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*Corresponding author: P Mandal, Department of Agronomy and Haor Agriculture, Sylhet Agricultural University, Sylhet-3100, Bangladesh, Email: [email protected] J. Sylhet Agril. Univ. 4(2):161-172, 2017 ISSN: 2308-1597 STATUS OF AGRONOMIC STRATEGIES TO CONTROL ROOT-KNOT NEMATODE (Meloidogyne spp.) IN POTATO P Mandal * and M S Hossain Department of Agronomy and Haor Agriculture, Sylhet Agricultural University, Sylhet-3100 (Available online at: www.jsau.com.bd) Abstract Root-knot nematode (Meloidogyne spp.) is considered as dreaded in potato production by growers. Because of its wide host range and higher reproduction rate, major control using chemical nematicides gradually phased out due to growing concerns about environmental safety. Therefore, alternative techniques are essential to control them effectively. Control of root-knot nematode through agronomic management practices are very important aspects of organic movement in agriculture around the world. Till now, many plants were identified as antagonistic plants against plant parasitic nematodes. These plants reveal antagonistic nature through the production of secondary volatile and non-volatile exudates in their different parts viz., seed, leaf, flower, root and stem. A single technique may not effective enough to control root-knot nematode in potato as compared with sole chemical control measures. Consequently, combining two or more control techniques in a systemic way is very effective for root-knot nematode control. These techniques can establish confidence among the organic farmers to control root-knot nematode in order to increase yield of potato. This review is focusing some very important and effective agronomic techniques to control root-knot nematode in potato. Keywords: Potato, root-knot nematode, Meloidogyne spp., agronomic control. Introduction Potato (Solanum tuberosum L.) is a well-known vegetable crop throughout the world. After wheat ( Triticum aestivum L.), rice (Oryza sativa L.), and maize (Zea mays L.), it ranks fourth in the world in terms of both production and consumption (Bowen, 2003). To overcome the risk of increasing international food prices, potato is considered as an important food security crop in low-income countries. Apart from its importance in food security, the potato crop has considerable nutritional value, containing high values of carbohydrates, protein, vitamin C, vitamin B6, vitamin B3, potassium, phosphorus and magnesium (Cotton et al., 2004). As a carbohydrate rich food potato contain greater amount of dry matter and protein compared to cereals (Bamberg and Del Rio, 2005). In the human diet, potatoes are also a valuable source of health-promoting antioxidants. In addition, fat percentage in potato is very low. It is a good source of vitamins B1, B3 and B6, minerals and dietary antioxidants, which may play a great role in preventing diseases related to ageing. Carbohydrates and anti-diabetic factors such as antioxidants of potatoes contribute in prevention and management of diabetes (Camire et al., 2009). Potatoes contain a significant amount of fiber, which helps in lowering the total amount of cholesterol in the blood, thereby decreasing the risk of heart disease of human. So, potato plays a significant role for regular functioning of human health. Potato is now growing around 140 countries in the world (Haase, 2007). Unfortunately, the crop suffers from different fungal, bacterial, viral, viroid and parasitic nematode diseases under unfavourable conditions and causes huge losses its production. These invigorate the development of effective processes for sustainable assembly to control potato diseases and pests. Root-knot nematodes have drawn attention in importance in different parts of the world and cause severe yield damage (Karssen et al., 2013; Onkendi et al., 2014). Root-knot nematodes (RKNs) Meloidogyne spp. especially M. javanica, M. incognita, M. arenaria, M. chitwoodi and M. hapla the most harmful agricultural pests among the nematode species, are showing aggressive effects and heavy yield reduction (Anwar et al., 2007). Both tuber quality and quantity can be reduced by 12.2% as a result of nematode attack (Hadwan and Khara, 1992). RKNs (Meloidogyne spp.) impose an important barrier for potato production through their destructive activity. On a global scale, the distribution of nematode species varies greatly. There are more than 50 different species of RKNs (Jones et al., 2013). As potato being highly susceptible to M. javanica, M. incognita, M. arenaria, M. chitwoodi and M. hapla, which all adversely affect the efficiency of roots and yield. Nematodes are multicellular organism mainly found in the group Ecdysozoa, or organisms that are able to change their cuticle. The majority of them are said to be free-living and feed mainly on bacteria, fungi, protozoa and other nematodes, only a minority feeds on parasites, animals and plants (Bélair, 2005). They are motile organisms, with most

