全 文 ：儿茶素诱导的拟南芥根细胞膜脂变化*
郑国伟1, 陈摇 佳2, 李唯奇1**
(1 中国科学院昆明植物研究所中国西南野生生物种质资源库, 云南 昆明摇 650201;
2 云南瑞升烟草技术 (集团) 有限公司, 云南 昆明摇 650106)
摘要: 儿茶素是一种可以短时间内杀死植物细胞的植物毒素, 由于具有强的植物毒性, 儿茶素是开发除草
剂的理想化合物, 它可以诱导植物根系统的死亡。 为了研究植物根细胞膜脂对化学胁迫的响应规律, 我们
运用高通量的脂类组学方法检测了拟南芥根中膜脂分子的组成, 比较了儿茶素处理下拟南芥野生型 (WS)
及磷脂酶 D啄缺失突变体 (PLD啄鄄KO) 根中膜脂分子的组成情况、 膜脂含量、 双键指数及碳链长度值。 结
果发现, 儿茶素处理拟南芥根 90 min 后, 二半乳糖基二酰甘油 (DGDG)、 单半乳糖基二酰甘油 (MG鄄
DG)、 磷脂酰甘油 (PG)、 磷脂酰胆碱 (PC) 及磷脂酰肌醇 (PI) 的总含量在 WS 与 PLD啄鄄KO 植株根中
都显著下降, 磷脂酰乙醇胺 (PE) 和磷脂酰丝氨酸 (PS) 在 WS中下降, 在 PLD啄鄄KO中上升。 儿茶素处
理导致 PLD啄鄄KO植株的 PC / PE比值显著下降, WS植株 PS碳链长度显著增加。 上述结果说明儿茶素处理
后, 磷脂酶 D啄缺失突变体膜不稳定性增加, PLD啄鄄KO植株对儿茶素胁迫更加敏感。
关键词: 植物毒素; 脂类组学; 儿茶素; 拟南芥
中图分类号: Q 945摇 摇 摇 摇 摇 摇 摇 文献标识码: A摇 摇 摇 摇 摇 摇 摇 文章编号: 2095-0845(2012)04-383-08
Profiling of Membrane Lipids of Arabidopsis Roots
during Catechin Treatment
ZHENG Guo鄄Wei1, Chen Jia2, LI Wei鄄Qi1**
(1 The Germplasm Bank of Wild Species, Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650201, China;
2 Yunnan Reascend Tobacco Technology (Group) CO., LTD., Kunming 650106, China)
Abstract: Catechin is a kind of phytotoxin and can kill plant cells in an hour. It can be developed as herbicide to
kill weeds due to its strong phytotoxic activity. The main effect of this chemical is to trigger the death of the root sys鄄
tem. To understand the response of root cell membrane lipids to catechin stress, we used the lipidomics approach to
study the profiles of Arabidopsis root lipids molecules under catechin treatment. The changes of molecular species in
membrane lipids, content of membrane lipids, double bond index (DBI) and acyl chain carbon number of the fatty
acid were examined in Wild type (WS) and PLD啄 deficient mutant (PLD啄鄄KO) during catechin treatment. The re鄄
sults indicated that after 90 min treatment with catechin, the lipid contents of digalactosyldiacylglycerol (DGDG),
monogalactosyldiacylglycerol (MGDG), phosphatidylglycerol (PG), phosphatidylcholine (PC), and phosphatidyli鄄
nositol (PI) decreased both in WS and PLD啄鄄KO roots, lipid contents of phosphatidylethanolamine (PE) and phos鄄
phatidylinositol (PS) decreased in WS roots, but increase in PLD啄鄄KO roots, lipid contents of (phosphatidic acid)
PA increased at the begin of treatment and declined to the level of control in WS roots. The ratio of the two major
lipids in roots, PC and PE, declined significantly in PLD啄鄄KO plants, the acyl carbon number of PS in WS plants
植 物 分 类 与 资 源 学 报摇 2012, 34 (4): 383 ~ 390
Plant Diversity and Resources摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 DOI: 10. 3724 / SP. J. 1143. 2012. 12005
*
**
Foundation items: Fund of State Key Laboratory of Phytochemistry and Plant Resources in West China (0807B01211), West Light Founda鄄
tion of the Chinese Academy Sciences
Author for correspondenc; E鄄mail: weiqili@ mail. kib. ac. cn
Received date: 2012-01-12, Accepted date: 2012-02-15
作者简介: 郑国伟 (1982-) 男, 博士, 助理研究员, 主要从事植物逆境分子生理学研究。 E鄄mail: gwzheng@ mail. kib. ac. cn
increased. The results suggested that PLD啄鄄KO is more sensitive than WS during catechin treatment, and suppres鄄
sion of PLD啄 exacerbated membrane damage induced by catechin.
