Crop rotation sequencing to minimize yield losses of summer-irrigated lowland rice in Myanmar caused by the rice root-knot nematode Meloidogyne graminicola

Abstract

The effect of different rice-based crop rotation sequences on the population densities of Meloidogyne graminicola and on the yield of rice was evaluated in a microplot experiment under lowland rice field conditions. Ten treatments of cropping sequences were grown in four successive growing seasons: continuous growing of the susceptible rice cultivar Thihtatyin (rice-rice-rice-rice), four treatments of 1- season crop rotation sequences (rice-rice-chickpea-rice, rice-rice-black gram-rice, rice-rice-soybean-rice, rice-rice-cowpea-rice), and five treatments of 2-season crop rotation sequences (groundnut-rice-chickpea-rice, green gram-rice-black gram-rice, cowpea-rice-soybean-rice, sesame-rice-cowpea-rice, sunflower-rice-sesame-rice). Population densities of M. graminicola recovered from the rice plants and the nematode multiplication factors (Mf) in the soil in the continuous rice cropping sequence (rice-rice-rice-rice) was the highest among the ten cropping sequences. Lowest nematode Mf in the soil was observed in the 2-season crop rotation sequence sunflower-rice-sesame-rice and the highest was observed in the 1-season crop rotation sequence rice-rice-cowpea-rice among the nine crop rotation sequences. Highest rice yield was obtained in the 2-season crop rotation sequences green gram-rice-black gram-rice, sesame-rice-cowpea-rice and sunflower-rice-sesame-rice, which were about 2 times higher compared with the 1-season crop rotation sequences and about 3 times higher compared with the continuous rice cropping sequence.     

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.     

Pest risk analysis of rice root-knot nematode (Meloidogyne graminicola) for the Italian territory

Meloidogyne graminicola is one of the most harmful organisms associated with rice cultivation throughout the world. Until recently, M. graminicola was present only in Asia, parts of the Americas, Madagascar and South Africa. In 2016, it was detected for the first time in mainland Europe (Northern Italy) and subsequently added to the EPPO Alert List. In this study, the risk posed by this nematode to rice in Italy was assessed using an internationally developed pest risk analysis scheme. In the risk assessment section, information about biology, pathways for entry, the likelihood of establishment, spread and negative impact of M. graminicola are reviewed. Internationally, its spread is more likely through the movement of infested host plants, soil or growing medium and non-host plant parts that may have growing medium attached, from areas where this nematode occurs. Migrant waterbirds, machinery and travellers were also considered as possible pathways for entry. The probability of establishment is very likely due to the suitable environmental conditions and the large range of host plant species present in Italy. This and the nematodes’ ability to survive for long periods in low oxygen environments make the control of M. graminicola very difficult.     

氟吡菌酰胺不同施药方式对水稻拟禾本科根结线虫的防治效果

为评价41.7%氟吡菌酰胺悬浮剂防治水稻拟禾本科根结线虫Meloidogyne graminicola的应用潜力,将氟吡菌酰胺与吡虫啉种衣剂混合后采用种子包衣法及喷洒法研究其对水稻拟禾本科根结线虫的田间防效,并测定了不同施药方法对水稻生长和产量的影响。结果表明,氟吡菌酰胺4.2、8.3、12.5 g（a.i.）/kg种子与吡虫啉18.0 g（a.i.）/kg种子混合包衣处理,播种后35 d其根结抑制率和防效分别为41.0%~51.8%和47.4%~58.6%,土壤中2龄幼虫减退率为38.6%~40.4%,显著高于单施吡虫啉18.0 g（a.i.）/kg种子处理。水稻播种后连续3次以氟吡菌酰胺250.2、375.3、500.4 g（a.i.）/hm²进行土壤喷洒,最后1次施药后7 d,其根结抑制率和防效分别为81.0%~89.9%和65.9%~74.3%,土壤中2龄幼虫减退率为65.4%~73.4%,均显著高于对照药剂克百威1 800.0 g（a.i.）/hm²处理。氟吡菌酰胺各处理对水稻苗期生长均有较好的保护作用,能显著提高千粒重和有效穗数,产量比空白对照增加50.0%~61.2%,保产效果显著。水稻播种后35 d采用氟吡菌酰胺对稻田进行1次喷洒,对水稻具有一定的保护作用,保产效果不明显。表明采用氟吡菌酰胺与吡虫啉混合包衣种子处理及在水稻苗期进行喷洒处理对水稻拟禾本科根结线虫防效显著,具有显著的保产效果。     

