The transmission of strawberry latent ringspot virus by Xiphinema diversicaudatutn (Nematoda)

Strawberry latent ringspot virus (SLRV) was found in diseased rose bushes growing in a glasshouse where the soil contained Xiphinema diversicaudatum (Micol). Adult female, adult male and juvenile X. diversicaudatum all transmitted the virus to cucumber seedlings, and nematodes kept without plants for 32 days after acquiring SLRV later transmitted it. When transferred to fresh plants every 2–4 days for 3 weeks, single nematodes transmitted up to three times; one nematode did not transmit until 19 days after the transfers began. One adult X. diversicaudatum, out of 141 tested, transmitted both SLRV and arabis mosaic virus. In all respects X. diversicaudatum behaved as a vector of SLRV as it does as a vector of arabis mosaic virus.     

Effect of nutritional stress on control of Longidorus elongatus by nematicidal chemicals

Studies were made to determine the effect of starvation on the susceptibility of Longidorus elongatus to nematicidal chemicals. In vitro studies, using stylet protraction as a measure of susceptibility, showed that the EC₅₀ for nematodes exposed to oxamyl for 5 min was 6-5 μg cm^-3 for starved nematodes and 7-7 μg cm^-3 for well-fed specimens. In pot studies aldicarb and oxamyl, applied at half the normal commercial rates, decreased the numbers of starved, but not of well-fed, L. elongatus. In a field study aldicarb, oxamyl and the fungicide quintozene decreased the numbers of starving and well-fed L. elongatus. Oxamyl was more effective than aldicarb and the numbers of L. elongatus were decreased more in plots previously fallowed than in those cropped with a grass/clover mixture. A significant interaction between oxamyl treatment and initial crop was found. Starved L. elongatus appear to be more susceptible to carbamate nematicides than well-fed ones. Quintozene decreased numbers of L. elongatus more, and for a longer period of time, than either of the systemic nematicides. The mode-of-action of the nematicidal chemicals is discussed.     

Tests for transmission of cherry leaf roll virus using Longidorus, Paralongidorus and Xiphinema nematodes

The ability of 10 nematode species to transmit three strains of cherry leaf roll virus (CLRV) was tested by three methods: (1) virus-infected source plants and virus-free bait plants were grown concurrently in nematode-infested soil, (2) as for (1) but virus source plants were removed before bait plants were planted, and (3) nematodes were extracted from soil after access to virus source plants, and were added to pots containing bait plants. The occurrence of galls on roots showed that nematodes fed both on source and on bait plants in all experiments and, in some experiments, CLRV was detected by direct assays (slash tests) of Longidorus elongatus, L. leptocephalus and Paralongidorus maximus. Although the nematodes readily transmitted control viruses, for which they are known to be vectors, CLRV was detected by root assays in only a few bait plants exposed to L. elongatus, L. macrosoma, Xiphinema diversicaudatum or L. leptocephalus + X. vuittenezi in tests by method 1. The recovery of CLRV in these tests is interpreted as being due to contamination. These results add to the increasing circumstantial evidence against the involvement of nematodes in the transmission of CLRV. Other possible mechanisms of spread are discussed.     

A comparison of the effectiveness of the four British virus vector species of Longidorus and Xiphinema

The frequency with which the four virus-vector species of longidoroid nematodes occurring in Britain transmitted their associated plant viruses were compared in a series of experiments using a standard procedure. In these tests Xiphinema diversicaudatum proved an effective vector of British isolates of arabis mosaic virus and strawberry latent ringspot virus and Longidorus attenuatus of an isolate of tomato black ring virus from England. In comparison, isolates of raspberry ringspot virus and tomato blackring virus from Scotland and of raspberry ringspot virus from England were transmitted much less readily by their respective vectors, L. elongatus and L. macrosoma. These differences in ability to transmit virus were not related to differences in feeding access on the virus source- or bait-plants, in the extent to which virus was retained within the nematode feeding apparatus or in the frequency with which virus was recovered from Longidorus in concurrent slash tests. Three Scottish isolates of raspberry ringspot and tomato black ring viruses were transmitted equally infrequently by two populations of L. elongatus and the frequency with which virus was transmitted was not greatly increased when the species of source or bait plants was changed.     

