Faba bean necrotic yellows virus
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- Risk of Introduction
- Hosts/Species Affected
- Host Plants and Other Plants Affected
- Growth Stages
- List of Symptoms/Signs
- Biology and Ecology
- Means of Movement and Dispersal
- Vectors and Intermediate Hosts
- Detection and Inspection
- Similarities to Other Species/Conditions
- Prevention and Control
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Faba bean necrotic yellows virus
Other Scientific Names
- faba bean necrotic yellows nanavirus
Taxonomic TreeTop of page
- Domain: Virus
- Unknown: "ssDNA viruses"
- Unknown: "DNA viruses"
- Family: Nanoviridae
- Genus: Nanovirus
- Species: Faba bean necrotic yellows virus
Notes on Taxonomy and NomenclatureTop of page
Four species are assigned to the genus Nanavirus: Subterranean clover stunt virus (type species), Banana bunchy top virus, Milk vetch dwarf virus and Faba bean necrotic yellows virus (FBNYV). The species name of FBNYV is derived from the symptoms induced on faba bean, the natural host from which the virus was first isolated. The genus name is derived from the Greek noun nanos meaning dwarf, and refers to the observations that these plant viruses have the smallest known virions and genome segment sizes, and cause dwarfing in their hosts.
DescriptionTop of page The average diameter of FBNYV particles is 18 nm. They have an angular outline, but no obvious substructure (Katul et al., 1993).
Analysis of the genome of an FBNYV isolate from Syria (Sy) revealed the presence of 10 circular ssDNA species. All 10 components range from 985 to 1014 bases in size, contain one major open reading frame in the virion sense, and have a non-coding region containing a highly conserved sequence possibly forming a stem-loop (SL) structure. The SL also contains the conserved nonanucleotide sequence (AGTATTACC) which, as in geminiviruses, may be the origin of a rolling-circle replication mechanism (Koonin and Ilyina, 1992; Laufs et al., 1995). The 10 circular DNA components (C1-C10) found associated with the FBNYV genome potentially encode four distinct replication-associated proteins (Rep) and six non-Rep proteins (Katul et al., 1995, 1997, 1998). The presence of four Rep and six non-Rep components in the FBNYV-Sy genome is further supported by sequence and PCR data on another virus isolate from Egypt (FBNYV-Eg), as well as by the data of Sano et al. (1998) on milk vetch dwarf nanavirus (MVDV), a close relative of FBNYV. Thus the FBNYV and MVDV genomes appear strikingly similar, not only in the total number of identified components but also in the number and types of homologous non-Rep components present in each genome.
In addition to the two Rep components (C1 and C2) identified earlier (Katul et al., 1995, 1997), the sequences of two further Rep-encoding components (C7 and C9) in two FBNYV isolates have recently been determined (Katul et al., 1998). The presence of four genome components potentially encoding distinct Rep proteins in two FBNYV isolates and in MVDV is a most puzzling phenomenon, and is in contrast to geminiviruses which possess only one (begomoviruses) or two (mastreviruses) Rep genes (Lazarowitz, 1992). Tentative evidence suggests that the Rep protein encoded by FBNYV-C2 may play a pivotal and indispensable role in FBNYV replication, whereas the three other Rep components (C1, C7 and C9) may actually be non-integral parts (satellite DNAs) of the FBNYV genome, as suggested for some Rep components of banana bunchy top nanavirus (Horser et al., 1996).
The four Rep and six non-Rep components identified so far may represent the complete FBNYV genome. However, the possibility that the genome contains further, as yet unidentified, components cannot be eliminated. Conclusive evidence for the complete identification of all integral parts of the FBNYV genome will only be obtained in infectivity tests with cloned components, in order to reproduce a disease whose causal agent is indistinguishable in all its biological properties from field isolates of FBNYV.
DistributionTop of page
FBNYV was first isolated from Vicia faba near Lattakia on the Syrian coast. It is endemic in a number of countries of West Asia and North Africa. It caused a serious epidemic on the faba bean crop in Middle Egypt which led to a crop failure during the 1991/92 growing season. In addition to the countries shown on the map, it has also been observed in Sudan (K. Makkouk, ICARDA, personal observation).
