The role of sinigrin in host plant recognition by aphids during initial plant penetration

Plant penetration behaviour (probing) of the cabbage aphid, Brevicoryne brassicae, and the pea aphid, Acyrthosiphon pisum, was studied on excised leaves of broad beans, Vicia faba, kept in water or in a 1% aqueous solution of sinigrin. Using the DC EPG (Electrical Penetration Graph) technique it was shown that the cabbage aphid on sinigrin-untreated bean leaves showed numerous short probes into epidermis and mesophyll. None of these aphids showed either phloem salivation or ingestion waveforms on untreated leaves. In contrast, on sinigrin-treated bean leaves, 35% of the probing time was spent on phloem sap ingestion (E2) and almost all aphids reached phloem vessels and started feeding. The duration of phloem salivation before phloem ingestion and the mean duration of phloem ingestion periods were similar on a host and a sinigrin-treated non-host plant. However, the total probing time by B. brassicae was 10% longer, the total phloem sap ingestion time was twice as long, and the time to the first phloem phase within a probe was three times shorter on the host plant compared to sinigrin-treated broad beans. Acyrthosiphon pisum also responded to the addition of sinigrin to broad beans, but in this case sinigrin acted as a deterrent. On sinigrin-treated leaves, A. pisum terminated probes before ingestion from phloem vessels, and none of these aphids showed phloem salivation and ingestion on treated leaves. Glucosinolates were detected in the mesophyll cells of the brassicaceous plant, Sinapis alba. Based on this finding and in addition to the foregoing EPG analysis of aphid probing on these plants and broad beans, our hypothesis is that aphids may recognise their host plants as soon as they probe the mesophyll tissue and before they start ingestion from phloem vessels.     

Characterisation of bird cherry-oat aphid (Rhopalosiphum padi L.) behaviour and aphid host preference in relation to partially resistant and susceptible wheat landraces

The bird cherry-oat aphid (Rhopalosiphum padi L.) is a major pest of wheat (Triticum aestivum L.) and can cause up to 30% yield losses. Heritable plant resistance to aphids is both an economically and ecologically sound method for managing aphids. Here we report how the behaviour and performance of R. padi differs on two resistant, one susceptible wheat landrace and a susceptible elite wheat variety. Feeding behaviour differed among the genotypes, with aphids on resistant lines spending longer in the pathway phase and less time phloem feeding. These behaviours suggest that both inter- and intracellular factors encountered during pathway and phloem feeding phases could be linked to the observed aphid resistance. Locomotion and antennal positioning choice tests also revealed a clear preference for susceptible lines. Although feeding studies revealed differences in the first probe indicating that the resistance factors might also be located in the peripheral layers of the plant tissue, scanning electron microscopy revealed no difference in trichrome length and density on the surface of leaves. Aphids are phloem feeders and limiting the nutrient uptake by the aphids may negatively affect their growth and development as shown here in lower weight and survival of nymphs on resistant genotypes and decreased reproductive potential, with lowest mean numbers of nymphs produced by aphids on W064 (54.8) compared to Solstice (71.9). The results indicate that resistant lines markedly alter the behaviour, reproduction and development potential of R. padi and possess both antixenosis and antibiosis type of resistance.     

Effect of methoxyphenols on grain aphid feeding behaviour

Methoxyphenols might be important in the resistance of cereals to aphids. Electrical penetration graph (EPG) recordings were used to determine the effect of caffeic, ferulic and sinapic acids, and scopoletin on the feeding behaviour of the grain aphidSitobion avenae (F.). Aphids on wheat seedlings treated systemically with these phenols showed reduced ingestion of phloem sap and salivation into sieve elements in most cases. The earlier pathway phases of probing were prolonged. Moreover increase in number of probes as well as reduction of total time of probing was observed. In addition, no O-demethylase activity was found in homogenates of aphids fed on moderately-resistant (phenolic rich) or susceptible (phenolic poor) wheat varieties. The significance of these results for understanding the resistance of cereals to aphids is discussed.     

