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PubMed2004

Genetic architecture of wing morphology in Drosophila simulans and an analysis of temperature effects on genetic parameter estimates.

The Drosophila wing has been used as a model to investigate the mechanisms responsible for size and shape changes in nature, since such changes might underlie morphological evolution. To improve the understanding of wing morphological variation and the interpretation of genetic parameters estimates, we have established 59 lines from a Drosophila simulans laboratory population through single pair random matings. The offspring of each line were reared at three different temperatures, and the wing morphology of 12 individuals was analyzed by adjusting an ellipse to the wings' contour. Temperature, sex and line significantly affected wing trait variation, which was mainly characterized by longer wings having the second, fourth and fifth longitudinal veins closer together at the wing tip. As for the genetic parameter estimates, while the cross-environment heritability of some traits, such as wing size (SI), decreased with an increasing difference between the temperatures at which parents and offspring were reared, wing shape (SH) heritability did not seem to change. Since we found indications that neither an increase in the phenotypic variation nor the occurrence of genotype-environment interactions could fully explain the low heritabilities of SI estimated by cross-environment regressions, we discuss the importance of other effects for explaining this discrepancy between the SI and SH heritability estimates. In addition, although the genetic matrix was not entirely represented in the phenotypic matrix, several correspondences were identified, suggesting that the observed patterns of wing morphology variation are genetically controlled.

Animals↗
PubMed1994

Blistered: a gene required for vein/intervein formation in wings of Drosophila.

We have characterized the blistered (bs) locus phenotypically, genetically and developmentally using a set of new bs alleles. Mutant defects range from wings with ectopic veins and intervein blisters to completely ballooned wings where the distinction between vein and intervein is lost. Mosaic analyses show that severe bs alleles behave largely autonomously; homozygous patches having vein-like properties. Developmental analyses were undertaken using light and electron microscopy of wild-type and bs wings as well as confocal microscopy of phalloidin- and laminin-stained preparations. bs defects were first seen early in the prepupal period with the failure of apposition of dorsal and ventral wing epithelia. Correspondingly, during definitive vein/intervein differentiation in the pupal period (18-36 hours after puparium formation), the extent of dorsal/ventral reapposition is reduced in bs wings. Regions of the wing that fail to become apposed differentiate properties of vein cells; i.e. become constricted apically and acquire a laminin-containing matrix basally. To further understand bs function, we examined genetic interactions between various bs alleles and mutants of two genes whose products have known functions in wing development. (i) rhomboid, a component of the EGF-R signalling pathway, is expressed in vein cells and is required for specification of vein cell fate. rhove mutations (lacking rhomboid in wings) suppress the excess vein formation and associated with bs. Conversely, rho expression in prepupal and pupal bs wings is expanded in the regions of increased vein formation. (ii) The integrin genes, inflated and myospheroid, are expressed in intervein cells and are required for adhesion between the dorsal and ventral wing surfaces. Loss of integrin function results in intervein blisters. Integrin mutants interact with bs mutants to increase the frequency of intervein blisters but do not typically enhance vein defects. Both developmental and genetic analyses suggest that the bs product is required during metamorphosis for the initiation of intervein development and the concomitant inhibition of vein development.

Animals↗
PubMed1996

Distinct spatial and temporal functions for PS integrins during Drosophila wing morphogenesis.

At the onset of pupariation in the Drosophila wing, the PS1 and PS2 integrins are expressed preferentially on the dorsal and ventral wing epithelia, respectively. Clonal analysis experiments have indicated that integrins are required to maintain the tight association of the wing surfaces. Surprisingly, we find that even in clones of cells lacking integrins the wing layers become apposed early in metamorphosis. However, following the normal period of wing separation, large integrin mutant clones do not become re-apposed in the pupa, and integrins are not organized in basal plaques in cells opposite a mutant clone. Paradoxically, our experiments indicate that at least one integrin function requires different integrins on the dorsal and ventral wing surfaces, however in some cases both alphaPS subunits can function to some degree on each wing surface. Finally, overexpression of an alphaPS subunit throughout the wing leads to a dominant wing blister phenotype, and the critical period for this phenotype is the beginning of pupariation. These data indicate that integrin requirements in wing morphogenesis can be separated into early (prepupal) and late (pupal) functions. The late function seems to reflect the traditional view of integrins as cell-matrix adhesion proteins. The early requirement, which probably requires dorsoventral segregation of PS1 and PS2, suggests functions for PS1 and PS2 in signaling events that regulate morphogenesis.