Key words: Phytotoxin; Lipidomics; Catechin; Arabidopsis
Abbreviation: electrospray ionization tandem mass spectrometry, ESI鄄MS / MS; phosphatidylcholine, PC; phos鄄
phatidylethanolamine, PE; phosphatidylglycerol, PG; phosphatidic acid, PA; phosphatidylinositol, PI; phosphati鄄
dylserine, PS; digalactosyldiacylglycerol, DGDG; monogalactosyldiacylglycerol, MGDG; PLD, phospholipase D
摇 Catechin is a common compound in plants which
has antioxidant activity (Lee et al., 2002; Lu et al.,
2011; Ma et al., 2003; Meyer et al., 1998; Ono et
al., 1998; Su et al., 2002; Vuong et al., 2011;
Yang et al., 2003 ). However, recent researches
were focused on the phytotoxic and antibacterial ac鄄
tivity of catechin that released from some plants to
help them compete with surrounding plants. For ex鄄
ample, Inderjit et al. ( 2008) found that catechin
can significantly inhibit root growth of Bambusa and
Koeleria seedlings; Bais et al. (2003) reported that
catechin can inhibit seed germination of 6 kinds of
weeds and crops; a study by D忆Abrosca et al. (2006)
displayed that ( -)鄄catechin can inhibit green alga
Selenastrum capricornutum growth; (+)鄄catechin from
seed coat of Sesbania virgata and velvetleaf can inhibit
root elongation of Arabidopsis, cress, radish and soy鄄
bean (Paszkowski and Kremer, 1988; Sim觛es et al.,
2008). Weir et al. (2006) reported that catechin treat鄄
ment could cause Arabidopsis lipid peroxidation and in鄄
hibited plant growth. Catechin may be developed as a
powerful herbicide to kill weeds due to its strong phyto鄄
toxic effects on roots. However, little is known about
precise mode of action of catechin and the response of
plant cell membrane lipids to catechin stress.
Membranes, particularly plasma and chloroplast
membranes, are sensitive to environmental stimuli,
the membrane lipids is crucial in plants response to
stresses. PA is an important second messenger, it
involves in plant response to various stresses (Tes鄄
terink and Munnik, 2005; Wang, 2004, 2005b;
Wang et al., 2006), and its level increases within
minutes under these stresses (Munnik, 2001). PC and
PE are related to the membrane stabilization (Welti
et al., 2002; Yeagle et al., 1976). We found that
through remodeling of membrane lipids plants re鄄
spond to frequent alterations between high and low
temperatures (Zheng et al., 2011). Phospholipase
D (PLD) hydrolyzes phospholipids to generate PA.
PLD啄 is one of the 12 PLDs in Arabidopsis, and it
involves in plants stress response and PLD啄 increa鄄
ses during stress (Wang, 2005a). Zhang et al.
( 2003 ) found that PLD啄鄄null cells displayed in鄄
creased sensitivity to H2O2 鄄induced cell death.
PLD啄鄄mediated hydrolysis of phospholipids plays a
positive role in the plant response to oxidative stress.
Li et al. (2004) found that PLD啄鄄KO plants exhibi鄄
ted less tolerance to freezing injuries whereas PLD啄鄄
OE plants exhibited more tolerance. PLD啄 and PA
signaling may involve in the response of plants to
drought and salinity (Hong et al., 2010). However,
under chemical stresses, like catechin, the change
of lipid and the effects of PLD啄 during this stress
have not previously been described.
Plant lipidomics based on ESI鄄MS / MS can tell
us the 11 classes of lipid changes under certain con鄄
ditions (Welti et al., 2002). The purpose of this
study was to use lipidomics and Arabidopsis PLD啄 mu鄄
tant to determine: (1) harmful effects of catechin to
Arabidopsis root membrane lipid composition and (2)
involvement of PLD啄 in root membrane lipid profiling.
