Status of Rice Nematode Research in India.pdf - Rice Knowledge ...

Status of Rice Nematode Research in India 

Prasad, J. S 1 , Somasekhar, N 2 and Varaprasad, K.S 3 

1 Principal Scientist, Entomology, Directorate of Rice Research, Rajendranagar, Hyderabad-30 

2 Senior Scientist, Entomology, Directorate of Rice Research, Rajendranagar, Hyderabad-30 

3 Project Director, Directorate of Oilseeds Research, Rajendranagar, Hyderabad-30 

For more Information contact: Visit Rice Knowledge Management Portal http://www.rkmp.co.in 

Rice Knowledge Management Portal (RKMP) 

Directorate of Rice Research, 

Rajendranagar, Hyderabad 500030. Email: naiprkmp@gmail.com, pdrice@drricar.org, shaiknmeera@gmail.com 

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Introduction 

Rice (Oryza sativa L.) is major staple food crop of India. It is cultivated in about 42.5 million 

ha with a production of about 95 million tons meeting food requirements of over 50% population 

of the country. At the current rate of population growth and per capita consumption, rice 

requirement by 2011- 12 is estimated to be around 100 million tons. This essentially means that 

the country has to produce an additional 10 million tons of rice by the end of XI Plan period to 

meet the food requirements. Keeping this in view, the Government of India has planned to launch 

the National Food Security Mission to achieve the production target of additional 10 million tons 

of rice (Viraktamath, 2007). 

Rice is the only crop grown in all the agroclimatic zones from 49 ° North in Czechoslovakia 

to 35 ° south in New South Wales of Australia. In India, rice is grown in diverse environments as 

sole crop (rain fed, irrigated or deepwater) or as a major component in various cropping systems 

besides in problem soils under saline and alkaline conditions. Rain fed and deepwater rice are 

grown under most unfavourable environments, therefore, the yields are very low. Since, both 

these important systems cover large area under rice in India, even a smallest pest problem would 

have great impact on yield and farmers’ income. About 300 nematode species belonging to 35 

genera have been reported infesting rice. Among them, nematode species from about ten genera 

are economically important in rice production. Rice grown in different environments is attacked 

by different nematode species. Ufra (Ditylenchus angustus) and root-knot (Meloidogyne spp.) 

nematodes are major pests of deep water rice. In irrigated rice, infections by Hirschmanniella spp. 

and Aphelenchoides besseyi are common where as upland rice is invariably infested by 

Meloidogyne and Pratylenchus species. As a consequence of diversion of increasing proportion of 

the available water for human usage, diminishing and erratic rainfall resulting in depletion of 

ground water resources, the availability of water for rice is becoming reduced year after year. 

Hence, water saving irrigation technologies such as aerobic rice and System of Rice Intensification 

(SRI) are receiving renewed attention from researchers and farmers (Prasad & Somasekhar, 2009). 

Studies conducted in Brazil and China, however, revealed that the high yields of rice obtained in 

such systems in the initial years are difficult to sustain and yields may decline after 3-4 years of 





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continuous cropping. Although the causes for slow decline in yield are not yet fully understood, 

the build-up of soil-borne diseases, including nematodes, toxic substances etc. are likely to be the 

factors (Bouman, 2002). 

Besides causing direct crop loss, nematodes also inflict indirect monetary losses resulting 

from export/trade restrictions imposed due to the presence of quarantine nematode pests. 

Globalization aimed at enhancing the world trade is providing equal opportunities to compete for 

export, facilitating exchange of huge volumes of rice to a tune of 25.3 million tons annually across 

the world. As a consequence, several countries sensitized by the Sanitary and Phytosanitary (SPS) 

agreement of World Trade Organization revised their regulations and included several pests in the 

regulatory lists. Among the important nematode species that attack rice, Ufra and white-tip 

nematodes find a place in regulatory pest lists of several countries (Varaprasad et al., 2006). 

Nematode problems have received relatively less attention in the past due to incipient 

damage in vast areas and difficulties in investigations. Most of the times the losses caused by the 

parasitic nematodes in rice are just accepted mainly due to unawareness, poor economic 

condition of the rice growers and subsistence farming of the crop. However, importance of 

nematode pests has increased in the recent years due to the changes in cropping systems and 

introduction of new production technologies that favour nematode multiplication and spread to 

new ecosystems in several rice growing countries. Various aspects of important nematode pests 

infesting rice with special reference to India are discussed in this article. 

Stem or ufra nematode (Ditylenchus angustus (Butler, 1913) Filipjev, 1936) 

History 

Ufra nematode, D. angustus was first reported from Naokhali, Tippera and Dacca districts 

of East Bengal (now Bangladesh) over 90 years ago (Butler, 1913a). The disease was reffered as 

Dak pora in local language, as the damage resembles lightening struck field. During the personal 

discussions with the scientists from Bangladesh it was told that the disease was first observed in 

the rice fields of a farmer, Uftur Rahman and the disease was named after him (first two alphabets 





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of two parts of his name: Uf and Ra = Ufra). D. angustus is a major pest of deepwater rice besides 

yellow stem borer and rodents. 

With the recent free movement of rice germplasm, the risk of spread of this nematode to 

new areas through seed has greatly increased (Prasad and Varaprasad, 2001). Further, changes in 

the government agricultural policy and the introduction of new methods of rice production in 

developing Southeast Asian countries may influence the nematode problems in rice. For example, 

in Vietnam, a Government decree in 1986 allowing the farmers to lease the land provided an 

incentive to the farmers to commercialize their production. The irrigation facilities were improved 

and modern cultivars were adopted. This reduced the rice area planted to floating rice in Mekong 

delta of Vietnam which subsequently reduced the infestation of D. angustus (Prot, 1994a). 

Distribution 

D. angustus is widely distributed in deepwater rice tracts of India, Malaysia, Philippines, 

Egypt, Thailand, Myanmar, Vietnam, UAE and Madagascar. In India, this nematode was initially 

reported from deep water rice in Assam, Uttar Pradesh and West Bengal (Roy, 1977; Singh, 1953). 

The symptoms of the ufra damage were also observed in Orissa but the organism was not isolated 

(Rao et al., 1986a). Later, the nematode was recorded from irrigated rice in Maharashtra (Patil, 

1998) and Andhra Pradesh (Prasad et al., 2005), however, there are no further reports on its 

establishment. Presently the nematode distribution is limited to North Lakhimpur District of 

Assam. 

Host range 

Cultivated rice is the important host of D. angustus. Besides this wild rice species viz., 

Oryza perennis (Moench), O. glaberrima (Steud), O. cubensis (Ekman) , O. officinalis (Wall ex Wall) 

, O. meyriama (Zoll et Mor ex Steud), O. latifolia (Desv), O. eichingeri (A. Peter), O. alta (Swallen), 

O. minuta (J.S. Presl ex C.B. Presl), O. nivara (Sharma et Shastry), O. rufipogon (Griff) and O. 

spontaneae (Roschev) (Hashioka, 1963; Miah and Bakr, 1977; Sein and Zan, 1977) and weeds like 

duck weed (Hygroryza aristata Retz.), swamp rice grass (Leersia hexandra Sw.) (Miah and Bakr, 





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1977; Voung 1969; Sein and Zan, 1977), Sacciolepsis interrupta, Echinochloa colona L. (Cuc, 1982), 

Paspalum scrobiculatum L. (Pathak, 1992) also serve as hosts of this nematode. 

Symptoms of damage 

In the vegetative stage, injury due to D. angustus feeding results in a mosaic or chlorotic 

discolouration of emerging or emerged leaves; yellowish or pale green splash-patterns on affected 

leaves and leaf sheaths; appearance of brown to dark brown spots on leaves and leaf sheaths; and 

leaf margins become contorted. At the reproductive stage of crop, nematodes reach the space 

between the inner sides of imbricate whorl of leaf sheaths to feed on the ear primordia and 

developing ear heads. As a result, ear heads emerge in a twisted and crinkled manner with empty 

spikelets or do not emerge at all (Padwick, 1950; Miah and Bakr, 1977). The collective symptoms 

are known as ufra disease. In case of early infestation, panicles may fail to emerge. The symptoms 

of injury appear within one week in young plants and in 10-15 days in plants at or nearing 

flowering stage. Branching of infested ears and development of three to four ear heads enclosed 

in a single leaf sheath may occur (Rao et al., 1986a). Based on the emergence of the panicle the 

disease has been classified as ufra I (where the panicle fails to emerge), ufra II (where there is 

partial emergence of panicle) and ufra III (where there is complete emergence of the panicle) (Fig. 

1) (Cox and Rahman, 1980). 

Fig. 1. Panicles affected by Ufra nematode, Ditylenchus angustus. 





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Life/disease cycle 

D. angustus is an obligatory ectoparasite that survives and multiplies on living rice plants. 

The nematode can swim actively. In dry situations, it remains coiled in a dormant state on the dry 

soil or dried plant material and regains activity when comes in contact with flood water or rain. 

The developmental cycle of rice stem nematode from second stage juvenile (J2) to adult takes 15 

days, from J2 to egg 21 days and from J2 to J2 stage takes about 24 days. Atmospheric temperature 

of 28-30°C and more than 80% relative humidity are favourable for infection, disease development 

and reproduction (Miah and Bakr, 1977; Ou, 1985). The nematode reproduces inside the host 

plant between the months of May or June and November i.e. tillering to milky stage of the crop. 

During this period, three generations were recorded (Butler, 1913b). The fourth stage larva (J4) of 

the nematode is thought to be the resting stage that helps in dissemination of the nematode 

(Butler, 1913a). However, Ibrahim and Perry (1993) observed that though J4 is the predominant 

and constantly superior stage, the J3 and adults also show similar survival attributes. 

Host-parasite relationship 

D. angustus feeds ectoparasitically on the inner surface of unemerged leaves, sheaths, 

buds and developing panicles and causes ufra disease in cultivated and wild rice, and weeds at all 

stages of growth. The presence of viable, anhydrobiotic juveniles and adults of D. angustus on 

freshly harvested rice seeds may be of importance for the dissemination of this species. The 

presence of nematode in the germ portion of the seed was also observed (Prasad and Varaprasad, 

2001). 

