CC BY-NC-ND
0©2013 Parasitological Institute of SAS, Kosice DOI 10.2478/s11687-013-0101-y
VERS IT/C
Helminthologia, 50, 1: 3 - 14, 2013
Review Article
Organic amendments of soil as useful tools of plant parasitic nematodes control
M. RENCO
Institute of Parasitology, Department of Environmental and Plant Parasitology, Slovak Academy of Sciences, Hlinkova 3, 040 01 Kosice, Slovak Republic, E-mail: renco@saske.sk
Summary
Use of organic soil amendments is a traditional agricultural practice for improving physical and chemical soil properties, soil structure, temperature and humidity conditions as well as nutrients content which are needful for plants growth. Application of organic materials to soil can cause a change in soil microflora and microfauna including soil nematodes. Nematodes, are the most ample and varied group of soil fauna. They are ever-present habitants of all soil types with high population densities. The changes in soil nematodofauna can results in an increase in the number of beneficial nematodes such as bacterial or fungal feeders and decrease and/or suppression in the occurrence of economically important plant parasitic nematodes. A variety of organic amendments, such as animal and green manure, undecomposed (raw) or decomposed materials (compost) are used for this purpose. Generally, plant parasitic nematodes have been controlled mainly by chemical soil fumigants and nematicides, agricultural practices or resistant cultivars. However, organic amendments can provide an environmentally friendly alternative to the use of chemical nematicides, which are often expensive, of limited availability in many developing countries and above all environmentally hazardous.
Introduction
The enhancement consideration to the environment protection and to human and animal health, according to the recent European Legislations (Reg. CE 396/2005; 1095/2007; 33 and 299/2008 and 1107/2009) which have deeply delimited and revised the use of pesticides on agricultural crops, is stimulating investigation to find new alternative control strategies that are environmentally sound and economically convenient at the same time. Therefore, research on small environmental impact alternatives to chemicals has received a strong impulse and considered a wide range of options including agronomical
and physical methods (green manures, crop rotations, soil amendments, the use of resistant cultivars and arbuscular mycorrhyzal fungi, soil solarization and steam), the use of natural products from plants and biological control agents (Sasanelli, 1992; Vannacci & Gullino, 2000; Atungwu, 2005; Castillo et al, 2006; Riga et al, 2007; Sasanelli et al., 2009; Riga, 2011).
Among these alternative control strategies the soil organic amendments is particularly interesting because of their low cost and the more general positive agronomical effect on plant growth and on physical, chemical and biological properties of the soils (Davey, 1996). Moreover, it can suppose the management of large amounts of wastes generated by urban settlements and agro-industrial processes, after their transformation by composting process, and in addition it can improve plant resistance and plant protection by stimulating root development by recycling plant nutritive elements (De Bertoldi, 2008). Many organic material wastes represent an important resource of nitrogen, phosphorous, calcium and other elements as zinc, copper, magnesium essential to plant growth (Tester, 1990). There are sufficient data to indicate that organic materials have positive effect on soil structure, improve plant growth and yields and reduce disease impact caused by a wide range of plant pests including bacteria, soil-borne pathogens and phytoparasitic nematodes species (Akhtar & Mahmood, 1996; D'Addabbo & Sasanelli 1996a, 1997, 1998; Abawi & Widmer, 2000; Renco et al. 2007, 2009, 2010; Atungwu, 2012). In particular, the suppressive effect of soil amendments, with a wide range of composted waste materials, on plant parasitic nematodes was largely and frequently docu-mented, although an inconsistent nematode control or variable effects were also described in literature (Szczech et al., 1993; McSorley & Gallagher, 1995; Akhtar & Malik, 2000).
