Competition between Pratylenchus coffeae and Meloidogyne incognita

Plant Pathology Bulletin 17: 271-278,2008

五{nita

Bie Yun Tsai

Department of Plant Pathology and Microbiology,National TaiwanUniversi吟， Taip凹 ，Taiwan E-mail:b問yntm@ntu.edu.tw:Fax: +886-2-2363-6490

Accepted for publication: February 22,2008

也STRACT

Tsai,B.Y.2008. Competition betweenPraty{enζ !Ius C~妒。e HI泌 Me/oidogy附 incog n ita .Plant Palhol Bull.17: 271-278.

Pratylenchlls co

.ffeae

ar吋 Meloidogyneincognitawere mutually antagonistic on mung bean roots

The penetration of both P.cofJ切eand [v!. incognita was significantly reduced when they were inoculate吐simultaneouslycomparing to when they were inoculated alone. In the simultaneous inoculation the number of sausage-shaped M. incognita was significantlylow盯thanwhen M

incognitawas inoculated alone,howey凹 ，the percentage of M. il1ζog川的second叫agejuveniles

developed into sausage shape was not significantly different from that of the single inoculation Inoculation simultaneously with the ratio of P. corfeae and M. illcognitα 7 5 0: 250 or 250 : 750, resulted insigni日cantsuppression of penetration of both nematodes comparing to when they were inoculated alone. Two and 4 days prior inoculation ofP.coffeaedid not influence the late coming M

incognita,but prior inoculation of 111. incognita inhibited the penetration ofP.c(~ffi叩e.The substance

secreted by .M. incognita or produced by the host in response to infection might have been involved in the latter case. Elucidating the substance呵呵 pon s iblefor the inhibition of P. coltcae might lead to a novel way of controlling the nematodes

Key words:antagonistic可competltlO口 ，Prary/ellchuscojt切e ，Meloidogyne incognita

INTRODUCTION

Comoetition between ofQanismsexists widely in

nature. It has been known to exist among plants(7日 insectsl只 " entomopathogenicnematode:、 仆的 ， bacteria 付 " and even between spenns Competition between plant parasitic nematodes were also observedII.:.4.:\， ~，lO,12. 14. 1:\,16,17

In a field situation,it is notuηusualtohave more then one species of plant-p 的asiticnematodes. When the nemat仁des have overlapping host ranges,competition for food can be expected if the host plant tissues are insufficient for theexistin芷nematodepopulations. When there is sufficient host root resource,or nematodes have

different spatial aggregationniches， 吐iffen己nt nemato吐es

species may coexist in a field. Bell and Watson(2)reported

that no competition between Paratrichodorus minor and

Paratylenchusl1allUSpopulations was found in a grazed

pasture in New Zealand. Additionally,Brinkman,et al.{在j found no competition betweenHeteroderα αrenαnα an d

A1eloidogyne maritime on the natural dune grass

J1mmophila arenaria.Howev凹 ，there wen己morereports on

compet!訂on 出thanlack of competition between nematodes

Gay and Bird(12)found that population of M. incognita

was inhibited by the presence of Pratylenchusbrαζhyurus

2721 直物病理學會刊 第 1 7 卷 第 是 期2∞8

of Scutel!onemabradyswas greatly inhibitedI乃lP.coffeae

on Guinea yam. O'Bannon. et 01.(恥 foundthat Radopholus SIr.叫 /isand P. cojfeae inhibited each other on citrus. In addition,Yang,et al.'υ d的coveredthatIV!.incognitaand

Hoplo/aimllS ga/calUSwere mutually antagonistic on

cotton. Brinkman‘ e f al.(-1ireported that P. penetrans

suppressed theabun丘之lI1ceof H. arenaria on the natural dune grass

The interaction of different species of nematodes may not be always antagonistic to each other.Gay and Birdi>L

reported that l\i1.incognitaincreased the population ofP.

hrachyuruson cotton when inoculated simultaneously 01

when AI. incognita waS inoculated first.The effect of nematodes on the host plant may be altered when two species of nematodes attack the samepI剖1tsimultaneously Griff的studiedthe interrelationship of lvi.hap的 an d

