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STUDIES ON STUNT NEMATODES, TYLENCHORHYNCHUS SPP
DISSERTATION SUBMITTED TO THE ALIGARH MUSLIM UNIVERSITY, AUGARH
IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF
iiasiter of ^!iilo£iop{)p IN
BOTANY
MEHAR BANO
DEPARTMENT OF BOTANY ALIGARH MUSLIM UNIVERSITY
ALIGARH
1986
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!;r:ADER
DEPARTMENT OF BOTANY
ALIGARH MUSLIM UNIVERSITY ALIGARH-202001 {INDIA)
Dated .20., 4 . 198.7.
C_. •'LR„T_.I_F__I_C_A_T_E
This i s to c e r t i f y t h a t Miss Mehar Bano Is
v/orking in t h i s Department as a r e s e a r c h s tuden t
under my superv i s ion and guidance. The d i s s e r t a t i o n
e n t i t l e d "Studies on s t u n t nematodes. Ty 1 enchorbynchus
s p p . " has come t o sHape due t o the genuine and s i n c e r e
e f f o r t s made by her* The con ten t s of t h e d i s s e r t a t i o n
a r e u p - t o - d a t e and o r i g i n a l . She i s allowed t o submit
t h e d i s s e r t a t i o n t o t h e Aligarh Muslim Univers i ty for
c o n s i d e r a t i o n of t h e award of t h e degree of Master
of Philosophy in Botany,
( M. MASHKOOR AIAM )
I express iny deep sense of g r a t i t u d e t o
Dr* Me Mashkoor Alam# Reader# Department of Botany#
Al igarh Muslim Univers i ty* Aligarh for sugges t ing t h e
problem and prov id ing guidance wi th unceasing encourage
ment and keen i n t e r e s t dur ing t h e p r epa ra t i on of t h e
manuscr ip t .
I a l s o wish t o thank the Chairman* Department
of Botany* Al igarh Muslim Universi ty* Al igarh fc r
p rov id ing necessary f a c i l i t i e s ,
I am highly thankful t o Dr. M.A, S i d d i q u i ,
Mr» S a r t a j A, Tiyagi* hJrSo Anjum Ahmad* Miss Suha i l
Anver* Dr. R.P. Singh* Dr. Kusurn Kumar! and m/ o the r
co l l eagues and f r i ends for t h e i r he lp and c o - o p e r a t i o n .
( MfflAR BANG )
C O N T E N T S
1 . INTHUDUCTION AND REVIEW OF LITERATURE . . 01
2 , MATERIALS AND METHODS „» . . . . 29
REFERENCES
- oOo -
1. INTRODUCTION A JD REVIEW OF LITERATURE
Nematodes are large ubiquitous group of invertebrates
and adapted to almost every environment and are found where
ever there is- moisture^available^ More than 5000 species
of plant and soil nematodes are known today and of these
nearly 2000 belong to genera that contain plant parasites.
Plant parasitic nematodes capable of causing well
manifested diseases, usually occur in poly-specific commu-
ties and majority of them are polyphagous. On the basis
of their feeding habits they may be divided into migratory
and sedentary endo-parasites and ecto-parasites.
The nematode body plan is simple* They are long,
tubular organisms, un-segmented, usually circular in cross-
section, and basically bilaterally symmetrical. The
females are generally larger than the males and adults of
the two sexes are easily distinguished by morphological
differences. The females are often more adapted to
parasitism than the males and in some cases may swell to
become saccate or spherical.
Nematode, is regarded as phylum or class. It is
usually divided into two sub-classes: Adenophoria
(Aphasmidia), and Secernentia (Phasmida). Most of the
parasitic nematodes belong to the sub-class Secernentia.
The genus Tylenchorhynchus, the focal point of the
present dissertation, was established by N.A. Cobb in 1913,
• 2 »
The common name 'Stylet nematode' was proposed by
G« Steiner in 1949. It is also known as 'Stunt nematode' ,
Siddiqi (1986) has placed this nematode in the following
classification s
Order
Suborder
Superfamily
Family
Sub-family
Genus
Tylenchida
Tylenchina
Dolichodoroidea
Dolichodoridae
Tylenchorhynchinae
Tylenchorhynchus Cobb, 1913
Syn. Divithus Jairajpuri, 1984>•
Marasinema Javed, 1984;
Tessellus Jairajpuri and
Hunt, 1984.
Stunt nematodes are ectoparasites and feed primarily
on epidermal cells of roots in the region of elongation
but occasionally they are observed partly or totally embed
ded in the host tissue. This results in retardation in the
overall size of the roots, but the roots are not marked by
lesions. Top growth of parasitized plants is stunted which
results in severe loss in yield. The host range is quite
wide. They feed on plants ranging from herbaceous annuals n.
to woody perenials and are associated with most of the
crops, medicinal plants, conifers, and natural vegetation
etc.
: 3 :
The first record 'of Tylenchorhynchus spp, in India
was published as an abstract of a paper by Siddiqi and
Basir (1959). Later in 1960, T. dactylurus Das, 1960,
and To digitatus Das, 1969 were described from roots of
Capsicum annum and Ricinus communis respectively from
Hyderabad, South India.
In the present treatment it is proposed to work on
the pathogenicity of the stunt nematode, Tylenchorhynchus sp.
and its inter-relationship with other soil biota with the
ultimate goal of its satisfactory control. An overview
of the work carried out on these aspects is as follows :
I. PATHOGENICITY
Each host-parasite association has its own charac
teristics, A population that damages the hosts in one
association may not cause visible symptoms above ground in
another. In pot tests J. claytoni reproduces on pine
seedlings without causing visible abnormalities beyond
disturbances of root cap cells<, In alfalfa, this species
feeds on root epidermal cells without damaging its host.
However, the same species causes a serious disease of
azaleas, and high population of another species, J. dubtUs
es depress/growth of bent grass and Kentucky blue grass.
Although considerable work has been carried out on
the pathogenicity of several species of Tylenchorhynchus >
• 4 •
Usw-ev-er'; little is known about T. brassicae a species
commonly associated with cabbage and cauliflower in India.