Behavior and Population Development of <Emphasis Type="Italic">Phorodon humuli</Emphasis> (Schrank) (Homoptera: Aphididae) on Two Hop Cultivars of Different Susceptibility

The two hop cultivars “Hallertauer Magnum” (HM) and “Spalter Select” (SE) are regarded by growers as extremely different in their susceptibility to the damson-hop aphid Phorodon humuli (Schrank). To investigate these anecdotal observations, spring migration and initial population development of P. humuli were monitored on the two cultivars in 1998 and 1999 in an experimental hop garden. Numbers of migrant aphids on SE were significantly lower, comprising 18.8 and 30.2% as compared to HM in 1998 and 1999, respectively. Population development of apterous aphids on these two cultivars differed significantly. At the end of the monitoring period numbers of aphids on SE were 7.5 and 14.2% as compared to HM in 1998 and 1999, respectively. In behavioral studies of P. humuli alates released on glasshouse plants, those on SE spent significantly more time in motile behavior patterns than aphids on HM. In the glasshouse, population development also differed significantly and the number of aphids developing on SE was 12.9% of that on HM after 28 days. It is concluded that SE exhibits a certain repellent effect on P. humuli and, compared to HM, is possibly nutritionally less suitable to the aphid.     

Isotopic enrichment in a phloem-feeding insect: influences of nutrient and water availability

The isotopic enrichment between an animal and its diet can vary among and within living systems, but the sources of variation are not yet fully understood. Some studies have found that diet quality or an animal’s nutritional status can influence the degree of trophic enrichment, while others have dismissed nutrition as a contributing factor. We evaluated the effects of nutrient and water availability on carbon and nitrogen isotopic enrichment in a specialized plant–herbivore system. Aphids are largely sedentary and rely exclusively on nitrogen-poor phloem sap of their host for nutrition. We grew potato aphids [Macrosiphum euphorbiae (Aphididae)] on an accepted host, pumpkin [Cucurbita pepo L. (Cucurbitaceae)], in a glasshouse environment. Twelve pumpkin plants growing under high- and low-watering regimes were inoculated at 4 weeks of age with aphids. During the course of the experiment we collected leaves, phloem sap, aphids and honeydew (i.e., aphid exudates). We found no trophic enrichment between aphids and their phloem sap diet, but significant carbon enrichment of honeydew relative to aphids (2.5‰) and phloem sap (2.1‰). Honeydew was also enriched in nitrogen compared to the phloem sap (1.2‰). Watering treatment had a substantial impact on trophic enrichment. Correlations among tissues, an indication of uniform trophic enrichment among samples, were significant only for the carbon isotopic composition, and then only for plants and aphids grown in the low-water treatment. Diet quality also influenced the degree of isotopic enrichment; trophic enrichment for both carbon and nitrogen isotopic composition increased as diet quality (C/N) declined. We conclude that the degree of trophic enrichment is variable due, in part, to diet quality, but that the scale of variation is small.     

Probing behaviour of Diuraphis noxia on five cereal species with different hydroxamic acid levels

The feeding behaviour of Diuraphis noxia (Kurdjumov) (Homoptera: Aphididae) was electronically monitored on five cereal species (Triticum aestivum L., T. turgidum L., Secale cereale L. and x Triticosecale Wittmack) containing different hydroxamic acids levels and on Hordeum vulgare L., lacking these compounds, by means of a DC-System. With increasing DIMBOA (2,4-dihydroxy-7-methoxy-1,4-benzoxazin-3-one) levels D. noxia showed less probing particularly before reaching the phloem phase and a lower percentage of aphids achieved sustained phloem ingestion. However, no significant correlation was found between DIMBOA levels and the total time of phloem feeding. These findings indicate that factors from the mesophyll and vascular tissues appear to be involved in the effects of resistance. The underlying mechanism, however, remains unclear. Thus, DIMBOA seems to be only part of the chemical defenses of the cereal plants that might be implicated in D. noxia resistance.     

