REVIEW OF LITERATURE - Shodhgangashodhganga.inflibnet.ac.in/bitstream/10603/28726/12/12_chapter 2.pdf · Review of Literature 36 2.1.2. Biofumigation The term biofumigation is used

Review of Literature

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REVIEW OF LITERATURE

Phytonematodes are an extremely important limiting factor in vegetable

production, therefore crop protection is an integral part of food production and must

be considered within the context of modern agriculture and sustainable development.

Effective crop protection is essential both to combat the threat of widespread diseases

and to provide the effective pest management programmes. The majority of the plant

species, which account for the major world’s food supply, is susceptible to attack

from phytonematodes which are capable of causing sustainable economic losses in the

quantity and quality of the crops (Jain et al., 2007; Berry et al., 2008). The crop losses

caused by phytonematodes in economic terms estimated about $ 157 billion annually

to the world agriculture (Abad et al., 2008). Yield losses due to root-knot nematodes

(Meloidogyne spp.) range from 35.0 to 39.7% (Reddy, 1985; Jonathan et al., 2001). In

India the losses of agriculture by phytonematodes estimated at about Rs. 210 crore

annually (Jain et al., 2007).

Phytonematodes are severely destroying the roots and other parts of various

crops. Roots damaged by the phytonematodes are not efficient in the utilization of

available moisture and nutrients from the soil resulting in reduced functional

metabolism. Visible symptoms of nematode attack often include reduced growth of

individual plants. Furthermore, damaged and weakened roots by nematodes are easy

prey to many types of fungi and bacteria, which invade the roots and accelerate root

decay. These deleterious effects on plant growth result in reduced yields and poor

quality of crops. To overcome this effect, management of nematode is, therefore,

important for higher yield and quality that are expected from the higher cost of crop

production (Vadhera and Shukla, 2002; Kumar and Ja in , 2007).

Once nematodes are established in the field, the possibilities of complete eradication


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are exceeding remote and impractical on field scale due to soil inhabitant nature of

nematodes. However, several measures are adopted to decrease nematode population

up to an acceptable level. The main objectives of phytonematodes management are

usually a matter of reducing the nematode population by one or more methods or

integration of one or more methods to a low level so that the damage is negligible or

at an economically acceptable level (Rajvasnshi and Sharma, 2007).

Integrated Nematode Management (INM) evolved as a philosophy and

technique for the alleviation of real, potential or perceived problems associated with

nematode management programmes (Allen and Bath, 1980; Akhtar, 1997; Sikora et

al., 2005). The integrated nematode management strategy uses a combination of

different disease control methods to decrease disease, increase yield, minimize

environmental damage, prevent the buildup of resistant pathogen strains and produce

high quality products (Widmer et al., 2002).

Efficient management of plant parasitic nematodes requires the carefully

integrated combination of several methods. Although each individual method of

management has a limited use, together, they help in reducing the nematode

populations in agricultural soils or in plants. Integrated pest management (IPM)

provides a working methodology for pest management in sustainable agricultural

systems (Oka, 2010). One of the main objectives behind this work was to observe the

application of current methods for the management of plant-parasitic nematodes

within the guidelines of IPM. Integrated nematode management can broadly be

divided in cultural, chemical, biological, physical, host resistant and integrated crop

protection system, in which the best combination of resistant cultivars, crop rotation,

organic amendment and soil solarization can be utilized with minimum use of

nematicides.


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2.1. CULTURAL CONTROL

Control of nematodes by cultural practices is more economical or practical

especially on vegetable crops. Numerous cultural practices can be beneficial by

reducing population densities of plant-parasitic nematodes. These practices include

fallowing, cover crops, trap crops, antagonistic plants, weed management, flooding,

conservation tillage, green manuring, cropping systems etc. This operation is feasible

and can be adopted and carried out without extra expenses. Secondly the product can

be consumed at any time after harvest, as there are no residual effects of the chemical.

2.1.1. Crop rotation

Seasonal rotations of susceptible crops with non-host or poor-host crops in the

same area of land remain one of the most important techniques used for nematode

management worldwide (Viaene et al., 2006). By this method, the populations of

phytonematodes are reduced to a minimum level. This process should be repeated for

several years depending upon the initial population and decease rate of population.

The rotation must also provide economically useful crops. Choosing rotation, care

should be taken to avoid a new set of pathogens in place of the one to be controlled. A

number of crops and other plants have been found resistant to phytonematodes

(Stefanova and Fernandez, 1995; Gomez and Rodriguez, 2005; Rehman et al., 2006).

Use of Witchgrass in a peanut rotation has beneficial effects on soil, reducing

parasitic nematode populations (Kokalis-Burelle et al., 2002).

Similarly, rotation crops, such as beans, bahiagrass, maize and cabbage that

support extensive growth of the nematophagous fungus, Pochonia chlamydosporia in

their rhizosphere but support only limited reproduction of root-knot nematodes, are

used to maintain the abundance of the fungus in the soil whilst suppressing

populations of the nematode (Timper et al., 2001; Puertas and Hidalgo-Diaz, 2007).


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Hence, growing an approved crop in the rotation to maintain populations of natural

enemies on roots is another alternative to improve the efficacy of nematode

management programmes based on crop rotations (Rodríguez-kábana and Canullo,

1992).

The crop rotation may provide a short-term suppression of nematode

population densities (Starr et al., 2002). However, due to the polyphagous nature of

the pest as well as the relatively low economic value of some recommended rotational

crops, control of root-knot nematodes by crop rotation becomes very limited (Waceke

et al., 2001). It has been reported by several workers that different cropping sequences

reduce the populations of some harmful phytonematodes to the levels that do not

cause economic losses (Alam et al., 1981; Idowu and Fawole, 1989; Singh et al.,

1997; Haider et al., 2001; Haider and Pathak, 2001).

The crop rotation to a non-host crop is often adequate by itself to prevent

nematode population from reaching economically damaging levels. However, it is

necessary to positively identify the species of plant-parasitic nematodes in order to

select appropriate crops, which should be poor hosts or non-hosts for the prevailing

nematode species. The population of phytonematodes suppression by crop rotation

has been reported by many workers (Kluepfel et al., 1993) and it can also be induced

by crop rotation with antagonistic plants such as velvet bean (Mucuna deeringiana)

(Vargas et al., 1994) and switchgrass (Panicum virgatum) (Kokalis-Burelle et al.,

1995). Various cover or trap crops and antagonistic plants are useful for reducing

nematode populations as well as conserving soil and often improving soil texture

(Nusbaum and Ferris, 1973; Alam and Jairajpuri, 1990; Abawi and Thurston, 1994).

Haider et al. (2004) reported that the intercropping two rows of yellow sarson

(Brassica campestris var. sarson) with sugarcane were very effective to control the

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nematode disease. Similar results of inclusion of mustard, a poor host for several

nematodes, in different cropping sequences for reducing nematode populations have

been reported by several other workers (Singh and Sitaramaiah, 1993; Kumar et al.,

2006).

Some of the selected non-host plants that can be effectively used in the crop rotation

practice against various plant-parasitic nematodes are listed below.

List of some non/poor hosts that can be included in crop rotation (Minimum of two

year rotation)

Target nematode Non/ poor host

Globodera Wheat, strawberry, cabbage, cauliflower, peas,

maize, beans

Heterodera avenae Pea, maize, carrot, fenugreek, gram, mustard

Meloidogyne, Pratylenchus,

Tylenchorhynchus,Rotylenchulus,

Radopholus, Heterodera

Crotalaria spectabilis, C. striata. Tagetes spp.

Heterodera glycines Maize, cowpea, potato, tobacco, most vegetables

H. schachtii Alfalfa, bean, clover, maize, onion

H. zeae Wide range of crops

Globodera rostochiensis Maize, green beans, red clover

Hoplolaimus indicus Cabbage, chilli, eggplant

Meloidogyne javanica Cotton, groundnut, sorghum, velvet bean

M. hapla Maize, cotton, grasses, lettuce, onion, radish

M. incognita Fescue, orchard grass

Meloidogyne spp. Crotalaria spectabilis, millet, oats, wheat

Paratrichodorus minor Maize, Crotalaria spectabilis

Pratylenchus leiocephalus Groundnut

P. penetrans Alfalfa, beet, fescue, marigold, oats, rye

Pratylenchus spp. Lettuce, onion, radish

Radopholus similis Crotalaria spectabilis

Tylenchorhynchus mirzai Wheat

T. brassicae Potato, tomato

Xiphinema americanum Alfalfa, maize, fescue, tobacco

Source: Trivedi and Barker (1986)

Prasad et al. (2004) found the highest linseed equivalent when linseed was

intercropped with mustard followed by gram. The decrease in nematode populations

by intercropping mustard could be attributed to the presence of 2-propenyl

isothiocynate in mustard having nematicidal activity as reported by Kowalska and

Sonalinska (2001).