In 1972 Siddiqi _et _al» reported the host range and varietal
resistance of certain crucifers against T. brassicae. Out
of 22 vegetables and 10 ornamentals tested only cabbage,
cauliflower and tomato proved to be highly efficient
hosts; sesbania, radish^sugar beet, lettuce, Nemesia,
Callistephus and Verbena moderately efficient and remain
ing were unfavourable hosts of J. brassicae. Almost all
the varieties of cabbage and cauliflower used in the test
were highly susq^tible to T. brassicae. The nematodes were
found mostly confined to the cortical tissues of both
cabbage and cauliflower and at times were also observed
in stelar region. These observations confirmed the find
ings of Steiner (1937). The work carried out on the host
range of Tylenchorhynchus sppe is summarized as under :
S.No. Nematode Plant References
1. Tylenchorhynchus sp,
2. Tylenchorhynchus sp.,
3. Tylenchorhynchus sp.
4. Tylenchorhynchus sp.
Sugarcane, B^echfield and Rice(va- ^^^^i" ( 956) riety Zeninth), Sweet potato (variety Porto Rico)
Fruit trees Yadav et ail.(1970)
Rice Das/ and f|V (1971)
Peach,Apple, Kyrou (1972) Pear,Cherry orchards
• 5 •
S.NOe
5.
6.
7.
8.
9*
10»
11.
12.
i.O a
14.
15.
16.
Nematode
Tvlenchorhynchus
Tylenchorhync
Tylenchorhync
Tylenchorhync
T. aqri
T. aqri
T. aqri
T, aqri
T. brassicae
T. brassicae
T» brassicae
T. brassicae
bus
hus
hus
X
sp.
sp.
sp.
sp.
Plant
Turf grass
Cabbage
Corn
Conifers
Kenland red clover
Leguminosae, Gramineae
Redelover, Kentucky blue grass
Trifoliuffi pratense, Poa paraten-sis, Triti-cum aestivum
Crucifereae
Cabbage, cauliflower knol-khol, Chinese cabbage, broccoli, tomato
Maize
Calendula, Corn flower, Black gram, Okra, Bengal gram, Cotton tomato. Green gram. Egg-plant
References
Fushtey and McElroy (197?)
Hirling (197S),
Caswell (1981),
Mancini et al.^ (1981)
Amosu and Taylor (1974)^
Coates e_fc a_l.(1978)
Coates (1979)
Coates _et _al.(1982)
Khan £t al, (1970)^
'Siddiqui et al. (1972)
Rai (1969)
Alam al. (1979),
• 6 •
S.No.
17.
18,
19.
20.
21.
^ ^ 9
23.
24.
25.
26.
27.
28.
29o
T.
!•
!•
!•
Tc
!•
To
!•
I»
I*
!•
To
I*
Nematode
brassicae
brassicae
brassicae
brevilineatus
brevilineatus
claytoni
claytoni
claytoni
claytoni
claytoni
claytoni
claytoni
claytoni
Plant
Ornamental plant
Turnip, CandyTuft
Toria (B. rapa var. Toria), laha(B. junceal Chinese cabbage (B. rapa cv. Petsai),brussels sprouts (B. ele-raceae cvT Sutton Express)
Groundnut
Pea-nut.
Alfalfa
Cranberry (Vaccinium macrocarpon Ait)
Redpine(Pinus resinosa Ait)
'Southern stock'azaleas
Turf grasses
Creeping red fescue
Long leaf pine (Pinus palus-tris)
Japanese Holly (Ilex crenate)
References
Sarosh and Saxena (1984)
Khan et ail. (1985)
Khan et ail. (1986
ICRISAT Ann. Report (1982) '
Reddy ejt al.. (1984)
Krusberg (1959)^
Zuckerman (1961)
Sutherland and Adams (1964).
Barker and Gayleworf (1964)^
Troll and Rohde (1965)
Troll and Rohde (1966)
Ruehle (1973)
Aycock et £1.(1976)
3 cultivers Helleri,Convene, Rotundifolia
: 7 s
S,No. Nematode Plant References
30. J. claytoni
31, T. claytoni
32o
33.
34,
I«
J.
I*
clarus
clarus
clarus
35a T. contractus
36. T. dubius
37. T. dubius
38. T. dubius
39. T. dubius
40. T. dubius
Ilex cornula cv. Barker _ejt al,( 1979^ Rotunda,!_. vomi-toria var. nana.
Rice
Alfalfa
Krishnaprasad and Krishnappa (1982)
Noel and Lownsbery (1977)
Cotton(Gossy- Kheir jet .( 1977) pium barbadense)
Rye grass, Sweet corn. Parsley,Tomato (cv. VoF. 145, Yellow Dock), Alfalfa (cv. Lahonton).
Tobacco
Sitka spruce (Piceae s lichens is.)
Rice,Sorghum, Perenial Rye grass, oats, millet, summer rye, corn, barley,cauliflower, turnip, radish, gram, pea, spinach, beet, carrot
Lownsberry (1980)
Katalan Gateva and Budurova (1981)
Gowen. (19717
Sharma (1968)
' Toronto*creep- Laughlin^and, ing bent grass, Vargas j3fi/(l972) 'Merlon'Kentucky blue grass
Barley
Brassica rapa var. silvestris
Saynor (1972)
Wyss (1974)
: 8 :
S.No.
41.
42,
43.
44 o
45.
46.
47.
48,
49.
50.
51.
52,
^53)
54^
55,
^56,/
yj 1 <>
I» T.
T.
!•
I'
1" T J.'
!•
I'
Tc
T,
! •
!•
T,
!•
Nematode
dubius
dubius
dubius
dubius
dubius
indicus
Plant
Rye grass
Ornamental plants
Strawberry
Dianthus Caryophyllus
Maize
Pearl millet
lamelliferus Rye qrass
martini
mashoodi
maximus
nudus
nudus
vulgaris
vulgaris
vulgaris
vulgaris
vulgaris
Bermuda grasses
Ornamental plants
Rye grass
Kentucky Blue grass (Poa pratensis)
Sugar cane
Zea mays.
Maize
Gram (Cicer arietinumj
Maize
(CencMJS ciliaris cv.
References
Bridge and Hague (1974) d' '
Krishnappa et al. (1980)
Szczygiel (1981)
Katan-Galeva and Milkova (1982) <-
Hellinga et al./-(1983)
Nandkumar and Khera (1970) -^
Bridge and Hague (1974) 0
Johnson (1969)
Sarosh and Saxena (1984)
Bridge and HaqjUe (1974) C
Smolik and Malek (1973)
Haider et a2.(1985)
Upadhyay and Swarup (1972)^
Upadhyay et al (1972) ^.- "~
Gill and Swarup (1977)
Jain (1982)„
Azmi and Singh (1984)
: 9 :
II. INTERRELATIONSHIPS
A 'healthy' crop is not necessarily a crop without
pathogens. It is rather a crop in balance with its compe
titors including the pathogensc When one species of pathogen
reproduces to excess, then the crop suffers. But in nature,
the pathogens including the nematodes rarely occur in mono
specific communities. The nematode communities are rather
dynamic and their members are constantly interacting with
each other as well as with other organisms, plants and the
environment. Under favourable conditions there may be several
kinds of interactions leading to various disease complexes,
however, with considerable overlappings. Hence, it is logi
cal to assume that the net result of plant damage caused by
different disease complexes may be different from those
caused by the individual pathogens. Moreover, the nature of
the disease complexes caused by certain sets of pathogens
may not be the same under different ecosystems.
