Insect wing polyphenism enables a single genome to produce distinct wing morphs in response to environmental cues, yet its underlying cellular determinants remain elusive. Here, we perform single-cell RNA sequencing of long-winged- and short-winged-destined wing buds of Pyrrhocoris apterus and Nilaparvata lugens, identifying six conserved cell types with comparable proportions between the two morphs. RNA interference-mediated silencing of 51 marker genes indicates that wing-patterning genes En (epithelial-like cells) and bs (tracheal cells), and cell-cycle genes Anln, CycB3, and cdk1 (neuron cells), are essential for long-winged development, among which En exhibits a specific temporal requirement. Flow cytometry analysis shows that long-winged formation mainly relies on an extended duration of cell proliferation. Cross-species comparisons indicate shared wing cell identities. Our findings indicate that hemipteran short-winged morphs may evolve from ancestral long-winged forms via precise regulation of wing-patterning and cell-cycle gene expression in epithelial‑like, tracheal, and neuron cells. This provides insights into the developmental plasticity of insect tissues at single‑cell resolution.
The flight capability of Aedes aegypti underpins global spread of vector-borne diseases (e.g., dengue, Zika, chikungunya), making it a key control target. Decapentaplegic (Dpp), evolutionarily conserved in insect morphogenesis, has unclear spatiotemporal regulatory networks in Ae. aegypti wing bud/haltere development. Here, we identify a wing bud/haltere-specific long non-coding RNA (lncRNA), AAEL025449, forming a competing endogenous RNA (ceRNA) axis to regulate Dpp-mediated development. Spatially, AAEL025449 acts as a microRNA-281 (miR-281) sponge, relieving Dpp transcriptional repression in wing buds. Temporally, it tunes Dpp dynamics during critical apoptosis windows, enabling Dpp to modulate apoptotic timing via the canonical c-Jun N-terminal kinase (JNK) pathway. Disruption of this axis results in shorter wings, indirectly compromising flight capacity and reducing mating success, thereby linking molecular perturbations to fitness-related traits. Additionally, it further clarifies the key temporal-spatial nodes where Dpp regulates wing bud cell apoptosis via JNK, with the spatial node differing from conventional Dpp regulation of wing bud development and manifesting as a gradient diffusion from anterior to posterior wing bud margin during the critical apoptosis period in Ae. aegypti. Our study uncovers a novel spatiotemporal regulatory mechanism for wing bud plasticity, advancing insights into insect wing bud adaptive evolution. This ceRNA axis provides a foundation for selecting targets to suppress mosquito populations and mitigate disease transmission.
The Peking Opera hat-wing technique is an intangible cultural heritage. This study adopted a single-subject case study design to characterize the movements of this technique, facilitating its intergenerational preservation and transmission. Biomechanical analysis was conducted to investigate the left-wing toss movement in the hat-wing technique, aiming to reveal the intrinsic biomechanical mechanism of the movement and provide a basis for its inheritance and training. An infrared motion capture system was used to collect kinematic parameters of the markers, and a three-dimensional force platform was employed to gather force data from the actor's left and right feet. Three sets of data-including Center of Pressure (COP) position, head movement parameters, and knee joint angles-were analyzed to elucidate the operating mechanism of the left-wing toss movement. Based on the results, the left-wing toss movement was theoretically segmented, and the initiation principle of the movement was determined. Additionally, factors influencing the sustained movement of the left-wing toss and those required to achieve a relatively static state at the end of the movement were identified. The primary hypothesis was that the phase relationship between the L3 Z-axis displacement-time (L3 Z-T) curve and the head tilt angle-time (α-T) curve can effectively distinguish the initiation, maintenance, and termination stages of the left-wing toss movement. Quantitative analysis of five repeated trials showed high movement repeatability, with a coefficient of variation (CV) < 15% for all key kinematic metrics. Among them, the phase difference (CV = 6.0%) and peak angular velocity (CV = 10.6%) exhibited excellent repeatability (CV < 11%), and other indicators also met the repeatability requirements for single-subject case studies (CV < 15%). During the initiation stage, the peak head tilt angle reached 6° ± 0.8°, and the peak angular velocity was 29.3°/s ± 3.1°/s. Throughout the movement, the COP excursion in the anterior-posterior direction was 4.2 ± 0.5 cm, and the medial-lateral excursion was 2.1 ± 0.3 cm. These findings provide a scientific foundation for the standardized training and intergenerational preservation of the Peking Opera hat-wing technique, highlighting the value of biomechanical analysis in traditional performing arts heritage.