Interaction with other organisms and disease complexes 

The spots on the sheath of D. angustus infected plants serve as the sites for secondary 

invasion by Fusarium, Cladosporium and Sclerotium. The nematode infected plants become 

susceptible to blast (Pyricularia oryzae), sheath rot (Sarocladium oryzae) and bacterial leaf blight 

(Xanthomonas oryzae) diseases (Voung, 1969). Rathaiah (1988) reported leaf and nodal blast in 

ufra infested plants from Assam. 





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Effect of environmental factors 

Deepwater rice has a distinct pest complex due to prolonged deep flooding, extended 

growth duration and a complex environment. Flooding smothers weeds, prevents population build 

up of some pests and diseases and stimulates new growth which may compensate for early 

damage (Catling and Islam, 1999). D. angustus is highly adapted to these aquatic conditions and 

exploits the succulent growth and mild weather during this period. Maximum incidence of D. 

angustus occurs in early sown crop and gradually decreases as the sowing is delayed. Ufra 

nematode infection usually occurs with flood waters and tidal water helps spread of ufra 

nematode. The broadcasted crop often suffers more ufra infestation in comparison to 

transplanted crop. Most of the ufra-prone areas grow broadcast ‘aman’ rice (April-November), 

followed by ‘boro’ rice (November-March), thus facilitate nematode survival throughout the year. 

Repeated cultivation of highly susceptible cultivars contributes to the nematode build up leading 

to development of ufra disease. Das and Bhagawati (1992) observed that maximum infection of 

the nematode occurred in early March transplanted crop that gradually declined in November 

transplanted crop. Zinc deficient rice plants had less fertile tillers, fewer filled grains and expressed 

more severe symptoms of D. angustus infection than plants supplied with added Zinc (Mondal and 

Miah, 1984). Further, Mondal and Miah (1985) observed that plants grown in soils having 220 ppm 

or greater level of K2O express resistance to infection by D. angustus. 

Yield losses 

The yield loss due to D. angustus may vary from year to year depending on the variety, 

time and degree of infection, and the environmental conditions prevailing during the crop season. 

In India, yield losses due to ufra were reported as 5-50% in U.P. (Singh, 1953); 10-15% in West 

Bengal (Rao et al., 1986a) and 30% - 100% in hot spots for this nematode in Assam (AICRPN, 1986). 

In southern region of Thailand 10-90% loss was observed (Hashioka, 1963). Khuong (1983) 

observed most severe and conspicuous damage by D. angustus in 50,000 ha flooded fields with 

50% yield loss in the Mekong Delta and Dong-Thap Province of Vietnam. 





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Management 

Management of D. angustus is difficult because of the nature of the deep water rice 

ecosystem. difficulties in application, contamination concerns and the low economic returns 

restrict nematicide use. Varietal resistance and cultural management are the most effective 

options available in this ecosystem. 

Host plant resistance 

In North Lakhimpur, cv. IR63142-J8-B-2-1 recorded the least (3.6%) ufra-infested panicles 

in comparison to 81.8% infested panicles in susceptible cv. Rangabao (Sarma et al., 1999). Rice 

cultivars viz. B-69-1 (Sein, 1977), Rayada 16-06, CN 540, NC 493, TCA 55 (Rahman and McGeachie, 

1982), Brazil-65, Rayada 16-05, Rayada 16-06, Rayada 16-07, Rayada 16-08, Rayada 16- 

011,Rayada 16-013, Ba Tuc (IRRI, 1986; Rahman, 1994), AR 9, IR 13437-20-P1, IR 17643-4 (Pathak, 

1992), and a wild rice species O. subulata (Sein, 1977) were found resistant to D. angustus. The 

rice cultivars Padmapani (McGeachie and Rahman, 1983) and Digha (Mondal and Miah, 1987), 

which mature early, completely escape infestation. In the breeding programme for resistance to 

ufra nematode in Bangladesh, it was found that all the resistant entries identified had either 

Bazail-65 or Rayada 16-06 as one of the parents (Rahman, 1994). Mondal and Miah (1987) 

reported that cultivar Khalni was moderately resistant and Rayanda (Rayada) and Keora were 

resistant to ufra nematode. However, in the resistant cultivars identified, elongation capacity was 

low in comparison to established deepwater varieties. Incorporation of resistance to ufra into 

established cultivars utilizing the modern biotechnological tools may be a better option. 

Cultural control 

Burning of infested stubbles on community basis in an organised manner may greatly help 

in destroying the infection loci. Similarly, preventing flood water from the river, the source of ufra 

nematode infection, into the fields by strengthening the bunds could be beneficial (Sein and Zan, 

1977). The best way to control ufra is by completely drying fields when they are fallowed, 

ploughing to destroy loci of infection in stubbles and rotation with non-host crops. The early rice 





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cultivars Padmapani and Digha completely escape infection. To take advantage of this, McGeachie 

and Rahman (1983) suggested lengthening the critical overwinter period of D. angustus by sowing 

deepwater rice later than normal or transplanting much later as a control measure against ufra. 

Growing a non-host crop such as jute in rotation with deepwater rice and rotation of transplanted 

rice with mustard, a non-host crop was found effective for the management of ufra nematode 

(McGeachie and Rahman, 1983). 

Chemical control 

Several nematicides were found effective in controlling D. angustus in field. Ethoprophos, 

carbofuran and isazofos treated plots yielded 0.9, 0.82 and 0.08 tons /ha respectively, more than 

untreated plots when applied at transplanting. Soil incorporation of mocap and carbofuran 

appeared to be effective against the stubble borne ufra nematode (Rahman and Miah, 1989; 

Mondal et al., 1990). Rahman and Taylor (1983) reported that the nematode can be controlled 

with carbofuran 1.5 kg a.i./ha. Rahman (1993) observed that application of carbofuran @ 0.75 kg 

a.i./ha at transplanting was most effective in reducing ufra infestation and consequently 

increasing rice yield in comparison to split or late application at 4-6 weeks after transplanting or 

control. Das (1997) recorded lowest infestation of the nematode in treatments with 2 sprays with 

carbosulfan 40 EC at 0.2% followed by 2 sprays of triazophos 40 EC at 0.2% as compared to the 

untreated control with significant increase in grain yields. 

Effective and adoptable recommendations 

Burning diseased stubbles, straw, followed by field spray with carbosulfan 40 EC at 0.2% 

and 2 sprays with triazophos 40 EC at 0.2% gives effective control of the disease and significant 

increase in grain yield. Burning diseased stubbles, straw, followed by several ploughings and 

growing early maturing varieties like Padmapani or Digha seem to be viable options for the 

management of the nematode. 





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White-tip nematode (Aphelenchoides besseyi Christie, 1942) 

History 

Kakuta (1913) and Tanaka and Uchida (1941) attributed disease symptoms in rice with plant 

parasitic nematodes in Japan. In USA, Jodan (1935) described a disease which he named as white- 

tip and attributed it to the deficiency of iron which was supported by Tullis and Cralley (1936) and 

Jones et al. (1938). But Martin (1939) and Martin and Alstatt (1940) thought it to be due to 

magnesium deficiency and magnesium and calcium imbalance, respectively. Christie (1942) 

identified and described the organism parasitizing strawberry as A. besseyi. Later, Yokoo (1948) 

described the nematode infecting rice under the name A. oryzae. Yoshii and Yamamoto (1950) 

compared the diseases of rice and millets and established that A. oryzae was responsible in all the 

cases. Allen (1952) compared the American and Japanese rice nematodes and established that 

both were identical to A. besseyi described by Christie (1942). So far the movement of processed 

or milled rice is taking place between countries. Presently, several multinational companies 

(MNCs) are taking up seed production in India for use in India and neighboring countries. In such 

cases, presence of this nematode in seed could create quarantine problems, if it is not prevailing 

in the receiving country. This is evident from the recent revisions made in the quarantine pests 

lists of some countries. For example, in Italy, A. besseyi is considered as an exotic plant pest 

posing a potential threat to the Italian agriculture and environment (Greco and Inserra, 2008). 

Distribution 

White-tip nematode, A. besseyi is widely distributed and now occurs in most of the rice 

growing areas of the world (Ou, 1985). The known distribution of A. besseyi on rice includes 

Australia, Ceylon, Comoro Islands, Cuba, El Salvador, Hungary, India, Indonesia, Italy, Japan, 

Madagascar, Mexico, Pakistan, Philippines, Taiwan, Thailand, USA, former USSR and in most 

countries of central and West Africa (Franklin and Siddiqi, 1972). 





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Host range 

Rice, strawberry and tuberose are the major hosts of the white-tip nematode. The host 

range encompasses more than 35 genera of higher plants (Fortuner and Williams, 1975). The wild 

rice species viz., Oryza breviligulata A. Chev et Roehr, O. glaberrima; common weeds like Cyperus 

iria L., Setaria viridis (L.) Beauv., Panicum sanguinale; food crops such as maize, bajra and Italian 

millets (Dave, 1982), Dioscoria trifida L., Ipomoea batatas (L.) Poir., Allium cepa L., Zea mays L. and 

Colocasia esculenta (L.) Schott, were reported as hosts of A. besseyi. In addition, many saprophytic 

fungi also serve as good hosts for this nematode. A. besseyi can survive but can not multiply on 

rice blast fungus, Pyricularia oryzae (Rao, 1985) but it can feed and reproduce on stem rot fungus, 

Sclerotium oryzae (Iyatomi and Nishizawa, 1954). Other host plants of A. besseyi include 

Polianthes tuberose, Hibiscus brachenridgii, Vanda orchid, hydrangea, chrysanthemum and several 

other flowering plants in Hawaii (Holtzmann, 1968; Raabe and Holtzmann, 1965; Sher, 1954); 

Boehmeria nivea in the Philippines (Fortuner, 1970); Ficus elastica and the wild grass, Sporobolus 

poirettii in USA (Marlatt, 1966; Marlatt and Perry, 1971); Setaria, Panicum and Cyperus iria (Yoshii 

and Yamamoto, 1950) and Peennisetum in Japan (Hashioka, 1964). Hockland and Eng (1997) 

recorded Capsicum annuum v. longum to be a host to white tip nematode. 


