Response of rice genotypes to rice root-knot nematode (Meloidogyne graminicola) infection under varying temperature regimes

The rice root-knot nematode (RRKN), Meloidogyne graminicola, is one of the major pests of the rice–wheat cropping system. Resistance against M. graminicola in rice could be most valuable in alleviating this problem. The host response of 75 Oryza genotypes was examined at three day/night temperature regimes, 29/26°C, 34/31°C and 38/35°C, in Pluronic gel as well as in soil. Out of the 75 selected rice genotypes, only Zhenshan 97 B exhibited high resistance to this set of temperature regimes, with the least number of galls/root system. At 34/31°C, more second-stage juveniles (J2s) were hatched and J2 population densities in roots of the susceptible rice genotypes increased significantly compared with those of plants grown at the 29/26°C. In resistant genotypes, only a few J2s penetrated roots and developed into mature females. The histopathological studies revealed that in susceptible rice genotypes at high-temperature regimes, the multinucleate giant cells were well developed. The results presented in this study revealed that an increase in temperature had a significant effect on the resistance of rice genotypes and resistance appeared more pronounced in genotype Zhenshan 97 B. This resistant genotype can be used in marker-assisted selection to develop resistant elite cultivars.     

Reaction of a heterozygous tomato hybrid bearing the Mi-1.2 gene to 15 Meloidogyne species

Many root-knot nematode (RKN) species (Meloidogyne spp.) are polyphagous and cultivated tomato (Solanum lycopersicum) is one of their preferential hosts, leading to significant losses. It is known that the dominant Mi-1.2 gene in tomato confers resistance to the three most important RKN species—M. incognita, M. javanica, and M. arenaria, and minor species—M. ethiopica, M. hispanica, and M. luci. However, little information is available about resistance of tomatoes carrying this gene to other tomato-infecting RKN species. In this study, resistance conferred by the Mi-1.2 gene/locus was evaluated against populations of 15 Meloidogyne species, employing tomato cultivars Santa Clara (homozygous recessive mi-1.2/mi-1.2, susceptible) and Debora Plus (heterozygous Mi-1.2/mi-1.2, resistant). Debora Plus was susceptible only to M. enterolobii and M. hapla, and was resistant to the other Brazilian populations of M. arenaria, M. ethiopica, M. exigua, M. hispanica, M. incognita, M. inornata, M. izalcoensis M. javanica, M. konaensis, M. luci, M. morocciensis, M. paranaensis, and M. petuniae. Mi-1.2 is located on tomato chromosome 6 within a cluster of seven homologous genes of the nucleotide-binding site leucine-rich repeat (NBS-LRR) family; further research is required to confirm if this multiple Meloidogyne spp. resistance phenotype is controlled exclusively by Mi-1.2 or by combined action of other closely linked genes. This evaluation of resistance of the Debora Plus cultivar to several Meloidogyne species suggests that the Mi-1.2 gene/locus may reduce losses induced by a wide range of Meloidogyne spp. Further studies using additional resistant cultivars and other populations of Meloidogyne spp. are needed to confirm these results.     

Resistance sources to Fusarium oxysporum f. sp. cubense tropical race 4 in banana wild relatives

Target trait evaluation in crop wild relatives is an important prerequisite for efficiently using the potential useful genes located in this valuable germplasm. Over recent decades, Fusarium oxysporum f. sp. cubense tropical race 4 (Foc‐TR4) has seriously threatened worldwide banana plantations. Breeding new resistant cultivars from wild banana species is expected to provide invaluable additional resources. However, knowledge on resistance to Foc‐TR4 in wild Musa species is very limited. In this study, eight genotypes of wild banana relatives (Musa acuminata subsp. burmannica, M. balbisiana, M. basjoo, M. itinerans, M. nagensium, M. ruiliensis, M. velutina and M. yunnanensis) were characterized for resistance to Foc‐TR4 in both greenhouse and field conditions. Most wild bananas showed higher resistance levels to Foc‐TR4 than the reference cultivars ‘Brazilian’ (AAA, susceptible) and ‘Goldfinger’ (AAAB, moderate resistance). Among the wild species, M. balbisiana showed the highest levels of disease intensity followed by M. acuminata subsp. burmannica. Some individuals of M. yunnanensis, M. nagensium, M. ruiliensis and M. velutina showed low levels of rhizome discolouration in greenhouse conditions, but were resistant in the field. No symptoms were observed on M. basjoo and M. itinerans, suggesting higher levels of resistance to Foc‐TR4. The results revealed different sources of resistance to Foc‐TR4 in banana wild relatives, which constitute a valuable genetic resource for banana breeding programmes aiming to produce cultivars resistant to fusarium wilt.     

