STUDIES ON THE EFFEC OTF CERTAI COMBINATIONN S OF …ir.amu.ac.in/364/1/T 2324.pdfAn X. brassica Wa...
Transcript of STUDIES ON THE EFFEC OTF CERTAI COMBINATIONN S OF …ir.amu.ac.in/364/1/T 2324.pdfAn X. brassica Wa...
STUDIES ON THE EFFECT OF CERTAIN COMBINATIONS OF STUNL ROOT KNOT AND RENIFORM NEMATODES
ON CAULIFLOWER, BRINJAL AND TOMATO.
A B S T R A C T
THESIS SUBMITTED TO THE ALIG XRH MUSLIM UNIVERSITY ALIGARH
IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY
IN B O T A N Y
By
RASHID MASOOD KHAN
DEPARTMENT OF BOTANY ALIGARH MUSLIM UNIVERSITY,
ALIGARH Nov. 1981
A A ^ l X J U L O .
1 . t(5mato seedlings w«r» inoculated with MBloidoovng
im.l^anltA o r IWyj^JHshUUM. r e n l f o j ^ ^ with four d i f f e r e n t
populationst i t was observed t h a t with an i n c r e a s « i n
i n i t i a l pofxilation t h e r e v ras an Incraasa i n f i n a l fiopula-
t i o n of the t m nmnatodas. s i m i l a r increaso was observed
on eggplant. Although increase i n ,iOi^lation of
Tvlgnchoriivnchus bF^^^ICaft, was also observed on both
plants however i t was not as hiqh as i n the case of othf«r
two ntifi.1d(tO«l9£i •
2 . Jn caul i f lower , on the other hand, several fold Irwraase
An X. brass icae Was observed but not t h a t of renifprmis
or ii. iDfiSaolM*
3 . In concomitant inoculat ion of tomato and eggplant seedl ings ,
the copulation of both th® iwmatodes v i i . , ineoonlta and
renlfoHBlg was reduced. The reduction i n th® population
of foiraer was aore than the l a t e r , iteduction i n dry weight
of plants was as high as with the high inoculum l e v e l of
nematodes in s ingle inoculat ion t e a t .
4 . In H. reniforvAt and X* brassicaa combination on tomato and
eggplant the >opulation of the l a t e r was adversely affec ted
while that of former remained the same, un c a u l i f l o w a r , on
the other hand the population of X* brassleae remain unaffecte
and of Ji. reduced.
5 , In J , ^neoatilta and brasalcae combination, the pomila-
t i o n of both the nviiftatod^s was redoced on tomato arKi
egcjplant, whrtreas on cauliflowrfr the posxilation of
i * bras sic.aa. rt'jji.-dned unaffrfctei and t h a t of incognita
was
In sequential inoculat ion t e s t s of J . Incoantta
li* g^ralfgr'4„g.i g^nilagaiff, - I * kcaasuULai i * "»
I * ^nd J . ii* * I . toggle
an the population i n a l l th«i cjnbin.ttion was
re^ucad, Advyjrs© a f f e c t wegy laor© jronounccU on m^Eiatoci©
vdth wJiich th© aatrdlin^is ivare inoculated l a s t .
whan saudlings of taraato and Q jgplant qvtsun i n s o i l amaniud
with na&mt c a s t o r t yjraund-mit cakss , thimet and f e r t i l i z e r
and l a t j r inoculated vdth liyrofinita - H. ranifogmisi
ii« X* juedsMstiS.; ItOCOTnilfa - brassicao?
M* I n m a a i a ^ * ygr^fffiadg - I * teific,^ combinations
tha ix»puiation of a l l the threff nematodes was redticad i n
a l l the amencfcaants. rtdvarse ©ffact of o i l cakes was g r e a t e r
on than ii. j w n l f w l g o r I . farasal&aft* IHe dr^
weight of alants a lso increased i n coni Carlson t o plants
qxxMn in unm^nded s o i l .
8 . Cone omit ant inoculat ions adversely af fec ted the penetration
of larvas and iE»nature females of ineonnita and
g , rh» penetration of intnatura ftsmaXafi of M. rgnlfnrnils was
not .jrvsverjtsJ in tha jraaanct* uf brassicaa,ii
penatration of J . i n o q . i i t a was raducRd,
iO. in CQncoTTiitant inoculation t a s t of incarynita. R^reniformls
<an<J penetration of bol-h d* incognita ani
g 'iTifor^nis. was raJucad.
iX, In concomitant inocul j t ion of ^ ^ ii* r^nlfor^ls,
•n t a a a t o , the 3iza uf ft*? lala, the riu. ibur of fc idli? v«as
raiucv^d. Alia fdCundU.ty uf r>iiAforchis a lso naducsJ.
.vioreovor, tha charact j r i^i t ic seiiiilunar tippejrjnce c f qinjit
calisv;aQ not obiiarveJ. i»iight ahif t i n fajJing sit-:} WjS
also noticed. iiov/avi.?r, in the jire.onccj of brasuicae tha
fecundity, feeding s i t a .jnti charact j r i s t i c sw^dlunar
arranno'Tifjnts of aiant v as not. disturbed.
12, In inqootTit3 T, brassica^i cotnMnation, thr» dGVolopvisnt
of Of nit a W 3S ialdyeKi aa a number of fourth st jqe
larvae and evan mdlias ware observed,
13. When tht? tomato saedlirKis were inoculated with tha ;«ii^urG
of tha three nt^natovies, adverse affec t an the davalopment
of root-knot and rani form neiatod® was obsorvad. tia niforw
nematode failed t o produce char<3ct-rist lc qlant c a l l s .
There was a tendancy for production of giant c a l l s in
c o r t i c a l region.
>4
14, In concomitant inoculation, there was ci tc^ndifncy fov
oroJuction of mora male a , Hiqb«?st numb-jr of rn.ilf
noted whara o i l cakas wara employed,
15, iVhen eq p a s s e s of ir^ogjiita m-rfi alX&^ed to htitch in
.araaes-Ka of r>miforiii.5 / £,. brasslcaa, or in Wc^tar in
which e i ther of th ^ above neincrtoie has b«en sus.T^indsd f o r
tifCQnty four hours, l a r v a l hatch vv s consiierably reduced.
STUDIES Oiy THE EFFECT OF CERTAIN COMBINATIONS OF STUNT, ROOT-KNOT AND RENIFORM NEMATODES
ON CAULIFLOWER, BRINJAL AND TOMATO.
THESIS SUBMITTED TO THE ALIGARH MUSLIM UNIVERSITY ALIGARH
IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY
IN B O T A N Y
By
RASHID MASOOD KHAM
DEPARTMENT OF BOTANY ALIGARH MUSLIM UNIVERSITY,
ALIGARH Nov. 1981
/U-iiiMJW UNIVcHoITY, .U^GAiiH - 202001 ( I ) ,
Ja t«d I Nov. 1981,
This 3.S t a c e r t i f y that '''ir. ifashid ^ .isoO'J i fian fids
worked in t h i s cieaartm^nt as a U»se.srch .scholar undar ray
su pervision and i|uidanciJ. llis vvork on tho "studies on the
e f f e c t of Cf?rt3in c'^mbinations of s tunt , root -knot md r»ni-»
form n«wJdtod« on caulif lower irinjaii an i torn sto" i s u ito date?
anci a r i g i n a l . Hfs i s allowai t o suissilt h i s t h a a l s f a r ths
considaration of the awanJ of the Jagree of Joctor of
losoF^y •
I /
(aSaAiT M, KHHN/ (/<4inn.)
F, j F , a , o , X , ;F,N,A, Professor of 3otarry,
IhQ authojp i s highly indabted t o J ^ f e s s o r Abrar 4. i\han
(.>1inn» J, F.N.i*, and take tha opportunity t o express his
aincara thanks with doap sariua of qratitud® f o r his viqi lant
.iuidanca, construct iva suqciastions throughout th« coursa of
these invaatigations arKi th^ help rendered i n preparation of
tha manusc^^t without vMch i t would hav« not been possible,
liis love dnd affect ion has al\ways been a source of inspi ra t ion .
Jirwara thanks aire also due t o sJr. .4, H<ijid Khan v/hase
help and encoura^jHraent w&r@ highly invaluable.
Htj i s a l so deeply qra ts ful t o /^ofessor ^iaKana,
Head uf xh'i .isjii irt fl iint of .Jotany for h i s inborn Qanerosity, c r i -
t i c a l sugjestians and for providing laboratory f d C i l i t i 0 s f o r
carrying out the so studies .
si near© thanks are also da© t o J r . M.F.
J r , Xrshad '" ahmood. ^ultanul Haq, i^r. ;ihabbir Ashraf,
Mr. Zakiuddin, -^r. Htfhanur Hehman, Ar. Akhtar Heseeb and othf?r
colldfgues and friends including Mr. < >azi Javed Ahmad, the
ty>:>ist) Mr. lahulam ^ujtaba and tdr. Q.H. Ansarif the l i to togra^ers
for t h e i r help in coia J let ion of the radnuscript.
The finaJTOial assis tance arovidad by Council of ^ i a n t i f i c
and Industrial rtasearch i s also grataful ly acknowladged.
I . ZfO'iluJUGriON iiiiVX W iJF LXT^it/O'a.ia . . . . I
IX. i4aTHOJt» 20
{ i ^ Inoculation of tomato c v . marr^Xobe
29
( l i ) Inoculation of eiq al^nt c v . ;>usa purpla long swadiiJig with i^gloiaonvr^ inc oanlta. ^tvl^nehulua rafl^ftfCTli^ and TYl?nghQtV. rhvnchus bras s ic ae alone ani i n liff^^rertt comM nations. «* . * • * . . 37
( i i i ) Inoculation of cauliflo&^er c v . early snowijall seadlings with Aaloidoovna laMgiaytii* 'MvltnchtfJ^Wi r^ntforrBlff
*Yl«nch^irhYmfnU,a brassIcao alone and in different combinations 46
( ivJ c f f a c t of s inqla , concomitant and sequan-t i a l inoculations of eqqplant soedlings with iatY^l^nchy^tfg
fflaaia and tyA^ ichffighYnc^yff yxatiicfli
(v ) iiffact of sinqla and concomitant irKkculations on tc^ato 8a<»dlinqs
irown in amanJaJ s o i l 63
( v i ) ciffect of sin<iil9 and concoiaitant inoculations on tf^qplant s^adlings qrown in amondod soi 1 80
(v i i ) a f fac t of concomitant inoculation of toiaato S99JlinQS on ponotration 91
( v i i i ^ afftfct of concoiaitant Inoculation of eggplant seedlings on penetration •• 98
-c'aflO
(ix^ Histopathology of tomato m o t s rasultlng froffl %lnqlfi and concomitant i n f a c t i o n •• 104
( x ) ttff€ct of concomitant infec t ion on facundity of Incoqnilfi and •MyAftosnyAm •
( x l ) Hatching i l 3
IV. ij I a C u .J a i sj 119
M, ii a F u ii J iv U u a 129
i^lants roots are cunstantiy ax >033i t o blot ic as well
as dOiotic fac tors , i i a t i c f^jct ara inciuJa rayriads of dLcro-
org^nisms which irtfiabit s a i l , Th<?se raicroorriinisms (Jav«?lo >
various Kinis of intorraldt ianshi ^ amom tho isslves as ivtJll as
v/ith hlih fr slants. Thi y may be beaef ic ia l or deleterious,
/ccorjirh? t o . takman and Harrjr th« r'^'iatianshicomi
•ith«r fje c>coiaqical or syrj,bIotic. Tlje fijra-'r can '.?ith''-r be
n a u t r a i i s t i c , ant ibiot ic or ayn-^riiitic ind tho l a t t p r can -je
eith Jr neutral , nutuaii s t l c or .mt a 3 ini a t i c .
/Uthauuh ill Jilt Mr a s i t i c nfj';!.3ta'ias aiona ara ca ' ja la of
Cdusinq sev ra J issase syiaptoms, howavir, i t tha jrss. aca of
oth r;r -ivjii fiiicroorqani;:! na iaadga at time Of?coma« devest at iaq,
Funqal ni?':iato le int rac t ion v, ,3 re )ortei f a r tho f i r s t time by
Atkinson {ifa92j, in which he founi that cotton slants infected
by root~knot na.natoJe anhancecl the savari ty of fusari Jm wilt in
cot ton, iactf^ridl dnd niMUtodfs, virus and nanatodo, neniatode
and nenatoi® int>?rralation8hioa wero rs Hjrtad for th > f i r s t time
by (i9Ui.), Hawitt a l ( 1 9 5 8 ; , uhapmn itoss
(1959) res >ectivaly. Iha l i t rature on t h i s as loct of n«-iiatology
has bosjn reviewod by rJitchfar ( i963 , 1965, 1978;? ^owell (1963,
1971 a ,b, 1979) ; - t i l l er (1965 j, v.eischir ( i 9 6 8 ; , irozeski (1970 ; ,
3er:)6aon (1972 J, Hirano ( i ''75). ^^tch >r (1965) while reviewing
th© infoiraaition regarding the i n t e r a c t i o n batwei'n ularit l a r a s i t i c
narnatodes and othi?r jathoqans has suq,jested s i x diffsrerrt
iriQchanisias»
t : r l a l w i l t of jlarrts (Lucas J.954; Uixridn 1 9 6 4 ) .
gstablishfiiaTtt oncQ i n cant act with the for
iteUillM, (-^toinbuch, ±79 Uhitwood, 1935? i M s t i m J ^ ^ m
ialag At a nasof f , 1925) and of f^gyy llQbafft tay yr l^jSitiCl.
(l.'utchinsonj iurtiay ^ {Carne, i 9 2 6 j VjsuJeva Hi-i'V-arani
19521 Uu;jta ami ^ a r u i , 196B) and < tiij Ivji-ai ioi dea fra-iari aQ
(iUtztiuabosa) of f jsc^ans, C l i l f a r d j Jaws on
( i ^ t c h . r and Cross, 195^1 l t c h : ? r , 1 9 6 3 h
QstabHahment unless introduced balow the e ddairoia f o r
example Xi.Cinema iridex Cobb, 1913 f o r viii® farv l e a f virus
(Hewitt 1 9 5 8 ) , fffffho^Pr^S, Qhl^^^f 1957 f o r
tobacco r a t t l © v i r u s , (Walkinshaw g t , 1961) and Lonnidgrua
ft^ftiqatua 4«n (Harrison fll^i., 1961) for toraato black rinq virus.
( 4 ) .^ematod^s as providers of i n f e c t i o n necrt:>tic courts f o r
exam )1®, Pratvlenchus )Qnatrans (Coijb, 1913) F i l i >jav and Stekhoven
1941 i n root ro t of ^each ( iountain and i^atrick, 1959) and wil t
of egg plant (fountain and <4ckeen, 1 9 6 2 ) .
3
( 5 ) as qoc^f^ers of substrates t - f a r e x ^ p l a , root*
knot nora<3todt? and coaulex in cot ton (>1iriton and yintoa»
melavlQaft ia faaajLCQAfl oQf^ a a l a a l and
incognita (Kofaid and ^Mt®, i 9 i 9 j Chitwood, 1949
codiJlax in tQr7j,jtQ roots (.^olden and \faii GunJy, 1972), Ifeterodera
p,;ycl,n^A Xchinohe, 1962 and Pii . adim » >• GomAsx i n soybaan
(iioss, 1965) .
foroxarniie
i n root-Knot and F>js. gltaa. s;». complex i n ccjtton (At^dnsont iB 2 ) ,
; - 4 g s u a C -fcUjiioohi ja-is^ia jfiJ iSMlmshm
(ruihn, JUB57), FiU.^QV 1936 i n lidXt r^mlstant v a r i ^ y
of a l f a l f a (Haton and Hanna, 1967),
Th«3 KJ JUlatiqn dynafoics of namdtodas i s qavcrned bcrt.h
biot ic and abiot ic f a c t o r s (»*all«cei 1963* 19731 forton, 1 9 7 8 ) ,
ijut of the diff«rertt b i a t i c fac tors the role of )r9daci0U8
nematodes such as loncntshua a i, Bastiant 1866 j qinlQciaster sp,
^chultz* i857 j in reducing oojxilation of >1 j it p a r a s i t i c nefBatodas
has b«en wall estabUshad (Coba, 1918, 1920; >atain^r and Helnly,
1922* Thorn®, 1927) , asa&r (1963) , while working with predacious
nematod®» obs<»rv&d th .t c^^rtain s oeciea of plant :>arasitic
nematodes like Hoololaimus Jaday, 190Q;
3ft.l.an9JiflifB«a lonaioaudatua Hau, 1952; and QriLgftn^^ aflii.te, S
Taylor, 1931, wer<s r e s i s t a n t t o jredation while vteloidodaga
,Uftrt<^tnajLat chitwood, » aar»on and Esser , 1956? ^atvlenchus
agnftranff* xuiaus. Allan and Jensen, 1951 ; .^aratYifnghui
curvl ta tua v . d . Und, 1938 and .telolJonyne sp, ware s u s c a o t i b l e ,
Joosal is and .4ankau (^965) reported t h a t poriulation of Tbornia so ,
ijTcreasQd i n Jrasanca of Tvlanchulus samloa net rang Cobb, 1913,
without i l l e f f e c t on tha h o s t . They, howssver, f a i l e d t o
d^irmlna the ootimal conditions for predatian, -several spcjcies
of ^»ainura Fuchs, 193i hava been ra Jurted t o feed on /icihaJ^Qnehiis
tiaatian, 1965 j /4 anenchoidas r:)ari9tirei8 (tJastlan, 1865)
*it3in(ir, 1932; A'.Ayle^pchus s 3 . , J i tvlenchua J i a ^ ^ i and
flaterodara t r i f o l i i Uoffdzt, (l.achl.-3r, 19631 Hachlsr and Taylor ,
1966^,
<Uthough i t i s wall establishad t h a t i n the rhizosrtjere
of slants a v a r i e t y of plant Paras i t i c nenatodas are found vdth a
considr?rai3lQ ovarlaiping of ho3t rjiTfa (utostanbrink, 1 9 6 6 ) ,
h the as^ectii dealinti with t h a i r a c o l o g i c a l relationship^
rtJtiainad unaxalorad. C r e i i t t o i n t i t i a t e work on t h i s as >act
qoas t o Chaprnan ( 1 9 5 9 ) , who ra JortaJ t h a t on inoculat ion of
red c l o v a r and a l f a l f a root® with a mixture of oenetrana
and Tvlenchorhvnchus flUCyjai. Fieldinq, 1956, Jlant qrowth
remained almost tha same as on inoculatinq with e i t h e r of the
two alone. Howavdr, i n mixed inoculat ion the tx>3Ulation of
X* i^ ' rtini was reduced, .4alek and Jenkins (1964) studied t h a t
i n some combinations of MaloidQavtia hapla Ohitwood, 1949,
SMPr aXm ^nd T. p h j i s t l a resul ted i n decrease
i n the pojulation of n<jmatodes. .^illtsr and vdrheim (1968)
ra w r t a i th jt whan tobacco was inoculated with Heteroderf)
5
tafaacupi Lowns and Lcmhs, i.934; <^ratvlenchus aeriotfany and
Tvlanehorhvnchua s p , , t h a i n f e c t i o n aeil s u w i v a l of a l l the
t h r e e namatodas was r s Juced . i^rton (1969) found t h a t about
twice Xlsjiittetna aBericaniia Gobb^ 1913 spacimaTowers nresenft In
tha absence of t-^IoidotiynQ fta;?la than I n I t s irasance i n
a l f a l f a f i e l d s , iiikora ^ (^972) r.? ^ortai t h a t combination
of feloidlotivne fiaasi Franklin and T, on c r s a dnq b Jrit
grass resulted i n Irihibition of t h e po^julatian of the forner ,
ififhereas t h s popul j t i o n of aori. and -jenetrtjns. rersialnad
unaffected i n a l l tha c saibiii-jtlons. Toborjimanz (19T3)
stutiiad the oufiibimd a f f a c t at li-fyfltvlenchus rrdnvu^, and
Allen, X9S3, Tvl3iTChDghvnchua ^^bius (iJuliscJii, IST?) F i l l i jev,
1936, and lacroaasthonlA c u i v a t a an j^mitMs. carvo ^ l l u s
d ' alpimjB and obs irvcji t h a t jo j u l a t i o n of rnitwus was
raducad on i)oth plants i n tha presence of oofxilation
of d* on i n the presence of P. t^jlTYm*
on a . carvQahvllus and ^n f>r<»sence of both T.
and a^nyus. aird aJt J i i . (l >74) ro >ort®d that under f i e l d
conditions {jQfifJ.gj.fljif vta, CgiimBl>U& IflCaaniU s igf i i f i cant ly
i r t i ibi ted th® poiuldtion of conccaraitaot S;>5cie» on c o t t o n .
Population a.uild ua of H. indleug was retarded i n presence of
ineoanita on tomato r o o t s (Hasan and Alara, 1 9 7 5 ) , vtlllar
and 4elntyre (1975 ) , iSclntyre and ^ l l o r (1976) reported that
Tvl«fichQjFhvnehu« c lavtont i i ta iner , prevontad the entry of
^flitYdLfnghtta penetrans i n tobacco roots both in ^rior and
18
simultanaous inoouliitions. iiimllarly the ra iroduotion of
iavanlca (TreufaJ Chltwood, i 9 4 9 , Was inhibited on tomato
i n the presence of -^atvlenchus acrgbnagi. ut®in®r, 19451
I * f h r t s t i f l and Hemicvclioohora aiTgnayia Haski, i 9 5 8 (Van uundy
and iCrikpjtrick, i 9 7 5 ) . jRliochat Sk Ml* U ' M ) re )ort©d theft
mixture of aCiliim and iSMim, ^"hiirn© and
All<in» i950» caused g r o a t a r reduction i n qroswth than f i t h ^ r
alone and t h e .KJi^ilation of was reduced i n presence
of vulnus. i M l a and Aiejendro (1976) r e jorted t h a t i n
simultaneous inoculat ions of yan with |fe^ratvlanchus c o f f e a e
{iAmmt 1908) F i l i p . & u t e c k . , i94A» and ^utellonema, bravdis
( S t a i . ii t o Hew, i 9 3 3 i Andr.t i 9 5 8 , the J0:iul,iti0n of l a t t e r
was reduced and i l l e f f e c t on garosvth was a l s o not iced. Yang
e l (1976) reported an i t i t a r a c t i o n of J . i n e a a n i t a .
d. Jroi^oic^^udat^l and iJ.. aaleatua Gobb and absQXVBd t h a t f i n a l
pajTulation of incoonit i and aaleatus was r e luced i n
concoodtanit i n f e c t i o n , iianto and dolander (1977) re xjrted
that ^^rooosthonia Loof and Oe Urisse
decreased the areproduction of ji. ha^la on gra^e. The presence
of c r t g g n m a i t o . 9sm%m Haski, X958, HqalgAaUffly, t^iattgMi
and rvlenehorhynehua s o . reduced the s e v e r i t y of root-knot
on b r i n j a l (^ is ra and Uas, i 9 7 7 i j H. eoluiabua ^ e r , on soybean
(Fortnura and l.ewl8, 1978 h i , vulaarln U^iadhvava a l j l . , 1972,
on .^ffnnlLiltyiB t v iioides (Vaishnav and o e t h i , 1978)2 tey,iiffiff.aS
a i d d i q i , 1961, on tomato (Khan fil jyL»t 1 9 7 8 ) ; Par at r ich odorua
7
^ n o r and aarl on craeptnq iMint grass ( ^ k o r a ai , .»X979) .
Un the other hand, Incognita suppressed th® ix>pulatlon of
brachvurtan ( i i t a i n e r ) Andrag OH c o t t o n and eolumfautt su »p-.
rassed tha laojaaldtion of J , Inco^jtUta (ichmidt and Ji-evds, i 9 B i ) ,
The r e s u l t s iwrtainlng t o i n t e r a c t i o n of t n o endo*-
p a r a s l t e s are rc^viavsed belcmt
Hoss (i959» 1964) r e lortaJ th,it the jo-Hilation of
ineoani ta was sU!>prs8S9d by H, alvelrw^ on soybean «)her©as
E* l^irc'iely remain unaffaqtad ^tyofar^^tA but r o o t -
knot najsidtodo vxa^ulation c u r t a i l e d t h a •irodaction o f c y s t
noraetodo i n ©arly i>a3 t of 8€?ason, hossevsr^in the l a t e r i>art
of season the jopulatlon of cyst namatode t^as highar i n ,:)lots
containing ooth naaiataies than c y s t mmiatoda a Ion®, fiuerout
(1968) reported t h a t i n new ^Jlantation of :3in©-appl®,
Insftq^uirfl dominant over faraBhYmaUg, C^^odfrey,
1929) F i l i o j a v and ;itafc. 1941, while i n old plantation
braehvurus was dominant, f i l l e r (197Q) while studyinq the
i n t e r a c t i o n of H^-^^gy^^ga ii* .agflgtxina, o" tobacco
observed that at l U r a t i o , the r^opulation of former increased
only at moderate r a t e and at f a s t e r r a t e v4ien i t s fKipulation
was high. In the former case the pO!:HJlation of penetrans
was reduced and i n the l a t t e r case aenetrans was comoletely
eliminated, /«iiosu and Taylor (1974) r a x i r t e d t h a t at moderate
Inoculum iovsl of p»ngtrans and 4 , Incognita on rad c lovar ,
th© fX»puiation of pangtrdins was r^Jucsd, hotoevsr» at hinher
inoculum density of ijenirtr^i^^ tail f irmation was'ra ducad.
uatoras ( i 9 7 1 j , i^storas and a>en (1972) stuiied tho Interact ion
ijetwean Qg^chvurus and 4 , incQordt.^ on toraa?t:o arid obsarvad
Inhiiiition In ra >roducti jn of aoth iiGoidtodas, Taj rmy and
Cha raan (1972) observed the riiduction i n ;3Qnetration of
ii* iqcQinit^ m r«d c lover an-l a l f a l f a i n ths aros -.ica of hiqti
i30,3Ulation of penetrans. Chapman afvd Turner 1979)
afiiixjrted thiit tha iiresanc® of ha on red clovisr advarssly
aff£Jcti;J t h e fticundity of jjgjJElXstia* Frucb-iun and Clid-mn
(1972)^ on tho othv^r. hanJ^oasawad t h a t v4-ten rad c lover ^ as
inoculatad v^lth i.?Q/' atrans and H* the invasiveniss
of aith r r@i?5air^naffacted, (lay and iiird (1973) reaorted
that prior inoculation of tofodto vdth ineoqinit}^ su ^pressed
ii» brachvurua, hc»^9Vart no such i l l e f f e c t was notlc«d on
a l f a l f a and tobacco. Weischar (1974) re >orted that on tobacco
leaves th® population of d^t^aei was reduced in the aras^nce
of A.jhalenchQidgs ritzemabosi (ochwartz) ^tainor, 1932, howover^
on steam and p9tiol« ths population of rltzmnabQai v.-na
enhanced where i t does not develop normally, J a t a l a and
Jensen (1972, 1976) inoculated ji, vulmarls vdth a mixture of
haSiXJL and Heterodera schaeht^ .*chsildt, 1871, and observed
that when l a t e r preceded h^pl^. a marked reduction in galling
occurred! when foiraer preceded the l a t s r a five fold increase
n
i n c y s t fonaation occurred, hmvavert on inoculating aimulta-
naousiy no s i g M f i c a n t chango i n xipulation of s l t h r of the
namatodn was recorded, ^arraa and i^atbi (1976) noted t h a t i n
cancomitanft inocula t ion of coirfjea with Hotanadgra calan;! Coshyt
1967 and incocini t^. the sjopulation of both tha iwmataJgs was
auripressed* »jdLiidlar r e s u l t s war© obtainad by .J*i3annon e t (bI.
