全 文 ：Influences of Three Arbuscular Mycorrhizal Fungi on Growth 、
Mineral Contents and Drought Resistance of Bahia Grass
WU Xue-jun , 　XIAO Jia-xin , 　YANG An-na , 　An Jing ,
CHEN Ying-ying , 　HU Cheng-yin
(Provincial Key Laboratory of Biotic Envi ronment and Ecological Safety in Anhui , College of Life Sciences , Anhui Normal University , Wuhu 241000 ,
China)
Abstract:The ef fects of three arbuscular myco rrhizal(AM)fungi Glomus aggregatum(G.a), G.
etunicatum(G.e)and G.intraradices(G.i)colonization on grow th , mineral nut rient status and
several physiological indices of bahia grass(Paspalum notatum Flugge)under drought st ress w ere
investigated.AM inoculat ion significantly increased root leng th and shoot dry weight , the order of
mycorrhizal dependeny of plants w as:G.a >G.i >G.e.Root P , K , Mg and Cu contents in G.a
and G.e inoculated plants , root P , Mg and Ca contents in G.i inoculated plants , root Ca content in
G.e inoculated plants w ere significant ly higher than those in non-AM plants , whereas , shoot P , K ,
Mg , Ca and Cu contents w ere low er in G.e and G.i inoculated plants than in non-AM plants.When
drought stress for 4 days , POD activit ies in G.a and G.e inoculated shoots , proline content in G.a
inoculated shoots were significantly higher than those in non-AM shoots , when drought stress fo r 8
days , no significant differences were observed on SOD and POD activities among treatments , while
AM inoculation enhanced proline content notably , MDA contents were significantly lower in G.e and
G.i inoculated shoots than in non-AM shoots.In summary , among the three AM fungi G.a
inoculat ion played a posi tive role in the enhancement of grow th and drought resistance of bahia g rass ,
while the ef fects of G.e and G.i inoculation were relatively slight .
Key words:arbuscular mycorrhizal fungi;bahia g rass;mineral nutrient;drought stress
CLC Number:Q945 ,S666　　Document Code:A 　　Paper No:1001-2443(2011)05-0461-06
　　Drought is one of the most common environmental st resses on plants.Arbuscular myco rrhizal(AM)fungi
can establish symbiotic relationship with approximately 80% of the land plant species , contributing to improve
the resistance of host plants.Document show s that inoculat ion wi th AM fungi like Glomus mosseae can enhance
the drought tolerance of host plants such as cit rus[ 1-3] , korshinsk peashrub[ 4] , wild jujube[ 5] , whereas ,
information about the benef icial roles of different AM fungi in baiha g rass(Paspalum notatum Flugge)under
drought stress is scarce.
　　Sod culture in o rchard not only can effectively avoid w ater and soil erosion , but also can enhance organic
mat ter , mineral content in soil[ 6] , hence , i t is thought to be an important planting technology in modern eco type
o rchard.Presently , bahia grass is widely applied to sod culture in orchard of China[ 7](Li et al ,2005), which can
increase f rui t t ree roo t colonizat ion by AM fungi , and improve nutri tio nal level and frui t quality[ 8-10] .However ,
the ef fects of dif ferent AM fungi on g row th and drought resistance of common grass of sod culture in o rchard
such as baiha grass is still unclear.The purpose of this study is to evaluate the effects of AM fungi Glomus
Received date:2011-04-20
Foundation item:Supported by S pecialized Research Fund for the Doctoral Program of Higher Educat ion(20103424120002), China Postdoctoral
S cience Foundation(20080430172)and Anhui Provincial Natural S cience Foundat ion(070411004).
Author s brief:Wu Xue-jun(1988-), f emale , master , mainly study on plant physiology and ecology.
第 34卷 5期
201 1年 9月 　　 安 徽师 范大 学 学报(自然科学版)Journal of Anhui Normal University(Natural S cience) Vol.34 No.5Sept .2 01 1
DOI :10.14182/j.cnki .1001-2443.2011.05.007
aggregatum , G.etunicatum and G.intraradices on grow th , physiological character of bahia g rass underwater
stress conditions , w ith the aim of finding out the ecological effects of sod culture in orchard.The result w ill play
a theoretical and basic role in taking practical steps for production in the area.