Transmission of the hop strain of arabis mosaic virus by Xiphinema diversicaudatum*

Hop plants became infected with the hop strain of arabis mosaic virus (AMV(H)) when grown in hopfield and woodland soil in which infected plants had been growing. Infection occurred in soil infested with the dagger nematode Xiphinema diversicaudatum, but neither in uninfested soil nor in soil previously heated to kill nematodes. X. diversicaudatum transferred direct from hop soils transmitted AMV(H) to young herbaceous plants and to hop seedlings; some of the hop seedlings developed nettlehead disease. A larger proportion of plants was infected using X. diversicaudatum obtained from a woodland soil and then given access to the roots of hop or herbaceous plants infected with AMV(H). AMV(H) was transmitted by adults and by larvae, in which the virus persisted for at least 36 and 29 wk, respectively. Difficulties were encountered in detecting AMV(H) in infected hop plants, due partly to the delay in virus movement from roots to shoots. Infection of hop shoots was seldom detected until the year after the roots were infested and sometimes nettlehead symptoms did not appear until the third year. Isolates of arabis mosiac virus from strawberry did not infect hop. The results are discussed in relation to the etiology and control of nettlehead and related diseases of hop.     

Ingestion,Retention, and Transmission of two Strains of Raspberry Ring-spot Virus by Longidorus macrosoma

The transmission of two strains of raspberry ringspot virus (RRV) by small numbers of nematodes was compared. A strain of RRV from Scotland (RRV-S), originally found in the field associated with Longidorus elongatus, was transmitted frequently by L. elongatus but only once by L. macrosoma. A strain from England (RRV-E) associated with L. macrosoma in the field was transmitted infrequently by each species of nematode. The reasons why L. macrosoma infected only a small proportion of bait plants with virus were investigated, and it was found that most of the nematodes tested had fed on the source plants and many had ingested virus. Most nematodes exposed to RRV-E or RRV-S had fed on the roots of the bait plants and, when thin sections were examined by electron microscope, had retained particles (thought to be those of the virus) in the region of the anterior odontostyle, Thus, most nematodes seem to have had ample opportunity to transmit virus, and the low frequency of transmission may have been due to a failure of the virus particles to be released from the site of retention or to a lack of infectivity of the virus when L. macrosoma was the vector and Petunia hybrida was the host.     

Observations on the occurrence, pathogenicity and control of Pratylenchus vulnus, P. thornei and Xiphinema diversicaudatum associated with glasshouse roses

About half the nurseries and glasshouses in the Lea Valley of Hertfordshire were found to be infested with either or both Pratylenchus vulnus and Xiphinema diversicaudatum. The latter nematode probably occurred naturally in the soil on which the glasshouses were built but P. vulnus is thought to have been imported with rose rootstocks and is not known to occur outdoors in Britain. Both nematode species can cause decline of roses and even small numbers seem harmful. Numbers of P. vulnus can increase greatly, especially on rootstocks of Rosa canina, although numbers were often smaller when roses were severely damaged than on healthier crops. Numbers of X. diversicaudatum increased more slowly and R. canina was a better host for it than R. chinensis. X. diversicaudatum seemed to respond less quickly than P. vulnus to reduced host-plant vigour. The two rootstocks exhibited different host-status for the two species of nematode and cuttings of various rose cultivars showed different host-status to P. thornei, another species of lesion nematode which is not known to be pathogenic to roses. Two applications of dibromochloropropane liquid at the rate of 70 l/ha in a large volume of water maintained nematode densities at an acceptably low level, and growers who adopted this treatment as a supplement to pre-planting steam sterilisation and/or DD (dichloro-propane: dichloropropene) prolonged the productive life of their crops for several years.     

Host status of some plants for Xiphinema diversicaudatum (Micol.) and their susceptibility to viruses transmitted by this species

Forty plant species were grown in pots containing viruliferous Xiphinema diversicaudatum (Micol.) for 15 wk to assess the host range of the nematode in relation to infection with arabis mosaic (AMV) and strawberry latent ring-spot (SLRV) viruses. Host status for the nematode was determined mainly from changes in total populations, but the presence of eggs in the uteri of females and changes in the numbers of adults provided additional criteria. The nematode multiplied on relatively more woody perennials than on herbaceous crop plants or weeds. Chrysanthemum coronarium was the only plant on which numbers declined significantly below those on the controls. Most plant species became infected with either AMV or SLRV. Neither virus was detected in eight out of thirteen species of trees and shrubs although four were good hosts for the nematode. Galling or distortion of the terminal region of fine feeder roots, associated with X. diversicaudatum feeding, was seen on many of the experimental plants.     