Ditylenchus dipsacion the resistant and susceptible alfalfa

andfaun吐asynergistic weight depression on Ranger,a cultivar susceptible to bothnematodes 可withsimultaneous inoculation of the two nematode species

p.C(~f(e a e(Zimmermann) Filipjev& Schuurmans Stekhoven and the root-knot nematode A1. incognita (Kofoid& White)Chitwoo吐areboth important plant-parasitic nematodes in Taiwan. Un丘erstandinghow they interact with each other is important for a successful nematode ma缸1gementstrategy. The purpose of this study was to investigate the effect of simultaneous and sequential inoculation of P. cqjfeae and M.incognitα on host penetration and nematode development under controlled conditions

JVIATERIALS AND METHODS

Preparation of Nematodes

The lesion nematode P. cof

t'eae

was originally collected from the experimental station of the National Taiwan University and cultured on excised tomato roots in Gamborg's B5 medium (GlBCOBRL)following surface sterilization withH20!(3%) for 2 min. The nematodes from one-month old culture were extracted withmodi在ed Baermann funnels‘collecteddaily and kept at 15°C.They were used within three days

A population ofIv!.incog川的 W恥 alsocollected from the experimental stationof 桂1eNational Taiwan University

and cultured on mung bean(Vignαradiala (L.)\九!ikze k )

seedlings grown in Seed-Pack Growth Pouch (Mega International of Minneapolis). Eggmasse污werεdissected from roots and hatched in a hatchimz chamberν ill;.Fresh second-stagejuveniles 以ierecollected every day to keep at 15"C,and were used within three days

Preparation of seedlings

Mung bean seeds were surface sterilized with 70(}(, ethanol for 4mit日utes‘rinsed th閃e tllne后withtap water. and sown in Seed-Pack Growth Pouch (Mega International of MinneaDolisL Another batch of mung:bean seeds. after surface sterilization,was sown in sterile sands in plastic cups. All of the above were kept at 28GC

in a growth chamber with 16 h phOloperiod.The seedlings were used five days after sowing

E

!Tect

Oil penetratioll

ExperiInent 1~Inoculation with equalnUIubeI有of both nematode species (500 nematodes each)

For the simultaneous inoculation.P.c(彷'caeand M lJ1 cogm tα i n2ml suspension each were inoculated to a mung bean seedling in the growth pouch or in the cup with sterile sand. Two ml sllspension of each species plus 2 ml of distilled water were inoculated as the control (single inoculation).FOI 時quential inoculati凹 ， 戶 川汀ωewas inoculated 2 and 4 days priortothe inoculation ofAt

incognitato the mung bean seedlings in the growth

pouche~弘orvice versa.The controls (single inoculation) of the prior species were inoculated at the same time as the sequential inoculation. The controls (single inoculation) of the later species were inoculated to the seedlings of 2“and

4-days older than those for the sequential inoculation,so that the aQC of the seedlilH!s \vas the same as for the sequential inoculation

Experiment 2- Inocula吋onwith unequal numbers of nematode species (P. cojj-'eae:M. incognita::::::750

250 or 250 : 750)

The two species were inoculatedsimultaneous妙 ，01 singly as the control,to the mung bean seedlings in the gro\vth pouches

All of the seedlings in experiments I and 2wer它kept in the 28''( growth chamber for 3 clays after inoculation,01 3 days after the final inoculation in the sequential

inoculation treatments,and then the roots were stained with acid-fuchsin16> The nematodes in each rootsy叫em were countedun吐er a 吐issectingmicroscope (Olympus SZ6l) .There were four replicates for each treatment.Both experiments were repeatedt丸NIce

Effect on development and reproduction fhe inoculation was done the same way as in the above for the simultaneous inoculation with equal numbers of each species to seedlings in戶 the 凹owthDouches. The plants were kept in the growth chamber for 10 days for the development test and 30 days for reproduction. After 10 days 吋theroots of the seedlings were stained with acid

'、J

fuchsin. The numbers of vermiform juveniles and the sausage-shaped juveniles of I\ll.incognitawere counted sep

‘Irately.