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。     

Rice nematode problems in Nigeria: Their occurrence,distribution and pathogenesis

Abstract

In a survey of plant parasitic nematodes associated with or affecting rice throughout Nigeria, some important nematode pests, especially the white tip disease nematode, Aphelenchoides besseyi and the rice root nematodes Hirschmanniella spinicaudata and H. oryzae were identified from seed, soil and root samples from swamp rice fields respectively. The sugarcane cyst nematode, Heterodera sacchari occurred in swamp rice fields only around the major sugarcane estates in Nigeria. The root‐knot Meloidogyne incognita and the root lesion nematode, Pratylenchus brachyurus were also encountered in upland (rainfed) rice fields. The white tip disease nematode, A. besseyi occurred at low levels in soils and rice seeds throughout the country. High population levels of H. spinicaudata and H. oryzae were encountered especially in areas where monoculture of rice is practised. General chlorosis, poor tillering and significantly reduced yield have been observed due to H. spinicaudata. Rice plants attacked by H. sacchari also showed intense chlorosis, delayed and reduced tillering and reduced grain yield. The roots of attacked plants were twiggy, very necrotic and blackened. The root‐knot M. incognita and the root lesion nematodes P. brachyurus have both been observed to reduce rice yields. Rice cultivars screened for reactions to the nematodes showed varying degrees of susceptibilities. Some varieties were however resistant to the root‐knot nematode, M. incognita.     

Differential behaviour of sheath blight pathogen Rhizoctonia solani in tolerant and susceptible rice varieties before and during infection

This study characterized the early infection and establishment of the sheath blight pathogen Rhizoctonia solani on a tolerant rice variety, Swarnadhaan (IET 5656), and a susceptible variety, Swarna (MTU 7029). Assays using whole plants showed that disease severity was higher in Swarna than Swarnadhaan. In a detached leaf assay, Swarnadhaan showed a disease index that was 50% less than that with Swarna. Rhizoctonia solani exhibited different growth behaviour in the tolerant and susceptible varieties. The pathogen showed more hyphal growth in the susceptible host than in the tolerant variety. It also showed profuse branching, making intimate contact with the host surface to form more inter‐ and intracellular structures, and greater sclerotial development in the susceptible host compared to the tolerant one. Using light and scanning electron microscopy, it was observed for the first time that the pathogen could intercept host surface structures and use these for anchorage or penetration. Transformed R. solani, expressing green fluorescent protein, was observed using confocal laser scanning microscopy to investigate pathogen behaviour, including the formation of infection cushions and subsequent colonization of the host tissues. This is the first ultrastructural report to characterize the differential behaviour of the sheath blight pathogen in the vicinity and within tolerant and susceptible rice plants.     

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.     

Virulence structure of theMagnaporthe grisea rice population from the northwestern Himalayas

The virulence structure of theMagnaporthe grisea rice population from the northwestern Himalayan region of India was deciphered on 24 rice genotypes harboring different blast resistance genes. Matching virulences appropriate to all the rice genotypes, except Fukunishiki (Pi-z, Pi-sh) and Zenith (Pi-z, Pi-a, Pi-i), were present in the pathogen population. Moreover, a very low percentage of isolates were virulent on Tetep (Pi-ta, Pi-k ^h, Pi-4^b) and Tadukan (Pi-ta/Pi-ta ²). Although virulence was recorded on most of the lines tested, none was susceptible to all of the isolates. Three pairs of genotypes, namely, C101LAC:C101A51; K-1: Dular; and Dular: HPU-741, exhibited complementary resistance spectra as no isolate combined virulence to both the members of each of the three pairs of genotypes despite the fact that individual members were susceptible to a major portion of the pathogen population. The blast resistance genesPi-z, Pi-k ^h, Pi-l andPi-2 and their various combinations were construed to provide broad spectrum and durable blast resistance in Himachal Pradesh. Pathotype analysis revealed the existence of extremely high pathotypic diversity in the pathogen population. Based on the observed population structure forM. grisea, it was not possible to designate a minimum set of pathogen isolates that could be used in blast resistance screens to identify effective sources of blast resistance. The overall results suggested that the pathotype analysis alone is insufficient to describe the existing pathogenic variability, especially when this information has to be used for guiding the breeding programs aimed at developing durable blast resistance. However, population genetics approach of studying pathogenic specialization by monitoring the frequency of individual virulence genes and analyzing virulence gene combinations for their association or dissociation might generate useful information for developing durable blast resistance. http://www.phytoparasitica.org posting May 14, 2006.     

