 0.15) and (1.73 ± 0.17) mg g−1 FW,
mean ± SE] than W6154s [(2.095 ± 0.04) mg g−1 FW]
and its corresponding mutant 8077s [(2.04 ± 0.15) mg
g−1 FW]. After bentazon treatment, leaf Chl contents in
both wild type varieties decreased over the first 2 d,
and then increased gradually, which was similar to
those of untreated controls at 5 d after bentazon treat-
ment (Fig. 1). In contrast, Chl contents in the two mu-
tants decreased continuously during the 5-d period of
observation, and N8m was more affected than 8077s.
On the fifth day after treatment, Chl contents in N8m
and 8077s were 16.09% and 39.61% of their corre-
sponding wild type varieties. MDA content highly
increased starting at 3 d in both mutants, while it re-
mained relatively low in the wild types during the
whole period of treatment (Fig. 2).
2.2 Superoxide radical
Level of 2O
− in leaves of the wild types in-
creased rapidly at 1 d after bentazon treatment, fol-
lowed by an continuous decrease and approached con-
trol level at 3 d (Fig. 3). The level of 2O
− in the two
mutants increased quickly and could not recover over
the 5 d period. On the fifth day after treatment, the
parameter in N8m and 8077s was increased by 89.06%
and 74.33% compared with those in their correspond-
ing wild type varieties, respectively.

Fig. 1 Effect of bentazon (400 mg L−1) on the chlorophyll (chl) content in control and treated leaves of different rice varieties
CK means treatment with distilled water containing 0.25% (V/V) of Tween 20. Each value is the mean of three replicates and is representative
of three independent experiments. The vertical and horizontal bars indicate standard errors of means.

Fig. 2 Effect of bentazon (400 mg L−1) on the malondialdehyde (MDA) content in control and treated leaves of different rice varieties
CK means treatment with distilled water containing 0.25% (V/V) of Tween 20. Each value is the mean of three replicates and is representative
of three independent experiments. The vertical and horizontal bars indicate standard errors of means.
1618 作 物 学 报 第 34卷


Fig. 3 Effect of bentazon (400 mg L−1) on the level of -2O in control and treated leaves of different rice varieties
CK means treatment with distilled water containing 0.25% (V/V) of Tween 20. Each value is the mean of three replicates and is representative
of three independent experiments. The vertical and horizontal bars indicate standard errors of means.

2.3 Photosynthesis
The average Pn of control leaves was significantly
lower in N8 and its mutant N8m [(17.31 ± 0.74) and
(16.9 ± 0.87) μmol m−2 s−1, mean ± SE] than in
W6154s and its mutant 8077s [(20.52 ± 1.19) and
(18.68 ± 1.11) μmol m−2 s−1]. The average Pn of the
wild types decreased rapidly and became undetectable
(zero) at approximately 0.5 h after bentazon treatment,
followed by a continuous increase and approached
control level at 1 d (Fig. 4). The two mutants could not
recover from the bentazon damage and Pn remained
near zero over the 5 d period. It was noted that after
treatment, Pn of W6154s recovered more quickly than
that of N8.

Fig. 4 Effect of bentazon (400 mg L−1) on the net photosynthetic rate (Pn) (at 1 000 μmol m−2 s−1) in control and treated leaves of
different rice varieties
CK means treatment with distilled water containing 0.25% (V/V) of Tween 20. Each value is the mean of three replicates and is representative
of three independent experiments. The vertical and horizontal bars indicate standard errors of means.

2.4 Chlorophyll fluorescence
Figure 5 presents the changes in the minimal
fluorescence yield (Fo) and the maximal efficiency of
PSII photochemistry (Fv/Fm) in dark-adapted state in
control and treated leaves. The average Fv/Fm showed
a significant reduction in the first 2 h after treatment.
The mean value was 12.3% for N8 and N8m, and
10.3% for W6154s and 8077s (Fig. 5-B). After the
initial reduction, the Fv/Fm value increased gradually
in the two wild type varieties. One day after treatment,
the N8 had 92.5% of Fv/Fm compared with control and
W6154s recovered completely (99.9%). For the two
mutants, the Fv/Fm decreased, and the average Fo in-
creased continuously over the 5 d observation period
(Fig. 5).
The above results indicate that bentazon treat-
ment might have a significant impact on the primary
photochemistry of PS II in dark-adapted leaves of both
mutants. However, the leaves of the wild type varieties
were able to recover from bentazon injury over time.
We further investigated whether bentazon treatment
could induce any significant changes in PS II photo-
chemistry in light-adapted leaves at 2 and 24 h time
period after treatment. In the control leaves without
bentazon application, the two genotypes exhibited
similar changes (ФPSII and qp decrease) with increasing
第 9期 WANG Yun-Sheng et al.: Difference of Bentazon Tolerance in Wild Rice and Lethal Mutants 1619