Nematode infested plants show white-tip or whip-like malformation of the top third of the 

leaf blade (Fig. 2). In flowering tillers, chaffiness and abnormal elongation of glumes in some 

spikelets, rachii and rachillae occurs (Todd and Atkins, 1958; Rao, 1970). Infected plants show 

reduced vigor, height and weight of spikelets and number of grains. Abnormal elongation of the 

panicles (Rao, 1978) and chaffiness or scattered chaffiness in the florets also occurs in case of 

severe infestations (Fig. 2-3) (Prasad et al., 2007). In some rice cultivars, A. besseyi may produce 

only the symptoms of small grains and erect panicles, but not the typical leaf white tip (Liu et al., 

2008). 





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Fig. 2.White-tip nematode damage in leaves (a) floret (b), and spikelets (c). 

Fig. 3. Chaffiness in grains caused by Aphelenchoides besseyi. 


A. besseyi is bisexual and an ectoparasitic nematode. The duration of life cycle from egg to 

egg is about 6 to 7 days. In nature, the length of life cycle depends on the ecological factors and it 

may take 3 days (at 31.8°C) to 29 days (at 14.7°C) for completion of life cycle (Tikhinova, 1966). 

The nematode becomes active at the germination of the seed and starts feeding ectoparasitically 

on the leaf primordia. The nematodes feed and multiply in the leaf whorls and climb to the 

booting panicle through a fine film of moisture on the surface. Nematodes invade the florets 

through the tunnel below the apiculus where lemma and palea remain open (Fig. 2). As the grain 

ripens the nematodes become quiescent in dried tissues of panicles and straw. The nematode 





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emains viable in this state in dry tissues and under hulls of rice grains for up to 3 years. This 

nematode is disseminated with the infected material. 


A. besseyi feeds endoparasitically in the coleoptile for 7-10 days in the initial stages of 

development of rice plants and ectoparasitically within the innermost leaf sheath during other 

plant growth stages (Tsay et al., 1998). At late tillering stage, nematode numbers may increase 

rapidly, and reach a peak during the reproductive stage of the plant. Damage to the outer wall of 

the ovary causes partial filling of kernels and damage to the lodicules prevents closure of flower 

after anthesis, exposing the embryo to environmental stresses like desiccation (Rao and Rao, 

1979). Survival of A. besseyi is inversely related to the extent and rate of dehydration and the 

nematodes in larger aggregates were found to survive better than those in the smaller ones. The 

starvation adversely affects the ability of the nematode to survive dehydration. Larvae and adults 

of the nematode were found equally capable of withstanding desiccation (Huang and Huang, 

1974). Hoshino and Togashi (2009) observed that there was a trade-off between both the 

dispersal and competition of rice seeds and between dispersal and reproduction of white-tip 

nematodes harbored in the seed. Lighter seeds from nematode infested fields showed a larger 

mean degree of swelling than did those from non-infested fields and light seeds harboring many 

nematodes had a well developed endosperm. 


Damage to the lodicules by the nematode exposes the embryo to infection by Alternaria (= 

Trichoconis) padwikii and spikelet sterility (Rao and Rao, 1979). Pathogenic fungi Acrocylindricum 

oryzae and Dorticium sasaki invade the interveinal areas of nematode affected leaf (Rao and Rao, 

1979). Increase in humidity and delay in emergence of the panicle due to feeding of the nematode 

on the inner layer of the boot provides an opportunity for infections by opportunistic fungi such as 

Fusarium spp. (Prasad et al., 2007). 





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The nematode remains active at a temperature range of 13-42°C with ideal relative 

humidity above 70% (Tikhninova, 1966). When nematodes in desiccated condition in rice seeds 

were exposed to 70°C for 12 h, about 16% nematodes were survived and the germination of rice 

seeds decreased to 44%. However, at 60°C the survival of A. besseyi was 40% and there was no 

effect on rice seed germination. The nematode can survive for over 1 year in rice seeds between 

glumes and grain, and 53 days in water at 10°C. The minimum temperature for nematode activity 

is 4°C and the thermal death point is 49°C for 10 min. Nematodes die totally when exposed to 41- 

44°C in water for 1 hour. The temperature and precipitation during 10-15 days after sowing can 

influence disease severity. A. besseyi invades rice mainly during sowing to the 3-leaf stage. Low 

temperature and more precipitation cause the disease to become more serious (Qiu et al., 1991). 

Yield losses 

White tip disease is a serious problem in many countries. The loss in grain yield due to 

chaffiness of ear heads was 20% (Muthukrishnan et al., 1974) and the ear head damage due to 

partially filled grains ranged from 21-46% (Nandakumar et al., 1975). An epidemic of white tip 

occurred during 1979 on AICRIP Farm in Hyderabad in which 60% of the varieties grown were 

severely infested (Jayaprakash and Joshi, 1979). Yield loss of 44.9, 34.7 and 24.2% were recorded 

when infestation rate was 57, 34 and 18%, respectively (Tsay et al., 1998). In Brazil, 50% crop loss 

occurred to upland rice crops (Silva and da Silva, 1992). Rice seed with 80% infestation, when 

sown produced 97% infested and 67% diseased plants in greenhouse, and 54% infested and 31% 

damaged plants in field conditions (Popova, 1984). Rice samples (146 out of 1653) collected from 

four rice-growing areas in AP were found infected with A. besseyi (Savitri et al., 1998). The 

nematode was not recorded in the samples from Karnal, Kurukshetra, Kaithal and Ambala districts 

of Haryana (Dabur,1998). 





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Management 


The most effective control method for A. besseyi is the use of nematode free or resistant 

planting materials (Silva and da Silva, 1992). Genotypes Bluebelle, BR-IRGA 409 and IRGA 172 F4 

SS39 did not show symptoms of white tip when inoculated with A. besseyi in Brazil (Oliveira and 

Oliveira, 1989). In USSR, utilizing the progenitors of almost all the resistant varieties bred there in 

the last 40 years i.e. Fortuna, Nira, Rexoro and Bluebonnet, a notable derivative Bonnet 73 with 

multiple resistance to A. besseyi and various other diseases was developed (Zelenskii and Popova, 

1991). 

Physical control 

Storage of A. besseyi infested seeds in regulated gas medium (97.5% nitrogen and 2.5% 

oxygen) for 10 days at 25 o C gives the best control (Aleksandrova and Beloglazov, 1989). 

Treatments with either ethoprophos 20EC at 0.5% or hot water at 53-54 o C for 15 min reduced the 

infestation in seed to almost nematode-free level (Tacconi et al., 1999). In addition to hot water 

treatment, a combination of seed treatment (at 0.3% by seed weight) and spraying (at 2.5 g/dm 3 

at 1 or 15 days after transplanting) with benomyl protects rice plants from infestation by A. 

besseyi (Gergon and Prot, 1993). Sivakumar (1987) observed that soaking rice seeds in 1% 

potassium chloride or 1% sodium chloride for 20 h and then sun drying (40-41°C) for 6 h to almost 

original moisture level or sun drying for 6 h without pre-soaking were helpful in disinfecting the 

seed from A. besseyi to an extent of 87 to 97%. 

Cultural methods 

In irrigated rice, damage due to the nematode is less when rice was sown directly into 

water rather than flooded after sowing (Silva and da Silva, 1992). Damage due to this nematode 

can be minimized by thoroughly washing the pre-soaked rice seed with excess amount of water 





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efore sowing in the nursery. This treatment reduces the nematode inoculum in seed by removing 

the activated nematodes along with the excess water. 


Seed treatment with thiobendazole, benomyl or fenitrothion reduced nematode 

infestation to a very low level, but they were not completely eliminated (Silva and da Silva, 1992). 

Pre-soaking of seed with oxamyl or hot water treatment reduced the infestation rate and 

increased yields (Tsay et al., 1998). In USSR, pre-sowing treatment of rice with the organic mercury 

compounds Granozan and hydrogen peroxide reduced A. besseyi infestation of rice panicles, 

improved the density of the rice stands by 10-11% and increased the yields by 13-36%. Hydrogen 

peroxide quickly decomposed unlike Granozan which accumulates in the soil and in the grain and 

is toxic to mammals (Anikeev and Shabelnikov, 1980). Kumar and Sivakumar (1998) observed that 

monocrotophos sprayed @ 1000 ml/ha at the boot leaf stage, reduced the white tip incidence and 

grain chaffiness, and increased the yield of rice in field experiments. 

Regulatory methods 

In view of seed borne nature of the nematode, it is urged to enact legislation to prevent 

the sale of uncertified and infested seeds of rice (Silva and da Silva, 1992). In a bid to develop a 

quarantine treatment schedule, Prasad and Varaprasad (1992) tried sixteen treatment 

combinations in 10 rice entries heavily infested with white-tip nematode for its elimination. 

Soaking of the seeds in 0.2% solution of mancozeb and monocrotophos followed by vacuum 

fumigation (methyl bromide @ 32 g/m 3 ) for 2 h at 30°C successfully eliminated the nematode in all 

the test entries. Even when the vacuum fumigation was substituted with atmospheric fumigation 

(aluminium phosphide @ 9.3 g/m 3 ), the treatment was found equally effective. 





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Selection of disease free seed and seedlings; Addition of two volumes of boiling water to one 

volume of seed soaked in two volumes of water and keep stirring for 10 minutes; Destruction of 

weed hosts may be the practical solution for the management of this nematode. 

Root-knot nematode (Meloidogyne graminicola Golden and Birchfield, 1968) 

History 

Root-knot nematode was found infecting grasses Echinochloa colonum, Poa annua, 

Alopecurus carolinianus, Eleusine indica and oats in USA during 1965 (Golden and Birchfield, 

1965). Later this nematode was described as Meloidogyne graminicola (Golden and Birchfield, 

1968). This was followed by several reports of its association with rice in many countries. In India, 

M. graminicola is the dominant species infecting rice. M. triticoryzae infecting both rice and wheat 

including some monocot weeds is also reported from India (Gaur et al., 1993) and its occurrence is 

restricted to a few areas. 

