Breeding for resistance to whitefly-transmitted geminiviruses

Geminiviruses comprise a large and diverse family of viruses that infect a wide range of important monocotyledonous and dicotyledonous crop species and cause significant yield losses. The family Geminiviridae is divided into three genera, one of which is Begomovirus. Species of this genus are transmitted by the whitefly Bemisia tabaci in a persistent, circulative manner and infect dicotyledonous plants. Severe population outbreaks of B. tabaci are usually accompanied by a high incidence of begomoviruses. During the last two decades, there has been a worldwide spread of the B biotype of B. tabaci, accompanied by the emergence of whitefly‐transmitted geminiviruses. Control measures in infected regions are based mainly on limitation of vector populations, using chemicals or physical barriers. However, under conditions of severe whitefly attack, none of these control measures has sufficed to prevent virus spread. Thus, the best way to reduce geminivirus damage is by breeding crops resistant or tolerant to the virus, either by classical breeding or by genetic engineering. A number of begomoviruses have been the subject of much investigation, due to their severe economic impact. This review considers the most severe viral diseases of four major crops (tomato, bean, cassava and cotton). The approaches taken to breed for resistance to these viral diseases should provide a perspective of the issues involved in breeding for begomovirus resistance in crop plants.     

Prevalence of the major pests in transgenic and non-transgenic cotton genotype in response to climatic factors and their impact on yield

Abstract

The incidence of insect pests in cotton crop, such as Bemisia tabaci, Amrasca devastans and Pectinophora gossypiella (PBW) is dependent on climatic factors, such as maximum and minimum temperature, relative humidity and rainfall. Besides, cotton leaf curl disease (CLCuD) is also a major yield-limiting factor in the cotton crop. The results indicated that non-transgenic genotypes were heavily infested by whitefly, whereas jassid infested all genotypes with little difference. The population of whitefly and jassid acquired peak level in the month of July and August on all the genotypes. The correlation studies indicated the negative correlation of jassid, pink bollworm and CLCuD on yield component, whereas whitefly infestation was found positively correlated with CLCuD. Moreover, multivariate regression analysis (MRA) indicated the maximum and minimum temperature impacted the infestation of whitefly and jassid more than other factors, whereas maximum temperature greatly impacted the infestation of PBW in transgenic genotypes.     

Viruses Associated with Cassava Mosaic Disease in Senegal and Guinea Conakry

A survey in Senegal and Guinea Conakry established the presence and incidence of cassava mosaic virus disease (CMD) in both countries. CMD occurred in all the fields surveyed, although its incidence was higher in Senegal (83%) than in Guinea (64%). Populations of the whitefly vector, Bemisia tabaci, were low in both countries averaging 1.7 adults per shoot in Guinea and 3.2 in Senegal. Most infections were attributed to the use of infected cuttings, 86 and 83% in Senegal and Guinea, respectively, and there was no evidence of rapid current‐season, whitefly‐borne infection at any of the sampled locations. Disease severity was generally low in the two countries and averaged 2.5 in Guinea and 2.3 in Senegal. No plants with unusually severe CMD symptoms characteristic of the CMD pandemic in East and Central Africa were observed. Restriction fragment length polymorphism (RFLP)‐based diagnostics revealed that African cassava mosaic virus (ACMV) is exclusively associated with CMD in both the countries. Neither East African cassava mosaic virus (EACMV), nor the recombinant Uganda variant (EACMV‐UG2) was detected in any sample. These survey data indicate that CMD could be effectively controlled in both countries by phytosanitation, involving the use of CMD‐free planting material and the removal of diseased plants.     

Spatial and Temporal Spread of Cassava Mosaic Virus Disease in Cassava Grown Alone and when Intercropped with Maize and/or Cowpea

The spread of cassava mosaic disease (CMD) and populations of the whitefly vector (Bemisia tabaci) were recorded in cassava when grown alone and when intercropped with maize and/or cowpea. The trials were planted under conditions of high inoculum pressure in 1995 and 1996 at a site in the lowland rainforest zone of southern Cameroon. In the 1995 experiment, the maize and cowpea intercrops reduced the final incidence of CMD in the cassava cvs. Dschang White and Dschang Violet, but not in the more resistant cv. Improved. In the 1996 experiment with cv. Dschang Violet, the maize and cowpea intercrops grown alone or together decreased adult whitefly populations on cassava by 50% and CMD incidence by 20%. The monomolecular population growth model generally provided the best fit for disease progress. Areas under the disease progress curves (AUDPCs) and incidences expressed as multiple infection units were significantly (P<0.05) less for cassava intercropped with maize and/or cowpea than in cassava alone; times to 50% CMD incidence were significantly (P<0.05) longer in all intercrop systems. In 1995 the basic infection rates (r) were similarly low (0.010 per month) in the moderately resistant cv. Dschang Violet intercropped with maize and in all treatments in the more resistant cv. Improved. By contrast, rates were significantly higherfor cv. Dschang Violet alone or with cowpea and in all treatments for the less resistant cv. Dschang White (0.030–0.060). In 1996, r values in cassava grown alone (0.077) were significantly larger (P<0.05) than in the other cropping systems (0.042–0.052). There were no significant differences in the symptom severity in the different cropping systems. Disease foci were isodametric and more compact in plots containing cowpea than in other cropping systems.     