Fig. 5 Effect of bentazon (400 mg L−1) on the minimal fluorescence yield (Fo) and the maximal efficiency of PS II photochemistry
(Fv/Fm) in dark-adapted state in control and treated leaves of different rice varieties
CK means treatment with distilled water containing 0.25% (V/V) of Tween 20. Each value is the mean of three replicates and is representative
of three independent experiments. The vertical and horizontal bars indicate standard errors of means.

PFD (Fig. 6 and Fig. 7). After bentazon application,
the two parameters (ФPSII and qp) of the two wild type
varieties and mutants decreased to zero at 2 h after
bentazon treatment (Fig. 6). Differences between wild
type varieties and mutants were observed at 24 h after
treatment. In the wild type varieties, trend of changes
in these two parameters was similar between treatment
and the control. The only difference was that the ab-
solute values were significantly lower in the treated
leaves with a ФPSII of 46.0% for N8 and 20.1% for
W6154s. However, the parameters in the mutants re-
mained much lower compared to the wild type varie-
ties and the untreated control (Fig. 7).
2.5 Absorption and catabolism of bentazon
Bentazon content in plants increased very fast
and reached its peak [N8, (18.47 ± 0.56) mg g−1 FW;
N8m, (18.62 ± 0.57) mg g−1 FW; W6154s, (19.41 ±
0.15) mg g−1 FW; 8077s, (19.56 ± 0.50) mg g−1 FW,
mean ± SE] within the first 2 h after treatment. Dif-
ferences in reduction of leaf bentazon content among
the wild type varieties and their mutants were pre-
sented in Fig. 8. One day after treatment, the content
of bentazon dropped to (5.58 ± 0.32) mg g−1 FW in
N8m and (6.28 ± 0.31) mg g−1 FW in 8077s, and re-
mained at this level during the 5 d treatment period
(data not presented). In the wild type varieties, benta-
zon contents decreased much more dramatically, and
became barely detectable after 1 d of treatment. In
addition, bentazon contents reduced faster in W6154s
than in N8.

Fig. 6 The light response curves of actual quantum yield of PS II photochemistry (ФPSII, A) and photochemical quenching (qp, B) in
control and treated leaves of different rice varieties at 2 h bentazon (400 mg L−1) treatment
CK means treatment with distilled water containing 0.25% (V/V) of Tween 20. Each value is the mean of three replicates and is representative
of three independent experiments. The vertical and horizontal bars indicate standard errors of means.

1620 作 物 学 报 第 34卷


Fig. 7 The light response curves of actual quantum yield of PSII photochemistry (ФPSII, A) and photochemical quenching (qp, B) in
control and treated leaves of different rice varieties at 24 h bentazon (400 mg L−1) treatment
CK means treatment with distilled water containing 0.25% (V/V) of Tween 20. Each value is the mean of three replicates and is representative
of three independent experiments. The vertical and horizontal bars indicate standard errors of means.


Fig. 8 Changes of bentazon content in treated leaves of different
rice varieties in 24 h after bentazon (400 mg L−1) treatment
Each value is the mean of three replicates and is representative of
three independent experiments. The vertical and horizontal bars
indicate standard errors of means

