全 文 ：广 西 植 物 Guihaia 31(4)：520—525 2011年 7月
DOI：10．3969／j．issn．1000—3142．2011．04．019
薇甘菊可溶性蛋白和抗氧化酶活性对
田野菟丝子不同寄生密度的响应
刘梦佼1，2*，洪 岚 ，2*，沈 浩1 ，韦 霄3，叶万辉 ，曹洪麟
(1．中国科学院 华南植物园 ，广州 510650；2．中国科学院 研究生院，北京 100049；
3． 薯 广西植物研究所，广西桂林54106)
摘 要：为探求利用寄生植物 田野菟丝子对入侵杂草薇甘菊进行生物控制的有效措施 ，研究了薇甘菊对 0、1、
2、4和 8棵 田野菟丝子幼苗寄生在可溶性蛋白和一些抗氧化酶活性方面的响应。寄生后 30 d，1棵田野菟丝
子／株薇甘菊(以下简称棵／株)以上的寄生密度导致薇甘菊可溶性蛋白含量显著降低。和对照相 比，在寄生密
度为 1棵／株时，超氧物歧化酶(SOD)和过氧化物酶(POD)活性显著增强 ；但 随着寄生密度的加大而下降，而
且在寄生密度为 4棵／株时，SOD和 POD分别等于和小于对照 ，在 8棵／株时均显著小于对照。在各寄生密度
下 ，寄主的过氧化氢酶(CAT)活性均小于对照，而 SOD／CAT，SOD／POD和 SOD／(CAT+POD)比率均大于
对照。这些结果表明，田野菟丝子的寄生对薇甘菊可溶性蛋 白和抗氧化酶活性的影响依赖于寄生密度，在野
外利用田野菟丝子控制薇甘菊的最理想寄生密度是 4棵／株，从而可为野外利用 田野菟丝子控制薇甘菊的技
术体系提供参考。
关键词：生物防治；入侵杂草；寄生植物；抗氧化酶
CLC Number：Q945．79 Document Code：A Article ID：1000-3142(2011)04-0520-06
Responses 0f M ikania micrantha to parasitization
0f Cuscuta cam j tris‘ tot d soluble proteinC pes t in alSOUDI o t
O0ntent and activities 0f antioxidant enzymes
LIU Meng-Jiao1，2 ，HONG LanI，2 ，SHEN Hao1 ，
W EI Xiao3，YE Wan-HuiI，CAO Hong-Lin1
(1．South China Botanical Garden，Chinese Academy of Sciences，Guangzhou 510650，China；2．Graduate School of
the Chinese Academy of Sciences，Beijing 100049，China；3．Guangxi Institute of Botany，Guangxi Zhuang
Autonomous Region and the Chinese Academy of Sciences，Guilin 541006，China)
Abstract：To develop efficient biocontrol techniques for the invasive weed Mikania micramha，the use of the obligate
parasitic plant，Cuscuta campestris Yuncker as a biological control was investigated．In this experiment，whether the
impacts of the parasite on host soluble protein content and activities of some antioxidant enzymes were affected by the
density of the parasite was tested．The responses of M micrantha to parasitic densities of 0，1，2，4 and 8 individual
seedlings of C．campestris per host plant were examined．On the 30th day of parasitization。infection with more than 1
Received date：2010—11—08 Accepted date：2011 03 l9
Foundation items：Supported by National Basic Research Program of Ckm“973”Program(2009CBl19200)；the Knowledge Innovation Program of
the Chinese Academy of Sciences；the National Natural Science Foundation of China(30470345)；Guangdong Natural Scienee Foundation(05200701)
Bio~aphy：UU Meng-Jiao(1985一)，female，born in Anhui Province，master student，majoring in conservation ecology，(E-mail)shazi558(~163．com．
The tWO authors contributed equally to this work．
Author for correspondence，E-maiI：shenhao@sebg．ae．cn
4期 刘梦佼等 ：薇甘菊可溶性蛋白和抗氧化酶活性对田野菟丝子不 同寄生密度的响应 521
C．campestris seedling significantly lowered soluble protein content．Compared with control，activities of superoxide
dismutase(SOD)and peroxidase(POD)significantly increased at parasitic density of 1 parasite per host plant；but as
parasitic density increased，both SOD and POD activities decreased．M oreover，SOD activity was not changed but POD
activitv was significantly decreased at parasitic density of 4 parasites per host，and both SOD and POD activities were
significantly decreased at parasitic density of 8 parasites per host，compared with contro1．The infected plants had sig—
nifjcantlv lower catalase(CAT)activity but higher SOD／CAT，SOD／POD and S0D／(CAT+PoD)ratios than the
control at a11 parasitic densities．The results indicated that the effects of C．campestris infection on t^ micrantha were
densitv dependent，which provided a basis for refining this strategy for biological control of iV／．micrantha．The opti—
ma1 cost-efective number of parasites to control Mr．micrantha was 4 per host plant in the field．
Key words：biological control；invasive weed；parasitic plant；antioxidant enzymes
1 Introducti0n
Biological invasion is one of the major threats to
native biodiversity，economic development and biologi—
ca1 safety (Mack et a1．，2000；Pimentel et a1．，2005)
and it is an important element of global change(Vi—
tousek et口Z．，1 997)．The problems of invasive species
and their eontrol have been one of the most pressing
applied issues in ecology today (Hastings et a1．，
