Int.J.Curr.Microbiol.App.Sci (2018) 7(8): 4006-4023
International Journal of Current Microbiology and Applied Sciences ISSN: 2319-7706 Volume 7 Number 08 (2018) Journal homepage: http://www.ijcmas.com
Review Article https://doi.org/10.20546/ijcmas.2018.708.415
Arthropods as Vector of Plant Pathogens viz-a-viz their Management
Ravinder Singh Chandi, Sanjeev Kumar Kataria* and Jaswinder Kaur
Department of Entomology, Punjab Agricultural University, Ludhiana-141 004, Punjab, India
*Corresponding author
ABSTRACT
An insect which acquires the disease causing organism by feeding on the diseased plant or by contact and transmit them to healthy plants are known as insect vectors of plant diseases. Most of the insect vectors belong to the order Hemiptera, Thysonaptera, Coleoptera, Orthoptera and Dermaptera. Homopteran insects alone are known to transmit K e yw or ds about 90 per cent of the plant diseases. About 94 per cent of animals known to transmit plant viruses are arthropods. On the basis of the method of transmission and persistence in Insect vectors, Plant pathogens, the vector, viruses may be classified into three categories viz. non-persistent, semi Management persistent and persistent viruses. Irrespective of the type of transmission, virus-vector relationship is highly specific and spread of vector borne diseases also depends upon Article Info potential of vector to spread the disease. Also for transmission of virus, activity of insect vectors is more important rather than their number. There is a high degree of specificity of Accepted: 22 July 2018 phytoplasma to insects and interaction between these two is complex and variable. A number of plant diseases caused by bacteria are known to be transmitted by insects Available Online: 10 August 2018 because many of these insects are actually attracted by the sugars contained in the bacterial exudates. There are several insects associated with the spread of fungal diseases. The
common sooty mold fungus grows on the honeydews excreted by several homopteran insects. For the management of various types of plant diseases transmitted by insects, integrated management strategies need to be adopted to manage the vectors. Introduction by contact and transmit them to healthy plants are known as insect vectors of plant diseases. Vector is the most important component of Insect vectors include mainly aphids, virus disease epiphytology. A vector is an leafhoppers, whiteflies, thrips, psyllids, organism capable of transmitting pathogens beetles, mealybugs etc. from one host to another. It helps in spread of virus from infected plant to the healthy plant. Plant pathogens are transmitted by a number Plant viruses can produce direct and plant- of ways like by contact, by contamination mediated indirect effects on their insect through soil or other biological agencies. Most vectors, modifying their life cycle, fitness and of the plant diseases are transmitted by insects behavior (Delafuente et al., 2013). All those and arthropods like mites and few of them are insects which acquires the disease causing transmitted by mechanical means or organism by feeding on the diseased plant or contamination of the soil. Insects damage
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Int.J.Curr.Microbiol.App.Sci (2018) 7(8): 4006-4023 plants in many ways, but the most common These insects do not cause wholesome type of damage is the removal of plant tissues, destruction of cells during feeding and viruses as in the familiar examples of caterpillars, require living cells for their subsistence and beetles, and grasshoppers whose feeding multiplication. creates noticeable holes or even the wholesale removal of leaves, fruits, or other plant parts. Viruses Insect having piercing and sucking mouthparts and biting and chewing mouthparts are A virus is infective agent that typically associated with disease transmission. Fereres consists of a nucleic acid molecule in a protein and Moreno (2009) reported that most of the coat, is too small to be seen by light plant viruses are transmitted by insect vectors, microscopy, and is able to multiply only thus depend on their behavior, transmission within the living cells. Viruses are responsible and dispersal capacity to move from plant to for many diseases in man (influenza, measles, plant and to spread to distantly-located mumps, polio, pox, etc.) and plants (mosaic, regions. Hemipterans, (aphids, whiteflies and leaf curl, etc). Plant viruses are minute hoppers), involve most of the phytopathogenic parasitic which infect plant cells, altering their virus vectors. Transmission of plant viruses is chemistry and causing a wide range of mediated by the piercing-sucking mouthparts symptoms including discolouration, distortion of these insects (stylets), when penetrate and loss of vigour and yield. Plant virus through the intercellular spaces and establish diseases have become more prevalent and feeding sites in phloem sieve elements destructive in recent years. This is mainly (Forbes, 1969). because of better recognition of the virus diseases, exchange of plant material from Most of the insect vectors belongs to the order region to region facilitating spread of the virus Hemiptera (aphids, leafhoppers, whiteflies and to new areas, and distribution of many insect mealybug), but a few others belong to vectors in new regions in the world. Thysanoptera (thrips), Coleoptera (beetles), Orthoptera (grasshoppers) and Dermaptera Phytotoxemia is a disease like plant condition (earwigs). Homopteran insects alone are produced by toxic substance injected by known to transmit about 90 per cent of the insect. It is the production of plant symptoms plant diseases. A number of plant disease of distress caused by the reaction of plant to caused by viruses, phytoplamas, bacteria and the toxins produced by insect feeding. The fungi are transmitted by insects. The salient feeding of insects and mites especially those features of homopterans (aphids, leafhoppers) that suck sap from the plant can cause which make them efficient vectors are as symptom of distress such as yellowing, follows: silvering, bronzing, necrosis, wilting and discolouration of the shoots and malformation They make brief but frequent probes with their of leaves, stem, roots, fruit and other plant mouthparts into host. organ or tissues.
As the population density reaches a critical The great majority of insect species level, winged migratory individuals are responsible for phytotoxic effect on plants produced. belong to Hemiptera. Some mites are also responsible for the phytotoxaemia and some In many species, winged females deposit a gall forming insects are also found in other few progeny on each of the many plants. insect order.