( i 9 7 6 ) on inocuXu^inci ciiarua with a tnixtura of iiadoaholu^
(Cobij, 1893) l-b.jrm, 1949 and .-^drt^vl^riehus coffga^y.
ujdff in (1976) th :t simultaneous inocula t ion of
"i^vada", i^ith g , d^psa^l and .4, ha jla^ f a i l e d t o break the
a!^sista}Tc@ of ^"rs-or inocula t ion of " / s m a l " aead-
lings with ^J, tH ysacL increased the qal l ing of haala and
pr ior inoculat ion of "Lahonton" with imsr^ased i t s
susce J t i b i l i t y t o djlosacf. ^imjh ( 1 9 7 6 ) observed th.it in
inixad i n f a c t l o n of Hotvlenchulus yanifoisnia Lin. u l i v . , 1940
and ^iQidoovno jnsgflnitfl on soybean th® popul^ion of th® former
was mora adv«rsaly -affected, Th© po Julatlon of H* raniforgda
ndl Tvl®nchulua aeml x»netrana was reducffd i n aiixdd i n f e s t a t i o n s
on gra^e vin® (Taha and aul tan , 1 9 7 7 ) . Khair and Osman (1977)
reported th at i n mixed i n f e c t i o n of H* renlfarmis and
M. incognita on tomato, the |)opulation of 4 , incognita
declined* The population of both J . incogni ta and g , sgjaiXoBBiA
was reduced i n mixed i n f e c t i o n on tcwiato (Khan flX,, 1979 )•
Martin (1978) re jortod th it root-knot nematode reduced the
1 0
3opulation of ir*rdtvlenehus oonatrang on potato , Balakhina
r&poxt&d an in ters ;>ac i f i c antagonism b©tw«sn Aohglenchua
M S i m , ^astian 1B63 and Arjielanchoid^a 3rforonhil^« F r a i ^ U n ,
i957 on wheat, whsre aatiroohllus v as th® dcjiainant s p e c i e s .
t i r i f f in and J d r l i n g (1964) suggested t h a t apiea^jcanua
h ^ a suppmsslng e f f e c t on Grlconemoldes xenQolex iUskl , i'352
i n ornamental spruca nursaxiea . Johnson and i^sbam (1^68)
rejjozted t h a t population of X* clarytonl was r e iuced i n th®
presence of e h r i s t i e on a l l v a r i e t i e s of corn exce pt "Tahua
Ha:ii" and "iiyn** wh-?reas t h a t of i . c h r i s t i ^ on ^Tehua Hop!'*.
Johnson (1-169, 1970) s t u d i e J the intciract ion of C^CQngmo^dQa
mm}m» a a i l l D l * A* on s i x
v a z i e t i e s of bent grass and noted a jre iuction i n f i n a l po-au-
l a t l o n of a l l the thz®® nticiatadas and out of thefs the io>uiatlan
of lonateaudatus was l e a s t a f f e c t o d , Hof ir'eduction of both
the noniatoda was r^ducad i n concaraitant inoculat ion of
jYA^fflihqghYnchtfg mLiua. i9S5 and M?UcflWknfihtfff,
leic^eohaiua on wheat (Smolik, H. aa laatus i n ^sresanca
of I . lonoieaudatus on cot ton (Yang , fil d i . , 1976)1
^agflt^Yltnchua, wrolffgl^Mi i n px«sance of figlggnwBgiafi ilffliUa.
(Cobb, 1918) Chitwood 1949 and HffUcatYl^nghtfg ggyy^Qgfffaua^Mg
(iAc Owaloy and Chapman, 1976)? i . . c lavtoni S t e i n a r 1957 and
U* ewtn^ttm Dass 1966 on r i c a (i^raaad and Rao, 1 9 7 7 ) and
X* dubiug fhome 1949 suopressad prgnatus on f i v e v a r l a t i a s
of tomato (Krishnappa and ^ a s a d , 1 9 7 9 ) .
1 1
Thus f a r limited araount of worfc has baen carr ied out on
tha i n t e r a c t i o n bo tweon diffarant specins of same genua, ttnjt
and ;»trlch Harajcl t^postsd an irrtragtjmjric dotainanc® of
one sfmci@s of MX9tt49Wm, w « r another. i^n^tgan?
tended t o reduce th® population of alleni^ on soybean (F&rrls
^ Al ' t 1967) . Chapman (1966) reported th^jt ndxed i n f e c t i o n
of id* inegonita, and h^ola on tOTato, 9Q» fafnalas of
ineoaMtai matured at modarata tsmpsratui®, howevert at
m l a t l v e l y low tsmKratura females of Ji* i^ncoanlt.ai taaturad,
Kinloch and /illsn (1972) also rerjorted the pradOTinanc® of
i . iMa^^SJ 0V -3P 4 . liaaM* /iccorviing t o s L Ml* (l 'J76)
nsi thar J . f necKinita nor J . lavanica caused reduction i n qrosifth
of tobacco c u l t i v a r ®i*J 95** and "Hicks**, howevar, mixed i n f e c -
t ion brought abtiut s ignif icant reduction i n plant s i z e , Sinqls
and jo int i n f e c t i o n , on th@ oth r hand, stimulates tha grsjwth
of "aurlay tobacco KY 9 " .
J a t a i a and Jansan (1976) whan inoculated 3<ita vulaar is
with Sa) larva« of i . ha jla or 500 larvae of Sfitlightli
in ona treatment and in another SOU larvae at two int rvals of
a i t h a r of the nasuatode, the s e l f int^ rac t ion resulted in an
increase in the ^Kipulation of both nematodes.
There are also instances where coinhabitant i n f e c t i o n ,
had resulted in an im:rease in the population of one or the
other nematodes. 3ird and Jonkins (1964) reported that vrfien
12
crane berry roots v\ara incx:uiated simuitanaously with
M I U Thorns, X95S and pgiggHTOQiafS,
thor^ Was an incraaso i n oopulation of t h e foimar, Johnson and
^ lusba»Jm ( i 9 6 e j ra xirted thart poijulation of ^ratvlenchua z«ae
was g r a a t a r on some corn v a r i e t i e s i n coaMnation with
rrlchodorua chr^st^^ than aXon«. Jut population of X«
increased on a l l var i t J t ias excast "Tahua Moiii", ^^cordinq t o
Johnson and (1970J tnixsd inocula t ion of ^ incognita,
and ii. ftrt^hvurus r«sultad i n siiqnificant incri?ase in the
population of the l a t t e r on r u s i s t a n t tobacco v a r i e t y , aikora
£ t (1972) rojortt^i t h a t tha xjpulation of i* iignatr-ans, was
incraasad i n preaerKJO of J , y^as^ on grass• J i m i l a r
iiKJraasa i n the population of brdchvurus Vi<a3 obtained on
c o t t a n whf n i t v- fas inoculated e i t h a r v^ith iiTCtxinita or
arenaria (uay and i i r i , 1 9 7 3 T o b a r j i r a n a z <1973) re lortad
t h a t on aloinu^i the raproJuction of ^acroaasthonia c u r v a t a .
^aaki, 1952 was t?nhanca»d i n pr«sffnce of mlnyua. Yam a l * t
(1976 j reported t h a t i n coixoraitant i n f e c t i o n of H. aalaatua. (3nd
A* lonolcaudatus. the population of l a t t e r incraased , oanto and
iolandar (1977) re jortad that oofsulation of Macroijosthonia xenoalex
increased i n presanca of ii . haala on grape, Valshnav and iiethi
(1978) observaJ an increase i n f i n a l population of Tvlaneh ir^vfichus
vulaarla in prasance of M* JLncgflnild ^n tXShflliiflLI..
^hmidt and Lawis ( I 9 8 i ) ra!:K>rted that t h e population of
Jeutallonama brachvurun ( ^ t a i n e r ) Andrassy, increased i n the
presence of U* columbua on c o t t o n .
1 . 1
aome useful informations have b«@n otrtained by a nwber
of wozkers i n complexos involving throe nematode s p e c i e s .
Johnson and :\i&beium U96G) observed t h a t tho presence af
I . fihrtst^a and H. f^l.tot^ftXA on soybean reduced th» iwpulation
of brachvurug. howevaXt on c e r t a i n v a r i a t i o s the population
ii* <4^hvst«ra was reduced. Johnson and iiisbaura (1970)
worked vdth thrac» species corabinatian v i z . t incoardt^ .
«3(nd P. ^y^cht^urufi on four vari<?ti0S of tobacco and
observed t h a t popylotion of brachvurus was do;3r»ss0d i n
thr»Q species combination on su»c®jjtibls v a r i e t i e s l ike •^fTlcks"
and 2326" and also on r e s i s t a n t v a r i e t y "iC 9 5 " . iimosu and
Taylor (1974) r«s)ortad an i n t e r a c t i o n bwtwean M. haol,^^
o«n«trans and J,, ^ori on r a i c l o v e r and found t h a t po!3ula-»
t i o n of ii. pgnatrans and X* agri was s i g n i f i c a n t l y reduced i n
thr»® spftcias combination, Th« dry weight of t o o and roots
was also rsduci^^J s i i r d f l c a n t l y , Toberjimnax S l a l « * ( l 9 7 7 )
worked with a combintftion of Pratvlenchua s p . , i^aratvlgnchus s p . ,
lYlfnBhOghYfWhtff 8tfACdtU8» s p . , Hgtarodey^ s p . ,
grt^ffMPfl ap. and Hgrnicvelioai^qy^ sp. on ixmiSL tetwtrum. Hprtttya
m s U i m * i * miimuLt and franKana cqgygfrpfj ^^d found t h a t
M s f f n t a a and Hf^fivcUgph^ra dlsaaoear^d from s o i l .
Tho isiportane* of nomaticidas and organic amanctoents f o r
th« control of plant p a r a s i t i c nematodas need not t o b«
ami^asixed. The l i t e r a t u r e pt^rtaininy t o th<9se aspects has
1 4
been reviewttd isy Oostantirink, (1960) ; oingh and ^Itaramaidh
(1970, 1973)J Khan ( l 9 7 l ) and Van aerkura and Hoestra (1979) .
iJropkin ( i95S , 1969)» Fielding(l959 J» ^ir thorst
(1961a) Krusberq (1963 ) , fountain (I96&)t (1974, 1979)»
iando (1971, 1975) , Baldwin (1978) hav» revlevswd the Utaratur©
on host sarasit© relat ionship of diffor©nt plant nar»isitic
nraatodos.
incoanita i s an andopara»it» and th® vahole 11 f® eye la
exca i t r'r®r>«wa8itic s®cond stag® larv<i» I s ccxaplated within
r o o t , a^ond stage noruially ontar i n host t i s s u e through
apical L'i »risteia (ifeaisc, 1910) . 3ut i t a l so mvjy antor throuqh
more mature t i s s u a (Chris t ie , 1 9 3 6 ) . Th© larvae move idthin
c o r t i c a l tissuQ both i n t r a and i n t e r c s l l u l a r l y felthout causing
any daauga t o root t i s s u e s (Chris t ie , 1936f Uroiskin and Jalson,
1960) . The c h a r a c t e r i s t i c feature of t h i s nematode i s the f o r -
mation of ^iant c « l i or syncytia near th« head rc»gion. vJropkin
and Joone (1966) sugsestei that telaiio^ivne s h a s two dis t inc t
s f f a c t on hos t : ( a ) syncytia develop f raa the c a l l s Which ar« in
incor;7orat9d as a r^^sult of c o l l wall dissolut ion! (b) c o l l s
of Mrlcycls show hypertrophy and hyperplasia, stimulus for
hypertrophy and hyperplasia reaches several hundred c e l l s in
the v i c i n i t y of nemitode head and thus g a l l formation occurs
1 r-
withln 24 hours a f t a r penetration of iarvao. with reqard t o
fcMcaaiiion of syncytiina Uiff^srsnt opinions have i»een forwardod.
^ n c y t i s davsiop usuaiiy i n th«? xylaas paxemhyma mav th» h^ad
region of famale Xtirvaa e i ther due t o dissoiution of c e l l la/all
(Kostoff and isUindall» i930j ( b r i s t l e , XV36; usvsrn and ^petoh,
19(A) i or du« t o snlarg^iaont of sinqlia c e l l {Ffuaoq and laqqentl,
i.96 Uj or both J isaoiut ion of c e l l wail an-i Qnlara<9m 'it
of sinrjlQ c a l l ( i i r i , 1972, 1973 ) . ^idJiqui I t (X974)
pMla tjorldnq with ^ nc oanita . on ^aenar^a la uc ant ha
m iort«d th'^t n^ iaatad© a f t e r gaintno «intry i n t o r o o t s , solect.ed
faedin'^ nita in fihloansi. .iianrt c « l l s havlrvj 2-.JL5 nuclei*
Clustar of ilant c a l l s surroundttd by a larga mjmbr»r of abnormal
xylmti» Tha elamerrts of reaction xylam .3oss»ssed no defini te
shapt? and sizo thay ware iianerally narrower and smallfiir in
dimansion as cociparad t ; normal on^aa. itoy (1976^ rsportai^^*'^
g g ^ n i c o l ^ f0«?ds on s t s l a r rat ions of roots of r i c e plant ,
iiiant c e l l foraation occurred in vascular parenchyma, ^ o l i -
feratad abnormal xylam elements; also present. Khan and
Tiaqi ( I977i studied th® isolated giant c a l l s and reported that
qiant c a l l s ware having the appendages of various s i s e . Nuclei
are usually crowded i n th® haustoria liko a )^}enda9es,
ah at nag ar ijJi. i l l . (1979) also reported thft giant c e l l s having
haustoria liko appendages, .^jrfaal xylem was c y l i n d r i c a l and
wider having jsits arranged i n r e t i c u l a t e pattern, wh«)reas
aOnojRBal xylum vessel was rounded or c h i s e l shaped.
I G
i>tudi»s on abnormal histology caused by ranlfornda
rsvaaled that fomale remain partly tmbedded In the r o o t ,
Unford and ul lv^ira (X940> observed that the feiaales were
part ly or ant i ra ly embedded in the cortex of cowpea r o o t ,
Peacock (1956) observed only necrosis around the nematode head
and h y p e r t r o j ^ o r break dcn /n of the c e l l s was not observe
3irchfi®ld (1962 j found no evidence of fe -iiir^g on eoldermal o r
c o r t i c a l t i s s u e but obs»rv0«i the females feeding on pholem
c e l l s , ^ n c e thare are number of publications on a v a r i e t y
of hosts where i t has been firmly established that faedinq s i t e
of renifoxsi n?»matode i s not cor tex but eith ar eadodeirais,
p j r icycle or phloeta. Further t h i s naraatode was capable of
causing of h y p e r t r o g i a n t c e l l s contairing e i t h e r on* or
more than one nuclei (Nath ^ 1969» 19761 Ha bios ©t
1970, i97Sj H r c h f l « l d , 1972j Oteifa and oalem, 1972| Jivakuniar
and ^ s h a d r i , 19721 Cohn, 1973, 19761 Heald, , 1975* Hazak
and svans, 1976f Jon®» and Jropkin, 1975| Yik and Birchfield,
1979 i* chain of events has claimed for another
species of ^ W l f UghuiUSt, namely fflflgr9^9ffat,18 (Coh 1976,
Cohn and /4ordechai, 1977) , In most of the cases i t has been
observed that uniser ia te j s r i c y c l e layer adjacent t o the erido-
ddmis responded t o infec t ion and became hypertroF^iied curved
sheet comprising 7 - 1 3 c e l l s ,
Fmdlni laechanlsai of different s >ecies of Tvltfnchorhvnchus
has b9®n studied. Heynolds and tivans (1953) reported that
1 7
dubiu^ feeds on eodLd irtBal t l s s u a , tfrfioreas Krusii rg (1959)
observed t h a t X, c lavtoni fed on a si dermal c e l l i n the region
of root <»lonqation and root h a i r s and namatodes were found t o
ba CQiapXetely eml>€»ddad i n root cap c e l l s and w&rQ not prdsont
i n o lder r o o t s . Similar claims hav© baen mad© by ivyss ( 1 9 7 3 ) ,
iaiddiqui ( 1 9 6 9 ) observad t h a t by and lartj® X,* byassicae, f«d on
epidarmal t i s s u e but i n cor t tdn cases neojatodes were observed
in c o r t i c a l t i s s u e and a t e l a r region .
Thus f a r considerable work on feedinc} and resul t ing
changes on host t i s s u e by indivi Jual nfcsmatodea has be-m c a r r i e d
out , Howavar, information on changes resul t ing from concOTfdtant
i n f e c t i o n are very meagre.
.^ankau and Unford ( i 9 6 0 ) obs^rvad the formation of
c h a r a c t e r i s t i c patholoqical t i s s u e of ii. t y i f o l i j . and ^aala
on red c l o v e r even i n concomitant i n f e c t i o n . tJteifa and Qsman
(1974) observed t h a t when ii. renifornds and J , incognita ware
present at the sdme i n f e c t i o n s i t e , the development of l a t t e r
Was inhibi ted by former, according t o J a t a l a and Jensen ( 1 9 7 6 ) ,
inoculat ion of M f l tUl^flrtS with a mixture of i , haola and
H, achacht i i i t was observed t h a t when feedirej s i t e of two
species was c l o s e enough each nematode produced i t s d i s t i n c t i v e
and char acta r i s t i c changes i n the i n f e c t e d t i s s u e . S imilar
claims were made by Taha and Kassab (1979) when they inoculated
X l a M aintnsl^ with a* yer^foCTli and locflaDili..
1 8
From th« foregoing i t i s c l e a r t h a t som® useful work
has bean c a r r i e d out on changes occurring in namatode pornilation
r»3iulting frtwj inoculation of two or wore nematoda, hommver,
thus f a r no c l o a r picture has eiattrged as t o how thsss assoc ia -
t ions influeiice tha population. Further very l i t t l e work has
baen Carried out i n India and oth^r v/arniar regions of the world,
Thar^fore, a t t w ; i t s have bean mado t o study th& foXloiwing!
I . Inoculation of tomato c v . niarglobei cfgq slant c v . pass
purpl.0 long and cauliflow^^r c v . a-irly snowball with |.|icoQnita.
d* y^^n^formis. tind i . or.^ssicag singly and i n combination as
given In Table'a*and t o doterifdnQ chaJiges i n jjopulatian, djcy
weightt s&x r a t i o and root-knot index.
XX. CksncoRdtarit and aoquential inoculat ion of agqolarrt c v . riusa
fjurple lonq se»alings with E . r&nifonai^ and
braatieatt as givan in table 'b 'and t o d«terfflin« changes i n
population, dry weight, s«x r a t i o and root-knot indox.
111 . Inoculation of tomato c v . mdirglobe seadlings grown i n
una»flended s o i l , and s o i l <^and«d with inorganic f o r t i l i z a r * ne»8it
c a s t o r ground-nut cakes and thimat with the thr«»o nematodas
alono and i n combination as giv«n i n t a b l e ' c ' a n d t o detamdne
changos in population, dry weight, sex r a t i o and root-knot index.
XV, Inoculation of aggplant cv , pusa purple long seedlings
grown in unamended s o i l and s o i l amended with inorg<^nie f e r t i l i z e r ,
1 9
ne^ra* c a s t o r , grounii-rwt and thiraot vdth the three
n^atodas alone and i n conblnation as given i n table c and t o
det<ax(nine changes i n population, drjr weight, sex r a t i o and
root-knot index.
V« Inoculation of tomato c v . niarglot3e seedlings grotsrn i n
wicropots with the neciatoies ilon® aiid i n combinations as
yiven i n t.iblQ'd'ferU to d&t^mlm the a f f e c t on nematode pene-
t r a t i o n .
VI, Incwulation of eggalant c v . i*isa Hinla long set*dlinqs
qrown i n micirtj 'ots ^ith the thra? n^matoUas alone and in cori-
iairiationsas qiv#ii i n table 'd 'and t o dstwrmints the offec t on
OfBaatod© janatr^ition,
VII . Mstoaatholoqical chaaffas result ing i n tottjato roots by
inocul itinq tham with ^ ineoanlta., Ji, renifowai^ and
X* braasicae singly and i n combination.
VIII , Effect of concomitant infliction on fecundity of
Ji* and yenlffttUui*
IX . ftffect of presence off g , iranifgmii brassle^a alone
and t h « i r mixture on hatching of incogni ta .
2 0
Thraa Rumstocies v i a . , laloiioavnQ 1 tic.k^riita (iCofoii and
white Chitwood, 1949, f^ptvltir^cf^u lu^ y^r^form!s Unford
and uliVQira JL94U and I v l a rysh orhv sichu^ bras si ca^ a l J Jioui i '6JL
v.-ere selected as t»28t nematodes and t h r e s va-^atable cr-a JS v i z , 9
timato to^PV^IsliPMa Cv. larqloba)! eq mlant
(^la.nura CTQlonggn^ U n n . Ov, i'usa .)ur:7le long) and c a u ' i f lower
(JCJaM&i SL^MS&m v a r . tiotryt^,^ U n a . c v . ^ o w b a l l ) used
<i5i t e s t plant?.
The iaticuiuiu of th« aisava thvoa neinatodss I'Jtis rvdaed
as jlvan b£?lowJ
J . j-ncjaiilt^t a s ingia aggmass af J . obtained fran
i n f «cteJ r a o t s vvas aurf aco a t a r i l i z e d i n Xii^J solution ^ t r h l -
§aw f ive minutt^s and washed t h r i c a in s t j r iUzed d i s t i l l -
ed water. The ogg mass was thsn allowed t o hatch in s t e r i l i z s d
d i s t i l l e d vvator. Tomato seedlings raised in autoclavsd s o i l
were inoculated vdth larva® thus obtainod.
renifortaiat imnjature fc' ialea of roiiiformis ware obtained
from the haavily infested f i e l d . Later the seedlings of tomato
plant (jrown i n autoclaved s a i l were inoculatad f o r maintaining
the cul ture of r^niforrn neraatode.
2 1
X « bPaasieaf < f o r ra is ing the pur® GUltmrs^ tha 8Q 3>dHinc|8 o f
Cauliflower wore inoculated with a sing la qravid fa.nala.
Throughout the studies tha progeny of c u l t u r e s maiotainffd
on tixnato jnd cau.llflowar i n jats x ian used,
^^a^atadas from s o i l wax^ i s o l a t a d Ijy using iost.^isbrink's
e l u t x i d t a r and i3aejcraann*8 funnel ami it tm roattsj ay warinqi
ijlendsr (atfjmarding, J . Uountin'. was Jan© with the of
Couitting dish (^uth3y» 1 9 7 0 K
The )i3n af if.'ork daal ini with concmdtdfit inocula t ion
s tudies o f t h e threo nanestoies on throo host plants i s givon
as ^Qlmti
Table i
I" inc'i'giTit.a li. K8i3L£azBidLa. L ' ^raaalcai .
100
- -
900 •
lOOO -
100 -
- 3 0 0 «M»
- 900 -
1000
2 2
Tabloe (ContdJ
.Jumaar of a jQciraans inoculafted
4, incoanita a . r^nifopTiis^ bras s i c ae
X90 « - 50-)
m» •• 9QQ
m iODO
mj iDO -
ioo 901 ••
5UJ 50 )
9QJ ioa LOO 9 0 )
- 5CX> 500
90i.J mt JUX3
IDO m TO
500 • I B 500
33 3 333 333
For studying tho i f fec t of sanaentlal Inoculation t e s t
on the nematode pojxjlation, of two or thraa nematodes the
following scheme was oinJloyadl
2 3
Tablo
Inoculation Ajiaber of s jejcitaens inoculaited ;^multaneou8/
days prior M* i i g w o l t a 11* ygnlifWills ! •
JLOJO
Kxn
^muXt.ineous
i5 days r^rior
i ^ t . 15 days pr ior
500
SOQ
500
5QJ
500
SOP
^imult sneoua
Hot, 15 dijys prior
Tyl . 15 days prior
500
501
500
500
500
50T
oirauXatn^ous
Mai, 15 days pr ior
Tyl. 15 dfiys prior
5 0 1
500
500
500
500
50^1
^raultan^ous 333
Mel, 15 days prior 333
Hot. 15 days prior 333
Tyl. 15 days p r i o r 333
Mai & Hot. 15 days prior 333
« o t . & Tyl .15 days prior 333
Mol, & Tyl ,15 days prior 333
333
333
333
333
333
333
333
333
333
333
333
333
333
333
M«i, » ifTgoanilii; Rot. m g , ranifomdai Tyl. a X>
2 4
^eaJllngs of th@ t e s t aXants wez^ raised in autocXavtd
s o i l t ransferrad t o 6" c l a y jjots, Ttvanty four hours aft r trans-
plariiatdon 2<-3* deaa holas we a made around the roots of oach
seedlings. Tha calouXat»d mimbrir of namatodles aith alone or
i n coraiJination were pAppoted i n t o holas iucj around the r o o t s .
'Latcjr tha holes ware plugnad and seedlings sparsely wat??red.
Thercj wora f ive r e p l i c a t e s f o r each treatment . The exaarinjents
war© terrainated days a f t a r inoculat ion, i'lants vmv& c a r e -
ful ly uprooted« tho roots ware washed in slosn? runrdng
The excess of water was r ^niovad Ijy placinq tha Jlants on
biottin^3 pap'^r. Tho dxy ivoight of :?XarTts w-.is deti^rminad i n
rj. t^ata on root-knot indox of J . ^ncoanit^ v#a3 rc3cor«i@d
according t o seal© iro losad by Khan ( i 9 7 i ) , /fematodes
froni s o i l wer« i so la ted lay enjplcjying uostenbrink © l u t r i a t o r
and daarraann funnal and numiiar of each sp»clf?s recorded par
kg s o i l , ^'opulation from roots was dotaxniinfiid by macarl atlng
the whole root syatara in waring blender (stomerding, X96a) .
Coyntinfj was done with the halp of counting dish (Southeyt
X970).
For datezQiining the a f fec t of concoiaitant inoculation
on nemaftoda multiplicat ion whan seadlimjs waro raised in s o i l
amandod with neani* c a s t o r , ground-nut cake^at the r a t a of i gm
nitrogan par kg s o i l , inor}anie f e r t i l i z e r and thlmat ^ . 0 2 gm
activa ingradiants par kg s o i l } . The oiX cakes were added X5
2 5
days p r i o r t o inoculat ion , and inorganic f e r t i l i z e r and thimat
were added ono day p r i o r t o inoqulat ion, Tha schoiae of inocu-
l a t i o n i s cjiVGn as bsiofe?;
Table c <
i-imiber of s,)ecimam» io jcuZateJ
I * teas,fig,ag,
5CX) JDiXi
5C).)