1摇 Materials and methods
1. 1摇 Plant materials and chemicals
A PLD啄鄄knockout (PLD啄鄄KO) mutant isolated
from Arabidopsis ( Wassilewskija ecotype ( WS))
was from Dr. Xuemin Wang爷s Laboratory previous鄄
ly; the loss of PLD啄 was confirmed by the absence
of the transcript, protein, and activity of PLD啄
(Zhang et al., 2003). (依)鄄catechin and (+)鄄cate鄄
483摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 植 物 分 类 与 资 源 学 报摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 第 34 卷
chin were purchased from Sigma鄄Aldrich ( C1788,
C1251 respectively).
1. 2摇 Plant growth and treatments
Seeds of two Arabidopsis genotypes were steri鄄
lized with ethanol (75% ) for 2 min and sodium hy鄄
pochlorite (5% ) for 2 min, and then rinsed three
times with sterile distilled water. Surface鄄sterilized
seeds were cold stratified for 2 days at 4 益, and
then sowed on 1 / 2 MS medium (Murashige and Sk鄄
oog, 1962) for hydroponic culture as described by
Tocquin et al. (2003). The conditions of the growth
chamber were 23 / 18 益, a 12 / 12 h light / dark cy鄄
cle, and 120 滋mol·m-2 s-1 photosynthetic photon
flux density.
We used 40 day old hydroponic seedlings to test
the effects of catechin on Arabidopsis roots. Catechin
was added to the hydroponic solutions to the concen鄄
tration of 150 mg·L-1, and hold for 0, 10, 30, 90
min. Roots were harvested for lipids analysis at each
time.
1. 3摇 Lipid extraction and ESI鄄MS / MS analysis
The process of lipid extraction, ESI鄄MS / MS a鄄
nalysis, and quantification was performed as de鄄
scribed previously with minor modifications (Devaiah
et al., 2006; Welti et al., 2002). Briefly, the roots
were cut at sampling time and, to inhibit lipolytic ac鄄
tivities, were transferred immediately into 3 mL of
isopropanol with 0. 01% butylated hydroxytoluene at
75 益. The roots were extracted with chloroform /
methanol (2 颐 1) two additional times with 3 day of
agitation each time. The remaining plant roots were
heated overnight at 105 益 and weighed. The weights
of these extracted and dried tissues were described as
“dry weight冶 of the plants. Lipid samples were ana鄄
lyzed on a triple quadrupole MS / MS equipped for
ESI. Data processing was performed as previously de鄄
scribed (Devaiah et al., 2006; Welti et al., 2002).
1. 4摇 Data analysis
Statistical analysis was performed using Origin
7.0 (OriginLab Corporation, Northampton, MA, USA).
Double bond index (DBI) were calculated with the
formula: DBI=[移(N1伊mol% lipid)] / 100, where
N1 was the total number of double bonds in the two
fatty acid chains of each glycerolipid molecule
(Zheng et al., 2011). Average carbon number (C)
of acyl chains of lipid classes were calculated by the
formula: C = [移(N2 伊mol% lipid)] / 100, where
N2 was the total acyl carbons in each lipid molecule.
2摇 Results and discussion
2. 1 摇 Root lipids profiling of Arabidopsis during
catechin treatment
To test the effects of catechin on Arabidopsis
root membrane lipids, we used a lipidomics ap鄄
proach to profile changes in molecular species of
membrane glycerolipids in Arabidopsis during cate鄄
chin treatment. We identified and quantified about
120 glycerolipids molecular species including 11
species of lipids in Arabidopsis roots during catechin
treatment (Figs. 1 -3). The lipid profile of root is
different from that of leaf, where PC and PE are the
main lipids of root. The major lipid molecules of
MGDG were 34 颐 6MGDG and 36 颐 6MGDG, in roots
the content of 36 颐 6MGDG was a bit more than 34 颐
6MGDG, but in leaves the content of 34 颐 6MGDG
was much more than 36 颐 6MGDG (Li et al., 2008)
(our unpublished data).
