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1 Meloidogyne Species - a Diverse Group of Novel and Important Plant Parasites 1 Meloidogyne Species - a Diverse Group of Novel and Important Plant Parasites Maurice Moens1, Roland N. Perry2 and James L. Starr3 1 Institute for Agricultural and Fisheries Research, Merelbeke, Belgium and Ghent University, Ghent, Belgium; 2 Rothamsted Research, Harpenden, Hertfordshire, UK and Ghent University, Ghent, Belgium; 3 Texas A&M University, College Station, Texas, USA 1.1 Introduction I 1.2 Impact 2 1.3 History of the Genus 2 1.4 Current Trends in Species Identification 2 1.5 Life Cycle 3 1.6 Diversity in Biology 6 1.7 Major and Emerging Species 8 1.8 Interactions with Other Plant Pathogens 11 1.9 Management and Control 11 1.10 Conclusions and Future Directions 12 1.11 References 13 1.1 Introduction out showing external symptoms on the harvested products, e.g. symptomless potato tubers. The Root-knot nematodes are members or the genus rapid rate of developmem and reproduction on Meloidogyne (Gbldi, 1892), Meloidogyne is of Greek good hosts results, in the majority or species, in origin and means 'apple-shaped female'. They are several generations during one cropping season, an economically important polyphagous group or leading to severe crop damage. Damage may highly adapted obligate plant parasites, are dis• consist of various degrees of stunting, lack or vig• tributed worldwide and parasitize ncarly every our, and wilting under moisture stress. Secondary species of higher plant. Typically thcy reproduce infection by other pathogens often results in and feed on modified living plant cells within extensive decay of nematode-inlected tissues. The plant roots, where they induce small to large galls common explanation for these above-ground or root-knots, hence their vernacular name. The symptoms is that i\lefoidogyne inlection affects above-ground symptoms arc not readily apparent water and nutrient uptake and upward transloca• and may be similar to those produced on any tion by the root system. By disrupting the host plants having a damaged and malrunctioning plant physiology, root-knot nematodes may not root system. Hosts may be heavily infected with- only reduce crop yield but also product quality ©CAB International 2009. Root-knot Nematodes (eds R.N. Perry, M. Moens and J.L. Starr) 2 M. Moens et al. (e.g. of potatoes and carrots) and therefore are of is given in Hunt and Handoo, Chapter 3, this great economic and social importance (see Greco volume; a brief summary is included here as and Di Vito, Chapter II, this volume). part of the introduction to the genus and its basic biology. The first illustrated report of root-knot disease appeared during the middle 1.2 Impact of the 19th een tury when the clergyman Miles Joseph Berkeley (1855) first attributed galls detected on glasshouse cucumber roots to nem• Damage and yield losses caused by plant patho• atodes. The first description of a root-knot gens, including plant-parasitic nematodes, are, on nematode was made by Cornu (1879); it was average, greater in tropical than in temperate based on nematodes found in root galls that regions because of greater pathogen diversity, more favourable environmental conditions for were detected on sainfoin (Onobrychis sativus Lam.) in the Loire valley, France. In 1887, pathogen colonization, development, reproduc• Gbldi briefly described and illustrated a root• tion and dispersal, and lack of human, technical and financial resources to combat infections knot nematode from coffee plants in Brazil and named it M. exigua. Although the 1887 publica• De Waele and Elsen, 2007). Se,"erity of damage tion was an advance copy or preprint of the full caused by l\1eloidogyne can be species-specific and also vary by host, crop rotation, season and soil article subsequently published by Gbldi in 1892, the 1887 article meets the requirements to type (Greco et al., 1992; Potter and Olthof, 1993). establish the actual publication date for the Similarly, economic thresholds "ary, primarily genus and type species as 1887 (see Hunt and depending on these same factors. Damage thresh• Handoo, Chapter 3, this volume for a full olds have been established for several crops, where account of this decision). the average is approximately 0.5-2 juveniles/g of The name Heterodera marioni was widely used soil (or from the lower limits of detection, over for root-knot nematodes until 1949, when 1000 individuals/500cm3 of soil; see Greco and Chitwood removed the root-knot nematodes Di Vito, Chapter II, this volume. In addition to direct costs, root-knot nematodes cause indirect from the genus Heterodera bccause they differed from cyst ncmatodes. Since the oldest name for costs because of the quarantine status of some the genus was Gbldi's Meloidogyne, that name had species of ,Ueloidogyne in se'"eral countries or precedence. Chitwood redescribed lV!. arenaria, regions. For example, M. chitwoodi is increasingly ,\1. exigua, 1\1. incognita and N1. javanica, and regulated because it is a serious pest of potato and described M. hapla and a variety of N1. incognita he other economically important crops