As thecountir嚕 。fdifferent stages ofP.c(iff削 in the roots was toostrenuous可thenumber of eggs in the~~ roots was counted along with total number of nematodes instead. After 30 days‘ t h enumbers of galls and egg

masses were counted. The number of adult females per root system was counted after teasing the root tissue apart to expose the females in the galls. There were fOUl replicates for each treatment.The experiment was repeated tWIce

One-way analysis of variance was carried out with

SAS9.1 software (SASInstitu缸 ，Cary,NC,USA) and treatment means were compared with t-test at the 95% level ofconfi 吐ence.Arcsine square root transformation wasperforme吐forpercentages before statistical analysis. In the repeated experiment,the data weresimilar‘so 吐ata from two experiments were pooled in the results

Competition betweenPratγlenchllsand l

w.eloidogyne

273

RESULTS

The two nematodes species inside the mung bean roots could be separated by the ditTerentshapes of theil tails after staining with acid-fuchsin (Fig. 1).The 恤ilofM

incognitatapered gradually into a very pointeden且while

P. cofleaehad a blunter tail. Therefore 可thenumber of

nematodes of each species could be counted accurately inside the same root system in the experiments

In experiment 1,the 間mberofP.C(~tleaepenetrated the roots was significantly reduced whenP. ω!fleaeand M

incognitawere inoculated simultaneously comparing to

when P.c(~tleaewas inoculated alone(Tat了Ie I). The average of 175 P. cofleaepenetrate吐theroots in the single inoculation compared to only 58.3 nematodes penetrated the roots in the concomitant inoculation with equal numbers of M. incognita. The number of lv!. incognita penetrated the roots was significantly reduced whenP.

cojleae and M. incognita wereinoculated 叩叫taneou向

comparing to when M. incognita was inoculated alone (Table I). There was no significant difference in penetration by P. cqfleae when P. cofleae was inoculated 2 days or 4 days prior to the inoculation of M. incognita. However,when M. incognita was inoculated 2 days or 4 days prior to the inoculation ofP. 叫fieaethe inhibition on penetration ofP. 叫 fleaewas significant (Table 1). There was no significant 副fferencein penetration ofM

incognitawhen M. incognita was inoculated 2 days or 4

days prior to or after the inoculation of P. cor[eae In experiment 2,inoculation with unequal numbers of the twosp叫 成 7 50P.coβ加 +250 M. incognita or 250 Rω>[[ea e

+

750 M.incogni的 ，simultaneously resulted in

Fig. 1.The Pratylenchus cofleae hadfi月er-liketail shape (的 an dMeloidogyne incognitahad tapering shape (B),

274 t古自物的經學 會 刊 第 1 7 谷 第 4 1的 2008

Table 1,Effect of simultaneous and sequential inoculation

with equal numbers of Pratylenchus co/feae (P) and

Me!oidogync incognita(主任)on penetration in mung bean

roots Treatment!

αe

P+M [9 58.3 P [75 加1 77.7 t-test 2dP+M 77 2[8 2吐P 244.7 M 90 t-test ns ns 2dM+P lOS [72 2dM 139.7 P 249 t• t e s t ns 4dP+M 98 207 4dP 220 孔f 116 t-test n5 n5 4dM+P 105 159 4dM 121 P 231 t-test ns

IP+M: simultaneous inoculatio口 ; 2dP+M: P. coj{eae

inoculated 2 days prior to M. incognita; 2吐 泌 + P :M

lllCOg叫fainoculated 2 days prior to P.coli:ωι 4dP+M: P

ωFωeinoculated 4 days prior to M.in印gnita:4d M+P:/vi

itlωgnitainoculated 4 dayspri肘to P. ω肘。r Two meanswit吋hi叩neachpa剖[叮rare SIg釘111缸C Ie叭、el a缸ccor吋吐i吐ding t岫o t卜一扭S泣z