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Page 1: STATUS OF AGRONOMIC STRATEGIES TO CONTROL ROOT-KNOT ... · Root-knot nematodes have drawn attention in importance in different parts of the world and cause severe yield damage (Karssen

*Corresponding author: P Mandal, Department of Agronomy and Haor Agriculture, Sylhet Agricultural University,

Sylhet-3100, Bangladesh, Email: [email protected]

J. Sylhet Agril. Univ. 4(2):161-172, 2017 ISSN: 2308-1597

STATUS OF AGRONOMIC STRATEGIES TO CONTROL ROOT-KNOT

NEMATODE (Meloidogyne spp.) IN POTATO

P Mandal*and M S Hossain

Department of Agronomy and Haor Agriculture, Sylhet Agricultural University, Sylhet-3100

(Available online at: www.jsau.com.bd)

Abstract

Root-knot nematode (Meloidogyne spp.) is considered as dreaded in potato production by growers.

Because of its wide host range and higher reproduction rate, major control using chemical nematicides

gradually phased out due to growing concerns about environmental safety. Therefore, alternative

techniques are essential to control them effectively. Control of root-knot nematode through agronomic

management practices are very important aspects of organic movement in agriculture around the world.

Till now, many plants were identified as antagonistic plants against plant parasitic nematodes. These

plants reveal antagonistic nature through the production of secondary volatile and non-volatile exudates in

their different parts viz., seed, leaf, flower, root and stem. A single technique may not effective enough to

control root-knot nematode in potato as compared with sole chemical control measures. Consequently,

combining two or more control techniques in a systemic way is very effective for root-knot nematode

control. These techniques can establish confidence among the organic farmers to control root-knot

nematode in order to increase yield of potato. This review is focusing some very important and effective

agronomic techniques to control root-knot nematode in potato.

Keywords: Potato, root-knot nematode, Meloidogyne spp., agronomic control. [

Introduction

Potato (Solanum tuberosum L.) is a well-known vegetable crop throughout the world. After wheat (Triticum aestivum

L.), rice (Oryza sativa L.), and maize (Zea mays L.), it ranks fourth in the world in terms of both production and

consumption (Bowen, 2003). To overcome the risk of increasing international food prices, potato is considered as an

important food security crop in low-income countries. Apart from its importance in food security, the potato crop has

considerable nutritional value, containing high values of carbohydrates, protein, vitamin C, vitamin B6, vitamin B3,

potassium, phosphorus and magnesium (Cotton et al., 2004). As a carbohydrate rich food potato contain greater amount

of dry matter and protein compared to cereals (Bamberg and Del Rio, 2005). In the human diet, potatoes are also a

valuable source of health-promoting antioxidants. In addition, fat percentage in potato is very low. It is a good source

of vitamins B1, B3 and B6, minerals and dietary antioxidants, which may play a great role in preventing diseases

related to ageing. Carbohydrates and anti-diabetic factors such as antioxidants of potatoes contribute in prevention and

management of diabetes (Camire et al., 2009). Potatoes contain a significant amount of fiber, which helps in lowering

the total amount of cholesterol in the blood, thereby decreasing the risk of heart disease of human. So, potato plays a

significant role for regular functioning of human health. Potato is now growing around 140 countries in the world

(Haase, 2007). Unfortunately, the crop suffers from different fungal, bacterial, viral, viroid and parasitic nematode

diseases under unfavourable conditions and causes huge losses its production. These invigorate the development of

effective processes for sustainable assembly to control potato diseases and pests.

