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.

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