Modification of native grasses for biofuel production may increase virus susceptibility

Bioenergy production is driving modifications to native plant species for use as novel biofuel crops. Key aims are to increase crop growth rates and to enhance conversion efficiency by reducing biomass recalcitrance to digestion. However, selection for these biofuel‐valuable traits has potential to compromise plant defenses and alter interactions with pests and pathogens. Insect‐vectored plant viruses are of particular concern because perennial crops have potential to serve as virus reservoirs that influence regional disease dynamics. In this study, we examined relationships between growth rates and biomass recalcitrance in five switchgrass (Panicum virgatum) populations, ranging from near‐wildtype to highly selected cultivars, in a common garden trial. We measured biomass accumulation rates and assayed foliage for acid detergent lignin, neutral detergent fiber, in vitro neutral detergent fiber digestibility and in vitro true dry matter digestibility. We then evaluated relationships between these traits and susceptibility to a widely distributed group of aphid‐transmitted Poaceae viruses (Luteoviridae: Barley and cereal yellow dwarf viruses, B/CYDVs). Virus infection rates and prevalence were assayed with RT‐PCR in the common garden, in greenhouse inoculation trials, and in previously established switchgrass stands across a 300‐km transect in Michigan, USA. Aphid host preferences were quantified in a series of arena host choice tests with field‐grown foliage. Contrary to expectations, biomass accumulation rates and foliar digestibility were not strongly linked in switchgrass populations we examined, and largely represented two different trait axes. Natural B/CYDV prevalence in established switchgrass stands ranged from 0% to 28%. In experiments, susceptibility varied notably among switchgrass populations and was more strongly predicted by potential biomass accumulation rates than by foliar digestibility; highly selected, productive cultivars were most virus‐susceptible and most preferred by aphids. Evaluation and mitigation of virus susceptibility of new biofuel crops is recommended to avert possible unintended consequences of biofuel production on regional pathogen dynamics.     

Mechanisms underlying the impacts of exotic annual grasses in a coastal California meadow

Biological invasions can impact the abundance and diversity of native species, but the specific mechanisms remain poorly discerned. In California grasslands, invasion by European annual grasses has severely reduced the quality of habitat for native forb species. To understand how introduced grasses suppress native and exotic forbs, we examined the response of a Southern California grassland community to factorial removals of live grass and the litter produced in previous seasons. To examine the role that belowground competition for water plays in mediating the impact of grasses, we crossed grass and litter removal treatments with water addition. Our results show that forbs were almost equally suppressed by both competition from live grass and direct interference by litter. Water addition did not ameliorate the effect of grass competition, suggesting that water was not the resource for which plants compete. This evidence is consistent with the susceptibility of forbs to light limitation, especially considering that litter does not consume water or nutrients. Interestingly, despite different histories of co-occurrence with annual grass dominants, native and exotic forbs were comparably suppressed by exotic grasses. Our results indicate that suppression by both live and dead stems underlie the influence of exotic grasses on forb competitors.     

Barley yellow dwarf virus: effects on carbohydrate metabolism in oat (Avena sativa) during cold hardening

Barley yellow dwarf virus (BYDV) causes significant losses in yield and in overwintering ability of winter cereals. Mechanisms by which the physiology of plants is affected by the virus are not clear. To see how carbohydrates in the crown of winter cereals were affected by BYDV, fructan isomers of degree of polymerization (DP) 3–5, fructan DP>6 and the simple sugars, glucose, fructose and sucrose, were measured before and during cold hardening in three oat ( Avena sativa L.) cultivars, 'Wintok', 'Coast Black' and 'Fulghum'. On a fresh weight basis fructan DP>6 decreased by 50% in infected 'Wintok' and 'Coast Black' and by 25% in 'Fulghum'. Two DP3, one DP4 and one DP5 isomer were significantly higher than non-infected controls. The percentages of simple sugars in infected crowns were significantly higher than controls in all three cultivars in every week except the first week of hardening. Crude enzyme extracts from BYDV infected plants incubated with sucrose suggested higher invertase and lower sucrose-sucrosyl transferase activity. When incubated with 1-kestose and neokestin, no significant difference was found in fructose fructosyl transferase or in hydrolase activity. The activity of unidentified enzymes catalysing the synthesis of larger (DP>5) fructan was altered by BYDV. The decrease of carbohydrates in the crown induced indirectly by BYDV may alter the plant's capacity to regenerate tillers in the spring. The ability of plants to prevent or tolerate carbohydrate fluctuations induced by BYDV infection may be an important genetically regulated characteristic for developing virus-resistant cultivars.     