2006)．Therefore，many researchers have tried to de—
velop effective strategies to control invasive species in
affected areas(Zavaleta et a1．，2001；Taylor & Has—
tings，2004；Buhle et a1．，2005；Culliney，2005；Hulme，
2006)．These strategies include cultura1，mechanical，
chemical，and biological contro1．Among these，biologi—
cal control is permanent，energy-efficient，non-poilu—
ring，inexpensive and ecologicaly safe relative to other
methods(Culliney，2005)． Thus，it has been widely
recognized as one of the most promising methods of
controiling invasive weeds(Culliney，2005)，especially
when native species are used to control exotic invasive
weeds(Shen et a1．，2005)．
M 是n 口micrantha is a fast-growing perennial
climbing vine of the family Asteraceae，and is native to
Centra1 and South America (Holm et a1．，1977)．
However，in its palaeotropic exotic ranges such as
moist tropical forest zones of the Pacific region，India，
and Asia，particularly South-east Asia，it has become a
horrific invader and a notorious weed，severely dama—
ging forestry and plantation crops (Zhang et a1．，
2004)．It climbs up on other plants to reach the cano—
PY for better sunlight，and smothers other plants，thus
it is named“plant killer”in China．It sprawls out rap—
idly in spring and summer，showing a vigorous，fast and
rampant growth habit，that is why it is named“mile-a—
minute weed” (W aterhouse，1994)． Moreover，the
weed can reproduce vigorously by both vegetative and
sexual reproduction (Zhang et a1．，2004)．It has been
listed as one of the i00 worst invasive alien species in
the world(Lowe eta1．，2001)，and is ODe of the top 10
worst weeds in the world(Holin et a1．，1977)．M ．mi—
c，—Ⅱ tha entered So uth China after 1910．Since the
1 980s，it has spread and invaded widely，causing severe
damage to many ecosystems，and leading to significant
economic lOSS in South China including Hong Kong，
Guangdong，and Hainan (Feng et a1．，2002；Zhang et
口Z．，2004)．Thus，it has been listed as one of the 16
most invasive species by the State Environmental Pro—
tection Administration(Zhang et a1．，2004)．
To biologically control M．micrantha，we have
tried to use the obligate parasite Cuscuta campestris
Yuncker in So uth China(Shen et a1．，2005，2007)．W e
found that C．campestris infection significantly reduced
total biomass and photosynthesis，changed the biomass
allocation patterns and completely inhibited flowering
of M ．micrantha plants．At the community level，field
studies(Lian et a1．，2006)showed that C．campestris
significantly reduced biomass and cover of M．micran—
tha，but had only minor effects on the growth of other
plants．Thus，C campestris might reduce the harmful
effects of M ．micrantha，and hence increase species di—
versitv and help reestablishment of native species．
These results indicated that the use of C．campestris
could be a potentially effective and relatively safe way
to control M micrantha．
522 广 西 植 物 31卷
In this study，the effects of parasitic density of C．
campestris on M ．micrantha were investigated in the
field in South China． In order to test whether the
effects of the parasite on host soluble protein content
and activities of some antioxidant enzymes are affected
by the density of the parasite，we examined the respon—
ses of M micrantha to parasitic densities of 0，1，2，4
and 8 individual seedlings of C campestris per host
plant．The work will provide useful and practical in—
formation for the deployment of C campestris for the
biological control of the invasive species M micrantha．
2 Materials and methods
2．1 Experimental materials and design
The study was conducted during the May-Decem—
ber 2004 growing season at the field station of So uth
China Botanical Garden(23。10 N，113。21 E，elevation
40 m a．s．1．)in Guangzhou，Guangdong Province，Chi—
na．The region is characterized by a typical south sub—
tropical monsoon climate．On 3 1 May 2004，whole M ．
micrantha plants were collected from a M micrantha
population in the suburb of Dongguan，Guangdong