Root-knot nematodes have drawn attention in importance in different parts of the world and cause severe yield damage

(Karssen et al., 2013; Onkendi et al., 2014). Root-knot nematodes (RKNs) Meloidogyne spp. especially M.

javanica, M. incognita, M. arenaria, M. chitwoodi and M. hapla the most harmful agricultural pests among the

nematode species, are showing aggressive effects and heavy yield reduction (Anwar et al., 2007). Both tuber quality

and quantity can be reduced by 12.2% as a result of nematode attack (Hadwan and Khara, 1992). RKNs (Meloidogyne

spp.) impose an important barrier for potato production through their destructive activity. On a global scale, the

distribution of nematode species varies greatly. There are more than 50 different species of RKNs (Jones et al., 2013).

As potato being highly susceptible to M. javanica, M. incognita, M. arenaria, M. chitwoodi and M. hapla, which all

adversely affect the efficiency of roots and yield.

Nematodes are multicellular organism mainly found in the group Ecdysozoa, or organisms that are able to change their

cuticle. The majority of them are said to be free-living and feed mainly on bacteria, fungi, protozoa and other

nematodes, only a minority feeds on parasites, animals and plants (Bélair, 2005). They are motile organisms, with most

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Mandal and Hossain (2017)

162

of them being able to move through the soil, covering an estimated distance of a single meter in their entire lifetime. In

addition, they can also move through nematode infected plants, seeds, plant tissue and bulb. Management of RKNs is

complex due to their wide host range and high reproduction rate.

Various methods are using for controlling RKNs, such as chemical control, organic amendments, resistant varieties,

soil solarization and biological control (Randhawa et al., 2001). But, current research is focusing on biological control

for managing plant diseases due to the adverse effect of chemical pesticides on human health and their persistence in

the surrounding environment. Besides, cultural practices can be employed by subsistence and small holder farmers in

order to manage nematodes, with many of these practices being a sustainable option for long term use. In addition,

cultural controls ensure unfavorable environmental conditions for pest growth, survival, reproduction and dispersal. In

a wide range of environmental conditions, the natural density of nematode population can be maintained by agronomic

strategies. Besides, it will be an ecofriendly option for plant protection with great potential for promoting sustainable

agriculture. In review study focused on agronomic approaches can be used for controlling RKNs in potato. This will

provide applied knowledge on RKNs of potato and diversified ecofriendly control measures.

Materials and Methods

To encapsulate and analyze existing scientific knowledge on a specific topic, a literature review is very helpful means.

The reviewed topic is too broad to be answered by a single or few experiments and since many studies have been made

by different scientists and are available in scientific journals, a literature review seems to be the most suitable method

for this work. This review study was conducted by searching Google, Google scholar, Scopus.nl and information

obtained from Wageningen University library by using the keywords ‘cultural control’, ‘Nonchemical control’, ‘root-

knot nematodes (Meloidogyne spp.)’ and ‘potato’. Many different combinations were tried to avoid missing useful

information.

Life cycle of RKN Meloidogyne spp.

The life cycle length of Meloidogyne spp. depends on temperature and varies from 4-6 weeks (summer) to 10-15 weeks

(winter) (Ploeg, 1999; Curto et al., 2005). The optimal temperature for development and reproduction ranges from 21

to 27°C. As compared to other plant parasitic nematodes, RKNs have a massive capacity in reproduction, as an

individual female is able to produce 200-1500 eggs in a gelatinous matrix known as egg-mass (Luc et al., 2005). The

first juvenile stage (J1) appears from the egg and, after the first moult, gives rise to the second juvenile stage (J2).

Second stage larvae move through the soil, to enter or parasitize a new root. The nematode then ingests the cytoplasmic

content of giant cells, acting as a metabolic drain that diverts nutrients from the plant to the nematode. Roots start to

swell within a few days after entering nematode into the root. Second juvenile stage (J2) then develops through forming

a flask shaped body, which will gradually increase in width and body size of a female larva. In the developmental step

juvenile 3, the larvae enter the moulting phase (Fig. 1). During this period the stylet and the median oesophageal bulb

disappears. In the final stage (J4) the life cycle of the adult nematode completed by the formation of a uterus and vagina

(Bengtsson, 2015).

