全 文 ：Studies on Mechanisms Underlying Promotion of Arbuscular
Mycorrhizas on Drought Resistance in Bahia Grass
WU Xue-jun , AN Jing , XIAO Jia-xin ,
YANG An-na , HUA Jun , YANG Bo
(Key Laboratory for the Conservation and Utilization of Im portant Biological Resources in Anhui Province , Colleges of Life Sciences , Anhui Normal
University , Wuhu 241000 , C hina)
Abstract:A pot culture experiment w as conducted to study the phy siological responses of bahia g rass(
Paspalum notatum )to arbuscular myco rrhizal(AM)fungi , Glomus mosseae , G.versiforme or G.
mosseae +G.versi forme colonization under ample water or drought st ress.AM inoculation especially
G.mosseae +G.versi forme significant ly alleviated the grow th reduct ion of drought st ress.Nitrogen
(N), phosphorus(P), potassium , calcium and magnesium contents in shoots , N and P in roots of AM
plants were higher than those in non-AM ones under drought stress.AM inoculat ion increased acid
phosphatase and catalase activities , and total glomalin content in rhizosphere soils.Whether drought
stress o r not , AM inoculation increased shoot peroxidase act ivity and decreased shoot malondialdehyde
content , although no signif icant difference w as observed on shoot superoxide dismutase activity.Under
drought st ress , AM plants had higher soluble sugar and protein contents , but had low er proline
content than those in non-AM ones.Results demonstrated that the benefi t of AM colonizat ion
especially G.mosseae + G.versiforme was due to the enhancement of soil enzymes , nutrition ,
osmoregulation and antiox idase matters in plants , leading to improvement of plant g row th under
drought stress.
Key words:arbuscular mycorrhizae;paspalum notatum ;drought resistance;physiological response
Classification No.:Q945 Document code:A Paper No.:1001-2443(2014)01-0047-06
Drought is considered to be one of the most impo rtant abiotic st ress limiting plant g row th and yield ,
part icularly in arid and semi-arid areas[ 1] .Arbuscular mycorrhizae(AM) is one of the most abundant
underground symbiosis , since AM fungi could colonize approximately 80% of the higher plant species[ 2] .AM
fungi play an impo rtant role in the maintenance of ag roecosystem stability and sustainable agricultural
development.For example , they can improve w ater status of host plant such as cit rus[ 3 , 4] , and pistachio[ 5] .
Clearly , the practice of sowing grasses into no-tillage soil w as developed to reduce soil erosion and costs of
agricultural no-tillage conditions , improve AM levels , fungal diversity and soil quality[ 6 ,7] , and i t is desi rable to
select sod culture as the best practice in hilly cit rus orchard in order to enhance AM fungi benefit s and promote
sustainable ag riculture[ 7 , 8] .Presently , bahia g rass(Paspalum notatum Flǜgge)is w idely applied to sod culture
in orchard of China , improving nutritional level and fruit quali ty[ 9 ,10] , and AM fungi such as Glomus mosseae
inoculat ion can improve g row th and drought resistance of bahia g rass[ 11 ,12] .In general , AM plants maintained
more no rmal w ater relations[ 13] .Potential mechanisms mainly include enhanced w ater absorption by external
Received date:2013-10-06
Foundation i tems:National Natural S cience Foundation of China(31372014);Anhui Provincial Natural Science Foundation(1308085MC37);the
Provincial Natural S cience Research Program of Higher Education of Anhui Province(KJ2012A128).
Author s brief:Wu Xue Jun(1988-), f emale , born in Wuhu , Anhui province , master , Current work unit:Cuiw en Middle School Af filiated to
Anhui Normal University.C orresponding author:Xiao Jia Xin.
引用格式:吴雪俊 ,安静 ,肖家欣 ,等.丛枝菌根提高百喜草抗旱性的机制研究[ J] .安徽师范大学学报:自然科学版 , 2014 , 37(1):47-52.
第 37卷 1期
201 4年 1月 安 徽师 范大 学 学报(自然科学版)Journal of Anhui Normal University(Natural S cience) Vol.37 No.1Jan .2 0 1 4
DOI :10.14182/j.cnki .1001-2443.2014.01.005
hyphae[ 14] , stomatal regulation through ho rmonal signals[ 15] , indirect effect of improved phosphorus[ 16] , and a
g reater osmo regulation in AM plants[ 17] .Whereas , information about the responses of baiha grass plants to
Glomus mosseae and/or G.versi forme under ample w ater or drought st ress is scarce.