The root-knot nematode is making its importance felt in allmost all the rice growing areas. 

Recent observations on the susceptibility of main crop to root-knot nematode in the rice based 

cropping system such as wheat (Chandel et al., 2002), onion (Gregon et al., 2002) and banana 

(Reversat and Soriano, 2002) contribute to the accentuation of the problem. In Philippines, 

economic reasons and the decrease in water supply have induced the large scale adoption of 

direct wet seeding, chemical weed control and intermittent irrigation that favour the development 

of M. graminicola and have drastically increased its economic significance. Kreye et al. (2009a, b) 

observed that root-knot nematode was one of the important factors responsible for the poor 

plant growth and yield failure in aerobic rice in Philippines. Sudden out break of M. graminicola 

infestation in 1500 ha area in Mandya (Karnataka, India) during kharif, 2001 stands as an example 

for our limited understanding of this nematode (Prasad et al., 2001). 





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Distribution 

M. graminicola is distributed in the countries of S.E. Asia, Burma, Bangladesh, Laos, 

Thailand, Vietnam, India, China, Philippines, Nepal and USA. M. oryzae has been found in Surinam 

on irrigated rice, M. incognita in Costa Rica, Cuba, Egypt, Ivory Coast, Nigeria, South Africa and 

Japan, M. javanica in Brazil, Egypt, Comoro Islands, Nigeria and Ivory Coast, M. arenaria in Nigeria, 

Egypt and South Africa and M. salasi in Costa Rica and Panama on upland rice (Bridge et al., 

1990). In India, M. graminicola has been found infecting rice in Assam, Andhra Pradesh, Karnataka, 

West Bengal, Orissa, Kerala, Tripura and Madhya Pradesh (Prasad et al., 1987). Root-knot 

nematode, M. graminicola is a serious pest of upland rice and nurseries world over in well-drained 

soils (Rao et al., 1986b). The nematode was reported on irrigated rice in Andhra Pradesh (Sharma 

and Prasad, 1995) and Karnataka (Prasad et al., 2001). The nematode can infect and multiply on 

semi-deep (Prasad et al., 1985) or deepwater rice also (Bridge and Page, 1985). Occurrence of M. 

triticoryzae is reported from Delhi, Uttar Pradesh and Haryana (Gaur et al., 1993). 

Host range 

M. graminicola has a wide host range with rice being a major economically important 

host. It was initially found on barnyard grass, Echinochloa colonum (Golden and Birchfield, 1965). 

Subsequently it was found that it readily attacks several grasses, bush bean, oats (Golden and 

Birchfield, 1965), Ranunculus pusillus, Cyperus compressus L. (Yik and Birchfield, 1979), Panicum 

miliaceum L., Pennisetum typhoides (Burm. F) Stapf and C.E. Hubb and Glycine max (L.) Merr (Roy, 

1978), Echinochloa crusgalli, E. colona, Eleusine indica, Paspalum sanguinola, Eclipta alba, 

Grangea madraspatensis, Phyllanthus urinaria, Fimbristylis miliacea, Blumea sp., Vandellia sp., 

Jussieua repens, Andropogon sp., chillies, tomato, wheat, Panicum spp. (Rao et al., 1970), Cyperus 

deformis (Bajaj and Dabur, 2000), Banana (Reversat and Soriano, 2002) and onion (Gregon et al., 

2002). 





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Root-knot nematode affected plants show depletion in vigor, stunted growth, chlorotic and 

curled leaves in nurseries (Fig. 4) and main field. The nematode infection is characterized by the 

formation of small galls near the tips of the roots (Fig. 5 and 6 ). Excessive branching of affected 

roots occurs. The crop damage depends on the density of egg masses/second stage juveniles in 

the soil. 


M. graminicola completes its life cycle in 26-51 days in different periods of the year (Rao 

and Israel, 1973). The second stage juveniles upon entry into the roots, establish at a point in the 

stele and start deriving their nutrition from the giant cells induced by the nematode secretions. 

The second stage juvenile undergoes successive moults to become an adult. The males are 

vermiform with weak stylet. The females are saccate and pear shaped with a bent neck which 

remains inserted in the stelar tissues. Each root-knot may contain one or more females. The eggs 

are laid in a gelatinous matrix and each egg mass contains 150-300 eggs. The females of M. 

graminicola remain embedded in the root cortex and eggs are laid inside the roots, unlike other 

root-knot nematodes where the females protrude out of the cortical tissues of the root (Jena and 

Rao, 1977). The root galls that are formed are white and look initially pearl like and turn dark 

brown as the nematode matures. The eggs get dispersed in the soil due to root decay and the 

juveniles hatch from eggs to invade the fresh plant roots if available or wait for the following 

season. 





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Fig. 4 . Root-knot nematode, Meloidogyne graminicola infested rice nursery. 

A B 

Fig. 5 . Meloidogyne graminicola infected (A) and healthy (B) rice seedlings. 





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Fig. 6 . Root galls on rice roots due to root-knot nematode, Meloidogyne graminicola. 


The root-knot nematode, M. graminicola is an obligate parasite and a major pest of rice. 

Infective second stage juveniles of M. graminicola select a point for entry into the root, usually in 

the meristematic zone. The juveniles cause disruption, hypertrophy and hyperplasia of cortical 

cells by intracellular migration and releasing oesophageal gland secretions. The nematode incites 

development of 5-8 giant or transfer cells in phloem. Around the giant cells abnormal xylem 

proliferation occurs that causes swelling in stelar tissue. Hypertrophy of cortical cells around the 

sites of establishment of the nematode is responsible for the formation of galls or root-knots. 

Inorganic nitrogen application to infected crop gives a temporary greenish appearance and the 

plants turn yellow within a week after application, due to the inability of roots to intake and 

transport the nutrients. Infection by the nematode results in reduction of N, P, K and increase in 

total sugars, amino nitrogen and DNA in plants. Increase in RNA in shoots and roots to an extent of 

20 and 80%, respectively were recorded due to excessive hyperplasia and hypertrophy and 

inhibition of protein metabolism (Rao et al., 1986c). The larval migration in cortex and 

establishment of giant cells in stele takes about two days in susceptible rice cultivars TN-1 and TN- 





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4 while it takes about 12 days in resistant cultivar TKM 9 (Senthilkumar et al., 2007). Rao and Israel 

(1972c) observed a higher rate of reproduction of M. graminicola at low levels of inoculum 

possibly due to the abundance of food, lack of competition and the ability of the host to support 

the population. The low rate of reproduction obtained at high levels of inoculum is considered to 

be due to crowding. The growth and development of the rice root-knot nematode population is 

thus dependent on its population density. 

In deepwater rice, root-knot nematode infected seedlings remain stunted, unable to grow 

above flood water and perish due to continuous submergence. In Surinam, glasshouse trials 

revealed that the rice yield was 15% lower when M. oryzae inoculated in flooded soil and 9% less 

when nematode was inoculated in soil without standing water in comparison to nematode-free 

pots (Segeren and Bekker, 1985). 

Mishra and Mohanty (2007) observed an increase in phenolics by 28-104%, phynylalanine 

ammonia lyase by 16-35%, tyrosine ammonia lyase by 9-54%, decrease in amino acid tyrosine by 

2-36% and amino acid tryptophan by 14-28% in rice cultivars Annapurna, Manika and Ramakrishna 

and suggested that these reactions could be used in rating of the resistance of cultivars to M. 

graminicola. Shrestha et al. (2007) detected a total of six significant or putative QTLs for root-knot 

nematode tolerance and observed that the partial resistance to nematode establishment was 

related to nematode tolerance. They opined that it may be possible to breed plants with greater 

tolerance. 

Singh et al. (2006) demonstrated the relationship between the biomass of M. graminicola 

developing in rice roots and the expression of disease symptoms. The biomass of invading second 

stage juveniles (0.09 μg) increased to 33 μg on day 16 when adult females were in advanced egg 

laying stage, with an increase of approximately 360-fold. Initiation of leaf yellowing was related to 

the ratio between total nematode and total root biomass of rice seedlings. Plants with nematode- 

to-root biomass ratios above 1: 161 did not show any symptom while those with ratios between 

1: 138 and 1: 121 exhibited yellowing. Plants with nematode-to-root biomass ratios between 

1: 115 and 1: 60 showed moderate stunting while those with ratios between 1: 43 and 1: 20 





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exhibited severe stunting. The wilting symptoms occurred at or below 1: 14 nematode-to-root 

biomass ratio. The yellowing of leaves in seedlings inoculated with graded inocula was expressed 

when the nematode-to-root biomass ratios reached to 1: 136 on day 15 at 1000 J2, 1: 138 on day 9 

at 3000 J2, 1: 134 on day 7 at 6000 J2 and 1: 129 on day 5 at 9000 J2 per pot. In rice nurseries, 

seedlings showing moderate stunting, severe stunting, wilting and wilting with single gall were 

recorded at nematode-to-root biomass ratios of 1: 92, 1: 20, 1: 12 and 1: 14, respectively. In 

severely stunted transplanted rice, the nematode-to-root biomass ratio ranged from 1: 84 to 1: 75 

(Singh et al., 2006). 


M. graminicola infestation causes reduction in phenols in the shoots and roots and it is 

thought to be the reason for greater susceptibility of nematode infected plants to rice blast 

pathogen, Pyricularia oryzae (Israel et al., 1963) and root fungus, Fusarium moniliformae 

(Hazarika, 2001). 


Rao and Israel (1972a) reported maximum hatching of eggs of M. graminicola in water at 

25 and 30°C. At 15 and 35°C hatching was reduced and at 20°C it was slightly less than that at 

25°C. Larval populations of M. graminicola in soil were large during December to February when 

soil temperatures were 20.9°C or less. Populations were small in March, July and August and very 

small in April, May and June when the soil temperature was 31°C. Maximum galls on rice roots 

were found during January to March and egg masses during February to March. Soil temperatures 

of 23.5°C or less were found most favorable for gall formation (Rao and Israel, 1971b). Larval 

invasion was greatest in soils at 32% moisture content; development and egg mass production 

were greatest at 20 to 30% soil moisture. Greatest larval invasion may occur at pH 3.5 but pH 

usually does not affect invasion, growth or development of the nematode to any significant 

extent. Drought conditions at the tillering stage and at both tillering and flowering stages favoured 

the development and reproduction of the nematode. Addition of nitrogen up to 40 kg/ha to the 





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soil resulted in increased reproduction. Application of additional phosphorus either alone or in 

combination with nitrogen also favoured nematode development (Rao and Israel, 1971a). 