To assess the role of PLD啄 in Arabidopsis re鄄
sponse to catechin, we employed the PLD啄 knockout
mutant Arabidopsis and compared its lipid profiles to
that of wild type plants during catechin treatment. In
wild type (WS) Arabidopsis plants, the content of
total lipids decreased with the time of catechin treat鄄
ment. After 90 min treatment with catechin, the
content of DGDG, MGDG, PG, PC, and PI de鄄
creased both in WS and PLD啄鄄KO roots, PE and PS
decreased in WS but increased in PLD啄鄄KO plants
(Fig. 1). PA is an important secondary messenger
in response of plants to abiotic stresses. It can in鄄
crease in minutes after stimuli and then decreases to
the levels of control (Wang et al., 2006). In out
experiment, PA rose at the begin of catechin treat鄄
ment and declined to the level of the control after 90
min treatment in wild plants. While in PLD啄鄄KO
5834 期摇 摇 摇 ZHENG Guo鄄Wei et al. : Profiling of Membrane Lipids of Arabidopsis Roots during Catechin Treatment摇 摇 摇 摇
plants, PA decreased during the catechin treatments
(Fig. 1). PLD啄 can hydrolyze phospholipids to PA
(Zhang et al., 2003), and suppression of PLD d
leads no PA increase during catechin treatment. We
thought that PLD啄 derived PA was very important for
Arabidopsis response to catechin stress.
The total content of LysoPLs was a bit different
in the two plants. Detailed analysis of the lipid pro鄄
files indicated that changes in the phospholipids and
galactolipids in the two plants were similar during cate鄄
chin treatment (Fig. 2, 3), except for minor differ鄄
ences in PE species 34 颐 2, 34 颐 3, and 36 颐 5 ( total
acyl chains: double bonds), in PA species 34 颐 2,
34 颐 3, and 36 颐 5 (Fig. 2), and in lysophospholipids
and lysoPC species 18 颐 2 and 18 颐 3 (Fig. 3).
2. 2摇 PC and PE ratio reduced in PLD啄鄄KO under
catechin stress
PC and PE are related to the stabilization of
membrane. Unsaturated PEs have strong propensity
to form hexagonal phases (Cullis and Hope, 1978)
which might lead to the formation of a nonbilayer lip鄄
id phase and disturb the membrane integrity and cell
function, whereas PC is a bilayer鄄stabilizing lipid
(Welti et al., 2002). The molar ratio of PC / PE,
which implies the membrane stabilization, tends to
drop in plants under cold and hydration stress
(Hazei and Williams, 1990; Welti et al., 2002).
To assess the membrane stabilization of Arabidopsis
during catechin treatment, we calculated PC and PE
ratio of this process. We found that both PC and PE
decreased during catechin treatment in WS, whereas
the content of PC decreased and PE increased during
catechin treatment in PLD啄鄄KO plants ( Fig. 1 ).
The ratio of PC / PE in WS did not change too much
during catechin treatment, however, the PC / PE ra鄄
tio in PLD啄鄄KO plants dropped from 1. 18 to 0. 92
after 90 min of catechin treatment (Table 1). The
results suggested that PLD啄 play an important role in
maintaining membrane stabilization in catechin in鄄
duced Arabidopsis membrane disturbance.
Fig. 1摇 Changes of each head group class and total lipids in WS and PLD啄鄄KO plants during catechin treatment. Values are means 依 S. D. (n=5)
683摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 植 物 分 类 与 资 源 学 报摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 第 34 卷
Fig. 2摇 Changes of the molecular species of membrane lipids in WS and PLD啄鄄KO plants during
different time of catechin treatment. Values are means 依S. D. (n=5)
Fig. 3摇 Changes of the molecular species of lysophospholipids in WS and PLD啄鄄KO plants during
different time of catechin treatment. Values are means 依S. D. (n=5)
7834 期摇 摇 摇 ZHENG Guo鄄Wei et al. : Profiling of Membrane Lipids of Arabidopsis Roots during Catechin Treatment摇 摇 摇 摇
Table 1摇 Lipid ratio of WS and PLD啄鄄KO plants during catechin treatment. An asterisk indicates that the value
is different from control (P<0. 05) . Values are means ±SD (n=5)
Lipids Plants
Lipid ratio
0 min 10 min 30 min 90 min
PC / PE
WS 1. 00依0. 07 1. 04依0. 06 1. 02依0. 09 1. 06依0. 13
PLD啄鄄KO 1. 18依0. 12 0. 99依0. 08* 0. 95依0. 09* 0. 92依0. 09*
(PC+PE) / Total
WS 0. 75依0. 01 0. 75依0. 02 0. 76依0. 01 0. 76依0. 02
PLD啄鄄KO 0. 74依0. 02 0. 75依0. 02 0. 77依0. 01* 0. 76依0. 02
Glycolipids / Total
WS 0. 11依0. 01 0. 11依0. 01 0. 10依0. 01 0. 11依0. 01
PLD啄鄄KO 0. 12依0. 01 0. 11依0. 01 0. 11依0. 01 0. 10依0. 01*
2. 3 摇 The influence of catechin to lipid double
bond index and acyl chain lengths
Maintaining the integrity and optimal fluidity of
the membranes is very important to organisms.