近年水稻主要线虫病害的研究进展

目前,以拟禾谷根结线虫Meloidogyne graminicola、水稻干尖线虫Aphelenchoide besseyi、水稻潜根线虫Hirschmanniella oryzae、水稻茎线虫Ditylenchus angustus和旱稻孢囊线虫Heterodera elachista为主的水稻寄生线虫侵染水稻以及其它禾本科作物引致的线虫病害,已造成了严重的经济损失。近年来,对拟禾谷根结线虫的致病机制和寄主的防御机制等方面的研究已取得了显著进展,但对其它水稻寄生线虫与寄主互作机制的研究还存在较大空白。水稻寄生线虫的防治多依赖化学农药,抗性品种和生物防治等措施的实施存在一定的局限性。本文对上述5种主要水稻寄生线虫病的发生为害、致病机制以及防治方法等进行了综述,并对水稻寄生线虫致病机理的研究以及抗性品种、生物防治和诱导化合物的应用进行了探讨。     

Food preferences of the multi-mammate mouse,Mastomys natalensis,in irrigated rice habitats in Tanzania

We investigated the composition of the diet of the multi-mammate mouse, Mastomys natalensis, within irrigated rice and fallow field habitats at set time periods related to rice crop growth stages. In both habitats, vegetative plant material, i.e. leaves, stems and seeds, were the most abundant components of the rodent's diet, while other food types (invertebrates, fruits) were observed only in low quantities. We conclude that vegetative plant material and seeds were the main types of food consumed not only due to their relatively higher abundance in the environments under study but also because of the highly specialised herbivorous/granivorous nature of the dominant rodent species, M. natalensis. Thus, the introduction and expansion of continuous rice-cropping using irrigation in Tanzania is likely to be severely constrained by the presence of M. natalensis. In our opinion, field hygiene, including the removal of alternative food resources and nesting sites for M. natalensis near cropping areas, may help to both lower rodent population numbers and reduce immigration potential. Non-chemical rodent control methods such as trap barrier systems developed for lowland irrigated rice in south-eastern Asia should, we argue, be evaluated for their effectiveness under African conditions.     

Studies on the weed hosts of the rice root nematode, Hirschmanniella spinicaudata Sch. Stek. 1944

Weed hosts of the rice root nematode, Hirschmanniella spinicaudata were surveyed in a swamp rice field at Badeggi, Nigeria. Forty-five weed species, mainly grasses and sedges occurred very frequently and were very good hosts of the nematode. A wild rice, Oryza longistaminata was a preferred host. The importance of the weed hosts is discussed.     

Nematode quantitative resistance conferred by the pepper genetic background presents additive effects and is stable against different isolates of Meloidogyne incognita

Root‐knot nematodes (RKNs), Meloidogyne spp., are a major disease problem in solanaceous crops worldwide, including pepper (Capsicum spp.). Genetic control provides an economically and environmentally sustainable protection alternative to soil fumigants. In pepper, resistance to the main RKN species (M. incognita, M. javanica and M. arenaria) is conferred by the major genes (R genes) Me1, Me3 and N. However, RKNs are able to develop virulence, thus endangering the efficiency of R genes. Quantitative resistance (QR) against Meloidogyne spp. is expected to provide an alternative to R genes, or to be combined with R genes, to increase the resistance efficiency and durability in pepper. In order to explore the ability of QR to protect pepper against RKNs, five pepper inbred lines, differing in their QR level, were tested directly, or after combination with the Me1 and Me3 genes, for their resistance to a panel of M. arenaria, M. javanica and M. incognita isolates. The M. arenaria and M. javanica isolates showed low pathogenicity to pepper, unlike the M. incognita isolates. The QR, controlled by the pepper genetic background, displayed a high resistance level with a broad spectrum of action, protecting pepper against Me3‐virulent as well as avirulent M. incognita isolates. The QR was also expressed when combined with the Me1 and Me3 genes, but presented additive genetic effects so that heterozygous F₁ hybrids proved less resistant than homozygous inbred lines. The discovery of this QR is expected to provide promising applications for preserving the efficiency and durability of nematode resistance.     