Symptoms of infected potato plants

The most diagnostic RKN damage occurs in below ground of the plant, numerous symptoms can also be observed in

the above ground part (Fig. 2) of the plant. An infected potato plant by RKN, shows slow or stunted growth, yellowing

of leaves and plant wilting (even under adequate soil moisture) and collapse of individual plant. Highly infected plants

produce only a small amount of root, which leads to death. Nematodes induce the expansion of root cells, which leads

to the occurrence swelling or gall formation in the roots of infected plants. The galls vary in size from slight thickening

to lumps 5-10 cm across. Gall formation may also occur in stem and leaf. Compared to other species of Meloidogyne,

M. hapla produces small gall in the root. Gall in the root blocks the vascular system of root and impede movement of

water and nutrients through the roots. Potato plants that affected by RKNs are more susceptible to virus and bacteria

(Noling, 1997). Symptoms are more severe when plants infected with fungal pathogens such as Verticillium and

Rhizoctonia (Taylor, 1990). RKN affected potato tubers become unusual in shape which are less preferred by the

consumer.

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Agronomic strategies against root-knot nematode

163

Agronomic practices for controlling RKN

Root-knot nematode causes detrimental effects and severe losses to the crops. Preventing root-knot nematode is one of

the very challenging due to scarcity of effective control measures. Different agronomic management practices are

considered as the most efficient ways for root-knot nematode (Meloidogyne spp.) control (Ferraz and Mendes,1992).

Fig. 1. Life cycle of root-knot nematode (Mitkowski and Abawi, 2003)

Fig. 2. Above ground (a) and below ground (b) symptoms of root-knot affected potato plant

(Vovlas et al., 2005)

(a) (b)

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Mandal and Hossain (2017)

164

Crop rotation and fallowing

Crop rotation (alternating of different crops and/or fallow) is used to manage M. javanica, M. incognita, and M.

hapla successfully in many annual crops, including potatoes (Janati et al., 2018). As a sustainable strategy in the

management of plant parasitic nematodes crop rotation has enormous potential (Ball-Coelho et al., 2003). In a rotation

program, the incorporation of crops that have antagonistic effect on nematodes reduce the initial nematode population

thus help the subsequent crop to establish properly (Oka and Yermiyahu, 2002; Luc et al., 2005; Mweke et al., 2008).

However, Mitkowski and Abawi (2003) reported that nematode species which have narrow host range can be

effectively controlled by this method. Due to polyphagous nature of root-knot nematodes, there are relatively few non-

host plants available for controlling them through crop rotation, Crotalaria was recommended as an intercrop to control

Meloidogyne spp. on pineapple (Netscher and Taylor, 1979; Luc et al., 2005).

The basic principles behind the crop sequences for nematode control are to keep the nematode level below the

economic threshold level, to conserve beneficial organisms and to maintain long term fertility of soil (Kimpinski and

Thompson, 1990). Tagetes erecta lowered levels of nematodes when intercropped with a highly susceptible banana

crop (Wang et al., 2007). Rotation with grasses is very effective in minimizing populations of M. arenaria, M. hapla,

M. incognita and M. javanica (Netscher and Taylor, 1979). Rotations with a non-host crop for RKN, such as maize or

wheat, will minimize some Meloidogyne spp. damage. For example, barley can be used in rotations to reduce M. hapla

and chitwoodi infections (Belair and Parent, 1996). It is better to use crops in rotation which is resistant on most of the

nematode species instead of a single one. Crop rotation can be used as the primary means of management of M. hapla

on carrot (Gugino et al., 2006).

Leaving the soil totally bare for a certain period is known as fallowing. Fallowing ensures starving the nematode

population by preventing vegetation in the field. It is also essential to keep the crop area weed free to prevent

nematodes from surviving on alternate hosts (Kutywayo and Been, 2006; Thomas and Murray, 2005). One year of

fallowing will lower the nematode population to a level that favors in growing annual crops successfully. During the

fallowing the soil should be moist, because that allows hatching of nematode eggs and nematodes die due to starvation

of food. Theoretically this approach should be more effective in hot and arid climates as compared to cooler regions

because there has high rainfall. The best result is obtained when fallowing is maintained during hot and dry weather

(Johnson and Campbell, 1980). Soil in the fallowing can be turn around every two weeks interval in order to expose

nematodes to the sun.

