Probing by aphids in the leaf tissues of resistant and susceptible sugar beet

The probing of Aphis fabae and Myzus persicae in the leaves of sugar beet with inherited resistance or susceptibility to aphids was studied by microscopic examination of samples of whole leaves, prepared after 48 h exposure to adult aphids at approximately three aphids cm^-2.The density of saliva stylet-sheaths left by the aphids (cm^-2) and the proportion reaching phloem differed between sugar beet stocks and were inversely associated. Differences in resistance between stocks could not, however, be related directly to either. All beet stocks examined were probed freely. Seasonal differences in sugar beet grown in the glasshouse affected the proportion of sheaths reaching the phloem, but the differences between beet stocks were similar at all times.The densities of sheaths left by different clones of M. persicae corresponded with the aphids' response to sugar beet as a host plant. Among aphid clones which readily colonize sugar beet, the densities of stylet sheaths which reached phloem suggested that the adults of both A. fabae and M. persicae gained sufficient access to sieve tubes to satisfy their nutritional needs. The phloem of sugar beet from the glasshouse was always within the estimated maximum depth to which the aphids probe; but, in leaves from the field, it appeared that the phloem might be inaccessible to young M. persicae in the sugar beet crop during late summer.

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Zusammenfassung Das Proben von Aphis fabae und Myzus persicae in Blättern von Zuckerrüben mit erblicher Blattlausresistenz bzw.-anfälligkeit wurde untersucht durch mikroskopische Durchmusterung von Speichelscheiden in Proben von ganzen Blatt. Rübenblätter wurden mit genähert drei adulten Läusen cm^-2 besetzt und nach 48 Stunden quergeschnittene Streifen der Blätter in Alkohol fixiert, gefärbt und mit der Unterseite nach oben auf Objektträgern eingeschlossen.23890 Speichelscheiden wurden registriert. Die Dichte der Scheiden von M. persicae (cm^-2) und der Anteil der das Phloem erreichenden Scheiden (SRP) unterschieden sich signifikant zwischen den Rübenstämmen. Bei A. fabae ergaben sich entsprechende, aber nicht gesicherte Unterschiede. Scheidendichte und Prozentsatz SRP waren gegenläufig, zwei Rübenstämme zeigten eine hohe Scheidendichte, zwei andere hatten weniger Scheiden, aber einen höheren Prozentsatz SRP. Diese Gruppierung der Stämme korrespondierte aber nicht mit ihrer Blattlausresistenz. Aus der Scheidendichte ergab sich, dass M. persicae und A. fabae auf allen geprüften Rübenstämmen, resistenten und anfälligen, unbehindert probten, so dass jede Laus das Phloem durchschnittlich etwa viermal am Tag erreichte. Ein Klon von M. persicae, der sich an Rüben nicht entwickelt, hinterliess weniger Scheiden in den Blättern aller Stämme.Der Anteil von SRP war bei Prüfungen im März grösser als im November. Dieser Unterschied war besonders deutlich bei Scheiden von Larven, die im übrigen zu allen Zeiten das Phloem weniger oft erreichten als ihre Eltern. Messungen des Abstandes von der unteren Blattfläche zum Phloem ergaben, dass das Phloem den Läusen in Gewächshaus-Zuckerrüben immer zugänglich war. M. persicae-Larven konnten jedoch in Blättern von Freilandrüben das Phloem nicht erreichen.

     

Breeding for resistance to infection with yellowing viruses in sugar beet

Differences in resistance to infection with beet yellows virus (BYV) and beet mild yellowing virus (BMYV) have been observed in virus-tolerant sugar-beet breeding material. The results of glasshouse virus-susceptibility tests usually agreed well with those of field experiments in which plants were exposed to artificial, or natural, infestation with viruliferous aphids. Breeding lines and varieties, which showed resistance to BYV when Myzus persicae Sulz, was used as vector, generally showed a similar resistance to this virus when Aphis fabae Scop. was used. Varieties which were resistant to infection with one virus were not necessarily resistant to the other, although some showed resistance to both BYV and BMYV. Preliminary results suggest that resistance to infection may be controlled by recessive genes which occur widely in sugar-beet cultivars. The mechanism of this form of resistance is not understood, but it does not appear to be closely associated with resistance to the aphid vectors of the viruses. The observed differences in resistance to infection demonstrate the possibility of breeding a sugar-beet variety in which two forms of resistance to virus yellows, tolerance and resistance to infection, are combined.     

