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At least 73 records · Page 4Linked to original sources

Into thin air: Contributions of aerodynamic and inertial-elastic forces to wing bending in the hawkmoth Manduca sexta.

During flapping flight, insect wings must withstand not only fluid-dynamic forces, but also inertial-elastic forces generated by the rapid acceleration and deceleration of their own mass. Estimates of overall aerodynamic and inertial forces vary widely, and the relative importance of these forces in determining passive wing deformations remains unknown. If aeroelastic interactions between a wing and the fluid-dynamic forces it generates are minor compared to the effects of wing inertia, models of insect flight that account for passive wing flexibility would be far simpler to develop. We used an experimental approach to examine the contributions of aerodynamic and inertial-elastic forces to wing bending in the hawkmoth Manduca sexta. We attached fresh Manduca wings to a motor and flapped them at a realistic wing-beat frequency and stroke amplitude. We compared wing bending in normal air versus helium (approx. 15% air density), in which the contribution of fluid-dynamic forces to wing deformations is significantly reduced. This 85% reduction in air density produced only slight changes in the pattern of Manduca wing deformations, suggesting that fluid-dynamic forces have a minimal effect on wing bending. We used a simplified finite element model of a wing to show that the differences observed between wings flapped in air versus helium are most likely due to fluid damping, rather than to aerodynamic forces. This suggests that damped finite element models of insect wings (with no fluid-dynamic forces included) may be able to predict overall patterns of wing deformation prior to calculations of aerodynamic force production, facilitating integrative models of insect flight.

Animals↗

Myogenic differentiation in early chick wing mesenchyme in the absence of the brachial somites.

A controversy exists in the literature over the ability of wing mesenchyme of somatopleural origin to form skeletal muscle. Experimental approaches used in such studies leave open the possibility of postoperative accessibility of the experimental wings to somitic cell invasion. In the present study wing somatopleural tissue was isolated from HH stage-12 to -21 chick embryos and grown either in organ culture (OC) or on the chorioallantoic membrane (CAM) of host chicks, conditions under which postoperative entry of somitic cells is impossible. In the presence of axial and somitic tissues of the brachial region, the wing territories from all four stages underwent comparable growth and tissue differentiation. However, isolated wing regions showed a stage-dependency in the differentiation of skeletal muscle but not of other limb tissues. The incidence and amount of skeletal muscle was markedly reduced in HH stage-12 and -15 isolated wing regions while myogenesis in HH stage-18 and -21 wing buds was not affected by the absence of the somitic tissues. These results are consistent with reported stages of somitic cell migration into wings with the exception of HH stage-12 explants which should have been muscleless if somitic cells are the sole source of wing myofibres. The possibility that somitic cells had been included in these explants was investigated by testing the myogenic potential of lateral plate tissue adjacent to the wing, altering the dissection procedure for isolating wing territories and using antibodies to skeletal muscle myosin and actin to detect myotubes. The results from this series of experiments illustrate the need for extraordinary care in the isolation of wing regions when investigating limb-somite relationships and suggest that the myogenic capacity attributed to wing somatopleural cells in the past can be accounted for by either postoperative entry of somitic cells into experimental wings or inadvertent inclusion of somitic cells in the primordia when dissected. Overall, the results show that somitic cells are the sole source of wing myofibres for, in their absence, somatopleural cells from all mesodermally-derived wing cell types except skeletal myofibres.

Animals↗

Development of the embryonic chick wing bud from stage 24 to stage 32.

