Studies on the transmission of groundnut rosette virus by Aphis craccivora Koch

Four strains of groundnut rosette virus were transmitted by a race of Aphis craccivora (Koch) from groundnut in Nigeria. Two of these strains, both from East Africa, were transmitted only by A. craccivora from Kenya. A fifth isolate, from Nigeria, was not transmissible by either race. The two races of aphids have been shown elsewhere to be distinct biotypes. Most A. craccivora needed longer than 24 h feeding on infected groundnuts to acquire virus, and many needed 2–3 days of feeding on healthy plants to cause infection, even after several days on infected plants. The delays partly reflect the slow uptake of virus and possibly a period needed for virus multiplication in aphid tissue but some is lost through resistance of the test plants to infection. In consecutive feeding experiments Natal Common variety could be infected soon after aphids had left the source of virus, but a more resistant Nigerian variety sometimes needed several more days. The frequency of inoculation by aphids, or the concentration of virus in the inocula or both, increased with time, but the times at which aphids were able to infect plants was also dependent on variety.     

Effect of Fungicides on the Viability of Phylophthora cactorum Propagules m the Soil

Vein-clearing followed by downward rolling and necrosis of leaves and severe stunting of groundnut (Arachis hypogaea) plants were caused by cowpea mild mottle virus (CMMV). The virus was readily transmitted by mechanical sap inoculations to groundnut and to 10 plant species belonging to Leguminosae, Chenopodiaceae and Solanaceae. Chenopodium quinoa and Beta vulgaris were good diagnostic hosts. Diseased sap remained infective at 10^–3 but not 10^–4, when stored 8 to 9 days at 25 °C; for 10min at 75 °C but not 80°C. In limited tests, virus was not seed-transmitted m groundnut or soybean. Virus was transmitted by Bemisia tabaci but not by Aphis craccivora or Myzus persicae. An antiserum for CMMV was produced and virus was serologically related to CMMV reported on cowpea and groundnut crinkle virus (GCV) from West Africa. Employing carbon diffraction grating replica as a standard the modal length of virus particles to be 610 nm. Infected cells contained large number of virus particles associated with endoplasmic reticulum.     

Peanut mottle virus in East Africa

A very common and widespread virus pathogen of groundnut and soybean in East Africa was identified as peanut mottle virus (PnMV)* on the basis of particle morphology, serology, host range and reaction, transmission and physical properties. Virus concentration adequate for serological tests was obtained from cowpea (Vigna unguiculata) cultured at 27 °C but not at 23 °C. Purified preparations from this source gave a single, specific light-scattering zone in sucrose density gradients. PnMV was purified using 0.5 M sodium citrate buffer containing 1% mercapto-ethanol; an antiserum made against such preparations had a homologous titre of 1/8192. Groundnut and soya isolates from N., N.E., N.W. and S. districts of Uganda, N.W. Tanzania, and W. and E. (coastal) districts of Kenya were serologically similar and varied, within narrow limits, in symptoms induced in certain groundnut and soya varieties. A serologically related but distinct virus was isolated from Voandzeia subterranea. PnMV was not related serologically to any of ten viruses of the PVY group. Glasshouse experiments simulating groundkeeper conditions in the field indicated 20% seed transmission in groundnut; PnMV was transmitted by Aphis craccivora in the non-persistent manner. All twenty-one varieties and breeding lines of soybean tested were highly susceptible. The prevalence of PnMV in East Africa and the reduction in yield caused in groundnut indicates the virus to be economically important, and groundnut and soybean improvement programmes should include routine PnMV susceptibility tests.     

Viruses associated with chlorotic rosette and green rosette diseases of groundnut in Nigeria*

Groundnut (Arachis hypogaea) plants from Nigeria with chlorotic rosette disease contained a manually transmissible virus, considered to be a strain of groundnut rosette virus (GRV(C)). GRV(C) infected nine out of 32 species in three out of nine families. It caused local lesions without systemic infection in Chenopodium amaranticolor, C. murale and C. quinoa, and systemic symptoms in Glycine max, Nicotiana benthamiana, N. clevelandii and Phaseolus vulgaris as well as in groundnut. Some ‘rosette-resistant’ groundnut lines were also infected. GRV(C) was transmitted by Aphis craccivora, but only from groundnut plants that were also infected with an aphid-transmissible second virus, which was not manually transmissible and was considered to be groundnut rosette assistor virus (GRAV). Plants infected with GRAV contained isometric particles c. 25 nm in diameter which were detectable by immunosorbent electron microscopy on grids coated with antisera to several luteoviruses, especially with antisera to bean leaf roll, potato leafroll and beet western yellows viruses. No virus-like particles were observed in extracts from plants infected with GRV(C) alone. A single groundnut plant obtained from Nigeria with symptoms of green rosette contained luteovirus particles, presumed to be of GRAV, and yielded a manually transmissible virus that induced symptoms similar to those of GRV(C) in C. amaranticolor but gave only mild or symptomless infection of N. benthamiana and N. clevelandii. It was considered to be a strain of GRV and designated GRV(G).     