Two means within each pair arc notsignificantly 丘ifferentat 5% level according to t-test

Table 2. Effect of simultaneous inoculation with unequal numbers ofPraf.vlenchus ω11;盯ze(P) and lvleloidog)'ne

lllCOg別的 (Ml an p叫1etrat的nin mung bean roots No,nematodes penetrated M. incognita P.colleae 9.6 143 229 Treatment 750P+250M 750P 250M 45.5 t心test 250P+750M 250P 750M 38 2J 35 『/ 吋/但一句 J 141.2 tHtest

?有omeanswithinωcllpairare 咿ificantlydifferent at 5% level according tot-test

significant suppression of penetration of both P. ω加ae

andlvI. incognita comparing to when theywe甜inoculated

alone (Table 2)

There were no significant difference between the penetration rates at inoculum levels of 250.500‘750、and 1000 nematodes foreither 后peciesof nematodes inoculated alone (data not shown)

The number of M. incognita developed into sausage shape in the simultaneous inoculation withP.cqffeaewas

sIgm日cantlylower than when M. incognita was inoculated alone (Table 3). However,because the penetration rate was lower in thes口nultaneousinoculation,the % development

oflvI. incognita (No. sausage-shaped juvenileslTotal no

nematodes penetrated) \vas not significantly different from that of thesin巴Ieν inoculation

The I口】叫u叫叮η叮前I缸jτ泊Z

S叫I草nt呵凶fica訂n叫1t叫tlylower in thes叫imultanηleou叫i拈s inocul拍已a叫i泣山t“101口1wIt廿血hi)甘f 111肌cogn叫it，的α t廿h】阻e臼n when}去'. coil，什Fαe w‘a站s mocu叫l‘扎叫!ted ‘a吐lone(Table 3). The total number of nematodes was also significantly lower in the simultaneous inoculation; there we出30.5% and 24.3 % reduction for eggs andnemat叫les ， 自spectively

The number of galls on the mung bean roots was significantly lower in the simultaneous inoculation than that in the single inoculation (Table 4),this was also true for the number of females and the number of egg masses

The simultaneous inoculation in soil resulted in significant reduction of the number of nematodes penetr正ltedthe mung bean roots for bothP.coffeaeand M

lJ7coglllfaco訂lparingto the respective single inoculation (Table 4)

DISCUSSION

P. co吋'If，戶e叫ιG肘t叩e a叩nc吐1 M . i刊川f川l C O gη川I叫t a were mt叫l t u叫.la叫11勾y

an叫tagom站st訂Ieonmu山mgbeanroo站Thq 已net叮rat山lionofbo叫th1

P.(叫01斤(eaeand i\1.I川n co句rμ11叫2

、w henthey were inoculated simultaneously comparing to when they were inoculated alone. Mutual antagonism between nematodes is one form ofcompe 位tion('I'.It has

also been found between other nematodes. Estores and Chen reported that P. pellerrans and M. incognlfα

depressed the population of each other in tomato O'Bannon,eraI, found that mixed inoculation of Rαdopholussimilisand P. cojfeae on citrus resulted in lower populations of each species than in separate inoculation

Competition betweenPratylenchusand 九1eloidogyl1 e 275

Although the use、of 盯owth且Douches for the experiments saves labor and space,it is an artificial environment for the nematodes. To verify the results obtained in growth pouches, P.cqffeae and lid,inζagnaa

were inoculate丘tomung bean seedlings grown in sands simultaneously and alone,Significant mutual antagonism

between the two species was found (Table 4). The resul臼 further confirmed the findiIH!:、4Sobtained in theνf2To \vth pouches

When there are enough penetration sites for all the nematodes encountering the roots simultaneous妙，mutual exclusion is not expected. Inoculation with 1000i11 incognita did not reduce the infection rate of M. incognita

comparingtoinoculation with 750,500 or 250 nematodes (data not shown),and similar results were also observed on P.C(~ffea e .These resultsin廿ieatedthat there was no shortage of infection sites for the nematodes inoculated in the experiments. Therefore,the reduction of penetration of