Animals↗
PubMed1999

The vestigial gene product provides a molecular context for the interpretation of signals during the development of the wing in Drosophila.

The vestigial (vg) gene of Drosophila plays a central role in the development and patterning of the wing: loss of vestigial results in failures in wing development and ectopic expression of vestigial leads to the development of ectopic wings. The wing-specific regulation of vestigial is mediated through two enhancers: (1) the Boundary Enhancer (vgBE) is early acting and becomes restricted to the wing margin, and (2) the Quadrant Enhancer (vgQE), acts later and is responsible for the expression of vestigial in the developing wing blade. These enhancers receive regulatory inputs from three signalling pathways: wingless, decapentaplegic and Notch/Suppressor of Hairless. Our experiments show that the vestigial gene product is also an input in the regulation of vestigial expression. In particular, Vestigial provides an important input for the regulation of the activity of the vgQE acting in concert with Wingless and Decapentaplegic. Our results suggest how interactions between vgBE and the vgQE mediated by Vestigial can explain the interactions between the wing margin and the wing blade during the growth of the wing. We further show that Vestigial and Notch collaborate with Wingless to subdivide and pattern the wing blade. These results lead us to propose a general role for Wingless during development in which it stabilizes cell fate decisions that have been implemented by other molecules.

Animals↗
PubMed2003

Aerodynamic force generation and power requirements in forward flight in a fruit fly with modeled wing motion.

Aerodynamic force generation and power requirements in forward flight in a fruit fly with modeled wing motion were studied using the method of computational fluid dynamics. The Navier-Stokes equations were solved numerically. The solution provided the flow velocity and pressure fields, from which the vorticity wake structure and the unsteady aerodynamic forces and torques were obtained (the inertial torques due to the acceleration of the wing-mass were computed analytically). From the flow-structure and force information, insights were gained into the unsteady aerodynamic force generation. On the basis of the aerodynamic and inertial torques, the mechanical power was obtained, and its properties were investigated. The unsteady force mechanisms revealed previously for hovering (i.e. delayed stall, rapid acceleration at the beginning of the strokes and fast pitching-up rotation at the end of the strokes) apply to forward flight. Even at high advance ratios, e.g. J=0.53-0.66 (J is the advance ratio), the leading edge vortex does not shed (at such advance ratios, the wing travels approximately 6.5 chord lengths during the downstroke). At low speeds (J approximately equal to 0.13), the lift (vertical force) for weight support is produced during both the down- and upstrokes (the downstroke producing approximately 80% and the upstroke producing approximately 20% of the mean lift), and the lift is contributed mainly by the wing lift; the thrust that overcomes the body drag is produced during the upstroke, and it is contributed mainly by the wing drag. At medium speeds (J approximately equal to 0.27), the lift is mainly produced during the downstroke and the thrust mainly during the upstroke; both of them are contributed almost equally by the wing lift and wing drag. At high speeds (J approximately equal to 0.53), the lift is mainly produced during the downstroke and is mainly contributed by the wing drag; the thrust is produced during both the down- and upstrokes, and in the downstroke, is contributed by the wing lift and in the upstroke, by the wing drag. In forward flight, especially at medium and high flight speeds, the work done during the downstroke is significantly greater than during the upstroke. At advance ratios J approximately equal to 0.13, 0.27 and 0.53, the work done during the downstroke is approximately 1.6, 2.8 and 4.2 times as much as that during the upstroke, respectively. At J=0 (hovering), the body-mass-specific power is approximately 29 W kg(-1); at J=0.13 and 0.27, the power is approximately 10% less than that of hovering; at J=0.40, the power is approximately the same as that of hovering; when J is further increased, the power increases sharply. The graph of power against flying speeds is approximately J-shaped. From the graph of power against flying speeds, it is predicted that the insect usually flies at advance ratios between zero and 0.4, and for fast flight, it would fly at an advance ratio between 0.4 and 0.53.

Animals↗
PubMed2003

Evaluation of a safety resheathable winged steel needle for prevention of percutaneous injuries associated with intravascular-access procedures among healthcare workers.