1　Materials and methods
1.1　Experimental materials
　　Seeds of bahia grass provided by the Center of Special Local Product Technology Popularization of Yiling
Region of Yichang City , Hubei province of China , were surface-sterilized w ith 70% alcohol for 10 min and
placed on sterilized moist filter paper for germination in darkness at 25℃.Seven days af ter germination ,
seedlings were transplanted to a plastic pot contained 3.0 kg of an autoclaved g row ing mix ture(fumigated w ith
0.5% fo rmaldehyde for 7 days)of yellow soil and quartz sand(9:1 , v/v), w ith the pH value 6.1 , 11.05 mg·
g -1 available phospho rus , 39.28 mg ·g -1 available potassium.The experimental po ts were placed in a
g reenhouse under natural condi tions w ithout any temperature control f rom April to August.
　　Myco rrhizal fungi inocula , provided by Bank of Glomales in China , Institute of P lant Nutrition and
Resources , Beijing Academy of Agriculture and Fo restry Sciences , consisted of spores , soil , hyphae and infected
root f ragments f rom a stock culture of Glomus aggregatum (No.BGC BJ07), G.etunicatum (No.BGC
GZ03C)and G.intraradices (No.BGC AH01).Twenty g rams of inocula w as inoculated per po t.The inocula
w ere placed 5cm below baiha g rass roo ts at the time of transplanting.Non-AM treatment also received the same
w eight of autoclaved mix ture.These seedlings did no t get additional nutrients during the entire experiment.
1.2　Experimental design
　　The present experiment consisted of a randomized block design w ith two factors:(1)AM fungi G.
aggregatum(G.a), G. etunicatum(G.e), G. intraradices(G.i) inoculation , and non-AM fungi
inoculat ion , and(2)water stress t reatments (cut of f w ater supply for 0 day , 4 days , 8 day s).Water st ress
treatment began after 75 day s (July 19 , 2010) of acclimation in natural conditions.Each of the twelve
treatments w as replicated three times , leading to a total of 36 pots.
1.3　Experimental methods
　　At the time of w ater st ress treatment , a f raction of the plants w as harvested and plant height , main root
leng th , dry shoot and root w eight were reco rded.Concentrations of P , K , Mg , Ca , Zn and Cu in the dried
shoots and roo ts w ere determined by inductively coupled plasma optical emission spect rometry (ICP-OES , Iris-
Advantage type , USA), according to the literature[ 11 ,12] .A fraction of fresh roots were carefully w ashed and
cut into 1 cm root pieces to fix in fo rmalin-acetic acid-alcohol solutions , these roo ts were cleaned with 10%(w/
v)KOH and stained w ith 0.05%(w/v)t rypan blue in lactophenol[ 13] .AM colonization was observed by
microscope and counted by the formula , AM colonization (%)=100 ×root length infected/ root leng th
observed , myco rrhizal dependency(%)=100×dry w eight of AM plant/dry w eight of non-AM plant.
　　A fraction of the plants w ere harvested at the w ater treatment t ime , samples w ere quickly freezed by liquid
nit rogen , and stored in ref rigerator fo r superoxide dismutase(SOD), peroxidase(POD)act ivit ies , chlo rophyll ,
proline and malondialdehyde(MDA)contents analy sis , according to the literatures[ 14 , 15] .Relative conductivity
w as determined by conductivity meter(DDS-307).
　　The data w ere subjected to analy sis of variance (ANOVA)by using the Statistical Analy sis System (SAS)
sof tw are.Fo r comparison of the means , the Duncan s multiple range test w as employed.
2　Results and analysis
　　Colonization was higher in G.a and G.e inoculated roo ts than in G.i inoculated roo ts.AM inoculat ion
markedly increased main root length and shoot dry weight , G.a and G.i treatments increased the root dry
w eight significantly , but no significant differences w ere found on root dry weight between G.e inoculated plants
462 安 徽 师 范 大 学 学 报 (自 然 科 学 版 ) 2011 年
and non-AM plants.The order of mycorrhizal dependency of plantsw as:G.a >G.i >G.e.