A shift of vector specificity acquired by a begomovirus through natural homologous recombination

Recombination is common in plant viruses such as geminiviruses, but the ecological and pathogenic consequences have been explored only in a few cases. Here, we found that a new begomovirus, tomato yellow leaf curl Shuangbai virus (TYLCSbV), probably originated from the recombination of Ageratum yellow vein China virus (AYVCNV) and tobacco curl shoot virus (TbCSV). Agrobacterium-mediated inoculation showed that TYLCSbV and AYVCNV have similar levels of infectivity on tomato and tobacco plants. However, the two viruses exhibit contrasting specificities for vector transmission, that is, TYLCSbV was efficiently transmitted by the whitefly Bemisia tabaci Mediterranean (MED) rather than by the whitefly B. tabaci Middle East-Asia Minor 1 (MEAM1), whereas AYVCNV was more efficiently transmitted by MEAM1. We also showed that the transmission efficiencies of TYLCSbV and AYVCNV are positively correlated with the accumulation of the viruses in whitefly whole bodies and organs/tissues. The key coat protein amino acids that determine their accumulation are between positions 147 and 256. Moreover, field surveys suggest that MED has displaced MEAM1 in some regions where TYLCSbV was collected. Viral competition assays indicated that TYLCSbV outcompeted AYVCNV when transmitted by MED, while the outcome was the opposite when transmitted by MEAM1. Our findings suggest that recombination has resulted in a shift of vector specificity that could provide TYLCSbV with a potential selective transmission advantage, and the population shift of whitefly cryptic species could have influenced virus evolution towards an extended trajectory of transmission.     

Cardinium infection alters cotton defense and detoxification metabolism of its whitefly host

Field monitoring revealed that the infection ratio of the bacterial symbiont Cardinium in the whitefly (Bemisia tabaci MED) was relatively low in northern China. However, the role of this symbiont and the symbiont–whitefly–host plant interaction mechanism are poorly understood. We investigated the influence of Cardinium on the competitiveness of the host whitefly and the physiological interaction between the host plants and host whiteflies. Cardinium-infected whiteflies were displaced by uninfected whiteflies after 5 generations, which showed that Cardinium infection reduced whitefly competitiveness. The defense response genes of cotton significantly decreased under infestation by infected whiteflies compared to uninfected whiteflies. The expression of detoxification metabolism genes, especially the uridine 5ʹ-diphospho-glucuronyltransferase and P450 genes, in infected whiteflies significantly decreased. These results demonstrated that Cardinium could inhibit the defense response of the host plant and decrease the detoxification metabolism ability of the host whitefly. The reduced competitiveness of infected whiteflies may be associated with the inhibition of the whitefly detoxification metabolism by Cardinium, resulting in the reduced performance of infected whiteflies. However, Cardinium infection can suppress plant defenses, which may benefit both infected and uninfected whiteflies when they coexist. This research illustrates the symbiont–whitefly–host plant interaction mechanism and the population dynamics of the whitefly.     

Cucumber Vein Yellowing Virus; Host Range and Virus Vector Relationships

Cucumber vein yellowing virus (CVYV) is transmitted by Bemisia tabaci, it has a narrow host range restricted to some cucurbitaceous plants including Cucumis sativus, C. melo, C. melo var. flexousus, Cucurbita pepo, C. foesti, Citrullus vulgaris, C. colocynthis and Lagenaria siceraria. Although a single whitefly can transmit the disease, the efficiency of transmission was low. At least 15–20 insects per plant were required to cause an infection of 55 % of inoculated plants. The minimum acquisition and inoculation feeding periods were 30 and 15 min, respectively. The latent period in the vector is about 75 min and the whitefly was infectious for not more than 5 h.     