California native and exotic perennial grasses differ in their response to soil nitrogen,exotic annual grass density,and order of emergence

Early emergence of plant seedlings can offer strong competitive advantages over later-germinating neighbors through the preemption of limiting resources. This phenomenon may have contributed to the persistent dominance of European annual grasses over native perennial grasses in California grasslands, since the former species typically germinate earlier in the growing season than the latter and grow rapidly after establishing. Recently, European perennial grasses have been spreading into both non-native annual and native perennial coastal grass stands in California. These exotic perennials appear to be less affected by the priority effects arising from earlier germination by European annual grasses. In addition, these species interactions in California grasslands may be mediated by increasing anthropogenic or natural soil nitrogen inputs. We conducted a greenhouse experiment to test the effects of order of emergence and annual grass seedling density on native and exotic perennial grass seedling performance across different levels of nitrogen availability. We manipulated the order of emergence and density of an exotic annual grass (Bromus diandrus) grown with either Nassella pulchra (native perennial grass), Festuca rubra (native perennial grass), or Holcus lanatus (exotic perennial grass), with and without added nitrogen. Earlier B. diandrus emergence and higher B. diandrus density resulted in greater reduction in the aboveground productivity of the perennial grasses. However, B. diandrus suppressed both native perennials to a greater extent than it did H. lanatus. Nitrogen addition had no effect on the productivity of native perennials, but greatly increased the growth of the exotic perennial H. lanatus, grown with B. diandrus. These results suggest that the order of emergence of exotic annual versus native perennial grass seedlings could play an important role in the continued dominance of exotic annual grasses in California. The expansion of the exotic perennial grass H. lanatus in coastal California may be linked to its higher tolerance of earlier-emerging annual grasses and its ability to access soil resources amidst high densities of annual grasses.     

The effect of soil nitrogen on competition between native and exotic perennial grasses from northern coastal California

The invasion of European perennial grasses represents a new threat to the native coastal prairie of northern California. Many coastal prairie sites also experience anthropogenic nitrogen (N) deposition or increased N availability as a result of invasion by N-fixing shrubs. We tested the hypothesis that greater seedling competitive ability and greater responsiveness to high N availability of exotic perennial grasses facilitates their invasion in coastal prairie. We evaluated pairwise competitive responses and effects, and the occurrence of asymmetrical competition, among three common native perennial grasses (Agrostis oregonensis, Festuca rubra, and Nassella pulchra) and three exotic perennial grasses (Holcus lanatus, Phalaris aquatica, and Festuca arundinacea), at two levels of soil N. We also compared the root and shoot biomass and response to fertilization of singly-grown plants, so we could evaluate how performance in competition related to innate plant traits. Competitive effects and responses were negatively correlated and in general varied continuously across native and exotic species. Two exceptions were the exotic species Holcus, which had large effects on neighbors and small responses to them, and competed asymmetrically with all other species in the experiment, and the native grass Nassella, which had strong responses to but little effect on neighbors, and was out-competed by all but one other species in the experiment. High allocation to roots and high early relative growth rate appear to explain Holcus’s competitive dominance, but its shoot biomass when grown alone was not significantly greater than those of the species it out-competed. Competitive dynamics were unaffected by fertilization. Therefore, we conclude that seedling competitive ability alone does not explain the increasing dominance of exotic perennial grasses in California coastal prairie. Furthermore, since native and exotic species responded individualistically, grouping species as ‘natives’ and ‘exotics’ obscured underlying variation within the two categories. Finally, elevated soil N does not appear to influence competition among the native and exotic perennial grasses studied, so reducing soil N pools may not be a critical step for the restoration of California coastal prairie.     