OBJECTIVE: To compare the percutaneous injury rate associated with a standard versus a safety resheathable winged steel (butterfly) needle. DESIGN: Before-after trial of winged steel needle injuries during a 33-month period (19-month baseline, 3-month training, and 11-month study intervention), followed by a 31-month poststudy period. SETTING: A 1,190-bed acute care referral hospital with inpatient and outpatient services in New York City. PARTICIPANTS: All healthcare workers performing intravascular-access procedures with winged steel needles. INTERVENTION: Safety resheathable winged steel needle. RESULTS: The injury rate associated with winged steel needles declined from 13.41 to 6.41 per 100,000 (relative risk [RR], 0.48; 95% confidence interval [CI95], 0.31 to 0.73) following implementation of the safety device. Injuries occurring during or after disposal were reduced most substantially (RR, 0.15; CI95, 0.06 to 0.43). Safety winged steel needle injuries occurred most often before activation of the safety mechanism was appropriate (39%); 32% were due to the user choosing not to activate the device, 21% occurred during activation, and 4% were due to improper activation. Preference for the safety winged steel needle over the standard device was 63%. The safety feature was activated in 83% of the samples examined during audits of disposal containers. Following completion of the study, the safety winged steel needle injury rate (7.29 per 100,000) did not differ significantly from the winged steel needle injury rate during the study period. CONCLUSION: Implementation of a safety resheathable winged steel needle substantially reduced injuries among healthcare workers performing vascular-access procedures. The residual risk of injury associated with this device can be reduced further with increased compliance with proper activation procedures.

Blood-Borne Pathogens↗
PubMed1995

Drag reduction by wing tip slots in a gliding Harris' hawk, Parabuteo unicinctus

The anterior-most primary feathers of many birds that soar over land bend upwards and separate vertically to form slotted wing tips during flight. The slots are thought to reduce aerodynamic drag, although drag reduction has never been demonstrated in living birds. Wing theory explains how the feathers that form the tip slots can reduce induced drag by spreading vorticity horizontally along the wing and by acting as winglets, which are used on aircraft to make wings non-planar and to spread vorticity vertically. This study uses the induced drag factor to measure the induced drag of a wing relative to that of a standard planar wing with the same span, lift and speed. An induced drag factor of less than 1 indicates that the wing is non-planar. The minimum drag of a Harris' hawk gliding freely in a wind tunnel was measured before and after removing the slots by clipping the tip feathers. The unclipped hawk had 70­90 % of the drag of the clipped hawk at speeds between 7.3 and 15.0 m s-1. At a wing span of 0.8 m, the unclipped hawk had a mean induced drag factor of 0.56, compared with the value of 1.10 assumed for the clipped hawk. A Monte Carlo simulation of error propagation and a sensitivity analysis to possible errors in measured and assumed values showed that the true mean value of the induced drag factor for the unclipped hawk was unlikely to be more than 0.93. These results for a living bird support the conclusions from a previous study of a feathered tip on a model wing in a wind tunnel: the feathers that form the slotted tips reduce induced drag by acting as winglets that make the wings non-planar and spread vorticity both horizontally and vertically.

Journal Article↗
PubMed2006

Ectopic expression of Tollo/Toll-8 antagonizes Dpp signaling and induces cell sorting in the Drosophila wing.

The wing imaginal disc of Drosophila consists of the primordia for the adult wing and the body wall. The zinc-finger transcription factor Teashirt (Tsh) is expressed in the region proximal to the wing primordium and regulates the formation of the wing-body wall boundary. Here, we report that Tollo/Toll-8, a member of Toll family transmembrane proteins, is also expressed proximal to the wing domain. Ectopic expression of Decapentaplegic (Dpp), a morphogen for wing development, represses tollo expression in the proximal domain. Likewise, misexpression of Tollo in the presumptive wing strongly antagonizes the effects of Dpp signaling. The extracellular domain of Tollo containing the Leucine-Rich Repeats (LRR) is required for the inhibition of Dpp signaling in the wing. Furthermore, clones of cells with Tollo overexpression are sorted out from the surrounding wild-type cells, resulting in the formation of epithelial folds around the clone boundaries. Tsh is ectopically induced at the border of Tollo-expressing clones. Despite the strong effects of Tollo overexpression on Dpp signaling and cell sorting, loss-of-function tollo mutants are viable with normal external morphology. Our data suggest that Tollo function might be redundant but is sufficient to antagonize Dpp signaling and induce sorting of Tollo expressing cells from the wing cells to develop proximal cell fate.

Animals↗
PubMed2005

The digits of the wing of birds are 1, 2, and 3. A review.

