The Gilded Wings appear to be two red split halves with a white neon color in the middle. On both sides, there are sets of golden brown rings. On the back there are two white rings that connect one golden beam each to the center of the band, making them look similar to wings.

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Cucumber mosaic virus (CMV) often accompanies a short RNA molecule called a satellite RNA (satRNA). When infected with CMV in the presence of Y-satellite RNA (Y-sat), tobacco leaves develop a green mosaic, then turn yellow. Y-sat has been identified in the fields in Japan. Here, we show that the yellow leaf colour preferentially attracts aphids, and that the aphids fed on yellow plants, which harbour Y-sat-derived small RNAs (sRNAs), turn red and subsequently develop wings. In addition, we found that leaf yellowing did not necessarily reduce photosynthesis, and that viral transmission was not greatly affected despite the low viral titer in the Y-sat-infected plants. Y-sat-infected plants can therefore support a sufficient number of aphids to allow for efficient virus transmission. Our results demonstrate that Y-sat directly alters aphid physiology via Y-sat sRNAs to promote wing formation, an unprecedented survival strategy that enables outward spread via the winged insect vector.

Satellite RNAs (satRNAs), which are associated with helper viruses, belong to the group of subviral agents. Cucumber mosaic virus (CMV) satRNAs ranging from 300 to 400 nucleotides in size are regarded as long noncoding RNAs1. CMV satRNAs typically reduce the viral titre and are transmitted with CMV by aphids. They rely on the helper virus CMV for their replication and encapsidation2,3. CMV satRNAs generally have a negative effect on the accumulation and vector transmission of the virus and often alter viral symptoms3,4. Y-satellite RNA (Y-sat) has been discovered in the fields in Japan in 19815 and still sporadically found in the fields. It induces bright yellow symptoms on Nicotiana species, although it can somehow attenuate CMV-induced symptoms6. In our previous investigation of persistence between four CMV satRNAs in mixed inoculation, we found that Y-sat is less competitive with other CMV satRNAs7. For the molecular mechanism for the yellow symptom induction by Y-sat, we previously demonstrated that Y-sat turned green mosaics induced by CMV to bright yellow (Fig. 1a) through the downregulation of the expression of the ChlI gene essential for chlorophyll synthesis by specific small RNAs (sRNAs) derived from Y-sat6.

What are we to conclude from this molecular crosstalk via Y-sat sRNAs that regulate the interactions among the four entities? We believe that our findings demonstrate an extraordinary role for noncoding RNA in viral persistence, especially in establishing an unusual interdependence (possibly a multilateral symbiosis) between a pathogen and its insect vector.

Thrips belong to the insect order Thysanoptera (fringe-winged insects, their wings consisting of fine hairs rather than a membrane) and TSWV vector species belong to the family Thripidae. Thrips are small, slender-bodied insects (around 1-2 mm long when fully grown, Fig. 1) that have become an important agricultural pest because of their ability to transmit plant viruses. Thrips-transmitted Tomato spotted wilt virus (TSWV), of the genus Tospovirus (Family Bunyaviridae), is the most serious disease caused by tospoviruses. TSWV is transmitted by multiple species of thrips (Ullman, 1997). Thrips responsible for spotted wilt epidemics in Georgia and Florida are western flower thrips, Frankliniella occidentalis, tobacco thrips, F. fusca, and possibly F. bispinosa (Salguero Navas et al., 1991; Riley and Pappu, 2000, 2004; and Webb et al., 1997, respectively). Other thrips species that have been reported as vectors of TSWV include Thrips tabaci, T. setosus, F. schultzei, F. intonsa, F. gemina, and F. cephalica (Riley et al. 2011), but the fore mentioned species are thought to be the most important in the Southeast.