Therefore the nematicidal effect of compost amendments is sorely predictable because of depending on the using of
raw materials, the type of composting process, the maturity of the final product incorporated into the soil, nematode species present and season or rate of application (Rodriguez-Kabana et al., 1987; Rivera & Aballay, 2008; Renco et al. 2007, 2010). Many mechanisms can be involved in this suppressive effect such as decomposition of the compost into the soil and ammonia production, stimulation of soil microbial biomass and release of biocidal substances which have nematicide activity (Oka & Yermi-yahu, 2002). The nematicidal activity, therefore, should be specifically assessed for different composts. This paper discusses about the use of different organic amendment on soil quality, plant growth as well as on control the most important plant parasitic nematodes and factors caused its response.
The organic amendments effect on soil quality and plant growth
It is necessary to mention that organic fertilizer has a great impact on soil fertility which is essential for good plant growth because of fertile soil providing nutrients necessary to grown plants. If absence of nutrients in the soil is obvious, we can add those to soil by the two types of soil amendment - organic (made from natural products) and inorganic (man made chemicals). However, organic manure compared with inorganic has many advantages because organic matter improve the physical properties of
soil, including water retention, water infiltration, permeability and aeration as well as increase soil fertility and provide a better environment for roots (Bulluck et al., 2002b; Mandal et al., 2007). The common organic amendments are cow, chicken and horse manure, compost or green manure (Riga & Collins, 2004). These increase the organic constituents of the soil thus providing favourable environment for bacteria and earth worms that enrich the soil. However, the content of nutrients in cow manure are generally lower than in chicken and horse manure. The inorganic amendments are man made and they include chemicals that are used for making the soil fertile. Though they are used for good output they will reduce the natural nutrients of the soil in the long run. On the other hand, organic manures often provide more than one of the many nutrients needed by plants but inorganic manures provide only one of the many nutrients needed by plants.
The soil amendments depend greatly on the soil requirements, types of crops that grow in the soil, texture of the soil and its salinity but from requirements of growers for soil improvement. Almost any kind of organic matter may be used as manure, but some kinds are better than others. Organic manures vary widely in the amount of plant nutrients that they contain. Some are more concentrated than others. That the various organic amendments improve plant growth and yield was confirmed in many studies (Table 1).
Table 1. Organic amendment possitive effect on plant growth and production
Amendment
Parameter
References
Poultry manure Chilli
Oilcakes of neem, castor, composted manure Pigeonpea Domestic garbage, dead vegetation, vegetable Tomato and fruit processing wastes, sugarcane trash, corn shucks, groundnut hay, groundnut hulls, paddy straw and husk, pressmud, spent tea, tobacco rettes
Oil cakes of cotton, flax, olive, sesame and Tomato soybean
Cattle manure, green manure , pigeon-dung, Tomato sewage sludge, horse manure and cotton root wastes
Cattle dung, bean and wheat straw compost Tomato
Vermicompost Chinesse
cabbage
Onion bulb envelope, dry leaves of sugar Banana
beet, fleabane and jojoba, filter cake or mud as sugar cane industrial residue and nile fertile mineral bio-fertilisers
Compost from poato waste, sawdust, beef Potato
manure; beef manure