Characterisation and serology of virus-like particles associated with faba bean necrotic yellows

A disease showing chlorosis, leaf rolling and stunting in Vicia faba and other legumes was observed in West Asia and North Africa during 1987–1988. The putative causal agent could not be transmitted mechanically, but could be transmitted by aphids, most efficiently by Acyrthosiphon pisum, in the persistent manner. Further studies revealed isometric virus-like particles (VLPs) closely associated with the disease, although their infectivity could not be demonstrated by membrane feeding. These particles, measuring c. 18 nm in diameter and containing a capsid protein of about 22 kDa and ssDNA of about 1 kb, are hereafter designated faba bean necrotic yellows virus (FBNYV). A high proportion of circular nucleic acid molecules of about 0.9 kb were visualised by electron microscopy. Hybridisation analysis of cloned viral DNA suggests that the circular genome is larger than 1 kb and consists of several components of similar size. An antiserum produced against FBNYV was used in ELISA, immunoelectron microscopy (IEM) and Western blot experiments for virus detection in aphids and field samples and for serological comparison with other viruses. Weak heterologous reactions between FBNYV and subterranean clover stunt virus (SCSV) were detected in IEM, but could not be confirmed in ELISA or Western blots. No serological relationship to banana bunchy top virus (BBTV) was detected. Using a direct tissue blot immunoassay (TBIA), FBNYV was detected in vascular tissue of infected faba bean leaves and stems.     

Groundnut eyespot virus, a new member of the potyvirus group

A virus causing ‘eyespot’ leaf symptoms in groundnut plants was transmitted by sap-inoculation and by Aphis craccivora in the non-persistent manner. It infected 16 of 72 species from five of 12 families and was easily propagated in Arachis hypogaea and Physalis floridana. The virus has particles c. 13 × 755 nm and is serologically closely related to soybean mosaic and pepper veinal mottle viruses, and more distantly to four other potyviruses. The virus differs in host range, in vitro properties and serological properties from previously described strains of soybean mosaic and pepper veinal mottle viruses. It seems to be a distinct member of the potyvirus group and we propose the name groundnut eyespot virus.     

Crimson clover latent virus - a newly recognised seed-borne virus infecting crimson clover (Trigolium incarnatum)

Crimson clover latent virus (CCLV) was detected in five seed lots of crimson clover (Trifolium incarnatum) from Europe and in one from the United States of America. Ninety-seven per cent of all crimson clover plants examined were found to be infected but were without symptoms. Keeping crimson clover plants at 32–38°C for 34 days failed to free them from CCLV. The virus was not transmitted by Myzus persicae, but was transmitted by inoculation of sap to Chenopodium album, C. amaranticolor and C. quinoa. Twenty-four other plant species from seven families were not infected. CCLV was best propagated in C. quinoa in which it caused stunting and systemic chlorosis. Sap from infected C. quinoa was infective after dilution to 10^-2 but not 10^-3, after 10 min at 60°C but not 65°C, and after 20 days at 20°C. In neutral phosphotungstate, CCLV had isometric particles c. 26 nm in diameter with a hexagonal profile. About 20 to 80 A_1cm,260 units of purified virus were obtained from 1 kg of infected C. quinoa or C. amaranticolor leaves by extraction in 0.5 M phosphate buffer, pH 7.5, containing 0.01 M ethylene diamine tetra-acetate and 0.4% 2–mercaptoethanol and clarification with chloroform-butanol followed by two precipitations with polyethylene glycol (mol. wt 6000) and several cycles of differential centrifugation. Purified virus sedimented as three components with sedimentation coefficients (s°_{20, w}) of 52S, 101S and 122S. The 101S and 122S components had buoyant densities in CsCl of 1.438 and 1.495 g/cm³ respectively. From these values the nucleic acid content of the 101S and 122S components was estimated to be 32–35% and 40–41% respectively. The virus contained a single protein with an estimated mol. wt of 52 000 and two single-stranded RNA species of estimated mol. wt 1.6 × 10⁶ and 2.2 × 10⁶. CCLV was serologically unrelated to 31 other morphologically similar viruses. Although its vector is unknown, CCLV seems to have affinities with nepoviruses. The cryptogram of CCLV is R/1:2.2/40–41 + 1.6132–35:S/S:S/*.     