Numbers of mononchid nematodes in soils sown to cereals and grasses

The distribution of mononchid nematodes Mylonchulus sigmaturus, Mylonchulus brachyuris, Clarkus papillatus, Prionchulus punctatus and Anatonchus tridentatus found in grassland and arable soils in England were studied. Mononchid nematode populations in excess of 5% of the total soil nematode population and in biomass more than 2% of the total soil nematode population were rare. No relationship was found between numbers of mononchids and those of other soil nematodes. In soils sown to spring barley there was no difference in the total numbers of female M. sigmaturus, C. papillatus or A. tridentatus found throughout the year. Juveniles of C. papillatus were most common while those of M. sigmaturus were least common. In grass leys and arable soil P. punctatus numbers were extremely low. Neither phorate nor oxamyl applied at 1.12 kg a.i./ha in March affected mononchid numbers. Although oxamyl at 11.2 kg a.i./ha did not affect the adult female numbers, the numbers of juvenile mononchids were reduced by 38% and 50% in June and November respectively.     

The geographical distribution of Xiphinema and Longidorus nematodes in the British Isles and Ireland

The distribution of Xiphinema and Longidorus species in the British Isles and Ireland were mapped by means of a computer programme from 5527 records. Longidorus caespiticola, L. elongatus, L. goodeyi and L. leptocephalus were found to be widespread among a wide range of vegetation types, particularly in grassland and arable crops; L. macrosoma and L. profundorum were recorded frequently in southern and central England, and L. attenuatus in East Anglia; L. vineacola was recorded once only in Eire. Paralongidorus maximus was recorded from two localities in eastern Scotland and one in southern England, on each occasion from intensively cultivated ground. Four species of Xiphinema were recorded but only X. diversicaudatum is widespread although apparently with a northerly limit in central Scotland; X. diversicaudatum was equally prevalent in arable crops, grassland and deciduous woodland; X. coxi and X. vuittenezi were recorded from a few scattered localities in England, and the few records for X. mediterraneum were all from Kent, south east England. All species were found in a wide range of soil types but mostly in light soils with a large sand fraction. Many of the species occurred together in mixed populations; L. caespiticola was most frequently associated with X. diversicaudatum and with L. leptocephalus and L. elongatus. X. diversicaudatum also was commonly found with L. goodeyi. Viruses were transmitted in laboratory bait tests in only eighteen of 325 soil samples containing X. diversicaudatum and four of 265 containing L. elongatus.     

Nuclear changes induced by the nematodesXiphinema diversicaudatum andLongidorus elongatus in root-tips of perennial ryegrass,Lolium perenne

Summary The DNA content and size of individual nuclei from galls of perennial ryegrass root-tips induced byX. diversicaudatum andL. elongatus were measured. Feeding byX.diversicaudatum increased the DNA content of the nuclei by varying amounts. No regular doubling pattern of the DNA content was discernible. The DNA values varied up to between 32–64C. Generally the size of the nuclei was not increased, although some were larger than control nuclei. The modified nuclei probably have an altered metabolic function, which increases the food value of the gall to the nematode. Some bi-nucleate cells were also observed, which probably result from mitosis without cytokinesis. A preliminary examination of nuclei from galls induced byL. elongatus revealed similar nuclear changes, but no bi-nucleate cells were found. Editor's note: Awarded the Viviane Maggi prize for the best paper presented by a beginner at the Annual Meeting of the Histochemistry and Cytochemistry Section of the Royal Microscopical Society in April 1981. This paper is a full report of that presentation.     

The association of Xiphinema diversicaudatum (Micoletsky) with strawberry latent ringspot and arabis mosaic viruses in a raspberry plantation

An outbreak of strawberry latent ringspot virus (SLRV) in a plantation of Mailing Jewel raspberry coincided with the greatest abundance of the nematode vector, Xiphinema diversicaudatum. Arabis mosaic virus (AMV) was not detected in the crop but was, together with SLRV, in many weed species present. AMV was transmitted through the seed of Poa annua, Capsella bursa-pastoris and Senecio vulgaris and SLRV through the seed of Mentha arvensis. X. diversicaudatum were more numerous within the rows than between them and vertical sampling showed that most occurred between 4 and 12 in depth in both locations. Monthly sampling showed that egg laying occurred from April to July; populations increased to a peak in late autumn but declined during the winter, resulting in about a twofold annual increase in numbers. Females, males and juveniles transmitted AMV and SLRV to cucumber seedlings, and in the absence of plants the nematode retained AMV for 112 days and SLRV for 84 days.     