Investigation of rice blast development in susceptible and resistant rice cultivars using a gfp‐expressing Magnaporthe oryzae isolate

In this study, an isolate of Magnaporthe oryzae expressing the green fluorescent protein gene (gfp) was used to monitor early events in the interaction of M. oryzae with resistant rice cultivars harbouring a blast resistance (R) gene. In the resistant cultivars Saber and TeQing (Pib gene), M. oryzae spores germinated normally on the leaf surface but produced morphologically abnormal germ tubes. Germling growth and development were markedly and adversely affected in leaves of these resistant cultivars. Penetration of host cells was never seen, supporting the idea that disruption of germling development on the leaf surface might be one of the resistance mechanisms associated with Pib function. Thus, this particular R gene appeared to function in the absence of host penetration by the fungal pathogen. Confocal laser scanning microscopy of M. oryzae‐infected susceptible rice cultivars showed the dimorphic growth pattern that is typically observed during the biotrophic and necrotrophic stages of leaf colonization in susceptible cultivars. The suitability of the gfp‐expressing M. oryzae isolate for further research on R‐gene function and identification of resistant genotypes in rice germplasm collections is discussed.     

云南省稻瘟病菌群体的致病性分析及交配型分布

为明确云南省不同稻区稻瘟病菌Magnaporthe oryzae的毒性频率及交配型分布,利用2007—2013年分离自云南省不同稻区的112株稻瘟病菌单孢菌株,对23个持有不同抗性基因的单基因系和持有Pi57(t)的水稻渗入系IL-E1454进行致病性测定。结果表明,稻瘟病菌对不同抗病基因的毒性频率存在很大差异,分离自粳稻区的稻瘟病菌菌株对持有Piz-t、Pi5、Pi9、Pi20和Pi57(t)这5个水稻品系的毒性频率分别为14.29%、5.36%、5.51%、5.36%和0;分离自籼稻区的稻瘟病菌菌株对持有Pik-h、Piz、Pita、Piz-5、Pita-2、Pi5、Pi7和Pi9这8个水稻品系的毒性频率分别为18.25%、9.13%、9.64%、7.50%、15.72%、0、13.05%和0;分离自陆稻区的稻瘟病菌菌株菌株对持有Pik-h、Pib、Pish、Pi1、Pi5、Pi9、Pi11和Pi57(t)这8个水稻品系的毒性频率分别为6.67%、3.33%、13.79%、13.33%、7.69%、6.67%、0和3.23%;交配型测定结果显示,陆稻区菌株可交配率为100.00%,...     

Genetic diversity of Meloidogyne spp. from rice and identification of multiresistant sources in Oryza spp. accessions

Recently a Meloidogyne species complex was detected parasitizing and causing damage to irrigated rice in southern Brazil, highlighting the need to study the genetic diversity of these species and their pathogenicity to Oryza spp. in order to select genotypes of rice with multiple resistance. This study compared the genetic diversity of Brazilian Meloidogyne spp. isolates from irrigated rice and evaluated the reaction of four wild accessions of Oryza species (O. glumaepatula, O. longistaminata, O. grandiglumis, and O. alta) and two cultivated species, O. glaberrima and O. sativa (control) to M. ottersoni, M. oryzae, and two variants of M. graminicola (Est G2 and Est G3). Genetic variability was assessed using RAPD and AFLP markers. M. graminicola and M. ottersoni showed high intraspecific variability: 83.76% and 41.14%, respectively. Cluster analysis showed a clear separation among rice root-knot nematodes (RKNs) into subclades according to their esterase phenotypes with 100% bootstrap. For rice resistance screening, plants were inoculated with 5,000 eggs, and the nematode reproduction factor evaluated 90–120 days postinoculation. O. glumaepatula, an American wild species, was highly resistant or resistant to all rice RKNs tested and is a valuable source of multiple resistance. Overall, the other rice species also showed different levels of resistance. Conversely, O. longistaminata exhibited low levels of resistance. M. graminicola Est G3 was the most aggressive isolate. Sources of resistance against RKN in wild Oryza genotypes, especially in an AA genome like O. glumaepatula, may be of great interest for future breeding programmes in cultivated rice.     