500 i o a j
500 333 1000
iOiJ 5CK3 5Qv)
IXXX)
SOJ
333
333 l o a j
333
5 0 1 lOQl
50^ mm
500 1000
5 0 1
333 looo
333
39
For stufiying tha e f f e c t of concomitant inoculation on
penetration, 15 days old seeJlinijs ware? t rdusfanrsi t o odcro-
pots containing s t r i l izeiJ SdiKJ, To such fjots caa ) la te . Long-
i^hton nutrient solution was addeJ VstianaV' r naed©'^, Tba scheme
of inoculation i s given as beXo^t
TaJjie d i
500
iDOO <m
500 «•>
100 n
5 0 ) 333
10 X) «•»
333
i'Ajmbsir of s;:)0cimans incwulatad
ii* g^^fUfarmla,
sua
lOUO 50 J 500
1000 mm
s>oo
333
tm
1000
333
50Q
10 )• 500
380
loeo 580
m
333
1000 333
'Giants from each treatmetit waro uprooted. The roots were
stained in lactoi^fienol-acid fuschin s t a i n (4c 30th et a l . ,
i 9 4 l ) and i a t s r each raot system was j>artially dissected and
nuttaber of larvae/inxnatura females enters J a f t a r 12 , 2 4 , 4 8 , 72
arKi 168 hrs of inoculaticjn ware countad. There f ee r e p l i -
c a t e s for each treatment.
For studies dealing with the histopathology of infactsd
r o o t s , iS days old ss^fdlings ware tra;isplaiift«d i n micro lots
containing autc^lavad s o i l . The scht>njo f o r inoculation i s qivan
as balowi
Table e *
MmihQT of sp©clBi»ns in£«:ulat©d
333
333
« • 333
333 3 i 3
333 333
333 « 333
333 333 333
2 8
war& uprooted four weeks a f t e r inoculation*
roots were washed and amali pieces of roots wora cut and fixed
i n F . A . / u and dahydrataciby T.iJ.A. s e r i e s and emoeddwd vmith
paraffin wax and the sect ions were cut at by rotary micro-
tome CJ'ohanson,i94J).
For deteimining tha fecundity of root-knot and renifoxin
nematodes t ten ©cjqmaases of equal s i s e ware selected f r o s each
o f the above treatment and each eggraass ^ms kept i n 2 pi^rcent
sodium hypochlorite solution. «ggs i n each egcTwiass w<?re counted
and mean was determined.
In order t o detoitaine the e f f e c t of ii. r9nifogHij.f and
and X* braasieae on the hatching of 4 . ^lyogr^t^g larvae* the
following schama was employed.
Table ^ t
of specimem
i * l»ncgqnitfl a* ^ffaasigag,,
Nematode aliquote (ft, penifQJtnia / I * braaaieaeJ .
5 eggoiass
» do «
- do «.
- do -do -
. do .
500
SOI
50Q
303
renifoimis)
(I* SEaaal&asL)
(a* jrynliQimli & L toialttflf ^
2 0
incognita* I * r^nifonAa ^nd x* fergaa^lctaft i|3,orw and
^^Ifg^Qnl (^abia « u figs. uio)t
i(hm tomato s e e i l i r ^ s wera inoculdted with iOO, 5CX),
9CXJ o r iOOu) ljrv.jo of J . incogni ta , the .aopulation 6a days
a f t a r Inoculat ion was 4 8 2 , 1928, 2148 and 350 J res:30Ctlv®ly,
The plant waifjht was B , 6 , 3 , 8 8 , 4,IB 'ind 2 . 9 g r©s a c t i v e l y as
against 1 2 , 0 g of urdncwulated >lant and mala/faraalo r a t i o was
. 0 7 / 1 , 0 . 2 2 / 1 , 0 . 2 8 / 1 and 0 . 3 0 / 1 r e s j e c t i v e l y . For corres.bond-
ing inoculum lave Is root-knot indox was 2 . 0 , 2 . 7 , 3 . 0 , 3 , 0
r0S jectivQly.
Un inocula t ion ^Lth 100, SO J, 900 or lOOJ immature
fwnalas of Ji. renifaHiiia t h e Jopulation 60 days a f t a r i n o c u l a -
t i o n was 650 , 1773, 2735 and 376B r e s p e c t i v e l y . The plant
waight for t h a c o r r e s jonding inocuiuBn l e v a l s was 5 . 1 1 , 3 . 8 0 ,
3 . 6 0 , 3 , 6 0 r e s p o c t i v e l y : raale/fmaale r a t i o was 0 . 4 3 / 1 , 0 . 5 B / 1 ,
0,61/1, 0 , 6 8 / 1 ros aoct ivoly .
On inocula t ion with 10 3, 5 0 ) , 900 or 1000 specimens of
I * iaraasicaa th# f i n a l po ,3ulation was 2 0 4 , 1196, U 1 5 and 1696
r9s ;>«ctively. The plant weight f o r c o r r e s sonding inoculim
l e v e l s was 1 0 . 8 J , 9 , 8 0 , 7 , 0 ) , 8 , 0 0 g r o s a c t i v e l y { taale/famale
r a t i o was 0 . 1 4 / 1 , 0 . 2 5 / 1 , 0 . 2 7 / 1 , 0 . 3 3 / 1 r e s p e c t i v e l y .
Fiq. i , 2 & 3s Growth of tomato c v . Marqlobe vi an inoculated
v^th and Iffit yj nchyAtfa
renlformes alons anvi with thv mixture of the
two.
Fig. 4, 5 & 6 t Qromth of tomato cv . Marglobe M en inoculatod
bra««ica« alone and with the mixture of thd
two.
Figs, 7 , 8 & 9 t Growth of tomato c v . Marglobe wh«n inoculated
vvith /^loidoQviw 4 new nit i and lytsnchoyhYmhMi
byaa^^eae alone and with th« mixture of th«
two.
Fig. JLOJ! Growth of tomato c v . Margiolie when Inoculatod with
ItifiaaolM.
and IvlanchQrhvnchus ||ra§,slfiat. alon® and with the
caixture of th« t h r » » .
3 5
How9v«irt vMhon tondto stttfdlinga w«ro inoculat«d simul-
tarwously with tho mixture of ^ i^r^onnlta and i^r^fani^K
In th« r a t i o 900 4 • JU) >B, the fiopulation of the foxiB«r waa
762 as against 2148 and t h a t of l a t t e r was 275 as against 650
when inoculatad singly. With 90QH * lOJM, the xj >uiation of
tha faxm^r was 1410 as a jdirist 2735 and tha l a t t e r was 197 as
a jainst 482 when inoculated sincily, hith 500 4 • SUDH r a t i o
the iJopulation of fona»^r was 677 as against 1928 and of l a t t e r
was 1348 as against 1773 whan inoculated s ingly . The plant
weicjht for the corras iondlr^ treatments was 4»80, 4 . 2 3 , 6,DO g
res . jactivelyi njala/f«nal« r a t i o of jl. i i y w a s 0 , 4 5 / 1 ,
1 . 0 / 1 , 0 . 6 2 / 1 res jectively and of Ji. ^nifogi^i^s, was 1 . 2 0 / i ,
0 . 8 9 / 1 , 0 . 9 9 / 1 respect ively ar*J the root-knot ind@K was 2 . 3 0 ,
i . o u , 1 . 9 a r^s-Hjctively.
Whan t«»a,ito seedlings were inoculated with the mixturo
of il. jTffn fgffiri..? and i . ^fiigflfiaff i n th^ r a t i o 90M • 10 IT the
oocAjlation of |i. ranifonats was 2673 as against 2735j tha
pomildtion of brdssicdiw was 92 as against 204 when inoculated
singly, with 900T JjOOft, th® population of foraer was 666 as
against 1115 and of the l a t t e r was 6J2 as against 650 wh@n
inoculated singlyi with 500a • 500T, the jjopulation of the
former was 1598 as against 1773 and of the l a t t e r was 321 aa
against 1196 when inoculated singly. The >lant weight for
corres;^K>nding treatments was 4 . 5 , 4 . 3 and 3 . 6 g res lectively
and Rtale/female r a t i o of H. reniforrais was 0 . ^ / 1 , 0 , 4 8 / 1 and
o G
0 . 5 8 / 1 and that of brassicae O . l B / i , 0 . 2 6 / i and 0 , 2 6 / 1
respec t ive ly .
un inoculation with 900 and iOQ, incoonita and
X* the ^ooulation of th < form'r ivas 1804 as aa.iinst
2148 and of tha l a t t e r was 183 as aqainst 204 wh@n incwulated
singly, uith 90i/r lOJM tha X)3uldtion of tho fom^'r was
1134 as aqainst 1115 and the population of tha l a t t e r was 283
as against 4B2 when inoculated ainrily, uith 5 0 J 4 5051 r a t i o ,
the tJOiJulatian of tha former vvas 1358 as arjainst 1928 and th©
rjo'TUlation of tha l a t t e r was 11X30 as ag i n s t 1196 whan inocu-
l a t a d s ingly . Tha ;>lant weight f o r tho c^rras-Jondinn t raa tnants
was 3,60, 8 . 4 0 , 7 . 7 0 g res lect ivaly c ials / fenale r a t i o of
iA* InCQQ'l^a Was 0 . 4 0 / 1 , 0 . 6 8 / 1 , 0 . 4 7 / 1 and of I . brassicas,
was 0 , 1 6 / 1 , 0 , 2 8 / 1 , 0 , 2 4 / 1 respec t ive ly . The root-knot index
was 2 , 5 , 1 , 8 , 2 , 0 resi ject ivaly .
Whan seedlings were inoculated with the mixture of the
three nematodes (333 saecimensof eachi the t o t a l ;X)i>uiation was
2689, out of which the oojulation of i^, inccKuiita was 905 ,
H, retiifoistiis was 1155 and X, oraaslcae was 629, The plant
waic^ht was 7 . 5 qii male/female r a t i o of incognita was 1 . 4 / 1 ,
of H. R^iUlVnjLa was i . 2 0 / l and of T. t faSSiCtilff was 0 . 2 4 / 1 ,
The root-knot index was reduced and w is 1 . 2 0 .
From the data given here i t can be concluded that the
population incrt^ase of both endo^arasite was adv^ rse ly affected
3 7
when prasent toqether* hoMOVert tho reduction was taoro i n case
of ,4.nC9 ntta, than a. in raniforra-atunt neraatod®
comfadnjftion tha population of tho form^^r rcnaained unaffectod
while that of Xafttert was roducad. In case of incoanlti^ and
X* bgaaaleae conibination, although th© tsoaulation se-ajned t o be
raiucsd at alX cc^abinationa but at hiqher incKulum iovai of both
tho n&matodas the raduction was not s i g n i f i c a n t . Th^rs was a
tendancy towards the [iroduction of mora malas i n a l l above
treatments. This tand&ncy was mora ironouncml i n ^^
and a» as eawparad t o I . brflg^jgj^*
d' JUmaafllMi H* toaalifida mA I n SLSffli IMyLans (Table « 2 j Fiqa. l jU20)»
aiigplant seedlings wara inoculatad with iOU, 900 ,
9 0 J and IQOO larva® of J . inconnit^. the population 60 days
a f t e r inoculation was 1968, 7022 , 7241 and 8670 r e s p e c t ! v a l y .
The plant weight was 4 . 0 , 1 . 7 7 , 2 . 8 7 and 1 . 2 3 g respect ively as
against 8 . 0 9 of uninoculated plants ; male/feoiale r a t i o was
O .oa/1, 0 . 2 4 / 1 , 0 . 2 6 / 1 , 0 . 3 2 / 1 r e s p e c t i v e l y . The root-knot
index for the corres xinding inoculum levels was 1 . 5 , 2 . 4 , 2 . 8 , 3,C
respect ively .
On inoculation with 100, 500, 900 and 1000 immature
females of ranlfomiis the f i n a l population was 788, 1264, 1994
Fiqs, JL1,12 ii 13« Hoot» of aggplant cv . ^Ba i\ir jla Long
vvhf?n inoc datad with inc jonl t^ . and
ii* y^niforniig aloiia and with the mixtura of
the? titfo*
Figs. X4, jys & i 6 t Hoots of oggiilant cv . Purpl«» i*ong
wh«n inoculated with gf^iUlglfflia
and with
tha mixture of the two.
Figs. 17 ,18 & 191 Boots of sgqalamt e v . ^ s a
when lnocuiat«d with I^Mdftayng I n O T r ^ l t
and and
with tho oiixtura of tha two.
Fig, 20i Hoots of egapXamt c v . i*u»a i\irpl« i-ong wh»n Inocu-
iatad with InfiffifiinAtJ*
and with the naixtur® of th® t h r w .
4 3
and 2062 r«Si>ectiv«ly. The Jlant weight for corras bonding
inoculum levels was 5.C]5» 4 . 4 , 3 . 9 g respectively and a a l e /
female r a t i o waa 0 , 3 i / i « 0,40/1, 0 . 5 0 / l t 0 . 5 0 / i r e s p ^ t i v e l y .
on inoculation with 10 J, 5(30, 900 and 1000 specimens of
X. braaaicae the f inal po.HJlation was 502 , 918 , 1086 and 1415
respect ively . The plant weight for corxespondir^ Inoeultmi levels
was 5 . 2 5 , 5 . 0 0 , 4 , 7 1 and 4«85 g resi>eQti\^alyi male/feiinale r a t i o
was 0 . 1 1 / i » ••14/1 , 0 . 3 1 / 1 , 0 . 3 6 / 1 respect ively .
Howevir, when seedlings were Inoculated with the miKture
of M. l-iyour^ta and ii. i^nifograis i n th© r a t i o 9 0 J i 10 7H,
the pomjlation of the f o m e r was 1966 as against 7241 and the
population of the l a t t e r was 643 as against 788 inoculated
8inr|ly. .dth 900H f- 1004, the population of the formar v as
1125 as against 1994 and the poi%ilation of the l a t t e r was 230
m against 1968 when inoculated singly, with 5014 • 500H.
combination, the po,:)ulatlon of the foxiaar was 1524 as against
7022 and the population of the l a t t e r was 773 as acjainst 1264
when inoculated s ingly . The ;olanit weight f o r the corresponding
inoculum levels was 4 . 1 2 , 4 . 5 5 and 4 . 7 0 g respectively^ male/
female r a t i o of J . incot^nitj^ was 1 . 1 1 / 1 , 1 . 2 0 / 1 , 0 . 6 8 / 1 respec-
t i v e l y and of a . ga»liforalis was 1 . 0 / 1 , 0 . 5 2 / 1 , 0 . 4 5 / 1
respectively ami root-knot index was 2 . 5 , 0 . 8 , and 1 . 8 respoc-
t i v e l y .
66
whan eggplant seedlings wera InocuiataJ with the mlKture
of a* and T. brasslcaa . In the r a t i o 90 H h- lOQT, th®
popul«ition of the foxraar was 1964 as against 1994 and the popu-
l a t i o n of th© l a t t a r was 53 as aqainst 502 vsfhen inoculatad
singly. <>n Inoculation with 90 JT * IQJU, the xjpulation of tha
former was 9 0 J as an .linst 1086 and the population of the l a t t e r
was 678 as againut 7tiG whon Inoculated ainrjly. »dth 50QH +• 50DT
mijcture, the ixipulation of th© former was 1233 as aqaimt 1264
and the population of the lat t i?r was 769 as against 918 wh«n
inoculateJ singly* The Jlant weiqht f o r corresoonlini corteo-
mitant inoculun l e v e l s was 4.6U, 5.U2 g raspnctivalyj
nials/fanalQ r a t i o of r9f4.fQy.4s. was 0 . 4 8 / 1 , • . 5 0 / 1 , 0 , 4 0 / 1
resaact ively and of X* ^rasaicaa was 0 . 1 2 / 1 , 0 . 2 4 / 1 , 0 . 1 5 / 1
ras lact ivaly.
Un inoculation with 9OJ 4.inconnita nd 100 T. bras s i c a® th®
population of the former was 6890 as against 7241 and tha popula-
t ion of the l a t t e r was 356 as aj^ainst 502 whan inoculated s ingly .
Ulth 9OOT+IOOM, tho po Julation of th 2 foritjar was 1(J9B as against
1066 and the po lulation of the l a t t a r was 1781 as aqainst 1968 when
inoculated singly. With 500i4 4- 500T r a t i o , tha population of
the formar was 5944 as ag .iinst 7022 and the population of tha
latt '^r was 608 as against 918 whon inoculated singly, Tha plant
wai<3ht for corrasponding concomitant inoculum levels was 2 . 9 3 ,
3 , 2 0 , 3 . 8 5 g respect ively ; raale/fetnale r a t i o of ineoordta
4 5
was 0 . 4 8 / 1 , 0 . 4 2 / i , 0 . 4 8 / i and of X* br^asicati was 0 . 1 3 / 1 ,
0 . 2 6 / l | 0 . 1 6 / 1 ms^metivoly and root-knot Index was 2 . 4 , 1 . 0 ,
2 , 0 raspact lvely ,
ifihan th© saedilncis were inoculated with the raixtura of
thrae namatodes (with 333 s >aciai»nsof each) , the t o t a l pomjlation
was 1852, out of ivhich tha wpulatlon of J . incog nit a was 660,
oi was 680 and of I . 512 . The plarrt
woiijht was 4.JU) gf faala/femal^ r a t i o of ^neapn^ta was 0 . 9 0 / 1 ,
of a* gonlfqnais Was 1 . 4 9 / 1 and of I . was 0 . 2 4 / 1 and
root-knot indfix was 1«80.
Xt appears from th@ data presented here that in r o o t -
knot and rani form n^matod® combination th© po/xilation of both
the natnatodes was adversely affaeted. ^ had bean the tond0ncy
for production of males aqainst thos® treatments v^ere sa^dlings
hav» b0»n inoculatcNi uith o i thar of the ntimatodessinqily. In
reniform-stunt mtmatode combination, on th» oth«r hand,
rwproAuction of th« l a t t a r was reduced and no adverse affact
was noted on th« population of tho forra»r. In root-knot—stunt
nematod* cgnbination, tha reproduction of th<$ former was
adversely affected at a l l conbination8,v«^er;»a8 the t>orj)ulation
of X* remained unaffected at hi<jh< r inoculum l e v e l .
In t h i s combination also male/feraale r a t i o was disturbed.
4 0
SIX* XngffmaUffin ,Qf c a y J i U m g cvt ^AUY. ^nwtoiXJL a f f ^ U m t
kUI i M* incvatUta* a* gyn^f^gmli iiosl I* Ugai^icw aiaai
iihon cauXiflobver soedlings wera Inoculated with iOU, SO),
9CK:i and XOUO sp^clmensof ^rMsiianita. the f inal poixilation 60
dii/s a f t e r inoculation was 34 , 62 , i 0 2 , 2W reS; )0ct i /Bly . Th©
plant waiqht for the 6orr»s Jonding inoculum l0vt»ls was 6 . 7 0 ,
6 . 0 3 , 5 , 4 0 and 4 . 9 2 g res lectivaly as against 7 , 3 0 g of
uninoculated alantst male/fetmale r a t i o was 0«20/X, 0 , 2 9 / l , U . 3 4 / i
and 0 , 3 8 / 1 ras.jsctivoly and root-knot index was 0 , 0 1 , O.iO, 0 , 1 5
and 0 ,18 res * ic t ively ,
un inoculaftion with 100, SQJ, 9 0 J and 1000 immature
feaialas of ii, renifortniff. the f ina l population was 110, 521 ,
1040, 1045 respect ively . The plant weight for the corresponding
inoculum levels was 4 , 3 2 , 3 , 8 7 , 4 , 1 0 , 4 , 1 8 g r e s m c t i v e l y as
against 7 , 3 0 q of uninoculated plants ; nale/female r a t i o was
1 , 2 0 / 1 , 1 . 2 0 / 1 , i . 3 0 / 1 , 1 , 3 5 / 1 respect ively .
^ith 100, 500, 90v3 and 1000 specimens of I* ^rassicae the
f inal population was 492, 1212 , 2996, 3386 respect ively . The
plant weight for the corresponding inoculiMi levels was 4 , 2 0 ,
3 . 0 5 , 1 ,58 , 1 .40 q respect ively ; inale/feniale r a t i o was 0 , 0 4 / 1 ,
0 . 1 0 / 1 , 0 . 1 5 / 1 , 0 . 1 7 / 1 respect ively .
However, v^en the se«dlin<js were inoculated with the
Fig. 2 i , 22 & 231 tirotesth of cauilfiowor c v . ©arly snmvbali whsn
Inoculatad with toloidoavno incoanitdj and
l^tvlanchulua g^nif jrmis alom and vdth tha mixtuE®
of thd two.
Figs* 24 , 25 & 2 6 : Growth of cauliflower c v . &arly snoivball
whan inoculated with gfnjLtQMBlt
and TYlgnff^ jy^vnfihya and with
tha mixture of the two.
Figs .27 ,28 S. 291 Growth of cauliflow«r c v . «ariy snowball
wh«n inoculatad with %),i?l,f|,i9Y.nff inCMnlta and
Tvlanchorhvnchu^ braaaicaft alone and with th«
fflixtwre of th@ two.
Fig. 30$ Growth of cauliflovwar c v . early snowball whon
inoculatdd with mi9l49SfYm lngM,nl.la» l^tY^^fWlttflM
renifaimig and ^iof^
with the mixture of the throe .
5 2
olxturo of ii. ^ncoanit^ r»nifcanalb in the r a t i o 9CX}M >
lOJH tha population of tho formar was 88 as a<3ainst 102 and
th« (Population of the l a t t e r twas Xii. as against 110 nvhan inocu*
lated s ingly . With 90Uii ^ Wui,thQ {oojulartion of the former was
1014 as against 1040 and tha i>opuiation of the l a t t e r was 20 as
against 34 when inoculated s ingly , va.th 5 0 ) 4 SOJH, the
population of tha fomer was 46 as against 62 and the population
of tha l a t t e r was 515 as a laitist 521 wh@n inoculatoJ s ingly ,
'fhe plartt vmhiht for tha corras:sonding inoculum levals was 3 . 6 5 ,
3 , 1 0 and 3 , 5 8 g raspectivelyJ male/fetnale r a t i o of J , incaar^lta
was 1 . 3 8 / 1 , 1 . 3 0 / 1 , 1 , 4 0 / 1 and of was 1 . 5 0 / i ,
1 . 1 9 / l t 1 . 6 8 / 1 ros-jectively and root..knot index was 0 . 1 4 , 0 . 0 1 ,
0 . 1 0 raspdctivoly.
Wten soeJlings were inocul^teJ vdth th@ mixture of
a . ggntfOgariLa and X. Igaaaicaft, i n tha r a t i o 90 U IOJT, tha
population of th« forraor was 923 as against lOAO and th® !3opu-.
l a t ion of th« l a t t e r was 478 as against 492 whwn inoculated
s ingly . With 900T • lOOH coiiibination_,tha poiulation of th«
foxm&r was 3126 as against 2996 and tha po xilation of tho
l a t t e r was 30 as against 110 whan inoculated singly. iMith
50)H > SOOT, the population of the foxsner was 126 as against
521 and the population of the l a t t e r was 1236 as against 1212
when Inoculated singly. The plant weight for corres >onding
inoculun levels was 2 . 5 3 , 1 , 6 0 , 4 . 8 2 g respect ively ; male/
femala r a t i o of fi, rerdfogais was 1 . 0 / 1 , 3 . 0 / 1 , 2 . 5 / 1 respect -
ively and of I . brasaleae was O . O 8 / I , 0 , 0 5 / 1 , 0 . 1 6 / 1 ar©specti\mly.
5 3
^han s0«<ilings wer^ inocuiata<i t»;ith the mixture of
i . Ing.gKanila and l . with 9 l » 4 • iX) TT r a t i o , tho po xu-
I ^ i o n of th0 former was 31. as a9airtst i02 and th« population
o f thi9 l a t t e r was 494 ds a ja i . i s t 492 inocuiatfifd s i n g l y ,
with 90JT * 10u4 raftio, t h » po^aulatian of tho forwmr was 2990
as aaain^t 2996 aiKi th® poaulation af th« l a t t a r v/m 28 as
against 3 4 wbsa incaculatad si-T^ily, ^ith 5Q ^ SajT, t h e
population of tha iormr was IS acjainst 62 and th© jo juiation
Dt th» l a t t a r was 1X96 js a j a i n s t 1212 when inoculatad sincily,
Tha '^Iditt wai:;|ht for th«i c o r r a s lo/idimi c o n c e i t ant irjocuXum
l e v e l s Was 5«2U, 3 , 5 5 anJ 3 #53 y r « s jectxveXyi m«3la/fa wie
r a t i o of ^neo^ir^t^ was 1 . 4 6 / 1 * 1 .4&/1« 1 . 5 0 / 1 r e s Tectively and
of X* iarjssic^a was 0 . 0 6 / 4 , 0 . U 6 / 1 , O.Q7/ i rm Kjctivviy and r o o t -
knot ind«x was O.CB, 0,01, 0 . 0 2 r e s j s c t i v e l y .
When seadlintjs wora inocylated w ith th® miKture of
thr«^ n^rnartodes (333 s 3»cim®nsaach )^the t o t a l >0 3ulaiti0n was
1232 out of which th« -copulation of J . incognita was 3 2 , of
r»t^faxwia was 90 md of ^rdsaicdg was l U O . l"he a.int
waight was 2 . 5 0 Bial®/f«H9ai« r«»tio of ^ ^neoonita was 1 . 4 0 / 1 ,
of ii. jcanlfgflajii was 2.SC)/1 and of I . vvas a . 0 6 / l f
roat .knot indax was 0 . 1 2 .
Fron the r e s u l t s prasantsd above, i t i s obvious t h a t
d* ^neaanLtM could hardly r a traduce on e<^uliflower, Q,* rayiifomds
was j u s t able t o maintain i t s popul^^tion and reproduction of
X* t|yaa«icaa was ^?rolifie. The r ^ production of ineoonita and
80
ganjfonala waa reduced I n combination. Hor9,too, th© tendency
f o r xcod^iction of more males was observed In case of ondo^arasltes .
XV. l iffact of s i n g l e , eoncqmlt^nt and sfiQaantlal iitocuXationsof
X* bg^ssleae (Tables 4-»7i F i q s . 3 1 - 3 3 ) 1
Th@ r a s u l t s of th® iittur^action of nsiaatodas when inocu-
l a t a d conjoint ly or s e i u » n t l a l i y ara summarlaod below.