Xylem ingestion by winged aphids

When aphids and their host plant are incorporated in a DC electrical circuit, phloem and xylem ingestion register as separate waveforms of the electrical penetration graph (EPG) signal. Aphids are primarily phloem feeders; xylem ingestion is seldom reported but can be induced experimentally by fasting the insects in desiccating conditions. In experiments with the black bean aphid, Aphis fabae Scop., young winged (alate) and unwinged (apterous) virginoparous adults were collected from their natal host plants (broad bean, Vicia faba L.) and allowed 3-h continuous EPG-recorded access to V. faba seedlings. Several aphids (47% of both morphs) showed ingestion from phloem sieve elements. Alate aphids also showed frequent xylem ingestion (60% of individuals), but no apterous aphids exhibited this activity. The EPG technique involves attachment of a fine gold wire electrode to each insect, a process that may affect normal behaviour at the plant surface. However, when the technique was modified to monitor the stylet activities of freely-settled aphids, high levels of xylem ingestion by alates were also recorded. The results suggest that the developmental physiology of winged aphids somehow predisposes them to xylem ingestion, possibly as a result of dehydration during the teneral period. Alate aphids may reduce their weight by fasting before take-off, giving aerodynamic benefits, but making rehydration, via xylem uptake, a priority following plant contact.     

Analysis of wheat resistance to the cereal aphidSitobion avenae using electrical penetration graphs and flow charts combined with correspondence analysis

The behaviour ofSitobion avenae (F.), was compared on resistant wheat lines ofTriticum monococcum (L.) and a susceptible variety ofTriticum aestivum (L.). Firstly, stylet penetration activities were monitored with the Electrical Penetration Graph (EPG) technique and subsequently analysed using flow charts combined with correspondence analysis. Plant resistance was shown to be associated with repeated penetrations without access to either the xylem or the phloem, and with numerous failures in starting a sustained sap ingestion (as represented by pattern E2). Access to sieve elements of the phloem did not seem to be much affected on resistant plants but it took the aphid three times as long to produce a sap ingestion pattern when maintained on the resistant lineT. monococcum n^o 44 (Tm44) as compared with aphids maintained on susceptible plants. As a result the total time spent in ingesting from sieve elements was reduced by 72% on Tm44. Secondly, direct observations of freely-moving apterous adults were performed. Aphids did not discriminate between resistant and susceptible wheat during the first 30 min of access to test leaves, but only 4 out of 25 aphids were still probing after eight hours on resistant Tm44. The relevance of these results to possible location of the resistance factor(s) are discussed. Although detection of plant resistance before sieve elements are reached can not be rigorously excluded, the factors involved inT. monococcum resistance toS. avenae undoubtedly occur within the phloem vessels.     

The resistance of lettuce (Lactuca sativa L.) toNasonovia ribisnigri: Bionomics ofN. ribisnigri on near isogenic lettuce lines

The intrinsic rate of increase (r_m-values), mean relative growth rates and mortality ofNasonovia ribisnigri (Mosley) (Homoptera, Aphididae) on different lines of lettuce (Lactuca sativa L.) were determined. Near isogenic resistant and susceptible lines, plus their ancestors, were used. Bionomics ofN. ribisnigri on the resistant lines (NrNr) with the dominantNasonovia resistance gene (Nr-gene) differed clearly from the susceptible lines (nrnr). Mortality was high, no larvae reached adulthood, and no reproduction nor honeydew production was seen on the resistant lines. Transfer of aphids to susceptible plants after a period of 2 days on the resistant lines showed no signs of intoxication of aphids. Apparently there is no feeding on the resistant lines. It is not clear whether the aphids can not reach the phloem or do not accept it on the resistant line.     

Electrical Penetration Graph analysis reveals population differentiation of host-plant probing behaviors within the aphid species Uroleucon ambrosiae

Uroleucon ambrosiae collected from the eastern and the southwestern United States were studied in relation to host-plant probing. In the field, eastern aphids are highly specific on Ambrosia trifida, while southwestern aphids feed on Ambrosia trifida plus many other species in the family Asteraceae. Electrical penetration graphs of insects on the principal host, A. trifida, and an additional host of southwestern populations, Heterotheca subaxillaris, revealed regional differentiation in host-associated aphid behaviors, specifically in the first phase of subcuticular probing, and in phloem finding activities. Eastern aphids used A. trifidamore efficiently than southwestern aphids, but were demonstrably less vigorous in their probing activities on H. subaxillaris.     