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Recently, Sundararaju (2005) reported that the maximum reduction in root-

lesion index and nematode population was observed where marigold (Tagetes erecta)

was grown as an intercrop and was at par with chemical treatment. Similar findings

were also reported by several workers who reported that intercropping marigold with

different crops can reduce the population of plant-parasitic nematodes thereby

exhibiting a better plant growth (Yen et al., 1998; Dhanger et al., 2002; Uma Shankar

et al., 2005).

Some of the cover/trap crops, which can be used for managing plant-parasitic

nematodes, are listed below.

Nematode species Trap crop Reference

Meloidogyne spp.

Crotalaria spectabilis,

Cowpea, English pea,

Periwinkle, Tagetes

minuta, Ricinus communis

Christie, 1959; Godfrey and

Hagan, 1934; Patel et al., 1991;

Owino and Waudo, 1995.

Heterodera avenae Oat Stone, 1961

H. schachtii Hesperis matronalis Moriarty, 1961

Globodera spp. Potato Carroll and McMahon, 1939

Vetrivelkalai and Subramanian (2006) observed that the population dynamics

of several plant-parasitic nematode species reduced sharply during the fallow period

in all the cropping sequences viz., sorghum-fallow, tomato-fallow, cotton-fallow and

black gram-fallow. The least population of M. incognita was observed during the

cropping period but not recovered during the fallow period in tomato-fallow and

cotton-fallow cropping sequences. Similar results were also reported by Wani (2005)

who observed that the cropping-sequence wheat-chilli-fallow caused the greatest

reduction in the nematode population followed in the descending order of efficiency

by chickpea-okra-chilli, mustard-mung-tomato and tomato-fallow-okra, however, the

extent of field ploughing also playing an important role and deep ploughing being

more effective than normal ploughing. Similarly, Cabanillas et al. (1999) reported that

sorghum-fallow and cotton-fallow reduced R. reniformis populations.


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2.1.2. Biofumigation

The term biofumigation is used when volatile substances are produced through

microbial degradation of organic amendments that result in significant toxic activity

towards nematodes or diseases (Bello et al., 1997). Generally, biofumigation is more

effective when there is an optimum combination of organic matter, high soil

temperature and adequate moisture to promote microbial activity. In Spain,

biofumigation has been largely applied successfully as an alternative to methyl

bromide in several crops (Bello et al., 2001).

Soil amended with fresh or dry cruciferous residues reduce significantly root-

knot nematode infestations, principally, due to isothiocyanates released in soil when

glucosinolates present in these crop residues are hydrolyzed (Stapleton and Duncan,

1998; Diaz-Viruliche, 2000; Ploeg and Stapleton, 2001; D’Addabbo et al., 2005).

However, the practical application of this approach is limited due to the large amount

of organic matter to be transported to the field or the cost of cover crops to be

incorporated into the soil, together with the plastic mulch and drip irrigation system

often necessary to improve the effectiveness of biofumigation. Also, the provision of

large amounts of nutrients to soils may affect the activity of facultative parasites of

nematodes. Among non-chemical alternatives, biofumigation, based on the use of

gasses resulting from the decomposition of organic matter, has demonstrated great

efficacy as an alternative to the use of methyl bromide, whose ban is imminent due to

its environmental impact (Bello et al ., 2003).

2.1.3. Tillage practices and Fertilizer application

Tillage practices play an important role in integrated nematode management

approaches, integration of tillage practices with other nematode management practices

viz., organic amendment, nematicides and crop rotation, soil solarization are very


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effective to the control of phytonematodes. Consistent effects of tillage practices on

nematode population densities have been reported by many workers (Minton, 1986;

McSorley and Gallaher, 1993) and greater effect was obtained from deeper ploughing

(Siddiqui, 2005, 2007; Ahmad et al., 2007b). Although tillage practices may be of

some benefit in nematode management, much more success has been achieved

through the design of effective cropping systems (McSorley and Gallaher, 1994).

Roget et al. (1987) have shown that the number of cysts of Heterodera avenae on

roots and the amount of damage caused by the nematode on wheat are reduced by

conservation tillage. Depth of ploughing influences the populations of plant-parasitic

nematodes; a greater reduction in nematode numbers was observed in deep-ploughed

than in normally ploughed agricultural soils (Tiyagi and Alam, 1995). Nematode

densities have shown to be greater in deep ploughed compared with normal ploughed

plots (Fortnum and Karlen, 1985; Jain and Bhatti, 1985; Mathur et al., 1991; Akhtar,

1997; Siddiqui, 2003, 2007; Anver, 2006). Siddiqui and Alam (1991) reported that the

depth of ploughing had a great influence on the population of plant parasitic

nematodes. The deep ploughing brought about a significant reduction in the

population of plant parasitic nematodes over normal ploughing treatment. The

combined effect of organic amendment/nematicides and ploughing reduced the

population level of phytonematodes (Siddiqui, 2007). The suppressive effect of deep

ploughing has been reported by several workers (Siddiqui and Alam, 1991; Anver,

2006).

It has been suggested that the deep ploughing disturbs the ecological set up of

nematodes which are exposed to external unfavorable conditions and thus their

population decline (Siddiqui and Alam, 1999). Siddiqui and Alam (2003) reported

that the integrated effect of ploughing, nematicides and organic amendment on the


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population of phytonematodes were very effective and it reduced to an acceptable

level. The population of Meloidogyne spp. decreased drastically when the field was

puddled every year over a 10 year period (Pankaj et al., 2006). Siddiqui and Alam

(1999) reported that the integrated effect of oil cakes, nematicides and ploughing were

very effective in reducing the population of phytonematodes and promoted the plant

growth characters.

Badra (1980) stated that ammonical nitrogen maintained in an amount up to

576 kg/ha at two intervals decreased damage caused by R. reniformis to tomato.

Additional experiments with urea, which is readily converted to ammonia by urease

present in the soil, showed that it is also a good nematicide if applied at levels in

excess of 300 mg/kg soil (Rodriguez-Kabana and King, 1980; Huebner et al., 1983).

Ismail et al. (2006) also reported that the soil and root population of R. reniformis

which reduced steadily with increasing the doses of nitrogen incorporated organic

amendments. Slight increase in infestation and invasion of R. reniformis was

associated with lower doses of potash. Incorporation of nitrogen and phosphorus was

promising for plant growth as well.

2.2. ORGANIC SOIL AMENDMENTS

Organic soil amendments can be successfully employed for the control of

plant parasitic nematodes. A variety of organic amendments, such as animal and green

manures, compost, nematicidal plants and proteinous wastes are used for the control

of phytonematodes (Oka, 2010). Application of organic amendments into the soil is

not only beneficial to nematode management but also improving the plant growth and

productivity (Adegbite and Adesiyan, 2005). On the other hand, application of

organic substrates leads to build up of beneficial micro flora around the rhizosphere,

which will help to reduce the plant parasitic nematodes in the soil (Oka, et al., 2007).


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2.2.1. Antagonistic crops and Composts

Plants antagonistic to nematodes are those that are considered to produce

toxic substances, usually, while the crops are growing or after incorporation into the

soil. In nematode management strategies the use of this approach relies on preplant

cover crops, intercropping or green manures. Marigold, neem, sunnhemp, castor bean,

partridge pea, asparagus, rape seed, crotalaria and sesame have been extensively

studied and used as antagonistic crops for nematode control (Wang et al., 2002,

2003).

Germani and Plenchette (2004) recommended the use of Crotalaria spp. as

precrops for providing green manure while at the same time decreasing the level of

root knot nematode and increasing the level of beneficial mycorrhizal fungi.

Incorporation of plant residues generally increases the number of free-living

nematodes, but increases in specific nematode genera may be affected by plant

residue type (McSorley and Frederick, 1999), which in turn may affect antagonistic

organisms, such as predatory nematodes and parasitic fungi. Incorporation of

sunnhemp (Crotalaria juncea) to soil increased nematode-trapping fungi, parasitic

fungi on R. reniformis eggs, vermiform stage parasites and bacterivorous nematodes

more efficiently than amendments with Brassica napus or Tagetes erecta (Wang et

al., 2001; Wachira et al., 2009). The population of R. reniformis was highly reduced

in growing mustard, wheat, fennel, carrot, sorghum, sesbania (Siddiqui and Alam,

2001)

Chrysanthemum coronarium significantly reduced root-knot nematode

infection of tomato roots and improved plant fresh weight (Bar-Eyal et al., 2006). The

use of soil amendments with organic matter to control nematode populations has been

reported by several researchers (Ferraz and Freitas, 2004; Halbrendt and Lamondia,


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2004). Similarly, the results of soil amendment with neem, Jatropha and castor leaves

are very effective (Kalaiarasan et al., 2007; Lopes et al., 2011).

Ritzinger and McSorley (1998) amended the soil with different quantities of

dry castor leaves and concluded that application at the rate of 0.5% was sufficient to

significantly reduce the population of M. arenaria Neal (Chitwood). The nematicidal

properties of castor plant have been attributed to ricin, a substance found only in the

seeds. Some of the plant parts such as roots, shoots, leaves, flowers and crop residue

are left intentionally in the field after harvest and ploughed deep into the soil, these

organic additives after their proper decomposition by the activity of several

microorganisms resulted in the suppression of many plant pathogens like plant-

parasitic nematodes (Johnson, 1971, 1972; Siddiqui and Alam, 1995, 1997, 1999).