Distribution TableTop of page
The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Iran||Present||Makkouk et al., 2002|
|Iraq||Present||El-Muadhidi et al., 2001|
|Jordan||Widespread||Katul et al., 1993|
|Lebanon||Present||Katul et al., 1993|
|Pakistan||Present||Makkouk et al., 1998a|
|Syria||Present||Katul et al., 1993|
|Turkey||Present||Katul et al., 1993|
|Yemen||Present||Makkouk et al., 1998d|
|Algeria||Present||Katul et al., 1993|
|Egypt||Widespread||Katul et al., 1993; Makkouk et al., 1994|
|Eritrea||Present||Kumari et al., 2008|
|Ethiopia||Present||Katul et al., 1993; Abraham et al., 2012|
|Libya||Present||Zidan et al., 2002|
|Morocco||Present||Katul et al., 1993|
|Sudan||Present||Makkouk et al., 2003|
|Tunisia||Present||Najar et al., 2000|
|Spain||Present||Ortiz et al., 2006|
Risk of IntroductionTop of page All reports published so far suggest that FBNYV is not seed transmitted. Accordingly, seed movement between countries does not pose an increased risk of introduction into new locations. However, since the virus is persistently transmitted by aphids, it can be carried in windborne aphids for hundreds of kilometres.
Hosts/Species AffectedTop of page
FBNYV has a relatively narrow host range, mostly restricted to leguminous species. Faba bean is the main natural host, but other legume crops such as chickpea, lentil, dry bean, pea and cowpea are also natural hosts of FBNYV (Makkouk et al., 1992; Katul et al., 1993; Franz et al., 1995; Horn et al., 1995). The virus also occurs naturally in the wild legume species Lathyrus sativus, L. gorgonei, L. annuus, L. hierosolyminatus, Medicago polymorpha, M. praecox, M. rigidula, M. rotata, M. scutellata, Melilotus officinalis, Tetragonolobus purpureus, Trifolium arvense, T. hirtum, T. lappaceum, T. subterraneum, Vicia ervilia, V. hybrida, V. palaestina and V. sativa, as well as in perennial species of the genus Onobrychis and in Medicago sativa; the virus also occurs naturally in some non-leguminous species including Amaranthus blitoides, A. retroflexus and A. viridis (Mouhanna et al., 1994a; Franz et al., 1997).
The virus can be transmitted experimentally by aphids to a number of plant species with a variety of systemic symptoms such as chlorosis, leaf distortion, leaf rolling, necrosis, reddening and stunting. Experimental hosts include Astragalus annularis IFAS 499; Coronilla reponda IFCO 85; C. rostrata IFCO 84; C. scorpioides IFCO 87; Glycine max cultivars Catlar 71, Clark, Crawford, Egyptian Local; Lathyrus cicera IFLA 536; L. hirsutus IFLA 47; L. ochrus IFLA 101; L. odoratus; Lens culinaris cv. Syrian Local; Medicago coronata IFMA 834; M. hispida [M. polymorpha]; M. intertexta IFMA 1156, Cilinaris; M. laciniata IFMA 4068; M. littoralis littoralis IFMA 5470; M. noeana IFMA 4067; M. orbicularis IFMA 135; M. polymorpha brevispina IFMA 5468; M. polymorpha vulgaris IFMA 4089; M. rigidula IFMA 282, v. cinerascens IFMA 3244; M. tornata IFMA 4065; M. truncatula [M. tribuloides] IFMA 4069; Melilotus indica IFME 25; Phaseolus vulgaris cultivars Black Turtle 2, Egyptian Local, Giza 3, Morgan, Great Northern 123, Redlands Greenleaf C; Pisum sativum cultivars IFPI 2903, Kleine Rheinlanderin, Koroza Onyx, Rondo Syrian Local; Stellaria media (Germany); Trifolium alexandrinum IFTR 799, Fahle, Gemmiza 1, Giza 6, Giza 10, Giza 15, Sakha 4; T. incarnatum; T. lappaceum IFTR 1297; T. pauciflorum IFTR 1683; T. resupinatum IFTR 88; T. spumosum IFTR 1793; T. subterraneum IFTR 699, IFTR 704, IFTR 1252, Karridale, Mt Barker; Trigonella foenum-graecum cultivars Beni Suef, Egyptian Local, Giza 2; Vicia mollis IFVI 3486; V. palaestina IFVI 2606; V. sativa IFVIC 4211, IFVI 4212, IFVI 3011, IFVI 652; Vigna unguiculata cultivars California Black Eye No. 5, Tvu 196, Tvu 1582, Tvu 3273, Tvu 3433 (Nigeria).
The virus also infects the following species symptomlessly: Hippocrepis multisiliquosa IFHI 108; Medicago polymorpha IFMA 4070; M. polymorpha var. polymorpha IFMA 1162; Scorpiurus muricatus IFSC 88 (Katul et al., 1993; Franz et al., 1997).