Identification of polypeptide markers of barley yellow dwarf virus resistance and susceptibility genes in non-infected barley (Hordeum vulgare) plants

Summary Barley yellow dwarf (BYDV) is a group a closely related viruses which cause economic losses in a wide range of graminaceous species throughout the world. Barley plants can be protected from the effects of BYDV by the Yd2 resistance gene. Plants which contain the Yd2 gene also contain a constitutively expressed polypeptide which was not found in any plants without Yd2. Conversely, BYDV susceptible plants contain another constitutively expressed polypeptide which was not found in any of the BYDV-resistant lines examined. These two polypeptides appear to have the same molecular weight (as assessed by SDS-PAGE) and only slightly different iso-electric points. They also appear to contain an extensive range of similar antigenic determinants. Both polypeptides were found in F₁ hybrids made from resistant and susceptible plants. We suggest that these two polypeptides are the products of two allelic genes. Analysis of near-isogenic lines showed that the locus which controls the Yd2 resistance gene and the locus controlling the synthesis of the two polypeptides may be within ± 9 cM of each other. We have developed a Western blot technique which allows assessment of barley lines, 4-days after seed imbibition, for the presence of the Yd2 gene.     

Capture and allocation of nitrogen byQuercus douglasii seedlings in competition with annual and perennial grasses

Summary The spatial overlap of woody plant root systems and that of annual or perennial grasses promotes competition for soil-derived resources. In this study we examined competition for soil nitrogen between blue oak seedlings and either the annual grassBromus mollis or the perennial grassStipa pulchra under controlled outdoor conditions. Short-term nitrogen competition was quantified by injecting¹⁵N at 30 cm depth in a plane horizontal to oak seedling roots and that of their neighbors, and calculating¹⁵N uptake rates, pool sizes and¹⁵N allocation patterns 24 h after labelling. Simultaneously, integrative nitrogen competition was quantified by examining total nitrogen capture, total nitrogen pools and total nitrogen allocation.Stipa neighbors reduced inorganic soil nitrogen content to a greater extent than didBromus plants. Blue oak seedlings responded to lower soil nitrogen content by allocating lower amounts of nitrogen per unit of biomass producing higher root length densities and reducing the nitrogen content of root tissue. In addition, blue oak seedlings growing with the perennial grass exhibited greater rates of¹⁵N uptake, on a root mass basis, compensating for higher soil nitrogen competition inStipa neighborhoods. Our findings suggest that while oak seedlings have lower rates of nitrogen capture than herbaceous neighbors, oak seedlings exhibit significant changes in nitrogen allocation and nitrogen uptake rates which may offset the competitive effect annual or perennial grasses have on soil nitrogen content.     

Abiotic constraints on the competitive ability of exotic and native grasses in a Pacific Northwest prairie