If a graft is placed in an early chick wing bud, the location of the graft after several days of further development cannot be predicted solely from the rate of proximal-distal outgrowth. The movement of the graft depends on the rate of outgrowth of the wing but also on morphogenetic tissue movements intrinsic to the wing and on accommodation to the growth and morphogenetic movements of the body of the embryo. Numerous experiments have been reported in which tissue grafted into ectopic sites in the wing causes abnormal wing development. These experiments have been discussed in terms of pattern formation or positional information. However, until the movement of wing tissue during normal development is understood, it cannot be known in what way the development of grafts placed in ectopic sites is abnormal. Previous experiments have demonstrated that carbon particles placed in the wing move in the same manner as grafts of wing mesenchyme, but the carbon particles do not affect normal wing development. Carbon particles were placed in the wing, dorsal to the base of the wing, and cranial and caudal to the wing, to plot the expected movement of a graft and to discover how this movement can be predicted from the tissue movements at the base of the wing. It is concluded that three tissue movements are responsible for the movement of a graft. These are outgrowth at a rate determined by the rate of cell division, formation of the shoulder through caudal movement of the tissues cranial to the wing, and ventral movement of prospective flank ventral to somite 19. These three tissue movements and their influence on normal wing development are discussed.

Animals↗

Aerodynamic yawing moment characteristics of bird wings.

The aerodynamic yawing moments due to sideslip are considered for wings of birds. Reference is made to the experience with aircraft wings in order to identify features which are significant for the yawing moment characteristics. Thus, it can be shown that wing sweep, aspect ratio and lift coefficient have a great impact. Focus of the paper is on wing sweep which can considerably increase the yawing moment due to sideslip when compared with unswept wings. There are many birds the wings of which employ sweep. To show the effect of sweep for birds, the aerodynamic characteristics of a gull wing which is considered as a representative example are treated in detail. For this purpose, a sophisticated aerodynamic method is used to compute results of high precision. The yawing moments of the gull wing with respect to the sideslip angle and the lift coefficient are determined. They show a significant level of yaw stability which strongly increases with the lift coefficient. It is particularly high in the lift coefficient region of best gliding flight conditions. In order to make the effect of sweep more perspicuous, a modification of the gull wing employing no sweep is considered for comparison. It turns out that the unswept wing yields yawing moments which are substantially smaller than those of the original gull wing with sweep. Another feature significant for the yawing moment characteristics concerns the fact that sweep is at the outer part of bird wings. By considering the underlying physical mechanism, it is shown that this feature is most important for the efficiency of wing sweep. To sum up, wing sweep provides a primary contribution to the yawing moments. It may be concluded that this is an essential reason why there is sweep in bird wings.

Aircraft↗

The genetic basis of the interspecific differences in wing size in Nasonia (Hymenoptera; Pteromalidae): major quantitative trait loci and epistasis.

There is a 2.5-fold difference in male wing size between two haplodiploid insect species, Nasonia vitripennis and N. giraulti. The haploidy of males facilitated a full genomic screen for quantitative trait loci (QTL) affecting wing size and the detection of epistatic interactions. A QTL analysis of the interspecific wing-size difference revealed QTL with major effects and epistatic interactions among loci affecting the trait. We analyzed 178 hybrid males and initially found two major QTL for wing length, one for wing width, three for a normalized wing-size variable, and five for wing seta density. One QTL for wing width explains 38.1% of the phenotypic variance, and the same QTL explains 22% of the phenotypic variance in normalized wing size. This corresponds to a region previously introgressed from N. giraulti into N. vitripennis that accounts for 44% of the normalized wing-size difference between the species. Significant epistatic interactions were also found that affect wing size and density of setae on the wing. Screening for pairwise epistatic interactions between loci on different linkage groups revealed four additional loci for wing length and four loci for normalized wing size that were not detected in the original QTL analysis. We propose that the evolution of smaller wings in N. vitripennis males is primarily the result of major mutations at few genomic regions and involves epistatic interactions among some loci.

Animals↗

Investigations for the diagnostic recording of nasal wing collapse.