The incorporation of mature dried residues of lespedeza, alfalfa, oats and flax into the

soil infested with M. incognita, significantly reduced the incidence of the nematode

on tomato (Johnson et al., 1967).

Cassava peelings, cocoa pod husk and rice husk significantly reduced

Meloidogyne spp. population infecting cowpea (Egunjobi, 1985; Egunjobi and

Olaitan, 1986). Cassava leaf and tuber rind applied as soil amendment @100g or 50

g/pot, significantly reduced the population of M. incognita and improved plant growth

parameters of okra (Ramakrishnan et al., 1999). Akhtar and Alam (1993) recorded

that vegetables, fruit processing waste and tobacco waste were most effective in

reducing the incidence of root-knot and the population of phytonematodes on tomato.

Soil amending with spent tea, wheat straw, paddy husk, sugar cane and domestic

garbage were beneficial in controlling the nematodes.

Some researchers reported that poultry refuse and mustard oil-cake were

effective in controlling root knot nematode and enhancing plant growth and yield of


41

tomato (Faruk et al., 2001, 2002) and many other crops (Bari et al., 2004a, 2004b).

The accelerated yield of tomato under field conditions has been reported by other

investigators using higher doses of organic soil amendments alone or their application

at lower dose mixed with Furadan 5G (Faruk et al., 2001, 2002). Soil amendment

with poultry refuse has also been reported to be effective against root-knot nematode

of okra (Bari et al., 1999), brinjal (Bari et al., 2004a). Organic amendments like

sawdust of neem and mango greatly reduced root-knot development and

multiplication of R. reniformis on tomato and eggplant and Tylenchorhynchus

brassicae on cabbage and cauliflower. The nematode control gradually increased with

increasing the dose of sawdust. Sawdust of neem was more efficacious than that of

mango. The combined effect of sawdust and ammonium sulphate was greater than

either of the separate both with respect to nematode control and to the improvement in

plant growth (Siddiqui and Alam, 1990).

The phytotoxicity of sawdust was effectively eliminated by supplementing the

sawdust with ammonium sulphate (Siddiqui and Alam, 1990). The soil amending with

different parts of neem/margosa (Azadirachta indica A. Juss.) is reported to be highly

effective in reducing the population of different plant-parasitic nematodes affecting a

variety of plant species (Hellap and Dreyer, 1995; Rao et al., 1996; Zaki, 1998;

Umamaheshwari and Sundarababu, 2001; Siddiqui and Alam, 2001; Oka and Pivonia,

2002; Yasmin et al., 2003; Shah et al., 2004; Raman and Venkateshwarlu, 2006;

Siddiqui, 2006a; Rather and Siddiqui, 2007a, b, c).

Although possessing a limited scope to be used as manure, the sawdust has

been suggested for the pest management and control of plant-parasitic nematodes. A

significant reduction in the intensity of root galling was reported (Srivastava et al.,

1971) when sawdust applied in a field planted with okra, eggplant and tomato.


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However, various other workers have pointed out the reduction in nematode intensity

by amending the soil with sawdust (Bora and Phukan, 1983; Singh et al., 1986;

Osunlaja, 1990; Akhtar, 1998). Consequently, it becomes more advisable to employ

sawdust in combination with other materials such as different oil cakes, sugarcane

bagasse, nematicides, urea, cow dung and other biocontrol agents, as a nematode

suppressant (Kushwaha et al., 1983; Acharya and Padhi, 1988; Akhtar and Alam,

1993).

Soil organic matter helps to retain nutrients, maintain soil structure and hold

water for plant use. The addition of various rice straw composts on the rhizosphere

soil microorganisms showed a high fertilizer value when applied @ of 5% (w/w).

Various rice straw composts @ 5, 7.5 resulted in reducing root-knot nematode

population of 79, 84% respectively and actualized prodigious depletion in egg

production (Rashad et al., 2011). Organic amendment viz., rice husk, saw dust, cow

urine, cow dung and neem cakes have been significantly effective against M.

incognita (Shurtleff and Averre, 2000; Singh and Khurma, 2007; Nagraju et al.,

2010).

Animal manures have been used since the beginning of agricultural food

production to improve soil fertility, recycle nutrients, improve biological and physical

properties of soil and increase crop yield (Rodriguez-Kabana et al., 1987; Sims and

Wolf, 1994). The research with animal manures amended in the soil, have shown that

they possess nematode-suppressive properties (Montasser, 1991; Kalpan and Noe,

1993; Opperman et al., 1993; Stephan, 1995; Oka and Yermiyahu, 2002). The mode

of action, however, has not yet been fully determined. The application of manure

enhances soil fertility, aids in controlling plant-parasitic nematodes and provides a

mean of disposing of the manure.


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Many previous reviews have focused on the use of organic amendments to

control plant-parasitic nematodes (Rodríguez-Kábana, 1986; D’Addabbo, 1995;

Akhtar and Malik, 2000; Oka, 2010; Thoden et al., 2011). Farm manure trials have

frequently involved poultry or cattle litter. Poultry litter appeared to be an appropriate

choice (Gamliel and Stapleton, 1993a), especially when combined with a sorghum

cover crop (Everts et al., 2006). Abubakar et al. (2004) reported that soil amending

with cow dung, urine and their mixture significantly reduced the extent of root-galling

and nematode multiplication of root-knot nematode, M. incognita and improved the

various plant growth parameters of tomato. Similar results of reduction in nematode

populations by soil amending with cow dung were also reported by Babatola (1990)

and Abubakar and Majeed (2000).

Chicken litter, a common form of poultry manure, consists of manure and pine

shaving beddings, contains significant quantities of N, P, K, Ca, Mg and

micronutrients and can be used as a substitute for commercial fertilizers (Ndegwa et

al., 1991). Several researchers have reported that the chicken litter when applied to

the soil as an organic amendment will lower the densities of plant parasitic nematodes

(Gonzales and Canto-Saenz, 1993; Owino and Waudo, 1995; Riegel et al., 1996;

Riegel and Noe, 2000; Ravichandra et al., 2001; Ribeiro et al., 2002; Ami and Al-

Sabie, 2004). This suppression of nematodes is probably a combination of enhanced

microbial activity and constituent toxicity. The majority of nitrogen in poultry manure

is in the form of uric acid that can be rapidly converted to ammonium nitrogen if

temperature, pH and moisture are suitable for microbial activity (Sims and Wolf,

1994). The ammonia produced has been shown to kill plant parasitic nematodes (Eno

et al., 1993). The presence of pine shavings in litter serves as a carbon source and


44

reduces phytotoxicity caused by the accumulation of ammonia and nitrates (Huebner

et al., 1983).

The addition of the agro-industrial wastes to the soil can exert a remarkable

suppressive action on phytonematodes (D’Addabbo, 1995). The release of toxic

compounds, performed or derived from the degradation of the wastes in the soil,

and/or the multiplication of nematode predators and/or parasites on the organic

substrate are supposed to be the fundamental mechanisms of this nematicidal action

(Stirling, 1991). Moreover, many of the currently available nematicides besides being

often costly offer no long-term suppression and having differential effects on the

species of nematodes as their activity is affected by many environmental factors

(Schmitt, 1986; Starr et al., 2002).

Akhtar and Mahmood (1996) reported that amending the soil, naturally

infested with different plant-parasitic nematodes, with cellulosic wastes and other

waste materials such as oil seed cakes, chitin, compost, livestock and poultry

manures, can be effectively employed against the damage caused by these plant

parasitic nematodes.

Addition of fly ash into the soil cause changes in its physical and chemical

characteristic and is supposed to increase concentration of carbonates and

bicarbonates (Khan and Khan, 1996). The reduction in the number of galls per plant

was very high with nematode inoculated plants grown at 10 and 20% fly ash levels, as

against M. incognita inoculated plants alone (Azam et al., 2007). Pasha et al. (1990)

reported decreased soil population of M. incognita at 10-100% fly ash levels. Niyaz

and Hisamuddin (2010) observed suppression in the morphmetrics of M. incognita

females and egg mass production with increasing fly ash levels.

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The soil application of organic amendments viz., vermicompost, neem cakes,

castor cakes, groundnut cakes, sunflower cakes and FYM have significantly reduced

the nematode population and increased the plant growth compared to inoculated

control (Jagadeeswaran and Singh, 2011). Compost is an organic matter resource

resulted from exploiting wastes through the controlled bioconversion process. It

seems to meet the objectives of the alternative agriculture system and the growing

consensus of both environmentalists and those concerned with the public health

through solving the waste disposal problem and its application in sustainable

agriculture instead of ecologically undesirable mineral fertilization (Ndiaye et al.,

2000; Madejon et al., 2001; Melero et al., 2007; Courtney and Mullen, 2008;

Chitravadivu et al., 2009).

Vermicompost is a new form of organic soil amendment that has considerable

potential in crop production. Vermicompost has large surface areas that provide many

micro sites for microbial activity and for the strong retention of nutrients.