Fermented pig manure commercially name Broccoli Difert
Fermented pig manure commercially name Sunflower Difert
Vermicomposts from food wastes, paper Peper
wastes and cattle manure
Vermicomposts from food wastes, paper Tomato
wastes and cattle manure
Vermicomposts from food wastes, paper Strawberies wastes and cattle manure
Yield T
Fresh weight, dry weight and height T Growth of plants T
Growth of plants T Growth of plants T
Yield T Growth of plants T
Yield T
Yield T Yield T Yield T
Yield, growth of plants, leaf areas, plant shoot biomass T
Yield T
Yield, leaf areas, numbers of strawberry suckers, numbers of flowers, shoot weights T
Khan et al, 2001 Akhtar and Mahmood, 1996 Akhtar, 1993
Radwan et al, 2009 Maareg et al., 2000
Korayem, 2003 Wang et al., 2010
Youssef and El-Nagdi, 2010
Kimpinski et al., 2003 Kovacik et al., 2008 Kovacik et al., 2010 Arancon et al., 2005 Arancon et al., 2003a Arancon et al., 2003a
The organic amendments effect on plant parasitic nema-todes
The control of plant parasitic nematodes is more difficult in comparison to other pests because they usually live in the soil and attacks the underground parts (roots) of the plant. A lot of nematode control strategies such as nemati-cides, resistant varieties, crop-rotation, non-hosts, antagonistic crops, biological control (predators and parasites of nematodes) have been used successfully but each method has some limitation to implementation (Akhtar, 1997). Probably for those reason, a wide variety of organic matters (incorporated to soil) have been tested as potential and alternative control of plant parasitic nematodes. The most studies worldwide are focused on root-knot nematodes of the genus Meloidogyne because of approximately 2000 plants are known to be hosts of these nema-todes from grass to trees where they cause the galls on roots. They are occurring mainly in temperate areas with short winters, especially in sandy soils. Because of crop rotation as a control tactic of these nematodes is rather difficult due to a wide range of their hosts, the alternative organic control is main investigated method for their regulation (Table 2). Sasser and Carter (1985) noted that Meloidogyne nematodes account for approximately 5 % of global yield loss.
The genus Meloidogyne includes more than 60 species, however four Meloidogyne species (M. javanica, M. arenaria, M. incognita, M. hapla) are considered as major plant pests' worldwide (Eisenback & Triantaphyllou, 1991). D'Addabbo (1995) found 160 literature sources on the effect of organic amendments on nematodes of the genus Meloidogyne under different host plants. For example, on tomato roots, the reduction of M. incognita population by the application of chicken manure was observed by D'Addabbo et al. (2000); by the application of water hyacinth compost, mustard straw, rice husk and asparagus compost (Sharma et al., 1997); by olive pomace (D'Addabbo & Sasanelli, 1996a; D'Addabbo et al., 2011); by grape pomace (D'Addabbo & Sasanelli, 1998) or pepper crop residues (Buena et al., 2007). On the other hand, not all types of organic amendments were beneficial in the suppression of root-knot nematodes. For instance, Bulluck et al., (2002a) observed that Meloidogyne incognita populations were not affected by amendments with swine manure or composts water extract prepared from bean or wheat straw, poultry or fish wastes (Korayem, 2003). Vermicompost showed no inhibitory effect on the number of M. hapla galls on cabbage and tomato roots; incorporation of compost consisted of cull waste potatoes, sawdust and beef manure had no efficacy on M. hapla populations in potatoes (Kimpinski et al., 2003). No suppressive effect of organic amendment was observed on the population of rice root-knot nematode M. graminicola (Gergon et al., 2001).