Prediction of cotton leaf curl virus disease and its management through resistant germplasm and bio-products

Cotton leaf curl virus disease reduces the cotton yield significantly every year and is transmitted by Bemisia tabaci. The study was designed to evaluate 15 varieties/lines against the disease. Multiple regression analysis was performed based on a-biotic environmental variables (maximum air temperature, minimum air temperature, relative humidity and rainfall) to predict disease incidence and its vector (Bemisia tabaci). Two bio-products were evaluated against the whitefly population to control the disease. Out of 15 cotton varieties/lines, no one was found highly resistant against the disease. Five varieties/lines (BT BT-980, BT-457, KIRAN, BT-666 and SLH-BT-6) exhibited moderately resistant response. Maximum air temperature (34–35.5 °C), minimum temperature (25.75–26.25 °C), relative humidity (64.14–66%), rainfall (1–2 mm) and wind speed (5.50–5.75 Kmh^?1) favoured the disease development. Maximum whitefly population was favoured by maximum air temperature from 34–35.5 °C, 25.8–26.2 °C minimum air temperature, 64.14–66% relative humidity, 1–2 mm from rainfall and 5.50–5.75 Kmh^?1 wind speed. Datura stramonium was found more effective as compared to Aviara (Homoeopathic) but not from the positive control (Acetamiprid).     

Zoophytophagy in the plantbug Nesidiocoris tenuis

1 The zoophytophagy of Nesidiocoris tenuis (Reuter) (Heteroptera: Miridae) was characterized in relation to prey availability and environmental factors by: (i) monitoring its population dynamics in tomato greenhouses; (ii) analysis of the influence of N. tenuis and whitefly density, temperature and humidity on the intensity of N. tenuis plant feeding; and (iii) laboratory assays under controlled conditions to determine the intensity of plant feeding in relation to prey availability, temperature and humidity. 2 A negative relationship was found between plant feeding and predated whiteflies in tomato greenhouses. Plant feeding was directly related to N. tenuis density and temperature and inversely related to whitefly density. The significance of prey availability and temperature was corroborated in laboratory assays. The intensification of plant feeding at low prey density indicates switching from zoophagy to phytophagy as prey become scarce. 3 Nesidiocoris tenuis showed a typical predator dynamic in relation to variance in prey density. Populations increased after whitefly outbreaks and decreased after whitefly had been depleted. The rapid decrease of N. tenuis populations after whitefly decreased, however, suggests that plants are a poorer nutrient source than whitefly for this species.     

Progress of cassava mosaic virus disease and whitefly vector populations in single and mixed stands of four cassava varieties grown under epidemic conditions in Uganda

Progress curves of cassava mosaic virus disease (CMD) and populations of the whitefly vector (Bemisia tabaci) were assessed using four cassava varieties grown alone and as a random mixture in two experiments established under epidemic conditions at a site near Kampala in southern Uganda. There were significant differences in final CMD incidence and in the areas under the disease progress curves between varieties when grown alone and as a mixture in both experiments. Variety Ebwanateraka had the highest incidence and SS4 the lowest, even though it supported the largest populations of adult whiteflies. The overall incidence of CMD in the mixture was similar to that in pure stands of the partially resistant Nase 2 and greater than in the resistant Migyera and SS4. Compared to pure stands, incidence of CMD in each component of the mixture was reduced significantly only in Ebwanateraka, whereas vector populations were less only in SS4 and Nase 2. On several observation dates the actual incidence of CMD and populations of adult whiteflies in the mixture were significantly less than expected values estimated from the results for the four varieties when each was grown alone. A highly significant positive relationship was established for each variety between peak populations of adult whitefly and leaf area index at the time. The implications of the findings and the scope for future research on the use of varietal mixtures for the management of CMD are discussed.     

Whitefly Bemisia tabaci (Homoptera: Aleyrodidae) infestation on cassava genotypes grown at different ecozones in Nigeria

Large-scale screening of cassava, Manihot esculenta Crantz, genotypes for resistance to infestation by whitefly Bemisia tabaci Gennadius, the vector of cassava mosaic geminiviruses, is limited. A range of new cassava elite clones were therefore assessed for the whitefly infestation in the 1999/2000 and 2000/2001 cropping seasons in experimental fields of International Institute of Tropical Agriculture, Ibadan, Nigeria. On each scoring day, between 0600 and 0800 hours when the whiteflies were relatively immobile, adult whitefly populations on the five topmost expanded leaves of cassava cultivars were counted. All through the 6-mo scoring period, there was a highly significant difference in whitefly infestation among the new cassava elite clones. Vector population buildup was observed in Ibadan (forest-savanna transition zone) and Onne (humid forest), 2 mo after planting (MAP). Mean infestation across cassava genotypes was significantly highest (16.6 whiteflies per plant) in Ibadan and lowest in Zaria (0.2). Generally, whitefly infestation was very low in all locations at 5 and 6 MAP. During this period, cassava genotypes 96/1439 and 91/02324 significantly supported higher infestations than other genotypes. Plants of 96/1089A and TMS 30572 supported the lowest whitefly infestation across cassava genotypes in all locations. The preferential whitefly visitation, the differences between locations in relation to whitefly population, cassava mosaic disease, and the fresh root yield of cassava genotypes are discussed.     