A controlled environment study of the effect of leaf physiological age on the movement of apterous Myzus persicae on sugar-beet plants

Apterous Myzus persicae were found to move frequently from leaf to leaf on sugar-beet plants in controlled environment conditions. It is suggested that aphid movement can be related to changes in the rate and content of translocate flow during leaf development. These changes make newly-emerged leaves nutritionally favourable to colonising aphids and make expanding leaves slowly wane in favourability during the process of ‘sink to source’ conversion leading to aphid dispersal from the leaf. Variation in temperature was not found to alter the rate of aphid movement or the period (measured in thermal time) that aphids spent on particular leaves. However, the lower temperature was found to increase the rate of aphid development, aphid size and fecundity; these effects could also be due to nutritional factors. This dispersal behaviour may be a tactic to maximise food intake by a polyphagous aphid and increase the probability that nymphs are deposited on nutritionally-favourable leaves. The implications of the interleaf dispersal of apterous M. persicae for within- and between-plant spread of beet yellows virus (BYV) and beet mild yellowing virus (BMYV) are discussed.     

Effects of sucrose sprays and darkness on aphid colonization of sugar beet and on aphid transmission of yellowing viruses

In the glasshouse, adult, apterous Myzus persicae (Sulz.) and Aphis fabae Scop, settled better and deposited more larvae on sucrose-sprayed sugar-beet plants than on water-sprayed plants. M. persicae settled badly and deposited few larvae on plants that were kept in the dark before or after infestation. The effects of darkness on aphids were reduced by spraying the host plants with 10% solutions of sucrose before infestation. Viruliferous M. persicae transmitted beet yellows virus (BYV) and beet mild yellowing virus (BMYV) less efficiently to dark-treated plants than to those grown in normal daylight. Spraying sugar beet with sucrose before inoculation with viruliferous M. persicae increased the proportion of successful BYV transmissions but only when the plants were dark-treated. The effects of sucrose and darkness on settling and larviposition of aphids and on virus transmission may be related to changes in the concentration of carbohydrates, particularly sugars, in the leaves.     

Glasshouse tests and field selection for heritable resistance to Myzus persicae in sugar beet

A technique is described to assess and select in the field for resistance to Myzus persicae in sugar beet. Lines developed by this method were aphid resistant when compared with commercial cultivars. The resistance was inherited by progeny lines. Variation between beet lines in glasshouse tests suggested segregation of a number of resistance genes with varying effects. Integration of selection for aphid-resistance and agronomic performance during beet breeding is discussed.     

Inherited resistance of sugar beet to aphid colonization

Results of glasshouse experiments have confirmed that inbred lines of sugar beet differ in each of three types of resistance to Myzus persicae Sulz. and Aphis fabae Scop., namely: resistance to settling, resistance to multiplication, and tolerance. Resistance to multiplication was not invariably associated with resistance to settling, although plants of some lines showed both forms of resistance. Plants that were resistant to settling of alatae were not always resistant to apterae of the same species, and there was not a close relationship between resistance to M. persicae and to A. fabae. The mechanisms involved in resistance to aphids in sugar beet are not understood. Progenies of plants, selected for resistance to aphids from inbred lines, were often more resistant than progenies of unselected plants. Inheritance of each type of resistance is probably polygenic. The potential value of the different kinds of resistance, in reducing direct feeding damage and controlling the spread of virus yellows in the field, is discussed. The ultimate breeding objective is to produce commercial varieties in which appropriate kinds of resistance to aphids are combined with resistance to virus yellows. The use of such varieties would reduce the need to control aphids in the field by applications of chemicals.     