Another most important worldwide are cyst nematodes. Therefore, their control is difficult because they are more resistant than endoparasitic, ectoparasitic and sedentary ectoparasitic nematodes, because of the presence of the layer of dead cuticle of females which serves to protect the
eggs and second-stage juveniles that are retained within (Zunke & Eisenback, 1998). In every case, several studies were focused on this group of plant parasitic nematodes with positive results (Table 2). For instance, Van der Laan (1956) found that development of Globodera rostochiensis on potato roots was delayed by application of organic material in comparison to untreated control. Similarly to it, Renco et al. (2007, 2011) observed reduced reproduction of females of G. rostochiensis patotype Ro1 and G. pallida patotypes Pa2 and Pa3 by use of nine type composts applied at four doses, when compared to the untreated control. Also, steer and chicken manures reduced the numbers of cyst G. pallida and resulted in increased yields of potatoes (Gonzalez & Canto-Sanenz, 1993). Contrary, vermicompost don't reduce Heterodera schachtii in study of Szczech et al. (1993). Kimpinsiki et al. (2003) observed an increase in the number of Heterodera trifolii juvenile in barley plots however this species parasitized on red clover crop. Similar increase in egg hatching of the species was observed by Kunelius et al. (1988). The authors attributed this support of hatching to increased aeration in conventionally tilled soil compared to non-tilled soil. Several studies were aimed at the control (reduction) of several other endo and ectoparasitic nematode species. The nematode suppression after and organic amendments was recorded, for example at Pratylenchus species (Khan et al., 1986; LaMondia et al., 1999; Abawi & Widmer, 2000; Kimpinski et al., 2003); Helicotylenchus species (Subba Rao et al., 1996; Khan & Shaukat, 1998) and many other (see Table 2). Contrary to that, no suppression of Pratylenchus sp. after the application of swine manure was observed by Bulluck et al. (2002a); sewage sludge (Weiss & Larink, 1991); yard-waste compost on vegetable crops (McSorley & Gallaher, 1995). Also, Xiphinema spp., Criconemella spp. and Paratrichodorus minor was unaffected by application of yard-waste compost on vegetable crops (McSorley & Gallaher, 1995). Soil treatments by solid waste compost did not affect the population densities of Criconemoides spp. and Paratrichodurus minor as well, however the population densities M. incognita increased in compost-amended plots (McSorley et al., 1997). Sewage sludge treatment produced no suppressive effect on Helicotylenchus dihystera (Sharma et al., 2000).
The organic amendments effect on nematode community changes
In addition to studies examining the impact of application of different organic materials on particular plant parasitic nematodes, during the last decades there has been an increasing interest deal with impacts of organic amendments on soil nematode community changes, in general. For example, Akhtar and Mahmood (1996) found a significant reduction of plant parasite and increase of predatory and free-living nematodes after application of all tested materials after an application of different rates of oilcakes of neem (Azadirachta indica) and castor (Ricinus commnunis), composted manure and urea, as well as using of composted manure combined with Tagetes erecta
Table 2. Documented suppressions of plant parasitic nematodes by different types of organic amendment
Nematode species Plant
Organic material
References
Meloidogyne incognita
Meloidogyne incognita
Meloidogyne javanica
Meloidogyne. hapla
Meloidogyne. arenaria
Meloidogyne sp.
Globodera. rostochiensis
Globodera pallida
Heterodera schachtii
Heterodera carotae Heterodera oryzicola
Heterodera avenae Heterodera sp.
Tomato Chicken manure
Chicken manure
Water hyacint compost, mustard straw,
rice husk, asparagus compost
Composted dry cork
Olive pomace
Olive pomace
Grape pomace
Olive pomace, straw, urea
Olive residues
Grape pomace
Multi varied compost
Municipal green wastes, sewage sludge, spent mushroom substrate