BACKGROUND: The inspiratory medial movement of the nasal wing at high flow velocities is a protective physiologic mechanism. If this collapse of the nasal wing occurs at lower flow velocities, it may result in nasal obstruction. "Nasal wing collapse" is generally a clinical diagnosis. However, in the pressure-flow relationship of rhinomanometry, the medial movement of the nasal wing can be documented in the inspiratory arm of the graph. The diagnostic impact of this hysteresis was investigated. METHODS: The pressure-flow curves of three box models and three nasal models with a moveable valve (analogous to the nasal wing) in the entrance area as well as three volunteers with unstable nasal wings were investigated. We recorded synchronously the pressure-flow relationship and by endoscopy the movement of the valve in the box models and the nasal wing in the volunteers on video. For evaluation, we used the frame by frame analysis of the tape. RESULTS AND CONCLUSIONS: The medial movement of the nasal wing causes a hysteresis in the inspiratory arm of the pressure-flow curve. At this point, the graph runs on or between two border curves, termed the "no collapse curve" (for the maximally opened valve or a stable nasal wing) and the "collapse curve" (for the subtotally closed valve or a collapsed nasal wing), respectively. Analogous to the nasal wing motion, the descending course of hysteresis runs in two phases, and the ascending course runs in three phases. The medial movement of the nasal wing is expressed by a deviation of the graph from the "no collapse curve." The flow, at which the graph leaves this curve, depends on the elasticity module of the nasal wing. The extent of nasal wing collapse is reflected by the approximation of the pressure-flow curve to the "collapse curve" of the graph. The hysteresis appears because of a late opening of the collapsed nasal wing.

Adult↗

The development of the posterior musculature of the embryonic chick wing.

It has been proposed that movements of cells at the base of the developing wing cause the wing to rotate (Yander and Searls, '80a,b). It was suggested that rotation of the wing caused the dorsal muscle mass of the wing bud to become the posterior musculature of the proximal part of the wing. Rotation of the wing was investigated by grafting dorsal myogenic tissue from a wing labeled with tritiated thymidine into the dorsal myogenic region of an unlabeled wing in its original orientation. Grafts were made both with and without their original ectoderm. The location and shape of the graft was recorded at the time of the operation. After a period of growth and morphogenesis, the host wing and adjacent body wall were fixed, sectioned, and prepared as autoradiographs. The location of the grafted cells was determined by reconstructing the host wing and adjacent body wall. It was found that the graft increased in length on the axis of the humerus and did not increase rapidly on any other axis. During stage 25, a deep groove began to form ventral to somite 19 because of ventral and cranial movement of the lateral body wall proximal to the base of the wing. The distal border of this groove was recognized as the posterior margin of the proximal wing. If the graft extended to the base of the wing, the graft participated in the formation of this groove. Grafted cells on the distal border of the groove were on the posterior surface of the wing at the level of the humerus.

Animals↗

Evidence for inherent morphogenetic properties of the myogenic regions of the embryonic chick wing.

The dorsal and ventral myogenic regions were isolated from the right and left wings of donor chick embryos. After removal of the ectoderm, the myogenic regions were grafted into the dorsal myogenic region of the right wing of host embryos in normal or reversed proximal-distal orientation. All grafts were made so that the surface of the graft originally in contact with ectoderm was dorsal. It was expected that myogenic regions grafted into the myogenic region would participate in the normal formation of the host wing. However, after 7 days of further development many of the host wings had developed abnormal cartilages. To investigate the interactions between the tissue of the host and the tissue of the graft and the source of the tissue giving rise to ectopic cartilages, the donor embryos were labeled with tritiated thymidine. The location and shape of the graft were recorded at the time of the operation. The host wings and adjacent body wall were fixed 1, 3, and 4 days after the operation, sectioned, and prepared as autoradiographs. The location of the grafted cells in the host wing was determined by reconstructing the host wing and adjacent body wall. Serial sections and the reconstructions were examined to discover the correlates of abnormal cartilage formation. The following observations were made: 1) Grafts from the dorsal region of the right and left wing in normal orientation participated in the formation of the groove at the base of the wing ventral to the nineteenth somite that produces the extensor surface of the elbow. Grafts from the ventral region of the right and left wing in normal orientation, and all grafts in reversed orientation, did not participate in the formation of the groove. 2) Grafts in normal orientation increased in length on the axis of the humerus and did not increase in width. Grafts in reversed orientation increased in both length and width. Increase in width caused the graft to extend into the proximo-caudal corner of the wing. 3) Wings with grafts from the dorsal regions of the right and left wings in normal orientation were not clearly abnormal 4 days after the operation. When the groove ventral to the nineteenth somite did not form normally, ectopic cartilage differentiated where the groove should have been from grafted cells of the myogenic region. Increase in width of the graft into the proximo-caudal corner of the wing produced an increased mass of graft cells ventral to the nineteenth somite and an increased amount of ectopic cartilage.