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About 94 per cent of animals are known to produces one of the most commonly seen galls transmit plant viruses are arthropods and 6 per found on the leaves of silver maple; this cent are nematodes. Nearly 99 per cent of the condition has attracted particular attention arthropods vectors are insects and 55 per cent because of the conspicuous galls and the wide of these are aphids. Most of the insect vectors distribution of the host plant in the various (90%) belong to the order Homoptera and states. The eruptions on the upper leaf surface about half of the insect vectors are aphids. The are initially green, later on becoming tinted main aphid vectors are Myzus persicae with pink and red colour (Drake et al., 2005). (Sulzer), Aphis gossypii Glover and Aphis craccivora Koch. In addition, whiteflies and Wheat streak mosaic virus (Family: leafhoppers are also responsible for Potyviridae; Genus: Tritimovirus) is a virus transmission of plant viruses. Whiteflies vectored by wheat curl mite (A. tosichella), mostly transmit mosaics and leaf curls in which is the most common virus infecting pulses, vegetables and other crops like cotton, wheat. The spider mite T. urticae is a tobacco and papaya (Table 1). parenchym cell content feeder; the species pierces parenchym cells and consumes their The leaf and plant hoppers transmit tungro, contents, producing significant leaf damage yellow-orange leaf, grassy leaf, grassy stunt (Kant et al., 2008). The Aceria mangiferae and ragged stunt in rice. Tomato spotted wilt (Family: Eriophyidae) produces distortion, is known to be transmitted by thrips. stunting and bud proliferation of new growth Mandibulate insects like grasshoppers, on plants, and the mites may transmit mango earwigs and chrysomelid beetles transmit malformation disease (Gamliel et al., 2009). turnip yellow mosaic. Several species of mites are also responsible for transmission of viruses Types of viruses of cereals and fruit crops (Table 2). There are thousands of species of mites and spider. The The transmission process of vector-borne mites are small arthropods, which can be red, viruses is categorized by two features: (1) the green brown or cream in colour. time period required by the vector for acquisition of the virus and inoculation of the They live in associations, typically on the virus, and (2) the retention time of viral lower side of leaves. Spider mites can particles in the vector (Ng and Falk, 2006). On replicate speedily in hot and dry climate. After this Basis, virus-vector relationships can be mating, females continuously produce as categorized as non-persistent, semi-persistent, many as 300 eggs completed in a twosome of or persistent. For non-persistent viruses, weeks. Mites can overwinter in more than a transmission can occur within minutes of few stages of development, like as eggs acquiring the viral particles (virions) and deposited near dormant buds and as adult particles are retained in the stylet or in the females under rough bark scales or ground alimentary canal of the insect (Ng and Falk, litter (Sarwar, 2014). Fascinatingly, high 2006; Uzest et al., 2007; Whitfield et al., humidity can really decrease mite numbers. 2015). Viral particles can be lost quickly in this transmission mode and multiple The wheat curl mite, which is an eriophyid, encounters with infected plants are required has been incriminated as the vector of two for vectors to remain viruliferous (Ng and viruses of wheat. Other species are also known Falk, 2006). Viruses can be of non- to transmit at least 10 other disease-causing persistence, semi persistence and persistence plant pathogens. The maple bladder gall mite type.
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Non persistent viruses Persistence viruses
These viruses are also called stylet borne Persistent viruses are those that persist longer viruses. These are those viruses which are within the infective agent. i.e. vector. These believed to be transmitted as contaminants of viruses, when acquired by a vector, pass the mouthparts. The method of transmission of through the midgut wall to the salivary glands this kind of viruses is mechanical. Acquisition from where they can infect new hosts. In case feeding time is very short, test feeding period of these viruses, insect feeds for longer time is very short and virus do not multiply in the on the source of virus. vector is a true about non-persistence transmission of virus. The insect becomes infective after a certain period of acquisition of virus ranging from These viruses do not persist longer within the several hours to 10-20 days, which is called insect vectors which can transmit them soon incubation period or latent period. after feeding on infected plant but the ability to transmit fresh infection soon disappear after Such viruses may multiply within tissues of a the insect feeds on healthy or immune plants vector, which retains the ability to transmit the e.g. mosaic-type diseases. Cucumovirus, virus for several days and in some instances, carlaviruses and potyviruses are aphid borne the rest of its life e.g. virus transmitted by stylet borne viruses. Aphids are the vectors of whitefly is persistent and circulative type. a great majority of such viruses which are carried only in their stylets. Following are the Mechanism of transmission main features of the non-persistence viruses: For inoculation of virus into a plant by Vectors are optimally infective when they sucking insects, the puncture is transmitted by have fed for approximately 30 seconds on the a number of forward and backward infected plant. movements of the inner pair of stylets. Fluid flows into them with forward movements and Transmission is improved if vector is starved saliva ejected during backward movements. for a period before an infective feed. Generally, an insect injects by feeding on any If the vector is starved after an acquisition, it part of the plant, but in some cases the virus is loses ability to transmit within 2 minutes. only found in the phloem and has to be injected into the phloem, the movement of Semi-persistence viruses which is perhaps controlled by the pH gradient between the mesophyll and the phloem. Many These viruses are carried in the anterior insects facilitate the entry of a pathogen into regions of the gut of a vector, where they may its host through the wounds and insects make multiply to a certain extent. Vectors do not holes in aboveground and belowground parts normally remain infective after a molt, of the plants (Agrios, 1997). presumably because the viruses are lost when the foregut intima is shed. Several of the Grasshoppers and beetles insects regurgitate leafhopper transmitted viruses fall under this with mandibulate during feeding. The category e.g. mosaic and yellows type regurgitated fluid containing the virus is diseases. Beet yellows virus is transmitted by brought into contact with the healthy plant, aphids, A. craccivora. thus transmitting the virus.
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Virus-vector relationship number of viruses can be transmitted through pollen to the seed (e.g. Barley stripe mosaic Irrespective of the type of transmission, virus- virus, genus Hordeivirus) while many that vector relationship is highly specific. cause systemic infections accumulate in Generally one type of virus disease is vegetatively propagated crops. Aphids transmitted only by insects belongings to one transmit viruses from many different genera, particular group, i.e. mosaic by aphids and leaf including Potyvirus, Cucumovirus and curl by whitefly. Single virus can be Luteovirus with the help of green peach aphid transmitted by more vectors. This depends M. persicae, the vector of many plant viruses, upon the specificity of vector, for example including Potato virus Y. The aphids, cucumber mosaic virus is transmitted by 2 Pentalonia nigronervosa Conquerel, transmit vectors i.e M. persicae and A. gossypii. banana bunchy top and cardamom mosaic. Sometimes one vector can transmit more than Similarly, the whiteflies transmit viruses from one virus. M. persicae can transmit different several genera like okra yellow vein mosaic, viruses like potato leaf roll virus, potato virus dolichos yellow mosaic, tomato leaf curl, Y. The capability of vector to transmit the papaya leaf curl, but particularly those in the virus is an important step to know its genus Begomovirus. B. tabaci, the vector of potential, as different vectors transmit the many viruses including tomato yellow leaf viruses in variable proportion. curl virus and lettuce infectious yellows virus. The hoppers transmit viruses from several Vector population and activity genera, including those in the families Rhabdoviridae and Reoviridae. Thrips The vector density must be affecting the virus transmit viruses in the genus Tospovirus and spread by influencing vector movement and Frankinella occidentalis, the western flower associated activity. The understanding of thrips that is a major vector of Tomato spotted vector abundance and vector movement is wilt virus. For transmission of virus, activity very important before monitoring of of insect vectors is more important rather than population density. Vector population depends their number. In case of aphids, it is the on environment factors, host type and growth activity and number of migrant insects that is stage. important in the efficiency of virus transmission rather than the number of Host seeking activity apterous individuals which are, of course, important in respect of their direct injury to Without host seeking activity by the vector, the crop. the virus cannot be carried to potential non infected host plant. Host seeking activity is Aphids very helpful in the transmission of virus. Host seeking activity involves finding a suitable The majority of viruses infecting plants are host plant, landing on the host plant and spread by insects, and aphids are the most probing the plant by feeding on parenchyma common group of virus vectors or carriers. All tissue. All the plant cells have a robust cell potyviruses (the largest group of plant viruses) wall and viruses cannot penetrate them are transmitted by aphids. Aphids are sap- unaided. Mostly plant viruses are therefore sucking insects and have piercing, sucking transmitted by a vector organism that feeds on mouthparts. Their mouthparts include a the plant or (in some diseases) are introduced needle-like stylet that allows the aphid to through wounds made during pruning. A small access and feed on the contents of plant cells.