Microscopic analysis of the effect of azoxystrobin treatments on Mycosphaerella graminicola infection using green fluorescent protein (GFP)‐expressing transformants

Green fluorescent protein (GFP)‐expressing transformants were used to investigate the effects of strobilurin fungicide azoxystrobin on Mycosphaerella graminicola infection. Azoxystrobin treatments (125 or 250 g AI ha^?1) were applied at various stages of the infection process under controlled conditions. GFP transformants showed conserved in vitro sensitivity to azoxystrobin and pathogenicity. Azoxystrobin controlled over 90% of M graminicola infections when applied before or during penetration of the pathogen (15% of the incubation phase). Azoxystrobin also impaired the growth of intercellular hyphae in M graminicola post‐penetration infection stages when applied at up to 50% of the incubation phase. Incubating infections observed in treated leaves were viable, but their growth was impaired and they did not induce necrosis under controlled conditions. Reduction by half of azoxystrobin dosage had little or no effect on azoxystrobin efficiency in controlling M graminicola. The contribution of post‐penetration fungistatic effect to azoxystrobin curative properties toward M graminicola in a field situation is discussed. © 2001 Society of Chemical Industry     

Physiological basis of bentazon tolerance in rice (Oryza sativa L.) lines

In the present study, 98.8 mm of bentazon was applied to 3‐leaf stage rice seedlings. Two tolerant lines, M202 and cv. TNG67, showed slightly visible injury, photosystem II inhibition, as well as a low level of lipid peroxidation 7 days after application compared with the susceptible lines. Further physiological study of the mechanism of differential tolerance among Japonica and Indica types indicated that, although the tolerant Japonica lines M202 and cv. TNG67 absorbed more ¹⁴C‐bentazon, most of the ¹⁴C remained in the treated leaf or translocated to older leaves. However, two susceptible lines, FSK (Japonica) and IR36 (Indica), absorbed less ¹⁴C‐bentazon throughout the experiment, and most of the ¹⁴C was translocated from the treated leaves to younger leaves, which might result in the death of developing tissues. In addition, more bentazon residue and less polar metabolites were detected in these two susceptible lines. It is proposed that the higher tolerance of lines M202 and TNG67 to bentazon could be mainly due to a higher rate of metabolism of this herbicide, and partially due to less translocation to developing tissues.     

Sublethal infection of different pine species by the pinewood nematode

Bursaphelenchus xylophilus, the pinewood nematode and causal agent of pine wilt disease (PWD), is a globally destructive pine pest. PWD is favoured by high temperatures and drought, but B. xylophilus can also persist in symptomless hosts in areas not prone to PWD. We compared seedlings of different Pinus species that were inoculated with B. xylophilus when well-watered and under moderate temperatures, conditions that favour sublethal infections. We compared isolates of B. xylophilus in south-west Europe (Portugal) and south-east USA (Louisiana). P. pinea, P. pinaster, and P. radiata were challenged in both areas with inoculations of local B. xylophilus. P. sylvestris in Portugal and P. palustris in Louisiana were also tested. Seedling mortality was low, as expected, but nematode establishment was common. There were higher nematode densities and more damage in seedlings of P. sylvestris and P. radiata, followed by P. pinaster, with P. pinea and P. palustris proving to be more tolerant. Life history of hosts was a better predictor of resistance to B. xylophilus than geographical origin, even under sublethal conditions. Fast-growing species (P. radiata and P. pinaster) appeared most likely to be symptomless carriers in areas not prone to the development of PWD. Cold-adapted species (P. sylvestris) may still be suitable hosts for the pathogen, even at the sublethal conditions of their distribution areas. There were more nematodes per gram of host using nematode isolates from Portugal. Nonetheless, our results opposed the hypothesis that North American pinewood nematodes have such low virulence that they are obligatorily saprophytic.     