Effects of initial nematode population density and water regime on resistance and tolerance to the rice root-knot nematode Meloidogyne graminicola in African and Asian rice genotypes

The rice root-knot nematode (RKN), Meloidogyne graminicola, is an important pathogen affecting rice production in South and Southeast Asia. Efficacy of resistance and tolerance in selected M. graminicola-resistant African rice genotypes TOG5674, TOG5675 and CG14 and -susceptible Asian rice genotypes IR64 and UPLRi-5 were examined under a range of initial population densities (Pi) and water regimes. Resistance to M. graminicola in resistant rice genotypes was not broken with increasing pathogen pressure (Pi = 15,000 to 60,000 J2/plant). Resistant rice genotypes were even tolerant to the damage and yield loss caused by high pathogen pressure. On the other hand, increasing Pi levels caused more damage on susceptible rice genotypes. Final nematode population densities in the root systems of resistant and susceptible rice genotypes were significantly lower under flooded conditions than under upland and drought conditions. TOG5674, TOG5675 and CG14 were more tolerant to M. graminicola infection even when grown under upland and drought conditions while IR64 and UPLRi-5 were highly sensitive.     

The threat of root‐knot nematodes (Meloidogyne spp.) in Africa: a review

Meloidogyne species pose a significant threat to crop production in Africa due to the losses they cause in a wide range of agricultural crops. The direct and indirect damage caused by various Meloidogyne species results in delayed maturity, toppling, reduced yields and quality of crop produce, high costs of production and therefore loss of income. In addition, emergence of resistance‐breaking Meloidogyne species has partly rendered various pest management programmes already in place ineffective, therefore putting food security of the continent at risk. It is likely that more losses may be experienced in the future due to the on‐going withdrawal of nematicides. To adequately address the threat of Meloidogyne species in Africa, an accurate assessment and understanding of the species present, genetic diversity, population structure, parasitism mechanisms and how each of these factors contribute to the overall threat posed by Meloidogyne species is important. Thus, the ability to accurately characterize and identify Meloidogyne species is crucial if the threat of Meloidogyne species to crop production in Africa is to be effectively tackled. This review discusses the use of traditional versus molecular‐based identification methods of Meloidogyne species and how accurate identification using a polyphasic approach can negate the eminent threat of root knot nematodes in crop production. The potential threat to Africa posed by highly damaging and resistance‐breaking populations of ‘emerging’ Meloidogyne species is also examined.     

Distribution and genetic diversity of root‐knot nematodes (Meloidogyne spp.) in potatoes from South Africa

A molecular‐based assay was employed to analyse and accurately identify various root‐knot nematodes (Meloidogyne spp.) parasitizing potatoes (Solanum tuberosum) in South Africa. Using the intergenic region (IGS) and the 28S D2–D3 expansion segments within the ribosomal DNA (rDNA), together with the region between the cytochrome oxidase subunit II (COII) and the 16S rRNA gene of the mtDNA, 78 composite potato tubers collected from seven major potato growing provinces were analysed and all Meloidogyne species present were identified. During this study, M. incognita, M. arenaria, M. javanica, M. hapla, M. chitwoodi and M. enterolobii were identified. The three tropical species M. javanica, M. incognita and M. arenaria were identified as the most prevalent species, occurring in almost every region sampled. Meloidogyne hapla and M. enterolobii occurred in Mpumalanga and KwaZulu‐Natal, respectively, while M. chitwoodi was isolated from two growers located within the Free State. Results presented here form part of the first comprehensive surveillance study of root‐knot nematodes to be carried out on potatoes in South Africa using a molecular‐based approach. The three genes were able to distinguish various Meloidogyne populations from one another, providing a reliable and robust method for future use in diagnostics within the potato industry for these phytoparasites.     