Host range and overwintering sources of bean leaf roll and pea enation mosaic viruses in England

Bean leaf roll virus (BLRV) and pea enation mosaic virus (PEMV) were each transmitted by Acyrthosiphon pisum (Harris) to fifteen of thirty species of legumes in the glasshouse; eleven species were susceptible to both viruses. The only biennial or perennial species infected by BLRV were hop trefoil (Medicago lupulina L.), lucerne (M. sativa L.) and red clover (Trifolium pratense L.), but naturally infected sainfoin (Onobrychis viciifolia Scop.) and white clover (T. repens L.) were found. The only perennial species infected with PEMV in the glasshouse was kidney vetch (Anthyllis vulneraria L.). Eggs of A. pisum, which seems to be the main vector of BLRV and PEMV in England, were found in winter on several species of cultivated perennial legumes, most on lucerne, fewest on white clover. In spring, more viviparae of A. pisum were found on lucerne than on other perennial legumes, and many on lucerne, but few on red or white clover, were infective with BLRV. Lucerne is probably the main overwintering source of BLRV in areas where lucerne is common, but elsewhere red and white clovers are probably as important. No aphid collected from perennial legumes between 1965 and 1968 was infective with PEMV, but this virus can overwinter in common vetch (Vicia sativa L.). Lucerne infected with BLRV was usually symptomless or showed only transient mild yellowing of young leaves. Lucerne plants showing vein-yellowing, similar to that previously reported as a symptom of BLRV, were possibly infected with an aberrant strain of BLRV but more probably with BLRV and another aphid-transmitted agent. Inoculations from lucerne with vein-yellowing symptoms sometimes caused vein-yellowing, and sometimes typical BLRV-symptoms, in crimson clover (Trifolium incarnatum L.).     

Observations on two races of the groundnut aphid, Aphis craccivora

Two clones of Aphis craccivora, one from Nigeria and one from Kenya, differed in their ability to transmit the GRVW₁ strain of the rosette virus (Okusanya, 1965). Comparison of body measurements of the two clones showed them to be two distinct but overlapping populations. The Nigerian clone formed colonies on Onobrychus viciifolia, Gomphrena globosa and young Soya max more readily than the Kenya clone. The fecundity of single viviparous females of the two clones on groundnut and bean does not differ significantly. Alatae produce half as many young as apterae.

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Zusammenfassung Zwei Klone von Aphis craccivora, einer von Nigeria und einer aus Kenia, unterscheiden sich in der Fähigkeit, Erdnußvirus-Stämme zu übertragen (Okusanya, 1965). Maße der Körpergröße und die Längen der Siphonen, Hintertibien, Antennenglieder und Vordertibien zeigen, daß sie zwei sich überlappenden Populationen angehören. Jungpflanzen von Soya max, Gomphrena globosa und Onobrychis viciifolia werden von dem nigerischen Stamm viel leichter besiedelt als von dem aus Kenia. Einzelne Aptere der beiden Klone produzierten an Erdnuß und Ackerbohnen ähnliche Nachkommenzahlen. Eine einzelne Aptere gebar an Erdnuß 230 Junge; die meisten Larven wurden während der ersten 12 Tage der Fortpflanzungsperiode geboren. Geflügelte beider Klone produzierten nur halb so viele Larven (im Mittel 57 und 52) wie Ungeflügelte (im Mittel 118 und 110). Die beiden Klone sind biologische Rassen von A. craccivora.

