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柑橘黑点病研究进展

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柑橘黑点病是由柑橘间座壳菌(Diaporthecitri)引起的一种重要柑橘病害,在全球柑橘主产区均有发生,主要危害叶片、枝梢和果实,严重影响柑橘的品质和鲜销商品价值.柑橘间座壳菌可以侵染所有当前栽培的柑橘品种,其中以葡萄柚和柠檬最易感病.喷施代森锰锌和铜制剂等杀菌剂是防控柑橘黑点病的重要方法,但柑橘间座壳菌在自然条件下有性生殖频繁,具有丰富的遗传多样性,需加强监测柑橘间座壳菌种群对代森锰锌等药剂的敏感性变化.近年来,关于柑橘间座壳菌的基因组信息、快速检测技术、遗传分化、致病机制以及防治方法等方面的研究取得较为显著的进展.就国内外近年来柑橘黑点病的危害症状与分布、病原种类、遗传多样性、生物学特性、侵染过程、致病机制、发生规律以及防治方法等方面的最新研究进展进行综述,并对柑橘黑点病的未来重点研究方向进行展望,以期为柑橘黑点病的防控策略提供科学依据.
Research progress in citrus melanose
Citrus(Citrus spp.)is the world's largest category of fruit,including many species,such as C.reticulata,C.sinensis,C.grandis,C.limon and C.paradisi,etc..They are deeply popular with con-sumers,because of their rich nutrition(vitamins,polysaccharides,organic acids,proteins,dietary fiber and antioxidants)and delicious flavor.The global citrus production for 2019 was estimated at almost 144 million metric tons.In China,the citrus production for 2021 was 55.96 million metric tons,and its annual value of production was more than 200 billion yuan.Therefore,it is the vital major industry of agriculture in China.However,in recent years,the citrus melanose has seriously occurred in major cit-rus production regions all over the world,including China,India,Brazil,Spain and Mexico,etc..In Chi-na,the citrus melanose was widely distributed in Guangxi,Hunan,Hubei,Zhejiang,Jiangxi,Fujian,Yunnan provinces and other major citrus production areas,and the disease incidences in some orchards were up to 100%.The typical disease symptoms including melanose,gummosis and stem-end rot form on leaves,shoots or fruits.The diseased fruits usually show many black to reddish-brown raised spots on the fruit surface or stem-end rot,which seriously affects the appearance and economic value of fresh fruit,causing major economic losses.Diaporthe citri(anamorph:Phomopsis citri)is the dominant spe-cies of citrus melanose pathogen all over the world,which could infect all citrus cultivars.At present,the genome information of D.citri has been sequenced and annotated,which provides an important reference for studying its infection mechanism and population genetic evolution,etc..The genomes of D.citri(MAT1-1 and MAT1-2 strains)contains 15 977-16 622 genes,including 1231-1287 putative pathogenicity genes,1837-1885 secretion proteins,and many carbohydrate-active enzymes(CA-Zymes),which may be associated with the pathogenicity of D.citri.The populations of D.citris are abundant in genetic diversity,due to its frequent sexual reproduction in nature.And the genetic differen-tiation of D.citris is closely related to geographic separation,whereas it has weak correlation with its host.The species-specific primers have been designed for the PCR method to distinguish D.citri from related Diaporthe species,based on the sequences of rDNA internal transcribed spacer,translation elon-gation factor 1-alpha,beta-tubulin,histone H3,and calmodulin gene,which contribute to monitoring and forecasting the citrus melanose in the field.It has been reported that the successful infection host by D.citris is related to its pectinase secretion,and the infection of leaves can cause an increase in the pop-ulation of antagonistic microorganisms in the citrus leaves.The RNA-Seq analysis results performed by Li et al.profiled the defense response pattern of citrus leaves against D.citri infection,including high induction expression of plant cell wall biogenesis-related genes at 3 days post infection(dpi),and high upregulation expression of the CYP83B1 genes,pectin methylesterase gene,and phytoalexin coumarin synthesis-related genes at 14 dpi.After the infected shoots became withered,a large number of alpha co-nidia(non-septate),beta conidia(long,slender)and a small number of ascospores(ellipsoid to cylindri-cal,septate)were produced on dead wood,using as the source of infection.Conidia were carried by raindrops and dispersed