Changing in the degree of lipid unsaturation and the
number lipid acyl chain carbon could influence the
integrity and fluidity of membranes. We got the
mol% content of each lipid molecule species based
on the data of nmol / mg dry weight, and calculated
the DBI and Carbon number of acyl chains of each
lipid species.
We found that unlike the DBI of leaves, total
lipid DBI of roots was about 3. 5, but in leaves the
DBI was about 4. 1 (Zheng et al., 2011) ( Table
2), because the main lipid constituent in root are
PC and PE which have less double bond than MGDG
and DGDG which was the main lipid in leaves. The
degree of unsaturation of root membrane lipids is less
than leaves, the DBI of PG in roots was about 1. 2
less than in roots, the DBI of MGDG, DGDG, and
PS was a bit less than that in leaves ( Li et al.,
2011) (Table 2). The DBI decreased as the catechin
Table 2摇 Double bond index (DBI) of membrane during catechin treatment in WS and PLD啄鄄KO plants. An asterisk indicates
that the value is different from control (P<0. 05) . Values are means ±SD (n=5)
Lipid species Plants
Double bond index
Control 10 min 30 min 90 min
DGDG
WS 5. 14依0. 04 5. 12依0. 03 5. 08依0. 05 5. 06依0. 02*
PLD啄鄄KO 5. 13依0. 04 5. 15依0. 04 5. 08依0. 05 5. 08依0. 06
MGDG
WS 5. 83依0. 02 5. 81依0. 07 5. 78依0. 03* 5. 77依0. 05*
PLD啄鄄KO 5. 79依0. 02 5. 80依0. 01 5. 77依0. 02 5. 77依0. 02
PG
WS 2. 02依0. 23 1. 96依0. 31 1. 92依0. 26 2. 04依0. 17
PLD啄鄄KO 2. 03依0. 20 1. 82依0. 32 1. 66依0. 37 1. 82依0. 18
PC
WS 3. 76依0. 02 3. 75依0. 11 3. 71依0. 04* 3. 71依0. 07
PLD啄鄄KO 3. 74依0. 01 3. 75依0. 02 3. 68依0. 05* 3. 65依0. 02*
PE
WS 3. 14依0. 01 3. 12依0. 05 3. 05依0. 06* 3. 04依0. 05*
PLD啄鄄KO 3. 11依0. 02 3. 11依0. 01 3. 04依0. 02* 3. 03依0. 02*
PI
WS 2. 69依0. 03 2. 71依0. 04 2. 66依0. 06 2. 67依0. 03
PLD啄鄄KO 2. 65依0. 01 2. 69依0. 03* 2. 64依0. 02 2. 63依0. 01*
PS
WS 2. 79依0. 04 2. 77依0. 10 2. 67依0. 11 2. 69依0. 10
PLD啄鄄KO 2. 71依0. 06 2. 69依0. 08 2. 68依0. 04 2. 67依0. 04
PA
WS 3. 02依0. 16 3. 06依0. 17 2. 99依0. 13 2. 99依0. 16
PLD啄鄄KO 2. 97依0. 07 3. 01依0. 22 2. 75依0. 25 2. 98依0. 09
Total Lipids
WS 3. 58依0. 01 3. 56依0. 08 3. 49依0. 07 3. 51依0. 05
PLD啄鄄KO 3. 58依0. 03 3. 55依0. 02 3. 49依0. 03* 3. 44依0. 04*
883摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 植 物 分 类 与 资 源 学 报摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 第 34 卷
treatment, and there was no difference in DBI between
WS and PLD啄鄄KO plants except for PG, which DBI in
PLD啄鄄KO plants was less than in WS when treated
with catechin ( Table 2). These results suggested
that catechin may induce membrane lipids oxidation.