In prairie ecosystems, abiotic constraints on competition can structure plant communities; however, the extent to which competition between native and exotic plant species is constrained by environmental factors is still debated. The objective of our study was to use paired field and greenhouse experiments to evaluate the competitive dynamics between two native (Danthonia californica and Deschampsia cespitosa) and two exotic (Schedonorus arundinaceus and Lolium multiflorum) grass species under varying nutrient and moisture conditions in an upland prairie in the Willamette Valley, Oregon. We hypothesized the two invasive, exotic grasses would be more competitive under high-nutrient, moderate-moisture conditions, resulting in the displacement of native grasses from these environments. In the field, the experimental reduction of competition resulted in shorter, wider plants, but only the annual grass, Lolium multiflorum, produced more aboveground biomass when competition was reduced. In the greenhouse, the two exotic grasses produced more total biomass than the two native grasses. Competitive hierarchies were influenced by nutrient and/or moisture treatments for the two exotic grasses, but not for the two native grasses. L. multiflorum dominated competitive interactions with all other grasses across treatments. In general, S. arundinaceus dominated when in competition with native grasses, and D. cespitosa produced the most biomass in monoculture or under interspecific competition with the other native grass, D. californica. D. californica, D. cespitosa, and S. arundinaceus all produced more biomass in high-moisture, high-nutrient environments, and D. cespitosa, L. multiflorum, and S. arundinaceus allocated more biomass belowground in the low nutrient treatment. Taken together, these experiments suggest the competitive superiority of the exotic grasses, especially L. multiflorum, but, contrary to our hypothesis, the native grasses were not preferentially excluded from nutrient-rich, moderately wet environments. Laurel Pfeifer-Meister and Esther M. Cole contributed equally to this work.     

The effect of temperature and inoculation access period on the transmission of barley yellow dwarf virus byRhopalosiphum padi (L.) and Sitobion avenae (F.)

Winged individuals of Rhopalosiphum padi and Sitobion avenae transmitted the PAV-like and MAV-like isolates of barley yellow dwarf virus respectively. Success of transmission after inoculation access periods of 2, 6, 12, 24, 48 and 72 h were examined and survival, reproduction and movement of the aphid vectors were recorded at these times. The experiment was done at four different temperatures: 6^oC, 12^oC, 18^oC and 23^oC. For both isolates the inoculation efficacy did not increase after a 24 h inoculation access period and there was no difference in inoculation efficiency at the three highest temperatures, that at 6^oC being significantly lower than at 12^oC to 23^oC. The results suggest that autumn temperature is a critical factor for BYDV epidemiology in Britain with a small increase in autumn temperature leading to greatly increased infection rates.     

Chickpea chlorotic dwarf virus,a new leafhopper-transmitted geminivirus of chickpea in India1

A disease of chickpea in India, characterised by chlorosis, severe stunting and phloem browning, was shown to be caused by a geminivirus. This virus was transmitted by the leafhopper Orosius orientalis from chickpea to chickpea and several other plant species. A method for purification of this virus was devised and a polyclonal antiserum produced. The majority of the purified particles were geminate. The size of the coat protein was shown to be 32 kD and the nucleic acid was shown to be circular ssDNA of 2900 nucleotides. By immunosorbent electron microscopy this virus was shown to be unrelated to the leafhopper-transmitted geminiviruses known to infect dicotyledons such as beet curly top, bean summer death and tobacco yellow dwarf viruses. On the basis of particle morphology, leafhopper transmission, host range and serology this virus was considered to be a new, hitherto undescribed, geminivirus and was named chickpea chlorotic dwarf virus.     

Variation in the establishment of a non-native annual grass influences competitive interactions with Mojave Desert perennials

Competition between native and non-native species can change the composition and structure of plant communities, but in deserts, the highly variable timing of resource availability also influences non-native plant establishment, thus modulating their impacts on native species. In a field experiment, we varied densities of the non-native annual grass Bromus madritensis ssp. rubens around individuals of three native Mojave Desert perennials—Larrea tridentata, Achnatherum hymenoides, and Pleuraphis rigida—in either winter or spring. For comparison, additional plots were prepared for the same perennial species and seasons, but with a mixture of native annual species as neighbors. Growth of perennials declined when Bromus was established in winter because Bromus stands had 2–3 months of growth and high water use before perennial growth began. However, water potentials for the perennials were not significantly reduced, suggesting that direct competition for water may not be the major mechanism driving reduced perennial growth. The impact of Bromus on Larrea was lower than for the two perennial grasses, likely because Larrea maintains low growth rates throughout the year, even after Bromus has completed its life cycle. This result contrasts with the perennial grasses, whose phenology completely overlaps with (Achnatherum) or closely follows (Pleuraphis) that of Bromus. In comparison, Bromus plants established in spring were smaller than those established in winter and thus did not effectively reduce growth of the perennials. Growth of perennials with mixed annuals as neighbors also did not differ from those with Bromus neighbors of equivalent biomass, but stands of these native annuals did not achieve the high biomass of Bromus stands that were necessary to reduce perennial growth. Seed dormancy and narrow requirements for seedling survivorship of native annuals produce densities and biomass lower than those achieved by Bromus; thus, impacts of native Mojave Desert annuals on perennials are expected to be lower than those of Bromus.     