( a ) i . incogni ta - r^fnlfomis i .Jn inocul<^tion of ogfj alaitt
seedlings vdth 1C3Q0 larva® of J . Incot^nitt.^. th® population 6Q
days a f t a r in(»ulat£on was 1331 and «clth WCD Invnature fanales
of ij. ronlfornais i t vva» 1655 . In simultaneous Inoculat ion with
5 0 J SiJQcimensof each* the x>pulation achieved wm 613, out of
which t h e population of ^ " c a a i ^ t a was 192 and of r tniformis
Was 4 2 1 . the s e e i l i n y s w@ir® Inoculatad vdth root-knot
nematode 15 days ^srior t o raniforoi mmatod® (with 5CKJ s::^ci-:ien5
each it t h e population achieved was 614, out of vi^ich th®
.copulation of M- inCQtinlti and of a* ffftn^kigmig was 3 5 4 .
^ e n roots were inoculated, on the other hand, with H* f e n i f o m i ^
15 days p r i o r t o ^ ineoonita the t o t a l population was 483 , out
of which t h e population of ^ ineoanlta was 51 and of reniformis
Was 4 3 2 . itioth concomitant and sequential inoculations adversely
a f f e c t e d the plant weight. The tendency f o r production of more
males was noted i n a l l concomitant and sequontial inocula t ion tes ts
H
MM
5 O <0
I
ffi tf o> a c
u « m A
•a
H
¥
« e
s S I
i : a "
I
M V
. 1
3
8 . § rJ ' - !
3 9 ' o o
oa a o
s s e • • • -I C M
I
lf> -I § 9
• % g
-I 3 C
8 S • •
IS 9 O O
H S 6 o
H £ 01 oJ
i
ii w
•-f in
HI ^ « TJ
•s 1 % mmi
a » 9
e • o u a §
• • o o
• • o o
3 3 • • o o
• •
t t
• * vQ O
SB
• O
K a
0 •
it • •
• • • •
5 5
£
ts to ffi % u o „
e %
JZ
H
I 8 jb
K ^ H ® R o •
ja m g
« > aj^f
•o 1 •H « O 4* I f ' I a •8 c o
* c o •H i-i Sf R
g •-4
«
ffi OJ JO
^ % to Hi ^ S m m
i i o " "a 0
g, o
1 1 M
3 M
» » « iH <0 ^ i « OL CJ» t-« e> «B « a
in
i
H
•H 2
a l a
A-
® jn^
Mm
a c
m n s §
ri Q
« C a
§
F fS
' §
o
• • • o rt
I I
3 o
0 o
s ? ^ to
* o
o
CM
I
§
8 TS ct III
tJ
r I
o
9 g o
cn
M O t c.
TS
tt %
5 : 1 • • o o
• • OO
8 ? • •
I I
s s
s x
II HH
o a • • • •
I ttt
I
a
5
a>
I K &
i § til
JO
ti
ns
I I ^ « > JC • 1
5G
s o m 4*
t i 1
IT •o
s H 1B
•H E §
« 11 x
•p
« HI
o u o
1
1
® -g 1 « 3 JC 0 ^
IS
I s » JC « M
4J 9 <M •H "S I flj « « • 8
m
£
1 ^
5 ft f
H :
o
l4
JO
TJ
3 ® §
p-4 C
R m c
OM
n
K
-it
t* HJS
If
«
•-I
1-4
1
8
• 1
O
• • • « o
I I
I
8 S 3
o
o
3 3 S C4
2i
en
•H a
a
-I
2S o
9 !!;
$ «
I
I «t
r> tj S o
9
-4
>0 -4 O
o
§
1
•n
-I 3 U §
•5-
• • OH
9Ri • • o o
• • O O
-I
O-l
I I
-lb. • *
NOCh
HQ
I S •HH
• • 1 Q • • • •
1 IS « w 15 u 21 rs %
% 51
x:
5
57
(9 n OS t a Hi •n t "
C o
•S <A i« JO « •H HI
I at •
Fig, 3 i J iirowth of ogciplarrt c v . pusa aurple long when inoculated
alono or th j i r mixture siraultanoously anJ sonuentially •
Fi j . 32i Growth of eq ji^lartt c v . pusa i^urala lonq whan inoculated
with ,r?n4ff?gr4g. ana
jyy.^^sic.fg alon© or t b a i r niixtura ai aultanaously and
aenuontially.
Fig, 331 urawth af agq dant c v . - usa purple lonrj whan inoculated
alone or t h a l r mixture simult jneously and s a n i s n t l a l l y .
500M+500R 500M+500R M INCOGNITA SIMULTANEOUS -nrULATED, ATION vc PRIOR reniformis INOCULATED
« 15 DAYSPRiOR
FIG. SI
fIG.32
_ 500M+500T SOOM'iSaE i S - i f ®
F1G.33
Fiqs, 34 & Growth of aggplant c v . pusa purpl® lonti when
inoculat&d v«ith jmCMfAU* %lY
j^rAfgir t f Iyl^n^?^9y^Yr^chu8 tosslsit aion®
or t h e i r mixture simultanAously and sequentially.
• B B 333M«33R+m T i33M+333R + 333T 333M+333R+333T TpmiSSlcaF lSniV r lOOOT I ^MSffiO'JS ; M incognita INOCU- RRENIFORMISINOC nPSTm S; niwcH » riOOOTJ INOCULATION • WFO15 DAYS PWR ' ULATED15 DAYS FTORJ 2 ^ ^
FIG.34
C O
F1G.35
G 1
and more were obsozved in tha nematodos with vi^ich s«9d«>
l ings mre inoculatod laiter» howevar, root-knot index was
reducod whan yanlfoy^ia wa» inoculated pr ior t o i , incognita .
(lli ranifagmia - brasslcaot wh n saedlings weare inocuiatad
t4 th iaX3 imniaturo faiaalss of y- 'i-Aforrais or v4th lOUQ s locimen
of X* tarasslCf^a. the f i n a l i^oijulation was 1880 and 1050
respsctiv»ly» In simultaneous inoculation (with 5iX) s KHsiraan
aach) tha .copulation was 1210, oat of viftiich tho population of
ii. JEHtUaDQla. was 841 and of iEiSSiiLa w is 369 . »»h®n
sdded A3 dtiys pHor t o ilEi&Siftii&tthe
pojMJiatian was out of which there were 870 s mciimna of
d* ^ r d formlm and 80 of bafassic.dQ. ivhan ss«iulings, on the
othv?r hand, ware inoculated with X» bras s i c aa f i r s t , tha copu-
la t ion V j3,907, out of which th^ i^opulation of ^ r^nlformla
^ ^ i * braaaicaa was 448 and 459 ras >®ctiV0ly, doth siisulta-
naoui and sequential inocula t iomsignif i cant ly raducad dry
weight, howwvar, thora was no difference betwaan the different
traditinants. Tha mala/faai^la r a t i o was not disturbed,
( c ) d ' Incognita - I . fhan seedlings wera inoculated
with iOUC) spaciraan of J , Incognita or X. bg.<s8tcae. the jopula-
t i o n was 1616 and 1025 rosoect ivaly . In simultanaous inoculation
tha population was 648, out of which the population of
ii' ineoanita and X« brassieae Was 336 and 312 rast^ectlvaly.
G 2
VVhen seedlings inoculated with incognita f i r s t , t h e popu-
lat ion was 744, out of which thare were 4S J s pecimen of 4«if|eoQnita
and 294 of X« braasicaa . In ray?r3o s i t u a t i o n , on tha other hand,
the oo Hilation was 616, out of which th© ,>Oi3ulation of J ,
i,ncoanita was 280 and of I . was 336 . The dry woiqht
of the plants was signlficiintly reducad both i n simultaneous as
well as sequential inoculation?* Hoot-knot index as well as
iaale/femalf» r a t i o was influ^ncsd*
W 4* incognita • ^^nifopa^a - X . QSmsXSM.* seedlings
v/erci inoculatod with iOX) specimans o i t h » r of incognita or
roniforsnis or X* tog^ssicae alona, tha f i n a l po'julation 6D d ays
aft jr inoculation was 1350, 1789 ond 1142 r©s ^©ctivoly. In
sii^ult snaous inoculation v^th 333 s jecimens of 0<sch, tha t o t a l
copulation was 781 , out of v#tich the potjulation of inc(3qnita>
and X. iZOasjJUS. was 5 5 , 648 and 78 re3.:>@ctively.
when M- was added 15 days »rior t o ii. and
braaaieatt. th® t o t a l >0 x i i j t ion was 4 8 7 , out of Vslilch the
copulation of i^ncoonita. H. jCfflAfggiBli, and T. l^yagglCi? Was
201, 220 and 66 resxvct ivoly . when rgniforwis was added 15
days )r ior t o ji. incrwinlta and X* braaiiie^a. th« t o t a l lulation
was 1127, out >f v\hich the i30;*ilation of ineoonltfl. ^^.reniformis
and X. brasaicae was 49 , 1011 and 67 respect ively . However,
I . b r a a a l e ^ Was added 15 days w i o r t o i , infigqnita and
li* renlfonala^ tha t o t a l ^^opulation was 1214, out of vyhich the
population of incog git a, gQtTjLf<?niti X* frffflialCiff, was
6 3
5 3 , 1026 and 135 r « s j ^ t l v o l y .
How0V»r,v^en ^ncQanlta anJ ii, ranlformift was inoculated
simultaneously and brasaicae was addad 15 days l a t e r , the
t o t a l lopulation was 1163, out of i^liich the iw ^ulation of
incQ^^nita. ^nd I . ferxa^s^ffaf, Wd® 4 7 , 1040 and
76 res^>ectivaly. when a* <jnd I , iMEiasifim were
inoculated sijraultaneously and J . Incocinita Has added l a t e r ,
the t o t a l population 1248, ;jut of which the poaulation of
d* f ncQiinit^, H. and ^^^ 5 0 , 1 BO and
l i e r e s K?ctiv<aly. «h«n inconnltd and iarassieae wejra addsd
aiinultaneously and JJ, r e n i f o r ^ s , 15 d ^ s l a t e r ^ t h e t o t d l loiula-
t i o n was 648 , out of which the poaulation of l .Tcoanlta.
ii- rar^formis and X. brassicde was IQB, 277 s -d 263 r e s J^ctlv^ly.
Tha dry weight of tha slants waS s i j n i f i c irrtXy reduced i n compa-
r ison t o uninoculati^d slants , Tho sex r a t i o was also disturbed
i n case of endio >arasitas, 4ore laal^s were oiassxved i n concomi-
tant as well as secuarrtial inocula t ion t e s t in com larison t o
treatments whera e i t h e r of the nematodeshas been inoculated
s i n g l y .
V. a f f e c t of Binale and concomitant inoculations on tomato
( a ) id. ineoanitfl « Ji. genjift^niitt (Table 8 { Fiq, 3 6 ) l Wh n tomato
seedlings grown i n unaraanded jutoclaved s o i l , w«re inoculated with
Fig, 361 uroivth of toirijto c v . nijrgloba aajJI inga qrjwn in iioil
auandad kith naam caka, on inocuiotlon m t h ^feloidociynQ
incognita and aotvlonchulus rani-forrfsis alona or t h o i r
:jiixture.
A sa Mended
U =3 Unamended.
95
5CX3 or JjOOO l^rva® of ineocifiita, th» oo^Hilation was 658 and
2543 respect!v«lyJ vdth 50D or 10 X) iraraature fataales of
ii* th0 ;Jopulation was 999 and 3338 r»Si>«ctiv»lyj
vd-th th» mixture of root-knot and reniforja nematod® (500 specimen
OdCh)» the jjosulation of root-knot and ranifor^ nera^todas wa8 4Q4
and 528 roselectively. Thus mixed inoculation brouciht about
raduction i n the po.julation of both th® n®aatod<ts» In s o i l
aiaend»i livith inorganic f e r t i l i a a r raduction in aoxilation occurred,
both i n 3in"|l0 and corwanitant inoculfitions,
Un inocuiiTfcina the soedlinis -jrjwn in s a i l arimndad with
mma cake at the r a t o of 1 g nitroyen/kr} s o i l stith or UXK)
larvae of |.ncoanita. the jjo HJlation um 66 , 28Q respectively?
th« corr®S'>onding figures for ii. rer^ form!s 153, 411 resrwc-
t i v e l y . In concomitant inoculation, the oopulation of
d* incognita was 62 and of y^rdformia was 153. Un inoculating
th^j seedlings grown i n s o i l awwndei »dth c a s t o r cak© the pooulation
of root-knot was 142 , 356 resiiectiv^lyj reniforra nematode was 401
and 8CX3 r»5 lectively and in laixed Inoctilation the population of
iJKffanl^a was 140 and that of r e n i f o r ^ s was 389. On
inoculating seedlinrjs grown in scrf.1 amended with groureinut cake
with 5 0 J or 1000 larv.ie of incognita, the population of root -
knot nematode was 287 and 302 res aectlvelyi with 50 3 or lOCX)
immature f«;^ales of r«aiforni nensatoJe the population was 507
and 693 respect ively . When inoculated with the mixture of
two nematodes (500 each) the po>ulation of ^ incognita was 190
G 7
and of ij, rantfonaia was 3 9 6 , In s o i l dinendiid with thlm®t th«
population of i * incognita was 497 and 6X0 respect ivelyi
H* gonlformlft was 795 and 1074 respect ively and i n the milted
inoculation the population of root«.knot n^matodo was 334 and
r e n i f o m nanatode was 480•
On inocuiatiny th« sof^dlir^s jr-vm i n unamnds^i s o i l ,
s o i l anior%t0d with inorgdnio f e r t i l i « @ r s , t^etn, ca3tor» grouncimit
cakes and thimot with SO J larvao of Incoonit^, the male/faniale
r a t i o was 0 . 2 0 / 1 , 0 . 2 5 / 1 , 0 . 8 2 / 1 , 0 , 8 0 / 1 , 0 . 7 B / 1 and 0 . 7 0 / 1
respact ivalyi t h e corrasjondii^ on inoculation v /ith SO^
innaatura faaalaa of reiiifopiiia w^r© 0 . 5 4 / 1 , 0 . 6 0 / 1 , 0 , B 2 / 1 ,
O . B V i t 0 . 7 8 / 1 and 0 . 6 2 / 1 iraspijctlvolyj with lOJO larvae ff
incoqf^t^ t h a ujals/famale r a t i o for th@ corrfjs londing
treatments was 0 . 3 0 / 1 , 0 . 3 9 / 1 , 0 . 8 9 / 1 , 0 . 8 2 / 1 , 0 . 7 5 / 1 and J . 7 2 / 1
r e s o e c t i v e l y ; and with lOJO ifatiature fa ia l ss of yerTlforra^p, i t
was 0 . 6 8 / 1 , O.TD/l, l , 2 D / l t 1 . 1 5 / 1 , 1 . 1 0 / 1 and 0 . 7 4 / 1 res^wctively,
vJn inoculation with the mixtura of two neraatodas (50 J spoclmens
oach)^tha (nale/female r a t i o of f o r carrespondlng
treatments was 0 . 6 0 / 1 , 0 . 6 0 / 1 , 0 . 8 6 / 1 , 0 , 8 4 / 1 , 0 , 7 7 / 1 and 0 . 7 2 / 1
resiMctively and of H, renlforml« w«s 0 . 8 2 / 1 , 0 . 8 1 / 1 , 1 . 2 5 / 1 ,
1 . 2 0 / 1 , 1 . 1 0 / 1 and 0 ^ 7 / 1 respec t ive ly .
At both tha inoculum levels used and i n conccxnitant
inoculation tha dry waight of slants was adversely affec ted ,
Howavar, i n case where plarrts were grown i n s o i l araanded with
o i l cakes, tha dry weight of plants increased s i g n i f i c a n t l y .
C 8
Although tha dry weight also inereasevi i n c a s * of plants grown
i n s o i l amanded with thiraet or f a r t i l i a e r but i t was l e s s i n
comparison t o o i l cakes.
( b ) . I . yenifomia « X. ^SMniUm, (TaijXe 9 , F ig . 37,38):iifh0n tomato
seedlings vjrown in unevnended s o i l were InocuX^ed with 'SfXi or
inmnatura femal»50f rentforrais. t h e population 60 days
af t r inoculation was 722 and 1286 rasat^ctively I with 50Q o r
iOOQ S3«ciman of I , braasieae th» population was 682 and X084
respect ively ; with the mixture of ri^nlfana and stunt-neraatode
(5CX) speciiaan each) , the population of ronifaroii nanatods was
622 und of stunt nnaatoda was 467 . -Slxed inoculation brought
about si^jnificant reduction i n tha i^fxilatian of stunt 'nematodo.
On inoculatlnn the seedlings gro^m i n s o i l blended with irjoraanic
f e r t i l i s e r , s imilar r e s u l t s war^ obtain'dt howevertthe population
in each treatment was l i t t l e lower than unamended s o i l .
On inoculatinej the seedlintjs grown in s o i l araerrded with
neeia cake with 500 or JUXJO iimsature females of reniformig.
the mraber of renifortais s jecimen i so la ted were 270 , 413
respect ively ; the corresponding figures f o r brassieay were
200 , 722 res- iect ively . In concomitant inoculat ion, the population
of ii. rfnifyffail was 279 and of I . tyra^.ajffdg 2 8 5 . material
diff0rences occurred vvhen t e s t s were made In s o i l amended with
c a s t o r and groundnut cakes. In s o i l a^aended with thltaet the
population of ii. reniformis was 498 and 840 res wct ively and of
Fig , 37J Growth of tomato c v . marqlobd seai l inqs eirown i n s o i l
amended with nmm cako, on inoculat ion with ^totyl/nehuli^s
ygnlfgrraiff, l i s m k ^ m ox* t h e i r
mirbure,
A a ^wended
U a Unamanded
Fi j, 381 Hoots of torsjato CV. raarqlob* ? saeJlingo rirc^'n i n s o i l
aaendsd with mtsm caks , on inoculat ion v/ith HotvI^nichul^B
r e n i f o r ^ i anJ TyJ^^ncf-'prhvnchus aXona or t h ^ i r
mixturo.
X* fryf^^caa was 510 and 789 r^siwctively and in mixed Inocula-
t i o n th« j>o;3ula(tion of roniforra ne«atod« w&s 432 and of stunrt
nesiatods was 301 .
Tha ^ a n J a e n t s infiuencad thw 5iial»/fanal«» r a t i o of both
tha nematodes. Un onoculating th® scj^dlings grown in unamondttd
s o i l , s o i l amondad *vdth inorganic f o r t i U z © r , neein, cas tor*
groundnut cakus and thimtrt v^ith StM iwaatur© faraales of
ii* rgnlforrain. the m^le/femala r a t i o was 0 , 3 0 / 1 , 0 , 5 5 / 1 , 1 , 1 4 / 1 ,
0 , 9 7 / 1 , 0 , 7 9 / 1 and 0 , 6 3 / 1 res 5€fctiV0ly. The corras londing
figures f o r X* wara, 0 , 1 6 / 1 , 0 , 1 7 / 1 , 0 , 2 5 / 1 , 0 , 2 2 / 1 ,
0 , 2 1 / i and 0 , 2 0 / 1 ras ^actively, i hon seedlinga ivere inoculatad
with 1030 spacitaons of e i t h e r of tho naaiatodej.^ an increase in
th® mal«/f0friala r . i t lo »K?as observed, the increase was mora in
case of 4 , |!^ni,fQrtnia i n a l l the treatments. In conccnaitant
inoculations with 5 0 J spacimej^ of oach, tho raale/faiaale r a t i o
of reniform nematode for corr«s Jondint-i troaftaients was 0 , ( ^ / 1 ,
0 . 7 0 / 1 , 1 , 2 2 / 1 , 1 . 0 5 / 1 , 0 , 8 1 / 1 , and 0 , 7 0 / 1 ras )ectiv«lys of
X. brassicaa was 0 , 1 8 / 1 , 0 , 1 8 / 1 , 0 , 2 8 / 1 , 0 , 2 4 / 1 , 0 , 2 5 / 1 and 0 . 2 2 / 1
r9sp«ct ively .
In a l l tho t roataants no«ai cak« was more a f fac t ive
follow«d by c a s t o r , (jroundnut, thimet and f e r t i l i s e r * Similarly,
the o i l cakes caused an increase in dry weight of plant in conpa-
rison t o the ilanta cjrawn in unamended s o i l .
7 2
( c J a . l n 9 9 g m i - i . (Table 10 , F i g . 3 9 ) 1 m&n tomato
seodlifKjs qirown In unaraendiodi s o i l were liioculate'i with 500 or
1000 larva® of 4 . Ineoainlta. the fK»pulation w B 836 , 2002
resaoctlvdlyj with 500 or 1000 soeciinan of brass lcaa . th«
population was 782 , 1257 respactiv^lyj with the raixtura of r o o t -
knot and stunt nomatode (with s i^sciinena each) , th® population
of root-knot nematoJe was 335 arhi of stunt nematode was 491 .
•^xed incK:ulti1:ion i^ruught about reduction i n the population of
iboth the minatode^. On inoculatin':} th» seedlings grown i n s o i l
amended viith Inoarrjanic f e r t i l i « i » r almost/^results were obtained,
howaver^the ijopailation at J i f f e r e o t inoculi»n l a v s l s was somewhat
low.
Un inoculating the seeiliniis grawn i n s o i l afn inded with
mm cake with 500 or lOOJ larvae of incog nit a . the t o t a l
number of |,ncoanit^ specimen isola ted were 156 and 457 resisec-
t i v e l y , tha corres Jondinq figures for 2.. braaaieae were 272 and 680
respect ively . In coneoraitant inoculation the x»pulation of
ineoanltq was 127 and of I . braasleae was 246, oirailar trend
was observed in c a s t o r and groundnut cakes. In s o i l amended with
thiroot, the ooiiulation of |f«Qanita was 474 and 614 respec-
t i v e l y and of T. brass icae 502 and 892 respect ively and i n the
mixed inoculation the population of root-knot nematode was 360
and the oofiulation of stunt nematod® was 3 8 5 .
In undEat nded s o i i , sioil amended with inorcjanic f e r t i l i i e r ,
neem, c a s t o r , groundnut cakesand thimet with 500 larvae of
Fl3» 39 (a b ) j lirofeth of toiaato c v , sadr globss qrofe/n i n a o i l
aTi^niaci with naea Cdka, un x .uculat ion with
dSMdaiiima i O S i i a i ^ byasslcaa
alona or t h n i r raixtura.
A a i^ended
U ct Unanendod.
106
tho male / f^aXa r a t i o f o r corrosi^ondlrK} ir tatraentt
Was 0 . i 5 / i , 0 . 1 6 / 1 , 0 . 8 0 / i , 0 . 7 0 / i , 0 . 6 8 / 1 and 0 . 6 0 / 1 r«s i»c t iv»ly ,
Th0 corresponding figures f o r X* brassleaa were 0 , 1 7 / 1 , 0 . 1 6 / 1 ,
0 , 2 2 / 1 , 0 . 2 1 / 1 , 0 . 2 1 / X and 0 . 2 2 / 1 res w c t i v a l y . When to®dlln(3»
were inoculatad with 1000 larvae of or lOCX) specimon
of T. bra«siCd« an increase i n maXeness Wds observed, ^hen
seadlinys wara inoculated with 10 XJ specimen consisting of 50 ^
speciman each of J . incoardta a \d b r a s s t h e aialo/female
r a t i o of root-knot nQfaatode for the correspondir^ troat?aents was
0 . 3 8 / 1 , 0 , 4 2 / 1 , 0 , 9 0 / 1 , 0 , 8 0 / 1 , 0 , 7 4 / 1 and Q , 6 9 / l respect ively
and of X. brassicae was 0 . 2 0 / 1 , 0 . 2 0 / 1 , 0 . 2 3 / 1 , 0 . 2 2 / 1 , U . 2 0 / 1
and 0 , 2 0 / i foop?ct5.vsly.
Asjendaants with o i l c . ^ e s brought about an ir^jrease in
tha dry weight of plants i n comparison t o unawended s o i , ard
a decrease in root-knot index was a lso observed.
incognita - a* - I * frgfligjCJff, (Table l l , F i g . 4 0 ) !
Sfth n t o ^ . t o q;eedlin<|s grown i n unamended scdl were inoculated
with 333 or lOOQ larvae of incognita , the population 60 days
af tor inoculation was B62 and 1516 r e s p e c t i v e l y ; with 333 o r 1000
inmature females of H. renlformis the population was 1286 and
2238 respect ively ; with 333 or 1000 Sfjecimen of X« brassicae the
population was 600 and 1380 res Elective ly and with the mixture of
ths three neaatodes at the r a t a of 333 s lecimen of each, the
population of root-knot nematode was 620, of reniformls was
Fig. 4 0 : Growth of tomato c v . m<]r9Xoj»o soedlllngis grown in s o l i
amendetJ with neem cak« on inoculation with i4s>loidQayne
l i o a a i i u a alono or t h e i r mixture. A SB v^anded
U Unamended.
109
1062 and of X* braaslca^ was 3 3 6 . r o /41xod inocula t ion brought
about a r®uuction i n t h » po 3Uiation of a i l th^ t h r « e n»»natodle«
but i t was more i n case of root-knot and stunt naraatod«$.
iidmilar retiuction was obtained whan seedlings wera qrcwn i n s o i l
aser^ed with inorganic f«.*rtiiiz«»rs«
Un inoculatinri the aeadlings grown i n s o i l amended with
m m cake with 333 or 1000 larva® of J . imoanltat* the t o t a l
nu!Bb0r of J . incognita spociraen i s o l a t e d 120 and 250
r e s jsc t ivolyi the cor ros ^ondinij fii^ures f o r yQnifQrTOi, ,
250 and 663 r0S;>ectiv0ly and f a r br^isaicao wer«f 261 and 500
ras .^activoly. In concoraitjnt inoculat ion tho potwlation of
imf^nar,^,,^ 126» of 258 and of I . b r a s s ^ c a e
was 260 , ^similar r a s u l t s vrare obtainad i n treatments receivino
c a s t o r and groundnut cakos* "hen seedlings w r a grown i n s o i l
anaonded with thim»t and inoculated with 333 and 10 )0 larvao of
id* i n c a a n i t a the f i n a l population Was 216 and 773 rasp«ct iv©lyj
the c o r r a s jonding figures f o r li. r^niforrois were 360 and 960 r a s -
p s c t i v e l y and for i , br^salcaa wara 336 and 795 r©s l e c t i v a l y . In
tha mixture of thr«(} nematodes (333 specioien of each) th» po{Mjl«*>
t i o n of ill. ^ncotinlta Was 2 0 6 , of rgnifoymls was 336 and of
I - togfiffflf was 340 .