Therefore , the purpose of this wo rk w as to evaluate the effects of Glomus mosseae , G.versi forme or G.
mosseae +G.versi forme on g row th , mineral content , osmoregulation and antioxidant matters of bahia g rass
under ample w ater o r drought st ress condit ions , as w ell as soil enzymatic act ivit ies , in o rder to evaluate whether
AM symbiosis helps bahia g rass plants to alleviate injury of drought stress , and further understand drought
resistance mechanisms in AM plants.
1 Materials and methods
Seeds of bahia g rass were surface-sterilized with 70% alcohol for 5 min and germinated on wet filter paper
in Petri dishes in darkness at 25℃.Seven day s af ter germination , seedlings were t ransplanted into plastic pots
(19 cm in depth and 21 cm in mouth diameter)containing 3.0 kg of autoclaved experiment mix ture (0.11
MPa , 121 ℃, 2 h)of yellow soil , quartz sand and vermiculite (3:1:1 , v/v/v), with the pH value 6.85 ,
o rganic mat ter 13.8 g·kg-1 , 0.24 g·kg-1 total nitrogen(N), 10.5 mg·kg-1 available phosphorus(P), 17.18
mg·kg-1 available potassium (K), 356.18 mg·kg -1 available calcium (Ca), 11.25 mg ·kg-1 available
magnesium (Mg), 0.32 mg·kg-1 available zinc (Zn), 0.112 mg·kg-1 available copper (Cu).An aqueous
solut ion of N (N2H4CO)300 mg·kg-1 , P(KH2PO4)150 mg·kg-1 , K(K2SO4)200 mg·kg-1 were added to
autoclaved experiment mixture in each pot.After thoroughly mixing , the mixture w as allowed to stabilize fo r 15
days.
Mycorrhizal inocula , provided by Insti tute of P lant Nutrition and Resources , Beijing Academy of
Agriculture and Forest ry Sciences , consisted of spores , soil , hyphae and infected roo t f ragments from a stock
culture of G.mosseae(No.BGC HUN01A), G.versi forme(No.BGC HUN02B), and propagated on maize
and white clover plants grown in a sandy soil for 12 w eeks.
The inoculated dosage w as 50 g of inocula per pot containing approximately 600 spores.The inocula w ere
placed 5 cm below roots at t ransplant time , non-AM treatment also received the same weight of autoclaved
mixture , all pots were placed in a g reenhouse for 6 months with natural day light f rom April to September , the
average day/night temperature w as 25/20℃.
The experiment w as laid out in a randomized complete block design.Experimental t reatments consisted of
factorial combinations of four myco rrhizal t reatments (G.mosseae , G.versi forme , G.mosseae + G.
versi forme , and non-AMF), with tw o w ater supply condi tions(ample w ater and drought stress).Six replicates
per treatment w ere done totalling 48 po ts.Water treatment began after 60 day s of acclimation in greenhouse
conditions , at which time ample water(AW)pots w ere controlled with 70%of relative soil w ater content , and
drought stress (DS)pots were w atered w ith 40% of relative soil w ater content .The w ater status in the
subst rate w as daily determined and the amount of water loss w as supplied to each po t to keep the designed soil
w ater contents.
Part f resh plants w ere separated into shoo ts and roots , and oven dried at 75℃for 48 h to reco rd shoot and
root dry weight.A fraction of f resh roots w ere used for mycorrhizal colonization determination[ 18] .Myco rrhizal
dependency w as defined as the rat io of the dry weight of the AM seedlings and non-AM seedlings[ 19] .
The dried shoots and roots were wet digested in HNO3-HClO4(4:1 v/v)before analysis of P , K , Ca , Mg
and Zn by inductively coupled plasma optical emission spectrometry (ICP-OES , Pekin Elmer Optimal 2100 DV)
acco rding to the literature
[ 20] .N in the digest w as determined by Kjeldahl analysis[ 21] .
Soil acid phosphatase w as ext racted by the sodium acetate buffer(pH5.0), and its activity was determined
using spectropho tometry , and defined as millig ram of phenol consumption in 1 g air-dried soil for 12 h , soil
catalase (CAT)activi ty w as def ined as the consumption of KM nO4(0.1 mol·L-1)for 1min in 1 g air-dried soil.
48 安 徽 师 范 大 学 学 报 (自 然 科 学 版 ) 2014 年
Total glomalin in soils w ere determined w ith the method of Wright et al[ 22] .