Animals↗

Changing distributions of extracellular matrix components during early wing morphogenesis in Drosophila.

A new monoclonal antibody, specific to an epitope in the carboxyl terminus of the Drosophila collagen IV molecule (basement membrane collagen) was identified. The distributions of collagen IV, laminin, and an additional extracellular molecule, the 2G2 antigen (2G2-Ag), were followed immunocytochemically during early wing development. In late third instar larvae, collagen IV and laminin surround the entire wing disc, whereas the 2G2-Ag is limited to the region of the future wing pouch. For the first few hours following eversion of the disc, all three ECM components line the basal surfaces of all epithelial cells in the wing pouch, both those destined to line the wing veins and those destined to become tightly apposed in the large intervein regions. Collagen IV and laminin persist on these cells during the two initial rounds of apposition of dorsal and ventral wing surfaces; later, they become restricted to the cells lining the veins. The 2G2-Ag disappears completely quite early in the pupal period. Collagen IV appears to be synthesized at least twice, once in the larva and a second time in the pupa; in between it is enzymatically cleaved and may be eliminated, probably by hemocytes. In an extreme allele of blistered the wing is ballooned to form a single internal space. Collagen IV and laminin line all basal wing cell surfaces early in pupal development as they do in the wild type. Later, however, they continue to line the entire cavity of the mutant wing rather than assuming a restricted distribution. In a completely veinless wing (rhomboidveinletvein), collagen IV and laminin are also present generally on basal surfaces at early times, but are completely absent between the tightly apposed wing layers later. The ECM distributions both in wild type wings and in mutants suggest that the matrix plays a role in the establishment of the wing venation pattern. One possibility, strengthened by recent findings regarding ECM receptors in Drosophila, is their involvement in dorsal-ventral wing layer adhesion. Our findings also lead us to suggest that certain sets of features which distinguish vein from intervein cells may be linked during cell differentiation and thus help to define these cell phenotypes. The features include cytoskeletal specializations and certain cell surface and ECM molecules.

Animals↗

The function of the frizzled pathway in the Drosophila wing is dependent on inturned and fuzzy.

The Drosophila epidermis is characterized by a dramatic planar or tissue polarity. The frizzled pathway has been shown to be a key regulator of planar polarity for hairs on the wing, ommatidia in the eye, and sensory bristles on the notum. We have investigated the genetic relationships between putative frizzled pathway downstream genes inturned, fuzzy, and multiple wing hairs (inturned-like genes) and upstream genes such as frizzled, prickle, and starry night (frizzled-like genes). Previous data showed that the inturned-like genes were epistatic to the frizzled-like genes when the entire wing was mutant. We extended those experiments and examined the behavior of frizzled clones in mutant wings. We found the domineering nonautonomy of frizzled clones was not altered when the clone cells were simultaneously mutant for inturned, multiple wing hairs, or dishevelled but it was blocked when the entire wing was mutant for inturned, fuzzy, multiple wing hairs, or dishevelled. Thus, for the domineering nonautonomy phenotype of frizzled, inturned and multiple wing hairs are needed in the responding cells but not in the clone itself. Expressing a number of frizzled pathway genes in a gradient across part of the wing repolarizes wing cells in that region. We found inturned, fuzzy, and multiple wing hairs were required for a gradient of frizzled, starry night, prickle, or spiny-legs expression to repolarize wing cells. These data argue that inturned, fuzzy, and multiple wing hairs are downstream components of the frizzled pathway. To further probe the relationship between the frizzled-like and inturned-like genes we determined the consequences of altering the activity of frizzled-like genes in wings that carried weak alleles of inturned or fuzzy. Interestingly, both increasing and decreasing the activity of frizzled and other upstream genes enhanced the phenotypes of hypomorphic inturned and fuzzy mutants. We also examined the relationship between the frizzled-like and inturned-like genes in other regions of the fly. For some body regions and cell types (e.g., abdomen) the inturned-like genes were epistatic to the frizzled-like genes, but in other body regions (e.g., eye) that was not the case. Thus, the genetic control of tissue polarity is body region specific.