Additionally, vermicompost have been reported to have outstanding biological

properties and have microbial populations that are significantly larger and more

diverse compared with those of conventional thermophilic composts (Edwards, 1998).

Vermicompost and goat manure were significantly better alternatives in eco-

friendly management of M. incognita and they induce the growth of seedling of

tomato at its early stages due to high amount of nitrogen (1.94% in vermicompost and

4.9% in goat manure) which is important for plant growth (Pakeerathan, et al., 2009).

The organic matter amendments significantly improved plant growth of groundnut

and reduced the disease infection by phytonematodes. Fresh Azolla ranked the best in

improving plant growth and reducing host infestation significantly. Dry Azolla is also

improving plant growth against plant parasitic nematodes (Joshi and Patel, 1995).


46

2.2.2. Oil cakes and Green manuring

Many neem preparations, including leaves, oil cakes and kernel oils have been

tested for their nematicidal activity (Akhtar, 2000a) and the insecticidal limonoids

from kernel oils, including azadirachtin, nimbin and salannin have been reported to be

the main nematicidal compounds as well (Akhtar, 2000a). A neem extract containing

10% azadirachtin was nematicidal to Meloidogyne javanica juveniles in alkaline

sandy soil (pH 8.5) at concentrations higher than 0.05 g kg-1

, or at higher than 0.17 g

kg-1

for its formulation (Oka et al., 2007).

Oka et al. (2000a) reported that twelve of twenty seven essential oils extracted

from spices and aromatic plants immobilized more than 80% of juveniles of root-knot

nematode, M. javanica at a concentration of 1000 l/litre and at the same

concentration most of these oils also inhibited nematode hatching. The essential oils

of Carum carvi, Foeniculum vulgare, Mentha rotundifolia and Mentha spicata

showed the highest nematicidal activity in vitro and those from Origanum vulgare,

O. syriacum and Coridothymus capitatus reduced root-galling of cucumber seedlings

when mixed with sandy soil.

Many essential oils of medicinal plants and herbs have been reported

nematicidal (Oka et al., 2000a; Park et al., 2005), but the effect of their plant

materials when incorporated into the soil has not been well studied. Soil amendments

with several oriental herbal medicines at 0.2% (w/v) reduced M. incognita infection

on tomato (Kim et al., 2003).

In reducing the nematode population associated with Pomegranate by oil

cakes, are in agreement with Singh and Sitaramaiah (1970). The oil cakes apart from

contributing to NPK are greater benefit in the agriculture, which none of the synthetic

fertilizer or pesticide can offer. They provide slow and steady nourishment protection;


47

create antagonistic conditions for pathogens including soil nematodes (Khan et al.,

2011). Oil cakes retard nitrification of the soil/urea and thereby increase N uptake by

the plants (Tiyagi et al., 2002) oil cakes containing 2-7% of protein N applied at @ of

4-10%suppress soil nematodes and improve plant tree health.

Abd-Elgawad and Omer (1995) explored the essential oils of four medicinal

plants for phytonematode control. All the oils inhibited nematode mortality but

Mentha spicata was generally more effective in reducing the number of active

nematodes followed by Thymus vulgaris, Majorana bortensis and Mentha longifolia.

Soil stages of the reniform nematode were more affected by the oil than those of the

ring and lance nematodes. The content of oxygenated compounds in these oils ranged

from 45.79% to 96.50% and may be partially responsible for the nematicidal effects.

Pandey (2000) reported the nematicidal activity of eight essential oils against

root-knot nematode, M. incognita at four different concentrations viz., 2000, 1000,

500 and 250 ppm. Maximum nematicidal activity was recorded in oils of Eucalyptus

citriodora, E. hybrida and Ocimum basilicum followed by Pelargonium graveolens,

Cymbopogon martini, Mentha arvensis, Mentha piperata and Mentha spicata oils

respectively, however, eucalyptus and Indian basil oils were highly toxic even at

lower concentrations (500 and 250 ppm).

Green manuring is an essential and age old practice of Indian farmers as well

as the farmers throughout the world, wherein the green plants/plant parts are ploughed

deep into the soil to rot and provide nutrient for succeeding crops, however, it has also

been found to reduce the populations of plant-parasitic nematodes. It was Lindford et

al. (1938), for the first time who reported that incorporating chopped pineapple leaves

@50-200 tonnes/acre into the soil significantly reduced the root-knot incidence in

cowpea. The infested soil when amended with chopped cabbage leaves (Brassica


48

oleracea capitata L.) @ 680g/0.61m2, reduced the cereal root eelworm (Duddington

and Duthoit, 1960; Duddington et al., 1961). Hutchinson et al. (1960) noticed that the

population of some plant-parasitic nematodes like Hoplolaimus, Pratylenchus and

Tylenchorhynchus spp. were significantly lower in the soil where pieces of pumpkin

(Cucurbita pepo L.) were allowed to rot compared with the rest of the soil. Mankau

(1968) found that the application of alfalfa (Medicago sativa) green manure in root-

knot infested field was found to be a good nematode suppressant. The application of

rapeseed green manure @200, 300 and 400 mg N/kg soil was more effective than

velvet bean green manure in reducing root-galling caused by M. arenaria in squash

roots (Crow et al., 1996). Sudangrass has been reported to suppress infection and

damage caused by M. hapla, when incorporated as a green manure (Widmer and

Abawi, 2000). This addition of green manures stimulates soil microbial activities and

increases accumulation of plant decomposition products and microbial metabolites

that can be deleterious to nematodes.

Alternatively, Djian-Caporalino et al. (2002) identified 39 species of green

manures that belong to 22 botanical families, including peanut (Arachis hypogaea),

basil (Ocimum basilicum), cotton (Gossypium hirsutum), sesame (Sesamum

orientale), oat (Avena sativa) and rye (Secale cereale). But the most efficient were

sudangrass and sorghum (Sorghum sudanense), cruciferae, like oil radish (Raphanus

sativus) and rapeseed (Brassica napus), ricin (Ricinus communis), marigold (Tagetes

erecta, Tagetes patula, Tagetes minuta), Juglans regia, and velvet bean (Mucuna

deeringiana) (Crow et al., 1996; Bridge, 1996; Al-Rehiayani and Hafez, 1998;

Widmer and Abawi, 2002; McKenry and Anwar, 2003; Everts et al., 2006).


49

2.2.3. Nematicidal activity of plant extracts

Many plants possess nematicidal and nematostatic properties in their roots,

shoots, leaves, flowers, seeds and their extracts, essential oil, oil seed cake and

derivatives. Some of the plant species and parts antagonistic to Meloidogyne spp. are

leaves and flowers of marigold (Tagetes sp.), leaves, roots and seeds of neem

(Azadirachta indica), leaves and seeds of chinaberry (Melia azedarach) (Rather et al.,

2007). Essential oils and plant extracts of sweet wormwood (Artemisia obsinthium),

thyme (Thymus vulgaris), peppermint (Mentha spicata), fennel (Foeniculum vulgare),

garlic (Allium sativum) and Eucalyptus spp., were toxic and reduced hatching activity

of phytonematodes (Ibrahim et al., 2006). Among these plants, marigold (Tagetes sp.)

is the most commonly studied. As marigold belongs to the Asteraceae, it is possible

that other members of the family may also possess antagonistic properties against

plant-parasitic nematodes (Tsay et al., 2004). Neem (Azadirachta indica) is one of the

most common nematicidal plants that were recognized in the middle of the 20th

century. Simple home made products like neem seed powder, neem seed kernel

powder, neem seed cake powder, dry neem leaf powder and the appropriate aqueous

extracts made from neem are available (Javed et al., 2006).

Plants appear to be a source of effective pesticide compounds and may be

regarded as an inexhaustible source of harmless pesticides having a low plant and

human toxicity and being easily biodegradable (Prakash and Rao, 1997).

Consequently, a large number of plants/plant parts/plant products have been screened

for their nematicidal activities (Pandey, 1990; Eyal et al., 2006). Although most

researchers have investigated the non-volatile constituents of the plants for their

nematotoxic potential (Sangwan et al., 1990; Ghosh and Sukul, 1992), but little


50

attention has been given to volatile constituents of essential oil-bearing plants

(Kochhar, 2006).

Meena et al. (2010) reported that acetone extracts of leaf, flower, roots and

stem of five different varieties of Tagetes were found to be highly effective in causing

mortality of juveniles and inhibition of egg hatching. Acetone extract of T. erecta cv.

Indian Yellow was found to be highly effective in suppressing M. incognita. The

medicinal plants are showing the nematicidal properties so they are widely used for

the control of plant parasitic nematodes.

Joymadidevi (2010) reported that the nematicidal effect of medicinal plants as

chloroform methanol extract on egg hatching and larval mortality of M. incognita

were very effective. The rate of hatching was directly proportional to the

concentration of extracts. Several plants, belonging to different botanical families,

contain principles possessing nematicidal or nematostatic properties (Grainge and

Ahmad, 1988). Khanna and Kumar (2006) reported that the effect of five neem based

pesticides on the mortality of juveniles and egg hatch of M. incognita. Recently many

workers used plant extract showing nematicidal properties for seed treatment or bare

root dip treatment. The applications of methanolic extract of botanicals are very

effective in the control of phytonematodes (Usman and Siddiqui, 2011a, 2012b).