The use of monoclonal antibodies to detect beet mild yellowing virus and beet western yellows virus in aphids

Information on infectivity of the aphids which invade sugar beet root crops each Spring is required for forecasting incidence and providing advice on control of virus yellows. Monoclonal antibodies, produced in the USA to barley yellow dwarf virus (BYDV) and in Canada to beet western yellows virus (BWYV), were used to distinguish between sugar-beet-infecting strains of the luteovirus beet mild yellowing virus (BMYV), and the non-beet-infecting strains of the closely-related BWYV in plant and aphid tissue. Totals of 773 immigrant winged Myzuspersicae and 124 Macrosiphum euphorbiae were caught in water traps in a crop of sugar beet between 25 April and 5 August 1990. Using the monoclonal antibodies and an amplified ELISA, 67%M. persicae and 19%M. euphorbiae were shown to contain BWYV; 8%M. persicae and 7%M. euphorbiae contained BMYV. In studies with live winged aphids collected from the same sugar beet field during May, 25 of 60 M. persicae and two of 13 M. euphorbiae transmitted BWYV to the indicator host plant Montia perfoliata; two M. persicae and two M. euphorbiae transmitted BMYV. In another study three of 65 M. persicae and one of three M. euphorbiae in which only BWYV was detected, transmitted this virus to sugar beet.     

Some factors affecting the concentrations of sugars in leaves of inbred sugar-beet lines

In the glasshouse, large differences in concentrations of glucose and sucrose were observed between leaves of inbred sugar-beet lines that are known to differ from each other in resistance to pests and diseases. Differences between these lines in concentrations of fructose, glucose and sucrose were more pronounced in the petiole than in the lamina. The concentrations of glucose and fructose were lower in the first two leaves (primary leaves) than in those produced subsequently (secondary leaves); both types of leaf contained similar concentrations of sucrose. Secondary leaves from plants that had been kept in darkness for 17 h contained less fructose, glucose and sucrose than those of plants kept in sunlight for 5 h. Longer period of darkness lowered the concentrations of monosaccharides in the leaves further but did not affect the sucrose content. Primary and secondary leaves from the same sugar-beet plant often differ in non-race-specific resistance to pests and diseases; and darkness can affect suscetibility of beet to downy mildew and to the aphid Myzus persicae. The results of the sugar determinations therefore support the hypothesis that the concentrations of certain carbohydrates in sugar-beet leaves are important in non-race-specific resistance to pests and diseases.     

Resistance induction and herbivore virulence in the interaction between Myzus persicae (Sulzer) and a major aphid resistance gene (Rm2) from peach

In gene-for-gene host–enemy interactions, monogenic plant resistance results from pathogen recognition that initiates the induction of plant defense responses. Schematically, as the result of the on/off process of recognition, phenotypic variability in enemy virulence is expected to be qualitative, with either a failure or a success of host colonization. We focussed on a major gene from peach conferring avoidance resistance against the green peach aphid Myzus persicae. Measurements of herbivore density and time-dependent aspects of resistance induction were examined, as well as variability in the aphid’s ability to exploit the resistant host. Varying densities of infestation did not provoke differences in the aphid’s tendency to leave a plant, and a single aphid was sufficient to elicit a response. Similarly, the duration of infestation did not affect the aphid response. A brief aphid feeding time of 3 h triggered induced resistance, which became effective between 24 and 48 h after the initial attack. Induced resistance decayed over time in the absence of additional infestation. Thirty aphid genotypes collected from natural populations were tested in the laboratory. No clone could colonize the resistant host, suggesting that all of them triggered the induction of effective plant defense responses. However, we detected significant quantitative variation among clones in the tendency of aphids to leave plants. These results improve our understanding of induced resistance as a dynamic phenomenon and suggest that the potential for aphids to adapt to a major plant resistance gene may depend on factors other than the mere capacity to evade recognition.     

Genetic approaches to sustainable pest management in sugar beet (Beta vulgaris)

Sugar beet (Beta vulgaris) is an important arable crop, traditionally used for sugar extraction, but more recently, for biofuel production. A wide range of pests, including beet cyst nematode (Heterodera schachtii), root‐knot nematodes (Meloidogyne spp.), green peach aphids (Myzus persicae) and beet root maggot (Tetanops myopaeformis), infest the roots or leaves of sugar beet, which leads to yield loss directly or through transmission of beet pathogens such as viruses. Conventional pest control approaches based on chemical application have led to high economic costs. Development of pest‐resistant sugar beet varieties could play an important role towards sustainable crop production while minimising environmental impact. Intensive Beta germplasm screening has been fruitful, and genetic lines resistant to nematodes, aphids and root maggot have been identified and integrated into sugar beet breeding programmes. A small number of genes responding to pest attack have been cloned from sugar beet and wild Beta species. This trend will continue towards a detailed understanding of the molecular mechanism of insect–host plant interactions and host resistance. Molecular biotechnological techniques have shown promise in developing transgenic pest resistance varieties at an accelerated speed with high accuracy. The use of transgenic technology is discussed with regard to biodiversity and food safety.     