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During feeding, aphids simultaneously ingest BYDV infected than on non-infected plants. sap contents and inject saliva, which can Conversely, the infection of soybean with contain viruses if the aphid has previously fed alfalfa mosaic, soybean mosaic and bean pod on an infected plant. Transmission of a virus mottle viruses, inoculated separately, had by insects is a specific biological process. A several negative effects in the aphid Aphis particular virus is transmitted by one carrier glycines performance, decreasing the aphid group only for example aphids or whiteflies, population growth rate and the aphid density not both. The two broad categories of aphid on infected plants (Donaldson and Gratton, transmission are: non-persistent and persistent 2007). or circulative. These terms relate to the length of time an aphid takes to acquire and then Cucumber mosaic virus (CMV) is one of the transmit a virus, and the length of time the most common virus pathogens of plants, and aphid remains capable of transmitting the infects over 1,000 species (Roossinck, 2002). virus. The genus of CMV is cucumovirus in the family bromoviridae (Palulaitis et al., 1992), Plant viruses may also interact directly with is a multicomponent virus consisting of three their insect vectors, altering their behavior to genomic single-stranded RNA’s each enhance their own spread. Recently, Stafford encapsidated individually in a 28 nm diameter et al., (2011) found that tomato spotted wilt icosahedral particle. CMV is aphid transmitted virus (TSWV) directly modifies the feeding in a stylet-borne non-persistent manner (Nault, behavior of its vector Frankliniella 1997). The components of the aphid stylet occidentalis increasing its ability to transmit interact directly with virus capsid protein to the virus. Ingwell et al., (2012) found that influence transmission efficiency (Chen and after acquisition of barley yellow dwarf Francki, 1990 and Perry et al., 1998), and do viruses (BYDVs) the settling behavior of not utilize a virus encoded helper component aphid Rhopalosiphum padi changed. Aphids protein, as do non-persistently transmitted that acquired the virus preferred to feed on potyviruses. non-infected wheat plants, while non-infective aphids preferred BYDV infected plants. Most CMV strains are currently divided between two major groups (subgroup 1 and BYDVs are economically important viruses subgroup 2) based on serological and that infect cereal crops worldwide, including sequence similarities (Andersom et al., 1995, wheat grown in China. These viruses are Hsu et al., 2000). Common strains of CMV transmitted only by aphids and include several typically infect solanaceous and cucurbit members of the luteoviridae (Du et al., 2007). crops, but not legume species. Legume- BYDVs are naturally transmitted by at least infecting strains of CMV isolated from 25 aphid species in a highly specific, infected peas were first described by Whipple circulative, non-propagative manner, and have and Walker (1941). These strains were not been found infecting agronomically important seed transmitted, but were transmissible by the small grains (Gray and Gildow, 2003). green peach aphid. Another virus-vector mutualism relationship was found when the aphid Sitobion avenae, Whitefly fed on BYDV infected wheat (Fereres et al., 1989). Aphids had a significantly shorter Whitefly, Bemisia tabaci (Hemiptera: developmental time, a greater fecundity, and a Aleyrodidae) a complex of morphologically greater intrinsic rate of natural increase on indistinguishable species, are vectors of many
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Int.J.Curr.Microbiol.App.Sci (2018) 7(8): 4006-4023 plant viruses (Polston and Capobianco, 2013). TYLCV-B. tabaci complex. Whitefly transmitted plant viruses included several genera (Begomovirus, Carlavirus, B. tabaci is a destructive pest of horticultural Crinivirus, Ipomovirus, and Torradovirus) and crops and ornamental plants worldwide also include several hundred species of (Brown et al., 1995 and Dalton, 2006) emerging and economically significant especially because of its role as a vector of pathogens of important food and fiber crops. plant viruses (Brown and Czosnek, 2002) and These viruses do not replicate in their vector (Hogenhout et al., 2008). Actually, B. tabaci but move readily from plant to plant by the is a species complex containing at least 28 adult whitefly by various means. For most of morphologically indistinguishable species (De these viruses whitefly feeding is required for Barro et al., 2011 and Liu et al., 2012). acquisition and inoculation, while for others Among these, the ‘Mediterranean’ putative only probing is required. species (Liu et al., 2012 and Sun et al., 2011) has been previously referred as the Q biotype Navas et al., (2011) found that most of the and is the most extended in Spain. begomoviruses (Family: Geminiviridae) are transmitted by whiteflies, although these are Mutualism relationships between plant viruses also vectors of criniviruses, ipomoviruses, and their insect vectors have been observed in torradoviruses and some carlaviruses. Rosen et some Hemipterans. In the case of B. tabaci al., (2015) studied that the begomoviruses biotype B and tomato yellow leaf curl China comprise an emerging and economically virus (TYLCCNV) interactions, Jiu et al., important group of plant viruses exclusively (2007) found that fecundity, longevity and transmitted by the B. tabaci in many regions population density of whiteflies on virus- of the world. Begomoviruses is one of the infected tobacco plants increased. Zhang et largest and economically important group of al., (2012) explained that this mutualism is a emerging insect vector. Begomoviruses are consequence of changes in plant defense exclusively vectored by B. tabaci in a responses after begomovirus infection. In fact, persistent circulative manner. the infection by TYLCCNV interferes with the synthesis of jasmonic acid that is utilized by Moreno-Delafuente et al., (2013) studied that plants against herbivory attack, resulting in B. tabaci female adults were subjected to an enhanced performance of B. tabaci. acquisition access period of 72 h in tomato Furthermore, Luan et al., (2012) found that yellow leaf curl virus (TYLCV) infected and plant defensive compounds named terpenoids, non-infected tomato plants to obtain were depleted by begomovirus infection, viruliferous and non-viruliferous whiteflies, favoring whitefly fitness. respectively. The results show evidence that TYLCV directly manipulates the settling, TYLCV, as other geminivirus, has some probing and feeding behavior of its vector B. pathogenic characteristics and could be tabaci in a way that enhances virus deleterious to its vector, B. tabaci. In fact, the transmission efficiency and spread. presence of TYLCVs in B. tabaci has been Furthermore, TYLCV and B. tabaci associated with a decrease in the insect interactions are mutually beneficial to both the longevity and fertility (Jiu et al., 2007, virus and its vector because B. tabaci feeds Rubinstein and Czosnek, 1997, Pusag et al., more efficiently after acquisition of TYLCV. 2012). Furthermore, a recent study This outcome has clear implications in the demonstrated an imbalanced nutrition of epidemiology and management of the whiteflies infected with TYLCCNV (Wang et
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Int.J.Curr.Microbiol.App.Sci (2018) 7(8): 4006-4023 al., 2012). However, limited studies of They feed specifically and selectivity on changes in the behavior of whiteflies due to certain plant tissues, which makes them infection with plant viruses such as TYLCVs efficient vectors of pathogens residing in these have been reported. tissues.