Many researchers used different plant extracts as a seed treatment or bare root

dip treatment for the control of M. incognita and other nematodes (Siddiqui and

Alam, 1988a, b; 1989a, b). Hussain et al. (1984) showed that root dip treatments of

eggplant seedling with Margosa and Marigold leaf extracts considerably reduced root

knot nematode development as compared to treatment with Cina, Piprazine citrate,

Chenopodium oil and Ground nut cake. Abid and Maqbool (1991) showed bare root

dip treatment in the leaf extracts and neem leaves significantly reduced root-knot


51

infection caused by M. javanica on tomato and egg plant. The damaging effects of the

nematode were masked by bare root dip treatments as shown by improved plant

growth in both the test plants. Usman and Siddiqui (2011a) evaluated the leaf extracts

of Murraya koenigii L. and Vitex negundo L. as bare-root dip treatment for the

management of M. incognita infecting tomato (Lycopersicon esculentum) and chilli

(Capsicum annum) plants. Significant reduction was observed in the root-knot

development caused by M. incognita on the experimental plants. Leaf extracts

of Murraya caused relatively higher inhibition in root-knot development and

nematode multiplication than Vitex.

Various compounds such as nimbin, nimbidin, azadirachtin, salannin,

thionemon and meliantriol occur in the seeds, leaves and bark of neem in high

concentrations (Kraus,1995) and are responsible for the antimicrobial and nematicidal

activity (Eppler,1995). Powder from the seed kernels and leaves has been found to be

suppressive against some nematodes (Alam, 1993). Tiyagi et al. (2009) studied the

effect of leaf extracts of two latex-bearing plants such as Calotropis procera and

Thevetia peruviana as a bare-root dip treatment for the management of

phytonematodes, M. incognita and R. reniformis infecting tomato and plants. A

significant reduction was observed in the root-knot development caused by M.

incognita and nematodes multiplication of R. reniformis on the experimental plants.

Khan et al. (2011) showed that the curative effect of plant extracts viz., neem,

tobacco, Aloe vera, chilli, clove, garlic and onion against M. incognita. Similarly

significant reduction was observed in the population of plant-parasitic nematodes, M.

incognita, R. reniformis and T. brassicae infesting eggplant and cauliflower, when the

seedlings were given the root-dip treatment in leaf extracts of Argemone maxicana

and Solanum xanthocarpum (Ajaz and Tiyagi, 2003).


52

In addition to this, a number of indigenous plants have been reported to

possess nematicidal/nematostatic properties and thus, are capable of managing the

populations of various plant-parasitic nematodes (Zarina and Shahina, 2010). Some of

these plants tested for their antinemic properties by different workers include Abutilon

indicum, Solanum forskalii, S. nigrum and Xanthium strumarium (Shaukat and

Siddiqui, 2001). Allium cepa, Aloe vera, Calendula officinalis, Capsicum annuum,

Syzygium aromaticum and Nicotiana tabacum (Khan et al., 2008a), Asparagopsis

taxiformis (Rizvi and Shameel, 2006a), Avicennia marina (Tariq et al., 2007).

Azadirachta indica, Fresh leaves, extract, ethanolic extract, dry leaves powder, seed,

seed decoction, oil cake, product, oil, derivatives and formulation (Javed et al., 2007a,

b; Jiskani et al., 2005), Azadirachtin, Neem formulation (Javed et al., 2007c).

Bauhinia alba, Datura alba, seed decoction and Eucalyptus alba, leaves, bud, stem

and fruit (Pathan et al., 2002a). Bauhinia purpurea, Calotropis procera, Datura

fastuosa and Azadirachta indica (Zarina et al., 2003). Calendula officinalis,

Helianthus annuus, H. bipinnatus, Tagetes erecta and Zinnia elegans (Siddiqui et al.,

2005). Calotropis procera, extract of flowers, leaves, stem root and cake (Pathan et

al., 2002b). Cassia fistula, Daucus carota, Fumaria indica, Heliotropium

curassavicum, H. tuberculosum, Hibiscus rosa sinensis, Justicia adhatoda, Lactuca

remotiflora, Melia azadirachta, Mimosa hamata, Withania somnífera, S. surattense,

Euphorbia hirta and E. tirucalli (Abid et al., 1997). Catharanthus roseus

(Husan Bano et al., 1999); Cystoclonium purpureum and Iyengaria stellata (Rizvi

and Shameel, 2006b); Eclipta prostrata (Shaukat et al., 2004; Zarina et al., 2006).

Eucalyptus sp. (Dawar et al., 2007); E. camaldulensis, E. pulcherrima, F. religiosa,

Ficus benghalensis, F. elastica and Opuntia imbricata (Zurreen and Khan, 1984);

Jania capillacea, Sargassum binderi and Solieria robusta (Ara et al., 1996); Lantana


53

camera (Ali et al., 2001; Shaukat et al., 2003); Nerium oleander (Aziz et al., 1995a);

Nigella sativa (Shajia and Shahzad, 1998); Ricinus communis (Ahmad et al., 1991);

Sargassum spp. (Ara et al., 1997); Schweinfurthia papilionacea (Khan et al., 1997,

Khurram et al., 1997); T. patula (Husn Bano, 1999; Siddiqui et al., 2005); Zingiber

officinale (Zareen et al., 2003); Linum usitatissimum and Brassica campestris (Butool

et al., 1998), Salvia spp. (Idowu, 1999), Parkia biglobosa (Umar and Jada, 2000),

Murraya koengii (Pandey, 2000), Catharanthus rosea and Ipomea fistulosa (Hassen

et al., 2003). Blechum pyramidatum, Stenandrium nanum, Furcraea cahum,

Ageratum gaumeri, Ambrosia hispida, Bidens alba, Calea urtricifolia, Acalypha

gaumeri, Croton chinensis, Tephrosia cinerea, Trichilia arborea, T. minutiflora,

Randia longiloba, R. obcordata and R. strandleyana (Cristobal-Alejo et al., 2006),

Parkia biglobosa and Hyptis spicigera (Jesse et al., 2006), Euphorbia tirucalli, E.

neriifolia, Nerium indicum, Thevetia peruviana and Pedilanthus tithymaloides

(Siddiqui, 2006b), Ficus benghalensis and F. virens (Ahmad et al., 2007a), Eclipta

alba, Phyllanthus niruri and Withania somnifera (Usman and Siddiqui, 2012c), P.

hysterophorus, N. plumbaginifolia, A. fatua, C. album, A. retroflexus, C. murale, A.

spinosus and O. corniculata (Usman and Siddiqui, 2012b) .

Recently, Bello et al. (2006) reported the inhibitory effect of water extract of

seed, leaf and bark of five plants viz., Tamarindus indica, Cassia siamea, Isoberlinia

doka, Delonix regia and Cassia sieberiana against the larval hatching of M. incognita.

The standard suspensions inhibited larval hatching by 97% while dilution of S/100

inhibited larval hatch by 3%. The solvent extracts of the plant species viz., Allophylus

cobbe, Lepisanthes tetraphylla, Sarcococca zeylanica and Hedyotis lawsoniae, were

among seven Sri Lankan plants which showed significant nematicidal activity against

M. incognita maintained on tomato plants (Jayasinghe et al., 2003).


54

2.3. PHYSICAL CONTROL

Soil solarization

Solarization traps solar radiation with transparent plastic films placed on the

soil to maximize conversion of heat. First reported by Katan et al. (1976) solarization

has been widely studied. Solarization increases soil temperature by 2-15 0C in warm

climate conditions. Its efficacy depends on the combination of soil temperature and

duration. M. incognita second-stage juveniles were completely killed in a water bath

heated above 38 0C; it took 48 h at 39

0C, but only 14 h at 42

0C (Wang and

McSorley, 2008). Soil solarization with plastic mulches leads to lethal temperatures

which kill plant parasitic nematodes (around 45 0C) and is being used mainly in

regions where high levels of solar energy are available for long periods of time

(Whitehead, 1998). The effect of this approach is reduced with depth, but solarization

for at least 4-6 weeks will increase soil temperatures to about 35-50 0C to depths of up

to 30 cm and, depending on soil type, soil moisture content and prior tillage, will

reduce nematode infestations significantly (Viaene et al., 2006). In Japan and other

East Asian countries, several farmers growing successive crops, such as tomato and

melon susceptible to root-knot nematodes have used solarization in plastic tunnels for

30 days in the summer as an alternative to methyl bromide fumigation (Sano, 2002).