In upland soil, which was well-drained and had 74-75% sand, larvae were observed up to a 

depth of 22-28 cm in nursery soil and 22 cm under transplanted crops (Rao and Israel, 1972b). In 

poorly drained lowland soil, larvae were observed up to a depth of 18 cm in both nursery and 

transplanted areas. In lowland soils the pore space was less and moisture content high so that the 

rice roots spread more laterally and nematode populations were greater at a depth of 2-6 cm 

compared with maximum density observed at a depth of 4-12 cm in upland soils. Coarse and 

medium soils with particles above 0.053 mm in diameter and sandy soils allowed free movement 

of infective larvae and invasion into roots of the rice plant. Clayey soils were less suitable to 

nematode infection. With an increase in the sand content of the test soils, there was an increase 

in root growth, root-knot development and egg mass production by the nematode, the 

relationship between the sand content and the activity of the nematode was linear (Rao and Israel 

1972d). 

Sandy or loamy, laterite soils or recent alluvial soils (in which the available soil nutrients 

range from moderate to low and water holding capacity is low) favour development of the 

nematode. It has been observed that waterlogged condition in the direct seeded rice or 

transplanted crop had no detrimental effects on the survival of the endoparasitic stages (Prasad et 

al. 1985). Temperature of 22-29°C was found to be suitable for the prevalence of the nematode 

(Rao and Israel, 1973). Factors such as nutritional deficiencies, poor drainage, and soil-borne 

diseases can conceal the presence of nematodes. 

Population density of M. triticoryzae declined in puddled soil. Puddling reduced the bulk 

density of soil and decreased the hydraulic conductivity in the upper layers but not in the deeper 

layers where soil aeration was reduced due to high moisture levels retained in the puddled soil. 

The invasion of the roots by the second-generation infective juveniles was reduced. The 

population density of the root-knot nematodes was higher in the non-puddled soil especially in 

unsubmerged condition compared to puddled and submerged soil. However, where the seedlings 





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were already infected before transplanting and submergence, the nematode could survive well 

and reproduce within the aerenchyma of the root (Chandel et al., 2002) 

Yield losses 

On upland rice, M. graminicola causes 16-32% loss in grain yield due to incomplete filling of 

kernels (Biswas and Rao, 1971; Chakrabarti et al., 1971; Rao and Biswas, 1973). With increase in 

the inoculum given to 10 days old plants of cv. IR 8 by one egg mass, the corresponding reduction 

in grain yield was computed as 2.6% (Rao and Biswas, 1973). The threshold level to cause 10% loss 

is 120, 250 and 600 eggs/plant at 10, 30 and 60 days age of plants in direct seeded rice (Rao and 

Biswas, 1973). Severe infestations of the root-knot nematode, M. graminicola on rice was 

observed in Mandya district of Karnataka state covering an area of 1500 ha. The seedlings 

exhibited profused galling on roots and depletion in vigour, yellowing, stunting and curling of 

leaves. Some of the farmers could not grow plantable seedlings even after raising the nursery for 

the third time. Several farmers simply ploughed-in the nurseries as the infested seedlings were not 

fit for planting (Prasad et al., 2001). 

Management 


Rice varieties Loknath 505 and M-36 were found to be highly resistant to the rice root-knot 

nematode, M. graminicola at Allahabad (Hassan et al., 2004). Senthilkumar et al. (2007) reported 

varieties TKM 3, TKM 7, TKM 8, TKM 9, MDU 1, MDU 2, TKM 11 and PY 1 to be resistant to the 

nematode. Simon (2009) evaluated the susceptibility of 53 rice genotypes to M. graminicola in 

field and pot experiments and observed that 13 cultivars were highly resistant to this nematode. 

Rice root-knot nematode, M. graminicola was reported to reproduce on all the 10 wild Oryza 

species tested. O. australiensis Domin and O. brachyantha Chev and Rochr showed by far the 

greatest infestation (5855 and 10,235 juveniles/g root, respectively) compared with O. officinalis 

Wall., which recorded the lowest infestation (240 juveniles/g root). O. latifolia Desv., O. ridleyi 

Hook. f. and O. rufipogon Griff. recorded

selections from IR36/RD25 crosses showed resistance to Meloidogyne spp. in Thailand 

(Arayarungsarit et al., 1985). Soriano et al. (1999) observed that one accession of O. 

longistaminata A. Chev. represented by two individuals (WL02-2 and WL02-15) and three 

accessions of O. glaberrima (TOG7235, TOG5674 and TOG5675) were resistant to M. graminicola 

whereas all the O. sativa accessions were susceptible. Poudyal et al. (2004) reported that all the 

cultivars tested were susceptible to M. graminicola except Masuli and Chaite-6, which were 

moderately resistant. Bose et al. (1998) conducted RAPD analysis on five rice cultivars including 

three highly resistant (Ramakrishna, Rasi and Kalarata) and two highly susceptible (Annapurna and 

Kiran) to M. graminicola. The highest polymorphism was recorded between Annapurna and 

Ramakrishna. They suggested that cross combination of Annapurna and Ramakrishna could prove 

useful for mapping the M. graminicola resistance gene in rice. 

Prasad et al. (2006) observed that irrespective of the recurrent parent background, the 

percentage of resistant lines was higher in lowland stress selection compared to that in upland 

stress and the resistance to M. graminicola is not monogenic and support the multigenic nature of 

inheritance. Shrestha et al. (2007) identified quantitative trait loci (QTLs) for partial resistance to 

M. graminicola using a mapping population based on two rice varieties, Bala (tolerant) × Azucena 

(susceptible). M. graminicola did not significantly reduce yield in Bala, but caused a yield reduction 

of almost half in Azucena, suggesting that the partial resistance to nematode establishment was 

related to nematode tolerance. A total of six putative QTLs for nematode tolerance were 

detected. For two of the QTLs detected, Azucena was the donor of the tolerance alleles, 

suggesting it may be possible to breed plants with greater tolerance than Bala. 

Biological control 

Maximum mortality (>96%) of M. graminicola juveniles was recorded when exposed to 

culture filtrates (100 and 50% conc.) of Trichoderma harzianum Rifai (Pathak and Kumar, 1995). 

Application of Pseudomonas flourescens @ 20 g/m 2 was found to be effective in reducing the 

nematode numbers and increasing the grain yields (ACRIPN, 2003). In in vivo screening tests, 

Bacillus megaterium significantly reduced nematode galling and J2 penetration compared with 





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uninoculated controls. Additionally, in in-vitro tests using culture filtrates of B. megaterium 

significantly delayed nematode egg hatch and reduced J2 mobility (Padgham et al., 2005). Isolates 

of endophytic and rhizosphere fungi viz., Fusarium and Trichoderma are the potential biological 

control agents against M. graminicola in rice (Le et al., 2009). 


Prot et al. (1994) positively correlated the nitrogen concentration in roots with initial 

population and the number of juveniles of M. graminicola recovered from the roots. They 

observed that nitrogen application increased growth and yield whether plants were infested by 

the nematode or not. However, since the percent of yield loss remained approximately constant 

for a given initial population across the range of nitrogen quantities applied, nitrogen applications 

do not reduce the relative nematode effect. 

Soil amendments with decaffeinated tea waste or water hyacinth compost (300 or 

600g/4.5 kg soil) reduced root-knot nematode infestation and increased plant growth (Roy, 1976). 

Rice-mustard-rice crop sequence, followed by rice-maize-rice and rice-fallow-rice were effective in 

reducing nematode development (Kalita and Phukan, 1996). A drastic decline of 98 and 94% in the 

population of Meloidogyne spp. and Hirshmanniella oryzae respectively, were recorded when the 

rice crop was rotated with brinjal (Ramakrishnan, 1995). Crop rotation with non-host crops viz., 

sweet potato, cowpea, sesamum, castor, sunflower, soybean, turnip and cauliflower inhibit 

nematode development (Rao et al., 1984; Rao, 1985). Polthanee and Yamazaki (1996) observed 

that in situ green manuring with marigold suppresses root galling and increases rice grain yield by 

46% over the untreated check. The increase in yield was attributed to a reduction of nematode 

densities in soil by marigold. In addition, marigold plant materials may serve as organic manure 

and provide nutrients for rice growth. Burning of 15 cm deep rice hulls significantly reduce M. 

graminicola populations in the soil (Gergon et al., 2001). 

Compared with continuous rice treatments (averaged over burning and mulching 

treatments), treatments with fallow or cowpeas in the previous year had 32% less herbaceous 





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weed biomass, 90% fewer A. conyzoides and over 99% fewer M. graminicola in field trials (Roder 

et al., 1998). In Philippines, rice based cropping sequences such as rice-mungbean, corn-cabbage- 

rice, rice-tobacco-rice, rice-watermelon-rice, rice-cotton-rice, have been found effective in 

combating root-knot nematode menace in rice (Davide and Zorilla, 1983). 


Carbofuran, phorate, isazophos, cartap, carbosulfan or quinalphos when given as soil 

application @ 1 kg a.i. /ha significantly reduces the root galling by M. graminicola (Panigrahi and 

Mishra, 1995b). Fademi (1994) suggested a dosage of 2 and 3 kg a.i./ha for early and late 

applications for best results for the control of M. incognita. Lopez and Salazar (1989) found 

fenamiphos (@ 6 kg a.i./ha) to reduce root-knot index of M. salasi in rice. Oxamyl @ 500 to 1000 

ppm when applied as foliar sprays were effective in reducing M. graminicola followed by soil 

application of phorate and carbofuran @ 1 kg a.i./ha. (Krishnaprasad and Rao, 1984). 