The nematode-feeding on plant tissues causes extensive
damage in the form of injuries which sometimes paves ways
to secondary invaders which are otherwise incapable of
doing so alone. Sometimes the nematodes act as vectors of
viruses, bacteria, etc. While in some associations the
nematodes act as breakers of resistance of plants to
different pathogens. Pitcher (1965) suggested the follow
ing possibilities in different interactions where the
: 10 :
nematodes may act as:
i. Vectors of pathogens capable of selfestablishment
once in contact with the host "
ii. vectors of pathogens incapable of self-establish
ment unless introduced below the epidermis '
iii» mechanical wound agents ',
iv. providers of necrotic infection-courts ^
V. modifiers of substrates -
vi. breakers of disease resistance ',
vii. deterrents of plants disease .
Interaction between microorganisms is well establi
shed, and this aspect with respect to nematodes has been
reviewed by various workers (Pitcher, 1963, 1965;
Powell, 1963, 1968, 1971, 1979; Norton, 1978; Dropkin,
1980). These reviews reveal that the literature on the
interaction of plant parasitic nematodes with other micro
organisms is quite exhaustive. In the present treatment,
however.the discussion will be confined to the inter
relationships of the stunt nematodes, Tylenchorhynchus
spp. with other soil biota. An overview of this aspect
is as under :
1. Nematode-Nematode Interaction;
A possible interaction between Tylenchorhynchus
martini and Pratylenchus penetrans on alfalfa and red
: 11 :
clover were reported by Chapman (1959). There was an
increase in the populations of J., martini in combination
with P. penetrans, while P» penetrans was not affected.
In another experiment Johnson (1970) claimed that root
damage caused by the interaction of T, martini and
Criconemoides ornatus and Belonolaimus lonqicaudatus was
unfavourable for the reproduction of other species on six
bermuda grasses.
Ross et al. (1967) reported an interaction between
Helicotylenchus dihystera, Tylenchorhynchus claytoni and
Pratylenchus brachyurus on soybean in which the population
development of H. dihystera and T. claytoni increased.
Where as Miller and 'vVireheim (1968) observed that Globodera
tabacum reduced the infection and survival of Pratylenchus
penetrans and J. claytoni on tobacco plants. Miller and
Mclntyre (1975) and Mclntyre and Miller (1976) noted that
simultaneous inoculation of Pratylenchus penetrans and
T. claytoni inhibited the population of P. penetrans in
tobacco rootso Krishna Prasad and Rao (1977) reported that
interaction of Tylenchorhynchus claytoni and Helicotylenchus
crenatus on rice resulted in significant reduction in
multiplication of H. crenatus. In another study it was
found that in concomitant inoculation, H. crenatus
inhibited population build-up of J. dubius on tomato
(Krishnappa and (Prasad, 1979).
12
Sikora _et aj.« (1972) noted inhibition of Meloidoqyne
naasi in an interaction between M. naasi, Pratylenchus
penetrans and Tylenchorhynchus aqri on creeping bent
grass whereas the presence of M. naasi did not affect the
population of To aqri. Amosu and Taylor (1974) observed
that Meloidoqyne hapla alone and in all combinations with
£• penetrans or J. aqri was highly pathogenic to red
clover, A high inoculum level of P. penetrans alone and
in combination with Jo aqri significantly reduced the
gall index of M. hapla«
Alam e_t a]^, (1975) reported that combined inocula
tion of Meloidoqyne incognita and Tylenchorhynchus
brassicae caused significantly more reduction in the
growth of tomato plants than caused by either of the
nematodes* When inoculated simultaneously with M.
incognita the population of J* brassicae decreased to
levels below the initial inoculum. J. brassicae, on the
other hand, did not materially affected the development
of the root-knot nematodes. Similar results were also
reported by Khan et al, (1978, 1986) and Khan and Haq
(1979)e
Combined effect of U„ incognita and J, vulgaris on
bajra plant was additive though J. vulgaris showed
antagonistic relationship with M^ incognita (Vaishnav
and Sethi J 1978a)*
13 t
Misra and Dass (1977) reported that three ectopara-
sitic nematodes, e.g. Criconemoides ornatus, Hoplolaimus
indicus and 1iylencho;piiynchu"s) nudus in combination signi
ficantly reduced the infectivity of M. incognita in
brinjal (Solanum melonqena Lo). In this operation dry
weight of shoot and root and recovery of nematodes from
soil was also affected. Smolik (1972) observed that the
reproduction of T. nudus was greater than Helicotylenchus
leiocephalus when either was inoculated alone on spring
wheat. Reproduction of both the species was suppressed
when they were inoculated together.
Upadhyay and Swarup (1981) examined the combined
inoculation of Merlinius brevidens and Tylenchorhynchus
vulgaris on wheat. The results indicated that M. bre
videns significantly reduced the population of J. vulgaris
Kaul and Sethi (1982) noticed that M, incognita was
adversely affected by the presence of Heterodera zeae and
1* vulgaris on maize. Population of J. vulgaris was
reduced by H. zeae and M. incognita. But T. vulgaris
finally enhanced the penetration of JA^ incognita.
Siyattand e_t . (1982) investigated the life cycles of
J* vulgaris, Pratylenchus thornei and Hoplolaimus indicus
individually and in combined infestations in maize and
observed that the life cycle of J. vulgaris was shortened
by 13-15 days in combined inoculation. However, no such
differences were observed in the case of P. thornei and
: 14 :
H» indicus*
2. Nematode-Funqus Interaction;
Stunt nematodes become more numerous around pea
infected with Fusarium (Davis, 1964) and declined in
the presence of Trichoderroa yirlde* Holderaan (1956)
revealed that the Tobacco stunt nematode, Tylencho-
rhynchus claytoni was capable of increasing the incidence
of wilt caused by Fusarium oxyspor~um f, nicotianae in
susceptible varieties of flue-cured tobacco. Wehunt
and Weaver (1972) reported more growth reduction in
peach seedlings in presence of Fusarium oxyspori:um and
either of Hoplolaimus qaleatus, Tylenchorhynchus
claytoni or Criconemoides xenoplax. Vargas and Laughlin
(1972) examined the role of Tylenchorhynchus dubius in
the development of Fusarium blight on Poa pratensis.