Population structure and species delimitation of rice white tip nematode,Aphelenchoides besseyi (Nematoda: Aphelenchoididae), in China

Aphelenchoides besseyi is a major nematode pathogen in rice known as the causal agent of white tip disease. In the present study, A. besseyi was recovered from 32 seed samples collected from the major rice-growing regions in China. Nine mitochondrial COI haplotypes and 95 ITS genotypes were identified, suggesting a high genetic diversity and endemism level of A. besseyi in China. Therefore, A. besseyi is likely to have experienced a long history of host–parasite coevolution in China, rather than being recently introduced from other countries. The results of population structure analysis indicated the presence of four clusters within Chinese A. besseyi, but these did not correspond to their geographic distribution. Haplotype H2 was found to be the most widespread. Haplotypes H9 and H13 were most genetically divergent and have been recognized as two cryptic species by molecular species delimitation methods, and with limited support from morphometric measurements. The COI-based phylogeny suggested the ability to parasitize rice has independently evolved at least four times in the genus Aphelenchoides.     

中国北方杂草稻幼苗对干旱胁迫的生理响应

以杂草稻heb07-2、wr04-6及巴西陆稻IAPAR9为研究材料,采用人工气候箱培养幼苗,利用20%聚乙二醇6000（PEG-6000）模拟干旱条件,研究了杂草稻及巴西陆稻幼苗叶片和根系生理特性标的变化。结果表明：干旱胁迫下,杂草稻heb07-2幼苗的叶片与根系具有较高的超氧化物歧化酶（SOD）、过氧化氢酶（CAT）、抗坏血酸过氧化物酶（APX）活力;杂草稻heb07-2的可溶性糖含量、可溶性蛋白含量高于其它材料。杂草稻heb07-2细胞膜系统保持完整,膜质过氧化程度小于其它材料,表明干旱胁迫对杂草稻heb07-2伤害较小,具有较强苗期抗旱性。     

Susceptibility to imazamox in Italian weedy rice populations and Clearfield® rice varieties

The introduction of imidazolinone‐tolerant rice varieties has made selective Oryza sativa (weedy rice) control possible. We hypothesised that Italian weedy rice populations have variable degrees of susceptibility to imazamox prior to imidazolinone‐tolerant variety introduction. To this end, 149 Italian weedy rice populations collected from fields never before cultivated with imidazolinone‐tolerant varieties were tested in a glasshouse‐based, whole‐plant response screening study. Imazamox was applied to all populations post‐emergence at a rate of 70 g a.i. ha^?1, resulting in 70–90% shoot biomass reduction in the majority of cases. The results prompted a second study of the seedling dose response of four weedy rice populations from the initial study group. Three imidazolinone‐tolerant and one conventional rice variety were also included. The seedling roots were cut six days after germination and exposed to different concentrations of imazamox. The root regrowth associated with each concentration‐exposure was then measured. Imazamox concentrations to inhibit weedy rice root growth by 50% varied by about two orders of magnitude, or between 0.0018 and 0.12 mm . Even with this result, imidazolinone‐tolerant varieties were at least 31.8 times less susceptible than weedy rice populations, suggesting that Italian weedy rice populations were not tolerant to imazamox before introduction of these varieties.     

Rice response to simultaneous bacterial blight and drought stress during compatible and incompatible interactions

Plant response to one type of stress can be affected by simultaneous exposure to a second stress, for example when abiotic and biotic stresses occur together. Ten rice genotypes comprising those with bacterial blight (BB) resistance (R) genes, drought quantitative trait loci (QTLs) plus a BB R gene, and BB susceptible genotypes, were subjected to mild and moderate drought stress and plants were inoculated with two Xoo strains (PXO99 and PXO145) to simulate the challenges rice crops face under simultaneous stress of drought and BB. Plant height and dry shoot biomass were significantly reduced by drought stress treatments. The BB disease lesion lengths varied according to rice genotypes and PXO99 Xoo multiplication and spread in planta was higher compared to that of PXO145, which generally decreased under mild drought stress. Rice genotype IRBB7 (Xa7) showed less Xoo spread and a reduced Xoo multiplication under drought stress compared to the well-watered control with PXO145. In contrast, in genotypes with a different BB R gene and/or drought QTLs [IRBB4 (Xa4), IR87705–6-9-B (Xa4 + qDYT _2.2 ), IR87707–445-B-B-B (Xa4 + qDYT _2.2  + qDYT _4.1 ) and IR87707–446-B-B-B (Xa4 + qDYT _2.2  + qDYT _4.1 )], Xoo multiplication and spread in planta was higher with drought stress. This study has shown that drought stress affected rice response to the BB pathogen and the response varied according to the rice genotype. It is concluded that evaluating rice varieties under combined abiotic and biotic stresses will be the best strategy to determine biotic stress resistance durability under climate change.     

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.