Avoiding excessive usage of nitrogen; Application of non-edible oil cakes to the nursery; soil 

application of carbofuran @ 1 kg a.i./ha to the nursery 7 days prior to uprooting of the seedlings; 

Crop rotation with non-host crops and educating the farmers about the biology of nematode are 

the viable options. 

Cyst nematode (Heterodera oryzicola Rao and Jayaprakash, 1978) 

History 

On receiving the reports on appearance of tungro virus disease symptoms in Kerala state 

During Kharif season, 1976, Dr. Y.S. Rao, nematologist from Central Rice Research Institute, 

Cuttack visited that area as a member of the multi-disciplinary team and found a few symptoms to 

be different from that of tungro disease. He isolated nematode cysts associated with the roots of 

diseased plants from the modan lands of Kerala. Later, this cyst nematode was described as a new 

species Heterodera oryzicola in 1978. Though occurrence of a cyst nematode species H. oryzae 





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was reported from India earlier to this (Rao, 1965), no further information was available on this 

species. Other cyst nematode species reported from rice include H. sachhari (Odihirin, 1975) and 

H. skohensis (Kaushal et al., 2000). 

Distribution 

H. oryzicola is widely distributed in Kerala, India (Rao and Jayaprakash, 1977; Kuriyan, 

1985; Raveendran et al., 1976; Venkitesan, 1979; Charles and Venkitesan, 1990). Koshy et al. 

(1987) recorded this nematode on banana in Goa. Gupta et al. (1977) reported occurrence of a 

cyst nematode in paddy fields in Bankura and Burdwan in West Bengal. H. skohensis, was reported 

from rice and wheat fields of Kangra valley in Himachal Pradesh (Kaushal et al., 2000). 

Fig. 7 . Cyst nematode, Heterodera oryzicola on rice root. 





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Host range 

H. oryzicola has limited host range with rice and banana being major hosts. 

Some weeds such as Cynodon dactylon and Brachiaria sp., are good hosts and Kyllinga 

monocephala Rottb., is a poor host of this nematode (Charles and Venkitesan, 1990). 


Browning and chlorosis of leaves, stunting, and early flowering by 10-13 days are 

common symptoms observed in the cyst nematode infected plants. The roots do not show any gall 

formation, but turn brown at the site of infection and show depletion in vigor (Rao and 

Jayaprakash, 1977). In advanced stages minute cysts can be seen on roots. 


Females of H. oryzicola deposit many eggs into large egg sac attached to the vulval cone 

(Fig.7 ). Juveniles in egg sacs hatch freely in water under the influence of exudates from rice roots 

(Jayaprakash and Rao, 1982). The embryonic development and the emergence of infective 

juveniles completes in eight days. Penetration into roots takes place in one day. Duration of 

development was 6 days for second; 4 and 8 days for the third stage male and female and 5 and 8 

days for fourth stage male and female juveniles, respectively. Endoparasitic juveniles developed in 

10 days into males and to white females in 20 days. Virgin females secreted a strong male 

attractant. Eggs are deposited in a gelatinous matrix secreted by the female in 22 days and the 

females turn into brown cysts 2 days later. A single female on an average produces 198 eggs in egg 

mass and retained 120 eggs inside the cyst. The life cycle is completed in 24-30 days, which allows 

multiple generations depending on the duration of the crop (Jayaprakash and Rao, 1982). 

Host parasite relationship 

H. oryzicola juveniles in egg sacs hatch freely in water but there is evidence that 

exudates from rice roots are required to stimulate the hatch (Jayaprakash and Rao, 1982). 

Hatching of eggs from cysts was significantly high in thiamine hydrochloride followed by rice root 





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deffusates. Potent root deffusate was produced at 45 days age or at maximum tillering stage of 

crop. Dilution of root exudates even 16 times was effective as a hatching agent (Jayaprakash and 

Rao, 1982). However, Ibrahim et al. (1993) observed markedly different hatching behaviours in H. 

sacchari and H. oryzicola. Irrespective of the age of the host plant producing cysts, H. oryzicola is 

dependent on root deffusates to induce substantial hatch. The dependence of H. sacchari on 

diffusates is less easily defined; it is only with cysts from the last two extractions that a small 

proportion of eggs were dependent on root deffusates for hatch and the total percentage hatch 

from these cysts was considerably less than from cysts collected from younger plants. 

H. oryzicola infection induces several biochemical changes in rice plants. The plants 

infected with H. oryzicola showed a significant reduction in the leaf chlorophyll, N, P, K and Fe in 

roots and shoots. Calcium and sodium content of plants increase to compensate the loss of K and 

maintain the cation balance due to impairment of photosynthesis. Total and soluble sugars and 

proteins were reduced and starch accumulated in both roots and shoots. Soluble amino acids 

increased and there was no change in DNA (Rao et al., 1988). 


Infection of Sclerotium rolfsii in roots was enhanced in the presence of cyst 

nematodes while the penetration of nematode into roots and cyst formation was inhibited in the 

seedlings inoculated with the fungus (Jayaprakash and Rao, 1984). Antagonistic interaction 

occurred between M. graminicola and H. oryzicola has been reported. M. graminicola establishes 

faster and suppresses the multiplication of H. oryzicola (Rao et al., 1984). 

Yield losses 

`Estimated yield losses due to H. oryzicola infestations varied from 21-42% (Kumari and 

Kuriyan, 1981). The threshold level to cause 10% loss was 85-100 infective juveniles per plant up 

to 30 days age of plant (Rao, 1985). 





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Management 


Very limited information is available on the sources of resistance in rice germplasm to cyst 

nematodes. Lalnakanda-41, CR 143-2-2, Ratna, Hamsa, Mtu-17 and Mtu-4 were found resistant to 

H. oryzicola (Jayaprakash and Rao, 1983). Out of 73 wild rice accessions screened, 15 of O. 

glaberrima Stevd. and 7 of O. barthii A. Chev. were resistant (Reversat and Destombes, 1998) to a 

Congolese population of H. sacchari. 


Crop rotation with non-host crops is very effective against cyst nematode as it has a limited 

host range. Coyne and Plowright (1998) reported that solarization technique worked well against 

H. sacchari. They opined that the temperature rise (5.75 o C) due to solarization would seem 

unlikely to kill eggs within cysts, but it may influence H. sacchari population densities by 

encouraging egg hatching in the absence of the host. Pot experiments have shown that maize, 

millet and sorghum, which are commonly cultivated in cropping systems with rice in West Africa, 

were poor hosts of H. sacchari, but would maintain cysts in soil (Coyne and Plowright, 1999). 

Rotations with soybean or sweet potato were found to be helpful in the management of H. 

elachista (Nishizawa et al., 1972). 


Soaking rice seed in 0.2% solution of oxamyl or carbosulfan @ 250 ppm reduce cyst 

development in H oryzicola (Rao, 1985). Soil application of carbofuran or phorate @ 1 kg a.i./ha, at 

7 and 50 days after planting reduce the incidence of the nematode by 70% and increase grain yield 

by 28% (Kuriyan, 1985). 





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Use of chemicals for the control of cyst nematode should be discouraged. Since the cyst 

nematode has a limited host range, crops other than rice or banana should be grown wherever 

the nematode is a problem. As Kerala state is rich in banana germplasm and the crop is good host 

of H. oryzicola, care should be taken to use certified and disinfested planting material to avoid the 

establishment and spread of the nematode to other parts of the country through infected planting 

material. In Kerala State, where the cyst nematode is a problem, land available for rice cultivation 

is reducing over years due to urbanization and land requirement for human inhabitation. Under 

continuous paddy cultivation, shift of the nematode populations on rice and other hosts should be 

critically determined to develop sustainable cropping systems. 

Rice root nematode (Hirschmanniella spp.) 

History 

Several species of Hirschmanniella have been reported in association with irrigated rice all 

over the world. Literature prior to 1968 may deal with mixtures. In India H. oryzae and H. 

mucronata are the dominant species infecting rice crop. Previously, H. oryzae was thought to be 

associated with a serious disease of rice called "Mentek."(Timm and Ameen, 1960; Van der Vecht, 

1953). Due to the practice of thorough puddling and levelling of soil prior to transplanting of 

irrigated rice, the rice root nematode populations get evenly distributed in the field. Hence, unless 

suitable treated (nematode free) plots are maintained side-by, the uniform retardation in the crop 

growth in the infested fields cannot be distinguished (Prasad et al., 1987). 

Distribution and host range 

Sympatric prevalence of two or more species of Hirschmanniella has been 

reported from irrigated, semi-deepwater and deepwater rice environments (Mathur and Prasad, 

1971; Sivakumar and Khan, 1982; Prasad et al., 1987; Varaprasad et al., 1992). Both H. oryzae and 

H. mucronata were collected from Cajanus cajan, Cicer arietinum, Pennisetum typhoides, Pisum 





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sativum and Triticum aestivum rhizospere. Several weeds grown in and around rice fields have 

also been reported to host Hirshmanniella spp. (Mohandas et al., 1979) especially from the family 

Cyperaceae and Graminae (Van der Vecht and Bergman, 1952). Kumar (1990) reported that 

Echinochloa colona, Sesbania aculeata, Cyperus rotundus, Boerhavia diffusa, Eclipta alba and 

Polygonum plebejum harboured H. oryzae. 


Infestation by the rice root nematode results in retardation of growth rate and reduced 

tillering in early growth stages and flowering may be delayed by 14-15 days (Muthukrishnan et al., 

1977). MacGowan (1979) observed that the infection by H. oryzae is not expressed by any 

recognisable field symptoms. Infected roots first showed a yellowish to brown colour which get 

darkened over time. Heavily infected roots eventually decay. Infected seedlings showed reduced 

survival, delayed emergence of tillers and discoloured older leaves. Rapid root regeneration often 

results in plant recovery. 

Fig. 8 . Hirschmanniella spp. in rice roots. 