Some seasonal factors shown to affect thrips population densities include temperature (Lowry et al. 1992, Morsello and Kennedy 2009), photoperiod (Whittaker and Kirk 2004, Chaisuekul and Riley 2005), humidity (Kirk 1997), rainfall (Morsello et al. 2010), and availability of suitable hosts for food and oviposition (Lewis 1973). Temperature is likely the major factor affecting the rate of development. The number of degree days in Celsius needed to complete one F. fusca and F. occidentalis generation from egg to adult is reported to be 234 (lower threshold 10.5C) and 253 (6.5C), respectively, (Lowry et al. 1992). Fourteen days is about the length of time needed for a western flower thrips to hatch and mature to adulthood at 25C, and slightly shorter for tobacco thrips (Lowry et al., 1992). Typically, F. occidentalis adults insert bean-shaped eggs into leaf, flower or fruit tissues (Hansen et al. 2003). Similarly, F. fusca eggs are laid singly and are usually inserted in plant tissue (usually flower parts or young leaves, Fig. 6). The eggs hatch in 3-4 days. Immatures are a cream to yellowish color (Fig. 7) and adults are either dark brown or yellow/orange, depending on the species. Two larval stages last about 10-14 days, prepupa and pupa stages last around 7 days (Lowry et al. 1992). Late second instars usually fall into the soil and pupate (Fig. 7), but some can remain on the plant (Broadbent et al. 2003). The non-feeding pupal stage is mostly immobile and has distinct and well-developed wing pads (Lewis 1973). Adults emerge within 3 days at 30C (Lowry et al. 1992). Adult F. occidentalis females can survive for 4-5 weeks at 30C and lay an average of 50 eggs (Reitz 2008). Thrips can reproduce parthenogenically. For example, fertilized F. occidentalis females are known to produce female biased sex ratios and unmated F. occidentalis females are known to produce male biased sex ratios (Lewis 1973, Moritz 1997). That is, if a female western flower thrips remain unmated then all male offspring are produced asexually. If thrips mate, they produce mostly female offspring. Adults usually have wings, although wings can be absent in some adult tobacco thrips (brachypterous form). When wings are present, they are narrow and fringed with long hairs (Figs. 2 and 3).

Most thrips vectors are assumed to have a relationship to TSWV similar to what has been well documented for western flower thrips (Ullman et al. 1997). First, immature thrips in the first and second instars acquire TSWV from infected host plants, the virus replicates in the vector as it matures, and subsequently viruliferous adults spread the virus when they move to other plants (Fig. 7). Virus multiplication in the thrips vector makes managing TSWV difficult because, once infected, adult thrips can migrate long distances to new host plants and quickly transmit the virus, often before thrips can be controlled. It is important to note that each generation of thrips must re-acquire the virus since the virus in adult females is not passed on to her eggs. Thus, the frequency of viruliferous adults can vary greatly over time. However, even low percentages of viruliferous thrips can cause significant yield loss. Viruliferous thrips can be

Thrips use piercing and sucking mouthparts to feed on plant tissue. They produce an initial opening by penetrating the plant cuticle with their single mandible after rocking the head capsule downward and upward numerous times. Once an opening or food channel is punched through the outer cell wall, then a pair of maxillary stylets is extended into the plant tissue. Saliva is injected into the plant and cellular contents are drawn up through a pumping action (Kindt et al. 2003). The adults transmit TSWV as saliva is injected into the plant tissue. Thrips feeding can damage plant tissue directly; however, the greatest loss to agriculture is caused by the species that vector (transmit) Tospoviruses like TSWV. Since TSWV transmission occurs through thrips feeding, understanding factors that affect thrips feeding is critical for managing this pest. For example, the older the plant age, the less likely that tobacco thrips will feed on the plant (Joost and Riley 2007). Also, treatment with an insecticide can affect thrips feeding behavior. For example, the insecticide imidacloprid has been shown to reduce feeding by tobacco thrips (Joost and Riley 2004). Unfortunately, imidacloprid has been shown to increase western flower thrips settling in peanut and feeding in tomato (Riley 2007), so it is important to note that insecticide effects can be specific to the thrips species.

High aspect ratio wings are potential candidates for use in atmospheric satellites and civil aircraft as they exhibit a low induced drag, which can reduce the fuel consumption. Owing to their slender and light weight configuration, such wings undergo highly flexible aeroelastic static and dynamic deformations that cannot be analyzed using conventional linear analysis methods. An aeroelastic analysis framework based on the absolute nodal coordinate formulation (ANCF) can be used to analyze the static and dynamic deformations of high aspect ratio wings. However, owing to the highly nonlinear elastic force, the statically deformed wing shape during steady flight cannot be efficiently obtained via static analyses. Therefore, an ANCF with a vector-strain transformation (ANCF-VST) was proposed in this work. Considering the slender geometry of high aspect ratio wings, the nodal vectors of an ANCF beam element were transformed to the strains. In this manner, a constant stiffness matrix and reduced degrees-of-freedom could be generated while capturing the highly flexible deformations accurately. The ANCF-VST exhibited superior convergence performance and accuracy compared to those of analytical approaches and other nonlinear beam formulations. Moreover, an aeroelastic analysis flow coupling the ANCF-VST and an aerodynamic model based on the unsteady vortex lattice method was proposed to perform the static and dynamic analyses successively. The proposed and existing aeroelastic frameworks exhibited a good agreement in the analyses, which demonstrated the feasibility of employing the proposed framework to analyze high aspect ratio wings. 2351a5e196

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