4ala/f0faal» r a t i o was profoundly influenced by di f ferent
t raatm^nts . On inoculatinq the s e t t l i n g s grown i n un^andad s o i l ,
s o i l amsiKled with inorg<inio f@rtil iz«}r» ne»iB, c a s t o r , groundnut
cakes and thiraat with 333 larvae of i n c o g n i t a , the malt/f^raals
110
r a t i o was 0 . 1 5 / 1 , 0 . 2 0 / 1 , 0 . 9 0 / X , 0 . 8 8 / X , 0 . 8 0 / 1 and 0 . < » / l
r tspactiveXy. The eorr«sponding figures for raniformls w»re
0 . S 2 / 1 , 0 . 5 8 / 1 , 0 . 9 0 / i , 0 . 6 6 / 1 , 0 . 8 0 / 1 and 0 . 8 4 / 1 r»apoctively
and for i . b r a s B i c ^ wara 0 . 1 6 / 1 , 0 . 1 8 / i , 0 . 2 5 / J l , 0 . 2 5 / i ,
0 . 2 2 / i and 0 , 3 5 / i respcsctivaly. un increasing tha i n i t i a l
population from 333 t o lODO, an increase i n snaio/female
r a t i o was odserved. «ti<?n seodUn^s woro inoculated vdt.h 10JO
specimen consistinij of 333 siieciraen of bach nstaatod©, male/
female r a t i o of ii. |,ncoanit^ was 0 . 9 0 / 1 , 0 . 9 2 / 1 , 1 . 0 0 / 1 , 0 . 8 6 / 1 ,
0 . 8 0 / 1 and 0 . 6 8 / 1 res.}$.'ctivelyt of ii« renlfo«aia v»as J l . 2 / 1 ,
X . 1 0 / 1 , X . 3 2 / i , i . a S / X , X.XU/X dnd X.2S/X i^spsct ivsly and of
1 . arasaicao was 0 . 1 8 / X , Q . i 9 / X , 0 . 2 8 / X , 0 . 2 9 / X , 0 . 2 7 / X aiKl
Q . 4 V i respect ively .
Frcsn th® data given aJJOve as well as in e a r l i e r chapters
i t i s obvious thot i n c r e ^ e in inoculuro l e v e l , mixe 5 inoculation
and .saendin,! the s o i l with o i l cakes have pronsoted the production
of niora males.
In seedlinqs grown i n amende ' s o i l and inoculated sinqily
or i n the mixture, there has been an increase in root mass as
well as in t o t a l dry weiqht. lijespite t h i s very high reduction
i n population occurred presumbly because the t o x i c e f fec t of
decomposition oroducts of o i l - c a k e s .
111
dffact of single and eoneaialtant inoculatlorisof^ nanolant
losLaaiilSa • s:?iUfami>la. (Taw© 12^1 whan se©dunqa of erjiplant ijrawn In unaraendod s o i l wera inoculatdJ with 5 0 J o r
1000 Itirvaa of incognita , th® population was 691 and 1218
rQSpoctively; vdth 50J or l a X ) im nature females of ^^ rgnlfarsiis
the i^opulation was 874 and 1453 res oectivaly | with th# mixtur©
of i^ot-knot and ranlfjxm naniatod® (with 5DJ speciraen tachJ , tha
population of root-knot neoijtoda was 380 and of roniform ne?matod»
was 573 • 4ixad incx:ulation thus ^rau^ht aijout r«JiM;tion in
po-Hilation of hath newatod©s» i n inocalatinrj tho scjedlinis nrawn
i n s o i l ^aaendai vdth inorganic fsrfei l izer reduction in oopulatlon
ttim obsorvQd,
Un inoouljtinci the sG«rdlings qr<»m in s o i l imanded vdth
mm cak® vdth 50Q or 10JO larvae of incamnlta. the number of
im^onit^, s»cimf»n i s o l a t e d if as 148 and 318 respect ively , the
cor ras jondini figures f o r ii* rardfonaia ware 265 a.id 430 respec-
t i v e l y , In coix:o®itsint inoculation the population of l^neoonlta
was 144 and of remfanal^ was 229 • ^ni i la r trend was observed
i n s o i l t raated vdth c a s t o r and groundnut cake^. In s o i l amended
with thirnat the population of incoanit^ on inoculating with
5 0 ) or 10 specimen was 424 and 761 respectively and of
ii* r»nifa«ai« was 513 andlli^ rospectlvely and In the mixed
inoculation, the population of ineoanlta was 337 an<l of
a* geiUfOCTtla was 483 .
113
Xt apieairs from the data that the population of both th«
enioparasites was roducad s i g n i f i c a n t l y i n conccxDitant inocula-
t i o n t«.'st in comparison t o singla inoculation i n unamandted s o i l
anii s o i l ataendad with inornanic f « r t i l i « e r . whon seedlings war®
grown i n s o i l auandsd with o i l cake and incxsulatod with ona o r
\(i.lth th« itdxtur© of tiKJmatodes, a highly sigsiificdnt r«^diiction
i n tho poijuldtion of both tha n®n«itod«9 mas not^d, oath i n
sin-lie ani co-KSomitdnt inoculation tffsts.
an inocvilatinr? tht> samJlings grown i n unamended s o i l ,
s o i l araended vtith inorg*»nic f w r t i l i x a r t iw^ ,, c a s t o r , qroundnut
cakes ani thim^ t with 50U larVd® of incoatTit^ the laals/feraal©
r a t i o was 0 . 2 2 / 1 , 0 , 2 4 / l , 0 . 8 4 / 1 , 0 . 8 2 / 1 , 0 . 7 9 / 1 and 0 . 3 2 / 1
resp©ctiv©lyi of il. rani.fcflRaia i^as 0 . 4 2 / 1 , 0 . 4 8 / 1 , 0 . 9 2 / 1 , 0 . 7 5 / 1 ,
U . B l / 1 0 . 7 0 / 1 Tils j o c t i v o l y . ITie raaifi/fanidis r a t i o of both
the neui.jtoi0sim5rada9d i»ith the incr^aso i n inoculura lev»l in
a l l the traatinants. In conccwdLtaot inoculdtions of both n®®a-
todds (with 5 0 ) spt»cira9n 0«achj tnale/fofaaltt r a t i o of root-knot
namatode was 0 . 4 8 / 1 , 0 . S 4 / 1 , 0 . 9 2 / 1 , 0 . 9 0 / 1 , 0 . 8 8 / 1 and 0 . 8 0 / 1
raspect ivalyj of Ji. roniformip was 0 . 6 6 / 1 , 0 . 6 6 / 1 , 1 . 3 3 / 1 , 1 . 3 0 / 1 ,
0 . 9 6 / 1 and 0 . 8 7 / 1 ro»p»ct iv»ly .
Tha dry w«i<3ht of plants grown in s o i l amended with o i l -
cakes was s iqnif icantly incr<»asad i n comparison t o plants qrown
i n unaoaanded s o i l both i n singla and concacoitant inoculation t«st£>.
ai iai larly tha root-knot indax was also reduced i n th» presonc®
of o i l cakos at a l l inoculum levels t e s t e d .
114
<i>> ii. rer^formla « I . ^mmlSSM (Tabid 1 3 w h e n «e«dUnq» of
agg alant qraun in unaoierKleci s o i l * ware Inoculat&d with 50^ op
AOOO iiamatur© f«r^ale8 of K. yenlformia the population 687
and 1389 respact ivalyj with 5DJ or iCWO speclmn of br^ssleaa
tha (lo'iulatlon tfuas 628 and 1102 Pdspectively f with the mixture
of rent form and stunt mimatode (5CKJ stjocimen of each) , th® nopu-
i a t i o n of renlform nmaatodo was 628 and th® !>o;>ulation of stunt
n<?ii2atodo was 484* Un incKiuiation of the seedlings grown in s o i l
araonded t?«ith inorganic f e r t i l i ^ a r , tha pofiuiation of both th®
nffiiiatodeswas r#duc@d both i n singlo and conco^iitant inoculation
t£?sts»
Un inocula t inj t h e seedlinqa grown i n s o i l ara«»nded vdth
r: ara cake with 5 0 J or ICXX) iniaature femalss of i^r^fgrrai^,
t h e f i n a l population was 4X0 and 590 r e s p e c t i w l y j the corres»
ponding figures f o r X» bgassieae were 170 and 493 resp«?ctiv®ly.
In conconitant inoculation the |K}pulation of reniforwig and
X* Was almost t h e same as when seedlings were inoculated
s ingly . Ihe sarae trend was observed in seedlings grown in s o i l
amended with the remaining two o i l cakes ami thiraet,
un inoculating the seedlings grcflwn i n unaniedded s o i l ,
amended with inorganic f e r t i l i z e r , neem, castor,groundnut cakes
and thimat with 500 imraatura fenales of reniforrals. the male/
female r a t i o was 0 . 4 9 / 1 , 0 . 5 0 / 1 , 0 . 9 9 / 1 , 0 . 9 6 / 1 , 0 . 9 3 / 1 and
0 . 7 1 / 1 respect ively . Tho corres xjndinj figures f o r X. brassieae
116
were 0 . X 4 / 1 , 0 . J15 / I . 0 . 2 9 / X , 0 . 2 7 / i , 0 . 2 1 / 1 respect ively .
Vi/hen seatilinqs wara inoculated with 1CX)Q specimen of H«reniformia
X» braasicae an increase in ra^^io was recarded.
In conc(xiiitant inoculation with 500 specimen of a ^ h t the inal®/
feraale rcttio of r e n i f o n nematode f o r th© corres x>ndinq treatments
was 0 . 5 9 / 1 , 0 . 6 7 / 1 , 1 . 2 0 / 1 , 1 . 1 0 / 1 , 1 . 0 ) / l and 1 . 0 0 / 1 respectively^
of I . iiK^ssicae was 0 . 2 7 / 1 , 0 . 2 9 / 1 , 0 . 3 9 / 1 , 0 . 3 6 / 1 , 0 . 3 4 / 1 and
0 . 3 5 / 1 res x?cti / 9 ly .
u i l cafces brought adout a s ignif icant increase i n the dry
v^eight of slants. i«era cake was laost a f f e c t i v e followed
c a s t o r and ground nut cake.
( c ) l a s a a i i M - 'I* ( table 14)1 whan Biq Uant.
seedlings gr jwn in unafaended s o i l ware inoculated vdth 501 or
100.3 l.=jrvae of incognita , the population was €98 and 1141
respect ively , with 5CD o r 1000 o;^Jcimen of T, braasieae^the
rxipulation was 706 and 1110 res a c t i v e l y j with the mixture of
root-knot and stunt nematode (with 500 sj^ecitnen each) the
population of root-knot naniatode v is 320 and of stunt nmaatode
was 562 . j^ixed inoculation thus brou4jht reduction in the
population of both the nematodes. On inoculatinq the seedlings
grown in s o i l amended 4«ith inorganic f a r t i l i i e r almost s imilar
resul t s were obtained, however, the population in different
treatments was low.
118
On incxsuXatinq th« saodJlrnis grown i n s o i l dtnand»d with
naaci cako with 50U or iCXX) larvae of ii* incoani t^ . tha population
o ' i n c o g n i t a was 248 and 403 r e s a c t i v e l y and of X* fara»&icae
was 4D4 and 567 r e s p e c t i v e l y . In concomitdnt inoculat ion the
ijopulation of 4 . incogni ta 228 and of X* bj^^sslcaf was 3 9 1 .
.iirailar trand i n population of both the neraarodes was oiiserved
when seadlinqa wara qrown i n s o i l amended with c a s t o r and qround-
nut c a k 0 § . In s o i l araiendad fdth th imat , the population of
d' incanni ta vms 348 ami OBO ras Ktctlv@ly and of T, brass icaa
was 5B2 and 919 r e s p e c t i v e l y . In raixod inoculat ion , the po;3Ula-
t i o n of iroot-kn'jt. namatoia was 519 and af stunt n«ma'o wa« 5 7 0 ,
un inoculat ing th ? 8QodUnf|S qriSin i n unaraendod s o i l ,
s o i l awjnd@d vvith i n o r janic f u r t i l i z « r , naajn.castor, groundnut
cakas and thimet «vith SOU larvae of 4 . incoardta . the male/
female r a t i o was 0 . 2 0 / 1 , U » 2 2 / l , 0 , 8 2 / 1 , 0 , 8 0 / 1 , 0 , 7 6 / 1 and 0 . 7 4 / 1
riispectivfeflyiand of i , brasuicae was 0 , 1 8 / 1 , U . I Q / I , 0 , 2 4 / 1 ,
0 , 2 2 / 1 , 0 , 2 2 / 1 and 0 , 1 9 / 1 res iQctively, Th^re was an increase i n
male/fifiadle r a t i o of 0 0 t h n«!uatod«fs when K>UO sp<icimfsna of each
neaaatode were used. In cancomitant inoculat ions with S<X3 specie
men of each nematode, the male / feuale r a t i o of incoor^ta wa»
0 , 4 5 / 1 , 0 , 4 8 / 1 , 0 , 8 8 / 1 , 0 . 8 6 / 1 , 0 , 8 0 / 1 and 0 , 7 6 / 1 r e s o a c t i v e l y
and of X , brass icae was 0 , 2 0 / 1 , 0 , 2 0 / 1 , 0 , 2 6 / 1 , 0 , 2 4 / 1 , 0 , 2 3 / 1
and 0 , 2 0 / 1 ras .oect ively ,
Ihe dry woi<|ht of plants was increased they were
Qrvjwn i n s o i l amended o i l cakes, i^endn^nt with inorqanic f e r t i -f\
119
U z ^ r and thimot also i^rouqht about an incroaso i n tho plant
w«ilht mxt lncr<»ase In t h i s was l e s s i n comparison t o
o i l oakas.
4* - - (Tabi® X5)t
vihcfn a.} jolant seadlin^js iruwn i n unasaended s a i l ani incKsulated
with 333 or iXXXJ larvae of ij^^anitajthe* population 60 days
iiftsr im)culation was 1^2 and 1430 respect ively i with 333 o r
1000 speciuian of Ji. ;rBnifQi?(Bl& the po|HJiation luas 1278 and 2070
respectively and vdth 333 or lOUO srieciman of bPtisai^c^g i t
was << 0 and 13W r«s jec t iva iy . saedlin is
inoculato't '.idth 333 sp'..?ciraens of e.^ch nemaT.ode^th© .Jcspulvition
of J . ingar^nl,;!;. wi, of ^ g^.r^jlfinpto ^^^ i i^e and of
X* bf.ji3sica# 290 . ^n inocul,3tian of soedlin^s grown i n
s o i l asT.and id inorjanic f tar t i l i ier^the population of nema-
todes was z%r Jucad«
un inoculation of the wedlings jrown in s o i l araondod
with n«@iii cak0 with 333 or lOX) larva® of J , incoonita . th»
f inal population was 306 and 445 raspec t iva ly . The carrosponding
fiqur»s for ygnifoianis were 442 and 816 resp«ctlvdly and f o r
l|JPa«sieaa i t Was 310 and 932 res sectivaly and with the mixture
of thre<» nematodes th« )opulation of Jt* ineoonita was 292 , of
H* CTAfaimll was 416 and of I , jicaiaififli 275 . The s imi lar
trend of reviuction in ^ p u l a t i o n was observeci in case of s o i l
amended with thimot*population,of iryocifiita was 633 and 8 6 0 ;
90
of ffinlfMMll, was 579 and X12B and of 'I. brdfajCfff was 432
and iC350 rsspact ivelyi with the rolxturo of three neroatodos th«
population of lncftqni,t,|, was 368 , of H. was 593
and of T, ty^paaaic^y was 4 i 0 .
Thus frota thesa data i t a that tho populcition of
®ach nsnatoda increased vdth th® increase i n InocaXm level
whsrf^as i n concocdtant inoculation t©sta th® population of each
B vfcies was roduced. J . ^neoanlt^ was most adv rse ly affec ted ,
Mhi?n thQ S00dlirt«j» grown in amended s o i l and ^ere inoculatad
with th® thraa n©matodas a i t h « r singly or copcori ' . - iMy,
population of a l l tha thraa n*Ha.?tod03 was reduced.
un inoculatinvj th® seedlings gxmn i n unaraend^J s o i l ,
s o i l ^endad vdth inorganic far t i l i jsar^ ttieen?, c a s t o r , qroundnut
cakes and thimet with 333 specimen th® ruale/female r a t i o of
incoonita was 0 . 2 ^ 1 , 0 , 2 6 / 1 ^ 0 , 8 4 / 1 , Q,82 /X . 0 . 8 0 / i and
0 , 2 8 / 1 respect ively , of iJ. a^aflUfiJCSaU ^as P . 5 0 / i , 0 . 6 0 / i , 0 . 9 8 / 1 ,
0 . 0 0 / 1 , Q . 8 0 / 1 and 0 . 7 6 / i respect ively and of X, brassicae was
0 . 1 5 / 1 , 0 , 1 8 / 1 , 0 . 2 2 / i , 0 .23/JL, 0 . 2 4 / i and 0 , 2 0 / 1 respect ively .
The male/female r a t i o of a l l the three nematodes increased with
the increase in inoculum level at a l l the treatments. In
concomitant inoculations of the three nematodes with 333
specimen each, male/female r a t i o of root-knot nematode wam
0 . 8 8 / 1 , 0 . 8 8 / 1 , 0 . 9 5 / 1 , 0 . 9 2 / 1 , 0 . 9 0 / 1 and 0 . 9 0 / 1 respect ively}
of ii. was 1 . 2 0 / 1 , 1 . 2 2 / 1 , 1 . 4 0 / 1 , 1 . 3 5 / 1 , 1 . 3 0 / 1 and
1 . 2 0 / 1 res pectively and of i . jim&lfiaS. was 0 . 1 7 / 1 , 0 . 2 2 / 1 ,
91
0 , 2 4 / 1 , 0 ,26/JL, 0 . 2 4 / i and 0 . 2 1 / 1 res >ectiv»ly. I t i s c l e a r
from the data .laraaanted here that mala/fajuale r a t i o of a l l th»
thr«d nematodos incx^asad both i n cor c omit ant inoculation t e s t
and in s o i l sanded tvith neeoit c a s t o r , qroundnut cakes and
thimot.
Tha dry iwaiqht of plants was a lso incraasad s i g n i f i c a n t l y
i n o i l caka omsndad s o i l . Tht? highast increase was obtained i n
nofinj cake, aimllarly tha root-knot index was a lso reduced in
o i l cake asnendsd s o i l .
d* pa op nit a «. t& rii f capral a tT ahla 1 6 ) I ivhiin two waek old
tomato se«^dlintjs grc»?n i n eutoclaved sand contained in micropots
and l a t o r inoculated with 500 larva© of J,, ^ficaanita. th® number
of larvae V( iich 3en®trated i n the root system 12, 2 4 , 48 , 72 and
168 hrs aft@r inoculation was 5 , 9 , 17 , 25 , nd 40 r®sj>ectiv®ly J
tha corres ponding figures on inoculation with 10X3 larvae ware
8 , 3 6 , 50 , 60 and 122 respect ively . Un inoculation of tomato
seedlings «dLth 500 imfflature females of Ji* rentformta tha number
of imnature females vi^ich entered the roots were 4 , 9 , 20 , 23 ,
and 4 5 ; 12 , 24 , 46 , 72 and 168 hrs a f t # r inoculation respec t ive ly ;
with 10X3 immature females the corresponding fiqures were 4 , 15 ,
16, 30 and 98 respect ively and with 500M + 501R, the number of
1 1 m « H4 •J
^ o g " - ^
3 w
c a ^ tit
o o ® ^
> 3 % ** A o
0 'ri
1 ^ 1 N f I $ t
u « « ^
2 a .H » u « c o
I a
« r - i
H
s X
g
•H tn
I I I m S) •-4 m ttx
a
t
I 1-4
o (4 JQ
(II » T» O %
z
n
2«H JO
s
p
SI
f2
9
« g R
' ' Si s a
3 g
S! 8
5 8
in CO
£1 3
CO
o -y S 3
I I
a
t sT)
I to
t N
I '•i
21 CO
« • CM to
iS 5 • • to in
• • fO
^ IM N Ov • •
g 3 • •
to to
B S • • Cl 03
S S • •
• •
1 1 tH f4
• •
• •
• •
92
M O lO 99
a
©
JC
JC
tn ^ s
«H a » O
fi
K ®
TCI
t c o
M ja
s ^
« i
t X
s I ISi t •rt
m
% u t m 03 r-» (8
3
'S t<«
O a I Hi
I
9
£1
a
3
fS
a
«« CH
g
® i>
>
I
I I I I I
I I I I I
I t 1 1 1
I I I I 1
I I $ 1 I
S 8
n s
a a
I I
§ 1
1 1
I I
I I
1 1
I I
o Si
s
s
a
9
I I
I 1
1 I
I I
I I
h Ch ^ • •
HI O • « N 3
M
S 3 • • N
N fc • • vi3 ^
I I
% ^
I I
% • c? « • .''» './I • • *4 .4
93
(0 0}
t» n. g
o
t>
N jQ w «, ti s •d
B •
-1 S
1
§ 1
i ] w a
•H
<<•11 •
H H JC *
1 1
S Oi t •H ® •H
I f4
6., «
2; m
0 »» JQ 1
4
8
« o
•h a 0 u
W-H JO
1
fs
9
S
2(
{8
fS
s
« >
J
I I I I I
I t I I I
I I I I I
I I I t i
I t I t I
H so rt g
5 q
M e
vO N
I I
O s
I 1
§
I I
1 1 N (M f • t « g
1 1 * •
1 1 CO • • « q
1 t m n S • •
n Si
I I
1 I
I 1
I 1
I t
• #
3 E;
§ 1 H 'H
a 3 X X • •
Q a • •
« • ^ >4
94
£:
®
(A » U n. n m s £
S I
^ V e? H
I , X 8>
J9
e 3. !H K «
M C
i t
| l
I i § ^
<9 S
8 ^
i : I
0
1 TJ •H «H o
o
o
rs 1 £
0 **
I •H X i
c i «H c o «>: « §
ee
2}
8 £
m
2 •rt lO
•i IS 14
r-f «B
I I IS
1 I O
M
1 t
il ti « « i a
0
c 1
It ? S
2
P
9
a
R
a
p
a i
s
HI
i
o s! « ^
I I
§
S 3
8
8
i I I
f9 f r»
t i
I t
I t
I I
I f
t I
a « ' • =1
I •
• f oi
rt CO I I «
t I
t i
I I
t I
I i
I I
f j
m
tn
«s
PI Q PI
I t I I
OiV • •
• •
if} 00
• •
tOOJ • •
ii • •
• • 01
II * • • * « •
£
iS 01 m % I e
c
% £
A
*
M
JS *
i 1 o 01 t =
I s c « 0
1 I M •H
t g, ^ « »
« f m «
96
root-knot n0t»«rtodo larvatt anterad the roots was X, 7, 3, 5 and 30
raspoct ivoiy ; and of gontforml^ 3 , XQ, 3 , 12 and 27
r®3^actively,
tioncomitant inoculation of tosiato SB&dllngs with the
fBixture of root»i:not and rcniform nematode adversely aff^ctod
th0 i)9iwtration of both the nmiiatodQS*
^IlLISBilS. • Qgdssicaft (Tdblo 1 7 ) i safdlinqs ware
inoculat ed with 50J imnidtur® for^ales of r^niforrqiB th® niraber
of itamaturo fmiales w^ich laonetrated tha roots^ 12 , 24 , 4 8 , 72
and 168 hrs aft-^r inoculation was IQ, 15, 22, 25 and 39 respec-
t i v e l y } th@ corresponding figures on inoculation with 1 0 ) 0
iiBRiatur© famal&s ware 6 , 12, 2Q, 30 and 50 t@s a c t i v e l y .
£• brassicae failB-» t o penetrate th® roots si tan inoculated
singly and even i n the treatments where i t was inoculated v«ith
ii. ranifarmim. however, the mmb&r of a . was 10 , 12,
20, 24 and 35 r^'spactively.
I t i s concluded that t>g>.ta3ieae neither f a c i l i t a t e s nor
inhibits the entry of Ji. rani formi^ in roots ,
( c ) i . incognita - X. toiffsigat (Table 18)s Jn inoculating with
500 larvae of -i* inconnit^. the number of larvae penetrated 12,
24, 4 « , 72 and 168 hrs a f t e r inoculation was 6 , 2 , 10, 3 1 , and 82
res >ectivelyi the corresaonding figures on inoculation with 1000
larvae were 7 , 18» 13, 126 and 273 res ^ectively. Here
97
too X* brasslcae fcjlleJ t o xjnetrato when Inoculated! singly or
i n comiai nation *vith Incognita . Tho nusib-r of i neon nit a
larva© on inocuiatinq v^ith the mixture v^are s innif icant ly reJuced
only 72 .and X68 hrs a f t ^ r inoculat ion. I t was 18 and 72. and
168 hrs a f t o r inoculation as against 31 and SZ v#ien inoculated
singly .
(ci^ id" incuoiTita - i . raniformia « T. brassicaa (lal>le 1 9 ) : i»hQn
tosaato ssej l imjs v^ar.j incKiulataJ with 333 s i»cimen of /vf«j.ncoani,;td
tha numijer of laarvae penetrated, 12, 2 4 , 4B, 72 and 168 hrs
: i f t9r inoculation teas 7 , 6 , 9, 4j and 45 mspsctivoly; the
corros bonding fiqures I'dth lOJU larVuQ of ji, incuanita vvarg 11,
40 , 52 , ii9 and 135 ras JoctivQly, Jn inoculation with 333 s lec i -
man of raniformia tha nuint>ur of imrnatur© fosJialtis counted ^12,
24, 4t>, 72 and 168 hra a f t e r inoculation was 3 , 4 , 6 , 2 U and 30 and
with 1GCX3 spQCiacfn vvas 8 , 3L0, 35 , 42 and 112 raapect ively . ito
specimen of X« brasaicae vvas found in roots , nhon tha seadlinqs
were inoculated with the mixture of thr®e na;aatodes (333 s o e c i -
iuon aach) tha t o t a l number of incoaai ta or ii. renifoyfnis
spaciman that panatrated the iroots was f a r bolow as against
those t reat^ants where roots wore inoculated with e i t h e r of the
two naaatodas singly.
From tha data present®J hare i t can be concluded that v^an
endo j a r a s i t a n d one ecto parasite are prasent toqether tha
penetration of both i s re Jucad, The reduction in penetration
was more in case of aJ. iPffMPt^^ than ii. reniforiala.
98
V m • a f f e c t of caneomitant inoculdftion of maaolamt 8«edliryi«
on ptnfftffitjign*
( a ) i n g p g n i u « ii- K^njLffflgmil (Table 2 0 ) 1 m^n eqaplant
seedlings were Inoculated Alth 500 laxva« of Incoanitfl.
the number of 1 irvd® that pu.Tetrdited th» r o o t s , 12, 2 4 , 48 ,
72 and 168 hrs a f t o r inoculation was 5 , 6 , ID, 19 and 4 6
TOSijectlvely» The cori^s iondii^ figures on inoculation with
1000 larvae of J , inco'inita were 6 , 8 , 19, 22 and 5 0 restjoctively.
Un iiioculcition idth 501 immature faj-tialns of roniforraift. the
nu!!b{?r of irarfiafturi? fa lalas that jwrn^trdited th«? r o o t s , 12 , 2 4 ,
4 8 , 72 and 168 hrs aft;?r inacuXaftion was 3 , 6 , , l i , 2 0 and
raso^ctivolyi and with lOOU imraature fe lales the c arres ponil nq
f igures wara 4 , 6 , 2 1 , 24 and 52 res oact lvely . However, when the
seedlinjs war© inoculdted simultaneously with the miKture of
i . and a - ro'tifoMais (50U spacinen of each) , the
number of incaoinitj^ Irirvae penetrated 12 , 2 4 , 4 8 , 72 and
hrs a f t a r inoculation was 5 , 3 , 6 , 10 and 2B reapect ively
and the numb r of iraraature feiaalas of g , rerdformis was 4 , 5 ,
5 , 8 and 30 respect ively .