Eggs of H. oryzae are deposited in the root cortex and hatching occurs 4-6 days after 

deposition (Mathur and Prasad, 1972). The life cycle is completed in 30 days. All stages feed on the 

cortical cells and central vascular region of rice roots (Fig. 8 ). H. oryzae completes one generation 

in Northern India (Mathur and Prasad, 1972) and two generations in Japan (Ou, 1985) and three 

generations in Senegal (Fortuner and Merny, 1979) in a cropping season. Mahapatra and Rao 





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(1980) reported 4 peaks of H. mucronata population i.e. during the last week of September, first 

week of November, 3rd week of December and 2nd week of February. The nematode was active 

particularly in the presence of standing crops. They observed positive correlations between the 

fresh weight of roots and soil temperature at 5 cm depth, and the build-up of the nematode 

population. Maximum root populations were recorded at tillering stage of the crop (Rao, 1985). 


The nematodes once within a rootlet proceed through the parenchyma toward the base. 

The distance of migration by an adult nematode in root tissue was estimated to be in the range of 

6.3 to 10.3 mm in rootlets from the first node. Goto (1973) observed that H. imamuri was fairly 

evenly distributed from the tip to the base in the roots, with a comparatively high frequency in the 

section 11 to 60% of the length from tip on rice rootlets in paddy fields. Besides direct damage 

caused by feeding, intra and inter-cellular migration in cortex of roots, H. oryzae and H. mucronata 

cause degeneration of physiological functioning of roots (Mahapatra and Rao, 1973). As a result, 

retardation of growth rate and decrease in tillering occurs in early growth stages and flowering 

can be delayed by 14 days (Muthukrishnan et al., 1977). At low levels of nematode infestation 

(below 50 nematodes/g root) there was no appreciable reduction in chlorophyll, starch and 

proteins in plants and, hence, H. oryzae and H. mucronata were found to be highly successful 

parasites causing least lethal changes in nematode-plant interface (Jayaprakash et al., 1981; 

Prasad et al., 1982). Consequently, symptoms are not discernible in foliage till large populations of 

the nematodes build up. Hirschmanniella spp. can survive between crops on weeds. Khuong 

(1987) observed that population densities of rice root nematodes in root systems were lowest at 

post transplanting and highest at heading stage. Number of nematodes in root tissue increased by 

20-22 times from transplanting to heading stage. Population densities of Hirschmanniella spp. in 

two-crop rice fields were more than twice those in one-crop rice fields. Ramakrishnan (1992) 

reported that populations of H. oryzae and H. mucronata were low at transplanting. After 30 days 

the nematodes multiplied rapidly and reached a peak 60 days after transplanting. The high 

nematode population remained highest up to 90 days after transplanting and declined thereafter. 





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Mohandas and Rao (1982) inoculated the sprouts of variety Jaya with H. oryzae at 0, 10, 

100, 1000 or 10 000/seedling and observed significantly less height and number of leaves of 

seedlings receiving 10, 000 nematodes on the 7th day. In addition, 10% and 60% mortality was 

recorded in the above treatment on the 14 th and 21 st day, respectively. The height and number of 

leaves of seedlings were slightly more in treatments receiving 10, 100 and 1000 nematodes than 

in controls, up to the 14th day. However on the 21st day, the number of leaves started to show a 

gradual reduction in these treatments compared with controls. Maximum number of tillers was 

recorded in the control followed by 10, 100 and 1000 inoculum levels, respectively with no tillers 

at the highest level. On the 29th day, there was a further reduction in height, number of leaves 

and tillers. Leaf area and fresh weight of shoot and root were also reduced. 


Gokulapalan and Nair (1986) reported highest sheath blight disease intensity in rice cv. 

Jyothi receiving the highest H. oryzae inoculum (1000 juveniles/plant) along with Rhizoctonia 

solani. Plant height was reduced at all levels of nematode inoculum (10-1000/plant) both alone 

and in combination with the fungus. The soil and root populations of H. oryzae were higher when 

fungus was inoculated. 


Youssef (1999) found a positive correlation between the root population of H. 

oryzae and prevalent soil temperature, whereas, negative correlation was observed between the 

soil nematode population and soil temperature. 

Yield losses 

Yield losses due to rice root nematodes range from 25 to 42% (Hollis and 

Keoboonrueng, 1984; Fortuner, 1977 and 1985). In pot experiments, 21-day-old rice cv. IR20 

seedlings, inoculated with H. oryzae at 1 and 10 nematodes/g soil caused yield loss of 27 and 40%, 

respectively (Jonathan and Velayutham, 1987). Cho-Hen et al. (1994) reported that H. oryzae may 





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educe the yield by 8.3% in old lowland areas, 9.4% in new lowland areas, but no losses in new 

upland areas. The nematode depressed tillering, root growth and shoot growth. Poussin et al. 

(2005) observed that the average single grain weight was the most affected yield component. 

Application of nitrogen @ 80 kg N/ha increased the weight of each grain, but this effect was 

largely reduced in the presence of H. oryzae. Even though nitrogen amendments were able to 

counterbalance the negative effects of H. oryzae, nitrogen applied at 80 kg N/ha level was not 

considered a sustainable alternative because it increased nematode populations. 

Management 


Rice cultivars belonging to the Japonica group were more susceptible to H. oryzae than the 

other groups (Youssef, 1999). Significantly higher populations of H. oryzae were found in Pakistan 

Basmati and Basmati 370 while Basmati 385 supported the lowest population (Randhawa et al., 

1992). Ramakrishnan et al. (1984) found rice cv. TKM 9 to be resistant at all growth stages against 

H. oryzae. Wild rice species, Porteresia coarctata showed the highest degree of resistance to H. 

mucronata (Panigrahi and Mishra, 1995a). Oryzae collina and O. nivara were moderately resistant 

to H. oryzae 

Biological control 

Jacq and Fortuner (1979) observed that H. oryzae population levels in the soil of rice fields 

were lower when the activity of sulphate reducing bacteria was high. They suggested that further 

reduction in nematode population is possible by artificially increasing bacterial activity after the 

harvest. This method is harmless to the rice, controls nematode population well and increases 

yield. Application of Pseudomonas fluorescens Migula strain Pf-1 as seed treatment @ 10 g/kg of 

seed was superior to the treatments as nursery soil application, separately and either with or 

without soil application of carbofuran 3G @ 1.3 g a.i./m 2 , in the management of H. gracilis and 

increasing crop yield (13%) over the control (Ramakrishnan et al., 1998 and 1999). 





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There was more build-up of H. oryzae in crops transplanted in mid June, while populations 

were minimal when transplanted in mid July (Randhawa, 1991). Hendro et al. (1992) reported that 

Sesbania rostrata Brem. and Aeschynomene afraspera L. can effectively control H. oryzae and H. 

mucronata populations. Prot et al. (1992) opined that as these two leguminous crops do not 

generate direct return, using them to control the rice-root nematodes was not economical, 

despite significant yield increase was obtained with their cultivation. Further, Prot (1994b) 

observed that S. rostrata when applied as a green manure reduced field populations of 

Hirschmanniella spp., however, it is a very good host for Meloidogyne graminicola when grown in 

non-flooded soils. Hence its cultivation as a green manure before rice in non-flooded soils infested 

by M. graminicola may increase their number considerably. It is suggested that under rain fed 

conditions S. rostrata should not be used and other leguminous crops resistant to M. graminicola 

should be used as alternatives. Presence of weeds resulted in suppression of H. oryzae 

populations (Coyne et al., 1999). Germani et al. (1985) reported that the dry weight of paddy, 

culms and leaves, and number of culms in rice following Sesbania were 214, 158, and 121% 

greater, respectively, than those following rice. Ripening also occurred earlier if rice followed 

Sesbania rostrata. They have attributed beneficial effects to the trap-crop action of S. rostrata 

against H. oryzae. 

Application of neem cake @ one t/ha and press mud at 10 t/ha (Johnathan and Pandiarajan, 1991) 

and castor oil cake and mustard oil cake (Khan and Shaukat, 1998) significantly reduced H. oryzae 

populations. The nematicidal activity against H. oryzae was more in methanol extract of Tagetes 

erecta L. than in the methanol extract of T. patula L. (Youssef, 1998). Rotations of rice with 

cabbage and tobacco reduced populations of H. oryzae by 83-88% in paddy field experiments but 

winter ploughing and fallow were effective to a lesser extent (Gao XueBiao et al., 1998). 





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Ramakrishnan et al. (1984) reported that phosphamidon and chlorpyriphos given as 

root dips @ 0.02% for 20 min before planting reduced H. oryzae population to 0.83/2g root 30 

days after transplanting. Bare root-dip tretment, application of chlorpyriphos after infestation of 

H. gracilis, and triazophos, UC51762, BPMC and phenamiphos given as prophylactic treatments 

were very effective in the management of nematodes (Balasubramanian and Palanisamy, 1983). 

Lahan et al. (1999) observed that carbosulfan and phosphamidon @ 0.3% as root-dip treatment, 

reduced the H. oryzae population by 46.2% and 40% and increased grain yield by 35.1 and 34.7 

q/ha respectively over untreated control. 

Application of carbofuran @ 1.275 kg a.i./ha to nurseries (Jonathan and 

Velayutham, 1984), significantly reduces H. oryzae populations up to 30 days after transplanting. 

Application of carbofuran @ 1 kg a.i./ha to main fields significantly decreased H. gracilis 

population in roots and soil with significant improvement in plant growth (Ahmad et al., 1984). 

Carbofuran treatments (1 kg a.i./ha) reduced nematode incidence and increased yield by 39.4%. 

Carbofuran application in nursery soil and in main field at 7 and 50 days after transplanting 

significantly reduced nematode population and increased yield by up to 42.8% (Prasad and Rao, 

1984). Using carbofuran to control Hirschmanniella spp., increased the rice yield by 23.6% at the 

turn green stage, 18.8% of rice yield in the beginning of tillering and 12.3% in the transplanting 

stage (Zhang and Ai, 1994). The application of chemicals in standing water or mud balls was 

inferior to soil incorporation in controlling the rice root nematode H. mucronata (Prasad et al., 

1986). 


Incorporation of Sesbania rostrata and Aeschynomene afraspera; incorporation of non- 

edible oil cakes in the nurseries; application of carbofuran @ 1 kg a.i./ha to nursery 7 days prior to 

uprooting and to main field 45 days after transplanting are good control measures against this 

nematode. 