Low frequency of horizontal and vertical transmission of two begomoviruses through whiteflies exhibits little relevance to the vector infectivity

Transmissions of plant viruses between individuals of their vector insects through mating are rare events. Recently, three begomoviruses were found to be transmitted between males and females of the whitefly Bemisia tabaci through mating, and two viruses were shown to be transmitted transovarially to progeny. However, results between reports were not consistent. Here we examined the horizontal and vertical transmission of Tomato yellow leaf curl virus (TYLCV) and Tomato yellow leaf curl China virus (TYLCCNV) by the B and Q biotypes of B. tabaci, using virus isolates and whitefly colonies established recently in China. Both TYLCV DNA and TYLCCNV DNA were shown to be transmitted horizontally and vertically by each of the two biotypes of the whitefly, but frequency of transmission was usually low. In transovarial transmission, virus DNA was detected in eggs and nymphs but not in the adults of the first generation progeny, except in the combination of TYLCV and Q biotype whitefly where 2–3% of the offspring adults contained the virus DNA. We also showed that the first generation adults, which developed from eggs of viruliferous whiteflies, were not infective to plants. These results demonstrated that for the viruses and whiteflies tested here low frequency of horizontal and vertical transmission can be expected but these two modes of transmission are unlikely to have much epidemiological relevance in the field.     

A simulation study of population interaction between the greenhouse whitefly,Trialeurodes vaporariorum Westwood (Homoptera: Aleyrodidae), and the parasitoidEncarsia formosa gahan (Hymenoptera: Aphelinidae) I. Description of the model

A simple simulation model was developed to simulate the population dynamics of the system of the greenhouse whitefly (Trialeurodes vaporariorumWestwood ) and the parasitoid Encarsia formosaGahan . On the assumption that temperature is constant, the whitefly population was described as theLeslie Matrix model. Parasitization and host feeding by the parasitoid population were modelled by means of a modified disc equation. The validity of the model was demonstrated by comparing the predictions of the model with the observed values obtained in greenhouse experiments.     

Yellow Vein Mosaic Disease in Okra (<i>Abelmoschus esculentus</i> L.): An Overview on Causal Agent,Vector and Management

Okra (Abelmoschus esculentus L.) belongs to the Malvaceae family and is one of the most essential and popular vegetables globally. It is rich in proteins, carbohydrates, and vitamins. Abiotic and biotic factors threaten okra productivity. Okra yellow vein mosaic disease (OYVMD) is the most destructive disease of okra. The causal agent, [(i.e., Okra yellow vein mosaic virus (OYVMV)] of this disease belongs to the family Geminiviridae and genus Begomovirus. OYVMV is a monopartite with additional ssDNA molecule. This virus has two components DNA-A for protein coding and DNA-B for symptoms induction. Whitefly transmits OYVMV in persistent manner. Characteristic symptoms of OYVMV infected okra plants are chlorosis, dwarfing, and yellowing of veins and fruits. High temperatures with moderate rainfall enhance the development of OYVMV disease and the whitefly population. However, high humidity with low temperature and rainfall has no significant role in developing the OYVMD and whitefly population. Moreover, the virus also affects the secondary metabolites in the infected okra plants. The virus can be managed through various strategies including the application of plant defense activators, the development of resistant varieties and by controlling its vector via pesticides and plant extracts. Various plant defense activators such as monopotassium phosphate (KH₂PO), salicylic acid, benzoic acid, and citric acid enhance resistance in okra against OYMVD. In addition, the resistance to OYMVD can also be achieved by successfully incorporating high yielding but resistant cultivars of acceptable quality. In this review, we have discussed history, economic impact, symptomology, disease development under a natural environment, genetics and management of OYVMV.     