They indicated that association of Fusarium roseum and
1' dubius caused the Fusarium-blight.
Kisiel _et jl. (1969) noticed that Tylenchus aqricola
and Tylenchorhynchus claytoni also contributed towards
the increase in root rot of corn caused by Fusarium
roseum,
Jones and Overman (1971) reported^Tylenchorhynchus
capitatus did not affect Fusarium or Verticj/glum wilt
development at temp. 22^ or 25°C.
15 :
Upadhyay and Swarup (1981) observed that the growth
of maize plants reduced further in combination with
Tylenchorhynchus vulgaris, Pratylenchus zeag and Fusarium
moniliforme compared with the reduction caused by J.
vulgaris alone.
According to Haglund and King (19&1) the severity
of common root-rot of peas caused by the fungus,
Aphanomyces euteiches was increased by the nematode,
Tylenchorhynchus martini. Increase in the disease
intensity was also found to be directly related to the
number of nematodes.
Sobun e_t a_l. (1975) reported the relationship
between Tylenchorhynchus sp. and a root-rot fungus from
gram (Cicer arietinum L,). This fungus delayed emer
gence of gram seedlings«
Moore _et _al» (1956) noted in some cases that the
black shank fungus, Phytophthora parasitica f. nicotinae
has broken the resistence of many varieties of tobacco
and the stunt nematode (Tylenchorhynchus claytoni)
appeared to have influenced the development of black
shank symptoms.
Daykin and Sanson (1979) observed that Rhododendron
obtusum plants when inoculated with both Phytophthora
cinnamomi and Tylenchorhynchus claytoni showed signifi
cantly poor growth than when it was infested with the
single pathogen.
: 15 :
Barham e_t al. (1974) reported that Tylenchorhynchus
claytoni was able to modify resistance afforded by Pine
host by fungus symLionts against Phytophthora clnnamomi.
Khan ejt a_lo (1971) reported that To brassicae alone
did not affect the percentage of emergence of cauli
flower seedlings. Reduction occured when the soil was
inoculated with the combination of R. solani and
J" brassicae.
Vaishnav and Sethi (1978b) reported that the co-
inoculum of Sclerospora qraminlcola and Meloidoqyne
incognita advanced the appearence of downy mildew of
bajra by 12-13 days incomparison to the treatment
receiving inoculum of S. qraminicola singly or in com
bination with J, vulgaris.
Miller and Anagnostakis (1977) studied the suppre
ssion of Pratylenchus penetrans and Tylenchorhynchus
dubius by Trichoderma viride because it was antagonistic
to these nematodes.
Overman and Jones (1970) revealed the effect of
stunt and root-knot nematodes and two isolates of
Verticillium. Increased wilt symptoms occured with
!• capitatus at 33* 0 and with M. incognita at 29°C.
Ndubizu (1977) found that association of Praty
lenchus penetrans^ M. hapla or Tylenchorhynchus claytoni
with Verticillium dahliae resulted in greater growth
: 17 :
reduction of cherry seedlings thVn they caused singly,
3. Nematode-Bacteria Interaction:
Ingham and Colaman (1983) reported that higher
bacterial (Pseudomonas paucinobilus) population was
found in the presence of Tylenchorhynchus claytoni in
both rhizosphere soil and non-rhizosphere soil.
4, Nematode-virus Interaction;
Naqvi and Alam (1975) found that Brinjal mosaic
virus (BiW) promoted the population build -up of
Tylenchorhynchus brassicae on eggplant. Greatest
increase of nematodes was observed in plants inoculated
with virus 3 weeks prior to the nematode inoculation.
It was claimed that the virus was responsible for certain
physio-chemical changes in the host which favoured
nematode multiplication.
III. CONTROL
Control of diseases is a branch of applied ecology
and is concerned with systems in which there are numer
ous interrelated factors, it follows that there are
likely to be many ways in which changes in such factors
might either be deleterious or beneficial. Knowledge
of the crop and its environment, including parasites
: 18 :
and soil factors, is essential if satisfactory control
is to be achieved without undesirable consequences. It
is generally agreed that the aims of such measures are
to reduce the amount of disease within the crop, rather
than in the individual plant.
The nematode control methods are not very different
from those used for other parasitic agencies. They may
be classified in four broad categories: physical, bio
logical, chemical and cultural.
1« Physical Method of Nematode Control:
Application of heat as a means of denematizing the
soil in limited areas of controlled environment is a
very useful control method. The effects of physical
factors such as steam sterilization of soil, hot water
treatment etc. have been reviewed by Jenkins {I960),
Southey (1965) and Cuany (1971). In 1967.Heald and
Wells obtained the control of ectoparasitic nematodes
associated with turf by hot water treatments of 55°C
for 15 and 30 minutes but not at 21* 0. This kind of
control of plant parasitic nematodes has its own limi-
tations(e/g. it cannot be used on large scale.
2. Biological Method of Nematode Control;
Biological control means the reduction of nematode
populations by stimulation or introduction of predators
: 19 :
and parasites such as nematodes, fungi and bacteria,
etc, (Sayre, 1971, 1980a, bj Norton, 1978; Mankau, 1980),
Karg and Grosse (1983) examined the possibility of using
predacious mites as biological control agent against
plant parasitic nematodes of the genus Tylenchorhynchus
in caraway crop. It is however generally accepted that
the biological control of plant parasitic nematodes is
still far from being practical under field condition.
3. Chemical Method of Nematode Control;
The use of nematicidal chemicals though very
effective but not very conducive especially in Indian
conditions, because of their high cost of application
and the inherent pollution hazards. There are several
reviews on the use of nematicides for the control of
plant parasitic nematodes (Peachy, 1965J Smart, 1969;
Van Gundy and McKenry, 1977; Van Berkum and Hoestra,
1979; Lamberti, 1979; Wright, 1981).
Considerable work has been carried out on the
control of Tylenchorhynchus spp, by means of nematicidal
chemicals. Defloor (1971) reported that the population
of Tylenchorhynchus was decreased by 73 to Q^yi in plots
receiving 20 g/m of granular Dazonieto Yields of the
vegetables were increased and the weeds were controlled.
Singh (1975) reported that chemicals, D-D, DD-MENCS,
Bunema, Oxamyl, Phenamiphos and DBCP controlled
: 20 :
Tylenchorhynchus spp, Nath et al_. (1976) found out that
after the application of Dasanit at 25 kg/ha, sugar cane
yields were increased in soil where the dominant nematode
was Tylenchorhynchus spp. Some aromatic petroleum
fractions viz., '400 N' and 'Solvent B' also reduced the
population of Tylenchorhynchus spp. (Saeed _et £l., 1976).