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Lesion nematode (Pratylenchus spp.) 

History 

Lesion nematodes Pratylecnhus species are reported on rice from many countries. About 

ten species of Pratylenchus were reported on rice among them P. zeae and P. indicus are most 

common in rice. Rice is grown under rain fed conditions in most of the states in India during the 

kharif season. The root lesion nematode invariably infects upland rice and mortality of the 

seedlings is not uncommon. However, the infection either goes unnoticed or attributed to some 

other causes mainly due to the lack of awareness about this nematode pathogenicity. Since more 

than70% rice area in North Eastern states is in uplands and hilly tracts, this nematode plays an 

important role in this region. 

Distribution 

Lesion nematodes are widely distributed in the world and mainly inflict damage to direct 

seeded rain fed rice. In India, Pratylenchus spp., particularly P. indicus and P. zeae have been 

recorded on rice in Andhra Pradesh, Assam, Gujarat, Kerala, Orissa, Madhya Pradesh, Rajasthan, 

Uttar Pradesh and West Bengal (Prasad et al., 1987). 

Host range 

Rice is a good host of P. indicus. Among 52 weed species recorded from rice growing areas 

of western Cuba, 12 were recorded as hosts of P. zeae. Cyperus iria and Eleusine indica (L.) 

Gaertn. were good hosts for both the nematodes (Fernandez and Ortega, 1982). Weeds viz., 

Cynodon dactylon, Amaranthus spinosus L., Dactylotenium aegypticum (Desf.) Beauv., Digitaria 

sanguinalis Scop. and Echinochloa sp. supported population of P. zeae (Fortuner, 1976) and 

Cyperus iria, C. haspan L., C. rotundus L., Euphorbia hirta L. and crop plants green gram, Bengal 

gram, groundnut and wheat were hosts of P. indicus (Prasad and Rao, 1986). 





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The nematode infected plants are stunted and even smothered resulting in patchy growth 

in fields (Fig. 9 ). Chlorosis of leaves and reduction in number of ear heads and grains is also 

common. The nematode infected roots appear swollen with water soaked lesions which develop 

in to black necrotic lesions on the root surface. In advanced stage of damage, the lesions coalesce 

and the whole root turns black in color (Prasad and Rao, 1981). The nematode infected roots 

decay, and when such plants are pulled without proper care, the infected root portions and 

associated population remain in the soil. 

Life/disease cycle : P. indicus completes its life cycle in 33-34 days and several overlapping 

generations occur in a single crop (Prasad and Rao, 1982). The nematode invades the plant roots 

at a selected point singly or in groups. After gaining entry the nematode feeds on the cortical cells 

and forms galleries (Fig. 10 ). Infected roots develop water soaked lesions and yet at times 

swellings are also observed. The necrotic patches coalesce together resulting in brown to black 

lesions (Prasad and Rao, 1981). The optimum temperature for P. indicus reproduction is 23-30°C 

and peaks of population are always preceded by rainfall (Prasad and Rao, 1979). P. zeae 

reproduction was greatest after flowering and the population increased as the crop proceeded 

towards grain maturity. P. indicus populations declined rapidly during fallow (Prasad and Rao, 

1978a) whereas, P. zeae survived in clean fallow up to 6 months. 





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Fig. 9 . Root lesion nematode, Pratylenchus spp.infested rice field. 

Fig. 10 . Root lesion nematode Prtaylenchus indicus inside rice roots. 


Crop damage is directly influenced by the initial nematode population at the 

germination. Usually, the root damage and resultant mortality occurs by 30-40 days after 

germination. Upland rice is taken up with the first showers and seedling survival and yields 

are completely dependent on the progress of rainy season. If the mortality occurs in the 

irrigated rice, gap filling is possible. Such chance is not available in upland rice and seedling 

mortality or root damage that disables the plant to utilize the meagre water resources 

affects the yield. 

Yield losses 

Yield losses caused by P. zeae and P. indicus in India are 13-29% and 33% 

respectively (Prasad and Rao, 1978b). The yield loss is mainly due to improper filling of 





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kernels, leading to reduction in the weight and protein content of grain (Rao and Prasad, 

1977). 

Management 


Application of carbofuran or phorate @ 1 kg a.i./ha soil in the affected crops reduces the 

nematode injury and avert losses in grain yield up to 48.5% (Prasad et al., 1988). However, as the 

yields are very low in uplands, the chemical treatment may not be cost effective. Further, as the 

lesion nematode damage can be known much late in the season, the chemical treatment may not 

be effective for the existing crop. 


Fallowing or rotation with Phaseolus radiatus decreased the root-lesion nematode, P. 

indicus populations in rice cv. Annapurna, but populations increased in rotations involving 

Carthamus tinctorius L. or Nicotiana tabacum L. (Prasad et al., 1983). Sahoo and Sahu (1994) 

examined relative efficacy of oil cakes of neem (Azadirachta indica A. Juss.), karanj (Pongamia 

pinnata L. Pierre), mustard (Brassica juncea (L.) Czern. and Coss), polanga (Calophyllum inophyllum 

L.), til (Sesamum indicum L.), groundnut (Arachis hypogea L.), mahua (Madhuca latifolia (Roxb.) 

Macb.) and cotton (Gossypium sp. L.). Neem cake gave greatest reduction in nematode 

population. 


Growing greengram or blackgram as inter crop or in rotation with rice help in reducing 

the populations of lesion nematodes. 

Ectoparasitic nematodes 

Several ectoparasitic nematodes viz., Aphelenchus avenae Bastian, 1965; A. maximus 

Das,1960; Aphelenchoides asterocaudatus Das, 1960; Basira elegans Patil and Khan, 1982; B. 





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graminophila Siddiqi, 1959; Boleodoroides oryzae Mathur et al., 1966; Caloosia exilis Mathur et 

al., 1969; C. heterocephala Rao and Mohandas, 1976; C. parlona Khan et al., 1969; C. paxi Mathur 

et al., 1969; Criconemella sp., Criconemoides spa.; Filenchus filiformis (Butschlii,1873) Meyl, 1961; 

Gracilacus janai Baqri, 1978; Helicotylenchus spp. Steiner, 1945; H. abunaamai Siddiqi, 1963; H. 

dihystera (Cobb,1893) Sher, 1961; H. indicus Siddiqi, 1963; H. crenecauda Sher, 1966; H. retusus 

Siddiqi and Brown, 1964; Hemicriconemoides cocophillus (Loos,1949) Chitwood and 

Birchfield,1967; Longidorus elongatus Thorne and Swanger,1936; Macroposthonia rustica 

(Micoletzky,1915) De Grisse and Loof, 1965; M. onoensis (Luc, 1959) De Grisse and Loof, 1965; M. 

ornata (Raski,1958) De Grisse and Loof, 1965; Paralongidorus beryllus Siddiqi and Husain,1965; P. 

citri (Siddiqi,1959) Siddiqi et al.,1963; P. oryzae Verma, 1973; Paratylenchus dianthus Jenkins and 

Taylor, 1956; Psilenchus hilarulus de Man, 1921; Rotylenchulus spp; Seinura propora Siddiqi et al., 

1967; Trichodorus sp.; Tylenchus devainei Bastian, 1865; T. hayati Luqman Khan, 1984; T. goodeyi 

Das, 1960; Tylolaimophorus sp.; Trichotylenchus sp.; Tylenchorhynchus sp.; T. annulatus 

(Cassidy,1930) Golden,1971; T. brassicae Siddiqi, 1961; T. claytoni Steiner, 1937; T. elegans Siddiqi, 

1961; T. indicus Siddiqi, 1961; T. mashhoodi Siddiqi and Basir, 1959; T. zeae Sethi and Swarup, 

1968; Xiphenema elitum Khan et al., 1976; X. insigne Loos, 1949 and X. arbum Siddiqi,1964 were 

found to coexist along with the endoparasitic nematodes in the rice rhizosphere (Prasad et al., 

1987). 

Rao et al. (1986b) observed Tylenchorhynchus spp. widely distributed in upland well 

drained soils and feeding by these nematodes debilitates the plant growth. The nematode 

population could be brought under check by crop rotation with Sesamum or Basella rubra. Rao 

and Mohandas (1976) described the ectoparasitic nematode Caloosia heterocephala feeding on 

rice plants (Fig. 11 ). Padhi (1979) reported that Helicotylenchus abunaamai could survive in the 

soils without hosts for a period of 7 months. Both these nematodes have wide host range. They 

could be effectively controlled by soil application of carbofuran or phorate @ 1 kg a.i./ha. Most of 

the ectoparasitic nematodes have wide host range. Their role as pests in the rice based cropping 

system needs further studies. 





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Fig. 11 . Ectoparasitic nematode, Caloosia heterocephala feeding on rice roots. 

Future prospects 

Future studies on rice nematodes should focus on the following aspects viz., developing 

precise distribution maps of important nematode species infecting rice, creation of 

awareness among farmers and extension workers about nematode pests, development of 

locally feasible, low cost and sustainable nematode management methods, development of 

sustainable rice based cropping systems with due consideration to 

susceptibility/tolerance/resistance of the component crops to rice nematodes, exploiting the 

antagonistic potential of fungal and bacterial endophytes for nematode management and 

developing effective low-cost delivery system for these microbes, incorporation of resistance 

in to agronomically superior cultivars using conventional breeding and 

biotechnological/transgenic approaches. Application of molecular techniques for 

understanding host parasite relationships. 





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References 

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Hooghly (West Bengal). Bull. Zoo. Sur. Ind. 5: 85-91. 

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Aleksandrova, I.V. and Beloglazov, A. D. (1989). Use of regulated gas medium in the control of the 

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Allen, M.W. (1952). Taxonomic status of the bud and leaf nematodes related to Aphelenchoides 

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Bajaj, H.K. and Dabur, K.R. (2000). Cyperus deformis, a new host record of rice root-knot 

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Balasubramanian, P. and Palanisamy, S. (1983). Evaluation of bare root dip with chemicals for the 

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Bose, L.K., Sahu, S.C., Mishra, C.D. and Ratho, S.N. (1998). Molecular polymorphism between rice 

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