Locating a resistance mechanism to the cabbage aphid in two wild Brassicas

Feeding behaviour of the cabbage aphid,Brevicoryne brassicae, was monitored electronically on two resistantBrassica species,B. fruticulosa andB. spinescens, and compared with a susceptible controlB. oleracea var.capitata cv. Offenham Compacta. Aphids, monitored for 10 h on the under side of leaves, performed recognizable feeding behaviour on all species. Electrical Penetration Graphs (EPGs) of aphids on resistant and susceptible plants showed no difference in behaviour for aphids on resistantBrassica species compared to susceptible until stylets penetrated the phloem sieve elements when a large reduction in the duration of passive phloem uptake (E₂ pattern) onB. fruticulosa was indicated. Although feeding behaviour on 6 week-old plants ofB. spinescens was similar to the susceptible controls, behaviour on 10 week-old plants was similar to that recorded forB. fruticulosa. The mechanism of resistance is thought to be located in the sieve element as the normal sieve element salivation (E₁) signal was either quickly terminated by withdrawal of the stylets from the sieve element or continued as a disrupted E₂ pattern. Analysis of secondary plant compounds in the threeBrassica species only identified significant differences in the glucosinolate profile. No reproducible differences were detected in the concentration of phenolics or anthocyanins. The major glucosinolate component ofB. fruticulosa andB. spinescens was gluconapin rather than glucobrassicin and glucoiberin as found in the susceptible host plant. However, both pure glucosinolates and glucosinolate extracts from all three species did not reduce aphid survival on chemically-defined artificial diets. These results suggest that the mechanism of resistance may be a mechanical blocking of the sieve element or stylets rather than a difference in the secondary plant chemistry of glucosinolates and phenolics.     

Effects of elevated atmospheric CO2 concentrations on phloem sap composition of spring crops and aphid performance

Abstract

The concentration and composition of free amino acids and carbohydrates in the phloem sap of wheat and oilseed rape (OSR) and the effects on the performance of aphids (Rhopalosiphum padi and Myzus persicae) were determined under atmospheric carbon dioxide (CO₂) enrichment. The analysis of phloem sap showed that carbohydrates and amino acid levels of the host plants were significantly affected by elevated CO₂ level. Among carbohydrate concentrations in the phloem sap, significant increases were observed in fructose and glucose in spring wheat under CO₂ enrichment, whereas no changes were observed in OSR. These changes in plant chemistry affected the performance of herbivorous insects (i.e. aphids) in varying ways, positively affecting the relative growth rate (RGR) of R. padi in spring wheat and negatively affecting the RGR of M. persicae on OSR.     

Sieve element occlusion provides resistance against Aphis gossypii in TGR-1551 melons

Feeding behavior and plant response to feeding were studied for the aphid Aphis gossypii Glover on susceptible and resistant melons(cv.Iroquois and TGR-1551,respectively).Average phloem phase bout duration on TGR-1551 was<7% of the duration on Iroquois.Sixty-seven percent of aphids on TGR-1551 never produced a phloem phase that attained ingestion(EPG waveform E2)in contrast to only 7% of aphids on Iroquois.Average bout duration of waveform E2(scored as zero if phloem phase did not attain E2)on TGR-1551 was<3% of the duration on Iroquois.Conversely,average bout duration of EPG waveform El(sieve element salivation)was 2.8 times greater on TGR-1551 than on Iroquois.In a second experiment,liquid nitrogen was used to rapidly cryofix leaves and aphids within a few minutes after the aphids penetrated a sieve element.Phloem near the penetration site was then examined by confocal laser scanning microscopy.Ninety-six percent of penetrated sieve elements were occluded by protein in TGR-1551 in contrast to only 28% in Iroquois.Usually in TGR-1551,occlusion was also observed in nearby nonpenetrated sieve elements.Next,a calcium channel blocker,trivalent lanthanum,was used to prevent phloem occlusion in TGR-1551,and A.gossypii feeding behavior and the plants phloem response were compared between lanthanum-treated and control TGR-1551.Lanthanum treatment eliminated the sieve element protein occlusion response and the aphids readily ingested phloem sap from treated plants.This study provides strong evidence that phloem occlusion is a mechanism for resistance against A.gossypii in TGR-1551.     