Skroch et al,. (1977) found reduction in Tylenchorhynchus
after the treatment with Vorlex orazide (as Na N3 or
KN3). Naqvi et al. (1978) found that 'Petkolin' a
chlorinated petro-product, was as effective as Neraagon
for controlling Tylenchorhynchus spp. Dolmans and Bunt
(1981) reported the control of Tylenchorhynchus spp, in
maize by seed treatments with Oxamyl. Overman and
Howard (1982) investigated the influence of Nemacur and
Vydate on stunt nematodes in fields of strawberry,
Nemacur applied at 6,72 or 13,44 kg/ha controlled
Tylenchorhynchus spp, through out the season* Vydate
controlled nematodes only after the application of
4 weekly foliar sprays,
Siddiqui and Khan (1973) evaluated some nematicides
for the control of Tylenchorhynchus brassicae, Nemaphos
and Thimet reduced the population while Solvirexand
Rogor were almost ineffective against J. brassicae,
Siddiqui and Khan (1974) in another study reported the
chemical control of Tylenchorhynchus brassicae by the
application of I>-D and Vapam both in pots and underfield
conditions. The growth of cabbage and cauliflower plants
was also found significantly high in pots and beds treated
with these neinaticides, Singh _et a^. (1980) found good
control of Tylenchorhynchus brassicae by the application
of some nematicides such as Carbofuran, Diraethoate and
Aldicarb (0»1 or 0.2 g nematicides/lO g seeds) in tomato
without any phyto-toxicity. Haq _et . (1984) examined
the effect of nematicides, DD, DBCP, Phorate, Fansulfothion,
Dimethoate, Aldicarb and Carbofuran on the sex ratio of
Tylenchorhynchus brassicae on cauliflower. The number of
males was highest in DD at all the levels of inoculum.
Phorate, Diraethoate and Aldicarb increased male populations
at the population levels of 1000 and 5000.
Miller and iCring (1970) studied the effect of sys
temic insecticides and soil fumigants applied for control
of nematodes and insects in potatoes in U.S.A. At harvest
time population of Tylenchorhynchus claytoni was 90j< lower
in Aldicarb treated plots and 60 to 80;< lower in Telone
and Carbofuran treated plots than in untreated plots.
Radewald aipl-Mairev (1972) found good control of the stunt
nematode, Tylenchorhynchus claytoni on azalea (Rhododendron
indicum) by the application of 30, 60 and 90 ppm DBCP with
no phytotoxic effects. The treatment enabled the plant to
recuperate. Rodriguez-Kabana and Ingram (1977) examined
the effect of treatment of potato seed pieces with Oxamyl
: 21 r,
at concentrations of upto 24 g/l which were then planted
in pots containing soil infested with Tvlenchorhynchus
claytoni* Treatments with the 6.0 g/l solution resulted
in 74^ fewer To -Claytoni than in control. Ingrain e_t aJL.
(1977) reported that the numbers of Tvlenchorhynchus
claytoni in soil declined sharply in response to the neraa-
ticides viz., Ethoprop and Fensulfothion at concentrations
in the range of 0-100 mg/kg with little additional control
by increasing the concentrations further. Rodriguez-Kabana
et als (1977) reported decline in numbers of Tylenchorhyn-
chus claytoni with systemic nematicides in wheat and rye
by the prior treatment with Oxamyl, Carbofuran or
phei^amiphos. Rodriquez-Kabana and King (1979) observed
neroaticidal activity of the fungicide Ethazole at rates
above 4.5 kg against Tylenchorhynchus claytoni on cotton.
Schmitt and Corbin (1981) examined the interaction of
Fesulfothion and Phorate with pre-emergence herbicides in
the control of Tylenchorhynchus claytoni. Fensulfothion
and Phorate alone and in combination with the pre-emergence
herbicide effectively controlled T. claytoni. DiSanzo
(1981) reported the control of Tylenchorhynchus claytoni
on maize and tobacco by foliar application of Oxamyl,
Carbofuran and a related compound 'PMC 3500'.
Embabi and Shohla (1978) examined the effect of some
nematicides on Tylenchorhynchus clarus and on the growth
of cotton. In Fumazone 75 EC at 15 kg/feddan, Temik 10 G
: 2: :
at 15 kg/f, Teracur 5-G at 30 kg/f and Vydate 10-G at
15 kg/f the reductions in nematode populations were 98,8,
81»1, 46.5 and 71,2>i respectively. However, Fumazone was
found to be highly phytotoxic. Laughlin and Vargas (1973)
reported the influence of benomyl on Tylenchorhynchus
dubius alone and in combination with 4 non-fumigant nema-
ticides. Benomyl was applied in single or split applica
tion. After the initial application, T. dubius were lower
in treated than untreated plots. After 7 months Benomyl
enhanced the activity of the 2 least effective nematicides
vizo, BAY 25141 (fensulfothion) and Oxamyl while it
suppressed the activity of 2 most effective chemicals viz.,
BAY 68138 (Phenamiphos) and CHEM 7375 IS,S di-isoprophyl
(2-methyl vinyl) phosphoro dithioate. In another stydy
Ethoprop controlled Tylenchorhynchus dubius by 8Qj<
(Miller, 1978).
Elgindi and Oteifa (1967) reported the nematicidal
effect under field condition against the cotton nematode,
!• la"t s, There was significant reduction in the nematode
population by the application of D-D at the rate of 4001b
per feddan (4,200 sqm). Todd and Claflin (1984) observed
significant reduction in population of Tylenchorhynchus
martini on sorghum by the application of insecticides ~
nematicides.
: ^ :
Chhabra and Mahajan (1976) reported that the applica
tion of granular nematicides, Fensulphothion at 15 kg
a,i./ha was significantly more effective in controlling
the nematode, Xylenehorhynchus mashoodi than all the
other treatments. Jain (1980) reported the control of
J. mashoodi on Berseem (Trifolium adenandrinum L.) with
fumigant and systemic granular nematicides,
Azmi _et _al.. (1984) reported maximum reduction in
1* vulgaris population in pots treated with Phenamiphos
followed by Aldicarb and Carbofuran.