Sewage sludge
Peper crop residues
Oil cakes of cotton, flax, olive, sesame and soybean Cotton Chicken manure
Banana Destillery sludge, vermicompost, neem cakee,
poultry manure
Onion bulb envelope, dry leaves of sugar beet, fleabane and jojoba, filter cake or mud as sugar cane industrial residue and nile fertile mineral bio-fertilisers Pea Neem cake
Tomato Composted dry cork
Corn Neem, mustard, water hyacinth compost
Cacao Poultry litter
Soybean Neem leaf powder
Neem and Sunshine organic fertilizer Chili Crop-residues of marigold, mustard, sunflower
Tomato Domestic garbage, dead vegetation, vegetable and
fruit processing wastes, sugarcane trash, corn shucks, groundnut hay, groundnut hulls, paddy straw and husk, pressmud, spent tea, tobacco rettes Tomato Broiler litter, cottonseed meal, feather meal,
soybean oilcake
Cattle manure, grape marc compost Dry cork, dry-grape marck
Sheep manure, cattle manure, horse manure, pigeon-dung , chicken manure, sewage sludge, green manure , cotton root wastes, sawdust and humic acid Lettuce coffee pulp compost
Groundnut Farmyard and poultry manure Peat Compost, cocoa bean peels
Tomato Compost tea, vermicompost tea
Lettuce Green manure by Sudan grass
Tomato Oil cakes, chicken litter
Chicken litter Chicken litter Cowpea Composted manure
Chili Poultry manure, pigeon manure, sawdust
Potato Organic material
Composts of different origin Freshly-crushed conifer bark Potato Steer and chicken manures
Sugar beet Compost
GFT compost Carrot Exhausted olive pomace
Rise Distillery sludge, vermicompost, neem cake, poultry
manure
Cereals Sewage sludge, lime, dehydrated pig slurry
Sewage sludge Cereals Municipal green compost
D'Addabbo et al, 2000, 2003
Lopez-Perez et al., 2005 Sharma et al., 1997
Nico et al, 2004
D'Addabbo and Sasanelli, 1996a D'Addabbo et al., 1997, 2000, 2003, 2011 D'Addabbo and Sasanelli, 1998 D'Addabbo and Sasanelli, 1997 D'Addabbo et al, 1997 D'Addabbo et al, 2000 D'Addabbo et al, 2006 D'Addabbo et al, 2011
Castagnone-Sereno and Kermarrec, 1991 Buena et al, 2007 Radwan et al, 2009
Riegel et al., 1996; Riegel and Noe, 2000 Sundararaju et al., 2002 Youssef and El-Nagdi, 2010
Pandey and Singh, 1990 Nico et al, 2004 Verma et al., 1997 Orisajo et al., 2008 Atungwu et al., 2009 Atungwu et al., 2011 Akhtar and Alam,1992 Akthar,1993
Oka et al., 2007
Oka and Yermiyahu, 2002 Nico et al, 2004 Maareg et al., 2000
Ribeiro et al., 1998 Jioshi and Patel, 1995 Coosemans,1982
Edwards et al., 2007
Viaene and Abavi, 1998
Mian and Rodriguez-Kabana, 1982
Kaplan et al., 1992
Kaplan and Noe, 1993
Olabiyi et al., 2007
Khan et al, 2001
Van der Laan, 1956
Renco et. al., 2007 Matveeva et al., 2002 Gonzalez and Canto-Sanenz, 1993 Schlang, 1993
Ryckeboer and Coosemans, 1996 D'Addabbo and Sasanelli, 1996b Sundararaju et al., 2002
Lopez-Robles et al., 2006 Weiss and Larink, 1991 Renco et al, 2009
Pratylenchus penetrans Bean Chicken manure Abawi and Widmer, 2000
Fallow plot Poultry litter Everts et al., 2006
Potato Poultry litter Conn and Lazarovits, 1999
Mushroom compost LaMondia et al., 1999
Maize Anaerobically digested slurry, cow, pig and Min et al., 2007
biowaste slurry
Pratylenchus coffeae Coffee Distillery sludge, vermicompost, neem cake, poultry Sundararaju et al., 2002
manure
Pratylenchus scribneri Poultry litter Khan et al., 1986
Pratylenchus zeae Rice Poultry litter Khan and Shaukat, 2000
Pratylenchus sp. Maize Garden waste compost Leroy et al., 2007
Maize Liquid dairy manure Timper et al., 2004
Paratylenchus sp. Cereals Municipal green compost Renco et al., 2009
Tylenchus mirus Poultry manure Khan et al., 1986
Psilenchus haki Mustard Poultry manure, pigeon manure, sawdust Hassan et al., 2008
Tylenchus sp. Maize Garden waste compost Leroy et al., 2007
Ditylenchus sp. Poultry manure Khan et al., 1986
Bitylenchus sp. Cereals Municipal green compost, penicillin residues Renco et al., 2009
compost
Helicotylenchus Banana Distillery sludge, vermicompost, neem cake, poultry Sundararaju et al., 2002
multicinctus manure
Helicotylenchus indicus Betelvine Sawdust, NPK, neem cake combination Subba Rao et al., 1996
Poultry manure Khan et al., 1986