Populations of Longidorus elongatus (Nematoda: Longidoridae) in two New Zealand soils under grazed pasture

Abstract

Information on the “needle nematode” Longidorus elongatus in New Zealand pastures has hitherto been limited. Monthly sampling of Pukepuke black sand and Manawatu fine sandy loam yielded L. elongatus populations up to 87 500 m^?2 (February) and 21 600 m^?2 (August). First stage juveniles appeared when soil temperature exceeded 15°C; females dominated populations every month and males were rare. Differences in abundance between soils may reflect the pore space available to these relatively large nematodes. Specimens survived 24 weeks storage at 5 and 15°C. While abundance tended to decline with depth, at 30–40 cm depth in Pukepuke sand, numbers increased, perhaps through the impact of groundwater levels on rooting patterns. In Pukepuke sand, plant species were associated with significantly different populations of L. elongatus, with Trifolium repens and T. subter‐raneum supporting more than Lolium perenne. More L. elongatus were found in grazed pasture with lower plant available P.     

Nematode vectors of plant viruses in New Zealand

Abstract

A survey of virus vector nematodes based on a total of 320 published and recent records revealed that 3 Longidorus, 1 Paralongidorus, 6 Xiphinema, 2 Trichodoms, and 3 Paratrichodorus species are present in New Zealand. Xiphinema diversicaudatum and Paratrichodorus minor are the most common and widespread species. Longidorus orongorongensis, an undescribed Paralongidorus species and Trichodoms cottieri appear to be endemic species, and Longidorus taniwha, Paratrichodorus lobatus and probably one (still unidentified) species of the X. americanum group are also indigenous to New Zealand. Most of the other species (e.g. Longidorus elongatus, X. diversicaudatum, and Trichodoms primitivus) were probably introduced from Europe. Maps showing the distribution of the 15 species in New Zealand, and illustrations and tables of the morphological characters for species identification, are presented. Several species are of economic importance through direct damage they cause to cultivated plants, and all can be considered potential vectors of plant viruses.     

Soil-borne viruses occurring in nursery soils and infecting some ornamental species of Rosaceae

Soil from twelve of twenty-six sites on ten nurseries growing ornamental trees and shrubs contained viruliferous nematodes, transmitting arabis mosaic virus (AMV) at eight sites, tomato black ring virus at three and tobacco rattle virus at one site. Tobacco necrosis virus was detected at two sites. Xiphinema spp. were found at nine sites, Longidorus spp. at sixteen and Trichodorus spp. at thirteen sites. Thirty-one ornamental species of Rosaceae were tested for virus infections and AMV was found in Spiraea bumalda, Spiraea japonica alba and Kerria japonica.     

Seed-transmission in the ecology of nematode-borne viruses

Virus-free populations of vector nematodes can acquire tomato black ring (TBRV), raspberry ringspot (RRV) and arabis mosaic (AMV) viruses from weed seedlings grown from virus-carrying seed. When soils from fields where nematode-borne viruses occurred naturally were air-dried to kill vector nematodes and then moistened, TBRV and RRV occurred commonly in the weed seedlings that grew, but AMV occurred only rarely. Similar tests did not detect tobacco ringspot, grapevine fanleaf or tobacco rattle viruses in weed seeds in the single soil studied in each instance, although these three viruses are also seed-borne in some of their hosts. Many weed species, when infected experimentally, readily transmit TBRV and RRV to their seed, but the viruses were much commoner in naturally occurring seed of some of these species than of others. These discrepancies between the frequency of seed-transmission of viruses from experimentally infected plants and the extent of natural occurrence of infected seed seem largely to reflect the host preferences of the vectors. Infective Longidorus elongatus kept in fallow soil retained TBRV and RRV only up to 9 weeks. When weed seeds in the soil were then allowed to germinate, the nematodes reacquired virus from the infected seedlings. Some weed species were better than others as sources of virus. Persistence of these viruses in fields through periods of fallow or fasting of the vector therefore depends on a continuing supply of infected seedlings produced by virus-containing weed seeds. This is probably less true of viruses like AMV and grapevine fanleaf, which persist for 8 months or more in their vectors (Xiphinema spp.). A few seeds containing TBRV and RRV were found in soils free of vector nematodes, suggesting that the viruses are disseminated in weed seed. This probably explains how TBRV and RRV have reached a large proportion of L. elongatus populations in eastern Scotland.     