To address the limited understanding of the aerodynamic characteristics of bird-inspired flapping-wing aircraft across different flight phases and the unclear flow field interaction mechanisms between the wings and tail, this study performs three-dimensional numerical simulations based on a self-developed prototype using ANSYS Fluent and the overset mesh method. The aerodynamic effects of key tail parameters under different flight conditions are quantitatively evaluated, and the mechanisms of bidirectional wing-tail aerodynamic coupling are investigated. The results show that tail twist has a negligible influence on instantaneous lift and thrust during level flight, with a maximum variation of only 0.2 N, but significantly affects the overall aerodynamic moments of the aircraft. When the tail twist angle increases from 15° to 20°, the pitching moment increases by 6%. In contrast, during climbing flight, the tail pitch angle has a pronounced effect on lift and thrust, and its aerodynamic influence depends strongly on the aircraft angle of attack. At an aircraft angle of attack of 15°, the difference between the maximum and minimum cycle-averaged pitching moments reaches 0.2 N·m. Further analysis of vorticity fields and pressure distributions confirms the existence of distinct wing-tail aerodynamic coupling. The tail not only directly modifies the aerodynamic forces and moments acting on the aircraft but also alters the wing-generated flow structures, while the wing wake simultaneously influences the aerodynamic effectiveness of the tail. This bidirectional wing-tail aerodynamic coupling plays a critical role in shaping the aerodynamic response of the aircraft under different flight conditions. These findings clarify the aerodynamic roles of key tail parameters and reveal the underlying flow field interaction mechanisms across different flight phases, providing a theoretical basis for motion-parameter optimization and precise attitude control of bird-inspired flapping-wing aircraft.
In plant-pre-dispersal seed predator interactions (e.g., flowerheads-flies), insect morphological differentiation is frequently driven by divergent host plants. However, wing morphological variation among flies associated with different flowerhead species has seldom been explored in natural alpine systems. In an alpine meadow of the Qinghai-Tibet Plateau, larvae of four host-associated morphological groups of Tephritis sp. cf. femoralis (1, 2, 3, 4) feed on four asteraceous species: Saussurea nigrescens (SN), Carduus nutans (CN), Leontopodium leontopodioides (LL), and Anaphalis lactea (AL). We examined the wing size and shape of these four groups using a landmark-based geometric morphometric approach. The host plants vary significantly in capitular diameter, dry weight, and depth. Within each host-associated group, female wing sizes (length, width and centroid size) were significantly larger than those of males. Wing size differed significantly among the four female and four male groups across host plants. Female ovipositor length varied in parallel with capitulum depth across the four host plants. Significant intersexual differences in wing shape were observed in SN, LL, and AL. Wing shape also varied distinctly among groups in both sexes across host plant capitula, enabling clear discrimination of host-associated groups. Linear discriminant analysis achieved high correct classification rates for both sexes, supporting clear morphological separation among host-associated groups. In conclusion, variations in wing size, wing shape and ovipositor length were associated with host traits. Overall, our study illustrates morphological differentiation correlated with host plants in flowerhead-fly pre-dispersal seed predator interactions in the alpine meadow system.
Foldable wings significantly improve the portability of flapping-wing micro air vehicles. This study presents a novel multi-stage bio-inspired wing design, inspired by the venation pattern and folding mechanism of the hindwing of Xylotrupes dichotomus, achieving a folding ratio of up to 3.44 while ensuring reliable folding operation. A flapping-wing structure integrating linkages and a membrane is designed with a high-stiffness strategy. A systematic theoretical kinematic analysis is conducted to investigate the motion relationships among the linkages during the folding process, and a kinematic model is established to ensure stable folding operation. A one-way fluid-structure interaction analysis is subsequently performed to verify the high-stiffness assumption and to establish an aerodynamic baseline for the specific corrugation pattern induced by the folding mechanism. Results indicate that the maximum elastic deformation is approximately 5.7% of the wingspan, validating the quasi-rigid treatment. Comparative analysis with a flat-plate model further demonstrates that the corrugation has a negligible effect on aerodynamic performance. The validated high-stiffness configuration thus provides a well-characterized rigid-wing reference for future stiffness-modulated investigations.