     

Properties and partial purification of infective material from plants containing groundnut rosette virus*

Groundnut plants with chlorotic rosette disease contain a manually transmissible virus, groundnut rosette (GRV), which is also transmitted in the persistent (circulative) manner by aphids (Aphis craccivora), but only from plants that are co-infected with a manually non-transmissible luteovirus, groundnut rosette assistor virus (GRAV). Strains of GRV from plants with chlorotic or green forms of rosette are called GRV(C) and GRV(G) respectively. An isolate of GRV(C) from Nigeria remained infective in Nicotiana clevelandii leaf extracts for 1 day at room temperature and for 15 days at 4d?C, but lost infectivity after 1 day at -20d?C or after dilution to 10^--⁴. Its infectivity and longevity in vitro were not altered by addition of 1 mg/litre bentonite to the extraction buffer. Infectivity in leaf extracts was abolished by treatment with 50% (v/v) ether, 10% (v/v) chloroform or 8% (v/v) n-butanol, but not by treatment for 30 min with RNase A at up to 100 ng/ml. In attempts to purify GRV(C), nearly all the infectivity from N. clevelandii extracts was found in the pellets from centrifugation at 65 000 g for 1. 5 h; infectivity also occurred in a cell membrane fraction that collected at the top of a 30% sucrose ‘cushion’ containing 4% polyethylene glycol and 0.2 M NaCI. However, no virus-like particles were found in either type of preparation by electron microscopy. Nucleic acid preparations made directly from GRV(C)-infected N. clevelandii leaves were very infective; this infectivity was totally inactivated by treatment for 30 min with RNase A at 10 ng/ml in buffers of both low and high ionic strength and was therefore attributed to ssRNA. When nucleic acid preparations were electrophoresed in gels no virus-specific bands were visible but the position of the infectivity indicated that the infective ssRNA has an apparent mol. wt of c. 1.55 × 10⁶. A similar mol. wt was indicated by the rate of sedimentation of the infective ssRNA in sucrose gradients. Preparations of dsRNA made from GRV(C)-infected N. clevelandii leaves contained a species of mol. wt c. 3.0 × 10⁶; in addition some dsRNA preparations contained an abundant component of mol. wt c. 0.6 × 106 together with several other components of intermediate mol. wt. Similar patterns of bands were observed in dsRNA preparations made from Nigerian-grown groundnut material infected with GRV(C) alone, or with GRV(C) + GRAV, or with GRV(G) + GRAV. The properties of GRV closely resemble those of two other viruses that depend on luteoviruses for transmission by aphids, carrot mottle virus and lettuce speckles mottle virus.     

Peanut stripe virus - a new seed-borne potyvirus from China infecting groundnut (Arachis hypogaea)1

A new virus, peanut stripe (PStV), isolated from groundnut (Arachis hypogaea) in the USA, induced characteristic striping, discontinuous vein banding along the lateral veins, and oakleaf mosaic in groundnut. The virus was also isolated from germplasm lines introduced from the People's Republic of China. PStV was transmitted by inoculation of sap to nine species of the Chenopodiaceae, Leguminosae, and Solanaceae; Chenopodium amaranticolor was a good local lesion host. PStV was also transmitted by Aphis craccivora in a non-persistent manner and through seed of groundnut up to 37%. The virus remained infective in buffered plant extracts after diluting to 10^-3, storage for 3 days at 20°C, and heating for 10 min at 60°C but not 65°C. Purified virus preparations contained flexuous filamentous particles c. 752 nm long, which contained a major polypeptide of 33 500 daltons and one nucleic acid species of 3·1 × 10⁶ daltons. In ELISA, PStV was serologically related to blackeye cowpea mosaic, soybean mosaic, clover yellow vein, and pepper veinal mottle viruses but not to peanut mottle, potato Y, tobacco etch, and peanut green mosaic viruses. On the basis of these properties PStV is identified as a new potyvirus in groundnut.     

The response of certainTrifolium spp. to calcium in sand culture

Summary The response to calcium ofTrifolium incarnatum, T. pratense, T. repens andT. hybridum was determined over the range 4 to 128 ppm calcium in sand culture.T. incarnatum showed the greatest response to calcium and was severely affected by low levels of calcium.T. hybridum was the least responsive to calcium and showed no significant response over the range tested, although a gradual increase in yield with increasing calcium was apparent. The responses ofT. pratense andT. repens were intermediate and differed significantly from those of bothT. incarnatum andT. hybridum. There was no significant difference between the response ofT. pratense andT. repens.The results are considered in the light of the ecologically observed edaphic tolerances of the species, and it is concluded that there is a marked relationship between response to calcium and edaphic tolerance of the species.In view of the differences in response to calcium recently demonstrated within the speciesT. repens, the magnitude of the differences between species shown here may not strictly apply to all varieties or populations of those species.     