Province，China．Similar-sized two-node segments were
selected from the middle of the stems to minimize the
influence of phenotypic maternal effects．The segments
were planted in 89 L pots filled with a mixture of pool
mud and paddy field clay(1：2，v／v)．The mixed soil
had pH 6．5± 0．1，organic mater content 3．3±
O．O9 ，total nitrogen 0．182±0．08 ，ammonia nitro—
gen 103．574-5
． 86 mg kg-1，available P 23．98± 1．36
mg· kg ，and available K 132．83± 6．32 mg kg。
(mean±SE)．In each pot，three segments were planted
with the lower node buried below and the upper one a—
bout 5 cm above the soil surfaca．The upper nodes be—
gan to sprout 3 d later．W hen the plants were about
350 cm in height(23 August)，they were thinned to
one per pot，and 100 individuals，similar in height and
stem diameter，were selected and placed at random in
an open field uniform in environmental conditions．Of
these，8O were randomly chosen as host plants(infec—
ted group)，leaving the remaining 20 uninfected (con—
trol group)．T0 prevent M ．micrantha from climbing
from one pot to another．pots were separated at least 1
m from each other and a bamboo cane about 5 m long
was placed vertically in each pot for M．micrantha to
climb on．
On 12 August，C campestris seeds were sown in
pots filled with sands at a depth of about 1锄 and they
completed germination on 20 August．On 26 August，
when M micrantha plants were about 370 crn in height
and the mean number of leaves was 450，and C
campestris seedlings were about 5 cm in height，C
campestris seedlings with wet sands were placed on the
soil surface of each M micrantha pot in the infected
group．Density treatments(1，2，4 or 8 C campestris
seedlings per host plant)were randomly assigned to
M micrantha plants in the infected group．20 host
plants per treatment density．By 29 August，aU the M
micrantha plants in the infected group had become in—
fected wi th C campestris stems．The experiment ended
on 27 De cember，120 d after parasitization (DAP)or
210 d after planting，when the uninfected M micrantha
plants had started to wither．During the experiment，
the pots were weeded when necessary and watered
twice daily wi th tap water at 600 h and 1800 h，except
on rainy days．No fertilizer was added throughout the
experimental period．
2．2 Soluble proteins and enzyme assays
On 30 DAP，the 8th fully expanded sun leaf from
four randomly selected M micrantha plants per infec—
tion density was collected for determination of soluble
protein content and enzyme assays，four samples of
fresh leaves per treatment． Approximately 0．5 g of
fresh leaves (midvein excluded) per sample was ho—
mogenized in 5 mL of ice-cold 50 mM potassium phos—
phate buffer(pH7．8)in an ice-cold mortar．The ho—
mogenate was centrifuged at 16 000 g for 15 min at 4
℃ (CR22G Ultracentrifuge，Hitachi，Japan)，and the
supernatant was collected and stored at 4℃ for soluble
protein determination and enzyme assays．Tota1 solu-
ble protein content in the samples was determined by
the protein dye-binding method of(Bradford，1976)，u—
sing bovine serum albumin as standard． Superoxide
dismutase(SOD)activity was determined by measur-
ing percent inhibition of nitroblue tetrazolium (NBT)
4期 刘梦佼等：薇甘菊可溶性蛋白和抗氧化酶活性对田野菟丝子不同寄生密度的响应 523
reduction by SOD (Giannopolitis& Ries，1977)．Ac—
tivity was expressed in units where 50 inhibition is
equivalent to one unit of SoD activity． Peroxidase
(POD)activity was determined spectrophotometrically
as described by Chance & Maehly (1955)． Catalase
(CAT)activity was assayed spectrophotometricaly by
following the decrease in absorption at 240 nm due to
the disappearance of hydrogen peroxide (H2 O2)
(Chance Maehly，1955)．
2．3 Statistical analysis
All tests were carried out at a一 0．05 level using
SPSS (version 11．5，SPSS Inc．，Chicago，IL，USA)．
One-way analysis of variance(ANOVA)was used to
determine statistical significance for the effects of infec—
tion density on plants for the variables of total soluble