The multiplication of Longidorus elongatus (de Man) on different host plants with reference to virus transmission

In pot tests, Longidorus elongatus (de Man) populations increased rapidly on Fragaria vesca, Stellaria media, Mentha sativa and Lolium perenne and slightly on Brassica rapa (turnip var. White globe) and on Ribes nigrum; but on Rubus idaeus populations declined at the same rate as in fallow soil. Raspberry ringspot (RRV) and tomato black ring (TBRV) viruses were readily transmitted by L. elongatus to S. media, TBRV only was transmitted to L. perenne, and M. sativa became infected with neither virus. RRV was transmitted to R. idaeus var. Mailing Jewel, on which the nematode fed but evidently did not reproduce. In a field experiment L. perenne and Fragaria ananassa (strawberry) vars. Huxley and Redgauntlet were shown to be good hosts for L. elongatus and R. idaeus (raspberry) vars. Lloyd George and Mailing Jewel were confirmed as non-hosts, together with Rheum rhaponticum (rhubarb). Two periods of egg laying were detected each year in the Redgauntlet strawberry plots and these coincided with the growth of young roots during the late spring and autumn. The wide host range of L. elongatus among cultivated plants and weeds, together with its ability to survive long periods without food, precludes the use of crop rotation as a means of control. Chemical soil sterilization is considered the most effective means of control because it can greatly decrease L. elongatus populations and as the nematode multiplies slowly, even on favourable host plants, several years are likely to elapse before populations become large enough to spread viruses effectively from infector plants to susceptible crops.     

Effects of Oxime Carbamate Nematicides on Development of Heterodera schachtii on Sugarbeet

Treatment of sugarbeet, Beta vulgaris L., with aldicarb, aldicarb sulfoxide, or aldicarb sulfone 10 days after plants were inoculated with Heterodera schachtii prevented development of the nematode, but second-stage larvae penetrated the roots. These chemicals had no measurable effects on nematodes in plants treated 15 days after inoculation. The tests established that soil treatments of aldicarb are directly or indirectly lethal to larvae developing within roots of sugarbeet. Heterodera schachtii failed to develop on root slices of red table beet grown in soil treated with aldicarb or aldicarb sulfoxide. Similar treatment of plants with aldicarb sulfone or oxamyl did not affect subsequent development of H. schachtii on root slices of treated plants.     

Studies on the relation between density of Longidorus elongatus and growth of sugar beet, with supplementary observations on Trichodorus spp

Longidorus elongatus attacks sugar beet on light sandy soils in the West Midlands. Severely damaged plants may die or recover, producing fanged roots. Up to 335 L. elongatus/200 g of soil were found around attacked seedling plants and were often visible to the unaided eye on the roots of freshly lifted plants. Five experiments were made and regressions computed of yield of beet, total numbers of plants and numbers of normal and fanged beet on L. elongatus numbers. Between 24% and 50% of roots per 100 L. elongatus/ 200 g were killed or became fanged. Corresponding figures for loss of plant were between 7.5% and 33%. Estimated loss of yield varied between 0.8 and 7.3 tons (2.0–18.3 t/ha)/acre/100 L. elongatus/200 g, the former where the potential yield was high and the latter where poor growing conditions hindered recovery. Large numbers of Trichodorus occurred in two trial sites and there is some evidence of competition between the two genera. Significant negative regressions for Trichodorus spp. were obtained in one trial suggesting a loss of 12% total and 17% normal roots per 100 Trichodorus spp./200 g.     

Direct and indirect effects of a thrips‐transmitted Tospovirus on the preference and fitness of its vector,Frankliniella fusca

Phytoviruses including tospoviruses are known to affect the behavior and fitness of their vectors both positively and negatively. In this study, we investigated the effects of Tomato spotted wilt virus (TSWV) (family Bunyaviridae, genus Tospovirus) infection on the fitness and feeding ability of tobacco thrips, Frankliniella fusca (Hinds) (Thysanoptera: Thripidae) using peanut, Arachis hypogaea L. (Fabaceae), as a host. Potentially viruliferous F. fusca laid more eggs than non‐viruliferous F. fusca. In contrast, fewer potentially viruliferous F. fusca developed into adults and required a longer developmental time than non‐viruliferous F. fusca, indicating a direct negative effect of the virus on thrips fitness. In addition, no‐choice feeding tests indicated that non‐viruliferous F. fusca fed more rapidly than potentially viruliferous F. fusca. Typically, phytovirus infections are known to enhance the availability of vital nutrients such as free amino acids in infected host plants and to affect other important physiological processes negatively. Free amino acids are known to play a vital role in egg production and development. Further investigations in this study revealed that leaflets of infected plants had ca. 15 times more free amino acids than non‐infected leaflets. TSWV‐infected leaflets were used to rear potentially viruliferous thrips. Higher amino acid levels in TSWV‐infected leaflets than in non‐infected leaflets could have contributed to increased oviposition by potentially viruliferous F. fusca compared to non‐viruliferous F. fusca. Taken together, these results suggest that increased concentrations of free amino acids in TSWV‐infected plants might serve as an incentive for thrips feeding on otherwise unsuitable hosts, thereby facilitating TSWV acquisition and transmission.