Resistance to attack by Brevicoryne brassicae among plants of Brussels sprouts

Because they remained almost uncolonized by the cabbage aphid (Brevicoryne brassicae (L.)) throughout the growing season, plants of Brussels sprouts were singled out in each of 4 years, from plots heavily infested with the aphid, as possibly being resistant to attack. Clones of these plants were established from cuttings and tested in a controlled environment by inoculation with B. brassicae and later, in the field, by natural infestation. The tests confirmed that some of the plants were resistant to the aphid, and the most resistant of those from the first year of the work proved at least as resistant as any subsequently found. The resistance was expressed as antibiosis, but in the field host non-preference was also shown by incoming winged aphids. The possibility that biotypes of B. brassicae might exist, to which the resistant sprout clones were not necessarily resistant, was investigated using B. brassicae collected from sprouts from each of several areas in England. Eight sprout clones, seven of which were known to be resistant, and the other susceptible, to B. brassicae from Wellesbourne were tested with these other B. brassicae. The results showed that biotypes of the aphid, with differing abilities to colonize respective sprout clones, existed in each area, and of the seven sprout clones resistant to the Wellesbourne aphid, only one appeared never to be fully susceptible to one or more of the other biotypes of B. brassicae.     

Some effects of spraying with thiabendazole on the susceptibility of sugar beet to yellowing viruses and on their vector, Myzus persicae (Sulz.)

In a field experiment fewer sugar-beet plants became infected with aphid-transmitted yellowing viruses in plots that had been sprayed with solutions of thiabendazole lactate than in water-sprayed plots, after exposure to natural infestation with aphids. Subsequent glasshouse tests showed that foliar sprays of o·o1 % thiabendazole lactate in water significantly reduced the proportion of inoculated sugar-beet plants which became infected with beet yellows virus (BYV) or beet mild yellowing virus (BMYV) after inoculation with viruliferous Myzus persicae (Sulz.). This effect on virus transmission was not apparently due to a direct insecticidal action of thiabendazole, because adult aphids usually survived equally well on sprayed and unsprayed plants. Treatment of test plants with thiabendazole did not affect the transmission of beet mosaic virus to them by M. persicae. The fecundity of M. persicae was greatly reduced by transferring them to plants which had been sprayed with thiabendazole or by spraying them with thiabendazole before transfer to unsprayed plants. The fertility of adult Aphis fabae Scop, was also reduced by spraying with thiabendazole. The mechanisms whereby thiabendazole affected fecundity of aphids and transmission of viruses are not understood.     

The effect of co-infestation by conspecific and heterospecific aphids on the feeding behaviour of <Emphasis Type="Italic">Nasonovia ribisnigri</Emphasis> on resistant and susceptible lettuce cultivars

Aphid saliva can suppress the blocking of sieve elements, a reaction that plants employ to inhibit aphid feeding, but aphid saliva can also elicit plant defence responses. Such plant responses might affect interactions between different aphid species and intraspecifically, e.g. among different biotypes. The objectives of our study were to investigate if feeding behaviour and performance of two biotypes of the lettuce aphid Nasonovia ribisnigri are affected by (1) feeding by the other biotype and (2) feeding by the green peach aphid Myzus persicae or the potato aphid Macrosiphum euphorbiae. Additionally the effect of feeding in a group was studied. All experiments were performed on both a resistant and an isogenic susceptible lettuce cultivar. Feeding or probing by conspecific or heterospecific aphids had different effects on Nasonovia ribisnigri biotypes. Aphids were only slightly affected by feeding or probing of the same biotype on both susceptible and resistant lettuce. N. ribisnigri virulent biotype Nr:1 suppressed the resistance against Nr:0 in the resistant cultivar. In contrast, defence was induced by Nr:1 against Nr:0 in susceptible lettuce. Co-infestation by M. euphorbiae and M. persicae had minor effects on Nr:0. Defence against Nr:1 was induced on both susceptible lettuce and resistant lettuce by Nr:0 and M. euphorbiae. Additionally, M. persicae induced defence in resistant lettuce against Nr:1. Effectors in the saliva of Nr:1 aphids are likely responsible for the defence suppression in lettuce. Identification of these effectors could lead to a better understanding of the mechanism of virulence in N. ribisnigri.     