4. Cultural Method of Nematode Control;
Nematodes are also affected to a great extent by
land management and cultural practices such as fallowing,
flooding, tillage, prevention of spread, selection of
healthy propagating materials, crop rotation etc. (Berge,
1971; Nusbaum and Ferris, 1973; Norton, 1978; Netscher
and Taylor, 1979; Bird and Thomason, 1980; Khan, 1981;
Davide and Castillo, 1981; Anderson, 1982; Noe, 1986);
use of resistant varieties (Moore, 1960; Howard, 1965;
Hunt and Peaden,1972); and the application of organic
matter (Oostenbrink, 1960; Brown, 1965; Singh and Sitara-
maiah, 1970; Sayre, 1971; Alam, 1976; Norton, 1979;
Muller and Gooch, 1982; Hornick e_t aj.., 1984; Siddiqui,
1986).
T
Barker and Nusbaum (1971) reported population changes
in eight years in plots infested with Tylenchorhynchus
claytoni. Nematode populations were sampled before plant
ing and after harvest. Tobacco supported reproduction of
1* claytoni* Both tobacco and corn were excellent hosts.
Cotton and peanuts were poor hosts. Murphy £t _aj.. (1974) q
found that J. claytoni populations greater than 100/500 cm
of soil before planting caused decline of azaleas.
Ferris and Bernard (1971) observed the crop rotation
effects on population densities of ecto-parasitic nematodes,
Tylenchorhynchus martini and jT, acutus. The crops included
in the study were maize, soybeans, oats, wheat, lucerne,
red clover and brome grass. Numbers of J. martini were
significantly less in the second year of maize than in the
first. Khan _e_t aj.. (1975) studied nematode control by
crop rotation method and found an increase in population
of Tylenchorhynchus brassicae following the mono-culture
of eggplant, tomato, okra, spinach, cauliflower, cabbage
and mustard.
Alam _et _a_lo (1977) reported the influence of different
cropping sequences of soil populations of Tylenchorhynchus
brassicae. All the cropping sequences tested brought about
reductions in the population of _J. brassicae except in one
cropping sequence. Broadbean and Kochia in addition to
marigold were found to suppress the population of the
: 4 :
nematode. Schott and Jacob (1983) evaluated the effect of
continuous cropping of maize. There was increase in the
population of Tylenchorhynchus spp, and decrease in
Meloidogyne population. In comparison with 1:4 rotation
scheme, maize yield after 2 successive cultivations was
reduced by about by. and after 3 successive cultivations by
about 10/<. Shafahak and Salem (1979) reported that there
were very dense populations of Tylenchorhynchus spp.
following monoculture cotton or after clover, field bean
or wheat reduced the nematode population in cotton more
than fallowing after these crops.
Growing margosa and marigold has also brought about
reduction in the population of J. brassicae. Root exu
dates of margosa (Azadirachta indie a) was found to be more
toxic than those of marigold (Tagetes erecta). Exudates
of margosa also inhibited hatching of M. inc0f2yii.t.a
(Alam _et al,, 1975). Khan et B],<, (1971) found out that
Tagetes erecta when grown with different varieties of
tom.ato and okra brought about a significant reduction in
the population of Tylenchorhynchus brassicae. Growing
radish has reduced populations of Tylenchorhynchus by
80; (Hirling, 1976) c
Alam and Khan (1974) found out that the population of
Tylenchorhynchus brassicae was reduced by the application
of oil-cake amendments. Sitaramaiah and Singh (1978)
t 2^ :
reported that high concentrations of margosa cake increased
fatty acids which in^activated larvae of Tylenchorhynchus
^PP» iD vJ-tro but short exposures to low concentrations
were effective or stimulatory. Saxena _e_t _al. (1977)
observed that the application of mustard oil-cake (500 kg/ha)
and margosa (Azadirachta indiea) leaves (300 kg/ha) was
effective in controlling Tylenchorhynchus sppo on grape
vines, Siddiqui e_t a_l. (1976) reported that the population
of Tylenchorhynchus brassicae around the roots of cabbage
and cauliflower were less in soil where oil-cakes, were
applied then the soil treated with inorganic fertilizers.
Singh _et _al. (1980) found out that when tomato seeds
(cv, Marglobe) were treated with oil-cakes of castor,
mustard, neem, mahua and ground-nut (2 g oil cake/10 g seeds)
and planted in soil infested with Tylenchorhynchus brassicae,
there was a marked reductions in the numbers of the nema
tode. Phenol levels in roots were more in oil-cake treated
plants o
Khan _et _al. (1979) reported that the application of
oil cakes of Azadirachta indica, Madhuca indica, Ajachis
hypogea and Ricinus communis at the rate of 1001b N/acre
has significantly suppressed the population Tylenchorhyn
chus brassicae in the rhizosphere of Abelmoscus esculentus.
Singh e_t a_l. (1984) found out that seed treated with oil
cakes of linseeds, castor, sunflower and peanut resulted in
an increase in yield of cauliflower and reduction in the
population of Tylenchorhynchus brassicae. Plants grown
from seeds coated with oil-cakes had higher amount of total
free phenols and aminoacids,
Alam je_t _al» (1982) have observed that water soluble
fractions (WSF) of oil-cakes were highly deleterious to
Tylenchorhynchus brassicae.
Alam (1986) has obtained a significant reduction in
the population of T. brassicae by the application of chopped
shoots of some weeds;Solanum xanthocarpum followed by
Calotropis procera^ Datura metel, Croton bonplandianum and
Arqemone mexicana. Miller ert _al_o (1973) has found out
that about 95/. population of Tylenchorhynchus dubius was
inactivated on killed with the extracts from leaves and
stems of bean (Phaseolus vulgaris Lo) and leaves of tobacco
(NicotiCana tabacum L.)• Miller (1977, 1978) in another
study reported that Tylenchorhynchus dubius population was
reduced by the application of leaf mold composts, mycelial
residue, gelatine at 400 kg/haj mulches (plastic film),
vineger, sucrose plus fertilizer or ethoprop» He got more
than 80/. control of To dubius. '/i/hite and Dickens (1984)
noticed that Tylenchorhynchus spp, population was lower in
bermuda grass where activated sewage sludge was the source
of N then were MH.NOo was applied»
PROPOSED PLAN OF 'vVORK;
The above survey of literature clearly shows that
considerable work has been carried out on different species
s 2^1
of the genus Tylenchorhynchus but Tylenchorhynchu-s bra-
sslcae is comparatively neglected oneo This species is
one of the most important nematode species causing heavy
crop losses every year in Aligarh and els€ where* There-
fore, it is proposed to investigate hitherto unknown
avenues of knowledge. The following aspects will form
the nucleus of the future studies :
1. To study the reaction and damage of some_ economically
k important plants in order to assess the pathogenicity
of Tylenchorhynchus brassicae and to locate hitherto
un-recorded hosts. Various crop cultivars v./ill also
be screened to locate resistance, if any.