Animals↗

Coordination of wingbeat and respiration in the Canada goose. I. Passive wing flapping.

The effects of passive wing flapping on respiratory pattern were examined in decerebrate Canada geese. The birds were suspended dorsally with two spine clamps while the extended wings were continuously moved up and down with a device designed to reproduce actual wing flapping. Passive wing motion entrained respiration over limited ranges by both increasing and decreasing the respiratory period relative to rest. All ratios of wingbeat frequency to respiratory frequency seen during free flight (Soc. Neurosci. Abstr. 15: 391, 1989) were produced during passive wing flapping. In addition, the phase relationship between wingbeat frequency and respiratory frequency, inspiration starting near the peak of wing upstroke, was similar to that seen during free flight and was unaffected by perturbations of the wing-flapping cycle. Removal of all afferent activity from the wings did not affect the ability of continuous passive wing movement to entrain respiration. However, feedback from the wings was required to produce rapid within-breath shifts in the respiratory period in response to single wing flaps. In conclusion, although feedback from the chest wall/lung may be more important in producing entrainment during the stable conditions of passive wing flapping, wing-related feedback may be critically involved in mediating the rapid adjustments in respiratory pattern required to maintain coordination between wing and respiratory movements during free flight.

Afferent Pathways↗

Shape, flapping and flexion: wing and fin design for forward flight.

Both kinematics and morphology are critical determinants of performance in flapping flight. However, the functional consequences of changes in these traits are not yet well understood. Traditional aerodynamic studies of planform wing shape have suggested that high-aspect-ratio wings generate more force per area and perform more efficiently than low-aspect-ratio wings, but these analyses may neglect critical components of flapping flight such as unsteady fluid dynamics and wing or fin flexion. In this paper, we use an unsteady potential flow analysis that incorporates wing flexion to test predictions of optimal wing shape under varying degrees of unsteady motion and wing flexion. We focus on forward flapping flight and examine the effects of wing/fin morphology and movements on thrust generation and efficiency. We test the model by comparing our predictions with kinematic data derived from the aquatic flight of the ratfish Hydrolagus colliei. Our analyses show that aspect ratio and the proportion of area in the outer one-fifth of the wing can characterize wing shape in terms of aero- or hydrodynamic performance. By comparing the performance of wings that vary in these two parameters, we find that traditional predictions of optimal wing shape are valid only under limited circumstances (when flapping frequency is low, wings are stiff or wings are tapered at the tips). This indicates a complex relationship between locomotor traits and performance and helps explain the diversity of wing kinematics and morphologies observed in nature.

Animals↗

Effects of applying Safe2O poultry wash to broiler wings on shelf life, Listeria monocytogenes, Pseudomonads, Staphylococcus species, and psychrotrophic bacteria levels after three, seven, and ten days of storage.