云南野生稻和地方稻资源抗白叶枯病分析

为鉴定云南稻种质资源对水稻白叶枯病的抗性情况,于孕穗期采用剪叶接种方法,用水稻白叶枯病强致病型代表菌株BD8438、CN9404和X1接种云南野生稻和地方稻种质资源,以病斑长度大于6 cm为感病分界线,对其抗感表现型进行调查分析。结果显示,共鉴定出来源于云南省不同种植生态区的186份地方稻抗性材料和22个野生稻抗病居群。野生稻对水稻白叶枯病的抗病能力较地方稻强,其中疣粒野生稻的抗性最强,抗病等级为0~2;药用野生稻次之,抗病等级为1~2;普通野生稻相对较差,抗病等级为1~5。地方稻抗性资源来自于云南省各个传统水稻种植区,抗性1级的材料占17%,抗性2级的占2%,抗性3级的占81%;按照稻种质资源亚种类型、粘糯性和水旱性分类,各类型地方稻抗性材料所占比例分别为粳稻占61%、籼稻占39%;粘稻占66%、糯稻占34%;水稻占83%、陆稻占17%。从利用抗白叶枯病基因培育新品种的角度评价,这些抗性资源具有潜在的发掘利用价值。     

Resistance to Root-knot Nematodes (Meloidogyne spp.) in Vegetable Crops

Abstract

The development of root-knot nematode resistant vegetable varieties has provided an alternative control method to chemical and crop rotation. The term resistance is discussed. Work on 18 vegetables is reviewed.

Thirty tomato strains resistant to one or more Meloidogyne sp. are listed, there was a marked absence of reports on varieties resistant to M. hapla. Several workers have observed resistance in some cultivated and wild Solanum spp. The resistance could be increased by further sib and backcrossing experiments. Eggplant varieties tested for resistance to Meloidogyne spp. showed varying degrees of resistance in about 14 varieties. Further backcrossing studies between Solanum melongena and S. sysimbrifolium could provide useful results. Tests on resistance to nematodes in pepper have revealed many resistant varieties. All the pepper varieties tested were susceptible to M. hapla. The nature of resistance in sweet potato has been studied. Some varieties of Cucumis spp. have been found to be resistant to Meloidogyne spp. Since no resistance was found in C. melo, these wild species could be used in the development of a commercial muskmelon variety. Tests with 83 watermelon varieties indicated that all varieties were resistant to M. hapla. It was difficult to find resistance to Meloidogyne spp. in Cucurbita, but tests on wild species have yet to be carried out. Work on lima bean, snap bean, pea, soyabean, cowpea and broadbean has produced a limited number of resistant vegetable varieties, which are described. Since resistance patterns in plants may change under different environmental or biological conditions, it is essential to test varieties under these conditions for a long period before they are released for commercial cultivation. The advantages of using resistant vegetable varieties, as compared with other methods of pest control, are outlined.     

Identity and variability of Pythium species associated with yield decline in aerobic rice cultivation in the Philippines

The cultivation of aerobic rice in the tropics enables farmers to save water without lowering productivity. Unfortunately, this system suffers from declining yields due to a disease complex involving nematodes, pathogenic Pythium spp. and nutrient deficiencies. Assessing the impact of each underlying factor can contribute to efficient disease control measures. This study therefore investigated pathogenic and genotypic variability among Pythium species from affected aerobic rice fields in the Philippines using pathogenicity assays and sequence information from the internal transcribed spacer (ITS) region and β‐tubulin gene. Three closely related Pythium spp., P. arrhenomanes, P. graminicola and P. inflatum, were recovered from affected aerobic rice fields. All P. arrhenomanes isolates reduced rice seedling growth, whereas only a few P. graminicola isolates and no P. inflatum isolates were pathogenic, indicating that P. arrhenomanes is probably the most important species affecting rice. Both P. arrhenomanes and P. graminicola isolates showed little genetic variation, despite the observed pathogenic variation within P. graminicola. Intraspecific variation was higher among P. inflatum isolates, but again no correlation was observed with phenotype. When screening P. arrhenomanes isolates from other hosts such as sugarcane, maize and several grasses, a link between pathogenic and genetic variability was detected. However, rice and maize isolates seemed to lack host specificity, and therefore crop rotation with maize might be a risky strategy to manage yield decline in Philippine aerobic rice fields.     