Flapping-wing flight offers a promising solution for aerial mobility in low-density environments such as the Martian atmosphere, where conventional rotorcraft faces significant performance constraints. However, the coupled aerodynamic and structural mechanisms governing lift generation at low Reynolds numbers remain insufficiently understood. This study investigates the aeroelastic and unsteady aerodynamic behaviour of a bio-inspired flapping wing using an integrated experimental-numerical framework. High-speed imaging is employed to extract representative wing kinematics, including flapping frequency, stroke amplitude, and rotational motion. A geometrically scaled wing model is developed based on Reynolds number similitude and analysed using finite element methods to characterise its dynamic response. Aeroelastic behaviour is evaluated through modal transient simulations, while aerodynamic performance is assessed using both vortex-lattice modelling and computational fluid dynamics. The results show strong coupling between bending and torsional modes, with the structural response highly dependent on excitation frequency relative to the natural modes. Near-resonant conditions lead to amplified deformation and distinct phase relationships, while aerodynamic simulations reveal vortex-dominated lift generation. These findings provide a physics-based framework for the design and analysis of flapping-wing systems operating in low-Reynolds-number and low-density flight regimes.
Commercial applications of flying wing aircraft, such as the Flying-V considered herein, can contribute to reducing carbon and nitrogen emissions produced by the aviation sector. However, because of the lack of a tail, all flying wing aircraft have reduced controllability. For this reason, the placement and sizing of the control surfaces along the wing is a nontrivial problem. The paper focuses on solving this problem using offline handling quality simulations based on certification requirements. In different flight conditions, the aircraft must be able to perform a set of maneuvers as defined by the certification specifications. First, offline simulations calculate the minimum control authority required from the elevator, aileron, and rudder to perform each maneuver. Then, based on the global minimum for all maneuvers, the control surfaces are sized and placed along the wings. The aerodynamic model employed uses a combination of Reynolds-averaged Navier-Stokes (RANS) and vortex lattice method (VLM) simulations. The control authority of the control surfaces is estimated with VLM and VLM calibrated with RANS simulations, showing significant differences between the two.
Conventional span-morphing wings are often constrained by structural complexity, heavy weight, and discontinuous aerodynamic surface. Although flexible honeycomb and lattice structures offer lightweight solutions, they usually require external loads to maintain the deformed configuration and often exhibit limited stability under large deformation. In this study, a span-morphing wing section based on multistable honeycomb structures is proposed. The multistable honeycomb acts as the core deformation-load-bearing module, enabling multistage reversible spanwise reconfiguration through the bistable transition of cosine curved beams and the support of honeycomb structures. An equivalent nonlinear force-displacement model is derived to describe the structural response. Finite element analysis and fluid-structure interaction analysis are conducted to evaluate its mechanical and aerodynamic performance, while prototype fabrication and bidirectional morphing experiments are performed to demonstrate its functional feasibility. The results show that the proposed wing section achieves prescribed multistage state transitions, effectively regulates lift through span variation, and maintains good structural strength under typical aerodynamic loads. These findings demonstrate the potential of multistable honeycomb structures for lightweight and stable span-morphing wing design.
During morphogenesis, cell divisions are precisely regulated in space and time. The biological objectives achieved by such regulation are not fully understood. Here, by applying a newly developed lineage-reconstruction pipeline to Drosophila pupal wing, we reveal that the wing is composed of distinct cell groups that differ in division number, timing, and spatial positioning relative to wing veins. We show that the frequencies of these lineages, together with their initial cell sizes and growth profiles, converge to achieve a highly conserved average cell size. Our data further suggest that distance from veins provides spatial information that biases where distinct lineages arise, and that loss of veins caused by perturbation of EGFR signaling suppresses a specific lineage and disrupts cell-size control. Finally, our results point to a multiscale organization of division patterns, in which vein-associated spatial information is integrated with local neighbor effects in a manner that would mitigate mechanical instability within the tissue. Together, these findings delineate a cell-size control mechanism based on coordinated divisions of distinct cell groups that supports robust morphogenesis and functional tissue design.