Garlic Pigeon and poultry manure Khan and Shaukat, 1998
Chili Pigeon and poultry manure, sawdust Khan et al., 2001
Helicotylenchus sp. Cereals Municipal green compost, penicillin residues Renco et al., 2009
compost
Maize Poultry litter Summer et al., 2002
Cowpea Decomposed and composed manure Olabiyi et al., 2007
Tylenchorhynchus Chilly Pigeon and poultry manure, sawdust Khan et al., 2001
curvus Rotylenchulus sp. Cereals Municipal green compost, penicillin residues Renco et al., 2009
compost
Geocenamus sp. Cereals Municipal green compost, penicillin residues Renco et al., 2009
compost
Melinius brevidens Garlic Pigeon and poultry manure Khan and Shaukat, 1998
Hoplolaimus seinhorsti Garlic Pigeon and poultry manure Khan and Shaukat, 1998
Hoplolaimus indicus Rice Poultry mnaure Khan and Shaukat, 2000
Hoplolaimus columbues Cotton Poultry litter Koenning et al., 2003
Xiphinema index Grapevine Olive and grape pomace D'Addabbo et al., 1999
Xiphinema spp. Cowpea Decomposed and composed manure Olabiyi et al., 2007
Hemicycliophora sp. Vegetable Solid waste compost McSorley et al, 1997
Paratrichodorus sp. Maize Poultry litter Summer et al., 2002
Liquid dairy manure Timper et al., 2004
(Akhtar, 1998). Dmowska and Kozlowska (1988) recorded the increase in bacteriovorous nematodes after using a pig and cattle manure, especially Rhabditida - saprobionts. The increase in bacteriovorous and fungivorous nematodes is apparently assigned to bacterial and fungal populations that appear in higher abundance after the organic amendments of soil and afford the food base for these nematodes (Griffiths et al., 1994). After and adding of organic matter to the soil, organic residues must be decomposed to release nutrients for plant uptake. This decomposition can be divided into two channels, a faster-bacterial channel and a slower fungal-based channel. Soil ecosystem types and nutrient forms (C:N ratio) determine the predominant decomposition channels (Ingham et al., 1985). As an extension of these decomposition channels, when the bacterivo-rous and fungivorous nematodes graze on these microbes, they give off CO2 and NH4+ and other nitrogenous compounds, affecting C and N mineralization directly (Ingham et al., 1985).
The suppression of plant parasitic nematodes and improving of number of free-living (beneficial) nematodes were
found also in many other studies (Akhtar, 1999; McSorley & Frederick, 1999; Valocka et al., 1999, 2000; Bulluck et al., 2002a; Arancon et al., 2003b; YingXia et al., 2003; Wang et al., 2006; Nahar et al., 2006; Yao et al., 2006). On the other hand, the application of chemical fertilizer decreased the number of genera of bacterivores and omni-vores-predators. The numbers of total nematodes, bacterivores, plant parasites and omnivores-predators were significantly positively correlated with the contents of total organic carbon, total N, alkali-hydrolysable N, available P and available K (Cheng & Zhi-Ping, 2008). However, Wasilewska (1995) stated that manuring increased the proportion of bacterivorous, fungivorous but also plant parasitic nematodes and decreased the number of omnivores and predators. Renco et al. (2010) found that bacterivorous nematodes decreased after sewage sludge and municipal green residues compost application, while at compost derived from penicillin production significantly increased the number of these nematodes. A significant reduction in number of plant parasitic nematodes in soil was observed in all tested composts compared to untreated
soil, with a similar behaviour of tested composts for root-fungal feeding nematodes. Garcia-Alvarez et al. (2004) found that the application of mature compost did not alter the structural diversity of the nematode population during the six-year study. The similar results were obtained by Biederman et al. (2008) where the application of untreated urban wood waste did not affect the nematode density, family diversity and family richness by the amendment treatments. However, the number of bacterivorous, plant parasitic, omnivorous and predatory nematodes significantly increased during a 3-year study.