Host Plants and Other Plants AffectedTop of page
|Amaranthus blitoides (spreading amaranth)||Amaranthaceae||Other|
|Amaranthus retroflexus (redroot pigweed)||Amaranthaceae||Wild host|
|Amaranthus viridis (slender amaranth)||Amaranthaceae||Wild host|
|Cicer arietinum (chickpea)||Fabaceae||Other|
|Lathyrus sativus (grasspea)||Fabaceae||Wild host|
|Lens culinaris subsp. culinaris (lentil)||Fabaceae||Other|
|Medicago sativa (lucerne)||Fabaceae||Other|
|Medicago scutellata (snail medic)||Fabaceae||Wild host|
|Melilotus officinalis (yellow sweet clover)||Fabaceae||Wild host|
|Phaseolus vulgaris (common bean)||Fabaceae||Other|
|Pisum sativum (pea)||Fabaceae||Other|
|Tetragonolobus purpureus (Asparagus-pea)||Fabaceae||Wild host|
|Trifolium subterraneum (subterranean clover)||Fabaceae||Wild host|
|Vicia ervilia (Bitter vetch)||Fabaceae||Wild host|
|Vicia faba (faba bean)||Fabaceae||Main|
|Vicia sativa (common vetch)||Fabaceae||Wild host|
|Vigna unguiculata (cowpea)||Fabaceae||Other|
Growth StagesTop of page Flowering stage, Fruiting stage, Seedling stage, Vegetative growing stage
SymptomsTop of page
One-week-old faba bean plants show retarded growth as early as 5 days after inoculation. At 2 weeks after inoculation, the plants are usually severely stunted. The leaves become thick and brittle and show interveinal chlorotic blotches starting from the leaf margins. The uppermost young leaves remain very small and cupped upwards, whereas the older leaves are rolled downward. New shoots, leaves and flowers develop poorly. About 3-4 weeks after infection, interveinal chlorosis usually turns necrotic and infected plants die within 5-7 weeks after infection. Similar symptoms were also observed in other susceptible host plants. Some infected Trifolium and Medicago species, however, develop leaf reddening instead of, or in addition to, chlorosis (Katul et al., 1993).
List of Symptoms/SignsTop of page
|Fruit / reduced size|
|Growing point / dieback|
|Leaves / abnormal colours|
|Leaves / abnormal forms|
|Leaves / necrotic areas|
|Leaves / yellowed or dead|
|Roots / reduced root system|
|Seeds / empty grains|
|Seeds / shrivelled|
|Stems / dieback|
|Stems / stunting or rosetting|
|Whole plant / plant dead; dieback|
Biology and EcologyTop of page
The virus is not infective in expressed plant sap. The genome is circular ssDNA. Ten DNA components have been isolated from virus preparations; individual molecules, each of which appears to be encapsidated in a separate particle, are approximately 1 kb in size.
Only the aphid species Acyrthosiphon pisum, Aphis craccivora and A. fabae have been reported as FBNYV vectors. The efficiency of transmission by aphids was found to be high for the first two species and very poor for A. fabae (Katul et al., 1993; Franz et al., 1995, 1998). The information on its aphid transmission indicates that FBNYV has features typical of a persistently transmitted virus which circulates but does not multiply in the vector insects. Aphids require long acquisition and inoculation feeding periods to become efficient vectors, and FBNYV persists in the aphids for almost their entire life (Franz et al., 1998). As with all phloem-limited viruses, FBNYV is not known to be transmitted by seed or mechanical means. Since all its hosts are propagated by true seed, the only method of natural spread of FBNYV is by aphid vectors.
Sources of Infection
The host range of FBNYV seems to be largely restricted to the family Fabaceae (Franz et al., 1996). Many food and forage legume crops such as Cicer arietinum, Lathyrus spp., Lens culinaris, Medicago spp., Pisum sativum, Trifolium spp., Vicia faba (many cultivars) and many Vicia forage species are susceptible to infection (Franz et al., 1996). More recently, the virus was found to naturally infect cowpea (Vigna unguiculata) and French bean (Phaseolus vulgaris) (Franz et al., 1995), both summer legumes widely grown in the West Asia and North Africa region. In addition to the above legume crops, leguminous weeds such as Melilotus officinalis, Tetragonolobus purpureus, Lathyrus spp., Medicago spp. and Trifolium spp. were also found naturally infected with FBNYV in Egypt and Syria (Franz et al., 1996).