Phytoplasmas Their feeding is nondestructive, promoting successful inoculation of the plant vascular Phytoplasmas (originally called mycoplasma- system without damaging the conductive like organisms) are non-cellular degenerate tissues and eliciting defensive responses. gram-positive prokaryotes closely related to mycoplasmas and spiroplasmas. The They have a prokaryotic and persistent phytolasmas are non culturable parasitic relationship with phytoplasmas. They have prokaryotes, the mechanism of dissemination obligate symbiotic prokaryotes that are passed is mainly by insect vectors. to be offspring by transovarial transmission, the same mechanism that allow the The common vectors or at least those best transovarial transmission of phytoplasmas known are members of the order Hemiptera, from the families Cicadellidae, Cixiidae, Mechanism of transmission Psyllidae, Cercopidae, Delphacidae, Derbidae, Menoplidae and Flatidae. In the majority of Phytoplasmas are important phloem limited, cases transmission is only transovarial and has insect transmitted pathogenic agents causing a only been demonstrated in a few species thousand diseases, many of which are lethal, (Beanland et al., 2000 and Hanboonsong et in hundreds of plant species (Weintraub, al., 2002). 2007). Only phloem feeding insects can potentially acquire and transmit the pathogen. Insect vectors primarily leafhoppers, planthoppers and psyllids, have been The single most successful order of insect identified for relatively few phytoplasm phytoplasmas vectors is the Auchenorrhyncha. diseases (Weintraub and Beanland, 2006). To They are efficient vectors of phytoplasmas be transmitted to plant, phytoplasmas must because nymphs and adults feed similarly and penetrate and accumulate at high levels in the are in the same physical location, often, acinar cells posterior o the salivary glands. similarly and are in the same physical location, both immature and adults can In the salivary glands there are three barriers transmit phytoplasmas are propagative and that the pathogens must overcome before they persistent in them. can be expelled with the saliva: the basal lamina, the basal plasmalemma and the apical Most phytoplasma vectors are members of the plasmalemma. This group collectively family Cicadellidae like brinjal little leaf, possesses several characteristics that make its apple proliferation, aster yellows, tomato big members efficient vectors of phytoplasmas. bud and pear decline (Table 3).
They are hemimetabolous, thus nymphs and The feeding duration necessary to acquire a adults feed similarly and are in the same sufficient titer of phytoplasm (acquisition physical location-often both immature and access period), may range from a few minutes adults can transmit phytoplasmas. to several hours, the longer the period, the greater the chance of acquisition.
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Table.1 List of important insect vectors of plant viruses
S. Insect vector Virus (Hosts) No. 1. Acyrthosiphon pisum (Harris) Alfalfa mosaic (alfalfa), bean mosaic (beans), onion (Hemiptera: Aphididae) yellow dwarf (onion), pea mosaic (peas), soybean mosaic (beans, peas) 2. Aphis craccivora Koch Alfalfa mosaic (alfalfa), cowpea mosaic (cowpea), (Hemiptera: Aphididae) onion yellow dwarf (onion), papaya mosaic (papaya) 3. Aphis gossypii Glover Alfalfa mosaic (alfalfa, potato), chilli mosaic (chillies), (Hemiptera: Aphididae) cucumber mosaic (cucumber), dahlia mosaic (dahlia, zinnia) lettuce mosaic (lettuce, sweet pea), papaya mosaic (papaya), sugarcane mosaic (sugarcane)
4. Bemisia tabaci (Gennadius) Cotton leaf curl (cotton), dolichos yellow mosaic (Hemiptera: Aleyrodidae) (dolichos), okra yellow vein mosaic (okra), papaya leaf curl (papaya), sesame leaf curl (sesame), tobacco leaf curl (tobacco), tomato leaf curl (tomato)
5. Laodelphax striatella Fallen Rice streaked dwarf (rice), rice stripe (rice) (Hemiptera: Delphacidae) 6. Myzus persicae (Sulzer) Alfalfa mosaic (alfalfa), beet yellows (spinach, (Hemiptera: Aphididae) sugarbeet), cabbage black ring spot (cabbage), cauliflower mosaic (cabbage, cauliflower), cucumber mosaic (cucumber), dahlia mosaic (calendula, dahlia, zinnia), lettuce mosaic (garden pea, lettuce, sweet pea), onion yellow dwarf (onion), pea mosaic (broadbean, garden pea, sweet pea), potato virus Y (potato, tobacco, tomato), soybean mosaic (soybean), sugarcane mosaic (maize, sorghum, sugarcane)
7. Nephotettix nigropictus (Stal), Tungro (rice), yellow-orange leaf (rice) N. virescens (Distant) (Hemiptera: Cicadellidae 8. Nilaparvata lugens (Stal) Grassy stunt (rice), ragged stunt (rice) (Hemiptera: Delphacidae) 9. Pentalonia nigronervosa Coquerel Banana bunchy top (banana), cardamom mosaic (Hemiptera: Aphididae) (cardamom) 10. Rhopalosiphum maidis (Fitch) Barley yellow dwarf (barley, oat, rye, wheat), onion (Hemiptera: Aphididae) yellow dwarf (onion), sugarcane mosaic (maize, sorghum, sugarcane) Source: Modified after Gillot, 2005
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Table.2 List of important mite vectors of virus diseases of plants
Sr. No. Mite vector Virus Host(s) 1 Abbacarus hystrix (Nalepa) Agropyron mosaic Wheat, switch grass (Acari: Eriophyidae) 2 Aceria ficus (Corte) (Acari: Fig mosaic Fig Eriophyidae) 3 Aceria tulipae (Keifer) (Acari: Wheat streak mosaic Wheat, oats, barley, maize Eriophyidae) 4 Aculus fockeui (Nalepa and Prunus necrotic ring spot Plum, peach, cherry Trouessart) (Acari: Eriophyidae) 5 Eriophyes inaequalis Wilson and Cherry leaf mottle Sweet cherry Oldfield (Acari: Eriophyidae) 6 Eriophyesinsidiosu Keifer and Peach mosaic Peach, nectarine Wilson (Acari: Eriophyidae) Source: Modified after Gillot, 2005
Table.3 List of important vectors of phytoplasma-transmitted diseases