In Cuba, root knot nematode infestations are reduced, in peri-urban and small

organic farm production, using solarization under sub-optimum conditions (Fernandez

and Labrada, 1995) but for subsistence agriculture, the cost of plastic sheeting may be

limiting. The length of time required for effective solarization is a great limitation too,

but it could be reduced when it is used with biofumigation. Infection of M. javanica

by P. penetrans was increased in naturally infested soils in a South Australian

vineyard treated by solarisation and decreased in soils treated with the nematicides


55

oxamyl or phenamiphos but the bacterium did not significantly reduce nematode

populations (Walker and Wachtel, 1988). Similarly, in a cucumber crop in a

glasshouse trial the use of solarization and P. penetrans had an additive detrimental

effect on M. javanica populations (Tzortzakakis and Goewn, 1994). This physical

management was used for the control of root-knot nematode when black and

transparent polyethylene sheet single and double layer is spread on the soil surface for

different duration (Javed, 1992; Javed et al., 1994, 1997).

Soil solarization showed the best potential for reducing Meloidogyne spp and

other soil borne pathogens as it is easy to apply, pollution free , economical and

inexpensive (Calabretta et al., 1991a, b). Soil solarization with different thickness of

polyethylene sheets was very effective in reducing nematode population in the soil

this could be due to the fact that soil covered with polyethylene sheets reduces the

heat convection and water evaporation from the soil to atmosphere results in

formation of water droplets in the inner surface of polyethylene sheets, its

transmittance to long wave radiation is highly reduced resulting in better heating of

the soil (Grinstein et al., 1979; Katan, 1981; Abdel Rahim et al., 1988; Mazza et al.,

1994; Reddy et al., 2001).

The soil solarization also increases the temperature appreciably and the

thinnest polyethylene sheet proves to be efficient to increase the temperature and

better in heating, radiation and transmittance than the thicker one (Aguilar et al.,

1990; Siddiqui and Saxena, 1992; Ostrec, 1993; Siddiqui et al., 1997, 1998).

Soil solarization has been done by heating the soil beneath clear plastic mulch

for 6 weeks so that it reaches temperatures detrimental to soil borne pests

(Katan et al., 1976). This method has been used successfully against plant-parasitic

nematodes and soil-borne pathogens in several crops and regions around the world,


56

especially in hot climates (Katan, 1976, 1981; Stapleton and Devay, 1983)

Approximately 14 hours at 42 °C temperature was sufficient to kill all root-knot

nematodes in sand tubes, but at sub lethal temperatures (40-42 °C) at least 46 hours

were required to kill them all (Wang and McSorley, 2008). Previous research verified

that sunnhemp consistently increases the number of beneficial free-living nematodes

in the soil (Wang et al., 2001, 2002, 2003) but that soil solarization temporarily

suppresses the beneficial organisms (Wang et al., 2006). However, integrating soil

solarization with a leguminous cover crop could reduce the negative impact of soil

solarization on beneficial soil organisms while improving the pest-suppression effect

achieved by cover cropping alone (Wang et al., 2006).

Integration soil solarization with other nematode management viz., organic

soil amendment, ploughing and chemical treatment may have a much more adverse

effect on nematode population (Siddiqui and Saxena, 1992). It has been reported that

the integrated effect of green manuring, organic amendment and soil solarization are

very effective in controlling phytonematodes (Wang et al., 2001; Hooks et al., 2006).

2.4. CHEMICAL CONTROL

The primary advantage of chemical control is that the nematode population is

reduced drastically to very low level within a matter of days after chemical applied.

Most crops are especially vulnerable to nematode attack during seedling stage when

the young root system is becoming established. Crops planted in treated soil develop a

good root system so that usually in case of annual, the crop is matured before the

residual population of nematodes has increased to a damaging level (Haydock et al.,

2006). Seed treatment and nursery treatment offer minimum application of

nematicides in to the soil and provides nematode free seedlings. These means of

chemical control are safer and can be coined as an integral component of integrated


57

nematode management (INM). Vegetable crops accounting for the greatest proportion

of nematicide use and Meloidogyne spp. as the target for approximately half of this

usage (Haydock et al., 2006).

2.4.1. Soil treatment in the main field/Nursery

Application of ethylene dibromide or ethylene dibromide + chloropicrin in

Phaseolus vulgaris, dibromo-chloropropane (DBCP) @ 6 kg in pigeon pea reduces

the soil population of R. reniformis (McSorley and Parrado, 1983; Sharma et al.,

1993). Application of Temik, thimet + nemaphose and nemaphose alone, Nemacur or

Nemacur-Disyston in cotton seedling effectively control R. reniformis population in

soil (Bost, 1985).

Application of fosthiazate 6 or 12 pt/acre before or after bed preparation of

fields of sweet potato increased the yield by 15.6 and 88.9 lb/acre (Mclean and

Lawrence, 1996). Hadian et al. (2011) reported that drenching of soil with bavistin

(0.1%) completely eradicate the nematode population. Combinations of deep

ploughing (up to 20 cm) and nursery bed treatment with aldicarb at 0.4 g per m2 and

main field treatment with aldicarb at 1kg a.i./ha proved effective in the control of

root-knot nematodes in tomato which also registered maximum yield (Jain and Bhatti,

1985). In tomato, application of aldicarb and carbofuran each 1 Kg a.i./ha and in

combination with neem cake and urea each at 10 kg /ha, at transplanting, produced a

maximum yield with lowest gall index (2.5) and nematode population, 90 days after

planting (Routaray and Sahoo, 1985).

Carbofuran has been reported to control nematodes (Di sanzo, 1973). Low

galling index in soybean plants treated with carbofuran by both soil drench and soil

drench + foliar application has been reported (Ajayi et al., 1993). Furthermore it has

been reported that the application of 700 ppm on soyabean plants as soil drench


58

reduced the number of M. incognita eggs that hatched into juveniles (Ajayi et al.,

1993). In an experiment where carbofuran 3G was applied at the rates of 0, 100, 200,

and 300 kg/ha to three hybrid yam varieties in southwestern Nigeria (Adegbite and

Agbaje, 2007) there was an increase in the yield of the three hybrid yam varieties,

which was significantly higher than in the control. The use of carbofuran at planting

was the most effective application timing to reduce M. javanica population levels

(Jada et al., 2011). Many workers (Patel and Patel, 2009; Jada et al., 2011) reported

the effect of nematicides alone and in combination with other management practices.

Similarly efficacy of carbosulfan as soil drench has been reported by Prasad and

Narayana (2000) in sunflower and, Garabedian and van Gundy (1985) in tomatoes

against root-knot nematode. Although chemical agents like carbofuran are efficient in

controlling nematodes (Adegbite and Agbaje, 2007), their persistence may pose

ecological problems (Li et al., 2008). The effectiveness of granular application of

carbofuran and phorate against root-knot nematode of okra has been reported by

several workers (Sitaramaiah and Vishwakarma, 1978; Khair et al., 1983; Sultan and

Singh, 1987; Sheela and Nair, 1988; Jadhav, 1989; Jain, 1990; Gul et al., 1991).

However, Ahmad and Sultana (1981) found that carbofuran was less effective than

aldicarb granules in gall formation. Next effective seed treatments were carbosulfan

and acephate which supported the findings of Patil (1991) and Kathirvel et al. (1992)

who reported its effectiveness as seed treatment and their impact on promoting growth

parameters. Similarly, the effectiveness of seed treatment with acephate was

promising in cotton as reported by Dhoot (1990) and in french bean (Mohan and

Mishra, 1993).

Mishra et al. (1987) also observed that carbofuran at 1 kg a.i./ha was effective

in reducing the nematode population (J2) and increasing fibre yield of jute.


59

Similarly, Senapati and Ghosh (1992) also recorded greatest suppression of root-knot

nematode infestation in jute with the application of carbofuran at 2 kg a.i./ha under a

jute-paddy rotational system. This treatment increased fibre yield by 28% even with

the highest initial soil population density of the nematodes (Khan, 2004).

Many workers tested various nematicides on M. incognita and on the growth

of tomato were in conformity with Minton et al., (1993), Lawrence and McLean

(1995), Giannakou et al., (2005) and Hafez and Sundararaj (2006). These authors

reported that fosthiazate provided excellent control of root-knot nematodes and

increased plant growth and yield. In addition, cadusafos and fosthiazate reduced M.

arenaria population on winter-grown oriental melon from 35 to 90 % compared with

control (Kim et al., 2002). However, fosthiazate was better than cadusafos and

fosthiazate pre-plant plus post-plant application and reduced nematode population

densities as much as 90 % and increased yield (Kim et al., 2002). Carbofuran gave

reduction in the incidence of root-knot nematodes infecting different vegetable crops

(Radwan, 1995; Stephan, 1995; Badawi and Abu-Gharbieh, 2000; Bari et al., 2004b;

Bhat et al., 2005; Singh et al., 2006). Giannakou et al. (2005) reported that oxamyl

provided some nematode control while cadusafos failed to provide adequate nematode

control, which may be attributed to the inability of the nematicide to reduce nematode

populations even at relatively high concentrations in soil.

Developing any new marketable nematicide is a long and expensive process

and reports state that no new widespread used nematicide has been developed in the

past 20 years (Starr et al., 2002). The worldwide phase-out of methyl bromide (one of

the effective and widely used fumigant nematicide (Oka et al., 2000b) and the

extreme cost of bringing new nematicides into the market triggered the need for the

alternative nematode control strategies that are economically feasible and


60

environmentally acceptable even if these strategies cannot compare to the 100%

efficacy of methyl bromide.