Vectoring ability of aphid clones of Rhopalosiphum padi (L.) and Sitobion avenae (Fabr.) and their capacity to retain barley yellow dwarf virus

Vectoring ability of four aphid clones, Rp-M and Rp-R26 of Rhopalosiphum padi and Sa-R1 and Sa-V of Sitobion avenae, to transmit barley yellow dwarf (PAV, MAV and RPV) luteoviruses (BYDV) was compared in controlled conditions. Significant differences between highly efficient vectors (HEV), Rp-M and Sa-Rl, and poorly efficient vectors (PEV), Rp-R26 and Sa-V, were found in transmission of their specific viruses with acquisition and inoculation access periods (AAP, IAP) of 5 days. BYD-RPV was occasionally transmitted by both clones of S. avenae. None of 150 tested apterous adults of the Rp-R26 transmitted BYD-MAV, while 10% of transmission was observed from those of the Rp-M in a parallel test. An improved ELISA and immuno-PCR were adapted to test for viruses in aphids. The results obtained by the improved ELISA indicated there was a good correlation between virus detection in single aphids of HEV clones after a 5 day AAP and virus transmission by them. In contrast, the percentages of virus-carrying aphids of PEV clones were generally higher than those of their transmission rates. BYD-MAV and BYD-RPV were also detected by the improved ELISA in single aphids of their PEV clones, with the exception of BYD-RPV in those of Sa-V. However, after a 2-day IAP, the improved ELISA in most cases failed to detect these viruses in single aphids of PEV clones. Detection by immuno-PCR demonstrated that all three viruses could be acquired and retained by the aphids of both HEV and PEV clones. But, as visualised from electrophoretic bands, after the 2-day IAP the amplified products from aphid extracts of PEV clones were reduced. The detection in a batch of nine aphids by the improved ELISA revealed that virus content in PEV clones decreased more rapidly than that in HEV clones during transmission. Thus, the difference in transmission efficiency of the aphid clones within species was not caused by an inability to acquire virus, but was determined by variation in vectoring ability between them. This was due to differences in ability to prevent the passage of virions from haemocoel to salivary duct and/or different capacities for the retention of BYDV.     

QTL analysis of tolerance to a German strain of BYDV-PAV in barley (Hordeum vulgare L.)

One hundred and forty six barley doubled-haploid lines (DH lines) were tested for variation in grain yield, yield components, plant height, and heading date after artificial infection with a German isolate of barley yellow dwarf virus (BYDV-PAV-Braunschweig). Of these 146 lines 76 were derived from the cross of the barley yellow dwarf virus (BYDV) tolerant cultivar ’Post’ to cv ’Vixen’ (Ryd2) and 70 from the cross of Post to cv ’Nixe’. Phenotypic measurements were gathered on both non-infected plants and plants artificially inoculated with BYDV-PAV by viruliferous aphids in pot and field experiments for three years at two locations. For all traits a continuous variation was observed suggesting a quantitative mode of inheritance for tolerance against BYDV-PAV. Using skeleton maps constructed using SSRs, AFLPs and RAPDs, two QTLs for relative grain yield per plant after BYDV infection, explaining about 47% of the phenotypic variance, were identified in Post × Vixen at the telomeric region of chromosome 2HL and at a region containing the Ryd2 gene on chromosome 3HL. In Post × Nixe, a QTL was found in exactly the same chromosome 2HL marker interval. In this cross, additional QTL were mapped on chromosomes 7H and 4H and together these explained about 40% of the phenotypic variance. QTL for effects of BYDV infection on yield components, plant height, and heading date generally mapped to the same marker intervals, or in the vicinity of the QTL for relative grain yield, on chromosomes 2HL and 3HL, suggesting that these regions are of special importance for tolerance to the Braunschweig isolate of BYDV-PAV. Possible applications of marker-assisted selection for BYDV tolerance based on these results are discussed. Received: 1 December 2000 / Accepted: 9 March 2001     