Sr. No. Insect Vector Mycoplasma(s) Host(s) References 1 Circulifer tenellus (Baker) Columbia basin Beet, potato, weeds Munyaneza et al., potato purple top 2007 2 Cestius phycitis (Distant) Brinjal little leaf Brinjal Gillot, 2005 (Hempitera: Cicadellidae) 3 Fieberiella florii Stal Apple Apple proliferation/16SrZ-A Tedeschi and Alma, proliferation 2006 4 Macrosteles sp. Lettuce 16SrI-B group Borth et al., 2006 5 Nephotettix nigropictus (Stal), N. Yellow dwarf Rice Gillot, 2005 virescens (Distant) (Hemiptera: Cicadellidae) 6 Macrosteles quadrillineatus Forbes Aster yellows Aster, barley, carrot, celery, Gillot, 2005 (Hemiptera: Cicadellidae) cucumber, wheat, rice 7 Neoaliturus fenestratus (Herrich- Lettuce phyllody Lettuce, wild lettuce Salehi et al., 2006 Schaffer) 8 Orosius orentalis (Matsumura) Garden beet Beets Mirzaie et al., 2006 witches’ broom 9 Orosius albicinctus (Distant) Sesame phyllody Sesame Gillot, 2005 (Hemiptera: Cicadellidae) 10 Recilia dorsalis (Motschulsky) Yellow dwarf Rice Gillot, 2005 (Hemiptera: Cicadellidae) 11 Stephanitis typical (Distant) Coconut root wilt Coconut Gillot, 2005 (Hemiptera: Cicadellidae) 12. Aster leafhopper Macrosteles Aster yellows Tomato, lettuce, carrot, onion, Thind, 2012 fascifrons potato, chrysanthemum, aster 13. Brown leafhopper, Orosius Tomato big bud Solanaceous vegetables and Thind (2012) argentatus lettuce 14. Leafhoppers, including Philaenus Apple Wild and ornamental apple Thind (2012) spumarius, Aphrophora alni, proliferation species, and possibly pear and Lepyronia coleoptrata, Artianus apricot interstitialis, and Fieberiella florii 15. Pear psylla, Psylla pyricola Pear decline Pear Thind, 2012
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Table.4 List of major vectors of bacterial diseases of crop plants
S.No. Insect vector Disease Host(s) 1. Seed corn maggot, Delia platura Bacterial soft rot of Potato and nearly all fleshy (Diptera: Anthomyiidae) potato vegetables are subject to bacterial soft rots. 2. Striped cucumber beetle Bacterial wilt of Cucurbits, including Acalymma vittatum and the cucurbits cucumber, muskmelon, spotted cucumber beetle, squash, and pumpkin. Diabrotica undecimpunctata (Coleoptera: Chrysomelidae)
3. Corn flea beetle, Chaetocnema Bacterial wilt of Sweet corn, Dent corn, pulicaria (Coleoptera: corn Popcorn, Field corn, gamma Chrysomelidae) grass, Teosinte 4. Aphids, leafhoppers, psyllids, Fire blight of pears, Pears, apples and other beetles, flies, and ants (More than apples and other rosaceous plants 200 species) rosaceous plants
5. Olive fruit fly, Bactrocera oleae Olive knot Olive (Diptera: Tephritidae) 6. Sharpshooter leafhoppers, Blue Pierce’s disease of European-type grapes, green (Graphocephala grape muscadine grapes, hybrids of atropunctata), Green European grapes and (Draeculacephala minerva), Red American wild grapes headed (Carneocephala fulgida), Glassy winged (Homalodisca coagulata) (Hemiptera: Cicadellidae) 7. Citrus psyllid, Asian pysllid i.e. Citrus greening Citrus Diaphorina citri and African disease psyllid i.e. Trioza erytreae. (Hemiptera: Psyllidae) 8. Leafhoppers Empoasca papayae Bunchy top of Papaya and E. stevensi (Hemiptera: papaya Cicadellidae) 9. Hylemya cilicrura (Rondani), H. Potato blackleg Potato trichodactyla (Rondani), (Diptera: Anthomyiidae) 10. Diaphorina citri Kumayama Citrus canker Citrus (Hemiptera: Aphalaridae) Source: (Gillot, 2005 and Thind, 2012)
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Table.5 List of principal vectors of fungal diseases of crop plants
Sr Insect vector(s) Diseases Host No. 1. Hylurgopinus rufips (Eichhoff) Dutch elm Elm (Coloptera: Scolytidae) 2. Melanoplus differetialis (Thomas) Cotton wilt Cotton (Orthoptera: Acrididae) 3. Several species of insect visiting Ergot of bajra Bajra flowers 4. Bees Blossom blight of red Red clover clover 5. Monilinia fructicola, M. fructigena, Brown rot of stone and Stone and pome and M. laxa. pome fruits 6. Fusarium moniliforme, Alternaria Boll rots Cotton tenuis, Aspergillus flavus, and Rhizopus nigricans 7. Rice stinkbug, Oebalus pugnax Rice mold Rice Source: Dollet 2001 and Gillot 2005
The time that elapse from initial acquisition to insect to acquire and subsequently transmit the ability to transmit the phytoplasmas phytoplasmas to plants suggests a high degree (latent period or incubation periods) is of specificity of phytoplasmas to insects. temperature dependent and ranges from a few to 80 days. Phytoplasmas during latent period However, numerous phytoplasmas are moves through and replicate in the vector’s transmitted by several different insect species. body. They can pass intracellularly through In addition, a single vector species may the epithelial cells of the midgut and replicate transmit two or more phytoplasmas, and an within a vesicle or they can pass between two individual vector can be infected with dual or midgut cells and through the basement multiple phytoplasma strains. It has been membrane to enter the haemocoel. found that leafhoppers do not acquire equally Phytoplasmas circulate in the haemocoel, phytoplasmas from different plant species. where they may infect other tissues such as the malpighian tubules, fat bodies and brain Bacteria or reproductive organs. To be transmitted to plants, phytoplasmas must penetrate specific Bacteria are microscopic single celled cells of the salivary glands and high levels organism that thrives in diverse environments. must accumulate in posterior acinar cells of They can live within soil, in the ocean and the salivary glands before they can be inside the human gut. A number of plant transmitted. diseases caused by bacteria are known to be transmitted by insects (Table 4). Bacterial wilt Vector phytoplasma relationship of corn caused by Erwinia carotovora pv. carotovora and transmitted by seed corn The interaction between insects and maggots. Potato blackleg, caused by Erwinia phytoplasmas is complex and variable. The carotowora, is transmitted by seed corn complex sequence of events required for an maggot, Hylemyia cilicrura (Rondani).