protein content，and enzyme activities．M eans for sig一
专董
§
2．5
_c ·。
。 _
．1 1，5
薹
箸 0．5
— 0．0
C0 C1 C2 04 C8
CO C1 C2 C4 C8
nificant ANOVA effects were compared using Tukey
post hoc comparisons．
3 Results
3．1 Soluble protein content
At 30 DAP，inoculation with 1 parasite had no
significant effect on soluble protein content of M mi—
crantha plants~however，infection with more than 1
parasite significantly reduced soluble protein content
(Fig．1a)．Soluble protein contents at 2 and 4 parasites
per host plant were not significantly different，but sig～
nificantly higher than that at 8 parasites per host．The
soluble protein content in the infected M I micrantha
plants with 8 parasites per host was Qnly about half
that of uninfected contro1．
30
20
10
0
40O
C
E 300
200
箸 1O0
0
CO 01 02 e_A C8
nfection dens ty
CO Ct C2 C4 C8
Fig．1 M eans(± SE， 一 4)of soluble protein content(a)，superoxide dismutase(SOD)
activity(b)，peroxidase(POD)activity(c)and catalase(CAT)activity(d)of Mikania
micrantha 30 days after parasitization by different densities of Cuscuta campestris
Bars not sharing a common superscript letter are significantly different(P，
1j)一 29．447，P< O．001；
b，F“，15)一28．095，P< 0。001；c，F(4，15)一21．815，PC8：infection density at 0，1，2，4 and 8 C campestris per M_micrantha plant．
3．2 Activities ofSOD，POD and CAT
Cuscuta campestris had significant effects OD ac—
tivities of SOD．POD and CAT of M ．micrantha plants
(Fig．1：Dd)．SOD activities were significantly higher
at parasitic density of 1 or 2 parasites per host than
that of the contro1．similar between 4 and control。and
significantly lower at 8 than that of the contro1．For
infected treatments，SOD activities decreased signifi一
一ul。 J口．暑lu n) 一>一 o 凸0∞
4 2 O 8 6 4 2 0
一^ 。
君山一
c 一 oL
524 广 西 植 物 31卷
cantly with increasing parasitic density．Compared with
control，P0D activity was not changed at parasitic den—
sity of 4 parasites per host，but significantly increased
at 1 and decreased at 4 or 8 parasites per host plant．As
parasitic density increased，POD activity decreased．
The infected plants had significantly lower CAT activi—
ty than the contro1 at all parasitic densities．CAT ac—
tivities at infection density of 4 parasites per host were
significantly lower than that at 1 but h~gher than those
at 2 or 8‘parasites per host．Co mpared with the con—
trol，infection with C campestris led to significant in—
creases in the ratios of SOD／CAT，SoD／POD，and
SoD／(CAr+POD)activities in M．micrantha plants
(Table 1)．
Table 1 Ratios，of superomde dismutase(SOD)activity
． to eatalase(CAT)activity，SO D activity to peroxidase
(POD)activity，and SOD activity to the sum of CAT
activity and POD activty in M micrantha 30 days after
parasitization by different densities of C．campestris
No．followed by the~trlle superscript in each row are not significantly
different，according to Tukey post hoc comparisons(P< O．05)．Refer to
Fig．1 for definitions of CO，C1，C2，CA and C8．
4 Discussion
4．1 Response of M．micrantha to infection density ofC
campestris in soluble protein content
It has been shown that Cuscuta can form a strong
sink to redirect the flow of host resources to itself，and
these resources include water，nitrogen，mineral nutri—
ents and 99 of the carbon that it uses (Jeschke et
a1．，1994a，b；Jeschke 8L Hilpert，1997)． Parasitic
plants generally have high rates of transpiration(Stew—
art Press．1990)which may predispose hosts to wa—
ter stress and thus stomatal closure (Frost et a1．，
1997)．There have been reports that parasite infection
results in water stress in host plants(Taylor et a1．，
1996；Frost eta1．，1997)．Soluble proteins play an im—
portant role in osmotic adj ustment；and high soluble
protein content can sustain low osmotic potentiaI to re—
duce damage caused by water stress (Yu & Tang，
1999)．In this study，infection with more than 1 para-
site per M _micrantha plant significantly reduced solu-
ble protein content of the host．This reduction in solu-