to nearby citrus,which contributed obviously to the citrus melanose severity in an orchard,whereas the ascospores were carried by the wind for a long distance spread.Under high hu-midity and warm climatic conditions(at 25 ℃),the young leaves,shoots and young fruits(within 12 weeks after flowering)were seriously infected by D.citri,however the mature citrus tissues were more resistant to this pathogen attack.Therefore,these phenological periods of citrus are also a critical stage for the prevention and control of citrus melanose.The copper fungicide can act as a good preventative against citrus melanose,but it is susceptible to rain erosion,and is phytotoxic to citrus plant when it is used in hot weather.The other pesticides such as mancozeb and strobilurin etc.,play a good control ef-fect on citrus melanose,but they are also facing the risk of increasing resistance to D.citris,because of long-term use of the same fungicides.Some antagonistic microorganisms such as Burkholderia gladioli,Pseudomonas pudia,P.fluorescens,Bacillus subtilis,B.velezensis,B.amyloliquefaciens,Trichoderma asperellum,and T.asperelloides etc.,all play a strong inhibitory effect on the mycelial growth or conid-ia germination of D.citrus,which could provide a reference for commercial application on management approaches of citrus melanose in the field.The above results indicate that,in recent years,some pro-gresses such as species identification and detection,genetic diversity,genomic information,infection cy-cle,pathogenic mechanism,occurrence patterns,prevention and control measures of D.citris have been made by many researchers at home and abroad,but the following issues still need to be further ex-plored.(1)Whether the D.citris forms a special infection structure to successfully penetrate the leaves and peels of citrus with a waxy layer.(2)The pectinase secreted by D.citris is an important virulence factor,but the types,encoding genes and functions of pectinase still need to be clarified.In addition,the presence of other important virulence factors such as toxins,effector proteins etc.,needs to be further analyzed.(3)The propagules of D.citris only form on dead wood,but not on non-dead branches,so the molecular regulation mechanism of asexual spores and ascospores development needs to be studied in D.citris.(4)The community of antagonistic microorganisms increases in the citrus leaves,when they are attacked by D.citris.So,how the citrus plant recognizes the molecular signals of D.citris to regu-late autoimmunity,and the molecular interactions between citrus and D.citris remain to be understood.In conclusion,an in-depth understanding of the infection structure,virulence factors,molecular mecha-nisms of sporogenesis of D.citris,and the molecular signaling pathway of the recognition of D.citris by host will help to provide resources for citrus disease resistance-breeding,and also provide new tar-gets for accelerating the development and application of fungicides for the prevention and control of cit-rus melanose.At the same time,the population of D.citris is abundant,and it is a heterogeneous fun-gus,with frequent sexual reproduction.So it is necessary to strengthen the monitoring of its sensitivity to pesticides such as mancozeb,etc..And these measurements such as mixing pesticides scientifically and reasonably,using biocontrol agents and plant resistance inducers,will reduce the case of fungicide resistance against D.citris,and improve the comprehensive prevention and control ability of citrus mel-anose.

Citrus melanoseDiaporthe citriGenetic diversityPathogenic mechanismOccurrence regularityDisease control

卢松茂、林秀香

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福建省热带作物科学研究所,福建漳州 363001

柑橘黑点病 柑橘间座壳菌 遗传多样性 致病机制 发生规律 病害防治

福建省属公益类科研院所基本科研专项

2018R1028-3

2024

果树学报
中国农业科学院郑州果树研究所

果树学报

CSTPCD北大核心
影响因子:1.486
ISSN:1009-9980
年,卷(期):2024.41(10)
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