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Bacterial wilt of maize, caused by Fungi Xanthomonas stewarti is transmitted by the flea beetle, Chaetocnema pulicaria Fungi are organism having no chlorophyll, (Meisheimer). reproducing by sexual and asexual spores, not by binary fission like bacteria and typically Role of insects in bacterial diseases of possessing a mycelium or mass of interwoven plants threads (hyphae) containing well marked nuclei. There are about 4300 valid genra of In most plant diseases caused by plant fungi and about 70000 species living as pathogenic bacteria (especially in those that parasites or saprophytes on other organisms cause spots, cankers, blights, galls, soft rots) on their residues. which are produced within or between plant cells, escape to the surface of their host plants There are several insects associated with the as droplets or masses of sticky exudates. The spread of fungal disease (Table 5). The ergot bacteria exudates are released through cracks disease of bajra caused by Sphacelia or wounds in the infected area or through microcephala is mechanically carried by natural openings such as stomata, insects that visit the flowers attracted by the nectarthodes, hydathodes and sometimes surgery secretions found on the fungus through lenticells present in the infected area. infected earheads. The common sooty mold Such bacteria are then likely to stick on the fungus (Capnodiuim spp.) grows on the legs and bodies of all sorts of insects such as honeydews excreted by several homopteran flies, aphids, ants, beetles, whiteflies, etc. that insects like insects, leafhoppers, mealybugs, land on the plant and come in contact with the whiteflies, etc. bacterial exudates. Management of vectors and diseases Many of these insects are actually attracted by the sugars contained in the bacterial exudates The interactions between vectors, plant and feed on it, thereby further smearing their pathogens and crops are not so simple. Many body and mouthparts with the bacteria pathogens may be transmitted by more than containing exudates. When such bacteria one insect vector, and on the contrary, there smeared insects move to other parts of the are some vector species that are able to plant or to other susceptible host plants, they transmit different plant pathogens. The carry on their body numerous bacteria. If the knowledge about insect vectors is therefore insects happen to land on a fresh wound or on crucial when deciding pest management an open natural opening, and there is enough strategies, especially on a wide scale area. moisture on the plant surface, the bacteria Among the various types of plant diseases may multiply, move into the plant, and begin transmitted by insects, virus diseases are a new infection. considered to be the most serious.
Numerous plant diseases could be listed Hence multi-pronged strategy needs to be among those in which bacteria are spread by adopted to manage the vectors and virus insects passively as described above, for diseases. Some of the important components example, the bacterial bean blights, fire blight of such a strategy should involve selection of of apple and pear, citrus canker, cotton boll healthy seed, cultural practices, biological rot, crown gal, bacterial spot and canker of measures, resistant varieties and use of stone fruits, etc. chemicals.
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Healthy seed transmitted by transient winged aphids, including species that do not necessarily Management of virus diseases starts with colonize the sugarcane crop, so pesticide obtaining healthy seed, cutting or plants. Care application is not effective. Fortunately, there should be taken to obtain only certified seed, has been success with mosaic resistant i.e. seed obtained from the plants which have cultivars. been inspected during growing season and found free of certain diseases. With heat Cultural Control treatment virus free stock can be prepared, i.e. growing plants at high temperature for a Several cultural practices have proved to be weeks or even months. The production of helpful in reducing the incidence of vectors virus free stocks can also be achieved by and vector borne diseases. Many of the taking advantage of the fact that some plants effective management practices for diseases grow and elongate faster than the virus can caused by vectored plant pathogens involve occupy the new tissue. Therefore, the virus some sort of cultural control such as adjusted can be eliminated by using meristem or tip planting date, pruning, roguing and removal cultured plants. Virus free stock is tested by of volunteer crop plants and other non-crop indexing (growing a part of the culturing or reservoir hosts of vectors or pathogens. plant in a pot or greenhouse and recording its Control of volunteer crop plants may limit or condition with respect to disease symptoms), eliminate primary inoculum for newly planted bioassays and/or serological assays. crops. Volunteer potatoes are important sources of virus inoculum in Idaho potato Host plant resistance seed production areas. Similarly, volunteer grain can be an important reservoir for aphids Growing resistant and tolerant verities is and barley yellow dwarf at planting time for another effective way of managing vectors winter wheat. and vector-transmitted diseases. The efficacy of plant resistance to vector depends on the The volunteer wheat provides a ‘green bridge’ means of resistance and the mode of for the viruses and their vectors between transmission. Resistance that prevents feeding harvest of one crop and emergence of the next or repels the insects can prevent transmission one (Halbert, 2011). Other cultural control of pathogens spread by feeding. If the measures include elimination of weed hosts of resistance merely prevents or slows vectors or pathogens, use of reflective population growth, it cannot prevent primary mulches and paints to repel vectors, and spread. It can, however, have some effect on various protective row covers. Adjusting secondary spread. Resistance to the pathogen planting dates can minimize crop exposure to may be the only means of management in vectored pathogens. Intercropping with a some cases. An example is sugarcane mosaic barrier crop has provided encouraging results virus. Sugarcane is propagated vegetatively, to reduce the incidence of several diseases. so the virus is propagated with the crop. Once e.g. the incidence of yellow vein mosaic of planted, a crop remains in the same field for okra is reduced by intercropping with several years of production, and planting is soybean. Similarly, intercropping of tomato staggered throughout the production area. with coriander reduces the incidence of Thus, even if clean planting stock could be tomato leaf curl virus in tomato. Plant spacing found, there is no possibility for an area wide such as close spacing reduces the incidence of crop free and virus-free period. The virus is French bean crinkle stunt disease.