Phylogenetic and morphological analyses of Pythium graminicola and related species

Isolates of Pythium graminicola and related species were differentiated using restriction fragment length polymorphism (RFLP) analyses of the internal transcribed spacer (ITS) regions of rDNA and the cytochrome c oxidase subunit II (COX II) gene. These sequences were used in subsequent phylogenetic analyses. Finally, the phylogenetic placement of species was compared to that determined from morphological characteristics. The 62 isolates tested were divided into seven groups, A–G, based on RFLP analysis of the rDNA-ITS region. In the RFLP analysis of the COX II gene, isolates were divided into groups similar to those based on ITS-RFLP. Groups A and B were each separated into two additional subgroups. Grouping of isolates based on RFLP analyses agreed with the morphological differentiation. Groups A, B, D, E, F, and G were identified as P. graminicola, P. arrhenomanes, P. aphanidermatum, P. myriotylum, P. torulosum, and P. vanterpoolii, respectively. Group C was closely related to group B based on phylogenetic analysis of the rDNA-ITS region and the COX II gene and is similar to P. arrhenomanes. Each of the other species occupied their own individual clades. Although P. arrhenomanes is morphologically similar to P. graminicola, our phylogenetic analyses revealed that it was evolutionarily distant from P. graminicola and more closely related to P. vanterpoolii. Our analysis also revealed that P. torulosum with smaller oogonia is more closely related to P. myriotylum with large oogonia than to P. vanterpoolii, which forms smaller oogonia and is morphologically similar to P. torulosum. P. aphanidermatum with large oogonia and aplerotic oospores was not related to the morphologically similar species P. myriotylum. Results suggest that P. graminicola and related species are phylogenetically distinct, and molecular analyses, in addition to morphological analyses, are necessary for the accurate taxonomic placement of species in this complex.     

Effect of residue management and fallow length on weeds and rice yield

Reduced fallow length in slash-and-burn rice (Oryza sativa L.) production systems of northern Laos increased weed pressure, labour requirement and the need for soil and moisture conservation. On-farm and on-station studies and on-farm surveys were used to evaluate the effect of residue management and cropping intensity on weed population, rice yield and nematode density. Residue loads were 2.3–4.4 t ha^?1 after a rice crop and 9.5 t ha^?1 after 1 year of fallow. Compared with farmers' traditional burning of crop and weed residues, mulching reduced rice yield by 43% in one out of four comparisons and increased weed biomass by 19–100%. Compared with continuous rice treatments (averaged over burning and mulching treatments), treatments with fallow or cowpea [Vigna unguiculata (L.) Walp.] in the previous year had 32% less herbaceous weed biomass, 90% fewer Ageratum conyzoides L. and over 99% fewer Meloidogyne graminicola Golden & Birchfield. Rice yield was negatively associated with A. conyzoides density (?0.62, P < 0.01) and M. graminicola number (?0.42, P < 0.05). Less striking effects of fallow period on A. conyzoides and M. graminicola, observed from on-farm surveys, demonstrate the limitations of on-farm studies because of undocumented effects of farmers' management decisions.     