Leaf supero xide dismutase(SOD)(EC 1.15.1.1)act ivity w as measured using the method of Giannopolitis
and Ries[ 23] .Guaiacol peroxidase(POD)(EC 1.11.1.7)activities were measured acco rding to the methods of
Chance and M aehly[ 24] .Malondialdehyde (MDA)concentration w as determinated using thiobarbituric acid
colorimetric method
[ 25] .Soluble protein w as found out by Bradford s procedure using bovine serum albumin as
the standard
[ 26] .Soluble sugar and proline w ere determined using the method of Wang[ 21] .
The data w ere subjected to analysis of variance (ANOVA)using the S tatistical 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 discussion
In our experiment , plants inoculated with mycorrhizal fungi especially G.mosseae + G .versi forme
showed higher plant dry weight , mycorrhizal development and dependency under ample w ater or drought st ress
conditions(Table 1), implying that the combination of G.mosseae and G.versi forme is more ef ficient in bahia
g rass.The increase in g row th has been att ributed to the improvement of w ater uptake result ing in the
enhancement of direct water uptake and transport via ex ternal hyphae
[ 27]
and P nutrition
[ 16 ,28 , 29] .
Table 1 Mycorrhizal colonization, shoot and root dry weight ,
and mycorrhizal dependency of bahia grass plants under dif ferent treatments
Water status
in soi ls
Inoculation
t reatm ent
Mycorrhizal
colonization/ %
Shoot dry
weight/ g·plant -1
Root d ry
w eight/ g·plant-1
Mycorrhizal
dependency/ %
Ample w ater
(AW)
Non-AMF 0.00d 0.12±0.02b 0.06±0.01bc -
G.m 58.60±0.03b 0.13±0.01ab 0.07±0.01abc 149
G.v 59.60±0.01b 0.13±0.00ab 0.07±0.02abc 150
G.m +G.v 70.00±0.03a 0.14±0.01a 0.09±0.01ab 203
Drought st ress
(DS)
Non-AMF 0.00d 0.12±0.02b 0.06±0.01bc -
G.m 55.90±0.04b 0.14±0.01a 0.07±0.00abc 153
G.v 48.20±0.04c 0.13±0.01ab 0.08±0.02abc 156
G.m +G.v 68.50±0.03a 0.14±0.01a 0.10±0.01a 180
No te:Non-AMF , G.m , G.v and G.m +G.v represent non-inoculat ion , inoculation with Glomus
mosseae , G.versiforme and G.mosseae +G.versi forme , respectively.Different let ters in the same column
mean signif icant differences at P<5%.The same is in the follow ing table and figures.
In the present investigation , myco rrhizal plants had higher levels of N , P , K , Ca , and Mg compared w ith
non-mycorrhizal ones (Table 2), which w ere consistent w ith those found by many invest ig ato rs , such as
Subramanian et al[ 30] , who found that mycorrhizal tomato plants had significantly higher uptake of N and P in
both roots and shoots regardless of intensities of drought st ress.Similarly , N , P , K and Ca contents in AM-
treated pistachio shoots were greater than those in non-AM shoots under drought st ress[ 5] , the higher levels of
K , Ca and Mg in leaves and roots w ere observed in AM tangerine seedling s than those in non-AM seedlings
under w ater st ress
[ 4] .These results indicated that myco rrhizal plants tended to increase nutrient acquisit ion
under drought st ress , which probably resulted from the greater absorption of the surface area provided by
ex tensive fungal hyphae[ 14 , 31] .The role of AM fungi on the uptake of P o r N is well-documented in the
literatures.Potassium plays a key role in plant drought tolerance and has been found to be the cationic solute
responsible for stomatal movements in response to changes in bulk leaf w ater status[ 14] , thus , the protection of
mycorrhizal plants against drought stress is part ially related to K uptake.On the other hand , plants accumulated
some inorganic ions such as K
+ , Ca2+ and Mg2+ , which resulted in a g reater osmotic adjustment in AM plants
and allowed AM plants to accumulate more carbohydrates and increase plant biomass.