2. To study the inter-relationships of Tylenchorhynchus
brassicae with other nematodes as well as some soil
inhabiting fungi attacking some economically important
plants,
3. To study the effects of various control measures viz.,
organic soil amendments and nematicides for managing
the population of Tylenchorhynchus brassicae attacking
some economically important plants.
2. MATERIALS AND METHODS
Three species of the genus, Tylenchorhynchus viz.,
L* brassicae, 1_. mashhoodi and To vulgaris commonly inhabit
the Aligarh soils, the first named species been predominant.
Attempts will be made to study the different aspects on
this species.
2.1 Maintenance of Nematode Culture;
Pure culture of the nematode, Tylenchorhynchus
brassicae will be maintained in microplots which will have
cabbage, cauliflower on tomato as supporting crops accord
ing to the season. Soil from these microplots will be
processed by using Cobb's sieving and decanting method
along with Baermann funnel technique (Southey, 1970) for
isolating the nematodes as and when needed. The nematode
suspension collected from the funnels will serve as ino
culum of the nematode,
liVater suspensions of the nematode will be stirred
gently for making homogenous distribution of nematodes and
then 5 ml suspension will be transferred to the counting
dish (Southey, 1970) and the nematodes in each sample will
be counted under stereoscopic microscope. An average of
five such counts will determine the density of nematodes
per unit volume of the suspension.
: 30 J
2,2 Pathogenicity Tests;
Xylenehorhynchus brassicae is known to cause patho
logical effects on many plants, most of them belong to the
family Cruciferae. Therefore some economically important
plants of the family Cruciferae as well as other families
will be selected for pathogenicity test.
Sandy loam soil, which is commonly found in Aligarh
will be collected from a f^Jlow field and further passed
through a coarse sieve (1 mm pore size) to remove stone
particles and debris etce Compost manure at the rate of
1 g N/kg soil will then be added and thoroughly mixed with
the soil. Fifteen cm clay pots will be filled with 1 kg
soils compost mixtures and then these pots will be auto-
claved ana used for all the studies.
Surface sterilized seeds will either be sown directly
to these pots or seedlings will be raised from surface
sterilized seeds in big plastic trays containing autoclaved
soil-manure mixture. At 3-wk age the seedlings will be
transplanted to 15 cm clay pots containing soil-manure
mixture. Surface sterilization of seeds will be done by
treating them with 0.1>< solution of mercuric chloride for
2 munites. Then these seeds will be washed with sterilized
distilled water to remove any traces of mercuric chloride
before sowinq.
' 31 •
When, plants will attain an age of 4 wk they will be
inoculated with 100, 1000 or 10,000 freshly isolated
specimens of Tylenchorhynchus brassicae* There will be 5
replicates for each treatment. Uninoculated plants will
serve as control. The pots will then be placed on a green
house bench in a randomized manner. Necessary weeding and
watering etc. will also be done as and when required.
Data 'lA/ill be collected 2 months after inoculation.
Plants will be uprooted and the roots will be thoroughly
and gently washed with running water. Plant growth (fresh
and dry weights and lengths of shoot and root) as well as
final population of the nematode per pot will be determined.
Before taking the fresh weight, excess amount of water will
be removed by putting the roots and the shoots between
blotting sheets, while for determining the dry weight,
shoot and root will be dried in an oven at 60 C and then
separately weighed.
Reproduction factor (R) of the nematodes will be cal
culated by the formula of Oostenbrink (1966) as follows :
(where P^ represents the final population and P. represents
the initial population of the nematode).
Statistical analysis of the datcl for critical differ
ences (CD) at P = 0.05 and P = 0,01 will be done as per
: 32 I
procedure described by Pansey and Sukhatme (1978).
2.3 Varietal Screening;
Different varieties of the test plants will be grown
as described in 2.2, and the plants will be inoculated with
freshly isolated specimens of the nematode as under:
i) Uninoculated (Control)
ii) 100 nematodes per plant
iii) 1000 nematodes per plant
iv) 10,000 nematodes per plant
There will be 5 replicates for each treatment . Un
inoculated plants will serve as control. Final dat^ with
respect to the plants growth and nematode population will
be collected as described in 2.2.
Aftercare of plants such as watering, weeding etc.
will be done as and when required,
2.4 Interaction Studies;
2.4.1 Nematode-nematode interaction;
Selected plants will be raised according to the pro
cedure described in 2.2 and will be inoculated with Tylencho-
rhynchus brasslcae and Meloidogyne incognita according to
the following schedule;
; 33 s
1. Uninoculated (control)
2. J» brassicae alone (100 specimens/plant)
3. " " " (1000 «' " )
4. 81 i: .8 (10000 " " )
5. M. incognita alone (100 specimens/plant)
6. " " " (1000 " " )
7. " *' " (10000 " " )
^* J' brassicae (100) + M. incognita (100)
9. " " (100) + " '• (1000)
10. '• " (100) + " " (10000)
1» I' brassicae (1000) + M. incognita (100)
12. " " (1000) + " •• (1000)
13. " " (1000) + " " (10000)
14. T. brassicae (10000) + M. incognita (100)
15. " " (10000) + " " (1000)
16. " " (10000) -f " " (10000)
The experiments will be terminated according to the
procedure described in 2.2. The recording of data with
respect to plant growth and nematode population will be done
as described in 2,2. While the water absorption capability
of roots will be determined by the method described by
Alam et al, (1974). The root-knot index in case of
M* incognita infected plants will be determined on 0-5 scale
of Taylor and Sasser (1978) as under:
: 34 t
Gall Ind or
Eggmass
• 0
1
2
3
4
5
ex (GI)
Index (EI)
Number of galls
Egg or
masses
0
1—2
3-10
11-30
31-100
>100
2*4.2 Nematode-fungus interaction;
Selected plants will be raised according to the pro
cedure described in 2,2 and inoculated with J, brassicae
and/or Rhizoctonia solani according to the following
schedule:
i) Uninoculated (Control)
ii) Nematode alone
iii) Fungus alone
iv) Nematode and fungus simultaneously
v) Nematode 1 week prior to fungus
vi) Nematode 2 weeks prior to fungus
vii) Nematode 1 week after fungus
viii) Nematode 2 weeks after fungus
: 35 :
The inoculum level for the nematode will be lOOO/plant
while for the fungus 1 g mycelium/plant. The inoculum for
the nematode will be obtained as described in 2.1, while
for the fungus it is as under.