Comparison of two short DNA barcoding loci (COI and COII) and two longer ribosomal DNA genes (SSU & LSU rRNA) for specimen identification among quarantine root-knot nematodes (Meloidogyne spp.) and their close relatives

Root-knot nematodes (Meloidogyne spp.) are important pests of numerous crops worldwide. Some members of this genus have a quarantine status, and accurate species identification is required to prevent further spreading. DNA barcoding is a method for organism identification in non-complex DNA backgrounds based on informative motifs in short DNA stretches (≈600 bp). As part of the EU 7th Framework project QBOL, 15 Meloidogyne species were chosen to compare the resolutions offered by two typical DNA barcoding loci, COI and COII, with the distinguishing signals produced by two ribosomal DNA genes (small and large subunit rDNA; SSU?≈?1,700 and LSU?≈?3,400 bp). None of the four markers distinguished between the tropical species Meloidogyne incognita, M. javanica and M. arenaria. Taking P ID (Liberal) values ≥0.93 as a measure for species delimitation, the four mtDNA and rDNA markers performed well for the tropical Meloidogyne species complex, M. enterolobii, M. hapla, and M. maritima. Within cluster III A (Holterman et al. Phytopathology, 99, 227–235, 2009), SSU rDNA did not offer resolution at species level. Both mtDNA loci COI and COII did, whereas for LSU rDNA a longer fragment (≥700 bp) is required. The high level of mitochondrial heteroplasmy recently reported for M. chitwoodi (Humphreys-Pereira and Elling Nematology, 15, 315–327, 2013) was not found in the populations under investigation, suggesting this could be a regional phenomenon. For identification of RKNs, we suggest the combined use of SSU rDNA with one of three other markers presented here.     

Differential response of phytohormone signalling network determines nonhost resistance in rice during wheat stem rust (Puccinia graminis f. sp. tritici) colonization

Stem rust caused by Puccinia graminis f. sp. tritici is one of the most devastating diseases of wheat. Breakdown of host resistance under field conditions triggered by the evolution of new pathogenic races and pathotypes is a perennial threat for wheat cultivation. Rice, often grown in a rice–wheat cropping system, is immune to rust infection. Our microscopic studies revealed that P. graminis f. sp. tritici, although displaying nearly identical uredospore germination, stomatal entry, and epi- and endophytic mycelial growth in rice and wheat, failed to sporulate to cause rust disease in rice. We identified 18 key defence signalling genes in rice and unravelled their elicitation dynamics in time-course studies during infection. ICS1, NPR1-3, PRs, EDS1, PAD4, FMO1 (salicylic acid [SA] signalling), and ethylene-related genes (ACO4 and ACS6) were strongly elicited in rice. However, genes from the jasmonic acid (JA) signalling pathway (LOX2, AOS2, MYC2, PDF2.2, JAZ8, JAZ10) showed a delayed response during colonization in rice compared to an early or no induction in wheat. However, the JA/ethylene marker gene PDF2.2 was strongly induced in wheat as early as 12 hr postinoculation. Furthermore, rice and wheat displayed specific profiles of accumulation of various phenolic acids during P. graminis f. sp. tritici 40A infection. We propose a model where a differential modulation of the SA/JA-dependent defence network may modulate nonhost resistance. A deeper understanding of the molecular mechanism governing differential elicitation of defence signalling may provide a novel resistance mechanism for the sustainable management of rust diseases.     

The occurrence of Phialophora-like fungi related to Gaeumannomyces graminis under various grass species and some characteristics of these fungi1

Phialophora graminicola and Phialophora sp. (lobed hyphopodia) are weakly or not pathogenic to the cereal and grass roots which they colonize and are also known to restrict the infection of these plants by Gaeumannomyces graminis. The occurrence of these fungi under 15 field-grown grass species has been studied using wheat as the test plant. P. graminicola occurred under all grass species tested but at different population levels. The lowest populations were found under Bromus erectus and B. inermis, and relatively low populations under Agrostis alba, Festuca ovina and Phalaris arundinacea. Festuca pratensis, F. rubra, F. arundinacea, Dactylis glomerata and Arrhenatherum elatius harboured the highest populations of P. graminicola. Phialophora sp. (lobed hyphopodia) was found at low population levels under 7 of the 15 grasses examined. The strains of P. graminicola isolated in these studies differed markedly in growth rate on wheat roots and in restriction of G. graminis on wheat grown in perlite (preliminary results).