Tobacco yellow vein,a virus dependent on assistor viruses for its transmission by aphids*

Tobacco yellow vein, a disease found in Malawi, is caused by a combination of two viruses transmitted in the persistent manner by aphids. One component, tobacco yellow vein virus (TYVV) is manually transmissible, but aphids transmit it only from plants also containing the other (assistor) component, which is not manually transmissible. Aphids also transmit TYVV from plants containing either of two other assistor viruses - tobacco vein-distorting and groundnut rosette assistor. A virulent isolate of TYVV infected Soja max, Arachis hypogaea and several solanaceous species. It infected plants already containing tobacco mottle or groundnut rosette viruses but not those containing a mild isolate of TYVV.     

Different variants of the satellite RNA of groundnut rosette virus are responsible for the chlorotic and green forms of groundnut rosette disease*

Groundnut plants with symptoms of rosette disease contain groundnut rosette virus (GRV), but GRV is transmitted by Aphis craccivora only from plants that also contain groundnut rosette assistor virus (GRAV). Two main forms of rosette disease are recognised, ‘chlorotic rosette’ and ‘green rosette’. GRV cultures invariably possess a satellite RNA and this is the major cause of rosette symptoms: satellite-free isolates derived from GRV cultures from Nigerian plants with chlorotic or green rosette, or from Malawian plants with chlorotic rosette, induced no symptoms, or only transient mild mottle or interveinal yellowing, in groundnut. When the satellite RNA species from GRV cultures from Nigerian green or Malawian chlorotic rosette were reintroduced into the three satellite-free isolates in homologous and heterologous combinations, the ability to induce rosette symptoms was restored and the type of rosette induced was that of the cultures from which the satellite RNA was derived. Thus different forms of the satellite are responsible for the different forms of rosette disease. Other forms of the satellite induce only mild chlorosis or mottle symptoms in groundnut. Individual plants may contain more than one form of the satellite, and variations in their relative predominance are suggested to account for the variable symptoms (ranging from overall yellowing to mosaic) seen in some plants graft-inoculated with chlorotic rosette.     

Further properties of peanut clump virus and studies on its natural transmission

Purified preparations of particles of peanut clump virus (PCV) had A₂₆₀/A₂₈₀ values (corrected for light scattering) of 1.00. They contained rod-shaped particles with sedimentation coefficients of 183 S and 224 S, and a density in CsCl of 1.32 g/ml. PCV infected 36 species in 8 plant families. No serological relationship was detected between PCV and barley stripe mosaic, beet necrotic yellow vein, Nicotiana velutina mosaic and tobacco mosaic viruses. PCV was seed-borne for two generations in groundnut (Arachis hypogaea) but was not seed-borne in great millet (Sorghum arundinaceum), Phaseolus mungo or Nicotiana benthamiana. Seedlings of groundnut, great millet and wheat (Triticum aestivum) became infected when grown in soil from groundnut fields with outbreaks of clump disease, and the infectivity of soil survived air-drying at 25°C for 3 months. Groundnut seedlings became infected when grown in sterilised soil contaminated with washed roots of naturally-infected S. arundinaceum but not in soil to which roots of naturally infected groundnut or shoots of infected groundnut were added, or in which mechanically inoculated groundnut seedlings were grown at the same time. The patchy distribution of PCV in a crop was related to the infectivity of the soil for groundnut and to the presence of Polymyxa graminis resting spores which could be detected in the roots of S. arundinaceum bait seedlings, but not in those of groundnut. The results indicate that PCV is transmitted by a vector that is resistant to air-drying and closely associated with S. arundinaceum roots. For these reasons P. graminis is thought to be the vector of PCV.     

In vitro culture and somatic embryogenesis of four Trifolium species

Callus cultures were induced from hypocotyl sections of Trifolium incarnatum L., T. vesiculosum Savi., T. ambiguum Bieb., and T. repens L. Callus production was the greatest for T. incarnatum and T. ambiguum on Phillips and Collins L2 medium with 0.06 mg/l picloram and 0.1 mg/l 6-benzylaminopurine and for T. vesiculosum and T. repens on Gamborg B5 medium with 1.25 mg/l 2,4-dichlorophenoxyacetic acid (2.4-D), 0.5 mg/l naphthaleneacetic acid (NAA), and 0.5 mg/l 6-furfurylaminopurine (kinetin). Proembryos or somatic embryos were formed in all four species though germination and subsequent plantlet formation only occured in t. incarnatum and T. vesiculosum. Mature plants were obtained via somatic embryogenesis for T. incarnatum. The regenerated plants were fertile and had the normal diploid number of chromosomes (2n = 14).     