Susceptibility of hop to Phorodon humuli

A portion of the USDA World Hop (Humulus lupulus L.) Germ Plasm Collection was screened for antibiosis to the hop aphid, Phorodon humuli (Shrank). Hop aphid development and reproduction on the retained accessions 58016, 21090M, 21211, 21240, and 60038 were then compared to hop aphids reared on the commercial hop cultivars (cvs) L-8, Perle, Chinook, Galena, Eroica, Willamette, and Cascade. In general, hop aphids matured faster and reproduced more on the commercial cvs than on 21090M, 58016, 21211, or 21240. Accession 60038, of note because it supported unusually high numbers of aphids during screening, was found to be the most susceptible of any hop tested. Mortality and natality schedules were then combined to construct matrix models of aphid population growth on 60038, 58016, Cascade, and Perle (the most and least susceptible noncommercial and commercial hops, respectively). Beginning with one one-day-old nymph per leaf, the models predicted that P. humuli would require 24 days to surpass an action threshold of 100 aphids per leaf on 58016. This was seven days longer than on the susceptible Cascade and five days longer than on Perle. A commercial hop with the antibiosis of 58016 would probably require fewer insecticide applications during the growing season, thus retarding the development of insecticide resistance in P. humuli and enhancing the effectiveness of integrated management programs by protecting beneficial insects.

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Zusammenfassung Ein Teil der Genbank des USDA von Hopfen (Humulus lupulus L.) wurde auf Resistenz gegen Phorodon humuli (Shrank) getestet. Dabei wurden die besonders widerstands fähigen Klone 58016, 21090M, 21211, 21240 sowie Klon 60038, der ungewöhnlich viele Blattläuse ertrug, ausgewählt. Anschließend wurde die Entwicklung und Vermehrung der Blattläuse auf den ausgewählten Klonen und auf den Handelssorten (cvs) L-8, Perle, Chinook, Galena, Eroica, Willamette, Cascade untersucht. Im Allgemeinen entwickelten sich die Blattläuse auf den Handelssorten reascher und vermehrten sich stärker als auf den resistenten Klonen. Klon 60038 war dagegen die anfälligste Sorte. Mortalitäts- und Natlitätsdaten wurden kombiniert um Matrixmodelle des Populationswachstums der Blattläuse auf 60038, 58016, Cascade und Perle (die am meisten und am wenigsten anfälligen kommerziellen und nichtkommerziellen Sorten) zu konstruieren. Die Modelle begannen bei 1 Larve pro Blatt und sagten voraus, dass die Aktionsgrenze von 100 Tieren pro Blatt auf 58016 nach 24 Tagen erreicht wird. Das ist 7 Tage später als auf den anfälligen Sorten Cascade und 5 Tage später als auf Perle. Eine kommerzielle Sorte mit der Anfälligkeit von 58016 würde vermutlich während der Vegetationszeit weniger Insektizidspritzungen benötigen. Damit würde die Resistenzentwicklung bei Phorodon humuli verlangsamt und die Wirkung von IPM Programmen durch Schutz der natürlichen Feinde verbessert.