2.4.2. Seed treatment

Seed treatments gave adequate initial protection from nematode larvae to

seedling to better plant growth and increased yield. Cotton seed treated with

nematicides viz., carbofuran, phorate and fensulfothion can effectively be employed

to reduce population of R. reniformis from 32.45 to 49.06% (Muralidharan and

Sivakumar, 1975). Abamectin is a mixture of macro cyclic lactone metabolites

produced by the fungus Streptomyces avermitilis, which used as a seed treatment to

control plant parasitic nematodes on cotton and some vegetable crops. Abamectin was

effective on both M. incognita and R. reniformis in tomato plants (Faske and Starr,

2006). Abamectin has also a nematicidal effect against M. incognita and R. reniformis

on cotton plants as a seed treatment (Faske and Starr, 2007).

Furthermore, abamectin proved highly activity against lesion nematodes

(Pratylenchus spp.) as a seed treatment on corn with reduction evaluated by 25-72%

(Cochran et al., 2007). Seed treatment with carbofuran @ 0.1% in cotton and

1kg/100kg in mungbean were effective in reducing soil and number of eggs/plant of

R. reniformis (Brancalion and Lordello, 1982; Patel and Thakar, 1986). Korayem et

al. (2008) found that abamectin at the tested concentrations significantly reduced most

nematode parameters and enhanced plant growth parameters.

The protective and curative application of the biopesticides especially

abamectin and then azadirachtin were found effective in reducing the invasion of the

J2 and their further development in the roots. Azadirachtin was able to induce defence

mechanism in roots of plants, which consequently delayed the nematode development

(Rehman et al., 2009).


61

Reduction in gall numbers and final nematode population with nematicides for

12 h seed soaking has been recorded (Das and Deka, 2002; Deka et al., 2003). Earlier

several studies have proved the efficacy of carbosulfan and neem seed kernel powder

as a seed treatment in suppressing the phytonematodes in many field crops viz.,

chickpea (Chakrabarti and Mishra, 2001). Many workers (Singh et al., 1980; Siddiqui

and Alam, 1988a) demonstrated the application of Azadirachtin as the coating agent

which significantly reduced the root knot development.

Applications of nematicides as seed treatment or seed-cum-soil application

were found to give better control (Khan, 2004). Among the nematicides, fenamiphos

2kg a.i./ha was the most effective as it increased fibre yield of jute by 68%, being

particularly effective when the initial population of the nematode in soil was

relatively low. This was followed by carbofuran at 2 kg a.i./ha, which gave up to 40%

and 47% increase of fibre yield when applied to soil alone and in combination with

carbosulfan as seed treatment respectively (Khan, 2004). Mahanta et al. (1992)

found that soaking jute seed in a solution of carbosulfan 0.2% reduced root galling

and egg mass production of M. incognita. However, combining carbosulfan as seed

treatment with soil application of phorate at 2 kg a.i./ha and sebuphos at 2 kg a.i./ha

provided higher yields when they were applied alone (Khan, 2004). The application

of Nimbin as seed dressing significantly reduced the root-knot development,

populations of R. reniformis and T. brassicae (Siddiqui and Alam, 1990).

2.4.3. Seedling bare root dip treatment

Nematode damage is more harmful to seedling than to older plants. Hence

treatments with nematicides applied only at seedling roots may, therefore, lessen

nematode damage and may be more economical. Bare root dip treatment of brinjal

seedling with aldicarb, carbofuran, and terbofos at 500 and 1000 ppm was effective in


62

reducing nematode population (Prasad and Krishnappa, 1981). Bare root dip treatment

with pyridoxine hydrochloride (vitamin B6) solution of 0.1, 0.3 or 0.5% concentration

for 30 minutes resulted in improved plant growth (Ahmad and Alam, 1997).

Chemicals tested in the bare root treatment were induced some resistance in tomato

and eggplant against M. incognita and R. reniformis (Siddiqui, 1998). The poor root

knot development could be attributed to poor penetration (Siddiqui and Alam, 1988a)

and later retardation in different activities of the second stage juveniles such as

feeding and /or reproduction as suggested by Bunt (1975). This phenomenon might

have also happened in the case of R. reniformis where the plants did not support the

multiplication of the nematode as freely as compared to those which were not

subjected to root-dip treatment (Siddiqui, 1998).

It is possible that the chemicals are either absorbed by the roots or there might

have been some chain reaction which has been triggered due to some factor

(elicitor/activator). The initiation of the cascade mechanism leading to the resistance

of cells against the invasion and development of pathogens has been earlier described

(Bunt, 1975; Bell, 1981; Giebel, 1982). Bell (1981) considered the role of nematode

and plant enzymes in resistant plant reactions and suggested that enzyme effected

changes in growth regulators, free bound phenols, amino acid composition, and

induced lignifications to limit nematode development. Efficacy of carbofuran on

Meloidogyne as root dip treatment was reported by Gowda et al., 1988.

Application of a nematicide is required in each growing season of a

susceptible crop. Nematicide persistence in soil for 6-8 weeks is desired for effective

initial plant protection (Karpouzas et al., 1999). A few nematicides exist and currently

are not used for control of nematodes in India. Several insecticides used for insect

control, such as carbofuran, carbosulfan, cadusafos, phorate, and triazophos, possess


63

nematicidal activity, of which cadusafos has been reported to be the most effective

(McClure and Schmitt, 1996; Karpouzas et al., 1999; Le Roux et al., 2000; Meher et

al., 2005).

2.5. BIOLOGICALCONTROL

A variety of organisms have been shown as a potential biological control agent

of phytonematodes. This includes fungi, bacteria and virus etc. The beneficial effects

of certain types of plant derived materials and microorganisms in the soil have been

attributed to a decrease in the population densities of plant parasitic nematodes

(Akhtar, 2000b). Plant associated microorganisms have important roles in natural and

induced suppressiveness of soil-borne diseases. Several culturable rhizobacteria have

been tested for their biocontrol potential against plant parasitic nematodes (Siddiqui

and Shaukat, 2002, 2003; Khan et al., 2008b; Son et al., 2008, 2009). Addition of

treatments with plant extracts, bacterial suspensions or Vydate into soil suppressed

root galling and final populations of M. incognita, and except for Vydate promoted

plant growth and yield (Abo-Elyousr et al., 2010). Biological control organisms can

be artificially introduced into the soil, but naturally occurring soil microorganisms can

be stimulated by the addition of organic materials. There are now enough reported

examples of natural biological control of nematodes to show that this is a widely

occurring and common phenomenon (Bridge, 1996; Ahmad and Khan, 2004).

Biocontrol agents improve the health of plants and thus contribute to overall

productivity. These agents are also self propagating under favourable conditions and

therefore, may remain in the soil for a long period. Therefore, biocontrol is suggested

to be a safer solution. Various fungal antagonists of nematodes have shown promising

results. These mainly include endoparasitic fungi, parasites of nematode egg and

nematode trapping fungi. The fungus, P. lilacinus, is an egg parasitic fungus which


64

infects by direct hyphal penetration. The hyphae branch grows across the egg shell

(Khan et al., 2006).

The egg pathogenic fungus P. lilacinus is one of the most widely tested soil

Hyphomycetes for the biological control of plant parasitic nematodes

(Atkins et al., 2004). Lara Martez et al. (1996) demonstrated that P. lilacinus

significantly reduced M. incognita soil and root populations and increased yield of

tomato. (Siddiqui et al., 2000) has reported the reduction of M. javanica infection on

tomato by P. lilacinus. Cannayane and Sivakumar (2001) reviewed the biocontrol

efficacy of P. lilacinus and listed several reports where root-knot nematodes and the

potato cyst nematode Globodera rostochiensis were successfully controlled by this

egg-pathogenic fungus.

The culture filtrate of the Trichoderma species was highly significant in

controlling both nematode genera on eggplant. T. harzianum, T. hamatum and T.

koningii culture filtrates gave a significant reduction in vitro and decreased the female

and eggmasses of reniform and root-knot nematodes (Muthulakshmi et al., 2010;

Usman and Siddiqui, 2012a). Trichoderma culture filtrate was more significant on

root-knot nematode, M. incognita (Priya and Kumar, 2006). Trichoderma controls

nematode genera by a direct effect on toxic metabolites and inhibits nematode

penetration and development (Bokhari, 2009). Root colonization by Trichoderma spp.

frequently enhances root growth and development, crop productivity, resistance to

abiotic stresses and uptake and use of nutrients (Harman et al., 2004).

The Trichoderma species had been previously reported in the literature as

strains of T. harzianum and were reidentified (Rocha-Ramirez et al., 2002). These

species were used before nematode biocontrol studies (Sharon et al., 2001). Haseeb et

al. (2005) reported that the M. incognita-Fusarium oxysporum disease complex can


65

cause severe yield losses in V. radiata as in other crops. Although chemicals viz.,

carbofuran and bavistin showed a significant effect in increase of growth parameters

and in suppression of the disease complex, these can be replaced to some extent by A.

indica seed powder and microbial antagonists viz., T. harzianum and P. fluorescens to

avoid the hazards of chemicals.