Cover crop and trap crop

Cover crop is a crop that is not harvested and is grown in the season between two harvestable crops. This practice

allows less RKN damage of potato production in the next season. The success of this technique depends on the type of

crops using as cover crop. With the presence of non-host cover crops, the nematodes cannot migrate to another field

because its short distance migration ability (Gill and McSorley, 2011). The wide host range of RKNs (Meloidogyne

spp.) limits the choices of potential cover crops that may be antagonistic or non-hosts for these pests. Some cover crops

such as rye and crimson clover have shown nematode-suppressive characteristics equivalent to aldicarb (a synthetic

chemical pesticide). Other plants that suppress nematodes through chemical properties of residues, especially when

grown as cover crops and tilled into the soil, includes castor beans, chrysanthemums, and sesame because they contain

allelochemicals that works as nematode-antagonistic compound. Sunnhemp is very popular for controlling RKN

because it acts as a poor host for nematode. Besides, sunnhemp favors to grow nematode trapping fungi in the

rhizosphere, thus it helps to control RKN (Wang et al., 2001). Marigold has a suppressive impact on nematodes has

been documented for over 50 years. In both greenhouse and field growing conditions marigold (Tagetes spp.)

successfully reduced Meloidogyne spp. (Ploeg, 1999; Ijani et al., 2000; Wesemael and Moens, 2008). The reasons

behind that are marigold exhibit allelopathy behaviors producing different compounds and essential oils, enhance the

activity of endophytic bacteria, and encouragement of nematode-antagonistic organisms (Zygadlo et al., 1994; Kimenju

et al., 2004; Sturz and Kimpinski, 2004). Consequently, nematodes do not damage or penetrate the rhizodermis of

marigold, it is more effective as a cover crop than as a soil amendment. Marigold plants produce a number of

potentially bioactive compounds like α-terthienyl combined with triplet oxygen that allows limited nematicidal activity

when added into the soil (Wang et al., 2007). Moreover, cover crops can maintain or increase organic matter content of

soil when their residues are returned to the soil. There are many crops that can be used as trap crops for controlling

RKNs.

Trap cropping is a very crucial management technique for controlling nematode since the late 1800s. Matured female

nematode enters the root of host and unable to leave the plant root. By this trap cropping nematode within the root all

the hosts are destroyed by tillage practices and all the nematodes trapped within the soil (Westerdahl, 2007). Trap crops

are grown to keep away nematodes from the cash crop. However, many plant species are good trap crops with

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Agronomic strategies against root-knot nematode

165

potentially undesirable characteristics, such as toxicity to domestic animals or weedy traits. In the soil trap crops roots

produce chemicals that enhance the hatching of the nematode egg. After hatching they move to the trap crop’s roots

and start to feed. However, the trap crops such as sugar beet and cereals are grown as it supplies insufficient nutrition to

mature RKN, reproductive cycle is hampered. In addition, decomposed trap crops stimulate the establishment of

saprophytic fungi, which parasitize the eggs and nematodes. Rucola (Eruca sativa) is effective against M. hapla with its

marketability for controlling nematode (Melakeberhan et al., 2006).

Flooding and weed control

Flooding technique for controlling RKN in potatoes is very effective where water availability and water pumping

equipment and dikeare available. Seven to nine months flooding can control RKN by reducing the amount of oxygen

availability for respiration and increasing concentrations of toxic compounds such as organic acids, methane, and

hydrogen sulphide. Compared to prolonged flooding, alternative flooding and drying is more effective for controlling

nematode (Noling and Becker, 1994). It was observed that flooding techniques works best in high temperature

condition of both soil and air. Flooding can reduce the population of M. incognita, but air temperature regulates the

period of optimum flooding (Rhoades, 1982). It should take into consideration that poorly managed flooding can make

matters worse, as water is also an excellent means of nematode dispersal. Weeds can act as hosts reservoir of nematode