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Manipulation in planting dates is another way Chemical control of reducing the disease incidence. Roguing also help in the removal of the source of The control of insect vectors by application of disease causing organism. Removal of weeds insecticides appears to be a difficult task as and alternate hosts of viruses and vectors few survivors would be able to transmit the helps to reduce the incidence of diseases. disease. The timely application of insecticides restricts the spread of the disease by reducing Biological control the vector population. Several systemic and non-systemic insecticides have been reported Biological control is also one of the most to control the insect vectors. important tools for the management of vectored pathogens. Sometimes, the presence References of natural enemies evokes scatter responses in vector prey. This can actually cause an Agrios, G.N. (1997). Plant pathology (4th increase in pathogen transmission. Biological edn.). Academics Press, San Diego, control in simpler island ecosystems may California. have a better chance to work than in more Alma, A., Tedeschi, R., Lessio, F., Picciau, complex settings. For example, the L., Gonella, E. and Ferracini, C. (2015). introduction of psyllid parasitoids into Insect vectors of plant pathogenic Reunion Island dramatically reduced the Mollicutes in the Euro-Mediterranean transmission of the bacteria that cause citrus region. Phytopathogenic Mollicutes greening disease. However, the same 5(2): 53-73. parasitoids are present in Viet Nam, and citrus Anderson, B.J., Boyce, P.M. and Blanchard, greening disease is a major limiting factor in C.L. (1995). RNA 4 sequences from citrus production there. Sometimes disruption cucumber mosaic virus subgroups I and of these may increase the transmission of a II. Gene 161:293-294. plant pathogen if the existing biological Beanland, L., Hoy, C.W., Miller, S.A. and control does not work efficiently. Nault, L.R. (2000). Influence of aster yellows phytoplasmas on the fitness of Increased spraying for late blight in potatoes aster leafhopper (Homoptera: was linked to a major increase in numbers of Cicadellidae). Ann. of Ent. Soc. America green peach aphids. Evidently, populations 93: 271-276. had been controlled by an aphid pathogenic Borth, W.B., Fukuda, S.K., Hamasaki, R.T., fungus that was killed by the fungicide Hu, J.S. and Almeida, R.P.P. (2006). applications for late blight. The result was Detection, characterization and increase in incidence of potato leafroll virus, transmission of X-disease Phytoplasm transmitted by the aphids (Halbert, 2011). by the vector Colladonus montanus to Arabidopsis thaliana, a new Biopesticides experimental host plant. Pl. Dis. 89: 1121-1124. The use of biopestcides such microbial and Brown J.K. and Czosnek H. (2002). Whitefly plant extracts is an ecofriendly approach to transmission of plant viruses. In: Plumb manage the vectors and vectors borne RT, editor. Advanced in Botanical diseases. Several neem based formulations Research. Plant Virus Vector have provided effective control of B. tabaci Interactions. New York: Academic on cotton. Press. Vol. 36. 65-100.
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Brown J.K., Frolich D.R. and Rosell R.C. dwarf virus. Environ. Entomol. 18: 388- (1995). The sweet-potato or silverleaf 393. whiteflies - Biotypes of Bemisia tabaci Forbes, A.R. (1969). The stylets of the green or a species complex. Ann. Rev. peach aphid, Myzus persicae Entomol. 40: 511–534. (Homoptera: Aphididae). Can. Entomol Chen, B., and Francki, R.I.B. (1990). 101: 31–41. Cucumovirus transmission by the aphid Gamliel, A., Freeman, E.S., Sztejnberg, A., Myzus persicae is determined solely by Maymon, M., Ochoa, R., Belausov, E. the viral coat protein. J. Gen. Virol. and Palevsky, E. (2009). Interaction of 71:939-944. the mite Aceria mangiferae with Dalton, R. (2006). Whitefly infestations: The Fusarium mangiferae, the causal agent Christmas Invasion. Nature 443: 898– of mango malformation disease. 900. Phytopathology 99:152-159. DeBarro, P.J., Liu, S.S., Boykin, L.M. and Gillot, C. (2005). Entomology. Springer, Dinsdale, A.B. (2011). Bemisia tabaci: Dordrecht. The Netherland. a statement of species status. Ann. Rev. Gray, M.S. and Gildow, F.E. (2003). Entomol. 56: 1-19. Luteovirus-aphid interactions. Ann. Rev. Dollet, M. (2001). Phloem-restricted Phytopathology 41: 539-566. trypanosomatids form a clearly Halbert, S.E. (2011). Management of insect- characterized monophyletic group vectored pathogens of plants. Florida among trypanosomatids isolated from Department of Agriculture and plants. Int. J. Parasitol. 31, 459-467. Consumer Services, Division of Plant Donaldson, J.R. and Gratton, C. (2007). Industry Gainesville, Florida, USA. 1-5 Antagonistic effects of soybean viruses pp. on soybean aphid performance. Hanboonsong, Y., Choosai, C., Panyim, S. Environ. Entomol. 36: 918-925. and Danmark, S. (2002). Transovarial Drake, C.S., Gary, L.H., Frederick, E., transmission of sugarcane whitefly leaf Gildow, K.M.H. and Roy, F. (2005). transmission in insect vector Plant Virus HC-Pro Is a Determinant of Matsumuratettix hieroglyphics Eriophyid Mite Transmission. J. (Matsumura). Insect Mol. Biol. 11: 97- Virology 79(14): 9054-9061. 103. Du, Z.Q., Li, L., Liu, L., Wang, X.F. and Hogenhout, S.A., Ammar, E.D., Whitfield, Zhou, G. (2007). Evaluation of aphid A.E. and Redinbaugh, M.G. (2008). transmission abilities and vector Insect vector interactions with transmission phenotypes of barley persistently transmitted viruses. Ann. yellow dwarf viruses in china. J. Pl. Rev. Phytopath. 46: 327–359. Path. 89(2): 251-259. Hsu, H.T., Barzuna, L., Hsu, Y.H., Bliss, W. Fereres, A. and Moreno, A. (2009). and Perry, K.L. (2000). Identification Behavioural aspects influencing plant and subgrouping of Cucumber mosaic virus transmission by homopteran virus with mouse monoclonal insects. Virus Res. 141: 158-168. antibodies. Phytopathology 90: 615- Fereres, A., Lister, R.M., Araya, J.E. and 620. Foster, J.E. (1989). Development and Ingwell, L.L., Eigenbrode, S.D. and Bosque- reproduction of the English grain aphid Perez, N.A. (2012). Plant viruses alter (Homoptera, Aphididae) on wheat insect behavior to enhance their spread. cultivars infected with Barley yellow Sci. Rep. 2: 578.