Blockage of stylet tips as the mechanism of resistance to virus transmission by Aphis gossypii in melon lines bearing the Vat gene

Aphis gossypii is the main virus vector in muskmelon crops. The melon gene Vat confers resistance to non‐persistent virus transmission by this aphid. The mechanism of this resistance is not well understood, but no relationship has been detected between resistance and the probing behaviour of aphids on resistant plants. Results presented here suggest that temporary blockage of aphid stylet tips preventing virus particle release may explain the resistance conferred by Vat gene. We performed experiments in which viruliferous aphids were allowed to probe different sequences of resistant (Vat‐bearing) and/or susceptible melon plants. The results demonstrated that A. gossypii inoculates Cucumber mosaic virus (CMV) efficiently in susceptible plants having previously probed resistant plants, showing that the resistance mechanism is reversible. Furthermore, the infection rate obtained for susceptible plants was the same (25%) regardless of whether the transmitting aphid had come directly from the CMV source or had subsequently probed on resistant plants. This result suggests that virus is not lost from stylet to plant during probing of resistant plants, supporting the temporary blockage hypothesis. We also found that the ability of Myzus persicae to transmit CMV is noticeably reduced after probing on resistant plants, providing evidence that this aphid species also responds to the presence of the Vat gene. Finally, we also found that in probes immediately after virus acquisition M. persicae inoculates resistant plants with CMV more efficiently than susceptible plants, perhaps because the Vat gene product induces increased salivation by this aphid.     

Association between Aphis gossypii genotype and phenotype on melon accessions

Development of molecular markers has allowed the characterization of several host–aphid interactions. We investigated the usefulness of microsatellite markers to characterize the plant resistance interaction in the model Aphis gossypii/Cucumis melo. Six aphid clones, collected in different localities and years and belonging to two multilocus genotypes (MLGs) based on eight microsatellite markers, were phenotyped on a set of 33 melon accessions, some of them known to carry the Vat gene. Three parameters were used: acceptance of plant, ability to colonize the plant and resistance to virus when inoculated by aphids. Concordance and correlation analyses showed that aphid clones sharing a same MLG exhibited a very agreeable phenotype on the set of accessions for acceptance of plant and resistance to virus when inoculated by aphids. From host point of view, melon accessions were grouped into four clear categories, resistant to aphids of both MLGs, only resistant to the NM1 MLG, only resistant to the C9 MLG, susceptible to both MLGs and another group of unclear characteristics. The four categories revealed different patterns of virulence for NM1 and C9 MLGs that are likely controlled by a single avirulence gene in accordance with a gene for gene interaction. In contrast, the ability to colonize the plant appeared slightly variable among clones sharing a same MLG. We hypothesize it is due to the putative polygenic control of this aphid trait. Because the phenotypic variability of A. gossypii matched the genetic variability revealed by eight microsatellite markers, these markers could be used to infer the frequency of biotypes in field experiments and help to elucidate the allele diversity of melon resistance genes.     

Antibiotic resistance in potato cultivars to the aphid Myzus persicae

Commercially-grown potato cultivars were assessed for antibiotic resistance to Myzus persicae in a culture room at 19°C with a 2°C range and a 16 h photoperiod. The resistant rankings obtained were consistent for the following aphid metrics: maturation time, teneral adult weight, embryo complement and the production of nymphs. Intrinsic rates of increase (r_m) correlated with the above parameters of aphid growth and reproduction but this was not always the case for nymphal survival rate. The cultivars Ulster Tarn, Record and Maris Piper were relatively resistant to M. persicae and cultivars Desirée and King Edward were the most susceptible. When the aphid clone or the culture plants were changed there was no significant effect on the relative resistance of the potato cultivars under test. The relationship between the number of embryos in teneral adults and the production of nymphs by aphids was examined by grouping aphids in weight batches. There was a strong relationship between these two variables.     