From the data presented i t i s concluded that in mixed
inoculation the root-knot rwmatode prevents the errfcry of
renifoxm nematode i n t o the roots and v i c e - v e r s a .
(b ; ii. renlfQimlfl - tigaiffiCfli (Table 21 > I li/hen etigplant
seedlings were inoculated with 500 ira^nature fetnales of
« (c «a
I i U
n %
1 ^
•ti ft
c o g
w t tz •H l/i
«N n o J m p. fi
tN o U JO
"O © ts
Cv © c ? «
o G •H » i J m a
fS
IS c o
3
3
«
*
g x:
01 H ro a «M
i
<H o u %
1
3 -4
3
fS
S!
s
1 i i i .Hi •h 3
t I
» I
i t
t I
« 8
^ a
a ^
^ ca
I i
§ 1
9 » «
S
^
I t
I I
I t
t t
9
t f
I I ^
CO
iO
in
n
n
tn
f2 ^ ch t • •
3 S • • (M "it
«M VO • • O CO
5 s * • DJ XT
8 P • • to
& 5 • • ^ %
^ 0)
s g • • VO
IS & • « n to'
s t at «
• • ri o • • 1 fJi • •
(A <0
rf: 4->
tf)
o;
•s a X
o 3 4->
T3
iS 01 •H S S.
M
I J iti •
9 9
s I 1
%
01
c
% u
o o
^ 1
i : 8
(H
I ? Hi* O
u JQ Q
£ §
I I
c Oft
3 o
I
u
-51
s
l i s .H •sg
§1 H
{S
9
£1
I
fS
i
I I I I I
I I I I t
I I I I t
I t i t I
1 1 I t I
g 9
3 55
3 2i
v3 fs.
I I
t I
1 I
t I
( 1
» 1
t I
3
iO
t I
I I
t I
1 1
I 1
N * •
C3S «
• •
a 3 • •
» •
-t • *
I I
• • •3 O • * tn '.I • •
-I
« 2
Ml
e, a m nj U
&
JZ *> T? ,'Si'
fi fi t » X • ®
P 36 »
C o
«0 x> M
t CCt «
1 J tij •
n « a CP o t
I I
1 a % t . " ^ £ ^ % s ^^
- I ^ O 83
« 5 1
c
WC
S'
1 1 <A O iJ «
I I
Z %
X
S
•H
t
«M Q U « 1
V •IM n
i Hi
O
-4
N
S -I
a
§
a
i
I t
t I
t I
I t
I I
3 S
0)
^ 3
CM tf)
8 1
I t
t I
t f
I 1
I t
-4 ISi t r 't . t
V , t r • •
00
H 1 t •
CM
! ! sO H vC S ! * *
fN
* i 1 Os
H CO ( 1 H
• CM
I I
t i
I I
t I
t t
I I
101
I <t> •-i
(-a A
3
a «
-4
x; M
to It % u
% g
5
1ft
SS
'/I 01
I Nl
»< es o
C o
9 w <a 4J
% .a cn
c 0 o a
S -I
. 1
§ a>
<M C M o c
m
8 s
B m •rt
tn % n
I I % m S V § «w
t 5S
% 5 •H <H O N
2
Ht
%
c c
a
s
a
S
p 9
S
SI
e 9
o § . 1
i
a a
If) 00
^ vO
I I
I I
U 9
S i
I t
i I
t I
I i
I I
t
» • s
t I
I I ^
t t
I t (O
I «
I I N
I I tn
I I ^
I t «
?9 I I Ci r i
I I
t I
I I
I i
t t
t • • •
• •
m • »
• • ifieo
I I i I 3 ft
•88 • «
Si • •
a a
11 -I <H
* • QQ • • • •
Hi
i-J
iA 0
ci %
I -JS
GS
1 ^ ^ J n. g «
*
s s , •S I «» •H s g.
g
m «
103
gonlfoTmii|. th« numbar of immatura females that perMtrat^d
i n the root systea^iJZ, 24t 48 , 72 an*i 168 hrs a f t e r inoculation
was 4 , 6 , iO, i8 and 30 r e s lact ivaly j the corraspondinq fic|ur»8
on inoculdftlon with iOtX) inunature females v/are 4 , 7t i2» 22 and
44 respect ively , fiowaver, vs ien sa^Jdlinqs were inoculated
together with SOQ spaciraens aiWJh of ii* reniformis and
to»8l.,ff,iag,t the numijar of ifffi.idture f^aal^^s of |i. gffnUffmli.
pen#trrjt«d^l2 , 2 4 , 4 8 , 72 and i6B hrs a f t e r inoculation was 5 ,
9 , 9 , 17 and 30 r e s MCtlvely.
(c> - ! • 2 2 ^ h a n seadlinqs of
egg )lant were inoculatad with 50D larvae of incoar^ita« th®
nunjbar of larvae that penetrated, 12 , 2 4 , 48 , 72 and 168 hra
a f t e r inoculation was 2 , 5 , 8 , 15,and 35 r e s lec t lvelyj th@
correspondincj figures on inoculation with ICX)! larvdie were 5 ,
IQ, 12, 29 and 58 respect ively , X,* braaaieae f a i l e d t o
penetrdte tha roots i r r e s p e c t i v e of inoculum density. However,
when tha sewJlimjs were inoculated sinsultaneously with mixture
of J . Incognita and braaaicae v^ith 50 J s jecimen of each, the
nuibar of ineoanit^ larvae penetrated in the root , 12 , 24 , 48 ,
72 and 168 hrs a f t e r inoculation was 1 , 2 , 6 , 9 and 24 respec-
t i v e l y . Thus presence of X» braasicae retards the entry of
root-krwt mwt.atode larvae ,
ifii d* Ineoardta - y^tUfQinaift * I' (Table 23^ On
inoculating eggjlant seadUntjs with 333 larvae of ii , l^m^oanlta.
1 0 4
th« number of larva® ;3an«trat«d,i2, 2 4 , 4 8 , 72 and i68 hr« a f t e r
inoculation was 4 , 5 , 9 , 1 2 and 25 ras wct ivoly j th« corrasoondlng
figurtJs on Inoculation with lOCJO Idrvao wear© 6 , 8 , 15 , 2? and
46 r»sp0Ctivaly, ^Ith 333 Hpacimen of r;onlfarmiB wera 4 , 6 ,
8 , 14 and 22 r a » p » c t i / 9 l y and thft c o r r a s jondinq fir|ur0s for
lOQJ ifflfiature fQjnales of H. yotiirfpiTOis war® 4 , 6 , 11 , 2 1 , and
42 resjaectivaly. brassic^^tt fa i l©J t o enter , Honfaver, wh n
th© saadlings inoculated with mixture of thrs© ne a a t ^ « s
with 333 s lecimen aach, tha t o t a l nurabir of 4 . incognita larva«
panatraftod at f iva i n t e r v a l s ware 35 as against 5 5 and of
Of rgntfortais was 40 as agaif^t 54 v^an inoculat»d s ingly .
Thorefor©, i t i s c l a a r that in combination of the three naraa-
todos, the panatration i s reduced,
tfl, ttwaft ffi jreaMlUna . twa 9imX%
Th« changott result ing on i n o c u l a t i i ^ of see Jlings with
d* iOBftflnl^it a* gtnllfQtwH or I . alon« or t h d i r
combination on th« dev»l0f3ffl»nt of f«n}ale, formation of qlant
c a l l s and anatomical changes in the root t i s s u e t h i r t y days
a f t e r inoculation are given bolowt
With 333 larvaa of ^ incognita al^n® th • farrsles w»r«
obaarv»<l with t h e i r haad in th® s t o l a r region and the raaln body
in the c o r t i c a l region. Hach fetaale contained large number of
Flqi, 4 i J Cross sac t ion of tomato c v . margloje root 30 days a f t e r
inoculat ion with incoanitd showint) giant
c a l l along the vascular bundle. 20Q),
JG a i i a n t c a l l j rC » i«jc ls i ; 4F =» /datura fa male
VJ Vascular bundio.
F i i . 42J Cross sec t ion of tomato c v , raarqlobe root 30 days a f t e r
inoculat ion v^dth ioloi io>.iyna ii-icoqnita. ^^ 2CX3).
a ;..i,itura fai i i ls j » dfjn mass.
Fig. 431 J i a n t c a l l of Llff.f-niMii,, . contjininq
l^rge nuRibar of noclai ( i i ) . (K TOO).
Fig, 44» Gros8ss9Ction of tomato cv . marglobo root 72 hrs a f t e r
inoculation with Rotvlenchulua rttnifoiniia showing disso-
lution of c o l l wall of c o r t i c a l c « l l (C) n«ar head
region of nematodi® (RF.), (X 1630)
Fig. 49» Cross section of toaiato c v . <4argloba root 30 days a f t e r
inoculation with RotvluiTchuXus showing
kidney shaped f m a U (HF). (X 200)
N « Neck; r» « i 'er icyclaj « tgg ©ass.
Fig , 46 (a & b)» Vascular region of Fig, 45 aft high magnification
showing hyportroihied isericycle c e l l s arrar^ed in
semilunar fashion (GCH. aL) . (X ^ 0 0 ) .
Flg8»47 & 48« Cross sect ion of tomato cv . marglobe root 30 days
a f t e r inoculation with th» laixture of ^XpjfitMYng
and ^tYA^fichuiya f?fUtqrfaia showing
t y p i c a l giant CdllsCoC) and fa^iale of J . incognita
(MF)j giant c e l l s of |i. r ^ m f o r a i s i n v iscular
raqion (UBH, and in c o r t i c a l region (<iGH, C)
and female of Ji. yaaifonais (HF) and absenco of
c h a r a c t e r i s t i c samilunar pattern of giant c a l l s of
Fig, 491 Vascular region of Fig, 47 at high»r oiaqnificatlon
showing giant c a l l s of M. incognita (GC) and of
H. raniformis (UC.H) containing nuclei (NC). 850 ) .
Figs. SO & Cross section of tom&to c v . marglobe root
•rfien Inoculatad with <49lQidoavna incoofiita and
IfvlenchorhYnchua sHoMinq giant c o l l s
and fourth stag® femala larva® (liW) pro-.
sant s t a l a r and c o r t i c a l region. (X 130)
Fig. 521 Cross sect ion of tomato cv« Marglobe whan ino-
culated concoroitantly with incognita and
T« brassica^ showinq giant c e l l s (GC) and
mature female (<^F) with smaller numbor of eggs
iiM) and mala (eA), (X 150)
Fig. 53 i tsnlarged view of the root inoculated concomitantly
w^th •r^ntf^rffi.i rv Tvl^nchorhvfTchua
orasslc-ae showing hypertroaied pericycle (GCH)
in semilunar pattern (SL), (XitiOO).
Fiq. 546! Cross soction of t(»)dto c v . '4ar9lob« lnoculat«d
simultaneously with /4«loi<ioQvne incooinita.
^r as s ic a* 30 days a f t a r inoculation showing niatur®
immatur* fenaalo of jfi. incognita (I/4F),fourth
stage raale larva (MlilV) and giant c e l l of
r^^UfoifBii ^nd d' f m o g M t i , /absence of
semilunar arrangement of giant c e l l s of
reniforalis t o be noted. (X 2CX)),
- tF " ii. GCH a ^ a n t c e i l of li, renlfarrais ciC a Giant c e l l of jtmihrtUtft-
Fi j , 54} Gross s w t i o n of tomato cv . mar^jilobe root when
inoculation simultaneously with incoonita .
H. ret i iforais ana I , showinq the giant
c e l l s of J . and of feflifftg^i.
(GCH). (X 2 0 0 ) . RF » H. sMlflCQlil*
Fig. 56i Cross sect ion of tomato cv . raarglobe root vA>en
inoculated simultaneously with j . ji,ncy^nita.
a . rQniforiOia and I . o r i ss icae showing pyramidal
shaded gidnt c e l l of rt. reiiifornis (iSCH),^)
in c o r t i c a l region. (X 200),
I l l
eqqs (Figia.42* )• HOout 8-JL2 Qiant c e l l s w ro pr9S9nt»
i'Sajorlty of tham had disjmo cytoilasia howavert th«r« werof faw
which w«r'J mnpty, dach jiant c e l l containad i8»26 nuclei
(F ig . 4 3 ) ,
>dth 333 iimaturs5 fa nale® of 4 . renifarw^ls- kicinoy
ffijoaiws v ersj .aroiucad, Th» egqs w r® contJined i n
g^ilatinous matrix , head of tha f^ idle reached UfJto 0ndo-
dermis and p x icyc l© r egion jjk 2CX-22 c d l l s und;?want cbaaqm
on iaoth sida of the mouth of fenala qivinq a semilunar apj^ajw
anca giarrt c ^ l l appeared t o have ba sn
formed aithf r iay onlarq^'-^^rrt of sinqie c e l l or duo t o dissolu-
t i o n of c e i l wail of thtf adjvacant c o l l s , .formally each ^jiant
c a l l containf?d one nuoleus but there Bcm':i giarrt c a l l s
\Mhlch containad more than anu naclfsus^
Un inocuiat inj with b^asaieaa aloiia, m^crot,ic t i s s u e
d«valopod her® and thar» on th« o?jid«rml8 of the r o o t , howavsr,
larvae or adult nmaatodos ¥f&r» not oOsifrvod in tha r o o t s .
Inoculation of roots with isixture of root-knot and
reniforra neraatoda with 333 «^)ecirs#n of «ach rasultod in produc-
t i o n of r a l a t i v e l y small^^r far^ialos containing raducad numbar of
aggs by tha former, ^ m i l a r l y tha nunib'>r and siza of giant c a l l
appaarad t o hava baan raiuKSod, Furth er aach giant c a l l con' ainad
raducad nunbar of nuclai ( 2 - 5 ) . ( F i 9 4 7 . 4 9 ) , In mixad inoculation
1 1 2
a group of giant e v i l s prootucad by r^niform n««matodlo did not
have c h a r a c t o r i s t l c semilunar appaaranc^t howavQr» each qiant
c o l l ytm almost of the sani^ s i s * as produced whon inoculatod
s i n g l y , i^raovart thera was a tend^»ncy for i^roductlon of giant
Ciills evan i n c o r t i c a l pare^nchyma ( F i g . 4 7 * ^ ) .
un inoculat ion with tho ifdjctura of ranifori^ii^, and
X* i^rasB^cae with 333 specimen o f each no ?nat»rial change occu-
rred i n tha of f m a l m of the former. Tissu«» changes
rwsultin.j f ro ! rerdfogiais v -jra almost vdth
a . iffliMfqwAa ( a g . 5 3 ) .
•n inoculatinc) tha soQdlimjs with ineo<inlta and
x « toaaaicaa, tho giant c®l l produced w r e r e l a t i v e l y of smaller
s i s e * howavar. t h © i r mor?:Siology was almost tha samo as i n
c o n t r o l . Moroovert the prasenc« of s a v e r a l faurth s tago l a r v a s
and mature male i n c o r t i c a l and s t e l a r region wera noticed
{ F i g . 50u32j .
Inoculation with the t h r e e neniatixios resul ted i n the
production of sfoall^r female« and r e l a t i v e l y smaller giant c e l l s
having fewer nuclei i n each giant c o l l . Similarly the giant
c e l l s of il. reniforialg were produced i n c o r t i c a l and endodermal
region . Fourth s tage male larvae of ineoanita was associated
with degenerated c e l l i n the c o r t i c a l region. Female of
li* renifoTiitdp was r e l a t i v e l y smaller i n s i z e . A char act f*rii»tic
pyramidal sha » c e l l was observed i n the c o r t i c a l regiont formed
1 1 3
as a r e s u l t of dissolution of o d l i wall of 4 - 5 c n r t l c a l and 2
0ndodeiraai c e l l s (F ig . 5 4 - 5 6 ) ,
» f f » c t of concoHiitant 1 nfoction on tha fecundity of
4* ^ li- mlfsaBal i . *
I t i s c l o a r from the (TaHio 27) that on inoculatinq tho
tcffiiato setidlin^s with Sajspeciuen of inc jqni ta or vdth 333
sni^cimen of genifor^la alone the averaq® mirsibar of eqqs
j^cord<id aar femalff was 246 and 5 6 r e s isJctiVRly. Howevort vshon
th© seedlings were Inoculated with a mixture of two nematodes
(333 sjecirnen ©ach), tha average nuniber of eggs aach foniale
contained was raJucnJ and was iS5 arWl 44 of root-hnot and
raniform neiaatode raspec t iva ly . tho seedlings woro
inoculated with tho Piixtur«» of ii* renifogfais and brassieae
the averaga number of a^igs i'uix fsuala of reriifortii nematode was
60 and with the mixture of iQeffiUlUauarei^. kEasajUtfl&i th»
numbt r of eggs reccarded par female of incognita was 2CXD,
i^ith the a ix ture of three nematodes the average number of eqgs
contained per female of iiTeonnita was 141 as against 246 and
of T-aniformia was 38 as against 56 vi en inoculated s ingly .
XI . tifltjBMjm*
Oat a as jiven in tables 24» 2S and 26 dotaonstyate that
hatching of root-knot nematode i s suppressed vthen eggmasses are
suspended in d i s t i l l e d water along with SOJ itnaature females of
1 1 4
Tabia No. 24t Larval hatch of as influeno<»d by the
presence of yanlfogBij.fi and watar i n iwhieh
geiiifoPBia hds been suspended f o r twenty four
hours•
Treatments Hatching a f t « r different I n t e r v a l s (hrs )
6 12 24 48 72
S l t i f l e d V l e ^ 60 78 iD9 260 528
ogq masses ^ ii* m l £ 2 S 2 L _ Dis t i l led water ^ . m l i s x E U * 29 33 74 202 326
11
laasses Water of nematode 34 43 105 255 387 suspension
L.S.U. at level 6 2 . 8 1 2 9 . 7 2 11 .99 13 .59 4 4 . 0 3
L.a.D, at l e v e l 104 .10 4 9 . 2 6 19 .87 2 2 . 5 4 7 2 . 9 7
iiach figure i s based on two exi^ezioients with three r e p l i c a t e s each.
I l f ]
TabX& No, 25 i Larval hatch of incoQ^t^ 98 influenced lay
thd presence of X* pr&a&lcao and Water i n which
I.* brassicae has boon for ttmnty four
hours.
Traatm@nt8 6 IT 24 48 72
isgg masses U i s t i l l e d watar
18 27 39 8 i 152
ogg masses fafdllifiM »
Dis t i l led water 7 U 25 39 9 6
Hgg masses f* Water of nematode suspension
16 22 33 73 122
L.^.D. at level 1 3 . 8 4 2 . 6 1 3Q,6i 13 .84 1 3 . 8 4
L . S . a . at i ^ level 22 •96 4 . 3 3 5 0 . 5 4 2 2 . 9 3 2 2 . 8 3
liach figure i s based on two exoaximents with three r e o l i c a t e s each.
I I G
Tablo 261 £«arval hdtch of incognita as Influ^nc^d bff the
pJT&senco of a . and And
w«iter i n which they havo be&n suap»nda>J f o r twanty
four hours.
Treatrnants Hatching aft-^r diffarant i n t e r v a l s f h r s )
i 2 2 4 48 72
S l t i U e ^ w t t s r 64 8 4 X04 ^ S28
Hgg masses 1 R. SMliflXalE AA 29 i U 202
I« teagffipat» Uis t i l led watur
H99 aasses 4* wstar of nematode 25 39 69 i99 290 suspension
X'.^.P. <t level i 6 . 3 4 3 1 . 7 3 6 ,58 i3»89 6 0 . 3 6
L,S.U. at level 27 .06 32 .59 10«9i 2 2 . 9 5 100 .05
laach figure i s based on two experiments with three r e o l i c a t e s each
1 1 7
Table No, 271 Fecundity of S inceani ta and H.
influenced by concomitant inoculat ions .
Ini t ia l , inoculi^ iNto* of «qg» of of eggs of
' c " " 4 ' ' 'J'"' ••f am a l t .4b1. Hot, Tyl.
333 - 246
~ 3 : ^ - ~ 56
333 333 «* m 4 4
333 333 « 60
333 333 200
333 333 333 141 3 8 . 6
* MbI. m incoanitat W . » ' L* tefSiCfli
** Each figur«» represents the average nvmbex of ©gga per ten fenale*
118
renlfoxm naniatod® or in water i n icisature femalas wero
e a r l i e r susTondl^J for twanty four hours and l a t s r ramavad,
s imilar re -uct ion in hatchin':| «vas oijserved wh«?n aqior as»»» wore
kapt i n distiXl0d water along with stunrt rasmatode car in watar
i n tefhjch «tunt ns latoda has bean suspended, Hirjhest suppression
i n hatching was observed ss-hf n both ygnifanai^ and y .brasa icae
war® present toqather .
n n
In natura no ono kliid of organisms can l ive by i t s « l f ,
f t haa t o shar« tfidth others olth r throu jhout i t s l i f e cycXa or
a p.3rt of i t , ;icHaQ orgordsas a f f e c t oth.^r organisms i^i l©
growing saparately Ijut c lose tOvisther, a>iii in s t i l l othf;rs th®
two or^aiTism cl«vQlop cert^iin degree of orqanic union. In tho
f ia ld of phytonamatolociy ooth symoiotic and lacological r s l a -
tionship betwean naraatodes ami oth i r organisms aT& f a i r l y ccmaon.
Tho l i t ?ratura covarinj iiffafrent «ss;MCts has bean reviewed frc»n
time t o time (i^avsll, 1963, 1971 a & b| 1979} e t c h e r , 19^3,
1965, 1978, 4111 r , 1965| dergaason, 1972) .
jQSfiito tha fact that namcjtololists 3uch as Cobb (1914) ,
iljortw (i94B, 1957i , ^t(sim>r (1950) , uostenbrink g l a l . (1956) ,
Uostanbrlnk (1966) pointad out tha occurroiiCQ of a number of
genera of plant p. irasit ic n«;aatod#s and sav#r«il s i!»ci®s of the
saroe qenus In and around roots of cul t ivated plants , yat tha
studies on the intviraotion b«tW0©n diff«rent nematodes has
received l i t t l e a t tent ion , Cha xsian (lQ59)t ^ osa (i^>59) were
the f i r s t t o I n i t i a t e research on t h i s aspect of Hematology.
An atterapt has been made t o study the population, changes
i n sex r a t i o of mjiaatodes arnl dry weight of plant when 15 days
old seedlings ere inoculated with different ccmbination of
nematodes, besides e f f e c t of concocaitant inoculation on
1 2 0
ponetrirt^ion and resul t ing hiBtc}r>athoXo(jlcaX ch ^ngsfi dev«lo.3ln9
In r o o t s h«vo been s tudied. In addition t o thase aspects th«
e f f e c t of conqmoit^ftt inoculat ion on aoedlings grown i n s o i l
a landed with o i l cakas has been s tudied .
iihen tomato c v . Aarjlobe and eg^jplant cv» i^sa r»urtjle
Lanci were inoculated with ^ LnCmnU^t or g^„n3i,fgml8
2U brassisea^ singly a savara l fa ld increaso i n t h e f i n a l
t i o n of root-knot and raniform n9«iatod« has occured i n tcraato,
eq i jlant and not on caul i f lower , ito auch inertias® i n
^ooulation of X* bras s i c ac observed (Tibles I and 2 ) . Un
caulif lower* on t h e other hand^such an increase i n po'Wilation
o f only X, braasicatt ^as obsarved (Tabl© 3 i , lEtji-an^ incroas©
i n i n i t i a l po «J la t ian thara has baen a corrospondinc} increaso
i n the f i n a l population. ^Uant daraaqa has been s i g n i f i c a n t on
tomato and ® j?] ilant by root-knot and reniforra neraatode and only
marginal by stunt neiaatoda. un cauliflowrort on the othar hand,
root-knot n«a>atodo has fa i lad t o mult iply ; reniforra nematode
has baen abls t o maintain i t s i n i t i a l inoculum l e v e l , while
X* has multiplied ^arolifically* Heduction i n dry
weight has baen s i g n i f i c a n t as a r a s u l t of inoculat ion with
I* brasaicae and only marginal with InCftflnUfl and a«mtiL£2BalaL«
i^an seedlings ware inoculated with mixture of root-knot
an<i renifana nataatode, t h e rx^pulation of both root-knot and
renifoxm nematode has bean considerably reduced i r r e s p e c t i v e of
the di f ferent r a t i o s of t h e two nematodes i n mixture (Table i 4 2 ) ,
121
Thtt present studltts add t o our knawl<»dga regarding
various explanartions that hav« baen advancod front time t o tinm
t o explain tha reduction i n ?}OpuIation of mmatode resulting
from concomitant inoculations (lioss* JL964; i^stores and Ch tn*
1972; Turner and Qia man, 1972| CJia ^an and Tumar, 19751
:iingh, 1976 J ^arma and »>0thi, 19761 i ^ a i r and iJsfaan, 1977) ,
th« v a r i a t i a s of tomato and ai?r! ilant usad in the ^rss©nt
atuiiias are highly sascfi stiblo t o doth root-knot and rcmifojm
n«ia«atode ani se^dliniis at the tiraa of inoculation had i3©en
f a i r l y youn^, th^refora, in canccBaitaJit inoculation the larva®
of root-knot and i®aaturQ f^iaalw of r^^nifarra naniatod© tend t o
a<iar«.iat® in a limit^'d region of rootb as i f th»y have los t
t h a i r a o i l i t y t o s e l - c t sit© of th^ i r iienstration, vlonsoquantly
aithor Jua t o compatition or kind of ant r iiism i n i t i a l
pOfMJlation declintis and population at the tira© of inoculatii>n
does not remain aara® with vvhlch ona presu®»a t o have inoculated
th® seedlings. This prasumption i s supported ijy th® fact that
in studies dealing with concomitant inoculation the penetration
of both netaatodes i s reduced s ignif icant ly (Tables 16 and 2 0 ) ,
A close iroxinaity of infection court of the two nematode
xylem oarenchyma in root-knot (Chris t ie , 1936} Bird, i9T4j
41 ^ endtxiortais, pericycle and x/lera parenchyma in reni -
farro netaatode (Heald, 19791 Heoios aJLAl** 1975$ drathwaite and
Juncan, 1974| Fig.^g^J^^r^inces the possibi l i ty of exchange of
t o x i c metabolites from one feeding s i t e t o anothor feeding s i t e
122
which In f inal analysis como5in tha way in the ©stablishment
of noriaal host parasite relationship of the two noi^atodes.