Most of these susceptible legume species are grown as winter crops, except cowpea and French bean which are probably important over-summering hosts of the virus. When aphid vectors are abundant, the virus can be spread from overlapping winter to summer legumes and again to winter legumes. Severe epidemics can occur when environmental conditions permit the build-up of the aphid vector populations which have access to virus sources.
Virus spread in any region depends not only on vector activity and population build-up and on virus titres in sources of infection, but also on the number and proximity of the source plants. Thus, a nearby infected crop provides a much higher inoculum pressure than a few infected weeds (Bos and Makkouk, 1994). In Egypt, an FBNYV-susceptible crop such as Egyptian clover (Trifolium alexandrinum), which is planted in large areas and often next to faba bean fields, probably plays a role in virus spread to the faba bean crop. Quantitative studies have not yet been conducted to determine the relative importance of such crops and weeds as sources of FBNYV infection. However, final infection pressure to which a given crop (e.g. faba bean) is subject is, of course, determined by inoculum potential and vector pressure.
Means of Movement and DispersalTop of page
The only method of natural spread of FBNYV is by aphid vectors. Acyrthosiphon pisum, Aphis craccivora and A. fabae have been reported as FBNYV vectors.
Vectors and Intermediate HostsTop of page
ImpactTop of page FBNYV-infected faba bean, chickpea or lentil plants produce few or no seeds depending on time of infection. Infection of young plants leads to 100% yield loss, whereas when older plants are infected significant yield loss occurs. High incidence of FBNYV has been reported on faba bean in Egypt, in the Jordan Valley of Jordan, in the coastal region of Syria and in some regions of Ethiopia. The FBNYV epidemic on faba bean in Middle Egypt during the growing season 1991/92 led to almost complete crop failure (Makkouk et al., 1994).
Detection and InspectionTop of page
Observation of symptoms induced by FBNYV on infected legume crops is not enough to confirm virus presence. There are a number of phloem-limited viruses which affect legumes (for example, Bean leaf roll virus (BLRV), Beet western yellows virus (BWYV), Soybean dwarf virus (SbDV) and Chickpea chlorotic dwarf virus (CCDV). ELISA (using either polyclonal or monoclonal antibodies) and dot-blot hybridization are very sensitive tests to detect the virus in infected plants as well as in its aphid vector (Katul et al., 1993, 1995; Franz et al., 1996). Serologically specific electron microscopy is also useful in detecting the virus in infected sap, and in differentiating it from viruses in the family Luteoviridae which induce similar symptoms on legume species, such as BLRV, BWYV and SbDV, or viruses of the Geminiviridae such as CCDV.
Similarities to Other Species/ConditionsTop of page
FBNYV is taxonomically very closely related to Milk vetch dwarf virus, and less so to Subterranean clover stunt virus. All three viruses cause similar symptoms in legume species (Grylls and Butler, 1959; Inouye et al., 1968; Chu and Helms, 1988; Sano et al., 1993).
Symptoms induced by FBNYV on the cool-season legume crops chickpea, faba bean, lentil and pea are mostly yellowing, stunting and poor pod setting. Such symptoms are very similar to those induced by Bean leaf roll virus (BLRV) (Duffus et al., 1990), which previously caused misidentification of FBNYV as BLRV in many countries of West Asia and North Africa, as such identification was based on symptoms alone. At present, it is possible to differentiate between all the above viruses by using serological as well as other techniques (Katul et al., 1995).
Prevention and ControlTop of page Introduction
Disease control is based on the results of ecological and epidemiological studies. Control measures may involve one or more of the approaches listed (Makkouk et al., 1998b, c).
Reducing Sources of Infection
A significant decrease in disease incidence can be expected if sources of infection are eliminated from within or near crops. In practice this is not easy, as it is impossible to eliminate all weed hosts of the virus. Prior to advocating such an approach for FBNYV control, more information on the relative importance of wild legumes as sources of FBNYV infection is needed.
Another measure to decrease disease incidence is to eliminate sources of infection within the crop. Roguing of FBNYV-infected faba bean eliminates or reduces primary infection foci. Reduction in virus incidence was noticed in a number of fields in Egypt when roguing was practised two or three times during the growing season (El-Hadi and Hilmi, personal communication). More experimentation on this aspect is still required to evaluate the economic impact of roguing on crop yield. On the other hand, a scattered distribution of initially infected plants throughout a crop was observed in some faba bean fields in Egypt, suggesting that FBNYV had been introduced from a distant source. This type of spread makes the reduction of infection through management of cropping patterns alone unlikely.