Our data revealed that there w ere increases in acid phosphatase and catalase activities of mycorrhizal soil
under well-watered and w ater stress conditions (Figure 1), which may be at tributed to alterat ion in root
phy siology and secretion of root exudates by mycorrhizal colonized plants.The higher acid phosphatase and
4937卷第 1期 吴雪俊 ,安 静 , 肖家欣 ,等: 丛枝菌根提高百喜草抗旱性的机制研究
catalase activit ies in mycorrhizal soil make P available and hydrogen pero xide elimination , respectively.Several
experiments have demonst rated that AM inoculation increased enzymatic activities in rhizophere of many plant
species
[ 30 ,32] .The intense enzymatic activities may be contributed to the enhanced availability of nutrients in
acco rdence w ith the result of mineral levels ment ioned above in mycorrhizal plants.In addition , total g lomalin
content was relatively high in rhizosphere soils inoculated wi th G.mosseae +G.versi forme under w ater st ress
(Figure 1).Glomalin , a glycoprotein copiously produced by AM fungi , is tight ly correlated with soil ag gregate
stability
[ 33] , and could influence soil carbon storage[ 34] .Wu et al[ 35] repo rted that glomalin can g lue smaller
macroaggregates into macroaggregates (water-stable agg regates), and the increased w ater-stable aggregate due
to mycorrhizal symbiosis lead to better soil st ructure and might alter soil moisture retention properties that , in
turn , lead to bet ter plant drought resistance.
Table 2 Nitrogen , phosphorus , potassium , calcium and magnesium contents
in shoots or roots of bahia grass under dif ferent treatments.
Water
status
in soils
Inoculation
t reatment
Nit rogen
/ g·kg -1DW Phosphorus/ g·kg
-1 DW
potassium
/ g·kg -1DW
Calcium
/ g·kg -1 DW
Magnesium
/ g·kg -1 DW
shoot root shoot root shoot shoot shoot
Ample
water
(AW)
Non-AMF 23.28±1.01c 22.21±1.21c 0.91±0.08bc 2.41±0.06c 18.01±1.46bc 6.82±0.44d 1.64±0.12bc
G.m 23.94±0.86bc 23.50±0.56bc 1.23±0.08a 2.52±0.10bc 22.56±1.24a 9.62±0.56ab 2.26±0.18a
G.v 25.16±0.42b 24.49±0.40b 1.08±0.21ab 2.53±0.12bc 19.79±3.61ab 8.84±1.48bc 1.84±0.35ab
G.m +G .v 27.39±0.63a 26.65±1.01a 1.11±0.17ab 2.67±0.11ab 19.40±2.26ab 8.34±0.97bcd 1.86±0.27ab
Drought
st ress
(DS)
Non-AMF 14.48±0.82e 13.22±0.62f 0.16±0.03e 2.16±0.13d 12.69±0.38d 3.70±0.48e 0.44±0.06d
G.m 17.41±0.58d 16.63±1.37e 0.59±0.10d 2.70±0.06ab 15.14±2.54c 6.76±1.23d 1.27±0.22c
G.v 19.12±1.22d 18.86±1.07d 0.79±0.13cd 2.64±0.13ab 18.81±2.29abc 11.03±1.51a 1.68±0.28bc
G.m +G .v 19.02±1.53d 19.07±0.94d 0.98±0.03bc 2.77±0.07a 19.61±0.77ab 7.66±0.29cd 1.77±0.08b
Figure 1 Acid phosphatase and catalase activit ies , and total
glomalin content in rhizosphere soils under diff erent t reatments.
AW and DS represent ample w ater and drought st ress ,
respectively.Bars w ith the same letter are not signif icant ly
dif ferent at p <0.05.The same is in the follow ing figures.
Figure 2 Superoxide dismutase(SOD) and peroxidase(POD)
activit ies , and malondialdehyde(MDA)con tent in leaves of bahia
grass under dif ferent t reatments.
50 安 徽 师 范 大 学 学 报 (自 然 科 学 版 ) 2014 年
Figure 3 S oluble protein , soluble sugar
and proline con tents in leaves of bahia grass
under diff eren t treatments
In this wo rk , AM inoculation especially G.mosseae +G.versi forme
increased POD activity , decreased MDA concentration in shoots under w ater
stress (Figure 2), which indicated that mycorrhizal infection increased
activi ties of defense enzymes , especially POD , involved in the elimination of
reactive oxygen species (ROS), and alleviate cell lipid peroxidation ,
protecting the organism against oxidative damage.In contrast to POD
activi ty , shoo t SOD activity did not vary signif icantly among the t reatments
under AM colonization(Figure 2), this agreed with previous reports obtained
from leaves of trifoliate o range under water stress[ 3] .Drought stress induced
an increase of SOD activity in AM and non-AM plants , result ing in great
drought tolerance of bahia grass plants reg ardless of AM fungi colonizat ion.In
addi tion , o ther defense mechanisms based on antio xidant small molecules may
protect cell f rom ROS[ 36] .Acco rding ly , myco rrhizal plants had higher
contents of soluble sugar and protein in shoo ts under drought st ress(Figure
3), indicating that AM infection might alleviate o r decrease RNA disassembly
and might enhance the ability of the non-enzymatic antioxidant defense
sy stem[ 5 , 37] .Addit ionally , proline w as low er in shoots of AM plants than in
non-AM shoots when exposed to drought stress conditions (Figure 3).This
result may be att ributed to either greater drought resistance of AM plants or
less injury in AM plants g row n under water st ress condition[ 4 , 5 , 38 ,39] .Thus ,
the accumulation of low molecular mass o rganic solutes such as soluble sugar or pro tein , proline in plants could
regulate the osmot ic potential of cells aiming at improving absorption of w ater under drought stress , leading to a
lower stress injury in the plant.