«^roiuctian of tn^ftabolitas such as pho lol ics , amino acids by
riarilfoffn nematode (<lahmaodl, 1980) and phanaiics, amino acids,
XfV% and IBA iiy root-knot nc aal ode (Hasan, 1977i Owens and
>»petch, 1966? dalasubramalvfamand Hangaswami, l^GZt Viqiiarchio
and Vu, has basn rafJorttd, <uid th«y by and larg® arrest
tha develoisaent and iMsouldtion build up of n-?ioatoii©3 ( ^asad
and oet ty , 1974i 4ahraood, 1980K /vidltional ©vidance has been
provided by th« studi«s carr iad out by Haa in (1977) , %#so
studied that in roots dipiad in hyiroquinon fm larv«}@ ontf»rod
ani th© v liich have a ch-nxj® t o antor d9V«»lop into stiall
coiTtaininrj a fewer nufabt?r of ogqs. In such roots poly-
phanal oxidasa a c t i v i t y was ral^itivaly >oor*
/\n in<Jiriict ^^viJenca t o th'^ abova hy,>ath:^sia has b»«n
provided vnrfien histopathology of roots infect^^d with two n^^Kiatodos
has b«on studied, /to a conssquanee of concomitant inoculation
not only the host resiaonse had umlergon® changes but also the
two parasites in man/ respects with which roots have been
inoculated. The giant c e l l s of root-knot nematode in such roots
are smaller in s ize as i f coaelescing has been checked. Each
individual c e l l i s highly vacuolated and contain few nuclei .
Females produced jr® re la t ive ly smaller in size containing fewer
eqigs as aiainst tr-aatraents wh«re seedlings have been inoculated
by root-knot nematode alone (Fly . 47-49).;>irailarly the group of
123
giant c e l l s foimed oy reniform na^atode donot assume characteiv
i s t i c swailunar shale (Fig, 47*48} . Purthur production of
oyrataldai c e l l s in c o r t i c a l raglon In a J J i t i o n t o sndodaraitil
and 5^ricycl0 reijion at the feoding s i t e of renifoira neniatod©
(Fig . 56) havo been sean. uuch i9yra.nid«il c a l l s v^are rilso
observed by iiazak and livans (1976) t o which th«y at tr ibuted du©
tu presonca of raor ? than om? fa.nale at one infect ion s i t e , i t
i s iirasumed th«3t prosenca of larvae of root-knot a lso i n c i t e s
such a response, Heduction in fecundity of innaonlta i n the
prosancQ of calani (ohaxma, 1970) , oenotr-ina in the presence
of incoonita (Chaiaan an J Tumor, 1975) and reduction in
daveloiaaent of J . i i^oanit^ hdving vacuolated qi.int. c a l l in
presence of H. reniformis (uteifa and usman, 1974; v)taifa at a l . .
1974) has a i r e idy bracm reported.
In raniforTi-stanrt nematode Ciimbination, the copulation
of former has incraasaJ almost at tb« sasna r a t e as i n sirK^le
inoculation ts?st, wht?rs»as the population of l a t t a r has declined.
Ih® nonreduction in the population of H. ranlfortala in th ? combi-
nation can be ex plained by the fact that the two hosts are
congenial, and has >roJuc«d char - ic ter is t ic semilunar qrou i of
qiant c e l l (Fig , 5 3 ) and also th«? fac t that X.* brassic.ie has
failad to adversely affect the fecundity and i t s ^senetratlon in o
the roots (Tables 17, I and 2 7 ) . The decline in po;Julation of
X* bragalc.ja may be du® to the fact that the two host are not
congenial and the reniforoi nematode moves int c e l l u l a r l y causinn
1 2 4
disruption of c o r t i c a l c e l l s (Ficj. 4 4 j , ai^ th ?re by mducing
i t s successful host isarasitQ r»»l t t ionshia .
In root-knot stunt na iatode canliination, tha population
of th^ form ar has canjidjrably reducs»d d&s Jita f a c t th,3t
both tom^jto and agqplant ar& good hoDt f o r t h i s naniatode orasum-
ably because tha presanc© of stunt neiJdtoie t-jdvorsely o f f s e t s
the hatchinq ( labia 2 5 ) , ponetration ( l a b l a s IB and 2 2 ) ,
facundity (Table 27) and davalOi^ient of fe.i^'las. Thf prest^nce
of several fourth stage larva© (Fig , 5U,5X> i n addition t o rnaturo
fsaales i n such a s i tuat ion inwide aidit iondl prwof t o t h i s
contantion. Th© reduction in fscundity of incognita in
pr^saixs of Qal^ratus (Y^m gL d l . , 1976 ; and it.
(Jchmidt and i-ewis, 1981) has also been ra iortad, Schmidt and
Lswis (1981) also ra;xjrtad that fewar larvae of incoQiTitd
entered and raach#d u rto maturity i n presence of H. coluinbus
in cotton roots anJ also the giant c e l l s b^Jcaie necrotic and
l a t e r abort ic . The rcjduction i n population of endoparasites i n
coiicomitant inoculation by var ie ty of nematodes has been
reported by a number of workers (Chafwaan, 1959; Gureout, 1968;
oikora ^ 1972, 1979; Amo»u and Taylor, 1974; r»inochet
2 1 j i . , 1976; iAisra and Jan, 1977; Khan a l i i . . . 1978) . These
are, however, at variance with tha findings of Johnson and
iMusbaum (1968) , Toberiiimnaz {x973) , aanto and Jolander (1977) ,
*/aishnav jnd ^ethi ( i 9 7 8 ) who reported an increase in population
of one cohabitant.
1 2 5
Un c a u i l f l o v . j r , on tha oth-sr hanJ, thare has been a
m.iriced incrocise i n t.ha JODulation of sic^^ i n a l l t h a
treatrnants anJ a correa x>nJint| dacre aso i n t h e ;30.xil<3tion of
root-knot anJ reniform neaatod© (Table This i s unJerst^n-
dabla ainca tha vari^aty aar ly Jnowtjall i a a >oor h^st for the
two enJooarasittJs . Xhtfse rtisults sraihasiza t h j importance of
ho2t a f f a c t i n auch s t u d i a s . Tha r o l a of host s u i t a b i l i t y on
populdition chjnqas and ^jar^jtration i n concomitant i n o c u l i t i o n
studies has baen em phasized by jdy and dird <1973J, Turner and
Jhapman ( 1 9 7 2 ) .
The difforcnc33 in t h " J^ita as iiv.' n i n t^iolos
(4^7 I- ' .^ojulaticn from the data as given i n another
t a b l a s ( l , 2 a ) can be a t t r i b u t e d t o t h e fiict that tb^jso
studias carri-rdi out when glasi. h<ru;;.a t a rjarat-urt? wjs low,
IffiG data . j a r t a i n i n t ^ t o senuential inoculat ion f a s t ravoal
that by and lars^e g r a a t a r reduction i n the jo julation of the
neuiatodes has occurred with which the ssedlinqs h.iv® been
inoculated subsequently. This has oean t r u e in a l l the combi-
nations t a s t e d (Tablas 4 - 7 ) es jec ia l ly i4.incoQnita - H.reniformis
combination (Table 4}, This reduction occurs jrssij^iably duo t o
s p a t i a l and temjoral f a c t o r s which o::>erate i n such a s i t u a t i o n .
The former because the n^Jniatode with which seodlincjs have been
inoculated e a r l i e r induced c e r t a i n oiochomical and physioloqical
changes which may not be favourable f o r nematode with which
162
saadlings havo bean inocularted subaequotttlyi l a t e r bscause the
time f a l l s short t h a t i s required for caaple t ion of i t s l i f e
cyclo*
For ©xUorini] tha charges induced i n such a s i t u a t i o n
as s ta ted above his topatholo^icai s tudies are i n progress* These
r e s u l t s confirm t h e f indit^s of Oay and 3ird ( 1 9 7 3 ) , 4c Intyj:^
and ( f i l l e r ( i976)» idnochot ( 1 9 7 6 ) , and iiharma and
iiethi (1976) teflio a lso ob5i?rvQd t h a t pr ior invasion of seedlings
by one nematode reduced the entry or pOi»Jlation of ne^ iatode
inoculated subseouently. J a t a l a and Jensen (1976) reported
thdt when acf^achtif inoculated prior t o inconnl^ta. the f '
copulation of ^rKsoanit^ declined i n reverse s i t u a t i o n , on
the oth ' r hand, c y s t production of H, schacht^i increased .
^ interL'Sting example of bio predisposit ion has been
provided by Powell ij^ ^a , (1971.), who observed t h a t Trichoderma
sp. and Curvularia sp. ify iich are normally incapable of inducing
disease i n tobacco r o o t s , however, they cause axtansive damage
In roots already infact«i^d with root-knot nenatoda. Such a bio-
predisposit ion has not bean observed i n the present study both
i n concomitant and sequential disease complexes. There however,
has been reduction i n population and increase i n sex r a t i o .
s tudies dealinq with concomitant inoculat ion of seedlirv^s
grown i n s o i l amended with o i l cake and thimet reveal t h a t , there
i s several fold r e J u c t i o n i n the population of d i f ferent nematodes
1 2 7
i r r e s p e c t i v e of tha combination t -'stoi incluJinq whiiro the
seflKilinqs inoculatad with Jith r of the n aatoJessinqly as well
as in combination arrJ a corres ion Jinn incredis^? in plrint ^veinht
and pi'oduction of rualss than in the trt3<3t iont3 vvhera ssadlinffs
hjve aaen nrotm in unamanJlad aoi l (fables i i - JS) . This i s
unJarstandabla as i n i t i a l poiiulatian of tha n i-'toJss in
Ruestion iu reduceii in the rhizos "Jh -'re of root:i Qartiy Jue t o
k i l l i n j effuct of coiu )QunJ3 r-^lejsed as a resul t of oi l -cakes
or Uua t o n?Tu«ticiJ<3l a f f w t af thim'Jt (^injh and oitararaaiah,
1970, 197JJ ^iddiqi, i96-j!| r^dn, 1971 i and fiartly due t o r o a i s -
tdfica which jlant iavdlop as a resul t of >ib?iorjjtion of i®,]radinrj
products of o i l cakes (rJian, 1981 J .
vihan saedlinqs v.ara inoculat,jd, simultaneously or
sequentially, ha uex r a t i o of thi5 noiiidtoles used in the
mixture unJt?rgoos changes and thi-TJ i s .3 marked tendency of
production of lora malas (Taoles 1-7^. This e f f e c t i s more
pronounced in sequential t a s t s than in corwomitant inoculation
t e s t (Tables 4 , 6 and I t i a , therefore , concluded th.it due
to ir ior infect ion a i p r o j r i a t e nutrients are not available ,
consenuently th^-re ia a sh i f t in sex r a t i o . Jut of the
different exjeriaients radde,hiqhuit reduction in sex r a t i o has
taken place in seedlinjs which have been grown in - o i l aojended
with o i l cakas or thimet (Taoles 8 - 1 5 ) . in »uch a s i tuat ion
nonavailability of fooi and s t ress e;<jrtHd by decora Position
oroducts are both res jonaible for disbalancinq the sex r a t i o .
128
Thosa atudias t h 0 r«foro, for thef f i r s t tiiae >irovide avidtnco
that i n addition t o various raasons liven above with reaard t o
role of o i l cal««s i n roJucing the .lopuiation, the di«tur£ianc«
of SOX r a t i o i s aqually important. Thasa findin-is are i n agrea-
raant with thoso of Tyler (X933), iillenoy ( i 9 5 4 ; , TraintaiJj yllou
( i 9 6 0 ) , Glura and Viyiierchio (1966 ) , Trudgill (1967) , Javide.
and TraintaiJhyllou (1967 a , b ) , K^jestan (1969)» Hoss and
Trudgill (1969) , i4ian (1972) , ivoUopanoa and Trainta{:#iyUou
(1972) , Beria Sl, ( 1974) , who observed that s t j^sses due t o
unftTVourable environfuent hav® a masculanizing e f f e c t on nsraatod®.
Th© fjresonco of fourth atti^io larvae^ of malas and ©von
mature males in sect ions cut also coroborates t h i s f a c t (Fiq.
52 & 5 5 ) .
J 2 9
d f g ^ tj g c §
/tooau, J . U . and Taylor ( 1 9 7 4 } , I n t e r a c t i o n of Meloidoov^ haaXAt .^jratY^^nghna. M m t g a n a ^rui jmL on Kanland red clovar» h U M m Brat?nff« Jialjl.ftn i t o H L a l - . A24-131.
Atanasoff, a , ( 1 9 2 3 T h e .J^loahoaohor^ J l s a a s e of c e r e a l s . ^ v t o o a t h * . 1 1 - 4 0 .
/^kinson, vj,F. ( 1 8 9 2 i i o r t i ® disoases of c o t t o n , /^^bama. t^olvtgch. I p a t . ojsi^l. Aflt j m * i io. 4 1 , 61*65 .
•Jalakhina, sf.t". (1V79K Tba a f f e c t of ^i^alenehua avarue and Aahq^enchoides gatfy^fthjUMa on a a i r i l M fiJ^ffig^IIIB, i n f e c t i o n on sprinq v^04t . In i-»rintsipy i raathsody izucnaniya Viealmootnoshanii rnezhdu para atiticheskimi nsiaatodasai i rastardy<£ai Tar tu . U . ^ . a . a . , Akadaniiya iiauk Ustonskol aiiii JJUbX. 1979 J 144-148 (HuP Vl. t lJ UiGQW. U^H,
Jalasuisramaniaia, A, ar^J Hangaswatai ( l 9 6 2 j . of indole cotn rounds i n nuyatodo g a l l s , .^turt^ London, 1941 7 7 4 - 7 7 5 .
•Jalilwin, J . i . ( 1 9 7 8 ) . UiafTt c e l l s induced by na^natodes of Hatarodaridae, i^ia ta logy c i r c u l a r , divis ion of Plant Industry . iJeptt, of Agricultura and ConsiMiar 1978, 3 6 . iiiauraau of Me.^atolof|y, t^.v). 3ox 1269t Gainsvi l la ,
•sJartjasson, ( 1 9 7 2 ) . Concapt of nematoda fungus a s s o c i a t i o n i n >lant disease comalaxasi A r&viaw. a x a t . P a r a a i t o l .
3 0 1 - 4 1 4 .
•Barcja, J , 3 . , A. Jaliaassof and M.I, R i t t a r ( 1 9 7 4 ) , Influanca of tha host on ciavalopmarrt and sax datajcinaination i n plant p a r a s i t i c namatodas HiPii* Sgfflg^ffi,* ModUE ^SUSM.*
3hatnagar, A. ; T.K, lAukarjaa and 3 , Tiagi ( 1 9 7 9 ) , I s o l a t i o n and study of cjiant c a l l s and aonorraal xylara i n r<M>t g a l l s i n c i t a d by J . inc9ianlt,a i n liiilLia JL* m i a l a l « » J 2 . t
i i r c h f i a l d , w. ( 1 9 6 2 ) , Host paraa i ta r o l a t i o n s of |^otYl<>neht^lua ranlfoHnlfl on ^gtgYf^UB blBUtiaffi. i^Ytflga^^l» 862-865 .
130
a i r c h f i e l d . V*, ( i 9 7 2 i . J i f ferancas i n host c e l l res >onses t o renlfom naniatod®* i%vtooath.> 747 (Abs.) ,
• i i r d , A.F. <1972 j . Ceil wall braak-down durinq the formation of syncytia induced i n plants by root-knot nanatodes.
(1973) . ub8«»rvatioii on chrcxnosotrtes and nucleoli i n syncytia induced by SmMsA' ^ m i S i k ' Pfltiml. a* 387-391.
(1974) , ^ larrt resjoonse t o rocst-knot nematodes, Ann. Hey. ^^hytoaath..^t 69-^5.
(1979)» Histopathology and f^/siology of syncytia . i n I Hoot-Knot namatodes ( teloidoovne s n a . ) . uystsRiatiCB, biolotjy and control , ^icadsmic London, U.K. and j4aw Vork, pj , 155-170,
i^rdt and w.ti. Jenkins (1964) . Jccurrance. i>arasitism and pdthou macity of nu ff-stades associated v^ith cransharry,
Sl» 677-688,
J j.Ju. iJrooks and C.u, x^iry ( i 9 7 4 ) . Jynamics of concomitant f iald populations of c,9iVB?»Ua and lOSLaaOlla. si. .I^Hit,,!?!* t 190-195.
3oosdlis , and H, 4dnkau (1965) , ^^'arasitisni and oredation of so i l microorganisnj pp, 374-391 . Ini K.F, Jaker and V4.C, anyder, udits ,* bcology of doil borne plant pathoqens, Univ. Calif* t 'ress, aerkelay, C a l i f ,
• Brathwaitd, C, tit. 1} • and u« J . iXincan (1974) . Javelopment and histopathology of Ji. renl formic in sws^t potato,
imsSm^* 437-441 .
* Sreseski, M.W, ( i 9 7 0 ) . fha intarrelat ionships of newiatodes and other pathogiens in plant diseases, Paras i to l . 561 509-513,
• Carne, (1926) , carcockla (MjtoChlUl JUCLiifil) «nd bac ter ia l flftfl^monii t r t U g j , of wheat. I .
Viieat i ^ s t . (S^r. 2 ) 508 ,
1 3 1
Chapman, H.A. (1959) . iJevalopmont of ^atvlenchus ponetrana and Tv lonehorhvnehua qartinl on red clov ^r and a l f ^ f a .
357-359 .
(1966) . Infaction of slngla root system by larvae of two concid»nt so«k;10S of root-knot nejiatodes.
and J .H. Xurnsr (1972) . affact of entrant .^loldoavna incognita on ra aroduction of concoraitafTt Z S H i S u a ii^rFtgiaoa, c X o v r . i,. jSasatal* 221 ( / ^ s . ) .
and (197S). t f f a c t of ^loidoayna InccxaMta on re production of i " rad clover and a l f a l f a , Namatol. X* 6 - 1 0 .
C3"iristie, J .H. (1936) . The devalojiraent of root-knot nematode t jal ls . j^vtopath. . gjgiJ 1 - 2 2 ,
*Gobb, tJ.A. (1914) , J Jasnatodas and t h e i r r«?lritionships. yaar|3, U,^. l^tBtt,. iaEife. 19141 pp. 457-490.
(1918) . risinatodos of the slow sand f i l t e r e d bads of ftaiarican c i t i o s . Contributions t o a acl»mce of ffematalogy, m , pp. 189-212.
^ (l92i)) TransfarorKs of namatodos (mononchs) from plac« t o placd for economic r^rposos. science n . s . 5 l ( l 3 3 0 ) » 640-641.
Cohn, li. (1973) . Histology of the faading s i t s of Hotvler^hulus m l i f l X S U . I Kin tglftq Cfl, iSL< 455-458,
(1976) , Cellular changoa induced by two s iecles of th« J»nus ag^yl nghMAiUa. fiiffffl^^QiMifiJt i l * 169-173,
and M, <^rd«chai (1977) , Uninucleate ^jiant c e l l induced in soybean by nematode, Mflfig'y^Wfll&yi'
B5.93 .
Javide, and h ,C , Traintaphyllou (1967a) , Influence of the anvironroent on developnidnt and sex different ia t ion of root-knot netnatod«t, I . iiffect of infect ion density, age of host ilant and so i l temperature, NamatoloQiea. Jj^t 102-110,
I ^ o 1 t^'CL
iiavid0( H.U, and .uC. Tralnta{^yllou (i967b)« Influence of the environment on davalopraont and sex d i f f e r e n t i a t i o n of root-iknot nematodes, I I . e f f e c t of host n u t r i t i o n .
•uropkin, V.H, (1955) . Tho r e l a t i o n s between nwiaatodes and plants , ^ x o t i . i i a m l ^ a l . e 2 8 2 - 3 2 2 .
(1969) , Cel lular resijonses of plants t o neiiatode® i n f e c t i o n s , fin<i. Hey, ^ v t o - i a t h . . Z* 101-122,
and itelson (196U), The histopatholoqy of root-knot rwmatode inflictions i n soy jeans, /^vtopath. 221 442-447 .
argl vi.ii, 3oone (1966 ) , Analysis of host a a r a s l t e ralat ionships of root-knot nematodes by sinqle larvae inoculations of oxcised tcHnato r o o t s . i%^atoloaica.
225-236 .
allenby, C. (19S4) . sinvirorenttrital Uetamiinotion of the sex r a t i o of a slant p a r a s i t i c nematode, iiature l4>ndon, i H * 1U16-1017.
tindo, d . y . (1971) , Neaitode induced syncytia (giant c e l l s ) . Host oarasi te rtJl itionships of Hettjroderidae, Int ^'arasitic nematodes e d t s . J . 4. Zuckernan, W.F. 4ai , a .A, i<hoda, 91*»'117f iMew York, Academic j^ress* 347 pp,
(1975) , '^athoganasis of nera a''ode-infected plants . iM,* dticltittati!.! J^f 213-238 .
•ttsser, H,r», (1968) . Nematode i n t a r a c t i o n s in plates of non-s t a r l l e watar agar . **roc. ai sLL dQii itfXail* jfil*» ifiS.* fIS.*
121-138.
•tistores, H.A. (1971) . Int^^ractlon of and l^eloidQc J i a s . Abs. Int
Interac t ion of ^r^tyA^'nchui g?.notgarui S. i g ^ ^ ^ ^ t a s ^ o i w a b i t a n t s in tomato.
and T.A. Chen ( 1 9 7 2 ) . Interact ions of ^atvlenchus g^n^jgani ingfaflntia coinhabitants in tomato. Nematol.. 4 i 17D-174.
F e r r i s , V . a . j Ferr is and H,L. aornard (1967) . Relative competitiveness of two spei competitiveness of two species of *-*ratvlQnchua in soybean. . /.
13.1
•Fioliling, M.J, ( 1 9 5 9 ) . Neraatodas i n plant ( l lseasas. Ann. m c r o b i o l . . 2 3 9 - 2 5 4 .
Fortnuia, and Lewis ( 1 9 7 8 ) . Intoir«stion betw»en Qylindrocladium root ro t and lolys w c i f i c nematode popula-t i o n s on soybean, m h m a * dLtiEltB MSk' £!sadL. tiafilmsL. ^.Hamgait m q . JL97e. i a » 287 (Aba. ) .
Frockraan, J . i* . and CSiaioraan, ( 1 9 7 2 ) , Infec t ion of red c l o v e r secjdUngs by Heteroviera t r i f o l i i Oof f a r t and
il^fflttfia^,., 2 3 - 2 8 ,
Gay* C,.'!, and a i r d , ( 1 9 7 3 ) . Influence of cinccmdtant ^dtvlanchus ^nd >3>9idQaynft 3pp, on root jenatration and population dynamics, - i t o l . ,
2 1 2 - 2 1 7 .
"Oldanf J.i^. and o . J , i/an tiundy ( 1 9 7 2 ) . Influence of L i neon ni t a on root davelOixaant by ^Mgoff fflrtl m X m L
i n tomato. 2 2 5 .
G r i f f i n , ( 1 9 7 6 ) . dffect of tcj-i-K^rature on root-knot and stssia nwiidtode i n t e r s c t i o n on a l f a l f a , JssjL* 1576 .iba, ^a. 14 , ^jp,^,, tfkiXm^^k* tiStSi' a* 3J-7.
nd H, 4, Uarling, ( 1 9 6 4 ) , wcological study of Xiijhinoraa amaricanmB Cobb, i n an ornamantal saruc« nuraary, iSl* ^71,
•liuorout, H, ( 1 9 6 8 ) , Comoqtition Pratylenchua brachvurua i^loidQavna n. sp. dans l a s cul tures of anasde c e t « d* l w o i r « p 7 4 6 4 - 6 8 9 . In a coianans, 4.T, Franklint H i t t o r and .3, »»oi8Ch«r («d. ) i i iaht . I n t . jvmp. .^matol. ^Ql* Francs,
>iupta» and G, C^arup, ( 1 9 6 8 ) , On the e a r cockle and yellow s a r rot of I , Symptoms and his t o anthology, m x m d i i i s m ^ ' * 3 1 8 - 3 2 3 .
•Harr ison, a,D, j i4owat, and G,£ . Taylor , ( 1 9 6 1 ) , Transmi-ssion of a s t r a i n of tomato black ring vi rus by k^naigglVf fflgrwfltm ( ^ m a t o d a ) . Vir^looy J ^ t 4 8 0 - 4 8 5 .
1 3 4
Hasan, N. (i.977). uffact of o i l cakes sawdust and c e r t a i n chat^caX compounds on tho development of root-knot on eggplant and tomato. ^ . J . Thesis, J e p t t . of Botany, iUigarh Muslim University, aliqaxh.
and /Uaia, (1979) , Studies on the influanc® of WlQidooYi^e incognita on the iaoxilation of indicua around tomato r o o t s , Ueobios. X2i-i22«
Hawn» - . J . and 'Ud, Hanna, (1967) . Influence of starfj nonatode inf istation on bac ter ia l wilt reaction and foraa® yield of a l f a l f a . Can. J . tifi. • 203-208.
Meald, C. U (1975J. i'athogenicity and histopatholorjy of inflicting cantaloups. I ,
I t 149-152.
Hechl^r, H.G. (i.963), iJescription, develofxnarjtal bioloqy and feeding habits of .^ffjnvga (J8 4an), J . Uooiey, I960 (i^jematodalai^fiflonchoidida©), a nomatode nredatox. diB2$6» Helm, ^oc. 182-195.
and J . P . Taylor ( 1 9 6 6 ; . Tha l i f e h i s t o r i e s of rffifflgd saliZiUt ii* smim i^roc. iigia« ^QC' wash, g^s 7 1 - 8 3 .
Hdwitt, I J , J , Haski and a .C . Goheen, (195B), .te^atode vector of soil-5orne fanlaaf virus of grapevines, ^ g p a t h . > 5tio-595.
^Hirano, K. (i.979). Interrelat ionship between plant p a r a s i t i c nematodes and other plant pathogenic organisms, ^ev. PJLjML. ilftL. a m . 5 3 - 6 8 .
Huang, C.^. and A.B. Aagqenti, (1969) . Mitotic 'b>rr ; ' i jns and nuclear changes of developing giant c e l l s in ^/ipia faba. caused by root-knot nematode, i a M d a g n i f t idiXMca..
ytoaath** 52.» 447-455. * Hunger, P.W.T. (1901) . Hen bact«r ie -x iekte den tcKaaat. S.
Lands. **lant. /^ded. . 481 4 - 5 7 . ""
J a t a l a , and H.J . Jensen (1972) . Interrelationships of id* hasdA ffchachtlil populations on M a . xuiauttU. i . ijfiuaaifll.s 226.
35
J a t a l a , and H,J , Jensen, (X'i76h ^elf i n t ^rafCtlona of ^loidoavfia hflflla and t/n^-rQ^tfi S^fl^flUl on ,^tft vulaarla . J . 4^-48.
and • (1976h i'arasitistn of Beta vulgaris
a* 2(X3-a05,
— a n d ( i 9 7 6 i . fiistooatholo^y of Beta ii^laarls t o Individual and cancomitant infastions by
and [Mffyg. ^iCa s^hOffMil* tl« SEiSfll* t i 336 :341 :
Johans0n, J ,A, ( i 9 4 0 i , -'lant .licrotachni<i|u©, fcO^raw Hil l , Jook CcKQpany* NefW York and 1-ondon.