Fossil evidence documenting the evolutionary transition from theropod dinosaurs to birds indicates unambiguously that the digits of the wing of birds are digits 1, 2, and 3. However, some embryological evidence suggests that these digits are 2, 3, and 4. This apparent lack of correspondence has been described as the greatest challenge to the widely accepted theropod-bird link (Zhou 2004. Naturwissenschaften 91:455-471). Here we review the pertinent literature regarding the debate on the origin of birds and wing digital identity and the evidence in favor of a 1, 2, 3 identity of the wing digits. Recent molecular evidence shows that the expression of Hoxd12 and Hoxd13 in the developing wing supports the theropod-bird link. In the chicken foot and in the mouse hand and foot, digit 1 is the only digit to combine the expression of Hoxd13 with the absence of expression of Hoxd12. The same is observed in the anterior digit of the wing, suggesting it is a digit 1, as expected for a theropod. Nevertheless, Galis et al. (2005. J Exp Zool (Mol Dev Evol) in press), argue that Hoxd12 and Hoxd13 expression patterns in mutant limbs do not allow distinguishing the most anterior digit in the bird wing from digit 2. They also argue that constraints to the evolution of limb development support the 2, 3, 4 identity of the wing digits. However, the case put forward by Galis et al. is biased and flawed with regard to interpretation of mutant limbs, developmental mechanisms, stages observed, and the description of the evolutionary variation of limb development. Importantly, Galis et al. do not present evidence from wild-type limbs that counters the conclusions of Vargas and Fallon (2005. J Exp Zool (Mol Dev Evol) 304B(1):85-89), and fail to provide molecular evidence to specifically support the hypothesis that the wing digits are 2, 3, and 4. The expression of Hoxd12 and Hoxd13 in the developing wing is consistent with the hypothesis that birds are living dinosaurs; this view can lead to a greater understanding of the actual limits to the evolutionary variation of limb development.

Animals↗
PubMed2004

The role of the T-box gene optomotor-blind in patterning the Drosophila wing.

The development of the Drosophila wing is governed by the action of two morphogens encoded by the genes decapentaplegic (dpp; a member of the BMP gene family) and wingless (wg; a member of the WNT gene family), which promote cell proliferation and pattern the wing. Along the anterior/posterior (A/P) axis, the precise expression of decapentaplegic and its receptors is required for the transcriptional regulation of specific target genes. In the present work, we analyze the function of the T-box gene optomotor-blind (omb), a decapentaplegic target gene. The wings of optomotor-blind mutants have two apparently opposite phenotypes: the central wing is severely reduced and shows massive cell death, mainly in the distal-most wing, and the lateral wing shows extra cell proliferation. Here, we present genetic evidence that optomotor-blind is required to establish the graded expression of the decapentaplegic type I receptor encoded by the gene thick veins (tkv) to repress the expression of the gene master of thick veins and also to activate the expression of spalt (sal) and vestigial (vg), two decapentaplegic target genes. optomotor-blind plays a role in wing development downstream of decapentaplegic by controlling the expression of its receptor thick veins and by mediating the activation of target genes required for the correct development of the wing. The lack of optomotor-blind produces massive cell death in its expression domain, which leads to the mis-activation of the Notch pathway and the overproliferation of lateral wing cells.

Animals↗
PubMed1995

Homeotic genes and the regulation and evolution of insect wing number.

The evolution of wings catalysed the radiation of insects which make up some 75 per cent of known animals. Fossil evidence suggests that wings evolved from a segment of the leg and that early pterygotes bore wings on all thoracic and abdominal segments. The pterygote body plan subsequently diverged producing orders bearing three, two or just one pair of thoracic wings. We have investigated the role of homeotic genes in pterygote evolution by examining their function in Drosophila wing development and their expression in a primitive apterygote. Wing formation is not promoted by any homeotic gene, but is repressed in different segments by different homeotic genes. We suggest here that wings first arose without any homeotic gene involvement in an ancestor with a homeotic 'groundplan' similar to modern winged insects and that wing formation subsequently fell under the negative control of individual homeotic genes at different stages of pterygote evolution.

Animals↗
PubMed2003

Loss and recovery of wings in stick insects.