species including RKNs. Standing weed in crop field not only competes for light, space, water, and nutrients but also

can serve as reservoirs of pests, including plant-parasitic nematodes (PPNs) (Davis and Webster, 2005). Weed control

is useful in helping to reduce Meloidogyne populations. Control of Amaranthus spp. was found very effective for

limiting RKN, because this species is a good host for RKNs (Noling and Gilreath, 2002). Other direct effects of weeds

include protection of certain nematodes from pesticides or adverse environmental condition. It would appear that

unmanaged weed growth serves to prolong a nematode problem from one cropping season to the next, and quite

possibly, reduce the need for broad spectrum soil fumigants application for nematode control.

Soil solarization and organic amendments

Soil solarization can effectively suppresses RKN (Tisserat, 2006). But its prerequisites are sunny and warm weather,

because heat is trapped from the sun by clear plastic over ploughed land. Covering soil with plastic film for at least 2

weeks during mid-summer maximizes soil heating effects and this killed the egg of the RNK, thus reducing their

population. The most effective application of soil solarization takes place in heavier (loamy to clay soils) rather than

sandy soils (Noling, 1999). Wet soil with good water holding capacity capture heat better than dry soil and makes soil

organisms more susceptible to heat. It is important to use film with the appropriate physical properties, to enhance soil

thermal conductivity prior to solarization, with irrigation and tillage to avoid compaction (Scopa et al., 2008). To apply

this technique requires bright sunshine for long duration and applicable where exist plenty of solar energy for long

periods of time. However, the benefit of soil solarization effect is only bound to the top layer of the soils (Bello et al.,

2002).

Research studies have evaluated that RKNs could be controlled by adding organic matter to soil such as animal manure,

green manure from cover crops or crop residues. Cake from castor bean waste has proved effective in reducing

populations of Meloidogyne spp. due to combined effect of ricin and ricinine as well as nutritional compost properties

(Lopes et al., 2009). Organic soil amendments from Tagetes minuta, Ricinus communis, and Datura stamonium

increase crop yields significantly in nematode infested field through their nematicidal properties (Oduor-Owino and

Waudo, 1996; Kimenju et al., 2004; Akinyemi et al., 2009). Addition of some other manure such as wastes from wood

industries retain nematicidal properties for controlling RKN (Matveeva et al., 2010). In addition, Neam (Azadirachta

indica) can be used as a green manure by incorporating the leaf into the soil and as an extract for biological control.

The rate of organic amendment should be higher for effective control of nematode population (Van der Putten et al.,

2006). The organic amendments in soil improve physical and chemical properties of soil, and improve water holding

capacity that attracts other beneficial soil microorganism that inhibit RKN growth and spread in the field. Neem cake

has the potential to effectively control nematodes of Meloidogyne species on vegetable crops and cardamom (Ahmed

and Koppel, 1987; Bertrand and Lizot, 2000).

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166

Sanitation and tillage

Sanitation practices are important technique to minimize RKNs spread between planting sites or into new production

areas. It may include removal or destruction of infected plant materials, inspection and use of certified nematode free

potato propagating materials, cleaning of farm implements and heat treatment of potentially infected potato tuber.

RKNs can survive very short period in the residues of infected plant, until they consume their own reserves because

they are obligate parasite in plants (Ornat and Sorribas, 2008). Uprooting plants after every harvest as well as root

exposing to solar radiation is very effective to kill nematodes in root tissues and this practice is very common for

controlling RKN in the tropical areas (Bridge, 1996). This precaution could reduce Meloidogyne spp. populations about

90% instead of leaving residual roots in the soil. Tillage inverts and mixes soil and exposes deeper soil layers to the

sunlight. As nematodes depend on soil moisture; tillage is very effective for controlling root RKNs through desiccation.

On the other hand, minimum tillage allows to gather plant residues, which suppress nematodes through increase of

organic matter content in the soil (Widmer et al., 2002).