Johnsin, (1969) . ^^athcHiunicity .=ind ;x»xilttion davelapwj 'nt of oyn^uA. I . aajcljUiUA and 3,1.1909,1.g^^wa lenaicaudatus, singly anJ coiminedon bIx b^jSuJagrass .
mmtQl>> I t 294 <Ab»J
(1970 h ^^dthoqsnicity and tnt . rac t ion of thrao nematode specias on six b«Jrrnucid grasses , rferoatol..
36-41 ,
and C . J . / Jsbaum (1968) . The ac t iv i ty of iti sbJClsils:. ^
, _ . Lations on com and soyjean. lik^fltqlgQ^Cdt Id* 9 (Aba.),
• (1970) , Intoraction between incognita. MiMvfiirm tmls. ^nd
brachnuruM in tobacco. J . 2,1 334-340 .
*Jon«a, i4,G.K. and V.H, Oropkin (1975) , Cellular a l terat ions induced in soybeans by three endooaraaitic nematodes.
^gi^h?!.* 119-124 .
Ksrstan, U. (1969) . J i e 3eeinflussung des Goschlchchtrvefhatni-sses in der iattung F^oterodera. i l , 'ftniwallbensarum -selektive abater berate der geschlechtarg^ochlechter-verhaltnis (Heterodeya s c h a c h t i i ) . .>te?8atQloQica. jj^t 210 .228 .
i 3 G
iOiair, arKi a .A, Umiaits (1977) . Interac t ion of ^ l o l ir>«wnQ lng9ian|,td ^^ r^Mfffr^a on tomato.
Kharii 4U A* ( i 9 7 i ) . otudi«» on plant p a r a s i t i c nejaatodas associated ivith vafjataiile crops i n Uttar ^ a J o s h . Final Technical iirjport. Jejijtt, of Jcjtany, Aligarh 41^11® Univarsity, /Uigdrh,
(1981 )• Control of diaeasas caused by m»aatad#s by a.Jplicatian of oi l -cako laaJiures. Final Technical Ha )ort J « p t t , of J o t any, .Uio ith ^kjslim University, Aligarh.
iChan, F, (1972 j , Jitudi«s on the reniform nctniatod®, Uotvlet^hulus raniforiaia ii.nford and J l i v a i r a and on th© taxancxay of c e r t a i n other Tyienchids. Thi'sis, fiaptt, of iotany, iUigarh wiasllm University, «.li jaxtt,
l>han. It, 4 , , ftao, Jaxena, and i3ian ( i91B) , iiffect of coinhabitinq pooulations of .^aloidooyng irysoanita and
Mgafii>.g,af, on t h o i r mult ipl icat ion on tondto. Indian J . QJ
iOian, H, 4 , , It. 4, Khan, a.K, JaKsna and ;vltan (1979) . ilffoct of cor« omit ant population of .4eloiioavni» incognita and flg YAinffhMAMa, on t h a l r myl t ipl ica t ion . jfMqtp,
<\hdn, and (3. Tiaqi (1977) . ulant c e l l s in Laciarmaria lt»cunth.a rusby inci tad by tha root-knot n^matods, ^^lo^dpgyna 4rK?qnitfl (Kofoid & Mhit«)ChitMood. QffsUm SH 31
Ninloch, and /illon (1972j . In terac t ion of %loidoqvno haiilA and ,iaYarU>CJ Infactlnq tomato. JNjflaaLBJL.. 7 . 1 6 .
i^llopanoa, G, and IraintaF:4iyllou (1972) . e f fec t of infect ion density on »ax r a t i o of HatarodTa olveinga.
UL* 131 .137 .
•Kostoff , J . and J . tiandall ( ly3U). Cytolotjy of neraatoda q a l l i on <\aeatiana £pots . ^aa&jcali!!- iais^ajclfll* ^ar^igAtwnk* Hva. Abt. I I ^ t 86 -91 .
Jo ^ o 7
Krishnappdt r%» and v>arsad (1979 effec t of concoraltant inoculations of TYA^nc^ffirhYnrrNa ^kMm ^nd
on th<»ir oopulation build up on tomato, Mysore
«\ru3ii«rg, L.H, (1939 Xnv' s t iqations on th« 11 f» c y c l e , ra aroduction, feeding hatJits and host ranga of Ty nchurt Yighyi fiiaytgnl, .teTitoioa c.a. it 187-197.
(1963) . Host ras Jonse t o nanatoda i n f ^ t i o n . J b m M t i L . 219-240.
idbraanf U . J . j U, juaach and »V-lafa3 (1964^. :tol«3 of filant par^isitlc nea^itodas i n infect ion of tomatoes by ,>%eqd«Mionas .solan^apym, 151-153.
a n f a r d , -UJ. and J . 4. u l iv^rira {i94Q), liotvlgiichjIus t^qifogml^. nov. gi3n. n. s p . , a nematode wrasi ta of r o o t s , Hr:>c. Helni.
m h * X* 3 5 - 4 1 .
Uicas, iild, , J , J, ^ s s e r tjnd A, tCeliaan (1^54). Th«? afff?ct of root-knot nR.iatodgs on thf» axixressi^n of g n i n v i l l e wilt rasiatdnce. .^tosa^th, , 497 .
fiahmood, I , ( l 9 6 0 i , ^atudies on cart . i in aspocts of pathoqeniclty V of reniforfn maatod®, liotylenehulus ranifor^l^ on tomato
and eqgolant. Thasia, iJaptt. of dotany, Aligaxh Auslim University, Aligarh.
•.4alok, and Jvnkins (1964j , ^ispacts of hast parasite rdlationships of nomatodes and hairy vetch, a u l l . Ng^a Jersey iiaUC' iiSBl* ^ t a . . H131 31 p.
•Mankau, rt. and M.J. Llnford (1960) . Host parasite relationships of clov<»r cyst neraatode, Hglggodftfi t r t f o U j CJoff. M X ULL* M.* JSaSL* siiuA* JhjUUL* s^*
• lartin, G.C, (1978), i>k>t»s on Pratvlanehua oenetrana and othar ondoparasitic plant nematodes in re la t ion t o tubers of •iQlanum tuaerosum in ^outh^rn Aferica. 3rd Cong. acw;. Msk* M V l B i Z E y y u : W
vteaeth, C.^x.j A.L. Taylor and .\.L. jtaith (1941) . states on staining n^sndtodes in root t i s s u e s , ^ o c « tifiJLS* iSSL* SiSSll* 2 6 .
138
/^Clura, M.A,, and J.H. Vigliarchlo ( i 9 6 6 ) . The influenc® of host nutrition and intensi ty of infect ion on th« «ox r a t i o and development of i lgtt ffgyfTffi lOLaiinllaJLn s t e r i l e agar cultures of excised cucuaober roots , itenatoloaica. ^ t 248-238.
^ i ^ a l e y , i^.C. and U.A. Cha*aan (1976) , Javeloaiwjnt of corwoml-tant .copulation of HalicQtylaneh^« oseudorobustgg.
misfit MS and aj^Piilli on soybean. 296;
Iclntyre, J.Ju. and ^ 1. .Uller ( 1 9 7 6 C o i i i e t l t l v © interac t ion
in tobacco roots . ^vto ^atrv.. 1427-1430.
* ,Jillor» H.W. (1963) . Intu»rjction of nenatodes and other olant pathogens, i t e l * ^oil- isSaa- 310-32S.
.^ailort 4. (1970) . Hdte of increase of a low njorayldtion of t^kmum reduced by f'l^fizipm^m
s o i l . a s i z u . . b - 2 ^ .
and J . L . .fclntyr® (1975) . TvlG-xhorhynehua clavtonl feeding on tobacco roots re ucaa the entry of Pratvlenchus penetrans^ i tein^^ol. H ^27.
and ki/ii^aim (1968) . Iitual antagonism between Heterodera tabaeum and some other a a r a s i t i c nematodes.
J i a l Reptr. ^ t 57 -58 ,
4inton, N.A., and H.a, tALnton (1963) . Infection relationship between M* InCgqfU^i aSfflta ^nd Q ^ , 8 a o m f* ViMknfKXm in cotton, j f e ^ p a t h . .
««1inx, ci, and J . Strich-Harari (1959) . Inoculation experiment with a mixture of i^eloldocivne spp. on tomato r o o t s . Ktavioi. St» 275-279.
-4isra. C, and Jas (1977) . Interaction of sorae plant paras i t i c ne matodes on the root-knot development in b r i n j a l . Indian JL. iisaatfll. <6-53.
Aountain, ( 1 9 ^ ) . i^athogenesia by s o i l nematodes. Ini Ecology of ooil borne plant pathogens (Kdts. K.F, Saker and W.C. iinyder). Univ. of California ^ress Serkely pp. 283-301 .
139
Mountain, W.a, and G.O, -^iCaen (1962), uffact of \ /ar t ie l l l iua dahliao on the population of ^^rTtvlanehntt oenetrana^
and j^atrlck ( i 9 5 9 j . The pi?ach realant m prodleni in Untario VIZ, Tho pathoganicity of ^atvlartchu
i 9 i 7 ) Fiup, is, Jt9k, i 9 4 i . S o T Z T M .
Nath, J , Jwaawa and Hama jido ( i 9 6 9 ) , studies on the renif ul iverird the renifor a ne aatode^ iiotyl >tichulus ^iinif a m i s Linford and
t mkmSSEMST* 22.» 9 9 - i 0 4 .
t 4 , J . Haidajci i^htar and H. %rsad (1976) . ^tudias on tha ne iatodes of v»qetaales in B i h a r , ! , liffect of roni -form n^}.natodo, a . o" ^IffCla*
i M i a , A. and A, ^Uaj«ndaro, (JL976J. a f f a c t s of .^^^tyXgnchus coffaao and ^utaXloneaia bravdis alono and In canibin<itlon on Guinea Ym C^ioscpra^ yotadata ; . J. j^giaalaJL-t 3X5-317.
*,term?c, d, (1910 >. ' 'Jas ^ roblarn dar defruchtun^svox^anq» und amder® zytolooischo Fragan. Vx. Vielkarniqe mesenzellan i n H0terodera ia l len" . .Jp. 151-173, Uebru der dorntrager, 3orl in .
itorton, t».C. (1969) . %loidoavne haola as a f a c t o r in a l f a l f a d«clln® in Iowa. jHivtooath. 591 1824-1828,
,, (1978) , ideology of olant paras i t i c nematodes, JhorTiiiley and aonst York,
0*3annon, J .H,| J , J , Hadewaldi H,f , Tomerlin and ti, N. Insarra (1976) , Comparative influence >f RadojAolus airallia and Pratvlftnehu* Bftffejf. on c i t r u s , i T S S E ^ f T s S - ^ ,
oostanbrink, M, ( i 9 6 0 ) , jr'opulation dynatoics in re la t ion t o cropping* manuring and s o i l disinfect ion. IniNeniatology (Hdt,, Sasser, J .N, and i^.R, Jenkins), Univ, >lorth Carolina <*r9Si, oo, 439-442,
(1966) . .4aJor c h a r a c t e r i s t i c s of re la t ion between nenatode and plants . ^ •ffQtnl.na nt
1-46.
J 4 0
Oosteniarinkf l a J . J . s*Jacob and k , Kuipar (1956j , nn intc»r-pretation of some crop rotat ion ox.v^riences based on namatod© aurvoya and xi Julartion studies, ffe-natoXoaie^ ^t 202->215*
u t a l f a , and a, Usiian ( i 9 7 4 h Host par is l ta relationships of Hotvi«^nchijlus on Lypopfersjcffn C^bs,) ixa^asJU. AntflXtC^^/ kL j d m i a . ^ Jmalszr iQciQs. XfJ aaot . 1974; Uranada, a>ain, 23 pp.
and /A,/I. uaie i {1972), liolotjy and histopathogenesis of reniforui naaatod© itotvlenchulus reniforTiis on Bqyptlan cotton, iatiLaskOS.. r^YtOPfl f ' yntol* ^g^das .^rtuqai , 299-304 pp.
» t , . and Usman ( i 9 7 4 ) . Histoaathoqsno-s i s of and IficgqrAta a® coinhaoitants in roots of tom^o ^coograicon esculentum. ^iOi. y o u - ^airo, 2a.« 2 4 1 3 1 4 :
^CXsfans, 11.0. and H.^J. <ipotch (1964) . xioot-kiiot histofjanesis. ^ai^aik. M m toi.. W 47JL-489.
/snd . (i .966). diachoJicui ai t«rdtions inJucsd in host t i ssues ijy root-knot ne iatod©. Ckintrib. aovce Thotflpson. Xnst , . i 8 i - i 8 8 .
^ '"'aaz, J '4,a. -^cia and J . n , Meredith (jL976). (Individual and cofflbinad effect of 4 . InCoanilffl and UY.§niSA on four tobacco (j^d-CQtiana tabacum L . ; cult iVar i n Venezuela), tiff«cto individual Y cotabinado da 4 . incognita y J4, lavanica sobre cuatro c u l t i v a r s de tobacco tabacus^7 en. Venezuela. itefltlPglgJ* 68-76.
•s^eacock, F.C. (1956) . The reniforra nanatode in the Gold c o a s t . NetaatQloalcff. J^t 307-310.
flnochet, J . j J . J . Haski and A.C. lioheen (1976) . Effects of inde^ singly and ccKnbined
l i8iM2l . a* 330-335.
r l tch«r» (1963) . Hole of plant parasi t ic nematodes in bact -ir, U 9 6 3 ; . rtole of piant parasitK a r l a l dis<Jases. PhVtooath.. 22.; 35 -39 .
141
x^tchor» H. j , Xntarrelationships of nematodes and other pathogens of [>lants. Halm. ^iUastr. i.»X7.
. . (X978). l i t ter action of nautodes v dth oth<»r patho-qans. Int 31ant iiteajdtoXogy (udt, J . F , iiouthey) London, UK, H.4, s ta t ion , t i ff lce , p^. 63^77,
and J . a . Cross (1958) , atudies in tha r a l itionshlp of eeiworta and bacteria t o cart . i in ilant disaas08, I I , Furth'T andlysis of the s t r^iaerry cauiiflo»@r disaas® com 5lf4x, Naffjtoloaica. 2 4 4 - ^ 6 .
/ GWtf&Xi, i^.T, The roX® of iXdutt paras i t i c nematodes in fungus diseases, sfivtoaath.^ 2B-35,
(X97Xa), XiTtiJraction of aXant pardsi t ic nematodes vsrith oth ' r ilisaasa causing agtsnts. In J i^iant ^araaitic imaatoJss ( a d t , , ^duckarman, »«,F, .4aij and Hhode), .^ddenic *rf?83» 4 t w Yark and l^ndon, ^ol^ XI,pp, XX9~X36,
. (X97XbJ, Xntdraction bstweon ns latodes and fungi in disaaso c0?»i>Xax0S, ^na. iMX* *^vtoodth-. 2.J 253-274.
(1979), int&rnaX syn rqlsms diiong orgjnisnjs Inducin-a disease, In« i Xant tdsoasas ana Advanced treatis?? (iidts, , HorsfaXX, J , J and CowXinqJ, /«di9>nic rim V ork and London, VoX, X / , pj, XX3-X33,
J '-^X.L, 4eXdnd«z and C,*^, ia t ten (X97X)« Disedse complexes in tobacco involving MaloidooYnQ ineoonlta and c e r t a i n solX-oorne fungi, «f>ytopath,. X332-1337,
f^asad, and n.G.H, i»«tty (X974). Tho af fec t of plant
f roiwth substances as s o i l drench on root*knot nematode ^l9l49<xfm Ls&asailAi* mua i - . 152.154.
^»rasad, K.. and Hao (1977) , Interaction between TYlinchwhYnc^Mi ffliytani and H^4fi9tYignch.M,§, gamna^m. on r i c e , l o ^ i M I* SmSMU t 170,
Hazak, H,a, and A,A,F, livans (1976) . An intya^ellular tube associated with feeding by nattY^^IChuijli r^niformis on cowpea root , i>feiatnloqica. ^ t I82.»189.
M2
HaixLoB, H.V.I J . 4 . »ipps and ii.i:, Hartiwig (1970) , GorraXation of r«»8istanca in soybeans t o Heterodara alvcimis and
•'Mtwa^h,.* Wl* 695^700.
J 4ddden and i . J . kitlridg® (1975) . aome uXtra«-s t ructural changas induced in res is tant and suacoitibl® soybean roots f o l l >wing infec t ion by ftotvlenehulus raniformis, i i i todSfll . . X« A22-139,
iiaynolda, and 4. i. avans, (iyiV3). Tha a t y l a t namatod®, Tvlgnchorhvnchiia di^biua. a root oarasito of a c o n c ^ c i tanc» in Southwast. da i t r . 22* 940-544 .
Hoss, (1939) , int eraction of Aaloidoayi^t ir^coanita and ^lytopath., 49« 549 ,
U 9 6 4 ) . Interac t ion of alveiiies and ipoyng immnXl^g soyb»ans. f^ivtoaath.. J^J 304-307.
.. . ( I 9 6 5 i . ^Y0dis->ositioo of v>oyb(jaii t o Fusariuia wilt . ilYfiirei . and t m p a o l W £hxl8aaXli»
iioss, U . J , a , and J . L . frudijili ( i96 0 . Tha a f f e c t of iiopulation d«nsity on the sax r a t i o of Haterodara rostochlanaia - a two dimensional ^aod^l, i^matolQ^ica. 15i 601>.607.
Hoy, A.K, (1976) . Pathological a f f a c t s of ^^loidoovtig araminicola on r i c a and histopatholovjical studios on r i c « and maize, IfHttan dixlaaayiM 359^62.
aanto, ^.S. and dolandor (1977) . ;»9parate and concotaltant a f f a c t s of tmsi grapas. i . JijaadlajL-* 282-2 a f f a c t s of Mnmjllih and i . haalfl on concord
8 3 .
iicholdt, K..H, and ^ .n . Lewis (1961) . Jynaaics of concomitant populations of ^utellonema brachvui and :4ttloidO(ivna Ineoanit^ on cot ton. J . t^taatoI7 i ^ i 41-^
^iainhorst , J .w. (1961a) . r»lant-nenatode in ter re la t ionships , ^qq,. Sfijcaizlsl.. i77~i96.
N.K, (1970) . (as c i t e d by C.L, oethi in Complex r a l a t l o n i t o other pathogens - ^outh Hast /Viia i^ostgraduate Natnatology Course Lecture, Migarh 1979i*
14P>
Shaxnia* N.k. and C,L. ;iethl ( i ' 7 6 J . Xntaractlon betwoon ;4flgidffqYqtt Ififfff^ntta and SUtiLanl on cc^i^ea. Indian %mat.Q,l«t i - i a .
^ d d i a u i , /i.A. ( 1 9 6 9 ) . atudlas on Tvlenehorhvnchua brassicag uiddiqui, asbocidtad with the roots of cabbago and c a u l i -flower. ^^.iJ. ThiJSis, of riot any, Aligarh Musllra University, Allg arh•
;iiddlqul, h, i i shiJ j -U Yunus and U Uhuuse (1974) . atudits on reaction xyloi dc?v«?lo md djo to j^loldoavno lnc,gqnl^fl in the roots of ^^ygg'^ha.* MJlan d* Haia^iai.. 46 -52 ,
idkora, I J,r». Taylor, .lal ik and J . T , cdwrirda (1972) , Interaction of 49loldoovna n-aaalt penetrans <3nd Tvlgnchgyhvnchus dfflUSl on c rea )ing bent-grass. I . f^iaatol.. 4V l ^ - l ^ T ^
Mal&kt Taylor dnd iJ . I . udwajsls (1979 )• Haiuction of ^aloldoavnQ fiaasL Infection of craaaim? bont grass by Tvl^nch^y^n^nghW, m s L and !BlQS£*
ain^h, N.J. (1976) , Interact ion of j^Xg . qgynf, and /MyAengh^Atfff, ife.qAfffrwifi O-O soybean, ite^tfffipls^.^
jingh, H,ii, and K, ^tararaaiah (1970) , Uantrol of plant l a r a s l t i c nematodes with organic ainandments of s o i l . •'Ai 16 i 287-297.
and (1973) , Control of plant -parasi t ic nGi.itodes with orqanic amandraerjts of s o i l . Hasoarch aullet in
i ^ , d a i l . Jaiat, M y , oL mxXS, ^ I s d E S C T ^ a n t n a q a r ,
divakumart -and ii.ii, i»eshadil (1972) , Histajaathology of infection by the ranifjrra no latoda, r^tvlenchif^t^s rar^foipnis i4.n. ^ J l i v . , 1940 on c a s t o r , japaya and tomato. I n ^ a n
173-181,
iraolik, J . J , (1972) . jroduction of Tvle.lehorhvochua nudus and Halicotvlenchus laicaphalus on srarinci wheat and affect of X* on growth of spring wh^^at, fourth, pj k. /^ad. 1S3-159,
181
^ u t h y , J . F , (1970) . Laboratory methods for work with ulant and so i l nem<)tOiias. Tach. J u l l . g,, iijUjOa* ^ r i e . iXst,* food. Londoat 5 .
jtafetnan» i .G. dn«i J . tJ . i iarrar { i 9 5 7 } , ^Yincipals of Uant '^athology. fhe /Donald -^ess Ctorapany.
^tein>?rt '-J. (I'^Suj, ^-'lant nHridtolotiy ri?search in th« ^iureau of plant Industryt -joiis .and /Vjricuiturdl umjinsering, Ji . , J i s . il&n, 1951 463-470.
and H, Hainly (1922J, Tha possibil i ty of control
^^temerdinq, (1963 )i i:en mixer-watt^nfilter inathodeoea v r i j b e -weeqlijke end0;jarasitdire nematodan uit vvortela t e varxanaten. '/ors 1. . zi-3kt. Jlenst. waqenincien,j^$ 17(Jul7a.
Ta* at and ~»ultan il9T7), ^^opulation of 'iotvlenchulu^ rani for^is, and IVlaiyhulus a0CTicmn»i?trdns on grap# seedlinqs «s influenced by coincident infes ta t ion , i^tenatoloaia. ,4Qditerranaa. ^J 233-257.
and A. i . isassab {1979 Tbg histopatholoiical reac-tions of Viana sinensis t o s a jar at o and concomitant o^r,isitism bv ^Mloidoovne ^ nd iMxlSZlSlUlllUL ranifor-nia. J . NgrMfll-. iX^ iX7-123.
Thorne, 6 , (1927) . The l i f « history , habits and aconcxnic iaix>rtanc« of scwne mononchs. jJ. /Uir.Haa. 34t 2 ^ - 2 8 6 .
, (1948) . Nenatod«s as a d is t j rbaice factor in green-nouse, plot and f iald ax >jrir(5<?nts. J i s . ^ieptr. 22,z 473-475.
* (1957) . Plant paras i t ic njraatodas in s o i l biology, isss.' Mix. jfil* i ^ * il^
^ob/rjiranaa, A. (1973 K (isxoerinjijntal study of anta'^onisra b»tw««n plant >ara8itic nessatodas). Bstudio experimental sobra loss antagonism antra algunos namatodas aarasitos da vagatablas. ft^Ylj^i* ^ "ijfagUgjg^Agi ^ ^ 607-615.
* „ . ^^^ ^i^rnal (1977) . (studies on C^ulran 1967) . Hftvla^^
Oft W 191-203.
1 4 5
Triiifttaphyllou, /wC, ( i 9 6 0 ) , aex datarmination in Atloidt Ifyoanita Chitwood, 19491 and intersaxuali tv in M.layahlea (Troub, 188S) Chitwood 1949, i m t , iOal. .^YtWifhTjifnaHl > i ^ l 12-31.
Trudgiiit ( l 9 6 7 j . The e f f e c t of environment on sax deteiv mination in rg^lffffh^gnsjlM' Iffi ldl ffiigqjlgflt J^* 263-272.
Turnar, iJ.H. and a.A. Qiapman (19T2J. infi?ction of saedlings of a l f a l f a and red c l o v s r fay conc^nitant jxitxilations of i^^oidQcivnft ^ncffanlti, <3nd .f tfiiYir.i fTBbm JL* mm^aL^f 280-286 .
Tyl«r , J . ( i933aK Uavaioisment of the root-knot naroatode as affacted by ten ^arature. f^^gardia X* 391-415,
/aishnov, 4,U, and C.L. ^athi ( l « 7 8 j . Pathcxjenicity of '^loidooynq i M ivl.mcharhvix?f>us on bajra and t h e i r
Jhip. i a i i j " " interr^ldtionship. Indian J,, mm&itol,. 1 - 8 .
/an J0r^;uni, J .H, and H, Haestra (1979)» t a c t i c a s 3©cts of the charslcal control of noraatodas in s o i l , Ini a o i l J i s i n f e s -t a t i o n (ddts, .4uld«r, J .H, ac is i i t l f i c -•'ubl. Comn>,, tfEisterdBin, Jxford, i ^ Y o r k J .
Van i iundy, S.t), and J . J , .rikpatrick (I97'j) . I'tematode-netiatodo interact ions on tomato. J,, Mgrnatoi. X* 330-331 .
Vasudttvat H.ii. and Hingorani (1952) , i lacterial disease of wheat caused by l^yltlci (Hut), aergey s t i l * ^ t o o a t h . . 4 2 :
• ^^iglierchio, U.H. and Yu (1966) . .-'lant growth substances
and olant parasi t ic nematodes. 11. Host influences auxin content, iixnt. Par^aitpl.^ 88-95 .
Walkinshaw, C.lf.j Ci.J. l i r iff in and R.H. Larson { l > 6 l ) . Trlchodorus Chris t ie ! as a vector of potato carky rings pot (Tobacco r a t t l e ) t i rus* ^^vtooath.. fill 806-808.
Wallace, H.K. (1963) . The iaiologv of plant p a r a s i t i c nematodes, tidward Arnold Ajbl. Ltd. , LoiKlon.
(1973) . iNiematode ecology and plant diseases. iidward Arnold. iAibl. Ltd. , London.
14
V^«i9ch9r» {1968 )* vvBchselwlrkungen Zwischen n®raatod©n und andaran ^haderr&garn an liutsplarmen. I n t . ;»ynip. Naraatoj., B l l t i O y M * ^ P ^ -
( 1 9 7 4 ) , Int erspecif ic competition ijatween M. ritzeraabosi and ^J, .sfjp^aff; on tobacco. aiLiaSl*
X9T4, Ciranada, iipain.
t.ys® ( 1 9 7 3 ; . Feadlnn of IvU-;ichDrhvnchas dubius. , J^ l 125-136.
yang» M.j N.T. ^^well and sC.H. darker (1976) , Interdiction of concomitant soacies of namatodes and Fusartu^ Qjtvsg»oruni f . BP, vasinfaetum on c o t t o n . J,, tvien^atol.. 81 7 4 - 8 0 ,
Vfik, G..^ and J i r chf ie id ( 1 9 7 5 ) , iioot tis^iuo reac t ions of sweet iXJtato t o tha renifoxm ne -iatode, ji,
* Rafarencas not consulted i n o r i g i n a l .