     

Enhancement of resistance to aphids by introducing the snowdrop lectin genegna into maize plants

In order to enhance the resistance to pests, transgenic maize (Zea mays L.) plants from elite inbred lines containing the gene encoding snowdrop lectin (Galanthus nivalis L. agglutinin; GNA) under control of a phloem-specific promoter were generated through theAgrobacterium tumefaciens- mediated method. The toxicity of GNA-expressing plants to aphids has also been studied. The independently derived plants were subjected to molecular analyses. Polymerase chain reaction (PCR) and Southern blot analyses confirmed that thegna gene was integrated into maize genome and inherited to the following generations. The typical Mendelian patterns of inheritance occurred in most cases. The level of GNA expression at 0.13%-0.28% of total soluble protein was observed in different transgenic plants. The progeny of nine GNA-expressing independent transformants that were derived separately from the elite inbred lines DH4866, DH9942, and 8902, were selected for examination of resistance to aphids. These plants synthesized GNA at levels above 0.22% total soluble protein, and enhanced resistance to aphids was demonstrated by exposing the plants to corn leaf aphid (Rhopalosiphum maidis Fitch) under greenhouse conditions. The nymph production was significantly reduced by 46.9% on GNA-expressing plants. Field evaluation of the transgenic plants supported the results from the inoculation trial. After a series of artificial self-crosses, some homozygous transgenic maize lines expressing GNA were obtained. In the present study, we have obtained new insect-resistant maize material for further breeding work.     

Recombinant and classically selected factors of potato plant resistance to the Colorado potato beetle, Leptinotarsa decemlineata, variously affect the potato aphid parasitoid Aphidius nigripes

Different forms of crop resistance developed against majorpotato pests such as the Colorado potato beetle (CPB), Leptinotarsadecemlineata, may be variously compatible with biological controlof secondary pests such as aphids. We compared the performance of theparasitoid Aphidius nigripes developing in the aphid hostMacrosiphum euphorbiae, on five potato lines, including atransgenic `Superior-BT' line expressing the CryIIIA toxin ofBacillus thuringiensis, which is specific to Coleoptera; andthe `NYL 235-4' line derived from Solanum berthaultii,characterized by relatively unspecific resistance to herbivores based onmoderately-high density of glandular trichomes. The other lines testedwere a `Kennebec-OCI' transgenic line expressing the protease inhibitorrice cystatin I (OCI), a potential resistance factor against coleopteranpests; and the commercial cultivars `Superior' and `Kennebec' used ascontrols. Parasitoid immature survival and adult size were reducedcompared to controls when the wasps developed on aphids fed the`Superior-BT' potato. In contrast, parasitoid size and fecundityincreased above control when the wasps developed on aphids fed the`Kennebec-OCI' potato. Parasitoids reached the adult stage faster andwere more fecund on `NYL 235-4' than control lines. The results indicatethat the different forms of potato resistance currently developed mainlyagainst the CPB had various unexpected effects on aphid parasitoidfitness. These effects on the parasitoid were complex but were generallyinterpretable in terms of host aphid quality variation among potatolines used as food by the aphids during parasitoiddevelopment.     

Influence of Experience on the Response of Bathyplectes curculionis (Hymenoptera: Ichneumonidae), a Nonaphidophagous Parasitoid, to Aphid Odor

Bathyplectes curculionis (Thomson) is an introduced natural enemy of the alfalfa weevil in North America. The wasp requires carbohydrate foods as an adult. Adult wasps have increased longevity and fecundity when provided access to pea aphid, Acyrthosiphon pisum (Harris), honeydew in the laboratory, and adults respond positively to the presence of pea aphids in alfalfa fields. However, it is unknown how these wasps find aphid honeydew in the field. In a series of Y-tube olfactometer experiments, we evaluated the response of naïve and experienced adult female B. curculionis to odors from pea aphids, alfalfa, and pea aphids on alfalfa. Naïve adult females did not respond positively to pea aphid odor even when hungry. But adult females were able to learn aphid odor, and the mechanism of learning appears to be associative rather than by sensitization. Naïve females also showed no preference for alfalfa odor but learned alfalfa odor through sensitization. The wasps did not distinguish between alfalfa with aphids and alfalfa without aphids, even after exposure to aphids or alfalfa with aphids. However, they preferred pea aphid odor to alfalfa odor after a feeding experience in the presence of pea aphid odors. But after exposure to mixed odors of aphids and alfalfa while feeding, B. curculionis females preferred the odor of alfalfa to the odor of pea aphids. These results suggest that alfalfa odors mask or override aphid odors when aphids are associated with alfalfa (as happens naturally), thus interfering with the wasp's ability to respond to learned aphid odors. Therefore, although the wasps are capable of learning to find pea aphids and their honeydew in a simplified laboratory setting, it appears unlikely that they do so in the field.     