Plant resistance

Plant resistance is the most eco-friendliest and cheapest method for controlling RKN (Bridge, 1996; Dufour et al.,

2003). Plants can be a good source of naturally occurring compounds having nematicidal properties such as alkaloids,

fatty acids, thienyls, isothiocyanates, phenols, polyacetylens, glucosinolates and sesquiterpenes (Chitwood, 2002). For

example, few studies found existance of some metabolites (such as lantanosides and lantanone) in Lantana camara

against Meloidogyne incognita (Begum et al., 2000; Qamar et al., 2005). Most of the annual vegetable crops have

some resistant varieties against RKN. But very few species of potatoes are resistant against RKN. A number of

researches are on-going to develop crops with resistance genes against Meloidogyne spp. (Norshie et al., 2011). It is

expected that an increase in transgenic crops with resistance to Meloidogyne spp. can be anticipated in future. Firstly,

the resistance gene (Rmc-1) in some wild potato species positioned on chromosome 11 was found to confer resistance

against M. chitwoodi and other Meloidogyne spp. such as M. hapla and M. fallax (Brown et al., 2006). Several central

American Solanum spp. is resistant against to both M. Chitwoodi and M. fallax. Potato varieties Mc Cramick and

Golden were found as resistant variety and moderate resistance were recorded in varieties Oronek, ORA and Suzanna.

It is observed that few potato lines are also capable to show resistance against M. chitwoodi (Norshie et al., 2011).

Different crops such as beans (Phaseolus vulgaris), soya beans (Glycine max), peas (Pisum sativum L.), pepper

(Capsicum spp.), tomatoes (Lycopersicon esculatum), sweet potatoes (Ipomea batata) and cowpeas (Vigna

unguiculata) have the resistance to different species and races of Meloidogyne (Sasser and Kirby, 1979; Netscher and

Sikora, 1990). It is reported that potato species Solanum sparsipilum is a source of resistance against RKN. This

resistancy of S. sparsipilum germplasm acts against M. incognita, M. javanica and M. arenaria.

Botanical nematicides

Many plants, plant parts and their active components have been evaluated for their nematicidal properties worldwide

and these were found effective in reducing the nematode infection on plants (Saxena and Singh, 2001; Saravanapriya

and Sivakumar, 2005; Taniwiryono et al., 2009; Rehman et al., 2012). The presence of phytochemicals in the plant

parts acts as nematicides for sustainable management of nematode (Chitwood, 2002; Trifonova and Atanasov, 2009;

Du et al., 2011; Ojo and Umar, 2013). Effectiveness of certain plant extracts was also observed significant to kill

second stage juvenile of root-knot nematode (Saxena and Singh, 2001). Plants like Mucuna pruriens and Tithonia

diversifolia having antagonistic properties to nematodes considered to produce anti-helminthic compounds with

different modes of action (Quénéhervé et al., 1998; Pandey et al., 2003; Akinyemi et al., 2009). Marigolds (Tagetes

species) which exude polythienyls have been proven to be nematicidal for controlling nematodes (Wang et al., 2007).

Leaf extracts of Moringa oleifera, Ocimum gratissium and Azadirachta indica are very effective for controlling M.

incognita through 40-70% reducing the egg hatching due to its nematicidal properties, such as azadirachtin from A.

Indica (Claudius-Cole et al., 2010). Another study has proved that both leaf and seed extracts of A. indica containing

azadirachtin increase the juvenile mortality of potato RKN (Upadhyay et al., 2003). Tuber extract of Dioscorea

floribunda having anthelmintic properties prevented M. incognita egg hatching (Nath and Mukherjee, 2000). Fresh

leaves extract of Calotropis procera have good anthelmintic activity and showed nematicidal activity on eggs hatching

and juveniles of Meloidogyne incognita. Aromatic and culinary herbs contain the nematicidal compounds carvacrol and

thymol. Bawa et al. (2014) reported that extract of A. indica leaf, Capsicum annuum fruit and Parkia biglobosa seed

can control M. incognita population and it is due to some properties of these products such as neem (azadirachtin),

pepper (sincocin) and locust bean fruit husk (titerpine). In addition, Lantana camara contains 11-oxo triterpenic acid

(lantanolic acid, lantoic acid, pomolic acid and ursolic acid) and this compound was effective against M. incognita with

mortality rate 85 to 90% (Srivastava et al., 2006). The suppressing ability of P. lilacinus and Trichoderma viride and

botanicals (neem and castor cakes) against the root-knot nematode, M. incognita, in tomatoes (Rangaswamy et al.,

2000). Specific studies on RKN control of potato are very rare. There are many studies (Table 1) that found

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effectiveness of different plant extracts to control RKN. It can be a good option to evaluate their performance against

RKN in potato.