In the eight classes of lipids that calculated acyl
chain carbons, the acyl chain carbons of MGDG of
roots was longer than that in leaves (Table 3, our
unpublished data). The longest acyl chain was PS
which was about 39 carbons (Table 3). Under cate鄄
chin treatment, the acyl chain carbons of the lipids
change little except that of PS which in WS in鄄
creased 0. 4 and in PLD啄鄄KO decreased 0. 66 after
90 min of catechin treatment. Catechin could induce
change of length of lipid acyl chain, the most sensi鄄
tive lipid is PS.
Table 3摇 The acyl chain carbon of membrane lipids during catechin treatment in WS and PLD啄鄄KO plants. An asterisk indicates
that the value is different from control (P<0. 05) . Values are means ±SD (n=5)
Lipid species Plants
Acyl chain carbon (C)
Control 10 min 30 min 90 min
DGDG
WS 35. 54依0. 02 35. 53依0. 02 35. 56依0. 02 35. 54依0. 03
PLD啄鄄KO 35. 59依0. 02 35. 59依0. 02 35. 53依0. 03* 35. 53依0. 03*
MGDG
WS 35. 00依0. 03 35. 03依0. 09 35. 16依0. 11* 35. 15依0. 11*
PLD啄鄄KO 35. 10依0. 01 35. 08依0. 02 35. 18依0. 04* 35. 17依0. 04*
PG
WS 33. 47依0. 20 33. 46依0. 22 33. 40依0. 17 33. 47依0. 14
PLD啄鄄KO 33. 42依0. 09 33. 31依0. 23 33. 31依0. 25 33. 37依0. 19
PC
WS 35. 20依0. 02 35. 19依0. 05 35. 17依0. 01* 35. 16依0. 02*
PLD啄鄄KO 35. 20依0. 01 35. 18依0. 01* 35. 17依0. 03 35. 14依0. 02*
PE
WS 35. 22依0. 02 35. 21依0. 04 35. 16依0. 05* 35. 16依0. 03*
PLD啄鄄KO 35. 22依0. 02 35. 20依0. 02 35. 16依0. 01* 35. 15依0. 01*
PI
WS 34. 29依0. 03 34. 29依0. 02 34. 26依0. 01 34. 27依0. 02
PLD啄鄄KO 34. 29依0. 03 34. 26依0. 03 34. 28依0. 02 34. 24依0. 03*
PS
WS 38. 76依0. 12 38. 75依0. 18 38. 88依0. 45 39. 16依0. 38
PLD啄鄄KO 39. 48依0. 46 38. 89依0. 13* 38. 93依0. 55 38. 82依0. 31*
PA
WS 34. 45依0. 17 34. 50依0. 10 34. 45依0. 07 34. 53依0. 11
PLD啄鄄KO 34. 36依0. 15 34. 43依0. 13 34. 33依0. 20 34. 47依0. 13
Total Lipids
WS 35. 20依0. 02 35. 19依0. 04 35. 19依0. 05 35. 18依0. 03
PLD啄鄄KO 35. 19依0. 02 35. 19依0. 03 35. 17依0. 04 35. 16依0. 02*
摇 In conclusion, the results of this study suggested
that PLD啄 deficient plant was more sensitive to cate鄄
chin stress than wild plant. Catechin could induce
the root lipids change and lead to the membrane dis鄄
turbance. However, many questions, such as the
mechanisms of different lipid profiles in roots and
leaves; the role of PLD啄 play in Arabidopsis resist to
catechin stress; and the reason of changes of acyl
chain carbon length to catechin stress, remained un鄄
clear and would be further studied in the future.
Acknowledgement: We thank Dr. Hongyin Chen, Dr. Bo
Tian and Dr. Yanxia Jia for their critical reading of the paper.
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