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Jiu, M., Zhou, X.P., Tong, L., Xu. J. and Navas-Castillo, J., Fiallo-Olive, E. and Yang, X. (2007). Vector–virus Sánchez-Campos, S. (2011). Emerging mutualism accelerates population virus diseases transmitted by whiteflies. increase of an invasive whitefly. PLOS Ann. Rev. Phytopathol. 49: 219-48. ONE 2(1): e182. doi: Ng, J.C.K. and Falk, B.W. (2006). Virus- 10.1371/journal.pone.0000182. vector interactions mediating Kant, M.R., Sabelis, M.W., Haring, M.A. and nonpersistent and semipersistent Schuurink, R.C. (2008). Intraspecific transmission of plant viruses. Ann. Rev. variation in a generalist herbivore Phytopathol. 44: 183-212. accounts for differential induction and Palukaitis, P., Roossinck, M.J., Dietzgen, impact of host plant defences. Proc. R.G. and Francki, R.I.B. (1992). Royal Soc. B 275: 443-452. Cucumber mosaic virus. Adv. Virus Res. Liu, S.S., Colvin, J. and DeBarro, P.J. (2012). 41: 281-348. Species concepts as applied to the Perry, K.L., Zhang, L., and Palukaitis, P. whitefly Bemisia tabaci systematics: (1998). Amino acid changes in the coat How many species are there? J. Integr. protein of cucumber mosaic virus Agric. 11(2): 176-186. differentially affect transmission by the Luan, J.B., Yao, D.M., Zhang, T., Walling, aphids Myzus persicae and Aphis L.L. and Yang, M. (2012). Suppression gossypii. Virology 242: 204-210. of terpenoid synthesis in plants by a Polston, J.E., and Capobianco, H. (2013). virus promotes its mutualism with Transmitting plant viruses using vectors. Ecol. Lett. 16(2): 390–398. whiteflies. J. Vis. Exp. 8(81): e4332. Mirzaie, A., Esmailzadeh, H.S.A., Jafari, doi: 10.3791/4332. N.A. and Rahimian, H. (2006). Pusag, J.C.A., Jahan, S.M, H., Kwan-Suk, L., Molecular characterization and potential Sukchan, L. and Kyeon-Yeoll, L. insect vector a phytoplasma associated (2012). Upregulation of temperature with garden beet witches’ broom in susceptibility in Bemisia tabaci upon Yazd, Iran. J. Phytopath. 155(4): 198- acquisition of Tomato Yellow Leaf Curl 203. Virus (TYLCV). J. Insect Physiol. Moreno-Delafuente, A., Garzo, E., Moreno, http://dx.doi.org/ A. and Fereres, A. (2013). A Plant 10.1016/j.jinsphys.i.07.008. Virus Manipulates the Behavior of Its Roossinck, M.J. (2002). Evolutionary history Whitefly Vector to Enhance Its of Cucumber mosaic virus deduced by Transmission Efficiency and Spread. phylogenetic analyses. J. Virol. 76: PLOS ONE 8(4): e61543. 3382-3387. doi:10.1371/journal.pone.0061543. Rosen, R., Kanakala, S., Kliot, A., Munyaneza, J.E., Crosslin, J.M. and Upton, Pakkianathan, B.C., Farich, B.A., J.E. (2007). Beet leafhopper Magal, N.S., Elimelech, M., (Hemiptera: cicadellidae) transmits the Kontsedalov, S., Lebedev, G., Cilia, M. Columbia basin potato purple top and Ghanim, M. (2015). Persistent, phytoplasma to potatoes, beets and circulative transmission ofr weeds. J. Econ. Ent. 99(2): 268-272. begomoviruses by whitefly vectors. Nault, L.R. (1997). Arthropod transmission of Curr. Opinion Virol. 15:1-8. plant viruses: A new synthesis. Ann. Rubinstein, G. and Czosnek, H. (1997). Long- Entomol. Soc. Am. 90: 521-541. term association of tomato yellow leaf curl virus with its whitefly vector
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Bemisia tabaci: effect on the insect protein key to plant virus transmission transmission capacity, longevity and at the tip of the insect vector stylet. fecundity. J. Gen. Virol. 78: 2683-2689. Proc. Natl. Acad. Sci. U.S.A. 104, Salehi, M., Izadpanah, K. and Nejat, N. 17959-17964. (2006). A new phytoplasma infecting Wang, J., Bing, X.L., Li, M., Ye, G.Y. and lettuce in Iran. Pl. Dis. 90(2): 247. Liu, S.S. (2012). Infection of tobacco Sarwar, M. (2014). Some Insect Pests plants by a begomovirus improves (Arthropoda: Insecta) of Summer nutritional assimilation by a whitefly. Vegetables, Their Identification, Entomol. Exp. Appl. 144: 191-201. Occurrence, Damage and Adoption of Weintraub, P.G. (2007). Insect vectors of Management Practices. Int. J. phytoplasmas and their control-an Sustainable Agric. Res. 1(4): 108-117. update. Bull. Insectology 60(2): 169- Stafford, C.A., Walker, G.P. and Ullman, 173. D.E. (2011). Infection with a plant virus Weintraub, P.G. and Beanland, L. (2006). modifies vector feeding behavior. Insect vectors of phytoplasmas. Ann. www.pnas.org/cgi/doi/10.1073/pnas.11 Rev. 51: 91-111. 00773108. Whipple, O.C., and Walker, J.C. (1941). Sun, D.B., Xu, J., Luan, J.B. and Liu, S.S. Strains of cucumber mosaic virus (2011). Reproductive incompatibility pathogenic on bean and pea. J. Agric. between the B and Q biotypes of the Res. 62: 27-60. whitefly Bemisia tabaci in China: Whitfield, A.E., Falk, B.W., and Rotenberg, genetic and behavioural evidence. Bull. D. (2015). Insect vector-mediated Entomol. Res. 101: 211-220. transmission of plant viruses. Virology Tedeschi, R. and Alma, A. (2006). Fieberiella 47: 278-289. florii (Homoptera: Auchenorrhyncha) as Zhang, T., Luan, J., Qi, J., Huang, C. and Li, a vector of Candidatus hytoplasma M. (2012). Begomovirus-whitefly mali. Pl. Dis. 90(3): 284-290. mutualism is achieved through Thind, B. S. (2012). Phytopathogenic repression of plant defences by a virus prokaryotes and plant diseases Pp 327- pathogenicity factor. Mol. Ecol. 21(5): 476. Scientific Publishers, India. 1294-1304. Uzest, M., Gargani, D., Drucker, M., Hebrard, E., Garzo, E., Candresse, T. (2007). A
How to cite this article:
Ravinder Singh Chandi, Sanjeev Kumar Kataria and Jaswinder Kaur. 2018. Arthropods as Vector of Plant Pathogens Viz-a-Viz Their Management. Int.J.Curr.Microbiol.App.Sci. 7(08): 4006-4023. doi: https://doi.org/10.20546/ijcmas.2018.708.415
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