P. cqffeae andl'v1.incognita in the concomitant inoculation

was the l'ι、 ult sofcompeti訂onbetween the two species instead of lacking penetrations如Asto howP.c(~ffeae

and1\1.incognita interfered with each other when

inoculated simultaneously remained an interesting question. It is not easy to investigate the nematode activities on the surface of the roots without proper equipment.Itis common knowledge that animals fight fm food,The fightexist話evenbetween small organisms like 叮 i c ke t s .Nosil > r e p o r t e dthat housecricke脂 ，Acheta

domesticus,attacked each other for food by kicking,biting

(mandible lunge),head charges‘andwrestling. Whether microscopic animals like nematodes also attack each other physically on the surface of theroo站isunknown. It would be an interesting case of animal behavior if it could be confirmed with proper equipment

Competition of nematodes at penetration may not necessarily be linked to the subsequent development of nematodes,The data showed that the percentage ofM

incognitasecond心stagejuveniles developed into sausage

shape was not significantly different between the simultaneous inoculatioin and the single inoculation experiment (Table 3). Similarly,Diez,et01. found that the rate of 丘evelopmentofRotylenchu !I叮reniformiswas not affected by the presence ofM. incag川的 oncotton although they were capahle of inhibiting each other

24.3 214 299 430 76.6 81.1 6.3 21.0 24.2 85.1

Table3.Effect of simultaneous inoculation' ofMeloidogync incognita(M)and們的lench us c社ae(P) on the development of1\1.incogni旬 ，egg production and total number ofP 叫F叫eon mung bean roots

M. incognita P. ω加ae

% No. Total no. % %

Treatment

Sausage-shaped Penetration Development eggs nematodes Reduction Reduction of nematodes M+P

M P

t-test 和 牛ns'

1Eachspe臼eshad 500 nematodes

1Two means aresignificantly 吐ifferentat 5% level accordingtot-test,arcsine square root transformation wasp 叫formedfm percentagesbefo記statisticalanalysis

.' Two means are notsigni日cantlydifferent at 5% level accordingtot-test,after arcsine square root transformation

162.1 司長

In soil No. nematode penetrated

R c(~ffeoe M. incoglUta

67.0 24.5 89.9 14.5 38.8 No. galls 23.2 47.8 t-test

Eaeh species had 500 nematodes

Two means within each pair are significantly different at 5% level according tot 抬到

Treatment

Table4.The effect of simultaneous inoculation I withPro吵lenchμscojfeae (P) on Meloidogyne incognita(M)galling and reproduction in mung bean roots and the penetration of both nematodes on mung bean roots in soil.

In growth pouches

坐主必史!但…

No. females No. eggmasse 可

17.7 43.1 M+P

M P

276 植物f禹王申學 會 1 月第 l 7卷第 4 期 2008

initially. Melakeberhan and Dey also reported that competition betweenl-I.glycines and M. incognita had no

effect on nematode development.In the present studies, there were eggs ofP.colleae in the roots at 10 clays after

the concomitant inoculation (Table 3),indicating that reproduction ofP.cofleae occurred in the presence of!vi

incognita. The ratio of the total numbers ofnemato忌。sat

10 days after the concomitant inoculation to that in the single inoculation was higher than the ratio of those at 3 days after inoculation. This indicated that some eggs had developed into vermiform nematodes in the roots Although development ofP.coff削 wereobserved in the simultaneous inoculation,the percentage of development could not be calculated as forM. i月 c og n i ta(Table 3) because the initial inoculum was mixed developmental stag出of P. ι oJleae .

It is conceivable that when one species outnumbers the other soccics it would be of advantage in the competition. The data showed that inoculation simultaneously with unequal numbers of the two species, 750: 250,resulted in significant suppression of penetration of the lesser species comparing to when they were inoculated alone (Table 2). The present findings are coincident with the report of Diez. et01.(8) where

Ro!y/enchulus renz(ormis andM.incognita inoculated

simultaneously on cotton inhibited each other when the amount of inoculum of one species was higher than the athc!