Table 1　Root and shoot characters of AM and non-AMbahia grass plants
Treatments
Root
colonization(%)
Plant height
(cm)
Main root
length(cm)
Shoot dry w eight
(mg·plant -1)
Root dry w eight
(mg·plant-1)
Mycorrhizal
dependency(%)
non-AM 0c 1.12±0.33b 0.57±0.03c 101.5±6.17b 64.8±11b 100
G.a 56.00±2.64a 1.30±0.13ab 0.85±0.05a 137.1±5.1a 86.5±6.9a 159.6
G.e 60.33±4.73a 1.43±0.08ab 0.70±0.05b 135.7±6.06a 66±5.7b 136.9
G.i 25.33±2.08b 1.52±0.08ab 0.72±0.08b 136.3±12.08a 99±8.6a 155.7
　　Different letters in the same colum n mean significant dif ferences at P <0.05.
Figure 1　Phosphorus , K , Ca , Mg , Zn and Cu concent rations in shoots
and roots of AM and non-AM bahia grass plants
　　AM inoculat ion no tably increased roo t P , K , Mg and Cu contents (K and Cu contents in G.i inoculated
root excluded), G.e and G.i infection increased Ca content significant ly .Whereas , no signif icant dif ferences
w ere observed on P , K , Mg , Ca and Zn contents in G.a inoculated shoots as compared w ith those in non-AM
shoots , G.e and G.i inoculation markedly decreased P , K , Mg , Ca and Cu contents in shoots , and there was
no signifiant diference on Zn content betw een in AM and non-AM roots.
　　At the well w ater , SOD activities w ere significantly higher in G.a and G.e inoculated shoots than in non-
AM and G.i inoculated shoo ts , whilst AM inoculation significant ly increased POD activi ty.When drought
stress for 4 days , G.a and G.e inoculation increased POD activity significantly , while no significant dif ferences
46334卷第 5期　　　　吴雪俊 ,肖家欣 , 杨安娜 ,等:　三种丛枝菌根真菌对百喜草生长及抗旱性的影响
were detected on SOD activity betw een in AM and non-AM shoots , when drought stress for 8 day s , AM
inoculat ion didn t af fect the SOD and POD activities of shoots (Figure 2A , 2B).MDA and proline contents in
non-AM and AM inoculated shoots w ere increasing progressively wi th the increase of drought degree , when w ell
w ater , MDA and proline contents had no significant dif ferences among treatments , when drought st ress for 4
days , proline content in G.i inoculated shoo ts w as signif icantly higher than o ther t reatments , when drought
stress for 8 days , G.e and G.i inoculation notably decreased MDA content , and proline contents w ere
signif icantly higher in AM inoculated plants than in non-AM plants (Figure 2C , 2D).
Figure 2　SOD(A)and POD(B)activities , MDA(C)and proline(D)contents in shoots
f rom AM and non-AM bahia grass plants subjected to w ater stress
3　Discussion
　　The present study show ed that the inoculation wi th AM fungi significant ly promoted the g row th of bahia
g rass(Table 1).The contents of P , K , Mg and Cu in G.a and G.e inoculated roo ts , P , Mg and Ca contents in
G.i inoculated roots , Ca content in G.e inoculated roots were all significantly higher than in non-AM roots
(Figure1), which suggested that there would exist close relationship betw een the g row th promoted by AM
symbiosis and the increase of P and other related mineral elements uptake by plants.P is generally believed to be
removing poorly in the soil , the grand hyphal net fo rmed in the soil by AM can travel across P deficiency area
around the root , which can enhance the contact w ith the soil and turn unavailable P around root into available P
to roo t , the improvement of P , K , Ca absorption by AM , and the direct uptake and transpo rt of the ext raradical
hypha outside the root are all likely to increase the contents of P , K in plant , which is consistent wi th the
finding s that the field grass cultivation can effectively increase the contents of P , K in cit rus shoots and satsuma
mandarin[ 7 , 10] .The results mentioned above indicated that the g rass such as bahia g rass in orchard w ould highly
developed roo t system and be easily infected by AM fungi , which is beneficial to the activi ties of soil mineral
elements , and thus promote P , K , Ca uptake by roo t.However , in this study , G.e and G.i inoculat ion
decreased P , K , Mg , Ca and Cu contents in shoots.This result may be related to the increase of plant g row th
464 安 徽 师 范 大 学 学 报 (自 然 科 学 版 ) 2011 年
promo ted by AM resulting in “dilution” effect .It also show ed that the effects of AM symbiosis would depend on
mineral elements , AM fungi and host plant species.