Effect of Manipulations on the Host Selection Behavior of Sitobion avenae (Homoptera: Aphididae)

The study of aphid host selection and feeding behavior is difficult because aphids have to penetrate the plant to reach their feeding site, phloem tissue. The activity of the stylets, salivation or food intake, can not be observed externally and requires an indirect visualization technique such as the Electric Penetration Graph (EPG). The plant selection behavior of Sitobion avenae on potato varied depending on whether an ethological or EPG method was used to study it. A similar variation did not occur with Myzus persicae or Rhopalosiphum padi. The application of water-based silver conductive paint onto the thorax, as normally used for EPG, or onto the abdomen of Sitobion avenae alates resulted in increased duration and frequency of probing compared to results from ethological observations. Our results indicated that EPG manipulations might have different effects on different species of aphids and that a comparison of EPG and ethological data is required to confirm that the EPG method does not bias aphid feeding behavior.     

Analysis by DC–EPG of the resistance to Bemisia tabaci on an Mi-tomato line

The tomato Mi gene confers resistance to nematodes, Meloidogyne spp., and to the potato aphid, Macrosiphum euphorbiae (Thomas). Previous greenhouse choice assays with Bemisia tabaci (Gennadius) showed that tomato commercial varieties carrying this gene had significantly lower values of host suitability and whitefly reproduction than varieties lacking Mi. This indicated that Mi, or another gene in its region, could regulate partial resistance. In order to characterise this resistance, probing and feeding behaviour of Bemisia tabaci B-biotype was studied with DC Electrical Penetration Graph (EPG) technique on the near-isogenic tomato lines Moneymaker (without Mi) and Motelle (carrying Mi). Significant differences (P < 0.05) between tomato lines were found in EPG parameters related to epidermis and/or mesophyll tissues. On Motelle, a lower percentage of whiteflies achieved phloem phase and they made more probes before attaining first phloem phase, had a higher ratio (number of probes before first phloem phase)/(total number of probes), had a longer total duration of non-probing time, and a longer time before making the first intracellular puncture and before making the first phloem phase. In contrast, most of the parameters related to phloem phase were found not to differ significantly between these near-isogenic lines. The behavioural data strongly suggest that the partial resistance in the variety Motelle is due to factors in the epidermis and/or mesophyll that inhibit the whiteflies from reaching phloem sieve elements. However, once the stylets reach a sieve element, whitefly behaviour did not differ between the two varieties. Thus, phloem sap of the two varieties appears to be equally acceptable to the whiteflies. Further studies are necessary to provide a better understanding of these mechanisms of resistance to whiteflies in tomatoes.     

Hydroxamic acids affecting barley yellow dwarf virus transmission by the aphid Rhopalosiphum padi

2,4-Dihydroxy-7-methoxy-1,4-benzoxazin-3-one (DIMBOA), a hydroxamic acid (Hx) occurring in wheat, was shown to deter feeding by the aphid Rhopalosiphum padi (L.), and to reduce BYDV transmission to the plant. Dual choice tests with wheat leaves showed the preferential settlement of aphids on leaves with lower levels of DIMBOA. Electric monitoring of aphid feeding behaviour showed that in seedlings with higher DIMBOA levels fewer aphids reached the phloem and they needed longer times to contact a phloem vessel than in those with lower levels. When aphids carrying BYDV were allowed to feed on wheat cultivars with different DIMBOA levels, fewer plants were infected with BYDV in the higher DIMBOA cultivars than in the lower ones. Preliminary field experiments showed a tendency for wheat cultivars with higher Hx levels to be more tolerant to infection by BYDV than lower Hx level ones.