Factors affecting the relative abundance of two coexisting aphid species on sugar beet

1 The two most common species of aphid colonizing sugar beet Beta vulgaris L. are Myzus persicae (Sulzer) (Hemiptera: Aphididae) and Aphis fabae Scopoli (Hemiptera: Aphididae). 2 M. persicae colonizes sugar beet earlier than A. fabae but the population of the former also declines earlier. Despite similar numbers of each species migrating at the time of colonization, M. persicae is usually less abundant on the crop than A. fabae, suggesting differences between the species in their selection of, and performance on, sugar beet. 3 The intrinsic rate of increase of both species declines as sugar beet matures, however, at any given plant age the intrinsic rate of increase of A. fabae is one and a half times greater than that of M. persicae. This results in more rapid population growth and a later decline of the population. 4 Intraspecific competition appears to result in M. persicae becoming very restless, but there is no evidence for interspecific competition between the two species on this host. 5 A population growth model which takes account of the decline in host quality of sugar beet shows that the M. persicae population peaks 30 days before that of A. fabae, and, excluding differences in emigration rate, the maximum A. fabae population is 14 times greater than the maximum M. persicae population. These results are compared to field data.     

Aphid populations showing differential levels of virulence on Capsicum accessions

The green peach aphid,Myzus persicae,is one of the most threatening pests in pepper cultivation and growers would benefit from resistant varietices.Previously,we identified two Capsicum acessions as susceptible and three as resistant to M.persicae using an aphid population originating from the Netherlands(NL).Later on we identified an aphid population originating from a diferent gcographical region(Switserland,SW)that was virulent on all tested Capsicum acessions.The objeetive of the current work is to describe in detail diferent aspects of the interaction between two aphid populations and two sclected Capsicum acessions(one that was susceptible[PB2013046]and one that was resistant[PB2013071]to population NL),including biochemical processes involved.Electrical penetration graph(EPG)recordings showed similar feeding activities for both aphid populations on PB2013046.On acession PB2013071 the aphid population sw was able to devote significantly more time to phloem ingestion than population NL.We also studied plant defense response and found that plants of acession PB2013046 could not induce an accumulation of reactive oxygen species and callose formation after infestation with either aphid population.However,plants of PB2013071 induced a stronger defense response after infestation by population NL than after infestation by population SW.Based on these results,population SW of M.persicae seems to have overcome the resistance of PB2013071 that prevented feeding of aphids from NL population.The potential mechanism by which SW population overcomes the resistance is discussed.     

Assessment of the potential for and development of organophosphorus resistance in field populations of Myzus persicae

A survey of esterases in field populations of the peach-potato aphid, Myzus persicae was made during the spring of 1975. Assay was by electrophoresis of single aphid homogenates, and the known association between the activity of an esterase and resistance to organophosphorus insecticide (OP) was used to infer resistance in field populations. The resistant variant replaced the susceptible in populations which had been treated with OP and another variant with threefold (approximately) more esterase activity appeared to be replacing the resistant variant in populations which have been treated twice with OP. The significance of this for control of M. persicae is discussed. Differences in resistance between aphids in different parts of the same field, and the widespread association of these esterase variants in favoured combinations with two electrophoretic variants at another locus have also been investigated.     

Some factors affecting germination of Peronospora farinosa conidia in water

The germination of conidia of Peronospora farinosa f. sp. betae, collected from sugar beet and suspended in deionized water, was inhibited by dilution with 10% solutions of glycerol, glucose or sucrose and with sap from sugar-beet leaves. Germination was stimulated by diluting with deionized water but not with tap water or biological saline. Substances that diffused from excised buds of sugar-beet plants into deionized water also stimulated germination of conidia but diffusates from leaves did not. This may partly explain why buds are more susceptible to downy mildew than leaves in sugar beet. Germination of conidia was apparently stimulated more by diffusates from buds of seedlings than by those from buds of older plants; this may help to explain why sugar-beet seedlings are more susceptible to downy mildew than older plants. Diffusates from plants of four sugar-beet stocks, that differed from each other in susceptibility to downy mildew, had very similar effects on germination of P. farinosa conidia. Stimulation of spore germination on the surfaces of buds and leaves did not seem, therefore, to be an important factor in determining resistance or susceptibility to downy mildew in these stocks.