Among numerous organisms that have shown antagonism against root knot

nematodes, P. chlamydosporia (Kerry, 2000; Siddiqui, et al., 2009), P. lilacinus

(Jatala, 1985; Kiewnick and Sikora, 2006), and T. harzianum (Siddiqui and Shoukath,

2004; Bokhari, 2009) have been found to be highly suppressive to plant nematodes,

especially under greenhouse conditions (Khan, 2007). Combining P. lilacinus with

neem leaves can provide satisfactory control of root-knot disease in eggplant (Khan et

al., 2012).

Successful biocontrol combinations have been recorded against root-knot

nematodes. The combination of the bacterium Bacillus subtilis and the fungus

Paecilomyces lilacinus suppressed nematode populations beyond the individual

application of the agents (Gautam et al., 1995). The fungal bioagents viz., P. lilacinus

and Trichoderma viride alone or in combination with mustard cake and furadan

promoted plant growth, reduced number of galls/plant, egg masses/root system,

eggs/egg mass and nematodes reproduction factor as compared to untreated infested

soil (Goswami et al., 2006).

The maximum reduction in root galling and the soil population, occurred in

soil treated with both fungi in combination with mustard cake. T. viride used alone

responded least as compared to P. lilacinus which was also observed by Khan and

Goswami (2000). The fungi used in combination also increased the plant growth

(Goswami and Singh, 2004). Both the fungi along with mustard cake and furadan


66

showed least a reproduction factor (0.0) as compared to untreated infested soil

(1.783). Goswami (1993) obtained a significant reduction in root gall index where soil

was treated with P. lilacinus with castor leaves and fertilizer.

Reddy et al. (1996) proved that T. harzianum incorporated into oil cakes was

effective for increasing yield and reducing the nematode numbers in soil and roots.

Other researchers (Siddiqui et al., 1999; Haggag and Amin, 2001;

Haseeb et al., 2005) studied the effect of Trichoderma on the development and

growth of parasitic nematodes. Stephan et al. (2002) reported that T. harzianum and

animal organic matters reduced the numbers of root-knot nematodes. Faruk et al.

(2002) indicated the effectiveness of T. harzianum on the biocontrol of root knot

nematodes on tomato. Saifullah (1996a, b) showed the death of 100% of Globodera

rostochiensis and G. pallida by using poisoning compounds from T. harzianum on the

medium after 24 h of exposure. It is well known that T. harzianum produces several

poisoning and antibiotic compounds (DiPietro, 1995) that can protect plants from

pathogenic organisms in soil (Wu and Wu, 1998). Several studies (Spiegel and Chet,

1998; Susan et al., 2000; Haggag and Amin, 2001; Sharon et al., 2001; Howell, 2003;

Siddiqui and Shaukat, 2004; Santhosh et al., 2005) showed the use of Trichoderma

for inhibiting the growth of plant parasitic nematodes. The secondary metabolites of

Trichoderma include chitinase enzyme which is considered to be the most effective

component against pathogenic fungi. Chitinase enzymes degrade the fungal cell walls

which are composed of chitin (Lorito et al., 1993). Chitin comprises the outer shell of

nematode’s eggs so that nematode eggs are affected greatly by Trichoderma species

treatment (Haggag and Amin, 2001; Jin et al., 2005).


67

2.5.1. The combined effect of organic amendments and biological control

Large numbers of workers have demonstrated that use of organic soil

amendment with different biocontrol agents viz., T. harzianum, P. lilacinus etc.

Combined application of biocontrol agents with various oil cakes have been reported

to be an effective approach to minimize the loss caused by various phytonematodes

(Tiyagi et al., 2002; Borah and Phukan, 2004; Zareena and Kumar, 2005). Azam et al.

(2009) reported that the combination of leaf powder C. tora and P. lilacinus was

ascertained to be most effective in managing the root knot disease.

Ashraf and Khan (2010) evaluated the efficacy of biocontrol agents and

various organic amendments in the management of phytonematodes infecting

eggplant under glasshouse conditions. All the treatments effectively suppressed the

nematode population. P. lilacinus and green manuring of Zea maize and Sesbania

aculeata were very effective for the management of Rotylenchulus reniformis

(Mahmood and Siddiqui, 1993).

Fruit wastes of apple, banana, papaya, pomegranate and sweet orange @

20g/plant and the fungal biocontrol agent P. lilacinus @ 2g (mycelium+spores) /plant,

alone and in combination were very effective for the management of R. reniformis

(Ashraf and Khan, 2008). The combined application of P. lilacinus, carbosulfan,

poultry manure and FYM alone and in combination significantly increased the plant

growth parameters including yield and reduced the population of M. incognita (Das

and Sinha, 2005). The integrated application of certain botanicals and P. lilacinus is a

better approach in reducing the nematode population and increasing the plant growth

and yield when used alone (Zaki and Bhatti, 1990; Zaki, 1998; Walia and Gupta,

1997).


68

Application of P. lilacinus showed better results in improving plant growth

and reducing the nematode population build up as compared to oil cake treated plants

whereas neem cake gave better results than other oil cakes. However, integration of

neem cake with P. lilacinus gave the best result causing increased plant growth and

reduced population build up of reniform nematode (Anvar, 2003; Ashraf and Khan,

2005). Research on nematode trapping fungi has demonstrated that the enhancement

of trapping activity resulting from the application of organic matter in soil is

dependent on the fungal species and the type and amount of organic material added

(Jaffee et al., 1994; Jaffee, 2004).

It is clear from the literature that the benefits of a combined green manure and

a microbial agent depend on the soil, the type of green manure and the species of

agent. The rhizosphere of some plants antagonist to plant parasitic nematodes have

distinct microfloras that have physiological traits, which indicate that at least part of

the antagonism may be due to the bacterial and fungal community on roots (Kloepper

et al., 1991; Insunza et al., 2002).

Kumar et al. (2011) and Haseeb and Kumar (2006) also reported that the

treatment with T. harzianum, P. fluorescens, A. niger, P. lilacinus, neem seed powder

and farmyard manure alone significantly decrease the severity of the infestation of M.

incognita and F. solani on brinjal. Perveen et al. (2007) reported that application of T.

viride and P. fluorescens with farmyard manure effectively control M. incognita and

plant growth. Similar results were also obtained by number of workers on other crops

(Pant and Pandey, 2002; Kumar et al., 2009; Abuzar and Haseeb, 2010). Combined

application of biocontrol agents, organic amendment and nematicides treatment was

found to be best among all treatments which significantly increased the growth


69

parameters including yield and reduced the population of M. incognita (Das and

Sinha, 2005).

Goswami et al. (2006) observed that the maximum reduction in root-galling

caused by M. incognita on tomato plants, as well as the soil population occurred in

soil, treated with both fungi (T. viride and P. lilacinus) in combination with mustard

cake. However, mustard cake alone also showed adverse effects on the root-

nodulation. Bio management of root-knot nematode, M. incognita affecting chickpea

using non edible seed oil cakes is an effective and ecologically safer approach as a

substitute of nematicides for the pollution free and sustainable environment (Rehman

et al., 2012).

Integration of Paecilomyces lilacinus and carbofuran at 2 kg a.i./ha was found

to be effective in the management of reniform nematode, Rotylenchulus reniformis on

tomato (Reddy and Khan, 1988). Sundraraju and Kiruthika (2009) demonstrated that

the integration of P. lilacinus with neem cake or anyone of the botanicals viz.,

Tagetes spp., S. torvum, can be effectively used in the management of root knot

nematode, since the use of single bioagent or botanicals cannot be very effective in

the management of nematode induced disease complex. The combined application of

various oil cakes and biocontrol agents have been reported to be an effective approach

to minimize the losses caused by various plant-parasitic nematodes (Mahmood and

Siddiqui, 1993; Rao et al., 1995; Tiyagi et al., 2002; Borah and Phukan, 2004;

Zareena and Kumar, 2005).

Ahmad and Khan (2004) reported that the amendments of neem sawdust and

kail sawdust were more or less equally effective to M. incognita infecting chilli and

did not differ significantly. The combined application of neem sawdust with the

biocontrol fungus P. lilacinus was more promising in increasing the plant growth and


70

decreasing the reproduction factor and root-galling, however, the kail saw dust did not

significantly increase the efficacy of P. lilacinus as the parasitism of the fungus was

inhibited and resulted in lower infection only on the egg masses of root-knot

nematode, M. incognita. The inhibitory effect of kail sawdust on the parasitism of P.

lilacinus could in fact be due to the toxic principles contained in kail sawdust and

released in soil, which might have inhibited the activity of P. lilacinus.

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REVIEW OF LITERATURE - Shodhgangashodhganga.inflibnet.ac.in/bitstream/10603/28726/12/12_chapter 2.pdf · Review of Literature 36 2.1.2. Biofumigation The term biofumigation is used