Honey bees are commonly infected with viruses, including deformed wing virus (DWV-A and DWV-B) and sacbrood virus (SBV), which cause morphological symptoms and death in developing bees and primarily asymptomatic infections in adult bees. Co-infections occur regularly in colonies, but they have rarely been studied, especially in adult bees. In this study, we co-inoculated young adult honey bees with DWV by injection (simulating vectored transmission by Varroa) and feeding them with SBV (simulating oral transmission) before reintroducing them in colonies. Through the use of optical counters and regular sampling, we tracked their survival and behaviour, and quantified the dynamics of viral loads in treated bees as well as the expression of eight immune genes involved in honey bee anti-viral immunity. Here, we show that co-inoculations of DWV and SBV synergistically increase the virulence of DWV and conditionally promote the replication of SBV. We also show that SBV may play a role in the replication of DWV in specific contexts. Finally, our results show that immune responses in adult honey bees depend on virus genotype (i.e., DWV), their relative abundance and the pre-existing natural infections before virus injection). Together, these results confirm the existence of deleterious interactions between deformed wing virus and sacbrood virus, impacting honey bee health and colony dynamics.
Sphenoid wing meningiomas (SWMs) are located adjacent to critical organs of interest (OOIs) and present challenges for surgery and radiotherapy. Given the favorable overall survival in meningioma patients, preserving quality of life is important. Dose reductions within OOIs may lower adverse events. This study aimed to evaluate potential advantages of particle therapy regarding dose distribution and side effects for SWMs. Nine patients (eight females, one male; median age at radiotherapy: 55 years) with SWM received proton radiotherapy (PRT, 54 Gy (RBE), 1.8 Gy (RBE) per fraction). Comparative treatment plans were generated for volumetric modulated arc therapy (VMAT, 54 Gy, 1.8 Gy per fraction) and carbon ion radiotherapy (CIRT, 42 Gy (RBE), 3 Gy (RBE) per fraction). Target volumes and OOI dose guidance were maintained. OOI dose-volume parameters, normal tissue complication probabilities (NTCP), and risk ratios of radiation-induced secondary central nervous system (CNS) malignancies were assessed. Compared to VMAT, mean relative brain doses were reduced by -41.5% with PRT, and - 63.8% with CIRT. Particle therapy achieved significant dose sparing of bilateral hippocampi and lenses, and the ipsilateral inner ear. NTCPs indicated lower risks of ipsilateral hearing loss and cataract with particle therapy. The mean estimated risk of radiation-induced secondary malignancies was 1.7 for photons over protons, and 2.7 for photons over carbon ions. For SWMs, particle therapy demonstrated reduced dose exposure to several OOIs, and may lower the risk of side effects compared with VMAT.
Deformed wing virus (DWV) is the most prevalent and extensively studied Apis mellifera virus. The ectoparasitic mite Varroa destructor strongly influences DWV epidemiology and is associated with high colony mortality worldwide. In this study, the effectiveness of an organic (oxalic acid) and a chemical (amitraz) acaricide in reducing DWV load was evaluated. Twelve naturally infested colonies were divided into three groups: oxalic acid, amitraz, and untreated control. Varroa infestation levels and DWV titres were monitored every 1-1.5 months over an 18-month period, with DWV quantified using RT-qPCR. Both acaricides showed high efficacy (>90% efficacy); however, no significant reduction in viral load was observed during the acaricide treatment period. A general decline in viral levels was observed in all experimental hives during winter; however, untreated colonies exhibited higher levels, suggesting a subclinical infection. The re-emergence of varroa in the spring was followed by DWV titre increase in all groups. A positive correlation between mite infestation and DWV titres in adult bees was observed. These results highlight the complex DWV-varroa relationship and emphasize that continuous mite monitoring and control are essential to mitigate viral impacts and support colony health.