ble protein content may be the reason that C campes—
tris infection reduces gs and hence decreases P托 M。
rnicrantha host plants(Shen et a1．，2007)．
On the other hand，in Ca plants，Rubisco generally
accounts for 30—60 of the soluble proteins (Ellis，
1979)．The rate of photosynthesis and biomass accu—
mulation depend largely on the quantity and activity of
Rubisco (Lorimer，1981)．Therefore，in this study，the
reduced soluble protein content due to infection mi【ght
have resulted in reduced Rubisco content in the infec—
ted rnicrantha。and such a reduced Rubisco content
would have contributed to the reduced photosynthesis
of the host．
4．2 Response of M ．micrantha toinfectiondensity ofC
campestris in activities of S0D，PoD and C舡
In the present study，there was a complex rela—
tionship among activities of S0D，POD，CAT and infec-
tion densities．At low infection densities(1 or 2 para—
sites per host plant)，SOD and POD activities increased
or maintained，indicating the protective enzyme system
was relatively well balanced．However，at relative high
infection densities(4 or 8 parasites per host)，such a
balance was broken．These indicated that it’s possible
C campestris infection resulted in high levels of reac—
tive oxygen species(ROS)，and at low infection densi-
ty，the infected M micrantha plants in response in-
creased the activities of SOD to minimize the dam—
age from ROS，which promoted POD to get rid of
H2 02 and hydroxy1．However，when the infection
densities became too high，the infected plants were
unable to do so．
Furthermore，ratios of SOD／CAT，SOD／POD，and
SOD／(CAT+POD)were significantly higher in the
infected at al infection densities than in the contro1．
These ratios might be suggested as markers of H2 02
content(Shah et a1．，2001)．Together，these might re—
sult in an imbalance of antioxidant enzyme systems in
the infected M．micrantha plants，and such an imbal—
ance might lead to excessive H2 02．It had been shown
4期 刘梦佼等：薇甘菊可溶性蛋白和抗氧化酶活性对田野菟丝子不同寄生密度的响应 525
that if only SOD activity was increased，while the activ—
ities of other antioxidant enzymes sueh as POD and
CAT were not enhanced enough，plants can not prevent
damage caused by oxygen free radicals(Pitcher et a1．，
1 9 9 1)，or they might even suffer more serious damage
brought by hydroxyl(Gossett et a1．，1994)．Further—
more，the increased 1evel of H2 02 caused by UV-B ra—
diation increased the degradation of Rubisco by activa—
tion of proteolytic systems(He et a1．，2004)．There—
fore，in the present study，the C campestris infection
might have resulted in high level of ROS production
and imbalance of the antio~dant enzymes． These
would lead to Rubisco degradation and cell damage and
thus reduced net photosynthesis of the infected M ．mi—
crantha plants(Shen et a1．，unpublished data)．
In conclusion，the results indicated that different
levels of C campestris infection had significantly differ—
ent effects on the performance of M．micrantha plants．
Host soluble protein content was significantly de—
creased when infected with more than 1 C．campestris
seedling．For the infected treatments，as parasitic den—
sity increased，soluble protein content，activities of SOD
and POD decreased． The protective enzyme system
was relatively well balanced at low infection densities
(1 or 2 parasites per host plant)but unbalanced at rel—
ative high infection densities (4 or 8 parasites per
host)．Thus，the minimum number of C．campestris for
optimum control of M ．micrantha in the field was 4 per
host plant．
Acknowledgements W e are grateful to Mr．Chris
Parker for his research guidance．
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4期 王辉等：水稻花药愈伤组织诱导的多因子正交试验研究 549
伤组织形成 ，但效果相对较差，这可能是多种激素混
合形成“复合启动 因子”，对花药愈伤组织 的形成有
促进作用 。
3．4低温预处理时间对愈伤组织形态结构的影响
陈红等(2007)研究发 现 ，低温预处理 8 d左右
的材料诱导的愈伤组织 (结构紧密且略泛 白色)好于
低温预处理 10 d以上的材料花药培养 出的愈伤组
织(松散 ，成分散小米状)。本试验低温预处理 7～
10 d，结果两种材料试验所得 的愈伤组织大都松散
成小米粒状 ，转接时较为困难 。其原因是 否与处理
时间过长有关有待进一步研究 。
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