Nematode infested soil can be treated properly with several biological control agents (Sikora, 1992; Hallmann et al.,

1998). Biological control agents and organic amendments revealed successful ways to control nematode. Integration of

neem with fungal biocontrol agents Paecilomyces lilacinus and Verticillium lecanii for the integrated management of

RKN on tomato (Rao et al., 1998). P. lilacinus and Trichoderma viridealone or in combination with mustard oil cake

and carbofuran group nematicide promoted plant growth, reduced number of galls plant-1

, egg-masses root-1

against the

root-knot nematode, M. incognita infecting tomato (Goswami et al., 2006). Integration of Pasteuri apenetrans and

nematicides (carbofuran) and insecticide (phorate, a systemic insecticide) in the management of the root-knot

nematode, M. incognita, as an integrated approach, resulted in higher rate of parasitization also recorded higher plant

growth (Kumari and Sivakumar, 2005; Kumar and Khanna, 2006; Ferreira et al., 2010).

Table 1. Plant extract to control RKN

Plant Plants parts Effective against References

Areca catechu; Carica papaya;

C.gigantea

Seed and

Latex Meloidogyne incognita

(Saravanapriya and

Sivakumar, 2004)

Neem (Azadirachta indica) Leaf Meloidogyne incognita (Ntalli et al., 2009)

Nicotiana tabacum; Syzygium

aromaticum;Piper betle;

Acorus calamus

leaves Meloidogyne incognita (Taniwiryono et al., 2009)

Wild Sege

Lantana camara Leaf

Meloidogyne incognita

Meloidogyne chitwoodi (Ahmad et al., 2010)

Chrysanthemum coronarium,

Azadirachta indica,

Nerium oleander

Leaf and seed Meloidogyne javanica

(Ahmad et al., 2010)

Castor bean Seed Meloidogyne spp. (Adomako and Kwoseh, 2013)

Garlic (Allium sativum) cloves

and castor bean (Ricinus

communis)

Seed Meloidogyne incognita

(El-Nagdi and Youssef, 2013)

Rauvolfia tetraphylla Root, leaf and

fruit extract Meloidogyne incognita (Mandal and Nandi, 2013)

Spanish cherry

Mimusops elengi L Leaf Meloidogyne incognita (Ahmad et al., 2010)

Water Hyacinth Leaves Meloidogyne incognita (Umar and Mohammed, 2013)

Future thrust of research

This review is shouting about strategies of alternative and eco-friendly control process of RKN. Most of these

alternative techniques can control RKN in potato without affecting natural properties of soil as well as surrounding

environments. Although different types of experiments were conducted based on agronomic strategies, but its

suitability of application is very less in the field condition. Besides, a single agronomic practice may not work well to

suppress RKN infestation. The agronomic strategies that are discussed in this review article can be applied in different

steps of potato cultivation to minimize it at safe level (Fig. 3).

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Fig. 3. Agronomic strategies to control RKN in different steps of potato cultivation

Besides, more advance studies on different agronomic strategies combining with modern farming context are crucial

for controlling RKN in potato. This holistic approach is required to redesign the cropping systems according to the

goals of optimum production, pest management, and other services, like environmental conservation by reducing

chemical nematicides application. Besides, interest on this method by the farmer, chemical control process may

disappear, and synergistic combinations are discovered that increase their efficiency. The future of agronomic strategies

control of RKN will, of course, be closely related to integration into management programmes that involve reducing

pre-plant densities of Meloidogyne with conventional management tools and the use of microbial control agents that

more effectively target the susceptible stages in the life cycle of the RKN.

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