　　SOD and POD are the main ant ioxidant enzymes in plant , the important function of SOD is to remove
superoxide f ree radicals so as not to generate H2O2 , while POD can decompose H2O2 , these two enzymes wo rk
together to maintain the relatively low levels of the f ree radicals in plant , w ith an at tempt to controlling
membrane lipid peroxidation and reducing the membranes damage.When drought st ress for 4 days , POD
activi ties in G.a and G.e inoculated shoo ts were significantly higher than in non-AM shoots , whilst G.a
inoculat ion increased proline content no tably.When drought stress for 8 day s , proline contents in AM shoots
w ere significantly higher than in non-AM shoots , and proline content in G.a inoculated shoots was the highest
among the t reatments , inoculation with G.e and G.i decreased MDA content signif icantly.Proline is the ideal
o rganic matter of osmotic adjustment , and MDA is the breakdown product of membrane lipid pero xidation.As a
result , inoculation with AM fungi such as G.a and G.e can remove the accumulation of reactive oxygen caused
for the w ater st ress by improving POD activities , and adjust cell osmotic potential to protect the activities of
enzyme systems by promoting the accumulation of proline as w ell as P , K , w ith the aim of reducing the damage
caused by membrane lipid pero xidat ion , which in return enhance the drought tolerance of bahia g rass.
　　In summary , under drought st ress condition , AM fungi inoculation had a positive effect on grow th , osmotic
adjustment , and reactive oxygen metabolism of bahia grass , increasing osmoregulation mat ter(proline , P , K
etc.), protective enzyme (POD)act ivity in shoots or roots , and decreasing MDA content .The results
mentioned above indicated that the benef it of AM colonization under w ater st ress w as due to the enhancement of
osmot ic adjustment and higher antio xidant enzyme.Glomus aggregatum inoculation was most beneficial to the
tolerance of bahia g rass against drought stress , while the effects of G.etunicatum and G.intraradices
inoculat ion w ere ralatively sltig ht .
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46534卷第 5期　　　　吴雪俊 ,肖家欣 , 杨安娜 ,等:　三种丛枝菌根真菌对百喜草生长及抗旱性的影响
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三种丛枝菌根真菌对百喜草生长 、矿质元素含量及抗旱性的影响
吴雪俊 , 　肖家欣 , 　杨安娜 , 　安　静 , 　陈迎迎 , 　胡成银
(安徽师范大学生命科学学院 ,生物环境与生态安全安徽省高校省级重点实验室 ,安徽 芜湖　241000)
摘　要:研究了三种丛枝菌根(AM)真菌 Glomus Aggregatum(G.a)、G.etunicatum(G.e)和 G.
intraradices(G.i)对百喜草(Paspalum notatum)生长 、矿质元素含量及干旱胁迫下几种生理指标的影响.
3种AM 真菌处理显著提高了主根长和地上部干重 , 菌根依赖性依次为 G .a >G.i >G.e.G.a和G.e接
种株根部 P 、K 、Mg 、Cu含量 , G.i 接种株根部 P 、Mg 、Ca含量及 G.e 接种株根部 Ca 含量均显著高于对照;
G.a接种株地上部P 、K 、Mg 、Ca和 Zn含量与对照无显著性差异 ,而 G.e 和G.i处理的地上部P 、K 、Mg 、Ca
和 Cu含量均显著低于对照.干旱胁迫 4d时 , G.a和G.e处理的地上部 POD活性 、G.a处理的地上部脯氨
酸含量均显著高于对照.干旱胁迫 8 d时 ,各接种株的 SOD和 POD活性与对照并无显著性差异 ,而接种 AM
真菌显著提高了地上部脯氨酸含量 , G.e和 G.i 处理的 MDA含量显著低于对照.三种 AM 真菌以 G .a接
种处理一定程度上提高了百喜草的生长及抗旱性 ,而 G.e 和G.i的效果相对较弱.
关键词:丛枝菌根真菌;百喜草;矿质元素;抗旱性
466 安 徽 师 范 大 学 学 报 (自 然 科 学 版 ) 2011 年