3 Discussions
Previous researches showed that the primary ac-
tion of bentazon is to compete for the binding site in
the D1 protein with plastoquinone (QB) in the photo-
system II[14-15]. The herbicide will be converted
through the cytochrome P450 monooxygenase system
into inactive metabolites once it enters the cells of
tolerant plants[15-16]. Plant response to bentazon de-
pends on absorption, catabolism, and the affinity of the
herbicide with the target site of D1 protein in PS II.
Therefore, changes in Chl contents, MDA contents,
level of 2O
− , net photosynthetic rate, chlorophyll
fluorescence, and bentazon content in treated leaves
were compared using the two mutants and their wild
types in this study.
3.1 The mechanism of bentazon sensitivity in
mutants
Chlorophyll fluorescence provides information
about the state of PS II. It is an estimate of the extent
to which PS II is using the energy absorbed by chlo-
rophyll and the extent to which it is being damaged by
excess light. In this study, the changes in the chloro-
phyll fluorescence were monitored to explore the
mechanism whereby the Pn was reduced in treated
leaves of the mutants. The data showed that foliar ap-
plication of bentazon induced an irreversible inhibition
of photophythesis in both mutants. The inhibition, in-
dicated by a sustained reduction in Fv/Fm and an in-
creased Fo (Fig. 5), is associated with a decline in the
intrinsic quantum yield of CO2 assimilation. The dra-
matic and irreversible decrease in qp and ФPSII at 2 and
24 h time period after bentazon treatment may indicate
a significant increase in the proportion of the closed
PS II reaction centers or in the proportion of the re-
duced state of QA[9,17]. Bentazon competitively binds to
D1 protein and reduces the binding sites for QB thus
causing blockage of the electron transfer from QA to
QB[18]. It could be suggested that the accumulation of
the reduced state of QA induce the loss of Pn in the
bentazon treated leaves of both mutants.
第 9期 WANG Yun-Sheng et al.: Difference of Bentazon Tolerance in Wild Rice and Lethal Mutants 1621


The increase in the fraction of QA in the reduced
state suggests an increase in the excitation pressure on
PSII under the steady-state of photosynthesis, resulting
in abundant generation of 2O
− (Fig. 3). The abundant
generation of 2O
− can cause further photodamage on
chloroplasts or cells if excessive excitation energy is
not safely dissipated[19]. Changes in MDA and Chl
contents indicate that the lethal effect of bentazon in
both mutants is due to accumulation of oxidative stress,
which in turn acts on chloroplasts and membrane sys-
tem, causing Chl degredation and membrane lipid
peroxidation[20].
3.2 Bentazon resistant mechanism in the wild
type
When the values of Pn and the chlorophyll fluo-
rescence were compared among the two wild type
varieties and their mutants, no significant difference
was observed within 2 h after bentazon treatment. As
the fluorescence characteristics during the steady-state
of photosynthesis are representative of the efficiency
of electron transfer in PS II, the results indicate that
the affinity of the target site (D1 protein in PS II) to
bentazon is roughly equal in the wild type varieties
and their mutants. However Pn and chlorophyll
fluorescence parameters increased progressively and
approached the control for the wild types 1 d after
treatment. Therefore, the inhibition was significantly
alleviated during the process and very slight
photodamage has been incurred in treated leaves of
both original varieties.
Changes of Bentazon content were investigated to
define the correlation between bentazon accumulation
and the recovery efficiency of photosynthetic capacity
after foliar application of the herbicide. This study
reveals that wild type varieties absorbed bentazon as
equally as the mutants during the first 2 h, which is
approved by similar content of bentazon in the leaf
tissues. After 8 h, bentazon was metabolized into other
non-toxic molecules and became almost undetectable
in the wild type plants, but substantial amount of the
herbicide was retained in the leaf tissues of both mu-
tants. Based on the correlation between catabolism of
bentazon and changes in photosynthetic parameters, it
is proposed that the reduction of bentazon is the essen-
tial reason for the recovery of photosynthetic capacity
in the two wild type varieties.
In a previous study, it was found that N8m was
more sensitive to bentazon than 8077s[8]. The current
research detected that the Chl content decreased faster
in N8m than in 8077s after exposure to bentazon (400
mg L−1). When the two wild type varieties were com-
pared, N8 was significantly slower and more delayed
in the recovering rate of photosynthetic capacity and
the metabolizing rate of bentazon than W6154s. No
significant differences in MDA content, Pn, chloro-
phyll fluorescence and bentazon content were detected
between the two mutants. Therefore it is indicated that
bentazon sensitivity of the mutants could be affected
by the physiological characteristics.
4 Conclusions
By analyzing parameters associated with photo-
synthetic efficiency and bentazon catabolism, and
correlating them with the genotypes, we have identi-
fied the possible physiological mechanism for benta-
zon tolerance/sensitivity in rice wild types and their
bsl mutants. It is proposed that the herbicide might
inhibit rice photophythesis and accumulation of oxida-
tive stress with the treatment of bentazon in both mu-
tants. And a rapid catabolism of bentazon in the wild
type is the essential mechanism for their tolerance.

Acknowledgement: The authors thank Dr. Hong-Jun
Zheng for kindly providing the manuscript amend-
ment.
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