3 Conclusion
In conclusion , AM colonization especially G.mosseae +G.versiforme enhanced soil enzymes , antioxidant
enzymes , adjusting osmot ic and nutrient acquisition under drought stress , leading to higher g row th.
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52 安 徽 师 范 大 学 学 报 (自 然 科 学 版 ) 2014 年
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[ 28] Kyoung-Hee O , Dong-Hw an J , Young-Cheol C.Quanti fication of Toxigenic Microcystis spp.in Freshw aters by quan titative real-time PCR
Based on the Microcystin S ynthetase A gene[ J] .Jou rnal of M icrobiology , 2013 , 51(1):18-24.
[ 29] 郑春艳 ,张庭廷.鞣花酸对铜绿微囊藻和斜生斜藻的生长抑制作用[ J] .安徽师范大学学报:自然科学版 , 2008 , 31(5):469-472.
[ 30] HU W , ZHANG D , RAN YX , ZHANG TT.T he allelopahy and allelopathic mechanism of Mgriophyllum spicatum on freshw ater alage[ J] .
Journal of Anhui Norm al University(Natural Science), 2011 , 34(4):359-364.
The Research Progress of Microcystins s Harm and Its Analysis Techniques
ZHANG Ting-ting , ZHANG Sheng-juan ,
(The Key Laboratory of Biotic Environment and Ecological Safety in Anhui Province , College of Li fe Sciences , Anhui Normal Universi ty , Wuhu
241000 , China)
Abstract:Microcystins is a kind of algae toxins w hich occurs in high f requency , produces a mass of toxin
production , results in the most serious consequence to the pollution of cy anobacterial blooms.The fo rming
mechanisms has become a g lobal environmental issue w hich needs to be solved.Based on the latest related
literatures at home and abroad , this article not only summarized the production and mechanism of microcystins ,
phy sical and chemical propert ies , but also introduced the research progress of its analy sis methods as well as the
hazards of microcyst ins to org anisms (animals , plants , microorganisms and human)in detail.At last , the
development direction of determination analysis technology of microcy stins and its pollution control w ere
prospected.
Key words:microcystins;cyanobacteria;toxicity;HPLC;ELISA
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丛枝菌根提高百喜草抗旱性的机制研究
吴雪俊 , 安 静 , 肖家欣 , 杨安娜 , 滑 俊 , 杨 博
(安徽师范大学生命科学学院 ,安徽省重要生物资源保护与利用研究省级重点实验室 ,安徽芜湖 241000)
摘 要:通过盆栽试验研究了水分充足和干旱胁迫下百喜草(Paspalum notatum )对丛枝菌根(AM)真菌
Glomus mosseae 、G.versi forme 及 G.mosseae + G .versi forme 侵染的生理反应.AM 真菌尤其是 G.
mosseae +G.versiforme 接种处理显著减轻了干旱对生长的抑制.干旱胁迫下 ,接种株地上部 P 、K 、Ca 、Mg
及根部 N 、P 含量均显著高于未接种株 ,AM 真菌接种处理显著提高了根围土壤酸性磷酸酶与过氧化氢酶活
性及总球囊霉素含量.无论干旱胁迫与否 ,AM 真菌接种处理增加了地上部 POD 活性 ,降低了地上部 MDA
含量 ,而地上部 SOD活性并无显著性差异.干旱胁迫下 ,与未接种株相比 ,接种株可溶性糖与蛋白含量较高 ,
而脯氨基酸含量却较低.可见 ,AM 真菌尤其是 G.mosseae +G.versi forme侵染有利于提高土壤酶活性 ,
改善植株营养 、渗透调节能力与抗氧化能力 ,进而促进了干旱胁迫下植株的生长.
关键词:丛枝菌根;百喜草;抗旱性;生理反应
5737卷第 1期 张庭廷 ,张胜娟: 微囊藻毒素的危害及其分析方法研究进展