Density thresholds for Nesidiocoris tenuis (Heteroptera: Miridae) in tomato crops

Nesidiocoris tenuis (Reuter) (Heteroptera: Miridae) is common in vegetable crops of the Mediterranean area, with an increasing worldwide range of geographical distribution. This omnivore is a reputed predator of small arthropod pests, but also produces injuries on vegetative and reproductive plant parts. The aim was to estimate density thresholds based on N. tenuis and whitefly abundance for the management of N. tenuis in tomato crops. The assay was carried out in mesh-walled and plastic greenhouses in southern Spain during 2004 and 2007. The natural population dynamics of N. tenuis and whitefly were monitored, and impact on yield quantified. The economic injury level and intervention threshold were predicted based on the zoophytophagous response of N. tenuis and the yield compensation of tomato plants. The proportion of aborted flowers on the tomato plants was related directly to the abundance of N. tenuis and inversely to the interaction between the number of N. tenuis and the number of whitefly immatures. Over-compensation of fruit weight was predicted for flower abortion rates due to N. tenuis lower than 0.171. No yield reduction is expected for values <0.65 N. tenuis per leaf, independent of the whitefly abundance, nor for up to 5 N. tenuis and >26 whitefly immatures per leaf. For intermediate N. tenuis levels, the outcome depends on the prey density. The probability of N. tenuis producing yield loss in tomato crops increases at N. tenuis:whitefly ratios >0.168. Yield reduction is expected after N. tenuis population peaks, when whitefly numbers have been reduced.     

Migration, trapping and local dynamics of whiteflies (Homoptera: Aleyrodidae)

1 The ability to quantify whitefly migration provides a tool that can contribute to an improved understanding of the epidemic development of whitefly‐transmitted viruses. 2 In an attempt to develop a protocol for estimating whitefly immigration and emigration rates in an annual crop, new traps and sampling devices were tested in the field and models for population dynamics were developed. 3 An estimate of immigration rate was derived from the growth of a natural population of Trialeurodes vaporariorum (Westwood) in the beginning of a crop cycle before offspring of immigrants contributed to population growth. 4 A model for changes in whitefly density during an entire bean (Phaseolus vulgaris L.) crop cycle, including an immigration parameter, was also developed. 5 Non‐attractant window traps surrounding an annual field crop were assumed to intercept whiteflies immigrating into and emigrating away from the crop. Captures on these traps could not categorically be identified as immigrants or emigrants, but the cumulated captures nevertheless explained 66% of the variation in population density found within the field. Hence, window traps may be used as an efficient and reliable alternative to yellow sticky traps, aspirator methods and leaf‐turn methods, etc., for estimating whitefly densities in field crops.     

The effects of mullein plants (Verbascum thapsus) on the population dynamics of Dicyphus hesperus (Heteroptera: Miridae) in tomato greenhouses

The response of Dicyphus hesperus Knight (Heteroptera: Miridae) to whitefly populations in tomato greenhouses was measured in the presence and absence of mullein (Verbascum thapsus L.) as an alternative host plant. The dynamics of the D. hesperus population on tomato (Lycopersicon esculentum Mill.) and on mullein plants were followed through an entire growing season. In houses with mullein plants, more predators occurred on mullein when whitefly density was low on tomato. A mark-release-recapture experiment where rabbit IgG was used as an external marker showed that D. hesperus adults moved from mullein plants to tomato plants. D. hesperus was always more abundant in houses with mullein than in the houses with tomato plants alone. Movements between tomato and mullein plants are discussed as a strategy to optimize predator foraging. The use of mullein as an alternative host plant may contribute to the establishment of D. hesperus and help to preserve the predator population when prey on tomato crops is scarce.     

Tomato yellow leaf curl virus infection of tomato does not affect the performance of the Q and ZHJ2 biotypes of the viral vector Bemisia tabaci

Abstract To better understand the etiology of begomovirus epidemics in regions under invasion we need to know how indigenous and invasive whitefly vectors respond to virus infection. We investigated both direct and indirect effects of infection with Tomato yellow leaf curl virus (TYLCV) on the performance of the invasive Q biotype and the indigenous Asian ZHJ2 biotype of whitefly Bemisia tabaci. The Q biotype performed better than the ZHJ2 biotype on either uninfected or virus‐infected tomato plants. However, virus‐infection of host plants did not, or only marginally affected, the performance of either biotype of whiteflies in terms of fecundity, longevity, survival, development and population increase. Likewise, association of the vectors with TYLCV did not affect fecundity and longevity of the Q or ZHJ2 biotypes on cotton, a non‐host of TYLCV. These results indicate that the alien Q biotype whitefly, but not the indigenous ZHJ2 biotype, is likely to become the major vector of TYLCV in the field and facilitate virus epidemics.