Debate regarding the value of HIV screening in English colposcopy services is ongoing. With no national guidance provided by the NHSCSP, few colposcopy services offer HIV testing. Sheffield colposcopy introduced standardised HIV testing in January 2023. We assessed prevalence rates and acceptability to HIV testing in our referred population with moderate or worse dyskaryosis, CIN2 or worse. A retrospective service evaluation with interval analysis was conducted on 1st January to 31st December 2023 at the Jessop Wing Colposcopy Service, Sheffield, UK. All patients with moderate dyskaryosis or worse on referral cytology, CIN2 or worse on biopsy, were offered an HIV test. Of 367 eligible patients, six (1.6%) were known HIV positive. HIV prevalence rates in patients referred with moderate or worse cytology (16.34 per 1000) (95% confidence interval 6-35), and CIN2 or worse (12.46 per 1000) (95% confidence interval 3-28) were high in comparison with overall prevalence rates for Sheffield (2.14 per 1000). HIV testing was offered to 296 (82%) patients and accepted by 246 (83%). All HIV test results were reported as not detected. Sixty-five (18%) were not offered a test. Consultant colposcopists (p < 0.0001) were less likely to offer an HIV test as compared to colposcopy nurses. Acceptance of HIV testing in patients attending colposcopy is high, but willingness to offer the test varies at both local and national levels. Whilst there were no new HIV diagnoses in our sampled cohort, knowledge of HIV status is a key prognostic indicator and crucial for determining ongoing management. National societies must join forces and provide national guidance for the colposcopy community regarding HIV testing in colposcopy services. Audit registration number: STH 12112.
Superior orbital fissure syndrome (SOFS) is a rare, vision-threatening condition caused by injury or compression of cranial nerves III, IV, and VI. We report a 16-year-old male who sustained extensive craniofacial trauma, including a displaced left sphenoid wing fracture, after a motorized scooter accident. Initially, he had limited extraocular motility, but by hospital day 5, he developed complete ophthalmoplegia due to progressive nerve compression. Urgent surgical decompression of the superior orbital fissure and reduction of the fracture were performed via a pterional craniotomy on day 7. Postoperatively, the patient experienced full restoration of ocular motility and pupillary function, with no complications. This case highlights the potential for delayed neurological deterioration in sphenoid wing fractures and emphasizes the importance of close serial examination and timely operative intervention once SOFS develops, demonstrating that even delayed intervention can result in complete neurological recovery.
Organic molecular resistive memory offers a promising platform to overcome the von Neumann bottleneck. Here, we report four symmetric azobenzene-based small molecules with diverse terminal substituents (nitroimidazole, imidazole, carbazole, and triphenylamine) for memristive applications. By tuning the terminal groups, the devices exhibit tunable nonvolatile behaviors-ranging from ternary/binary WORM to bipolar nonvolatile resistive memory-all featuring high ON/OFF ratios, low operating voltages, and excellent stability. Mechanistic studies reveal that charge-transfer-induced conformational changes govern the twisted intramolecular charge-transfer states, dictating these distinct memory characteristics. Notably, the Cz-methylene-Azo memristor demonstrates continuous conductance tunability and essential synaptic functions (e.g., excitatory postsynaptic current (EPSC), paired-pulse facilitation (PPF), long-term potentiation/depression (LTP/D)). Furthermore, it serves as a versatile logic-in-memory unit, executing multiple logic gates (OR, AND, XOR, NAND, etc.), the half and full-adder circuits. Its applicability for in-memory computing is successfully validated via convolutional neural networks (CNN)-based image edge detection, highlighting its great potential for next-generation integrated organic neuromorphic architectures.