The effect of applied doses on nematode suppression There are several factors which determine the effect of organic fertilizer on plant parasitic nematodes. Besides the quality of applied organic matter (maturity), soil structure and target nematode species, much important factor for intensity of suppression is applied doses as well, because of at higher doses more efficacious substances are added. For example, Gutpa and Kumar (1997) stated that level of reduction of Tylenchorhynchus spp. and Helicotylenchus spp. in soil increased at higher doses and the longer periods of treatment by compost, fenugreek and chickpea straw or groundnut and mustard cakes. Renco et al. (2009) also found that relationship between applied doses and number of parasitic nematodes of genera Bitylenchus, Helicotylenchus, Heterodera, Paratylenchus and Rotylenchulus showed a significantly high negative correlation. That the applied doses can be consider as important factor for intensity of reduction of plant parasitic nematodes was confirmed in several other particular studies where increasing dose increase plant parasite nematode suppression (Akhtar & Mahmood, 1996; Nico et al., 2004; Renco et al., 2007, 2010; Rawdan et al., 2009)
Ammonium (N) and C:N ratio
Effectiveness of organic amendments suppression on plant parasitic nematode varies and depending mainly upon the nematode species and type of organic material (Akhtar & Alam, 1993). Although, the exact mechanism(s) of action of organic matter compounds is not exactly known at this time (Nico et al., 2004), (Oka et al., 2000) stated, that ammonia released from organic amendments during mi-crobial composition plays and important role in nematode control, though the environmental conditions, such as soil pH, temperature and humidity influence the nematicidal activity of ammonia.
Rodriguez-Kabana et al., (1987) pointed out, that the most effective amendments are those which have generally high nitrogen contents relative to carbon. Noe (1993) and Akh-tar and Mohmood (1996) also stated, that application of composted manure, oilcakes of neem and castor with low C:N ratios (6 - 10:1) and high ammonium nitrogen content resulted in a decrease of plant parasitic nematodes. Castag-none-Sereno and Kermarrec (1991) or Nico et al. (2004) also attributed the suppression of M. incognita and M. javanica to ammonia contained in sewage sludge treatments or dry cork compost (C:N < 20). Renco et al. (2010)
found, that composts with the highest native NH4+ content (C:N 20 - 23:1) suppressed plant parasitic nematode in great extent, and there was a positive relationship between plant parasitic nematode suppression and ammoniacal N content in the soil. However, the greatest extent of suppression of plant nematodes was recorded at compost with C:N ratio 4:1, though the NH4+ content were lower. In study of Agu (2008) plants of African yam bean treated with poultry and farmyard manures (C:N 1 - 2:1) gave significantly higher yields than those of other organic manures with wider C:N ratios. This was because rare root-galls caused by M. incognita occurred at those treatments. Otiefa and Elgindi (1962) wrote that plants with fewer root-galls would translocate more nutrients to vegetative organs than heavily galled roots. Miller and Donahue (1990) stated that organic residues with C:N ratios of 20:1 or narrower have sufficient nitrogen to supply to the decomposing microorganisms and also to release for plant use.