such as car• termed M. incognita var. acrita. The species were rot, and the known geographical distribution is separated from each other on the basis of peri• still relatively limited. It is on the list of prohibited neal pattern morphology, stylet knob shape, and pests of man)" countries (Canada, the EU, Mexico and other countries in Latin America, and the Far length of stylet and dorsal gland orifice. Since Chitwood's publication, many more lVleloidogyne East) (Rockland et al., 2006). ,\Ieloidogyne Jallax, species have been described (see Hunt and another pest of potato, is recognized as an impor• Handoo, Chapter 3, this volume, for a full list). tant pest by fewer countries (e.g. the EU), although Species descriptions gradually included increas• it poses a similar economic risk. In the future, ing numbers of features (observed by light micro• additional species of root-knot nematodes that scopy and/ or electron microscopy) of females, might be added to the list of quarantine species males and second-stage juveniles (J2) (see Jepson, include recently described species such as M. minor 1987, for a re'~e\V, and Eisenback and Hunt, in Europe and ,\1. citri in the USA. Chapter 2, this volume). 1.3 History of the Genus 1.4 Current Trends in Species Identification A more detailed account of the background lit• erature, progress in descriptions and classifica• Rcsearch on cytogenetics, isozymes and the tion, and authorities for the species of the genus genome of root-knot nematodes, mostly since the Meloidogyne Species 3 1970s, provided further evidence for the large approach requires some expensive equipment diversity of species within the genus. Importantly, (the PCR thermocycler) and supplies, and a several new technologies have provided tools to clean laboratory environment. None the less, assist in species identification. One of the most these approaches are likely to play an ever• important has been the use of isozyme pheno• increasing role in species identification (especially types, using PAGE of crude protein extracts and by those not well trained or experienced in clas• histochemical stains for non-specific esterases, sical morphometrics) and will be used extensively superoxide dismutase, malate dehydrogenase, by regulatory agencies. and glutamate oxaloacetate transaminase (Esbenshade and Triantaphyllou, 1985), with the esterases and malate dehydrogenase being the most useful for discriminating the four common 1.5 Life Cycle Meloidogyne species: M. arenaria, M. hapla, M. incog• nita and j\1. javanica. These enzymes have also The root-knot nematode life cycle is summarized been used successfully with less common species in Fig. 1.1 and Plate 36. Females lay eggs into such as M. enterolobii (= M. mayaguensis) (Brito et aI., gelatinous masses composed of a glycoprotein 2004), M. parti!Jla (Starr et al., 1996), and M. trijO• matrix, which is produced by rectal glands in the liophila (Mercer et aI., 1997). With the appropriate female, keeps the eggs together and protects equipment it is possible to make the species iden• them against environmental extremes and pre• tification on indi\-idual females. Ibrahim and dation. The egg masses are usually found on the Perry (1992) showed that PAGE combined with surface of galled roots, although they may also staining for esterases could be used with Meloidogyne be embedded within the gall tissue. The egg females in galled roots, thus obviating the need to mass is initially soft, sticky and hyaline but separate nematodes from the host tissue. It becomes firmer and dark brown with age. remains to be determined if these procedures will Surprisingly, there has been only limited analysis be able to discriminate all of the currently recog• of the glycoproteins (Sharon and Spiegel, 1993) nized species of this genus. or other components of the gelatinous matrix, More recently, much effort has been given despite its obvious importance. In addition to to the development of species-specific markers providing some protection to the eggs from envi• using variation in the genomes of the different ronmental extremes, it has been demonstrated species coupled with PCR techniques (see Blok that the matrix has antimicrobial properties and Powers, Chapter 4, this volume). There are (Orion and Kritzman, 1991). now many reports that describe protocols for dis• Within the egg, embryogenesis proceeds to tinguishing numerous species using species-spe• the first-stage juvenile, which moults to the infect• cific primers derived from the nuclear genome ive J2. Hatch of the J2 is primarily dependent on (Adam et aI., 2007) and the mitochondrial temperature and sufficient moisture, although genome (Powers and Harris, 1993). A major other factors, including root diffusate and gener• benefit of these systems is that they can be used ation, modifY the hatching response (see Curtis on an individual J2, the most common stage of et aI., Chapter 6, this volume) so that theJ2 hatch the nematode found in the soil, thus eliminating when conditions are favourable for movement the need to use mature females from roots of and host location.
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