Response of net ecosystem gas exchange to a simulated precipitation pulse in a semi-arid grassland: the role of native versus non-native grasses and soil texture

Physiological activity and structural dynamics in arid and semi-arid ecosystems are driven by discrete inputs or pulses of growing season precipitation. Here we describe the short-term dynamics of ecosystem physiology in experimental stands of native (Heteropogon contortus) and invasive (Eragrostis lehmanniana) grasses to an irrigation pulse across two geomorphic surfaces with distinctly different soils: a Pleistocene-aged surface with high clay content in a strongly horizonated soil, and a Holocene-aged surface with low clay content in homogenously structured soils. We evaluated whole-ecosystem and leaf-level CO₂ and H₂O exchange, soil CO₂ efflux, along with plant and soil water status to understand potential constraints on whole-ecosystem carbon exchange during the initiation of the summer monsoon season. Prior to the irrigation pulse, both invasive and native grasses had less negative pre-dawn water potentials ( _pd), greater leaf photosynthetic rates (A _net) and stomatal conductance (g _s), and greater rates of net ecosystem carbon exchange (NEE) on the Pleistocene surface than on the Holocene. Twenty-four hours following the experimental application of a 39 mm irrigation pulse, soil CO₂ efflux increased leading to all plots losing CO₂ to the atmosphere over the course of a day. Invasive species stands had greater evapotranspiration rates (ET) immediately following the precipitation pulse than did native stands, while maximum instantaneous NEE increased for both species and surfaces at roughly the same rate. The differential ET patterns through time were correlated with an earlier decline in NEE in the invasive species as compared to the native species plots. Plots with invasive species accumulated between 5% and 33% of the carbon that plots with the native species accumulated over the 15-day pulse period. Taken together, these results indicate that system CO₂ efflux (both the physical displacement of soil CO₂ by water along with plant and microbial respiration) strongly controls whole-ecosystem carbon exchange during precipitation pulses. Since CO₂ and H₂O loss to the atmosphere was partially driven by species effects on soil microclimate, understanding the mechanistic relationships between the soil characteristics, plant ecophysiological responses, and canopy structural dynamics will be important for understanding the effects of shifting precipitation and vegetation patterns in semi-arid environments.     

Juvenile Chinese mitten crabs (<Emphasis Type="Italic">Eriocheir sinensis</Emphasis>) in the Thames estuary: distribution,movement and possible interactions with the native crab <Emphasis Type="Italic">Carcinus maenas</Emphasis>

The Chinese mitten crab, Eriocheir sinensis is a successful invader. Whilst non-breeding adult E. sinensis have been associated with the destruction of riverbanks, little is known about the ecology of this species in its invaded areas. This is especially true of the juveniles which are a key migratory stage. Intertidal surveys along the Thames estuary indicated an increase in the population since the 1990s. Juvenile E. sinensis were abundant in the sampled upper tidal region of the Thames, refuging under boulders in the intertidal at low tide. Seasonal differences in sampled populations were observed, with a significantly lower abundance of crabs found during winter compared to summer. Mark-recaptures indicated movement in the intertidal occurring during high tide, with an influx of new crabs evident after a single tidal cycle. Endogenous rhythms were also apparent, with peaks in activity occurring corresponding with night-time high tide conditions. E. sinensis juveniles can successfully exclude similar sized native Carcinus maenas from shelters in the laboratory, regardless of which species originally inhabited the shelter. This may have implications for native estuarine Carcinus populations, which can rely on such intertidal shelters for refuges.     