Bacterial contamination of raw processed poultry continues to be of concern to consumers as well as regulatory and health officials. For many years wings were considered a low-value product; therefore, shelf life of wings was not a major concern. Due to changes in consumer attitudes and increases in the fast-food market, wings are now a valuable commodity. Because wings have a shorter shelf life than most other raw poultry products, acceptable intervention to decrease the population of associated spoilage organisms and human enteropathogens are needed. Safe2O Poultry Wash was evaluated as a postchill treatment to reduce microbial contamination and increase shelf life. Ninety-six carcasses were obtained from a local processor prior to final wash. On arrival at the research facility all carcasses were inoculated with 1 mL of a culture with 10(3) cfu/mL Listeria monocytogenes. After a 30-min attachment time, carcasses were subjected to a 4-s in-out final wash, hung for 3 min, and chilled in ice-water for 45 min. After the chilling, wings were removed by hand with a knife, pooled together, and subjected to a hand spray (4 mL/wing) with deionized water or Safe2O Poultry Wash. Two wings were then placed in each of 96 ziplock type storage bags, and wings were held at 5 +/- 1 degrees C for 3, 7, 10, and 14 d. On the day of sample, weep was decanted, and 100 mL of Butterfield's phosphate buffer was added to each bag. Three sets of wings were shaken by hand for 1 min, and total aerobes, Pseudomonads, Staphylococcus sp., psychrotrophic bacteria, and L. monocytogenes in the rinsates were enumerated. By using 7 log10 recovery of total aerobes from rinsates as a spoilage baseline, all wings were spoiled by d 10, but the wings treated with water were approaching spoilage counts on d 7, (log10 6.8), whereas only log10 5.5 bacteria were recovered from the wings sprayed with Safe2O Poultry Wash. Fewer Pseudomonads, Staphylcoccus sp., L. monocytogenes, and psychrotrophic bacteria were recovered from wings treated with Safe2O Poultry Wash and stored for 10 d. Log10 counts for the organisms were Pseudomonas sp., 8.2 and 6.9; Staphylcoccus sp., 5.5 and 4.9; L. monocytogenes, 5.2 and 4.6; and psychrotrophs, 8.2 and 6.9 for the water and Safe2O Poultry Wash treatments, respectively. Use of the Safe2O Poultry Wash as a postchill treatment on wings could increase the shelf life of wings by up to 3 d.

Animals↗

The moment of inertia of bird wings and the inertial power requirement for flapping flight

The agility and manoeuvrability of a flying animal and the inertial power required to flap the wings are related to the moment of inertia of the wings. The moments of inertia of the wings of 29 bird species and three bat species were determined using wing strip analysis. We also measured wing length, wing span, wing area, wing mass and body mass. A strong correlation (r2=0.997) was found between the moment of inertia and the product of wing mass and the square of wing length. Using this relationship, it was found that all birds that use their wings for underwater flight had a higher than average moment of inertia. Assuming sinusoidal wing movement, the inertial power requirement was found to be proportional to (body mass)0.799, an exponent close to literature values for both metabolic power output and minimum power required for flight. Ignoring wing retraction, a fairly approximate estimate showed that the inertial power required is 11­15 % of the minimum flight power. If the kinetic energy of the wings is partly converted into aerodynamic (useful) work at stroke reversal, the power loss due to inertial effects may be smaller.

Journal Article↗

Effect of feeding a raw winged bean seeds on gastrointestinal functions in rats.

The primary cause of the adverse effects of feeding of raw winged bean seeds in rats was investigated. In experiment 1, rats were fed on either a raw winged bean diet or a steamed winged bean diet for 10 days. Body weight gain of rats fed on 30% raw winged bean diet was significantly lower than that of rats fed on 30% steamed winged bean diet. The adverse effect of the feeding of 30% raw winged bean diet on growth was accompanied by disorders of the gastrointestinal tract including a significant reduction in intestinal sucrase activity, not being improved with feeding of the diet supplemented with methionine. In experiment 2, rats fasted for 2 days were refed on a 10% casein diet, a 30% raw winged bean diet or a 30% steamed winged bean diet, in which most of the carbohydrate component was sucrose, for 4 days. Although body weight gain and food consumption in rats refed on these winged bean diets were lower than those in rats refed on 10% casein diet, the effects of feeding of the raw winged bean diet on body weight gain and food consumption were extremely deleterious as compared with those of feeding of the steamed winged bean diet. Significant reductions in hydrolase activities localized in the brush border membrane of the small intestine were found in rats refed on the raw winged bean diet prior to the occurrence of apparent disorders in the gastrointestinal tract. These findings suggest that the primary cause of the adverse effects of raw winged bean seed feeding the disorders in the small intestine caused by lectin or similar substances in raw winged bean seeds.

Alkaline Phosphatase↗

Effect of slotted wing tips on yawing moment characteristics.