Pure cultures of the root-rot fungus, Rhizoctonia
solani will be maintained in culture tubes containing
potato-dextrose-agar (P.D.Ao) which will be prepared from
the following constituents:
Potato 25O«00 g
Dextrose 17.00 g
Agar 20cOO g
Distilled water 1000.00 ml
The inoculum of the root rot fungus, Rhizoctonia
solani will be raised on Richards' liquid medium (Ricker
and Ricker, 1936) having the following composition:
Potassium nitrate 10,00 g
Potassium dihydrogen phosphate 5,00 g
Magnesium sulphate 2,50 g
Ferric Chloride 0.02 g
Sucrose 50.00 g
Distilled water 1000.00 ml
One hundred ml of the above medium will be transferred
to 25 0 ml Erlenmeyer flasks which will be plugged with
cotton plugs covered with butter paper. Then the flasks
I 36 :
will be autoclaved. Small bits of the test fungus vail
be transferred to these conical flasks in an aseptic
chamber taking all the precautions prescribed for such
an operation. The fungus will be incubated for 15 days
in an incubator running at 28 + 2 C temperature. After
the incubation period the mycelial mats will be removed,
washed in distilled water to remove the traces of the
medium and then it will be gently pressed between sterile
blotting papers to remove the excess amount of water.
Inoculum will be prepared by mixing 10 g fungal mycelium
in 100 ml of sterilised distilled water and blending it
for 30 seconds in a waring blender (Stemerding, 1964),
In this way each 10 ml of this homogenate will contain
1 g of the fungus which will serve as the inoculum.
The experiments will be terminated 2 months after
the first inoculation. The final recording of the data
with respect to the nematode population and plant growth
will be done according to the method described in 2.2 and
for water absorption capability of roots as described
earlier in 2.4.1.
2.5 Control of the Nematode;
Attempts will also be made for control of the
nematode by organic soil amendments and nematicides.
: 37 :
2.5.1 Field trials;
Field heavily infested with J. brassicae will be
selected for the study. It will be thoroughly prepared
and divided into beds (2 x 2 m size) which will have
0,5 m wide buffer zone. The beds will be treated separa
tely in random^^ized manner with inorganic fertilizers
(N @ 110 kg/ha, P & K @ 55 kg/ha), oilcakes of castor and
neem/margosa (N #110 kg/ha), and nematicides (Aldicarb
and Carbofuran @ 1 kg a.i./ha).
Each treatment will have five replicates. Untreated
beds will serve as control. After two weeks of waiting
period seeds will be sown directly or seedlings will be
transplanted to these beds. Nematode population will be
determined before the treatment as well as at the time of
termination of the experiments (2-3 months after sowing
depending upon the crop) as per procedure described in
2.2. Plant growth will also be determined at the time of
termination.
2.5.2 Pot experiments;
Fifteen cm clay pots will be filled with 1 kg auto-
claved soil manure mixture in the same manner as described
in 2.2.
The pots will be treated with inorganic fertilizers
(N @ I g/pot, P 8c K @ Oo5 g/pot), Oil-ca^^Br4^^^=^^pot)
: 38 ;
and nematicides (@1 g a.i.y''pot). Each treatment will be
replicated 5 times. Untreated pots will serve as control.
After a waiting period of 2 wk seeds will be sown or
seedlings will be transplanted singly to these pots. The
plants will be inoculated with 1000 freshly isolated
specimens of T. brassicae. Final data with respect to
the nematode population and plant growth will be obtained
according to the procedure described in 2.2.
III. REFERENCES
Xlam, M.M.; N. Hasan and S.K, Saxena (1974)o Effect of root-knot nematode, Meloldoqyne incognita on water absorption capability of tomato roots, Indian J, Nematol,, 4 : 244-246.
Alam, MPM. (1976). Organic amendments in relation to nematodes. Ph.D. Thesis, Aligarh Muslim University, Aligarh : 146 p.
Alam, M.M. (1986). Possible use of weeds as soil amendment for the management of root-knot and stunt nematodes attacking eggplant. Agriculture Wastes, 16 : 97-102.
Alam, M.M« and A.M. Khan (1974). Control of phytonematodes with oil-cake amendments in spinach field. Indian J. Nematol., 4 : 239-240.
Alam, M.M. and S.K. Saxena (1975). Effect of Tylenchorhyn-chus brassicae on the water absorption capacity of roots. Geobios, 2 : 161-162.
Alam, M.M.; N. Hasan and S.K» Saxena (1975). Influence of concomitant populations of Meloidoqyne incognita ^'^'^ Xylenehorhynchus brassicae on their development and on the growth of tomato. Indian J. Nematol., 5 : 247-249.
Alam, M.M.; A. Masood and Sol. Husain (1975). Effect of margosa and marigold root exudates on mortality and larval hatch of certain nematodes. Indian J. Exp. Biology, 13 : 412-414.
Alam, M«M.; S,K. Sa xena and AoM. Khan (1977), Influence of differentjcropping sequences on soil population of plant par"asitic nem.atodes, Nematol, Medit., 5 : 65-72.
Alam, M.M,J AoM. Khan and S.K. Saxena (1979)o Mechanism of control of plant parasitic nematodes as a result of the application of organic amendments to the soil, V. Role of phenolic compounds, Indian J. Nematol.. 9 : 136-142.
Alam, M.M.; A.M. Khan and S.K. Saxena (1982). Relative toxicity of decomposed and undecomposed oil-cakes to Dlant parasitic nematodes. Acta Bot. Indica., 10 : 124-127.
: 40 :
Alam, M.M.I S.K. Saxena and A.M. Khan (1979). Suitability of crops to the reniform, stunt and root-knot nematodes. Recent Researches in Plant Sciences. Kalyani Publishers (New Delhi). ^ ^ e.'-v -V: >l
^ndersson, S. (1982), Colloque OEPP sur less nematodes a >. Kystes des ceroales. Bull. OEPP, _12 - 463-475.
i\mosu, J.O. and D.P. Taylor (1974) i?Stimulatlon of growth of red clover by Tylenchorhynchus aqri. Indian J. Nematol., 4 : 132-137o
Amosu, J.Co and D.P. Taylor (1974)vInteraction of Meloidoqyne hapla, Pratylenchus penetrans and and Tylenchorhynchus on kenland red clover, Trifolium pratense. Indian J. Nematol., A_ : 124-131.
,Aycock, R.; K.R. Barker and D.M. Benson (1976). Susceptibility of Japanese HoJly to Criconemoides xenoplax, Tylenchorhynchus claytoni and certain other plant parasitic nematodes. J. Nematol., 6 : 26-31o
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