Urea (330 kg N y ha-1) was also nematode toxic and improved plant growth when was applied alone or in combination with the neem-based products (Achook, Suneem G) because neem contains triterpene that acts by delaying the rapid transformation of ammonium N to nitrate N (nitrification inhibitor). This ensures slow and continuously available nitrogen during plant growth (Akthar, 2000). Wang et al. (2006) observed the increase of bacterivorous, fungivorous and occasionally omnivorous nematodes after incorporation of sunn hemp hay into soil, however the plant parasitic nematodes was suppressed relative to ammonium nitrate fertilizer. The abundance of bacterivores, fungivores and predatory nematodes, and total nematode abundance increased with the increasing dose of sunn hemp hay applied. In comparison to that, fertilization with ammonium nitrate increased the percentage of herbivores, but reduced percentage and abundance of omnivores. Suppressiveness of organic amendments on plant parasitic nematodes generally resulted from different contributory mechanisms (Stirling, 1991). It seems that ammonia is one best suppressive or toxic. But also other mechanisms of action are found in decaying plant material such as components directly toxic to nematodes (chitin, phenols, tannins, azadirachtin, ricinin, rutin, terpens) (Rodriguez-Kabana, 1986; Spiegel et al., 1987; Rich et al., 1989, Sasanelli & D'Addabbo, 1995; Atungwu et al., 2009; Maistrello et al., 2010; Renco et al., 2012).
Other alternative methods for plant parasitic nematode control
That organic amendments reduce the number of parasitic nematodes in the soil has been demonstrated many times, but first the producers would have to protect their land before the infection and transfer of parasitic nematodes from infected plots. Nematodes have a limited ability to move and spread their own activity and therefore are due to passive transfer of man-the most common way of spreading. Therefore, the most important protection against plant nematodes consider prevention, it means the use the
pathogen-free planting material from uninfected nematode soils (e.g potato tubers, plant transplants). In the event that we have in the garden soil infected with nematodes, we can either directly limit their ability to grow and reproduce, thus increasing population, an indirect support growth of crops. For example, the use of resistant cultivar, grafting, biological control (by naturally occurring antagonists such as nematophagous fungi, endoparasitic fungal parasites, AMF arbuscolar mycorrhizal fungi, the obligate parasite Pasteuria penetrans, predaceous organisms), soil solariza-tion (Sasanelli et al., 1997), biofumigation, nematicidal plants, soil steam sterilization, fumigants and non-fumi-gants nematicides (Sasanelli & D'Addabbo 1992, 1993; Sasanelli, 2009).
Conclussion
Farmers are constantly under pressure of naturalists, biologists, environmentalists or hydrologist to reduce use of pesticides (nematicides) and synthetic fertilizers. However, they must maintain the profitability of crops and crop quality. Annual global loss in agriculture due to damage by plant parasitic nematodes has been estimated as US$100 billion worldwide (Oka et al., 2000). For effective control of parasitic nematodes of plants in soil is necessary to choose an appropriate combination of several methods. Despite the fact that each control method has limited use because different developmental cycle of nematodes (e.g. cyst or root-gall forming nematodes) and varying range of host plants with different growth, an appropriate combination of methods can lead to successful reduction in the number of parasitic nematodes in soil and plants. It is clear, that organic soil amendments stimulate the development of population of soil microorganisms and reduce the plant parasitic nematode population or plant diseases as well as improve soil fertility, physical properties of soil, water retention, water infiltration, permeability, aeration and plant growth. The majority of studies have focused on the different types of organic amendments as suppressants of plant parasitic nematodes of economic importance, especially root-knot nematodes, because their hosts' range and reproduction rate is large. Because the price of pesticides (nematicides) and synthetic fertilizers is high, those can be related to increase of cost for cultivation of crops and may lead to large increases in food prices for example, in the poor harvest years (in the dry or very rainy). In addition to this, the pesticides are usually selective and destroying only targeted organisms. They react quickly and strongly (Deasaeger et al., 2011), but the protection of plants is not long, because their effect is short without some other positive effects on plant growth and soil quality. For this reason, the use of organic fertilizer as a tool for control of parasitic nematodes and other soil pathogens is beneficial because it is a natural material, low cost (usually made by the growers), from which nutrients and nematicidal substances are released gradually throughout the whole vegetation period.
Acknowledgement
The author acknowledge the support of the scientific grant agency VEGA (Grant N° 2/0079/13).
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Received October 24, 2012 Accepted December 5, 2012