In preliminary screening for FBNYV resistance using 200 pure lines of faba bean, no resistance was found (K. Makkouk, ICARDA, unpublished data). Further work, however, is planned to screen the remaining available faba bean germplasm.
When lentil lines were tested for FBNYV resistance under artificial field inoculation by viruliferous aphids, several lines such as ILL 213, 291, 6198, 6193 and 6245 were found to be highly resistant (Mouhanna et al, 1994b). Resistance was expressed by low virus incidence and extremely low (less than 5%) yield loss. These results were obtained in a single environment and require confirmation. Such preliminary data suggest that lentil genotypes vary widely in their reaction to FBNYV infection as compared to faba bean lines.
Novel approaches for virus resistance
Faba bean and FBNYV appear to offer a promising model for assessing the potential of pathogen-derived resistance to a DNA virus in faba bean by using various FBNYV genes. Since four potential Rep genes of FBNYV have been identified, and certain functions (e.g., capsid protein and cell-to-cell spread) have been definitely or tentatively assigned to some of the non-Rep genes of FBNYV (Katul et al., 1995, 1997, 1998), several viral genes appear to be suitable candidates to be used for transforming faba bean. However, transformation of faba bean with FBNYV genes depends not only on the availability of FBNYV genes but, more importantly, on an efficient transformation and regeneration protocol for Vicia faba. Since this species is very recalcitrant to tissue-culture techniques, such a protocol has not yet been established. Suitable and highly efficient promoters, supplied either by the FBNYV genome itself or by using well-established promoters, such as the CaMV 35S and the PR1 promoters, are available.
Although the molecular details of the replication mechanism of nanaviruses have not yet been elucidated, it appears valid to assume that their overall replication strategy is similar to that of geminiviruses (Lazarowitz, 1992). Several features of the genome structure and the Rep proteins of the nanaviruses (Harding et al., 1993; Boevink et al., 1995; Katul et al., 1995, 1997, 1998; Sano et al., 1998) are very similar to those of the geminiviruses: an inverted repeat sequence with the potential to form a hairpin-like structure; an AT-rich sequence within the loop of the hairpin, very similar to the conserved nonamer sequence that is the origin of geminivirus plus-strand replication; and putative Rep proteins sharing essential amino-acid motifs with the Rep proteins of geminiviruses. Therefore, the development of a pathogen-derived resistance strategy for FBNYV should concentrate on approaches that have already been shown to be successful for geminiviruses (Mettouchi, 1992; Aaziz, 1994; Desbiez et al., 1995). Hence, in vitro engineered Rep genes in particular appear to be the most promising candidates for generating transgenic faba bean plants with high levels of FBNYV resistance. This experimental approach is currently followed for generating FBNYV-resistant faba bean lines, by using as a model Medicago truncatula, an FBNYV-susceptible legume species, for which an efficient transformation and regeneration protocol is available.
Experiments conducted in Egypt during the period 1994-97 indicated that the use of one or two foliar sprays of a selective insecticide (pirimicarb) reduced virus spread significantly. In addition, experiments conducted at ICARDA showed that the use of a systemic seed treatment insecticide (imidacloprid) gave significant protection of lentil and faba bean plots against FBNYV infection for 2 months after sowing. Such treatment could prove useful in areas where infection with FBNYV is likely to occur early in the season.
In both Syria and Egypt, faba bean crops planted early in September are often severely attacked, leading to 100% FBNYV infection. In such circumstances, farmers plough the crop under and replant another crop. Delaying sowing until October or November often resulted in lower levels of virus infection, most probably due to lower population densities and reduced activities of the vector aphids. However, more studies are required to establish the most suitable sowing date.
Each of the control measures mentioned provides only partial control, but combining genetic resistance, cultural practices and chemical sprays is expected to lead to improvements. The use of host resistance, whether obtained by classical breeding or genetic engineering, and one or two well-timed sprays, coupled with optimal planting date and early roguing of virus-infected plants, could offer reasonable and economic control and stabilize cool-season legume production. However, to develop a sustainable package of integrated control, more work is needed on the ecology and epidemiology of FBNYV in the different regions where the virus causes damage to cool-season legumes. If we obtain a better understanding of the distribution, natural hosts and vector transmission of FBNYV, as well as the interrelationship between epidemic build-up and climatic conditions, better control strategies can be developed.
The seriousness of FBNYV infection of cool-season legumes, especially faba bean, in some countries of West Asia and North Africa requires a multidisciplinary approach, exploiting all possible control measures in order to achieve sustainable yields in areas where epidemics due to FBNYV infection are likely to occur.
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Distribution MapsTop of page
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