Theoretically occupying the territory ahead of the other species should give advantage to the preinvading

speCies， 也肘 ，timing was considered important to the outcome of competition between nematodes. Jatala and Jensen{i4)reported that no significant population changes

were observed for eitherM.hapla or Heterodera schachtii

when they were inoculated simultaneously to sugar beets, but when H. se)wehti was inoculated 10 days earlier than

M. hap的 ，the size of galls was significantly reducecL In addition. Gay andBir吐 【foundthat prior invasion byM

incognita suppressedP.brachyurus populations on tomato

and that previous inoculation of P.bl α ιhyurusalso inhibited root penetration byIv!.incognita. Umesh,er al

reporte吐thatin experiments on competition betweenP

neg/ectu.l,>andM. chillvoodi in barley,the species that

parasitized the roots first inhibited penetration of the latter species. The present findings showed that 2 and 4 days

prior inoculation ofP ω鼠?aedid not influence the late coming!vI .ineog凹的 .but prior inoculation of i'\1.incognita

inhihited the penetration ofP.coffeae (Tablel). It appeared

that the competition between the two nematodes occurred mainly on the surface of the roots,once one species has entered the roots,there was no longer interference on the root surface for the other species to utilize the infection sites. Similarly,Estores and ChenI 圳 re p o rt e d thatP

penetrans andIV!.incognita inhibited each other in tomato

but there were no significant difference in the numbers 01

P. penetrans when theywer亡inoculated10 days prior to or

after introduction of M. incognita. In the case where priot inoculation of M.incog川的 inhib i tedthe penetration of P

coffeae in the present studies,other factors might have

been involvecL Estores and Chen (]{) used split-root tomato plants to demonstrate that a transmissiblesubst乳nce.

secreted by M. i肌 o g n i raorproduce吐by the host in response to infection,was effective in reducing the penetration ofP.penetrans. In the present study,the effect of the prior inoculation of iV!.incognita onP.eoj]lωc penetration was effective in 5 days (2 days pτror inoculation plus 3 days after inoculation ofP.a拼叫 which was faster than those reported by Estores and Chen 110,where it took 22 days(15 days prior inoculation plus 7 days after inoculation ofP.penerrans). Elucidating

the substance responsible的 theinhibition of P. coffeaeol

P.penerrans,no matter it was secretedby 九1 .incognita 01

produced by the host in reaction to infection,mightlea丘to a novel way of controlling the nematodes

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總要

祭碧雲 2008 . 們 的 len chus cojIeae 與lv1eloidogyne incognita 之間 的競爭 織將 會 flJ17 : 271-278

(台北約立台灣大學織物病理與微笑物學系， ，在子郵件:bieyntm@ntu.edu.tw;傳真:+886

2-2363-6490)

在綠豆很內 P. cqffeae 與M. incog川的 之 f尚 存 在著 互 相抑制的 紛 (系 。 !司 將按1重雨1這線路i時 P coff叩 e 及 jll1.incognitaI之 侵入 率 皆比 單 獨疫手主持為 低 。 !可 將J射裝 的 種絞盡益的 發 育成香腸 H!~之 M

lIIcogmta 三約幼蟲敏 因 比單獨接種II幸少 ， 個2.JJ處發育1~ 1幸嗨Hj~之比率與 單獨 重的煎熬顯著策

獎。(古'JII剎那垂不約等敷衍的兩種線蟲，無論其佔有比率為何其侵入率背上七道 JmH去海將為低。先

;妥種 P.cofieoe爾夫或四天的，主;Jlv1 ,incognita 之侵入率無影響 ， 伯先按1重AI. incognita 雨大紋

的夭峙，可降低 P. coffeae 之侵入率 。 此種影警 J 可能牽涉到 EHlvI.incognita 分泌而來或是 的諾 :在

對受感染所產生之反!您的j來之物質。深入探討此等可抑制 P. coff叩c 侵入之物質 ， !得iJJ nr勛章rr

的線鐵防治方法之發展。