Pathology, soil transmission and characterization of cymbidium ringspot, a virus from cymbidium orchids and white clover (Trifolium repens)

Two strains of a virus, designated cymbidium ringspot virus (CyRSV), were isolated from cymbidium orchids and from Trifolium repens respectively in Britain. Experimentally infected cymbidiums developed slight chlorotic ring-mottle; T. repens developed flecks and mottling in the leaves, and slight stunting. Of 101 plant species tested, the cymbidium strain infected sixty-one (thirteen systemically) in twenty-three of thirty-five families; the clover strain infected sixty-four species (eighteen systemically) in twenty-two families. Both strains were propagated in Nicotiana clevelandii and assayed in Chenopodium quinoa. CyRSV was readily transmitted by inoculation of sap, and by foliage contact between plants, but not by the aphids Myzus persicae or Acyrtho-siphon pisum, nor through seed of T. incarnatum, Phaseolus vulgaris or N. clevelandii. Highly infective virus was released into soil from roots of infected N. clevelandii, and acquired by bait seedlings planted in such soil. Similar transmission occurred when purified virus was applied to the surface of sterilized soil containing bait plants; there was no evidence for any living soil vector. The virus was eliminated from 96 % of small cuttings taken from infected N. clevelandii plants grown at 35–37 °C for 9 wk. CyRSV was still infective in sap of N. clevelandii after dilution to 10?⁵-io–⁶ (only 2 × 10^_1 in cymbidium sap), or after 10min at 85–90 °C. It survived at least 10 months at c. 20 °C and more than 12 yr at 2 °C. Lyophilized sap was highly infective after over 13 yr at laboratory temperatures under high vacuum. Purified preparations made by clarification with n-butanol, followed by differential centrifugation and exclusion chromatography on controlled-pore glass beads, contained isometric particles c. 30 nm diam., with s°_20W= 137 S, and had a buoyant density in caesium chloride of 1–36 g/ml. The A 260/A 280 ratio was 1–55, and A max(26o)/A min(242) was 1–17. The virus contained c. 15 % of single-stranded RNA of mol. wt 1–7 × 10⁶; the nucleotide base ratios were: G27'8; A24/9; C2I-3; U26-I. There was one capsid polypeptide of mol. wt 43600. The virus was a good immunogen and a strongly reacting antigen in vitro; in Immunoelectrophoresis, each strain migrated as a single antigenic component towards the cathode. The cymbidium and clover strains were serologically closely related, although spurs were produced in immunodiffusion. No serological relationship was found to forty-three other isometric viruses, including eighteen tombusvirus isolates; CyRSV nevertheless shares many properties with tombusviruses, and we assign it provisionally to this group. The cryptogram is: R/r:1:7/15:S/S:S/O.     

Studies on the Relationship of Urdbean Leaf Crinkle Virus and its Vectors, Aphis craccivora and Acyrthosiphon pisum

Pre and Post-virus-acquisition starvation of Aphis craccivora Koch, and Acyrthosiphon pisum Harris resulted in appreciable increase in percentage of transmission of urdbean leaf crinkle virus. Highest transmission occured when aphids were starved for 90 min prior to virus-acquisition. A. pisum and A. craccivora on 20 and 80 min of post-acquisition starvation and 10 and 20 min of virus-acquisition, respectively inoculated more number of plants. Both the aphids transmitted the virus in probes lasting for one min. However, beyond 5 min of successive transfer on test plants loss in transmissibility was apparent. Viruliferous aphid off the plant retained the virus for a much longer period than on maize plant. The virus appears to be non-persistently borne in the aphids.     

RESOLUTION OF STRAWBERRY VIRUS COMPLEXES

Strawberry virus 4 produces vein chlorosis and necrosis on strawberry (var. Royal Sovereign) and slight chlorotic spotting on wild strawberry ( Fragaria vesca L.). No vector is known. Virus 5 produces leaf curling and vein necrosis on Royal Sovereign and F. vesca. It is transmitted by strawberry aphids ( Pentatrichopus fragaefolii Cock.) which have fed on an infected plant for 1 hr. or more and persists for about 1 hr. in the vector.
The names strawberry mottle, mild yellow-edge, crinkle, vein chlorosis and leaf-curl virus are proposed for strawberry viruses 1, 2, 3, 4 and 5 respectively.