This study aims to compare the complication rate, local recurrence and survival of different types of pelvic resections. Special interest is to compare the results from external and internal hemipelvectomies if the iliac wing resections are separately evaluated, as they are usually easier to operate and do not need any reconstruction, therefore they might disfigure the results from internal hemipelvectomy in positive direction, if involved in the same group. This non-randomized retrospective study included 75 cases with a mean follow-up time of 6.9 years. We divided them to three groups according to the type of surgery (external and internal hemipelvectomy, iliac wing resection). The oncological stages, surgical margins, local recurrences, complications and the survival rates were recorded for statistical analyses. A wide surgical margin (R0) was achieved in 73.3% after external hemipelvectomy, 47.6% after internal hemipelvectomy and 77.8% after iliac wing resection. We did not find significant differences in rates of local recurrence between the groups: the lowest was recorded after external hemipelvectomy (26.7%) and there was no difference between internal hemipelvectomy and iliac wing resection (33.3%). The 5‑year survival was 26.7% in external hemipelvectomy, 35.7% in internal hemipelvectomy and 61.1% for iliac wing resection. Cox regression analysis identified negative prognostic factors for survival as histological grade, local recurrence and patient's age but not the type of surgery. Internal hemipelvectomy is as safe a procedure as the external hemipelvectomy concerning 5‑year survival, complications and local recurrence, even if iliac wing resections are excluded from this group.
AbstractThe energy needed for insect flight varies with the size and body proportions of different species or individuals. Wingbeat frequency during flight usually decreases as size increases, though these changes depend on the proportions of the wings and flight muscles. Metabolic rate during flight often changes with wingbeat frequency. Nutrition during development plays a key role in determining an individual's size and body proportions. We used an intermittent feeding regime during the fifth larval instar of the hawk moth (Manduca sexta) to investigate how nutrition-induced size variation affects flight capacity and energetics. Adult body mass decreased with more severe intermittent feeding, widening the range of body mass variation to nearly fivefold. Changes in body mass influenced body proportions through allometric effects, with wing area and thorax mass scaling hypoallometrically, while abdomen mass scaled hyperallometrically. Sex and nutritional treatment further influenced these relationships. The flight wingbeat frequency increased in smaller-sized individuals, and the thorax mass-specific metabolic rate was higher. Smaller individuals from the intermittent feeding treatment also showed a lower flight success, apparently due to a lower thorax-to-abdomen mass ratio. Flight muscle metabolic enzyme activities did not differ with size or nutritional treatment. Overall, changes in body mass and proportions resulting from developmental plasticity influence flight energetics, leading to compromises in flight energy demands and success without notably altering flight muscle metabolic phenotypes, which could restrict muscle power during flight.
Climate change is a growing threat to global biodiversity, and so there is a pressing need to understand which traits impact species vulnerability, and the capacity of these traits to respond to environmental change. I quantify two thermal traits (thermoregulation, thermal tolerance) and functional traits (wing length, colouration, wing condition, sex) of butterflies across an elevational gradient in central Europe to investigate adaptation to local climatic conditions. There is no evidence of intraspecific variation in thermal traits, suggesting that these may be fixed within species. At low elevations, large species are better than small species at avoiding high body temperatures. Dark species have improved thermoregulatory capacities with decreasing elevation but ultimately have consistently higher body temperatures than pale species. This implies that small and dark species may be particularly vulnerable to extreme heat. I subsequently detect shifts from small dark species at high elevations to large pale species at low elevations. Finally, species with poor thermoregulatory capacity or larger wings have shown greater upwards elevational range shifts. This argues that butterflies are responding to climate change through redistribution of species rather than adaptation in place, and that thermoregulatory performance will be a key driver of ecological responses to climate change.
Unmanned aerial vehicles (UAVs) are increasingly used in precision pest management, yet their performance in operational forest settings remains underexplored. We evaluated the efficacy of SPLAT® SM-O mating disruptant applied using a UAV at a dosage of 14.8 for control of the spongy moth, Lymantria dispar dispar L. (Lepidoptera: Erebidae). One treatment plot received 11.4 g AI/ha because of a calibration deviation during application. Both treatments reduced trap catches by >90% for 10 weeks following the application, meeting the efficacy requirement set by the USDA's National Slow the Spread (STS) Program. One year after the application, trap catches continued to be reduced by 28% and 67% in plots treated with 14.8 and 11.4 g AI/ha, respectively. These levels of trap catch reduction in the year of treatment and one year after the treatment application are comparable to those reported following fixed-wing aerial treatments. These results indicate that UAV-applied SPLAT® SM-O meets STS requirements for operational use and is suitable for integration into the program for treating small or isolated blocks. These findings also have broader implications for the use of unmanned aerial vehicles to deploy SPLAT® formulations in forest pest management programs.