The aerodynamic yawing moment characteristics of bird wings with slotted tips are dealt with. Emphasis is placed on the effect of sweep which the separated feathers constituting the wing tips show and which can reach significant values. Reference is made to basic aerodynamic characteristics of wings with sweep which yields a stabilizing yawing moment significantly larger than that of unswept wings. Then, the yawing moment characteristics are determined for a wing, the features of which are considered as representative of bird wings with sweep in their slotted tips. A sophisticated aerodynamic procedure is used for obtaining results of high precision. It is shown that the sweep in the slotted wing tips yields a stabilizing yawing moment of significant magnitude, considerably increasing with the lift coefficient. To make the significance of wing tip sweep for the ability to generate yawing moments more perspicuous, a wing modification the slotted tips of which are unswept is considered for comparison. It turns out that this wing shows yawing moments which are substantially smaller. A physical insight into the effect of slotted wing tip sweep on the aerodynamic yawing moment characteristics is provided by showing the underlying mechanism. From the results presented in this paper it follows that the sweep in slotted wing tips provides a substantial contribution to the aerodynamic yawing moment and, thus, to yaw stability. It may be concluded that this is an essential reason why there is sweep in the slotted tips of bird wings.

Animals↗

Hemocytes are essential for wing maturation in Drosophila melanogaster.

Newly eclosed flies have wings that are highly folded and compact. Within an hour, each wing has expanded, the dorsal and ventral cuticular surfaces bonding to one another to form the mature wing. To initiate a dissection of this process, we present studies of two mutant phenotypes. First, the batone mutant blocks wing expansion, a behavior that is shown to have a mutant focus anterior to the wing in the embryonic fate map. Second, ectopic expression of protein kinase A catalytic subunit (PKAc) using certain GAL4 enhancer detector strains mimics the batone wing phenotype and also induces melanotic "tumors." Surprisingly, these GAL4 strains express GAL4 in cells, which seem to be hemocytes, found between the dorsal and ventral surfaces of newly opened wings. Ectopic expression of Ricin A in these cells reduces their number and prevents bonding of the wing surfaces without preventing wing expansion. We propose that hemocytes are present in the wing to phagocytose apoptotic epithelial cells and to synthesize an extracellular matrix that bonds the two wing surfaces together. Hemocytes are known to form melanotic tumors either as part of an innate immune response or under other abnormal conditions, including evidently ectopic PKAc expression. Ectopic expression of PKAc in the presence of the batone mutant causes dominant lethality, suggesting a functional relationship. We propose that batone is required for the release of a hormone necessary for wing expansion and tissue remodeling by hemocytes in the wing.

Animals↗

Determination of wing cell fate by the escargot and snail genes in Drosophila.

Inset appendages such as the wing and the leg are formed in response to inductive signals in the embryonic field. In Drosophila, cells receiving such signals initiate developmental programs which allow them to become imaginal discs. Subsequently, these discs autonomously organize patterns specific for each appendage. We here report that two related transcription factors, Escargot and Snail that are expressed in the embryonic wing disc, function as intrinsic determinants of the wing cell fate. In escargot or snail mutant embryos, wing-specific expression of Snail, Vestigial and beta-galactosidase regulated by escargot enhancer were found as well as in wild-type embryos. However, in escargot snail double mutant embryos, wing development proceeded until stage 13, but the marker expression was not maintained in later stages, and the invagination of the primordium was absent. From such analyses, it was concluded that Escargot and Snail expression in the wing disc are maintained by their auto- and crossactivation. Ubiquitous escargot or snail expression induced from the hsp70 promoter rescued the escargot snail double mutant phenotype with the effects confined to the prospective wing cells. Similar DNA binding specificities of Escargot and Snail suggest that they control the same set of genes required for wing development. We thus propose the following scenario for early wing disc development. Prospective wing cells respond to the induction by turning on escargot and snail transcription, and become competent for regulation by Escargot and Snail. Such cells initiate auto- and crossregulatory circuits of escargot and snail. The sustained Escargot and Snail expression then activates vestigial and other target genes that are essential for wing development. This maintains the commitment to the wing cell fate and induces wing-specific cell shape change.

Animals↗