The evolution of wings was the central adaptation allowing insects to escape predators, exploit scattered resources, and disperse into new niches, resulting in radiations into vast numbers of species. Despite the presumed evolutionary advantages associated with full-sized wings (macroptery), nearly all pterygote (winged) orders have many partially winged (brachypterous) or wingless (apterous) lineages, and some entire orders are secondarily wingless (for example, fleas, lice, grylloblattids and mantophasmatids), with about 5% of extant pterygote species being flightless. Thousands of independent transitions from a winged form to winglessness have occurred during the course of insect evolution; however, an evolutionary reversal from a flightless to a volant form has never been demonstrated clearly for any pterygote lineage. Such a reversal is considered highly unlikely because complex interactions between nerves, muscles, sclerites and wing foils are required to accommodate flight. Here we show that stick insects (order Phasmatodea) diversified as wingless insects and that wings were derived secondarily, perhaps on many occasions. These results suggest that wing developmental pathways are conserved in wingless phasmids, and that 're-evolution' of wings has had an unrecognized role in insect diversification.

Animals↗
PubMed2000

DER signaling restricts the boundaries of the wing field during Drosophila development.

Arthropod and vertebrate limbs develop from secondary embryonic fields. In insects, the wing imaginal disk is subdivided early in development into the wing and notum subfields. The activity of the Wingless protein is fundamental for this subdivision and seems to be the first element of the hierarchy of regulatory genes promoting wing formation. Drosophila epidermal growth factor receptor (DER) signaling has many functions in fly development. Here we show that antagonizing DER signaling during the second larval instar leads to notum to wing transformations and wing mirror-image duplications. DER signaling is necessary for confining the wing subregion in the developing wing disk and for the specification of posterior identity. To do so, DER signaling acts by restricting the expression of Wingless to the dorsal-posterior quadrant of wing discs, suppressing wing-organizing activities, and by cooperating in the maintenance of Engrailed expression in posterior compartment cells.

Animals↗
PubMed2005

The importance of antennae for pea aphid wing induction in the presence of natural enemies.

The pea aphid Acyrthosiphon pisum Harris has been shown to produce an increasing proportion of winged morphs among its offspring when exposed to natural enemies, in particular hoverfly larvae, lacewing larvae, adult and larval ladybirds and aphidiid parasitoids. While these results suggest that wing induction in the presence of predators and parasitoids is a general response of the pea aphid, the cues and mechanisms underlying this response are still unclear. Tactile stimuli and the perception of chemical signals as well as visual signals are candidates for suitable cues in the presence of natural enemies. In this paper the hypothesis that the aphids' antennae are crucial for the wing induction in the presence of natural enemies is tested. Antennae of pea aphids were ablated and morph production was scored when aphids were reared either in the presence or the absence of predatory lacewing larvae over a six-day period. Ablation of antennae resulted in a drastic drop in the proportion of winged morphs among the offspring, both in the presence and the absence of a predator whereas predator presence increased wing induction in aphids with intact antennae, as reported in previous experiments. The results show that antennae are necessary for wing induction in the presence of natural enemies. Critical re-examination of early work on the importance of aphid antennae and tactile stimuli for wing induction suggests that a combination of tactile and chemical cues is likely to be involved not only in predator-induced wing formation but also for wing induction in response to factors such as crowding in the aphid colony.

Analysis of Variance↗
PubMed2000

Quantitative trait loci affecting components of wing shape in Drosophila melanogaster.

Two composite multiple regression-interval mapping analyses were performed to identify candidate quantitative trait loci (QTL) affecting components of wing shape in Drosophila melanogaster defined by eight relative warp-based measures. A recombinant inbred line design was used to map QTL for the shape of two intervein regions in the anterior compartment of the wing, using a high resolution map of retrotransposon insertion sites between Oregon-R and Russian 2b. A total of 35 QTL representing up to 23 different loci were identified, many of which are located near components of the epidermal growth factor-Ras signal transduction pathway that regulates vein vs. intervein decision making and vein placement. Over one-half of the loci were detected in both sexes, and just under one-half were detected at two different growth temperatures. Different loci were found to affect aspects of shape in each intervein region, confirming that the shape of the whole wing should be regarded as a compound trait composed of several developmental units. In addition, a reciprocal backcross design was used to map QTL affecting shape in the posterior compartment of the wings of 831 flies, using a molecular map of 16 allele-specific oligohybridization single nucleotide polymorphism (SNP) markers between two divergent inbred lines. A total of 13 QTL were detected and shown to have generally additive effects on separable components of shape, in both sexes. By contrast, 8 QTL that affected wing size in these backcrosses were nearly dominant in their effects. The results confirm at the genetic level that wing shape is regulated independent of wing size and set up the hypothesis that wing shape is regulated in part through the regulation of the length and positioning of wing veins, involving quantitative regulation of the activity of secreted growth factors.

Animals↗
PubMed2004

DrawWing, a program for numerical description of insect wings.