Feeding behavior and transmission of barley yellow dwarf virus by Sitobion avenae on oats

Feeding behavior of Sitobion avenae F. (Homoptera: Aphididae) on oats (Avena sativa cv. Clintland 64) was electronically monitored, and waveforms corresponding to salivation, ingestion, and sieve element penetration described.During 90 min plant access, aphids ingested from phloem for 0–43 min (mean: 8.1 min) and non-phloem for 0–60 min (mean: 19 min). Only 65% of the aphids tested made phloem contact within 90 min, contacting phloem after 18–85 min (mean: 32 min). No significant difference was observed in the feeding behavior of aphids carrying barley yellow dwarf virus (BYDV) from that of non-viruliferous aphids.Penetration of a sieve element was a prerequisite for BYDV transmission but did not insure transmission. Penetration of one sieve element resulted in a 65% chance of transmission independent of the duration of phloem contact. The chance of transmission increased with increasing number of sieve element penetrations.Inoculation of oat seedlings with single, viruliferous aphids for 90 min is estimated to cause 54% of the plants to be infected. Also, it is estimated that no transmission can occur with plant access periods shorter than 17 min.

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Zusammenfassung Das Probeverhalten von Sitobion avenae F. (Homoptera: Aphididae) auf Hafer (Sorte Clintland 64) wurden elektronisch verfolgt. Drei Wellenformen (S=Speichelfluss, X-das Eindringen in ein Siebelement, I=Nahrungsaufname) wurden registriert. In histologischen Untersuchungen wurden diese Wellenformen mit der Position der Stechborsten korreliert. Wenn X-Wellen oder eine X-I-Folge registriert wurden, war die Stechborste immer im Phloëm; bei S-Wellen oder bei einer S-I-Folge, war die Stechborste nie im Phloëm.Die Blattläuse wurden auf gesunden, fünf Tage alten Pflanzen während 90 Min beobachtet. 65% der Blattläuse erreichten nach 18–85 Min (Durchschnitt 32 Min) das Phloëm. Innerhalb 90 Min nahmen die Blattläuse während 0–43 Min (Durchschnitt 8.1 Min) Saft aus dem Phloem und während 0–60 Min (Durchschnitt 19 Min) Saft aus dem Mesophyll auf. Keine signifikanten Unterschiede im Probeverhalten wurden bei Blattläusen mit und ohne Barley Yellow Dwarf Virus (BYDV) festgestellt.Blattläuse, die innerhalb 24 Stunden BYDV übermitteln können, wurden für Übertragungsversuche verwendet. Das Probeverhalten dieser Blattläuser wurde manipuliert, und die Leistungsfähigkeit der Übertragung mit einer immunologischen Technik (ELISA) untersucht. Um BYDV zu übertragen mussten die Blattläuse mit einem Siebelement in Kontakt kommen. Nach der Stechborstenpenetration in ein Siebelement wurden 65% der Pflanzen mit BYDV infiziert. Der Prozentsatz infizierter Pflanzen und der Virusiter in den infizierten Pflanzen waren mit der Dauer der Siebelement-penetration (Anzahl von X-Wellen) nicht proportional. Wenn die Blattläuse mit zwei oder drei Siebelementen in Kontakt kamen, wurde der Prozentsatz infizierter Pflanzen signifikant erhöht, während der Virustiter nich verändert wurde. Infektionsprozente niedriger als 100% nach Siebelement-penetration sind möglicherweise das Resultat von Unterschieden in den Siebelementen.Es wird geschätzt dass 50% Infektion eintritt, wenn Pflanzen während ungefähr 83 Min von einer einzelnen infizierten Blattlaus besogen werden. Keine Übertragung von BYDV kann eintreten, wenn die Probezeit weniger als 17 Minuten beträgt.