There is usually a pattern of veins on an insect wing. This pattern is species-specific and is used taxonomically. For example, the coordinates of some characteristic points on the wing are used to compare vein patterns. The characteristic points are often vein junctions or vein ends. A tool is presented that enables automatic identification of vein junctions. An image of an insect wing is used to determine the wing outline and veins. The vein skeleton is obtained using a thinning algorithm. Bezier splines are fitted to both the wing outline and the vein skeleton. The splines are saved in an encapsulated postscript file. Another output file in text format contains the coordinates of vein junctions. Both the program and its source code are available under GNU General Public License at [www.cyf-kr.edu.pl/~rotofils/drawwing.html]. The program presented in this paper automatically provides a numerical description of an insect wing. It converts an image of an insect wing to a list of coordinates of vein junctions, and a wing diagram that can be used as an illustration. Coordinates of the vein junctions extracted by the program from wing images were used successfully to discriminate between males of Dolichovespula sylvestris and Dolichovespula saxonica.

Animals↗
PubMed2005

Insect-like flapping wing mechanism based on a double spherical Scotch yoke.

We describe the rationale, concept, design and implementation of a fixed-motion (non-adjustable) mechanism for insect-like flapping wing micro air vehicles in hover, inspired by two-winged flies (Diptera). This spatial (as opposed to planar) mechanism is based on the novel idea of a double spherical Scotch yoke. The mechanism was constructed for two main purposes: (i) as a test bed for aeromechanical research on hover in flapping flight, and (ii) as a precursor design for a future flapping wing micro air vehicle. Insects fly by oscillating (plunging) and rotating (pitching) their wings through large angles, while sweeping them forwards and backwards. During this motion the wing tip approximately traces a "figure-of-eight" or a "banana" and the wing changes the angle of attack (pitching) significantly. The kinematic and aerodynamic data from free-flying insects are sparse and uncertain, and it is not clear what aerodynamic consequences different wing motions have. Since acquiring the necessary kinematic and dynamic data from biological experiments remains a challenge, a synthetic, controlled study of insect-like flapping is not only of engineering value, but also of biological relevance. Micro air vehicles are defined as flying vehicles approximately 150 mm in size (hand-held), weighing 50-100g, and are developed to reconnoitre in confined spaces (inside buildings, tunnels, etc.). For this application, insect-like flapping wings are an attractive solution and hence the need to realize the functionality of insect flight by engineering means. Since the semi-span of the insect wing is constant, the kinematics are spatial; in fact, an approximate figure-of-eight/banana is traced on a sphere. Hence a natural mechanism implementing such kinematics should be (i) spherical and (ii) generate mathematically convenient curves expressing the figure-of-eight/banana shape. The double spherical Scotch yoke design has property (i) by definition and achieves (ii) by tracing spherical Lissajous curves.

Air↗
PubMed2004

Swift laboratory thermal evolution of wing shape (but not size) in Drosophila subobscura and its relationship with chromosomal inversion polymorphism.

Latitudinal clinal variation in wing size and shape has evolved in North American populations of Drosophila subobscura within about 20 years since colonization. While the size cline is consistent to that found in original European populations (and globally in other Drosophila species), different parts of the wing have evolved on the two continents. This clearly suggests that 'chance and necessity' are simultaneously playing their roles in the process of adaptation. We report here rapid and consistent thermal evolution of wing shape (but not size) that apparently is at odds with that suggestion. Three replicated populations of D. subobscura derived from an outbred stock at Puerto Montt (Chile) were kept at each of three temperatures (13, 18 and 22 degrees C) for 1 year and have diverged for 27 generations at most. We used the methods of geometric morphometrics to study wing shape variation in both females and males from the thermal stocks, and rates of genetic divergence for wing shape were found to be as fast or even faster than those previously estimated for wing size on a continental scale. These shape changes did not follow a neat linear trend with temperature, and are associated with localized shifts of particular landmarks with some differences between sexes. Wing shape variables were found to differ in response to male genetic constitution for polymorphic chromosomal inversions, which strongly suggests that changes in gene arrangement frequencies as a response to temperature underlie the correlated changes in wing shape because of gene-inversion linkage disequilibria. In fact, we also suggest that the shape cline in North America likely predated the size cline and is consistent with the quite different evolutionary rates between inversion and size clines. These findings cast strong doubts on the supposed 'unpredictability' of the geographical cline for wing traits in D. subobscura North American colonizing populations.

Analysis of Variance↗